Veerasamy Sejian · Surinder Singh Chauhan ·

Chinnasamy Devaraj · Pradeep Kumar Malik ·

Raghavendra Bhatta Editors

Climate Change
and Livestock
Production: Recent
Advances and
Future Perspectives
Climate Change and Livestock Production:
Recent Advances and Future Perspectives
Veerasamy Sejian • Surinder Singh Chauhan •
Chinnasamy Devaraj • Pradeep Kumar Malik •
Raghavendra Bhatta
Editors

Climate Change
and Livestock Production:
Recent Advances
and Future Perspectives
Editors
Veerasamy Sejian Surinder Singh Chauhan
ICAR-National Institute of Animal Faculty of Veterinary and Agricultural Science
Nutrition and Physiology The University of Melbourne
Bangalore, India Parkville, Australia

Chinnasamy Devaraj Pradeep Kumar Malik

Centre for Climate Resilient Animal ICAR-National Institute of Animal Nutrition
Adaptation Studies, ICAR-National and Physiology
Institute of Animal Nutrition and Bangalore, India
Physiology
Bangalore, India

Raghavendra Bhatta
ICAR-National Institute of Animal
Nutrition and Physiology
Bangalore, India

ISBN 978-981-16-9835-4 ISBN 978-981-16-9836-1 (eBook)

https://doi.org/10.1007/978-981-16-9836-1

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Foreword

Livestock farming and allied practices provide a source of income to a major

proportion of people across the globe and contribute to the global agricultural
gross domestic product. Livestock are of prime importance for ensuring food
security and serve as the preferred protein source for the growing human population.
Climate change is a global challenge and its impact on the livestock has attracted the
attention of policymakers to work on suitable strategies and also to fund research in
this area. It is therefore necessary to propagate relevant information to all the climate
change researchers across the globe, thereby enabling the adoption of a multidisci-
plinary and multi-institutional collaborative approach to tackle climate change and
sustain livestock production.
I am informed that a workshop is being organized under the project, “Climate
Change and Livestock Production: Current Scenario and Way Forward” at ICAR-
National Institute of Animal Nutrition and Physiology, India, in association with the
University of Melbourne, Australia, to disseminate knowledge on various
established concepts of climate change impacts on livestock production and also to
inform the participants on the various advances in this field.
The organizers are bringing out the proceedings of this workshop in the form of a
book entitled “Climate Change and Livestock Production: Recent Advances and
Future Perspective.” I hope that the book will serve as resource material for the
policymakers to implement new policies to sustain future livestock production in the
changing climate scenario. It is expected that researchers, including students,
teachers, and other practitioners in the area will greatly benefit from the deep insights
provided by experienced authors in this book.

Department of Agricultural Research Trilochan Mohapatra

and Education, Indian Council of Agricultural
Research, Government of India,
Ministry of Agriculture and Farmers Welfare
New Delhi, India

v
Preface

Climate change is no longer a myth; its causes, outcome, and projected trends have
brought the entire globe to recognize this as a major concern. The rising global
surface temperatures, increasing incidences of environmental hazards, drought,
disease outbreaks, pandemics, severe economic losses, and future projections have
made it very essential to address this issue on priority. Livestock production is one
such area gaining immense importance for its significant role in supporting the
global agricultural gross domestic product, human livelihood, and food security.
This sector is also of huge relevance from a climate change perspective for its
significant impact on climate change through greenhouse gas emissions. Hence
there is an increasing interest for researchers in this subject.
Climate change and livestock production can be considered as a field of science
that is being established with various concepts and subsections. While some of the
classical concepts of impact of climate change on livestock production are well
established, there are also increasing advancements in this field of science.
Dissipating such knowledge among the widely spread scientific fraternity would
further enhance in the adoption, validation, improvisation, and further propagation
of the knowledge. This forms the basis of this volume entitled “Climate Change
and Livestock Production: Recent Advances and Future Perspectives.”
We believe that this volume serves as a mini-encyclopedia containing most of the
critical concepts of climate change and livestock production with the latest informa-
tion. Furthermore, it can also serve as a preamble for all the students, researchers,
scientists, and policymakers across the globe who are either established or beginners
in the field of climate change and livestock production. In addition, this volume also
contains classic reference of world-class standard involving the contributions from
nearly six scientists who have been listed in the recently published database on the
world’s top 2% scientists, released by Stanford University, USA. Moreover, the
authors of the chapters of this volume are distributed across the globe covering many
continents such as Asia, Oceania, Europe, Africa, and South America. The volume is
an exemplary compilation of 23 chapters which can be broadly categorized into six
parts. The first part on impact of climate change on livestock will introduce the
readers to an overview on varied sectors of livestock production that is severely
impacted due to climate change. This is followed by a list of chapters addressing the
impact of heat stress on different adaptive responses in livestock. From there, the
volume elevates to part III which lists out some of the methods to quantify heat stress
vii
viii Preface

response in livestock. Though there are a number of established methodologies to

assess impact of climate change on livestock production, this part brings information
on some of the recent advances in this field.
The volume also comprises a list of chapters covered in part IV that would
especially attract the varied policymakers as it contains the strategies to ameliorate
heat stress impacts in livestock. As mentioned earlier, livestock not only gets
severely affected due to climate change but also contributes to it. This unique
volume also has gathered some chapters by established researchers on the topic
pertaining to enteric methane emission and amelioration which is covered under part
V of this volume. Lastly, reserving the best for the last, eminent researchers have
channelized their vast experience to compile chapters pertaining to the adaptation
strategies and future perspectives of climate change and livestock production. The
chapters under this part have great promise to be of interest to a wide range of
readers, beginning from students to young budding researchers, established
scientists from multiple disciplines, and also the ultimate policymakers. Reference
materials pertaining to climate change and livestock production are scanty and
obsolete. Therefore, by addressing systematically and comprehensively all major
aspects of climate change and livestock production particularly concentrating on
some of the recent advances and future perspectives, this volume is a useful resource
material in understanding the various intricacies in this field of science.
The contributors of various chapters are world-class professionals with vast
experience in the chosen field supported by several peer-reviewed publications.
The Editorial Committee takes this opportunity to thank all the contributors from
different parts of the world for their dedication in drafting, timely submission, and
for sharing their rich knowledge and experience with others. The efforts of many
others, all of those cannot be individually listed, were also very pertinent in com-
pleting this relevant and important volume.

Bangalore, India Veerasamy Sejian

Parkville, Australia Surinder Singh Chauhan
Bangalore, India Chinnasamy Devaraj
Bangalore, India Pradeep Kumar Malik
Bangalore, India Raghavendra Bhatta
Contents

Part I Impact of Climate Change on Livestock

1 Impact of Climate Change on Animal Production and Welfare . . . . 3
Veerasamy Sejian, M. V. Silpa, Chinnasamy Devaraj, S. Trivedi,
P. Ezhil Vadhana, W. Ruban, R. U. Suganthi, A. Manimaran,
V. P. Maurya, and Raghavendra Bhatta
2 Effects of Heat Stress and Climate Change Induced Bushfires
on Beef Meat Quality . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Robyn D. Warner, Melindee Hastie, Paula A. Gonzalez-Rivas,
Surinder S. Chauhan, Minh Ha, Caitlin Pfeiffer, Allison Hilman,
and Brendan Cowled

Part II Heat Stress Impact on Adaptive Responses in Livestock

3 Adaptation of Beef Cattle to Heat Stress Challenges . . . . . . . . . . . . 29
J. B. Gaughan, A. M. Lees, and J. C. Lees
4 Behavioural Responses of Domestic Animals for Adapting to
Thermal Stress . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Vinícius F. C. Fonsêca, Edilson P. Saraiva, José D. C. dos Santos,
Larissa Kellen da Cunha Morais, Sheila T. Nascimento,
Cíntia C. de Melo Costa, Gustavo B. Moura,
Geni Caetano Xavier Neta, Kênia C. Bícego, Veerasamy Sejian,
Marcos Chiquitelli Neto, and Alex S. C. Maia
5 Heat Stress Associated Changes in the Immune System Related
Responses in Sheep . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
Mariangela Caroprese, Maria Giovanna Ciliberti, Marzia Albenzio,
and Agostino Sevi
6 Climate Change and Livestock Production: Significance
of Studying Multiple Stressors Impact in Cattle in the Changing
Climate Scenario . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
C. G. Shashank, M. V. Silpa, P. Ezhil Vadhana, Chinnasamy Devaraj,
M. Tiwari, and Veerasamy Sejian

ix
x Contents

7 Comparative Assessment of Thermo-Tolerance of Crossbred

and Indigenous Cattle Breeds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Gyanendra Singh, Hari Abdul Samad, K. Karthiga, K. Priyanka,
Lipika Sarma, Vikrant S. Chouhan, and V. P. Maurya

Part III Methods to Quantify Heat Stress Response in Livestock

8 Non-Invasive Methods to Quantify the Heat Stress Response
in Dairy Cattle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
Gundula Hoffmann, Mullakkalparambil Velayudhan Silpa,
Roman Mylostyvyi, and Veerasamy Sejian
9 Applications of Infrared Thermal Imaging and Rumen Boluses
for Quantifying Heat Stress in Cattle . . . . . . . . . . . . . . . . . . . . . . . 99
A. M. Lees, A. L. Wallage, L. Labeur, S. L. Sammes, J. C. Lees,
and J. B. Gaughan
10 Skin Based Novel Approaches for Establishing Climate Resilience
in Goats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113
M. V. Silpa, Veerasamy Sejian, S. Koenig, Chinnasamy Devaraj,
C. G. Shashank, A. P. Kolte, G. B. Manjunathareddy,
and Raghavendra Bhatta

Part IV Strategies to Ameliorate Heat Stress Impacts

11 Livestock Shelter Management: Climate Change Perspective . . . . . 129
Ravindra Kumar, Ankaj Thakur, Rakesh Thakur, and P. K. Dogra
12 Nutritional Amelioration of Thermal Stress Impacts in Dairy
Cows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
Frank R. Dunshea, Brian J. Leury, Kristy DiGiacomo,
Jeremy J. Cottrell, and Surinder Singh Chauhan
13 The Role of Napier Grass (Pennisetum purpureum Schumach)
for Improving Ruminant Production Efficiency and Human
Nutrition in the Tropics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151
M. R. Islam, S. C. Garcia, N. R. Sarker, B. K. Roy, N. Sultana,
and C. E. F. Clark
14 Strategies to Ameliorate Heat Stress Impacts in Sheep . . . . . . . . . . 161
Kristy DiGiacomo, Surinder Singh Chauhan, Frank R. Dunshea,
and Brian J. Leury
15 Strategies to Ameliorate Heat Stress Effects on Sheep
Reproduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175
Susan Robertson and Michael Friend
Contents xi

16 Applications of Genetic Selection in Breeding for

Thermo-Tolerance in Livestock . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185
Surinder Singh Chauhan, Richard Osei-Amponsah,
Veerasamy Sejian, and Frank R. Dunshea

Part V Enteric Methane Emission and Amelioration

17 Enteric Methane Mitigation in Livestock: An Indian
Perspective . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
Raghavendra Bhatta, Pradeep Kumar Malik, and Veerasamy Sejian
18 Heat Stress on the Rumen Fermentation and Its Consequence . . . . 213
Yutaka Uyeno
19 Comparative Assessment of Rumen Microbial Diversity in Cattle
and Buffaloes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 223
Pradeep Kumar Malik, S. Trivedi, A. P. Kolte, Veerasamy Sejian,
A. Mech, S. Biswas, A. V. K. Bhattar, A. Mohapatra,
and Raghavendra Bhatta
20 Metagenomics Application in Understanding Rumen Functions . . . 239
S. Trivedi, A. P. Kolte, Pradeep Kumar Malik, Veerasamy Sejian,
A. Mech, and Raghavendra Bhatta

Part VI Adaptation Strategies and Future Perspectives

21 Genetic Adaptation of Livestock to Heat Stress Challenges . . . . . . . 263
Concepta McManus, Andrea Queiroz Maranhão, Daniel Pimentel,
Felipe Pimentel, and Marcelo de Macedo Brigido
22 Goat as the Ideal Future Climate Resilient Animal Model . . . . . . . . 279
Veerasamy Sejian, M. V. Silpa, M. R. Reshma Nair,
Chinnasamy Devaraj, A. Devapriya, N. Ramachandran,
V. B. Awachat, C. G. Shashank, Vinícius F. C. Fonsêca,
and Raghavendra Bhatta
23 Future Vision for Climate Change Associated Livestock
Production . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 293
Veerasamy Sejian, Surinder Singh Chauhan, Chinnasamy Devaraj,
Pradeep Kumar Malik, E. Vadhana, M. V. Silpa, C. G. Shashank,
and Raghavendra Bhatta

Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 307
Editors and Contributors

About the Editors

Veerasamy Sejian MVSc, PhD, is a Principal Scientist at ICAR-National Institute

of Animal Nutrition and Physiology (NIANP), Bangalore, India. His central thrust
area of research is on “Climate Change and Livestock Production.” He is the team
leader in establishing the concept of “Multiple Stresses Impacting Small
Ruminants.” His current research focuses on identifying molecular markers for
different environmental stresses in small ruminants with the primary focus to
develop agroecological zone-specific thermotolerant breeds. He has published
more than 160 peer-reviewed articles, 99 book chapters, 285 invited/lead papers,
175 conference papers, and 22 technical manuals.
Further, he has published three International Springer books. He did his first
postdoctorate at The Ohio State University, USA. He was bestowed with Endeavour
Research Fellowship by the Australian Government to pursue a postdoctorate at The
University of Queensland, Australia. For his outstanding contribution to animal
science, the Indian Council of Agricultural Research (ICAR) has bestowed him
with the prestigious Lal Bahadur Shastri Outstanding Young Scientist Award in
2012. Further, Career360, an academic organization in India, had selected him to be
one of the top 10 scientists in India under the environmental sciences category
during 2017–2018. He is also listed in the world’s top 2% scientists by Stanford
University, USA, during 2020 and 2021. He is also listed in the world’s top 1% of
scientists by PubMed based on the last 10 years of publications in heat stress. In
addition, he is also serving as Field Editor in Springer’s International Journal of
Biometeorology, Associate Editor in Elsevier’s Small Ruminants Research, and
Academic Editor in PLOS Climate. He has developed four technologies and also
has one patent granted.

Surinder Singh Chauhan DVM, PhD, is the Senior Lecturer in the Department of
Veterinary and Agricultural Sciences, The University of Melbourne, Australia, and
has 12 years of research and teaching experience in veterinary and animal sciences in
three countries, including Australia, India, and the USA. He obtained his Bachelor of
Veterinary Science and Animal Husbandry and Master of Veterinary Science from
CSK, Himachal Pradesh Agriculture University, Palampur, India. He has served the

xiii
xiv Editors and Contributors

State of Himachal Pradesh, India, for more than 10 years and has provided his
services as Senior Veterinary Officer and Veterinary Officer in various Civil Veteri-
nary Hospitals and Livestock Breeding Farms. He has also served as Assistant
Professor in The Department of Veterinary Physiology and Livestock Production
and Management, College of Veterinary and Animal Sciences at the CSK, Himachal
Pradesh Agriculture University, Palampur, India. He moved to Australia in 2011
following his selection for the prestigious Australian Leadership Award (Ph.D.
scholarship) from the Australian Govt and completed his Ph.D. under the guidance
of Prof Frank Dunshea, Faculty of Veterinary and Agricultural Sciences, The
University of Melbourne in 2014. Later, he completed his Postdoctorate in Animal
Science (Muscle Biology and Meat Science) at The Ohio State University,
Columbus, USA. He has presented his research at various national and international
conferences, including 7 keynote papers, and has published more than 70 refereed
research papers. At The University of Melbourne, he is leading the Australia–India
Collaborations for heat stress mitigation and Sustainable Livestock production
funded by the Australian Government Australia–India Council.

Chinnasamy Devaraj MVSc, PhD, is a Scientist at the Division of Bioenergetics

and Environmental Sciences, ICAR-National Institute of Animal Nutrition and
Physiology (NIANP), Bangalore, India. His central thrust area of research is on
“Energy Metabolism, Environmental Physiology, and Livestock Production.” His
current research focuses on unraveling the physiological role of adipokines
associated with energy metabolism in small ruminants. The Indian Council of
Agricultural Research (ICAR) has awarded Junior Research Fellowship based on
All India entrance for pursuing M.V.Sc program (2004–2006) at National Dairy
Research Institute, Karnal, India. He has also received ICAR Award for Outstanding
Interdisciplinary Team Research in Agricultural and Allied Sciences (Animal and
Fisheries Sciences) for 2007–2008. He has published more than 160 research
articles, including 25 peer-reviewed articles, 5 book chapters, 45 invited/lead papers,
74 conference papers, and 7 technical manuals. Presently, he is serving as a reviewer
for the International Journal of Biometeorology and Journal of Agrometeorology.

Pradeep Kumar Malik MSc, PhD, is presently working as Principal Scientist at the
ICAR-National Institute of Animal Nutrition and Physiology, Bengaluru. He was an
Assistant Professor at Navsari Agricultural University, Navsari, and Senior Scientist
at NIANP. He has accomplished International Training at the CSIRO, Queensland,
Australia, and postdoctoral fellowship under the Endeavour Research Program of the
Australian Government at the University of Queensland, Australia, in 2014–2015.
His primary research area is enteric methane measurement and amelioration in
livestock. He is handling a multicentric project on “Estimation of Methane
Emissions under Different Feeding Systems and Development of Mitigation
Strategies.” He is also associated with the ILRI-ICAR collaborative project on
“Methane Emissions and its Mitigation” and the DBT-DFG Project on “Contamina-
tion of Feed and Fodders with Heavy Metals and Agrochemicals Impact on Milk
Composition, Rumen Microbes, and Methanogenesis in Dairy Cattle along with the
Editors and Contributors xv

Rural-Urban Interface of Bengaluru.” He has developed a state-wise national inven-

tory on enteric methane emissions from Indian livestock using primary data on
methane production potential. He has created two farmers’ friendly anti-
methanogenic products, “Harit Dhara” and “Tamarin Plus,” and filed four patents.
He has published more than 50 research papers in international and national journals
of repute, two international books by CABI and Springer in livestock production and
climate change, and 40 book chapters. He also has 264 NCBI submissions in his
name. He was also the recipient of the prestigious Australian Ambassador Award in
2015 by the Australian High Commission in India.

Raghavendra Bhatta MVSc, PhD, is currently the Director of ICAR-National

Institute of Animal Nutrition and Physiology (NIANP), Bengaluru, Karnataka,
India. During his research career spanning 27 years, he has done extensive research
work involving small ruminant nutrition concerning “plant phenolics.” He has
developed the All-India state-wise inventory on enteric methane emission based
on primary data and has evolved simple, eco-friendly strategies for enteric methane
reduction in livestock. He was awarded the prestigious Japan Society for the
Promotion of Science (JSPS) postdoctoral fellowship (2004–2006) at NILGS,
Japan. He was invited to present theme papers at the Greenhouse Gases and Animal
Agriculture (GGAA) Conference at Zurich, Switzerland in 2005; GGAA 2010
conference in Banff, Canada; and GGAA 2016 conference in Melbourne,
Australia. He is the Fellow of the National Academy of Agricultural Sciences, the
National Academy of Veterinary Science, and the National Academy of Dairy
Science, India. He is the recipient of several research awards, including the presti-
gious Sir CV Raman State Award from the Karnataka State Council for Science and
Technology, Govt. of Karnataka, and the Rafi Ahmed Kidwai Award for Outstand-
ing Research in Agricultural Sciences, 2019 by ICAR. He is recognized as one of the
world’s top 2% scientists by Stanford University during 2020 and 2021. He has
published more than 100 research articles in journals and presented more than
100 papers in conferences of national and international reputation and has authored
3 books, one each in Cambridge University Press, Centre for Agriculture and
Bioscience International, and Springer. He has written 45 chapters in books and
has two patents to his credit.

List of Contributors

Marzia Albenzio Department of Agriculture, Food, Natural Resources, and Engi-

neering (DAFNE), University of Foggia, Foggia, Italy
V. B. Awachat Centre for Climate Resilient Animal Adaptation Studies, ICAR-
National Institute of Animal Nutrition and Physiology, Bangalore, India
A. V. K. Bhattar ICAR-National Institute of Animal Nutrition and Physiology,
Bangalore, India
xvi Editors and Contributors

Raghavendra Bhatta ICAR-National Institute of Animal Nutrition and Physiology,

Bangalore, India
Kênia C. Bícego Department of Animal Morphology and Physiology, Sao Paulo
State University, UNESP/FCAV, Jaboticabal, Brazil
S. Biswas ICAR-National Institute of Animal Nutrition and Physiology, Bangalore,
India
Geni Caetano Xavier Neta Department of Animal Morphology and Physiology,
Sao Paulo State University, UNESP/FCAV, Jaboticabal, Brazil
Mariangela Caroprese Department of Agriculture, Food, Natural Resources, and
Engineering (DAFNE), University of Foggia, Foggia, Italy
Surinder Singh Chauhan Faculty of Veterinary and Agricultural Science, The
University of Melbourne, Parkville, Australia
Vikrant S. Chouhan Division of Physiology and Climatology, ICAR-Indian Vet-
erinary Research Institute, Izatnagar, Uttar Pradesh, India
Maria Giovanna Ciliberti Department of Agriculture, Food, Natural Resources,
and Engineering (DAFNE), University of Foggia, Foggia, Italy
C. E. F. Clark Livestock Production and Welfare Group, School of Life and
Environmental Sciences, Faculty of Science, The University of Sydney, Camden,
NSW, Australia
Jeremy J. Cottrell School of Agriculture and Food, Faculty of Veterinary and
Agricultural Sciences, The University of Melbourne, Parkville, VIC, Australia
Brendan Cowled Ausvet, Bruce, ACT, Australia
Marcelo de Macedo Brigido Universidade de Brasília, Instituto de Ciências
Biológicas, Brasilia, DF, Brazil
Cíntia C. de Melo Costa Innovation Group of Thermal Comfort and Animal
Welfare (INOBIO-MANERA), Biometeorology Laboratory, São Paulo State Uni-
versity, Jaboticabal, Brazil
A. Devapriya Centre for Climate Resilient Animal Adaptation Studies, ICAR-
National Institute of Animal Nutrition and Physiology, Bangalore, India
Chinnasamy Devaraj Centre for Climate Resilient Animal Adaptation Studies,
ICAR-National Institute of Animal Nutrition and Physiology, Bangalore, India
Kristy DiGiacomo School of Agriculture and Food, Faculty of Veterinary and
Agricultural Sciences, The University of Melbourne, Parkville, VIC, Australia
P. K. Dogra COVAS, CSK Himachal Pradesh Agricultural University, Palampur,
India
Editors and Contributors xvii

José D. C. dos Santos Animal Biometeorology and Ethology Group (BIOET),

Department of Animal Science, Federal University of Paraiba, Areia, Brazil
Frank R. Dunshea School of Agriculture and Food, Faculty of Veterinary and
Agricultural Sciences, The University of Melbourne, Parkville, VIC, Australia
Faculty of Biological Sciences, The University of Leeds, Leeds, UK
P. Ezhil Vadhana Centre for Climate Resilient Animal Adaptation Studies, ICAR-
National Institute of Animal Nutrition and Physiology, Bangalore, India
Vinícius F. C. Fonsêca Animal Biometeorology and Ethology Group (BIOET),
Department of Animal Science, Federal University of Paraiba, Areia, Brazil
Innovation Group of Thermal Comfort and Animal Welfare (INOBIO-MANERA),
Biometeorology Laboratory, São Paulo State University, Jaboticabal, Brazil
Michael Friend Office of the Pro Vice-Chancellor (Research & Innovation),
Charles Sturt University, Wagga Wagga, NSW, Australia
S. C. Garcia Livestock Production and Welfare Group, School of Life and Envi-
ronmental Sciences, Faculty of Science, The University of Sydney, Camden, NSW,
Australia
J. B. Gaughan School of Agriculture and Food Sciences, Animal Science Group,
The University of Queensland, Gatton, QLD, Australia
Paula A. Gonzalez-Rivas Faculty of Veterinary and Agricultural Science, The
University of Melbourne, Parkville, VIC, Australia
Minh Ha Faculty of Veterinary and Agricultural Science, The University of
Melbourne, Parkville, VIC, Australia
Melindee Hastie Faculty of Veterinary and Agricultural Science, The University of
Melbourne, Parkville, VIC, Australia
Allison Hilman Ausvet, Bruce, ACT, Australia
Gundula Hoffmann Department of Engineering for Livestock Management,
Leibniz Institute for Agricultural Engineering and Bioeconomy (ATB), Potsdam,
Germany
M. R. Islam Livestock Production and Welfare Group, School of Life and Envi-
ronmental Sciences, Faculty of Science, The University of Sydney, Camden, NSW,
Australia
K. Karthiga Division of Physiology and Climatology, ICAR-Indian Veterinary
Research Institute, Izatnagar, Uttar Pradesh, India
S. Koenig Institute of Animal Breeding and Genetics, Justus-Liebig-Universität
Gießen, Giessen, Germany
A. P. Kolte ICAR-National Institute of Animal Nutrition and Physiology,
Bangalore, India
Ravindra Kumar Bihar Animal Sciences University, Patna, India
xviii Editors and Contributors

L. Labeur University of New England, School of Environmental and Rural Sci-

ence, Armidale, NSW, Australia
A. M. Lees University of New England, School of Environmental and Rural
Science, Armidale, NSW, Australia
J. C. Lees University of New England, School of Environmental and Rural Sci-
ence, Armidale, NSW, Australia
Brian J. Leury School of Agriculture and Food, Faculty of Veterinary and Agri-
cultural Sciences, The University of Melbourne, Parkville, VIC, Australia
Alex S. C. Maia Innovation Group of Thermal Comfort and Animal Welfare
(INOBIO-MANERA), Biometeorology Laboratory, São Paulo State University,
Jaboticabal, Brazil
Pradeep Kumar Malik ICAR-National Institute of Animal Nutrition and Physiol-
ogy, Bangalore, India
A. Manimaran Southern Regional Station, ICAR-National Dairy Research Insti-
tute, Bangalore, India
G. B. Manjunathareddy ICAR-National Institute of Veterinary Epidemiology and
Disease Informatics, Bangalore, India
Andrea Queiroz Maranhão Universidade de Brasília, Instituto de Ciências
Biológicas, Brasilia, DF, Brazil
V. P. Maurya Division of Physiology and Climatology, ICAR-Indian Veterinary
Research Institute, Izatnagar, Uttar Pradesh, India
Concepta McManus Universidade de Brasília, Instituto de Ciências Biológicas,
Brasilia, DF, Brazil
A. Mech ICAR-National Institute of Animal Nutrition and Physiology, Bangalore,
India
A. Mohapatra ICAR-National Institute of Animal Nutrition and Physiology,
Bangalore, India
Larissa Kellen da Cunha Morais Animal Biometeorology and Ethology Group
(BIOET), Department of Animal Science, Federal University of Paraiba, Areia,
Brazil
Gustavo B. Moura Innovation Group of Thermal Comfort and Animal Welfare
(INOBIO-MANERA), Biometeorology Laboratory, São Paulo State University,
Jaboticabal, Brazil
Roman Mylostyvyi Department of Animal Products Processing Technology,
Dnipro State Agrarian and Economic University, Dnipro, Ukraine
Sheila T. Nascimento Innovation Group of Thermal Comfort and Animal Welfare
(INOBIO - MANERA), State University of Maringá, Maringá, Brazil
Editors and Contributors xix

Marcos Chiquitelli Neto Innovation Group of Thermal Comfort and Animal

Welfare (INOBIO-MANERA), Biometeorology Laboratory, São Paulo State Uni-
versity, Jaboticabal, Brazil
Richard Osei-Amponsah Faculty of Veterinary and Agricultural Science, The
University of Melbourne, Parkville, VIC, Australia
Department of Animal Science, University of Ghana, Legon, Accra, Ghana
Caitlin Pfeiffer Faculty of Veterinary and Agricultural Science, The University of
Melbourne, Parkville, VIC, Australia
Daniel Pimentel Universidade de Brasília, Instituto de Ciências Biológicas,
Brasilia, DF, Brazil
Felipe Pimentel CEUB, Brasília, DF, Brazil
K. Priyanka Division of Physiology and Climatology, ICAR-Indian Veterinary
Research Institute, Izatnagar, Uttar Pradesh, India
N. Ramachandran Bioenergetics and Environmental Sciences Division, ICAR-
National Institute of Animal Nutrition and Physiology, Bangalore, India
M. R. Reshma Nair Centre for Climate Resilient Animal Adaptation Studies,
ICAR-National Institute of Animal Nutrition and Physiology, Bangalore, India
Susan Robertson Gulbali Institute and School of Agricultural, Environmental and
Veterinary Sciences, Charles Sturt University, Wagga Wagga, NSW, Australia
B. K. Roy Bangladesh Livestock Research Institute, Dhaka, Bangladesh
W. Ruban Department of Livestock Product Technology, Veterinary College,
Hebbal, Bangalore, Karnataka Veterinary Animal and Fishery Sciences University,
Bidar, India
Hari Abdul Samad Division of Physiology and Climatology, ICAR-Indian Veter-
inary Research Institute, Izatnagar, Uttar Pradesh, India
S. L. Sammes Feedworks Pty Ltd, Romsey, VIC, Australia
Edilson P. Saraiva Animal Biometeorology and Ethology Group (BIOET),
Department of Animal Science, Federal University of Paraiba, Areia, Brazil
N. R. Sarker Krishi Gobeshona Foundation, Dhaka, Bangladesh
Lipika Sarma Division of Physiology and Climatology, ICAR-Indian Veterinary
Research Institute, Izatnagar, Uttar Pradesh, India
Veerasamy Sejian ICAR-National Institute of Animal Nutrition and Physiology,
Bangalore, India
Agostino Sevi Department of Agriculture, Food, Natural Resources, and Engineer-
ing (DAFNE), University of Foggia, Foggia, Italy
xx Editors and Contributors

C. G. Shashank ICAR-National Dairy Research Institute, Karnal, India

Mullakkalparambil Velayudhan Silpa Institute of Animal Breeding and Genet-
ics, Justus Liebig University Giessen, Giessen, Germany
Gyanendra Singh Division of Physiology and Climatology, ICAR-Indian Veteri-
nary Research Institute, Izatnagar, Uttar Pradesh, India
R. U. Suganthi Bioenergetics and Environmental Sciences Division, ICAR-
National Institute of Animal Nutrition and Physiology, Bangalore, India
N. Sultana Bangladesh Livestock Research Institute, Dhaka, Bangladesh
Ankaj Thakur COVAS, CSK Himachal Pradesh Agricultural University,
Palampur, India
Rakesh Thakur COVAS, CSK Himachal Pradesh Agricultural University,
Palampur, India
M. Tiwari Animal Biotechnology Division, ICAR-National Dairy Research Insti-
tute, Karnal, India
S. Trivedi ICAR-National Institute of Animal Nutrition and Physiology,
Bangalore, India
Yutaka Uyeno Shinshu University, Nagano, Japan
E. Vadhana Centre for Climate Resilient Animal Adaptation Studies, ICAR-
National Institute of Animal Nutrition and Physiology, Bangalore, India
A. L. Wallage School of Agriculture and Food Sciences, Animal Science Group,
The University of Queensland, Gatton, QLD, Australia
Robyn D. Warner Faculty of Veterinary and Agricultural Science, The University
of Melbourne, Parkville, VIC, Australia
List of Figures

Fig. 3.1 Distribution of cattle around the world (Cooke et al. 2020a, b) . . . 31
Fig. 4.1 Simple illustration of the thermal load thresholds delimiting
zone of least thermoregulatory effort. LCT lower critical
temperature, UCT upper critical temperature . . . . . . . . . . . . . . . . . . . . . . . 41
Fig. 4.2 Black Santa Ines newborn lamb exposed to direct solar radiation
in an equatorial semi-arid region. Thermal image taken by using
an infrared thermal camera (Fluke—Model TiX500 9 Hz; range:
range:
20  C and +650  C; accuracy: 0.1  C/1  C; emissivity:
ε ¼ 0.98, Everett, WA, USA) at a distance of 1.0 m from the
animal. Meteorological conditions at time which image was
taken: Ta: 35  C; RH: 35.3%; RS: 900 W m2; WS: 3.7 m s
1.
(Photo by J.D.C.S.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Fig. 4.3 Newborn goats within the source of supplemental heat (SSH)
and thermal load experienced (mean  SEM) by them within
the SSH and in open area. (Photo by V.F.C.F.) . . . . . . . . . . . . . . . . . . . . 44
Fig. 4.4 Shade-seeking behaviour of hair coat sheep during the hottest
hours of the day in an equatorial semi-arid region.
(Photo by V.F.C.F) . .. . .. . .. . .. . .. . .. . .. . .. .. . .. . .. . .. . .. . .. . .. . .. . .. . . 46
Fig. 4.5 Proposed daily routine adaptation for dairy cows raised
in equatorial semi-arid regions, according to the level of solar
radiation tolerated by cows (Oliveira et al. 2014). Levels of solar
radiation (RS, W m
2) recorded in the study were divided into
five classes: Class 1 (RS < 300 W m
2), class 2
(300 W m
2 < RS  500 W m
2), class 3
(500 W m
2 < RS  700 W m
2), class 4 (RS  900 W m
2),
class 5 (RS > 900 W m
2) .. . . . .. . . . .. . . .. . . . .. . . . .. . . . .. . . . .. . . . .. . . 47
Fig. 5.1 Percentage increases calculated to stimulated PBMC (CS) of
(a) cell proliferation, (b) IL-6, (c) IL-1beta, and (d) IL-10 levels
in stimulated PBMC treated with 100 ng/mL of cortisol (100)
and stimulated PBMC treated with 1000 ng/mL of cortisol
(1000) after acute stress (24 h), chronic stress (96 h),
normothermia (39  C) and hyperthermia (43  C) in vitro

xxi
xxii List of Figures

challenge. Stimulated PBMC were treated with

Phytohemagglutinin (PHA) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Fig. 6.1 Pictorial representation of summation effect of multiple stresses
on productive functions in cattle. (Source: Modified from
Sejian et al. (2012)) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
Fig. 8.1 Daily total lying time predicted under different heat load
conditions for the reference group, lactation 4 group, and DIM
(days in milk) >150 group. Reference group has normal milk
yield level, lactation state of 1, no pregnancy and estrus
(Heinicke et al. 2019) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
Fig. 8.2 Infrared thermal images in different body parts of a dairy cow . . . 93
Fig. 9.1 An example of infrared thermal (IRT) image using the FLUKE
software, showing an infrared thermal (IRT) image and regions
of interest consisting of the flank (a) and scrotum (b) showing
the use of the polygon tool to calculate minimum, maximum
and mean temperature of the selected area as described by
Wallage et al. (2017) . .. . . .. . . .. . . .. . . .. . . .. . . . .. . . .. . . .. . . .. . . .. . . .. . . 103
Fig. 9.2 An example of infrared thermal image interpretation and
analysis using the FLIR software, displaying the arrangement
of four regions of interest across the back of a lamb, being
(1) shoulder, (2) mid loin, (3) hips and (4) rump as described
by Labeur et al. (2017) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
Fig. 9.3 Diurnal rhythm using mean hourly (h) rumen temperature
(TRUM) of (a) shaded Angus steers over 128 days as adapted
from Lees et al. (2019a); (b) un-shaded Angus (UNSH AA),
un-shaded Charolais (UNSH CH), un-shaded Brahmans
(UNSH BH), shaded Angus (SH AA), shaded Charolais
(SH CH) and shaded Brahman (SH BH) steers over 130 days
adapted from Lees et al. (2018a); and (c) for three 6 day periods
(P1, P2 and P3), adapted from Lees (2016) . .. . . . .. . . . .. . . . .. . . . .. . . 105
Fig. 9.4 (a) Relationship between body temperature and panting score
at 06:00 h (AM), 12:00 h (MID), and 16:00 h (PM) and
(b) Relationship between body temperature (BT  SE;  C)
and respiration rate (RR  SE; bpm) at panting score
values of 0, 1, 2, 2.5, 3 and 3.5. Adapted from Gaughan and
Mader (2014) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
Fig. 10.1 Overview of skin based approaches to assess climate resilience
in goats . . .. . . . . . .. . . . . . .. . . . . . . .. . . . . . .. . . . . . .. . . . . . .. . . . . . .. . . . . . .. . . . . 118
Fig. 11.1 Livestock shelter designs for different agroclimatic zones of
Himachal Pradesh . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
Fig. 11.2 Inner view of typical animal house (a) and multispecies shelter
(b) in colder areas of Himachal Pradesh . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134
Fig. 13.1 Impact of increasing protein (Milk yield, L/d/cow ¼ 0.83  CP%
of grass + 1.63, R2 ¼ 0.69) and energy (Milk yield,
L/d/cow ¼ 3.49  ME of grass + 17.69, R2 ¼ 0.48) on milk
yield of cows using Napier grass (Source: Table 13.3) . . . . . . . . . . . . 157
List of Figures xxiii

Fig. 16.1 Variation of temperament, mastitis resistance, feed saved, and

fertility GEBVs with heat tolerance GEBVs of Holstein dairy
cows . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192
Fig. 17.1 Nation wise greenhouse gas emissions (WRI 2011) . . . . . . . . . . . . . . . 199
Fig. 17.2 GHG emissions according to different sectors .. . . . .. . . .. . . . .. . . .. . . 200
Fig. 17.3 Global agricultural GHG emission . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201
Fig. 17.4 Region wise enteric methane emission (modified from O’Mara
2011) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201
Fig. 17.5 Projections for CH4 emission (2010–2030) . . . . . . . . . . . . . . . . . . . . . . . . . 202
Fig. 17.6 Projections for N2O emission (2010–2030) . . . . . .. . . . . . . . . . . .. . . . . . . 202
Fig. 17.7 Species wise % enteric methane emission (modified from
Kamra 2014) .. . . .. . . .. . . .. . . .. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . . .. . . .. . . 203
Fig. 18.1 Effect of heat stress [ambient temperature and relative
humidity (RH)] on the population of specific bacterial groups
in the rumen of Holstein heifers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
Fig. 18.2 Effect of heat stress [ambient temperature and relative humidity
(RH)] on the rumen VFA level and the composition of Holstein
heifers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
Fig. 18.3 A schematic flow about how SARA occurs due to changing
feed . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . 217
Fig. 18.4 Schematic representation of fermentation of carbohydrates and
categorization of bacteria according to their demand
for H2 utilization/production . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
Fig. 19.1 Ruminal archaea community composition in cattle and buffaloes
both at (a) order level and (b) family level (Malik et al. 2021).
The inner-circle represents the methanogens distribution in cattle;
while the outer circle represents the distribution in buffaloes . . . . . 231
Fig. 19.2 Ruminal archaea community composition at genus and species
levels in cattle and buffaloes (Malik et al. 2021). Each genus is
represented by larger bars that are underlaid on the smaller bars
representing the abundance of all the species affiliated to the
corresponding genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232
Fig. 19.3 (a) Alpha diversity and (b) beta diversity of the ruminal
methanogens in cattle and buffaloes (Malik et al. 2021) . . . . . . . . . . 232
Fig. 19.4 Ruminal archaea representing the core microbiome at 50%
minimum prevalence in (a) cattle at genus level (b) buffalo
at genus level (c) cattle at species level (d) buffaloes at species
level (Malik et al. 2021). The colour gradient indicates
variability in prevalence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233
Fig. 20.1 Functions of rumen microbiome . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 243
Fig. 20.2 Different approached used for study of rumen microbiome . . . . . . . 246
Fig. 20.3 Schematic diagram of 16S rRNA gene showing the distribution
of the conserved and hypervariable regions within the gene . . . . . . 249
xxiv List of Figures

Fig. 21.1 Upset Plot (https://gehlenborglab.shinyapps.io/upsetr/) of the

number of genes linked to climatic adaptation cited in scientific
papers per species (based on 19,609 genes and 127 scientific
papers) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 268
Fig. 21.2 Functional analysis network from 1703 genes related to heat
response in cattle, sheep, goats and horses
(https://tools.dice-database.org/GOnet/) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 269
Fig. 21.3 Chromosomal localization of 1703 genes found in published
papers on the adaptation of cattle, sheep, goats and horses
worldwide (a) and a closeup (b) of one of the enriched regions
based on cattle genomic organization (individual genes in red,
enriched regions underlined in purple - these regions were
defined by chi-square tests using all 19,906 genes found in the
literature. http://bioinformatics.sdstate.edu/go/) . . . . . . . . . . . . . . . . . . . . 269
Fig. 21.4 A hierarchical clustering tree summarising the correlation
among significant pathways from the Enrichment analysis using
Reactome (a), KEGG (b) and Gene Ontology (GO) analyses (c).
Pathways with many shared genes are clustered together
(http://bioinformatics.sdstate.edu/go/). Numbers are False
Discovery Rate (FDR). Bigger blue dots indicate more
significant P-values . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 271
Fig. 21.5 Word clouds for Gene Ontology mappings directly annotated by
the source database (a) and Tissue expression locations
(b) (https://david.ncifcrf.gov/tools.jsp) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 272
Fig. 22.1 Model projections of extremely severe heat stress to be
experienced by different livestock species by 2050 and 2100
in East Africa (source Rahimi et al. 2021; Reshma Nair
et al. 2021) . . . . .. . . . . . .. . . . . . .. . . . . . . .. . . . . . .. . . . . . .. . . . . . .. . . . . . .. . . . . . 282
Fig. 22.2 Three indigenous goat breeds of South India . . . . . . . . . . . . . . . . . . . . . . . 285
Fig. 22.3 Various concepts associated with climate resilient goat
production . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 286
Fig. 22.4 Different biomarkers for quantifying heat stress response
in goats . . .. . . . . . .. . . . . . .. . . . . . . .. . . . . . .. . . . . . .. . . . . . .. . . . . . .. . . . . . .. . . . . 288
Fig. 22.5 Different adaptation strategies to sustain goat production in the
changing climate scenario . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 289
Fig. 23.1 Future strategies to sustain livestock production in the changing
climate scenario . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 296
List of Tables

Table 8.1 Overview of different non-invasive methods and their fields

of application in dairy cows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
Table 9.1 Modified assessment of panting score and description
of breathing/panting condition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108
Table 13.1 Characteristics of tropical and subtropical countries in the
world . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154
Table 13.2 Production of cattle meat and milk by region in 2017
(Source FAOSTAT 2018) in relation to world population . . . . . . 155
Table 13.3 Current cattle production from Napier grass and potential
to increase cattle production following examples of Kikuyu
grass . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 156
Table 17.1 Ameliorative measures for enteric CH4 mitigation . . . . . . . . . . . . . . . 205
Table 17.2 Estimate and projected emissions of methane and nitrous
oxide from manure management . . . . . . . . . . . . .. . . . . . . . . . . . .. . . . . . . . . 206
Table 17.3 Precautionary/ameliorative measures for reducing GHG
emissions from manure management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Table 21.1 Pathway enrichment analysis of 1703 genes
(from a total of 19,606) related to heat stress in at least three
species of livestock (cattle, sheep, goats or horses) . . . . . . . . . . . . . . 275

xxv
 Part I
Impact of Climate Change on Livestock
Impact of Climate Change on Animal
Production and Welfare 1
Veerasamy Sejian, M. V. Silpa, Chinnasamy Devaraj, S. Trivedi,
P. Ezhil Vadhana, W. Ruban, R. U. Suganthi, A. Manimaran,
V. P. Maurya, and Raghavendra Bhatta

Contents
1.1 Climate Variables Affecting Animal Production and Welfare . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
1.2 Effects of Climate on Production Parameters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
1.2.1 Yield . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
1.2.2 Quality . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
1.3 Effects of Climate on Welfare Parameters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
1.3.1 Behavior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
1.3.2 Physiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
1.3.3 Health and Life Expectancy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
1.4 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
1.5 Future Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

V. Sejian (*) · C. Devaraj · P. Ezhil Vadhana · R. Bhatta

Centre for Climate Resilient Animal Adaptation Studies, ICAR-National Institute of Animal
Nutrition and Physiology, Bangalore, India
M. V. Silpa
Institute of Animal Breeding and Genetics, Justus-Liebig-Universität Gießen, Gießen, Germany
S. Trivedi · R. U. Suganthi
Bioenergetics and Environmental Sciences Division, ICAR-National Institute of Animal Nutrition
and Physiology, Bangalore, India
W. Ruban
Department of Livestock Product Technology, Veterinary College, Hebbal, Bangalore, Karnataka
Veterinary Animal and Fishery Sciences University, Bidar, India
A. Manimaran
Southern Regional Station, ICAR-National Dairy Research Institute, Bangalore, India
V. P. Maurya
Division of Physiology and Climatology, Indian Veterinary Research Institute, Bareilly, Uttar
Pradesh, India

# The Author(s), under exclusive license to Springer Nature Singapore Pte 3

Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_1
4 V. Sejian et al.

Abstract

This chapter addresses in detail the various impacts of climate change on live-
stock production and welfare. Among all the environmental stressors, heat stress
is of major concern as it has the most deleterious impact on livestock production
especially in tropical countries which hold a major proportion of the world’s
livestock population. One of the primary responses exhibited by animals under
stress, especially heat stress, is decline in feed intake. This direct impact results in
a series of events that is responsible for significant reduction in the productive
potential of animals. The decreased feed intake in animals during heat stress has
been proved to explain nearly half of the milk yield reduction in dairy cattle. Meat
production is another critical economic trait which again is significantly altered
due to heat stress. In dairy cattle, heat stress has been proven to alter the milk
composition impairing its quality and ultimately the economic value. Reduction
in milk fat and protein content in sheep and goat milk due to heat stress negatively
influence the production of high quality cheese. Similarly, the meat quality traits
and its governing genes expression patterns are also compromised in heat stressed
animals. Further, climate change also induces changes in welfare variables
governing behavior, physiological, endocrine, and metabolic as well as immune
system related variables. In addition, climate change also affects the animal health
particularly the vector borne diseases. Thus, climate change and its impact on
animal production, health and welfare are of great relevance that needs to be
addressed efficiently to sustain animal production in the changing climate
scenario.

Keywords
Climate change · Health · Heat stress · Livestock · Milk production · Welfare

Abbreviations

% Percentage
ACTH Adrenocorticotropic hormone
ALP Alkaline phosphatase
ALT Alanine aminotransferase
AST Aspartate aminotransferase
BHBA β-hydroxy butiric acid
BUN Blood urea nitrogen
CAPN Calpain
CAST Calpastatin
CD Cluster of differentiation
CRH Corticotrophin releasing hormone
CRYA Crystallin alpha
DFD Dark, firm and dry
DGAT Diacylglycerol acyltransferase
1 Impact of Climate Change on Animal Production and Welfare 5

GH Growth hormone
GSH Glutathione
HPA Hypothalamo-pituitary-adrenal
HSP Heat shock protein
IFN Interferon
Ig Immunoglobulin
IGF-1 Insulin-like growth factor-1
IL Interleukin
LDL Low density lipoprotein
MDA Malondialdehyde
MSTN Myostatin
NEFA Non-esterified fatty acids
PSE Pale, soft and exudate
RVF Rift valley fever
SAM Sympathetic-adrenal-medullary
SOD Superoxide dismutase
T3 Triiodothyronine
T4 Thyroxine
THI Temperature humidity index
TLR Toll-like receptor
TNF Tumor necrosis factor
TSH Thyroid stimulating hormone
USD United States dollar
VFA Volatile fatty acid
WHC Water holding capacity

1.1 Climate Variables Affecting Animal Production

and Welfare

The global human population is prediction to increase by 33% from 7.2 to 9.6 billion
by the end of 2050. This would thereby rise the demand for agricultural products that
is predicted to increase by 70% (Rojas-Downing et al. 2017). However, there is no
increase in the total global land area for cultivation, rather it remains nearly the same
since 1991. This therefore reflects the intensification efforts initiated to increase the
production levels. Among the agricultural sector, livestock products are thought to
play a vital role in ensuring food security especially with the rising human demand
for animal protein. The livestock sector therefore has adopted several intensified
practices so as to boost their production levels. Livestock production however is
influenced by a number of factors, climate change being one among them. Apart
from having a direct impact, the indirect impact of climate change on livestock
production like reduction in fodder, pasture, water scarcity and increased disease
outbreaks, are also of great concern. Furthermore, these deleterious consequences
6 V. Sejian et al.

also impair the animal’s well-being thereby creating a global issue on animal
welfare.
Climate change is no longer a myth, in fact having known the impact of climate
change on the ecosystem, leaders from across the globe have formed numerous joint
ventures to ameliorate and mitigate the adverse impact of climate change. The
livestock sector holds a unique peculiarity for being both a contributor to climate
change and also affected by it. Air temperature, humidity, wind velocity and solar
radiation are among the prime climate variables that affect livestock production.
Furthermore, among all the environmental stressors, heat stress is of major concern
as it has the most deleterious impact on livestock production especially in tropical
countries which hold a major proportion of the world’s livestock population.
Animals possess innate abilities to combat the adverse climatic variabilities,
however these impose serious negative influence on their production potential.
Apart from production decline and economic losses, another vital aspect that is
often overlooked is the well-being of the animal. The impact of climate change on
animal welfare is a less explored area that needs much attention. The increasing
occurrences of climatic adversities, both in frequency and intensity, rapid spread of
diseases, lack of quality feed and fodder, along with the intensified production
practices to increase production, all of these together exert a lot pressure on all
livestock species. Hence it is the need of the hour to propagate the urge to ensure a
balance optimum production amidst the drastically changing climate scenario and
animal welfare.

1.2 Effects of Climate on Production Parameters

1.2.1 Yield

In the United States alone, heat stress was reported to cause an estimated financial
loss of nearly 900 million USD/year, 300 million USD/year and 300 million
USD/year in dairy, beef and swine industry respectively. This would quantify the
relevance to address the rising concern of climate change and livestock production.
One of the primary responses exhibited by animals under stress, especially heat
stress, is decline in feed intake. This direct impact results in a series of events that is
responsible for significant reduction in the productive potential of animals. Signifi-
cant decline in a number of economic traits like milk yield, meat yield, body weight,
body condition score, egg production and so on, as a consequence of heat stress have
been reported in livestock and poultry.
In a study conducted in Sub-Saharan African climate of Tanzania, the impact of
heat stress on milk production was assessed among the small holder dairy cattle
populations (Ekine-Dzivenu et al. 2020). Based on the results obtained, heat stress
was proved to cause a decline in milk production by 4.16% up to 14.42% across the
temperature humidity index (THI) groups (61–86). Furthermore, a W-shaped
response pattern of milk yield across THI scale was observed in this study. The
proportion of decline in milk yield and also its trend across THI may vary depending
1 Impact of Climate Change on Animal Production and Welfare 7

on the breed, region, management and several other factors. However, this is an
inevitable fact that heat stress results in a significant decline in milk yield and also its
components like fat yield, lactose, etc. Furthermore, the decreased feed intake in
animals during heat stress has been proved to explain nearly half of the milk yield
reduction in dairy cattle. Similar reduction in milk yield as a consequence of heat
stress has been reported in buffalo, and goats too.
Meat production is another critical economic trait which again is significantly
altered due to heat stress. This area however has been least explored in livestock
species. Unlike the milk variables, the impact of heat stress on meat production
cannot be measured immediately. Significant reduction in growth rate, feed conver-
sion efficiency, body weight, body condition score, carcase weight, omental fat and
loin yield, due to heat stress have been reported in beef cattle, sheep and goats.
Likewise, heat stress also has a deleterious impact on egg production in layers. The
impact of heat stress on meat production and body characteristics varies among
species, breed, sex and age. Nevertheless, the decline in production caused is severe
that needs to be addressed efficiently.
Based on the recently published report by Rahimi et al. (2021), the frequency of
dangerous heat stress events and the average number of days under heat stress for
livestock reared in East Africa was projected to increase by 2100. Additionally, the
report also suggested that with the projected rising global surface temperature, most
of East Africa would be unsuitable to support exotic swine, dairy cattle and poultry
production. This is of serious concern from the food security point of view and also
from the animal welfare perspective.

1.2.2 Quality

The livestock sectors stands strong for the massive economic returns obtained; this is
governed by several factors, the quality of product being one among the most vital
factors. Apart from negatively influencing the yield, heat stress also compromises on
the quality of livestock products obtained. In dairy cattle, heat stress has been proven
to alter the milk composition impairing its quality and ultimately the economic
value. In small ruminants, milk quality is a huge determinant for cheese production.
Reduction in milk fat and protein content in sheep and goat milk due to heat stress
negatively influence the production of high quality cheese. Moreover, the plasma
mineral imbalance caused during high ambient temperatures also impairs the quality
of milk and its products.
Studies on impact of heat stress on meat quality are less documented in most
livestock species like beef cattle, sheep, goat and pigs. Most of the studies on this
aspect have primarily focused on poultry meat and hence there is increasing research
on this line in other livestock species since the recent past. The impact of heat stress
on meat quality traits varies among the livestock species that could be possibly
attributed to the varied thermo-tolerance ability in these species and their breeds.
However, despite these variations, to summarize the impact of heat stress on meat
quality, animals subjected to acute heat stress prior to slaughter were reported to
8 V. Sejian et al.

stimulate muscle glycogenolysis. As a consequence of this impact, meat from such

animals usually undergoes a condition known as pale, soft and exudative (PSE) meat
having low water holding capacity (WHC). On the other hand, chronic heat stress is
reported to result in dark, firm and dry (DFD) meat in animals which is characterized
by high ultimate meat pH and high WHC. Irrespective of the type of heat stress, its
impact on animals negatively affects the quality of meat, be it PSE or DFD, reducing
its shelf life, consumer preference and food safety (Gonzalez-Rivas et al. 2020).
The impact of heat stress on meat quality in animals has been explained via a
number of physiological processes occurring in the muscles. However, advanced
studies have also reported the associations of a number of molecular mechanisms
which involve the expression of a number of genes. In a recent study reported by
Devapriya and co-workers (2021), to assess the meat quality of indigenous Kodi
Aadu goats subjected to heat stress, a number of novel genes influencing meat
quality were identified. The authors placed on record the vital role of myostatin
(MSTN), calpain 1 (CAPN1), calpain 2 (CAPN2), calpastatin (CAST), diacylglycerol
acyltransferase 1 (DGAT1), crytallin alpha (CRYA), heat shock protein 27 (HSP27),
HSP40 and HSP90 genes to influence the meat quality variables in goats on
exposure to heat stress.

1.3 Effects of Climate on Welfare Parameters

As mentioned earlier, alterations in the climatic conditions have an unequivocal

impact on livestock production. Though this is of major concern in the tropical
regions, it’s an alarming issue also for animals reared in the temperate regions
wherein the summers are getting relatively warmer over the years. Additionally,
the evolving human lifestyle is also exerting pressure regarding farm animal welfare.
Apart from intensification of livestock production, the outcomes of climate change,
especially heat stress, also significantly deter the welfare of animals. These can be
assessed based on several variables such as environmental indices for heat stress,
behavioral responses exhibited by animals under heat stress, physiological
alterations caused especially due to failure to cope with the deleterious heat stress
effects and also based on their health status (Sejian et al. 2011).

1.3.1 Behavior

Heat stressed animals exhibit a number of behavioral alteration so as to adapt to heat

stress. As a matter of fact behavioral adaptations are stated to be the first response
adopted by animals to combat the deleterious impact of climatic alterations. Among
the behavioral responses, shade seeking and reduced feed intake have been
documented as the quick and profound responses observed in heat stressed animals.
While shade seeking can be observed only in extensive rearing, reduction in feed
intake has been reported both in extensively and intensively reared animals (Sejian
et al. 2021). The primary objective of reducing feed intake during heat stress can be
1 Impact of Climate Change on Animal Production and Welfare 9

put forth as an adaptive response exhibited by heat stressed animals so as to reduce

their internal metabolic heat production.
Significantly higher water intake and increased drinking frequency are also the
classical response observed in heat stressed animals. This sign is of welfare issue
especially in extensively reared animals as they may have to travel longer distances
in search of water which adds on to the stress. The concept of multiple stresses
reported by Sejian et al. (2018) reflects a major animal welfare issue in animals
wherein heat stress, in addition with nutritional and walking stress, has a multifold
impact on the animals’ response and production when compared to any of these
stressors alone.
Alteration in standing time and lying time are also important behavioral responses
established in heat stressed animals. As a general concept, heat stressed animals have
been reported to spend more time standing which enables them to re-orient their
body so as to avoid the direct solar radiation and also from the surface radiations
emitted mainly from the ground. Similarly, altered urinating frequency, defecating
frequency and rumination time are among the other behavioral responses exhibited
by animals under heat stress. Additionally, wallowing is a unique behavioral
response observed only in buffaloes during heat stress. Furthermore, increased
aggression and restlessness are other behavioral signs indicating severe stress in
animals.
The expression of all the above mentioned behavioral alteration depend highly on
the type of species, breed, their age, stage of production and also management
practices followed. Furthermore, marked variations in the exhibition of these
responses have been reported in cattle, sheep and goats. Though there are a number
of causes for these variations, climate resilience is among the prime determinants
influencing these responses in animals.

1.3.2 Physiology

1.3.2.1 Endocrine
The endocrine system, involved in the production of hormones, play a significant
role in thermoregulation in animals. Stress induced especially due to environmental
alterations trigger the hypothalamo–pituitary–adrenal axis (HPA axis) which is the
predominant endocrine regulator for stress response in animals. The activation of the
HPA axis stimulate the production and release of a series of hormones comprising
the corticotrophin-releasing hormone (CRH), adrenocorticotropic hormone (ACTH)
that finally leads to the secretion of the classical stress marker, cortisol. Apart from
this, the thyroid hormones, triiodothyronine (T3) and thyroxine (T4), that are
associated with metabolic responses are also altered in heat stressed animals.
These hormones play a crucial role to maintain thermogenesis in animals especially
from the metabolic response point of view. Additionally, heat stress also leads to
altered hormonal profile in other homeostasis, metablosim and production associated
hormones like aldosterone, adrenalin/noradrenalin, melatonin, prolactin, oestrogen,
leptin, growth hormone (GH), insulin-like growth factor 1 (IGF-1) and many more.
10 V. Sejian et al.

Several studies have associated the alteration in such endocrine variable with heat
stress in livestock.
In a study led by Li et al. (2020) to assess the seasonal dynamics in buffaloes
under hot and humid climate, high THI was associated with significant increase in
serum cortisol concentration in non lactating Nili Ravi buffaloes. While the plasma
T3, ACTH, insulin and GH concentrations were observed to be lower in buffaloes
during the summer season when compared to spring, autumn and winter seasons. In
another study by Pragna et al. (2018), comparatively assessed the summer season
induced rhythmic alterations in metabolic activities in three indigenous goats breeds.
Based on their results heat stress was reported to significantly reduce the plasma
thyroid stimulating hormone (TSH) concentration in Malabari and Salem Black
goats exposed to heat stress when compared to their respective control. Additionally
the plasma T3 concentration was also significantly reduced in heat stressed Malabari
goats when compared to their control. Interestingly none of these hormone were
significantly altered in the third breed, Salem Black, thereby reflecting the better
climate resilience ability.
Several other studies in cattle, sheep, pigs and poultry depicted altered hormonal
profile in heat stressed animals. Moreover, a number of hormonal markers for heat
stress have also been established. Though there are no well established threshold
limits for the concentration of such hormones in livestock species, their relative
levels with respect to animals under thermo-neutral conditions can definitely reflect
the stress status of the animal.

1.3.2.2 Metabolic
The phenomenon of reduction in feed intake to reduce metabolic heat generation has
already been explained in the previous sections. Metabolic responses exhibited by
animals under heat stress can be assessed by screening their enzyme profile. This not
only provides representative information on the activity of other tissues and organs
during heat stress but can also reveal the wellness of the animal. Acid phosphatase,
alkaline phosphatase (ALP), aspartate aminotransferase (AST), alanine aminotrans-
ferase (ALT), non-esterified fatty acids (NEFA), β-hydroxybutiric acid (BHBA) and
volatile fatty acids (VFAs) are some of the established heat stress associated
metabolic markers in livestock.
The alteration of metabolic variables and impaired production during heat stress
was reported in lactating sows by He et al. (2019). The study revealed significant
increase in serum insulin, creatinine, BUN and plasma NEFA in heat stressed sows
(Landrace x Large White) that were during late gestational stage. The authors also
reported significant alterations in a number of metabolites involved in glycerolipid
metabolism, β-alanine metabolism, and pantothenate and CoA biosynthesis in preg-
nant sows. Their study summarized heat stress to elevate thermal response in late
gestational sows that significantly reduced their productive performance and most
importantly, enhanced lipid and protein catabolism.
In another study, Joo and co-workers (2021) assessed the alterations in blood
metabolites (biochemical metabolities, enzymes and minerals) in Holstein and
Jersey dairy cows exposed to heat stress. Apart from reporting an evident impact
1 Impact of Climate Change on Animal Production and Welfare 11

of heat stress on the metabolic profile in dairy cattle, the authors also reported breed
variations for some of the variables. Such variations were linked to differences in
productivity, metabolism and disease vulnerability between the two breeds. The
blood metabolities significantly altered due to heat stress included albumin, protein,
glucose, BUN, total cholesterol, LDL cholesterol, ALT, AST, creatine kinase,
calcium, sodium, potassium, chloride, magnesium and phosphorus. Such alterations
in the metabolic profile were assessed in several other livestock species. This thereby
highlight the severity of distress the animals would be undergoing as a consequence
of heat stress. Therefore, it is of utmost importance to work on ameliorative and
mitigation measures to combat the adverse effect of heat stress in livestock produc-
tion both from the production and animal welfare perspective.

1.3.2.3 Immunity
Impact of climate change on immune status in animals is another crucial yet least
explored concept. Heat stressed animals are prone to a number of diseases that
indirectly reduces their productivity and also adds up to the existing stress. Immune
suppression in heat stressed animal is mainly associated to the activation of the HPA
and sympathetic-adrenal–medullary (SAM) axes (Inbaraj et al. 2016). The final end
products excreted through these axes, glucocorticoids and catecholamines, suppress/
inhibit the synthesis and release of cytokines and other products having a vital role in
immune defense. Scientific studies exploring the impact of heat stress in immune
system of animals have been gaining more momentum over the past few years. A
number of immunity associated markers have been reported in livestock species.
Some of these include altered hematological profile, interleukin-2 (IL-2), IL-4, IL-5,
IL-6, IL-12, interferon γ (IFNγ), tumor necrosis factor-α (TNF-α), toll-like receptors
(TLR) 1–10, IgG, IgA, and so on.
In the previously stated study by Joo et al. (2021), the impact of heat stress was
also assessed on immune cells of lactating Jersey and Holstein cows. Similar to their
findings on the blood metabolite profile, an evident breed variation was observed
apart from heat stress influence on the differential immune cell population of
PBMCs. Heat stress was observed to significantly increase the proportion of B cell
(CD4–CD21+) only in Holstein cows, while that of monocytes (CD11b+CD172a+)
significantly reduced.
In a recent report, Xia et al. (2021) studied the heat stress induced mucosal barrier
dysfunction in boars. The authors studied the impact of short-term heat stress on a
number of variables which also included oxidation status and cytokine levels in pigs.
Based on this experiment, heat stress was proved to significantly reduce serum IL-8,
IL-12, IFNγ, and also the antioxidant activity which were reflected via increased
malondialdehyde (MDA) and glutathione (GSH) content in addition with lower
superoxide dismutase (SOD). Thus from the animal welfare point of view, the
impact of climate change on the immune status in animals should be considered to
be of alarming effect this can cause severe distress to the animal. In worst cases
extreme heat stress in association with compromised health status of the animal may
lead to increased mortality.
12 V. Sejian et al.

1.3.3 Health and Life Expectancy

Health status of animal is considered as one of the vital welfare variable. Impaired
health status in animals, especially due to climate change, not only reflects the
economic losses incurred but also the quantum of pain, discomfort or distress
evinced by them. In addition to the direct impact on immune response, the varying
environmental conditions lead to the emergence, re-emergence and rapid spread of a
number of deadly diseases. Global climate change has also resulted in huge
modifications on the micro and macroclimate of host and parasite in addition to
the significant alteration in host-microbe interaction. High temperatures along with
high relative humidity are ideal for the multiplication and survival of microbes,
parasites and also disease transmitting animal vectors. Moreover, the rapid surge in
livestock diseases during the warmer seasons further substantiates this concept of
climate change on livestock diseases.
As mentioned earlier, animals exposed to increasing and prolonged heat load
exhibit increased standing behavior. Though the primary purpose of this response by
animals is for heat abatement and cooling body surfaces, prolonged standing time
can predispose the animal to lameness. However, there are no direct studies
conducted to link heat stress and lameness in animals (Polsky and von Keyserlingk
2017). Apart from the direct and indirect impact of climate change on immune
response and diseases in livestock, there are also reports on association of some
climatic adversities with certain diseases. In eastern Africa, the outbreak of Rift
Valley Fever (RVF) has been often linked with El Niño. It may be noted that this
disease if of major concern for having serious impacts on livestock morbidity and
mortality.
Although the literature assessing the effect of climate change on livestock disease
is limited, the postulated impact on the health status in animals should not be
overlooked. This is of huge relevance from production, economic, animal welfare
and public health point of view. The pain and suffering undergone by animals due to
their deteriorating health status is grave. Moreover, their prolonged treatment also
impairs the future productivity of the animal thereby reducing its life expectancy.
Furthermore, most of the deadly diseases in livestock are zoonotic thereby risking
their spread among the human population. Therefore, climate change and its impact
on animal production, health and welfare are of great relevance that needs to be
addressed efficiently to sustain animal production in the changing climate scenario.

Learning Outcomes
• Climate change deleterious impact on livestock production and welfare is
primarily mediated by individual or combination of climate variables such
as temperature, humidity, rainfall, wind velocity, solar radiation etc.

(continued)
1 Impact of Climate Change on Animal Production and Welfare 13

• Climate change significantly reduces the economic traits like milk yield,
milk composition, meat yield, body weight, body condition, egg production
in livestock and poultry.
• The direct impact of climate change depends on its intensity and duration
and affects the health and welfare of livestock by suppressing the immune
system.

1.4 Conclusions

Livestock needs to be the priority focus in agriculture sector due to their high climate
resilience and are tipped to be playing a significant role in meeting future food
demands of growing human population. Although considered climate resilient, still
livestock production is hampered in order to facilitate adaptation in farm animals.
Thus the milk, meat, reproduction and immune system functions are compromised to
support vital life sustaining activities of farm animals in the changing climate.
Several welfare indicators covering farm animal behavior, physiology, hormones,
metabolites, immune and health related variables must be considered to assess the
animals’ adaptive potential.

1.5 Future Perspectives

A clear region specific database on the various impacts of climate change on farm
animal production needs to be developed. This will help the policy makers to
identify the breeds with minimal impact for dissemination to the farmers. Although
the various climate change associated environmental stressors impact was fairly
established in farm animals, still only predominantly heat stress impacts were known
in depth. Information particularly on assessing the impacts of climate change
associated water stress on livestock production and welfare have not been fully
explored. Such an effort would be very vital given the significance of water stress in
the changing climate scenario. Likewise the information pertaining to climate
change associated disease occurrences in livestock have not been fully established.
Therefore, establishing the impacts of climate change associated water stress and
sudden disease outbreaks on the performance and welfare of farm animals is the need
of hour. More intensified research efforts are also needed to identify more quantifi-
able welfare indicators related to animal health. In addition, efforts are also equally
needed to develop more cost effective individual-level sensors (e.g., accelerometers,
GPS, and RFID) and automated monitoring systems for group level animal monitor-
ing (e.g., cameras) needs to be developed to assess the welfare of grazing animals.
14 V. Sejian et al.

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Effects of Heat Stress and Climate Change
Induced Bushfires on Beef Meat Quality 2
Robyn D. Warner, Melindee Hastie, Paula A. Gonzalez-Rivas,
Surinder S. Chauhan, Minh Ha, Caitlin Pfeiffer, Allison Hilman,
and Brendan Cowled

Contents
2.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
2.2 Physiologic and Metabolic Consequences of Heat Stress . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
2.3 Effect of Heat Stress on Beef Meat Quality . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
2.4 Impact of Bushfires on Beef Meat Quality—A Case Study from Australia . . . . . . . . . . . . . . 21
2.5 Summary and Future Research . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

Abstract
Heat stress is considered a very stressful event for livestock, and it has detrimental
consequences for not only animal health and productivity but also product
quality. During bushfires, which are increasingly prevalent under climate change,
animals can be exposed to extreme heat stress events. Ruminants are very prone
to heat stress because of increased metabolic rate. Under the influence of chronic
heat stress, ruminants exhibit a reduction of muscle glycogen concentration,
which leads to increased, ultimate pH and WHC, producing the quality defect
of dark-cutting meat. In addition, heat stress causes extended protein and lipid
oxidation, oxidative stress and shelf life reduction of meat. A case study is
presented on the effects of the devastating bushfires in Australia in 2019–2020
on the quality of 450,000 beef carcases. This chapter focuses on the effects of
bushfire and heat stress on the ruminants and the consecutive changes produced

R. D. Warner (*) · M. Hastie · P. A. Gonzalez-Rivas · S. S. Chauhan · M. Ha · C. Pfeiffer

Faculty of Veterinary and Agricultural Science, The University of Melbourne, Parkville, VIC,
Australia
e-mail: [email protected]
A. Hilman · B. Cowled
Ausvet, Bruce, ACT, Australia

# The Author(s), under exclusive license to Springer Nature Singapore Pte 15

Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_2
16 R. D. Warner et al.

on the quality of the meat. Discussion on knowledge gaps, and future perspectives
are also provided.

Keywords

Beef · Bushfires · Cattle · Dark-cutting · Heat stress · Meat quality · MSA · PSE

Abbreviations

$ Dollar
% Percentage


C Degree Celsius
ANS Autonomic nervous system
AUS-MEAT Authority for the uniform specification of meat and livestock
cAMP Cyclic adenosine mono phosphate
cm2 Centimeter square
h Hour
HCO3 Bicarbonate
HGP Hormonal growth promotant
HSP Heat shock protein
kg Kilogram
L Lightness
MLA Meat and Livestock Australia
MSA Meat Standards Australia
NSW New South Wales
pH Potential of hydrogen
PSE Pale, soft and exudative
RH Relative humidity
ROS Reactive oxygen species
US Unites States
WBSF Warner Bratzler shear force
WHC Water holding capacity
β Beta

2.1 Introduction

The economic losses incurred by heat stress annually the U.S. beef industry is
estimated to be about 369 million U.S. dollars which quantifies the hazardous effect
of heat stress on a country’s economy (St-Pierre et al. 2003). Heat stress is consid-
ered one of the most expensive and stressful events affecting the productivity, health
and meat quality of the animal. The economic losses are associated with decreased
fertility, reduced rate of growth, inappropriate carcase composition and quality,
decreased carcase weights, leading to a surge in animal welfare issues and very
2 Effects of Heat Stress and Climate Change Induced Bushfires on Beef Meat Quality 17

high veterinary costs. Negative influences of heat stress on the economy have
resulted in the developing imperative to address climate changing scenarios. The
adverse effects of heat stress on animals’ health and productivity are likely to
continue in the upcoming years if the selection is more targeted towards improving
the traits related to production. It is to be noted that these production traits are
negatively correlated with traits associated with adaptability to climate change and
thermo-tolerance.
From the perspective of increased metabolic rate, higher growth rate, improved
production of basal metabolic heat and increased productivity; ruminants are highly
susceptible to the negative influences of heat stress. The heat produced during
ruminal fermentation is the major risk factor threatening the thermoregulation
capacity of ruminants. (Kadzere et al. 2002; Tajima et al. 2007). Pale, soft and
exudative meat (PSE), a condition characterised by decreased ultimate pH and
reduced water holding capacity (WHC), was commonly found in carcases of poultry
and pigs, (Kim et al. 2014; Warner et al. 2014). The reason for this condition is stress
or struggling of animals at slaughter, exacerbated by any acute heat stress, leading to
acceleration in the rate of muscle glycogenolysis and increased concentration of
lactic acid, rapid reduction in muscle pH, during early phases post-portem when the
carcase remains hot (Owens et al. 2009). In contrast to the PSE condition, animals
that are subjected to chronic heat stress had significantly less glycogen reserves and
subsequent reduction in lactic acid production, leading to increased, ultimate pH and
WHC producing dark-cutting meat in ruminants (Adzitey and Nurul 2011; Gregory
2010; Kadim et al. 2008; Mitlöhner et al. 2002). Moreover, during hot climatic
conditions, increased protein and lipid oxidation and reduced shelf life of meat were
also observed (Mujahid et al. 2007; Wang et al. 2009).
Bushfires have become increasingly prevalent with climate change in Australia
and overseas (Visner et al. 2021). Adverse health outcomes for cattle from bushfires
can include direct contact with fire, exposure to radiant heat, and inhalation of
bushfire smoke (Hillman et al. 2021). The potential effects of bushfires on meat
and carcase quality are diverse and multi-faceted, including those that may impact
carcase weight, condition of the animal and hence fat score, and marbling, meat pH,
and colour. These outcomes for meat quality from bushfires are likely caused by;
(i) heat stress from the radiant heat of the bushfire, (ii) reduced nutrient intake due to
heat stress and lack of feed/pasture availability, and (iii) impact of stress associated
with an unfamiliar environment, transport and more frequent movement of stock.
The effects of bushfires on carcase and meat quality will result in income loss for
cattle producers and have implications for animal welfare and productivity.
This chapter discusses the consequences of heat stress and bushfires for beef meat
quality. Data on bushfire effects on beef quality are lacking, and a case study is
presented on recent data which utilised animal and carcase data from the recent
bushfires in Australia. Gaps in the literature are identified, and directions for future
research are suggested. Knowledge gaps, scarcity in the literature and future
endeavours were also discussed in this chapter.
18 R. D. Warner et al.

2.2 Physiologic and Metabolic Consequences of Heat Stress

Response to heat stress is primarily mediated via the activation of the autonomic
nervous system (ANS) by catecholamines (noradrenaline and adrenaline), which
leads to increased body temperature, heart rate and respiratory rate, redirection of
blood supply to the skin from viscera for the process of thermoregulation, increased
utilisation of energy from body stores (Minton 1994) by muscle glycogenolysis
acceleration and energy reserve suppression (Afsal et al. 2018; Gregory 2010).
Muscle glycogenolysis acceleration is mediated via cyclic adenosine
monophosphate (cAMP) by initiating cascades of reaction in the muscle produced
by the action of catecholamines on β2-receptors (muscle). This process, in turn, leads
to glycogenesis inhibition and glycogenolysis activation by inhibiting glycogen
synthase (Roach 1990) and activating glycogen phosphorylase, respectively (Franch
et al. 1999).
As an adaptive mechanism to tackle the rise in metabolic heat production, animals
reduce their feed intake under high environmental conditions, which is considered
the major impact of heat stress on livestock (Kouba et al. 2001; Odongo et al. 2006;
Russell 2007). This feed intake reduction explains only 50% of the animals’
physiological and metabolic responses under heat stress, the rest of which can be
explained by the hormonal profile and energy partition variations (Baumgard and
Rhoads 2007). In beef cattle, the feed intake reduction starts at two combinations
of temperature and relative humidity (R.H.), one occurs at an ambient temperature of
about 27
C, and > 80% of R.H. and the other happens at an ambient temperature of
about 30
C and < 80% of R.H. (Bernabucci et al. 2010).
Also, fasting of steers for about 7 days was found to decrease the glycogen
concentration in muscle by about 30%. It is always important to provide adequate
nutrition to the cattle before sending them for slaughter to prevent the incidence of
(Knee et al. 2007; Pethick et al. 2005) dark-cutting meat, keeping in mind the fact
(Warner et al. 2006) that there is always a reduction in glycogen reserve of animals
sent for slaughter from the farm.
Providing ad-libitum water for heat-stressed animals is of paramount importance
to avoid dehydration and negative influences of heat stress. Usually, it is found that
there is a two-fold rise in the requirement of water for the animals experiencing heat
stress (Beede and Collier 1986), which is considered to be a fundamental mechanism
of thermoregulation to tackle evaporation of water through sweating (Marai et al.
2007) and panting. This is also proved to be a direct mechanism of cooling the
reticulo-rumen to decrease body temperature (Bewley et al. 2008). If the availability
of ad-libitum water to the animals is neglected under heat stress, this can cause
severe dehydration and enhance the adrenergic mode of responses to heat stress
(Matthews and Parrott 1991).
Increased respiratory rate under heat stress leads to respiratory alkalosis (Cottrell
et al. 2015; Odongo et al. 2006; West 2003). The body tries to compensate for this
situation via the urinary system by increasing the HCO3 excretion (Schneider et al.
1988), leading to metabolic acidosis. In this condition, the body’s metabolic depen-
dency shifts towards anaerobic conditions generating energy through the production
of lactate from pyruvate, similar to the early stages of post-mortem metabolism. As
2 Effects of Heat Stress and Climate Change Induced Bushfires on Beef Meat Quality 19

mentioned above, this results in PSE-like conditions (Wang et al. 2009; Gholamreza
et al. 2019) in meat.
Furthermore, there is also a surge in generation of reactive oxygen species (ROS)
and reduction in antioxidant capacity of the animal under heat stress, both of the
described conditions inducing oxidative stress to the animal (Chauhan et al. 2014a;
Liu et al. 2016; Mujahid et al. 2007; Mujahid et al. 2006; Mujahid et al. 2005;
Shakeri et al. 2018). This induced oxidative stress and acidosis in the tissue further
produces much more adverse effects such as a free radical-mediated chain reaction,
cytotoxicity, enhancement in lipid and protein oxidation, impaired health of animals,
all of which cause a reduction in the shelf life of the meat (Celi and Gabai 2015;
Chauhan et al. 2014b; Imik et al. 2012) and negative impairment of the meat quality.

2.3 Effect of Heat Stress on Beef Meat Quality

Heat stress is associated with dark-cutting in beef cattle (Gregory 2010).

Pre-slaughter stress depletes muscle glycogen reserves, resulting in a high pH
(>5.70 in Australia) that causes dark-cutting meat (AUS-MEAT 2005), and dark
colour of meat occurs mainly due to reduced light scattering in the muscle and
predominance of purple deoxymyoglobin in the meat surface. There is a multitude of
hypotheses to explain the mechanisms behind the dark colour of meat, and the main
ones are as follows. Mitochondria can thrive at high pH and increase their propor-
tional oxygen-consumption activity; as a result, deoxymyoglobin (purplish-red)
predominates, resulting in black meat (Faustman 1994; Suman and Joseph 2013).
Additionally, the high pH prevents post-mortem myofibril and muscle cell shrink-
age, minimising the meat’s light scattering capabilities measured by low lightness
(L*) values (Hughes et al. 2018). Finally, at high pH levels in meat, muscle proteins
above their isoelectric point bind more water, resulting in densely packed myofibrils,
and light scattering will be reduced (Abril et al. 2001).
Various reports suggest ruminants exhibit more dark-cutting during warm
weather than during cold weather. (Hughes et al. 2018; Kadim et al. 2004; Kadim
et al. 2008; McPhail et al. 2014; Węglarz 2010). Dark-cutting rates are higher in the
northern hemisphere between September and October because of the more frequent
occurrence of cold nights and hot days caused by unexpected weather changes and
temperature differentials, such as those that occur during late spring and fall break
(Boykin et al. 2017).
The increased incidence of dark-cutting in the summer can be attributed to a
combination of poor pasture quality (Knee et al. 2004) and reduced feed intake. Heat
stress affects feed intake, and liver glycogen cannot generally maintain energy
homeostasis in ruminants that rely primarily on volatile fatty acids as glucose
precursors; animals must rely on muscle glycogen as a glucose source during heat
stress (Gardner et al. 2014). Sheep exposed to handling and transport stress had
higher rectal temperatures associated with lower glycogen levels during slaughter
(Pighin et al. 2014). This suggests a relationship between increased body tempera-
ture, glycolytic stress response, and maybe elevated muscle pH and dark-
coloured meat.
20 R. D. Warner et al.

Feedlot cattle are the most susceptible to heat stress among ruminants. They are
frequently exposed to radiating surfaces, consume a lot of high-energy feed, and have
few opportunities to seek out shade, water, or ventilation for cooling purposes
(Gaughan et al. 1996; Holt et al. 2004; Mader et al. 1997; Renaudeau et al. 2012).
Pre-slaughter body temperatures in grain-fed cattle are higher than in grass-fed cattle
(Jacob et al. 2014), which is related to an earlier and stronger rigour mortis, resulting in
tougher meat (Warner et al. 2014). Additionally, ruminants fed grains that ferment
rapidly are more susceptible to heat stress (Gonzalez-Rivas et al. 2016; Gonzalez-
Rivas et al. 2017; Mader et al. 1997). Therefore, it is plausible to hypothesise that meat
from feedlot cattle will be more likely to exhibit poor quality due to heat stress. Shade
may be advantageous in feedlot conditions since muscle pH was significantly higher in
the loin of feedlot steers kept in the shade during the summer at 1 h (5.97 v. 6.03,
P ¼ 0.014) and 2 h (5.97 v. 6.03, P ¼ 0.043) post-slaughter. However, the shade had
no effect on hardness, texture, or drip loss of the meat (DiGiacomo et al. 2014).
A higher percentage (59%) of carcases classified as dark-cutting (here defined as
an ultimate pH > 6.0) were found in cattle slaughtered under heat stress conditions
(>30
C) than animals slaughtered during the cold season. Longissimus thoracis
muscles from hot-boned beef were found to have a higher pH (6.24 vs. 5.54), lower
WBSF (10.1 vs. 15.6 kg/cm2), and higher cooking loss (19.8 vs. 26.0%) when
processed during the hot season and removed within 60 min post mortem, compared
to those collected during the cool season (Kadim et al. 2004). For Omani beef that
was harvested in the hot season, the longissimus thoracis muscles had darker, less
red and less yellow meat than those collected in the cold season based on L*
(31.45 vs. 35.58), a* (18.53 vs. 23.19) and b* (4.16 vs. 6.40) measurements made
48 h post-mortem (Kadim et al. 2004).
Holstein-Friesian cross meat produced in winter exhibited a brighter red colour
and higher values of a*, b*, C and H values (redness, yelllowness, chroma, hue
respectively) than meat produced in summer; nevertheless, in summer, there was a
30% greater frequency of meat with an ultimate pH > 5.8 measured in the
longissimus thoracis at 48 h post-mortem (Węglarz 2010). A decreased incidence
of dark-cutting was seen in West Texas feedlot heifers housed in shaded pens during
summer relative to heifers in pens with no shade (8.3 vs. 19.8%), but no variations in
hot carcase weight, marbling score, or longissimus muscle area were observed
between the two groups (Mitlöhner et al. 2002). In contrast, shaded heifers in a
prior trial had heavier hot carcases (+16 kg) and thicker fat than unshaded cattle;
however, they found no change in meat quality or carcase yield (Mitlöhner et al.
2001). Animals under shade showed reduced aggressive behaviour, which is in line
with earlier studies suggesting that heat stress can exacerbate bullying behaviour by
depleting glycogen reserves, as reported by the authors (Kreikemeier et al. 1998).
These research established that providing shade to beef cattle effectively mitigates
the detrimental effects of heat stress on performance and meat quality.
In Korea, that Hanwoo beef cattle of different sexes slaughtered in summer had
significantly smaller ribeye areas and higher marbling scores than those slaughtered
in other seasons, but the maturity scores were higher (Panjono et al. 2009). Hanwoo
cattle were used in another experiment, and it was found that longissimus thoracis
from cattle slaughtered in summer had a higher pH at 48 h post-mortem than from
2 Effects of Heat Stress and Climate Change Induced Bushfires on Beef Meat Quality 21

cattle slaughtered in winter (5.46 vs. 5.36). This difference was attributed to the high
temperature and humidity of the Korean summer (Panjono et al. 2011).

2.4 Impact of Bushfires on Beef Meat Quality—A Case Study

from Australia

In Australia, the 2019/2020 summer period involved widespread and devastating

bushfires in southeast Australia. Hillman et al. (2021) investigated the spatiotemporal
relationships between exposure to bushfire and the Meat Standards Australia (MSA)
carcase index of 400,00 beef carcases. The study cattle population was slaughtered and
graded under the MSA program over the period November 2018–July 2020, and cattle
included in th study had a property of origin within 50 km of a bushfire and the
bushfire occurred between 2 and 180 days prior to slaughter. The MSA index is a
quantitative indicator of the expected eating quality and potential value of a beef
carcase. MSA grading data is calculated using only attributes influenced by
pre-slaughter production and reflects the impact on beef eating quality of management,
environmental and genetic differences between cattle at the point of slaughter (MSA
2021a). Geospatial data on the locations of all fires and farm properties were used to
determine the distance of cattle from the fire prior to despatch for slaughter and other
data on the fires. Other animals and carcase data were also collected. Data from the
‘Long Paddock’ database was used to estimate pasture biomass and pasture growth
(Stone et al. 2019) for the two months prior to slaughter. Consistent trends were
observed in the MSA index of beef carcases, with lower MSA index scores seen in
cattle slaughtered after being in close proximity to fire (Hillman et al. 2021). Hence
cattle closest to the fire produced carcases of lower predicted eating quality and lower
value, relative to cattle up to 50 km from the fire.
A subsequent study used MSA grading data from additional beef processing
plants in NSW and Victoria and utilised 450,000 carcases from cattle slaughtered
over the bushfire period (Warner et al. unpublished results). Modelling of loin pH
and incidence of dark-cutting (pH at grading >5.70) was conducted using all MSA
carcase grading data, as well as fire and pasture variables referred to above. And
step-wise regression was used to determine a parsimonious model. The number of
continuous days of fire within 180 km of a property, the closest distance to a
bushfire, the time since the bushfire, and interactions with animal factors signifi-
cantly affected the ultimate pH of the loin muscle as well as the incidence of dark-
cutting meat. The elevated loin pH and increased risk of dark-cutting caused by
bushfires were exacerbated in grass-fed cattle (relative to grain-fed), steers (relative
to heifers/cows), younger cattle (identified by ossification score at grading; MSA
2021b), lower Bos indicus content (identified by hump height at grading, mm; MSA
2021b) and cattle treated with hormonal growth promoters (HGPs, relative to
non-treatment with HGPs; MSA 2021b). The results will be used to create a better
understanding of what bushfires do to beef cattle and how to prepare and respond,
which will be an invaluable tool for producers living in high bushfire areas of
Australia (MLA 2021).
22 R. D. Warner et al.

Learning Outcomes
• Heat Stress can negatively influence beef production and meat quality and
this is attributed to the reduction in muscle glycogen and protein concen-
tration and depletion of fat reserves.
• The harmful consequences of heat stress on meat quality and production is
likely to continue with the current focus in breeding programs on produc-
tion instead of climate adaptatation or thermo-tolerance.
• Provision of high-quality feeds to cattle subsequent to heat stress and
bushfires will assist in ensuring quality carcases and meat.
• Bushfire remains a serious threat to livestock and to the quality and
economic value of carcases under the current climate changing scenario.
Therefore, identifying the risk factors and developing nutritional and
genetic strategies for ameliorating bushfire associated heat stress gains
significance.

2.5 Summary and Future Research

It is evident that cattle suffering under hot and humid conditions are associated with
reduced live weight and carcase weight and also the production of dark-cutting meat.
Under the acute and chronic influence of heat stress, the physiological and metabolic
responses are proven to have a profound association with meat quality by reducing
the protein concentration and glycogen reserve of meat and depletion and redirection
of fat reserves. These conditions also induce cellular damage, oxidative stress and
increased expression of HSPs, together hindering meat quality and producing
deterioration of the meat.
Climate change has also resulted in increased bushfire occurrence, and negative
consequences of bushfires for beef meat quality have recently been demonstrated.
Under the current trends of global warming, unfortunately, the adverse effects of
heat stress are expected to continue if the selection for traits associated with meat
production is prioritised over the traits associated with adaptability to climate change
and thermo-tolerance.
The development of nutritional and genetic strategies for ameliorating heat stress,
particularly in the period following a bushfire event, is important for the future
production of quality beef. Nutritional strategies are needed prior to or during a heat
stress event and in the months following, particularly where bushfires are involved.
Following the exposure of cattle to a bushfire event, in order to ensure muscle
glycogen recovery and reduced occurrence of high pH dark-cutting beef carcases,
producers should consider removing HGPs from beef production and providing high
energy grain-based diets, particularly for older cattle. In the longer term, in bushfire
prone areas or where heat stress events are increasingly common, heat-tolerant beef
breeds such as zebu, brahman, droughmaster and other Bos indicus based breeds are
recommended.
2 Effects of Heat Stress and Climate Change Induced Bushfires on Beef Meat Quality 23

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 Part II
Heat Stress Impact on Adaptive Responses
in Livestock
Adaptation of Beef Cattle to Heat Stress
Challenges 3
J. B. Gaughan, A. M. Lees, and J. C. Lees

Contents
3.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
3.2 Adaptation in Cattle: For a Stressful Environment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
3.3 Mechanisms of Beef Cattle Adaptation to Heat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
3.4 Constraints to Adaptation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
3.5 Beef Breeding Programs to Improve Tolerance to Harsh Environments . . . . . . . . . . . . . . . . . 33
3.6 Molecular Genetics the Way Forward? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
3.7 Advantages of Molecular Approach Over Traditional Approach . . . . . . . . . . . . . . . . . . . . . . . . . 35
3.8 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37

Abstract
The demand for animal protein is increasing with much of this demand in the
tropics and sub-tropics and beef consumption, which represent approximately
25% of global meat consumption, is expected to increase from 54 to 118 million
tonnes by 2050. Although heat stress in beef cattle is an increasing challenge in
many parts of the world there is sustained growth in the number of tropical cattle,
which represent more than 50% of all cattle worldwide. Beef production in the
tropics and sub-tropics is characterised by climate variability (heat and rainfall),
nutritional limitations, parasites, disease, and poor reproductive performance. If

J. B. Gaughan (*)
School of Agriculture and Food Sciences, Animal Science Group, The University of Queensland,
Gatton, QLD, Australia
e-mail: [email protected]
A. M. Lees · J. C. Lees
University of New England, School of Environmental and Rural Science, Armidale, NSW,
Australia

# The Author(s), under exclusive license to Springer Nature Singapore Pte 29

Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_3
30 J. B. Gaughan et al.

we are to meet the challenges of increasing animal protein production in the

tropics and sub-tropics, we will need to move away from traditional selection
methods to more advanced molecular and large-scale genetic evaluations that
consider the whole genome, not just a few specific traits.
This chapter will focus on what is required to improve the adaptive capacity of
beef cattle in harsh environments, some of the mechanisms of adaptation, and the
application, and challenges of selection and breeding strategies e.g., use of
molecular genetic tools to ‘speed up’ adaptation to a changing climate.

Keywords
Adaptation · Beef cattle · Thermal challenge · Breeding · Selection

Abbreviations

% Percentage
> Greater than
ABVg Australian genomic breeding value
DNA Deoxyribonucleic acid
e.g. Exempli gratia (for example)
GEBV Genomic estimated breeding value
HT Heat tolerance
N Nitrogen

3.1 Introduction

The demand for animal protein is increasing with much of this demand in the tropics
and sub-tropics (Cao and Li 2013; Ismail et al. 2020). Beef consumption, which
represent approximately 25% of global meat consumption, is also expected to
increase from 54 to 118 million tonnes by 2050 (Cooke et al. 2020b). Although
heat stress in beef cattle is an increasing challenge in many parts of the world there is
sustained growth in the number of tropical cattle, which represent more than 50% of
all cattle worldwide (Barendse 2017) (Fig. 3.1). Tropical cattle are a very diverse
category and include cattle that either arose in the tropics, are composites of tropical
cattle and temperate cattle, or were temperate cattle that were translocated to the
tropics and have been under natural or artificial selection for many generations
(Barendse 2017). Cattle that have evolved in harsh environments have developed
grazing patterns that allow them to be more nutritionally efficient especially with
low-quality forages (Russell et al. 2012; Cooke et al. 2020a, b), they are resistant to
internal and external parasites (Frisch and Vercoe 1984), and are quicker to respond
to environmental stimuli (Lees et al. 2018). Another adaptation that allows cattle to
grow and reproduce when ingesting low-protein diets is nitrogen (N) recycling to the
rumen (Silva et al. 2019). This is an important physiological mechanism allowing
3 Adaptation of Beef Cattle to Heat Stress Challenges 31

Fig. 3.1 Distribution of cattle around the world (Cooke et al. 2020a, b)

ruminants to grow efficiently when on low-protein diets (Silva et al. 2019). Selection
for improved N recycling may be a useful addition to selection programs. Globally
there are >800 cattle breeds, so selection for cattle that are heat tolerant is doable, but
there are potentially associated costs, which will be discussed below.
Beef production in the tropics and sub-tropics is characterised by climate
variability (heat and rainfall), nutritional limitations, parasites, disease, and poor
reproductive performance (Gaughan and Cawdell-Smith 2015). Cattle are therefore
exposed to multiple stressors such as droughts, floods, poor nutrition, and climate
stress (which can be both cold and heat stress). Even though our focus here is on heat
stress it should not be looked at in isolation. There is a need to move away from
traditional selection methods to more advanced molecular and large-scale genetic
evaluations that consider the whole genome, not just a few specific traits.

3.2 Adaptation in Cattle: For a Stressful Environment

One could argue that adaptation to, for example heat, occurs as natural selection in
beef cattle in the tropics and sub-tropics. And while this is true, this adaptation
comes at a cost (a trade-off) to other traits such as reproduction and growth (Frisch
and Vercoe 1984). The trade-off is essentially production verses survivability. It is
widely accepted that selection for heat tolerance, parasite resistance, and the ability
to do well where nutrition is poor, generally has negative impacts (from our
perspective not the animals) on fertility, growth, and milk production (Frisch and
Vercoe 1984; Fordyce et al. 1993). Longevity of adapted cattle is another key factor
that needs to select for (Cooke et al. 2020a). Our challenge then is to select animals
that are adapted to the stressful environment and still retain adequate production
32 J. B. Gaughan et al.

performance. It is likely that simultaneous genetic improvement of productive and

adaptive traits in tropically adapted breeds of beef cattle raised in tropical
environments is feasible without major impacts on adaptive or production traits
(Burrow 2012). However, further research is required to quantify the genetic antag-
onism between adaptation and production traits to evaluate the potential selection
response (Renaudeau et al. 2012).

3.3 Mechanisms of Beef Cattle Adaptation to Heat

Animal adaptation is a function of a number of factors, essentially it is a function of

Animal  Human  Resources. A large part of future adaptation will depend on how
well we use the available resources i.e., land, feed, water, financial, animals, and
people. A large part of this will be which genetic technologies we can afford to use.
Before we get to that we need to decide: What do we want from our animals? And
what adaptive traits should we be looking for? These traits could be any combination
that improve the survivability, fertility, production (growth, milk, meat), product
quality or animals that support economically and financially sustainable systems.
The needs will vary from location to location and will be influenced by operation
size and individual limitations.
Key to our understanding of a breed or species ability to adapt, is to define the
stressor(s), and define what we want the animals to adapt too (Gaughan et al. 2019).
We need to consider the following. How long does it take for adaptation to occur?
We should start breeding now for a predicted climate in 30 years’ time, but what if
the predictions are wrong? Can we breed animals (our current breeds and/or species)
for the level of tolerance we think they may need? What do we select for? How do
we select animals to use? Are controlled studies a good indicator of animal responses
in the field?
Some proposed strategies have been to identify stress tolerant animals within
non-tolerant breeds e.g., within the Angus breed, and the use existing already
adapted animals. There are large differences in thermal tolerance within and between
breeds, so the opportunity exists the use of molecular techniques to improve thermal
tolerance of beef cattle. However, further research is required to quantify the genetic
antagonism between adaptation and production traits to evaluate the potential
selection response. With the development of molecular biotechnologies, new
opportunities are available to characterize gene expression and identify key cellular
responses to heat stress. These new tools will enable scientists to improve the
accuracy and the efficiency of selection for heat tolerance. Epigenetic regulation of
gene expression and thermal imprinting of the genome could also be an efficient
method to improve thermal tolerance (Renaudeau et al. 2012). However, Monteiro
et al. (2016) reported long-term negative epigenetic effects (lower birth weight,
lower weaning weight and decreased performance through first lactation) when
dairy calves were exposed to heat stress in utero. Later studies (Ouellet et al.
2020) have confirmed that heat stress in late gestation negatively impacts the dairy
3 Adaptation of Beef Cattle to Heat Stress Challenges 33

cows’ productivity and health, that in utero heat stress alters mammary development
of the dam and the foetus, and importantly that in utero heat stress programs a lower
productivity phenotype in offspring. Furthermore, there is evidence that offspring
from prepartum heat-stressed cows have compromised passive immunity and
impaired cell-mediated immune function (Tao and Dahl 2013). It is not known at
that time if similar responses would be seen in beef cattle, but it is likely.
Traits such as coat colour (Brown-Brandl et al. 2006; Fanta 2017; Dikmen et al.
2017), heat shock protein expression (Collier et al. 2008), metabolic changes
(Collier et al. 2012; Baumgard and Rhoads Jr 2013; Aleena et al. 2018), maintenance
of feed intake (Verma et al. 2000) and performance have all been promoted as
selection options for improved heat tolerance. Genes affecting hair and skin colour,
hair length and density have been identified in cattle (Collier et al. 2008). Heat
tolerance in Bos taurus beef cattle with the ‘slick hair gene’ is well established
(Olson et al. 2003). However, the incorporation of these traits across breeding
programs has not had a large effect on improving performance of cattle in hot
environments, which highlights the difficulties in selecting for heat tolerance in
non-tolerant breeds.
We need to accept that productivity levels in the tropics will never match what has
been achieved in temperate regions. Selection must be realistic and in sync with the
available resources, the prevailing, and future climate. There are currently more
questions and unknowns than available answers.

3.4 Constraints to Adaptation

Probably the biggest constraint to adaptation is the genetic capacity of the animal.
Given time most species have capacity to meet the direct challenges of a changing
climate. Indirect effects such as habitat changes and human activities are more
difficult predict and breed for. All aspects of the environment are interlinked;
therefore, we cannot attempt to make a change in one area without understanding
the overall impact of the change on the affected species—including humans. A key
factor in animal adaptation is human adaptation and willingness to change.
The ability of animals to adapt to climate change will largely rest with how we
manage the change. The system is very complex!

3.5 Beef Breeding Programs to Improve Tolerance to Harsh

Environments

The use of natural selection, and traditional selection methods as the means of
adaption to heat stress is not logical. Given the rapid changes in climate that animals
are being exposed too we need to have methods that allow for rapid genetic change.
Especially considering the generation interval of these animals, we only have one or
two generations to make substantive change.
34 J. B. Gaughan et al.

Genetic improvement in animal populations through conventional improvement

programs mainly involves the selection of males and females that, when mated, are
expected to produce progeny that perform better than the average of the current
generation (Khare and Khare 2017). Performance is not (or at least should not be)
focused on a single trait but on a combination of multiple characteristics, or traits,
most of which are quantitative in nature. Quantitative traits that are controlled by
multiple genes (>100 to perhaps thousands) which are affected by environment
(Khare and Khare 2017). Therefore, selection for traits that are strongly influenced
by the environment and have low heritability e.g., fertility, are not only difficult but
require genetic the assessment of large numbers of animals (Hayes et al. 2019; Khare
and Khare 2017).
Traditionally the main criteria that has been used to identify individuals to be used
for breeding are estimates of their breeding values for the trait or traits of interest.
The breeding value of an individual is the sum of the additive effects of all loci that
contribute to the trait (quantitative trait loci), deviated from the population mean
(Falconer and MacKay 1996 cited by Khare and Khare 2017). The main source of
information for the estimation of breeding values of selected individuals are exten-
sive data bases of recorded phenotypes for the traits of interest, or for traits that are
genetically correlated to the traits of interest. These data bases are largely focused on
animals located in temperate regions and there is little information for breeds in the
tropics (Khare and Khare 2017).
Some traits of interest are only able to be recorded late in life (longevity), only on
one sex (milk yield), require animals to be sacrificed (meat quality), or require
animals to be exposed to conditions that would hamper the ability to market or
export their germplasm (disease resistance) (Dekkers 2012). These phenotyping
constraints limit the amount of genetic progress that can be made through conven-
tional selection and breeding method (Khare and Khare 2017).

3.6 Molecular Genetics the Way Forward?

Pollak (2005) outlined the numerous tools that can be applied to beef cattle breeding,
including: (i) reproductive innovations that focus on manipulating the physical
process of reproduction (e.g., artificial insemination, embryo transfer, cloning) or
reproductive cells (e.g. semen sexing, nuclear transfer); (ii) statistical methodologies
and computational algorithms designed to implement those methodologies; (iii)
‘mechanical’ inventions that provide mechanisms to capture new phenotypes, such
as for the capture of individual feed consumption, images for indicating tenderness,
and ultrasound measures of carcase attributes; and (iv) procedures that provide
information on DNA, generally characterised as laboratory detection of DNA
polymorphisms. While all of these have merit it is the use of molecular genetics
that is showing the best promise for rapid genetic change.
Molecular genetics is the field of science that combines molecular biology and
genetics. It is the study of the structure and function of genes, but at the molecular
level (Fulton 2008). Molecular genetics is the study of the sequence of genes as well
3 Adaptation of Beef Cattle to Heat Stress Challenges 35

as the structure of the protein it codes for. Molecular genetics allows for the
identification and mapping of genes and genetic polymorphisms i.e., whole genome
sequence analysis. Molecular genetics techniques make it possible to identify genes
that are involved in variety of traits, thereby allowing selection of animals on the
basis of their genetic makeup. The use of molecular information in selection
programs has the potential to increase productivity, enhance environmental adapta-
tion and maintain genetic diversity. The use of molecular genetics technologies
potentially offers a way to select breeding animal at an early age (even embryos);
to select for a wide range of traits and to enhance reliability in predicting the mature
phenotype of the individual (Khare and Khare 2017).
Molecular techniques have the potential for rapid changes in selection criteria for
animals used in breeding programs based on their phenotypes. Molecular techniques
offer the potential for selecting animals that are resistant to disease. Molecular
techniques allow selection for complex traits such as reproductive efficiency, and
feed efficiency in animals (Khare and Khare 2017; Mrode et al. 2019). The use of
molecular genetics techniques should be used in association with conventional
animal breeding tools (e.g., still need to assess for conformation faults) so to
optimize the animal breeding program (Singh et al. 2014). Molecular techniques
may allow the selection of animals and develop animal lines that are suited to
particular environments, and rapid genetic change in the face of climate change.
Genomic selection is an attractive alternative to traditional breeding as it will
allow young bulls (and heifers) to be selected on their heat tolerance genomic
estimated breeding value (GEBV) as well as on other traits (Nguyen et al. 2016).
The work of Nguyen et al. (2016) has resulted in the implementation plan of the
Australian genomic breeding values (ABVg) for heat tolerance (HT) (HT ABVg) for
Holstein and Jersey dairy cattle (Nguyen et al. 2017). However, the authors reported
that there has been a deterioration in the genetic trend of HT, and to moderate the
decline they suggested that the HT ABVg should be included in a multi-trait
economic index with other traits that contribute to farm profit (Nguyen et al.
2017). This highlights that there is still a need for traditional multi-trait selection.
The use of this technology is limited in many because most of the production
occurs in small holder systems which are characterized by small herd sizes, lack of
performance, and pedigree recording and therefore, the non-existence of conven-
tional genetic evaluation systems (Kosgey and Okeyo 2007; Nguyen et al. 2016;
Mrode et al. 2019). Therefore, there is a need for interventions that will allow these
societies to reap the benefits of the new genetic revolution.

3.7 Advantages of Molecular Approach Over Traditional

Approach

The following summary of the advantages of molecular genetics over traditional

breeding has been adapted from Khare and Khare (2017)1 and Barendse (2017).2
36 J. B. Gaughan et al.

• Genetic improvement of tropical cattle must move beyond the contemplation of

breeds and breed characteristics.2
• The use of genomic breeding values (GEBV) in tropical cattle will require
improvements in methods of genetic prediction for crossbreed cattle.2
• Assuming no genotyping errors, molecular genetic information is not affected by
environmental effects and therefore has heritability equal to one.1
• Molecular genetic information can be available at an early age, in principle at the
embryo stage, thereby allowing early selection and reduction of generation
intervals.2
• Molecular genetic information can be obtained on all selection candidates, which
is especially beneficial for sex-limited traits, traits that are expensive or difficult to
record, or traits that require slaughter of the animal (carcase traits).1
• Molecular genetic information enhances reliability in predicting the mature
phenotype of the individual and allows selection for a wide range of traits.1
• High-level genome sequence coverage of adapted breeds needs to be obtained
before the animals become rare, because there is the possibility of discovering
unique or unusual genes affecting adaptation to tropical areas.2
• DNA tests for GEBV have not reached commercial application for tropical cattle
owing to a lack of predictive accuracy and infrastructure to implement GEBV.2

However, there are still limitation in applying the technology in the field in many
locations. There is an urgent need to develop programs that will enhance genetic
improvement across both large-scale and small-scale beef production.

Learning Outcomes
• Beef consumption represents about 25% of global meat consumption and is
expected to increase from 54 to 118 million tonnes by 2050.
• Traits to be targeted in breeding program for climate resilience should be a
combination that improve the survivability, fertility, production (growth,
milk, meat), product quality or animals that support economically and
financially sustainable systems.
• Genomic selection is an attractive alternative to traditional breeding as it
will allow young bulls (and heifers) to be selected on their heat tolerance
genomic estimated breeding value (GEBV) as well as on other traits.

3.8 Conclusions

Livestock which are adapted to heat stress whilst still maintaining adequate produc-
tivity (growth, fertility etc.) is a challenge. Breeding strategies that will lead to rapid
genetic change are required to meet the challenge of rapid climate change. The
development of genomic breeding values is an important step forward but needs to
be considered as a tool alongside traditional breeding and selection strategies. The
3 Adaptation of Beef Cattle to Heat Stress Challenges 37

use of molecular genetics will play a major role in the selection of heat tolerant
animals however the inability to collect sufficient performance and pedigree data in
many regions will limit the use of tools such as GEBV for some time.

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Behavioural Responses of Domestic
Animals for Adapting to Thermal Stress 4
Vinícius F. C. Fonsêca, Edilson P. Saraiva, José D. C. dos Santos,
Larissa Kellen da Cunha Morais, Sheila T. Nascimento,
Cíntia C. de Melo Costa, Gustavo B. Moura,
Geni Caetano Xavier Neta, Kênia C. Bícego, Veerasamy Sejian,
Marcos Chiquitelli Neto, and Alex S. C. Maia

Contents
4.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
4.2 Body Size, Heat Exchanges and Importance of Thermoregulatory Behavior . . . . . . . . . . . . 42
4.3 The Role of Behavioural Thermoregulation: Precocial Domestic Neonates . . . . . . . . . . . . . . 43
4.4 Role of Behavioural Thermoregulation: Shade-Seeking Behaviour and Body Orientation
Adjustments for Livestock Grazing in Equatorial Semi-Arid Regions . . . . . . . . . . . . . . . . . . . . 45
4.4.1 Body Orientation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
4.5 Future Studies: Behavioural Plasticity of Free-Ranging Livestock . . . . . . . . . . . . . . . . . . . . . . . 48
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48

V. F. C. Fonsêca (*)
Animal Biometeorology and Ethology Group (BIOET), Department of Animal Science, Federal
University of Paraiba, Areia, Brazil
Innovation Group of Thermal Comfort and Animal Welfare (INOBIO-MANERA), Biometorology
Laboratory, São Paulo State University, Jaboticabal, Brazil
e-mail: [email protected]
E. P. Saraiva · J. D. C. dos Santos · L. K. da Cunha Morais
Animal Biometeorology and Ethology Group (BIOET), Department of Animal Science, Federal
University of Paraiba, Areia, Brazil
S. T. Nascimento
Innovation Group of Thermal Comfort and Animal Welfare (INOBIO - MANERA),
State University of Maringá, Maringá, Brazil
C. C. de Melo Costa · G. B. Moura · M. C. Neto · A. S. C. Maia
Innovation Group of Thermal Comfort and Animal Welfare (INOBIO-MANERA), Biometorology
Laboratory, São Paulo State University, Jaboticabal, Brazil
G. C. Xavier Neta · K. C. Bícego
Department of Animal Morphology and Physiology, Sao Paulo State University, UNESP/FCAV,
Jaboticabal, Brazil
V. Sejian
ICAR-National Institute Animal Nutrition and Physiology, Bengaluru, India

# The Author(s), under exclusive license to Springer Nature Singapore Pte 39

Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_4
40 V. F. C. Fonsêca et al.

Abstract

Animals use behaviour to alleviate the feelings of hot and cold, by avoiding
situations that arouse those feelings. That is the basis of behavioural thermoregu-
lation. By employing behavioural thermoregulation, domestic animals can
manipulate heat exchanges between body surface and surrounding environment,
thus reducing demands for evaporative water loss and heat production in order to
regulate body temperature. Availability of buffered microclimates and possibility
for animals to employ behavioural thermoregulation is of utmost importance for
those that have a high surface area to volume ratio. By observing the animal’s
behaviour, farmers can easily predict how animals are coping with thermal
challenges and to get insights about their preferences. This chapter briefly
discusses the importance of behavioural thermoregulation under the context of
domestic animals. Specifically, (1) the association between body size and rele-
vance of thermoregulatory behaviour; (2) evidences on how shade or shelter-
seeking behaviour and body orientation adjustments helps neonates and adult
animals to save water and energy; and ultimately (3) to prospect future studies
involving behavioural thermoregulation of free-ranging livestock, particularly to
better predict how they use some kind of behavioural plasticity to buffer direct
and indirect impact of climate change.

Keywords

Animal behaviour · Climate change · Heat stress · Thermoregulation ·

Thermoneutral zone

Abbreviations

% Percentage
> Greater than


C Degree Celsius
e.g. Exempli gratia
g Gram
h Hours
i.e. id est (that is)
LCT Lower critical temperature
m2 Square meter
SSH Source of supplemental heat
TRM Mean radiant temperature
Ts Surface temperature
UCT Upper critical temperature
W m2 Watt per square metre
4 Behavioural Responses of Domestic Animals for Adapting to Thermal Stress 41

4.1 Introduction

The voluntary changes in behaviour are the most powerful mechanism for animals to
regulate body temperature, mostly because it requires lower energetic costs and can
be sustained longer than autonomic responses (e.g., panting, sweating, or
non-shivering thermogenesis) (Bicego et al. 2007). The behavioural thermoregula-
tion can anticipate changes in core body temperature, and therefore decreases the
demand for evaporative cooling or heat production mechanisms. This anticipatory
control is due to a well designed peripheral-neural network that involves early
detection of temperature changes in the skin, integration with the central nervous
system through the preoptic area in hypothalamus, thus evoking voluntary
behavioural changes in order to conserve body heat or enhance heat dissipation.
By altering the behaviour, domestic animals manipulate heat exchanges between
body surface and their surrounding environment, which enhances their capacity to
regulate body temperature within a thermal zone of low thermoregulatory effort
(Fig. 4.1), namely thermoneutral zone. When an animal is outside of its
thermoneutral zone, costs of thermal homeostasis in terms of water use to cope
with heat stress or chemical energy use to cope with cold stress increases (Fig. 4.1),
and consequently less energy is available for animal production. The body size is one
the most important feature that determines the width of the thermoneutral zone, as
well as, environmental threshold to employ the thermoregulatory behaviour. For
instance, when exposed to high levels of solar radiation, due to a higher

Fig. 4.1 Simple illustration of the thermal load thresholds delimiting zone of least thermoregula-
tory effort. LCT lower critical temperature, UCT upper critical temperature
42 V. F. C. Fonsêca et al.

mass-specific rate of radiant heat gain and heat loss, newborn lambs are likely to
employ shade-seeking behaviour earlier than their mothers. In some cases, there
must be a trade-off between the thermoregulatory behaviour and there will be a
strong motivation of newborns to be in close association with mothers, particularly if
the shade or shelter resources are placed in large paddocks.
By observing the behaviour of animals, farmers can immediately predict how
they are coping with the thermal challenges and to get good insights about their
preferences. These insights may help farmers to design more sustainable environ-
mental management strategies in order to alleviate impacts of thermal stress on
animal welfare. For instance, dairy cows and sheep kept in open field will prefer to
graze at times with solar irradiance levels below 650 W m2 (Oliveira et al. 2014).
Sheep will experience lower radiant heat load when using shades of photovoltaic
panels than the shade of a shade-cloth structure that blocks 90% of solar radiation,
thereby explaining their preference to use shade provided by the photovoltaic panels
(Maia et al. 2020). Here in this chapter we briefly discuss about the following aspects
involving behavioural thermoregulation under the context of domestic animals:
(1) association between body size and relevance of thermoregulatory behaviour;
(2) evidences on how shade or shelter-seeking behaviour and body orientation helps
neonates and adult animals to save water and energy; and ultimately (3) to prospect
future studies involving behavioural thermoregulation of free-ranging livestock,
particularly to better understand how they employ some kind of plastic behaviour
to buffer direct and indirect impact of climate change.

4.2 Body Size, Heat Exchanges and Importance

of Thermoregulatory Behavior

Several studies have been postulated that small body sized animals have advantage
over the larger ones to cope with high heat load, by claiming that a larger body
surface area to volume ratio makes them efficient heat dissipaters in warm climates.
Yes, they are, as rates of heat transfer scales against body mass with exponent 0.55.
However, lets rethink about this statement by considering drivers governing heat
exchanges between the animals and its surrounding environment, e.g., physical
requirements for body heat dissipation is that the surface temperature > operative
temperature (i.e., equivalent temperature for heat exchanges by convection and long-
wave radiation), while the opposite represents requirements for body heat gain.
Free-ranging animals grazing in arid and semi-arid conditions normally experi-
ence operative temperature greater than their body surface temperature during most
part of daytime, which favors heat influx from the surrounding environment. Under
such circumstances, due to a greater mass-specific dry heat gain, smaller animals
would gain environmental heat more rapidly, per kilogram of body mass, than the
larger animals. Naturally adapted small body sized goats and sheep, when exposed to
levels of solar irradiance above 800 W m2 and operative temperature above to
40
C, can absorb near to 350 W m2 of radiant heat (Maia et al. 2015). This heat
load can represent almost ten times the amount of their heat produced by
4 Behavioural Responses of Domestic Animals for Adapting to Thermal Stress 43

metabolism. To maintain their thermal equilibrium, goats substantially increase the

rate of evaporative water loss, particularly through cutaneous surface. Again, due to
a greater mass-specific rate of evaporation, more rapid dehydration is expected for
smaller animals than larger ones, particularly if they do not have opportunity to
replace fluids or to seek for more buffered microclimate.
While high levels of solar radiation and air temperatures of arid and semi-arid
areas result in high rates of heat influx on animals during daytime, direction of heat
exchange between the environment and animal is usually reversed at night (Fuller
et al. 2021). The low operative temperature of clear night skies can result in
substantial amount of radiant and convective heat loss (Mitchell et al. 2018),
particularly for small body sized animals due to its high mass-specific radiant heat
loss. Small body sized sheep living in semi-arid regions of Brazil will have difficulty
in conserving body heat at night, with dry heat loss alone having the potential to
double the metabolic heat production (Fonsêca et al. 2019).
We may predict therefore that smaller animals are more susceptible to thermal
challenges than the larger ones. Indeed, it is, unless they have the opportunity to find
more buffered microclimate. Historically, larger animals were more vulnerable to
other past significant changes on earth climate (Mitchell et al. 2018), and that
vulnerability of larger animals is probably related to factors such as their inability
to easily access more buffered microclimates, and on the other hand, may explain the
advantage of small body sized animals.

4.3 The Role of Behavioural Thermoregulation: Precocial

Domestic Neonates

Although precocious neonates (e.g., newborn lambs, goats, calves) born with a well-
developed thermoregulatory system that help them to maintain body temperature
within a narrow range, high rates of neonatal mortality due to thermal stress are still a
prevalent problem to animal production around the world. Apart from the risk factors
to neonatal mortality such as poor maternal care, birth trauma, and infectious
diseases, maintenance of body temperature can be challenging for the survival of
newborns, mostly due to their poor insulation, small body size and consequent high
body surface area to volume ratio.
Depending on the level of cold exposure after birth, the brown adipose tissue
reserves and colostrum intake, which are the main sources of heat production, may
not be sufficient to offset the rates of heat loss of newborn lambs to the environment.
Rates of heat loss can be decreased if they have opportunity to employ thermoregu-
latory behaviour, by seeking a more buffered microclimate, e.g., a well designed
shelter can reduce heat loss by convection and radiation over the body surface of
newborns. Additionally, if supplementary heat is provided by using heat lamps, the
employment of thermoregulatory behaviour can also enhance radiant heat gain to
newborns. On the other hand, neonates delivered under an equatorial semi-arid
condition can also face high levels of radiant heat load over the year, e.g., levels
of impinging solar radiation up to 1000 W m2 and mean radiant temperatures up to
44 V. F. C. Fonsêca et al.

Fig. 4.2 Black Santa Ines newborn lamb exposed to direct solar radiation in an equatorial semi-
arid region. Thermal image taken by using an infrared thermal camera (Fluke—Model TiX500
9 Hz; range: range: 20
C and +650
C; accuracy: 0.1
C/1
C; emissivity: ε ¼ 0.98, Everett,
WA, USA) at a distance of 1.0 m from the animal. Meteorological conditions at time which image
was taken: Ta: 35
C; RH: 35.3%; RS: 900 W m2; WS: 3.7 m s1. (Photo by J.D.C.S.)

34

32
Black globe temperature, ºC

30

28

26 Source of supplemental heat

Open area
24

22

20

18

18 19 20 21 22 23 0 1 2 3 4 5 6
Time, h

Fig. 4.3 Newborn goats within the source of supplemental heat (SSH) and thermal load experi-
enced (mean  SEM) by them within the SSH and in open area. (Photo by V.F.C.F.)

60
C (Fig. 4.2). Under such circumstances, autonomic responses such as panting

and sweating may not be sufficient to offset the rates of heat gain of newborns from
the environment. The shade-seeking behaviour may therefore play an important role
to decrease heat load influx on neonates.
In maternity corral pens, we observed that newborn goats delivered under tropical
conditions voluntarily accessed a private source of supplemental heat (SSH) over
nocturnal periods, which efficiently allowed them to conserve body heat, and to
escape from low levels of radiant temperatures in the open environment. When the
newborns were within the SSH (Fig. 4.3), they were benefited with a lower thermal
gradient between its surface temperature and the mean radiant temperature (mean
Ts ¼ 30
C vs. mean TRMSSH ¼ 32
C), than if they were experiencing radiant
4 Behavioural Responses of Domestic Animals for Adapting to Thermal Stress 45

temperature of the open environment (mean Ts ¼ 28
C vs. mean TRM ¼ 20
C).

Over the daytime, newborn goats also employed shade-seeking behaviour, which
benefited them to avoid high levels of impinging solar radiation. Similarly, our
recent study with newborn lambs from two hair coat sheep breeds also showed
that behavioural thermoregulation as shade-seeking appeared to play an important
role in order to defend the body rectal temperature to rise.

4.4 Role of Behavioural Thermoregulation: Shade-Seeking

Behaviour and Body Orientation Adjustments
for Livestock Grazing in Equatorial Semi-Arid Regions

Grazing cows under tropical conditions may face incoming solar radiation up to
1000 W m2, which have potential to absorb as much as 640 W m2 of thermal
energy. If shade is not available, how a dairy cow would manage this amount of
radiant heat gain? By modeling a thermal balance, requirements for evaporative
water loss may be close to 900 g h1 to maintain their thermal equilibrium. High
levels of solar radiation also impose challenge for sheep naturally adapted to hot
conditions. We investigated the impact of solar radiation on the thermal equilibrium
of hair-coat sheep raised under natural conditions in an equatorial semi-arid region of
Brazil. When exposed to levels of solar radiation near to 850 W m2, sheep had
potential to absorb up to 350 W m2 of thermal energy by long and short wave solar
radiation. Under such circumstances, by accounting for heat generated through
metabolism (45 W m2), absorbed from thermal radiation (350 W m2), minus
that eliminated through the respiratory evaporation (20 W m2), and then solving for
the evaporative requirements to maintain their thermal equilibrium, sheep may need
to evaporate up to 500 g m2 h1 of sweat through the skin surface to offset the
accumulated heat.
Livestock that grazes unprotected against solar radiation will therefore need a
large amount of water to sustain high level of evaporative water loss (Mitchell et al.
2018), an increasingly limited natural resource, particularly for livestock species
living in arid and semi-arid regions. However, they can behaviorally manipulate
their thermal equilibrium by seeking shade (Fig. 4.4), thus reducing sensible heat
gain and requirements for evaporative cooling.
The solar radiation is the most important variable for predicting shade use by
livestock in the tropical environment (Oliveira et al. 2019). The critical level of solar
radiation that motivates dairy cows to stop grazing and seek shade is in the interval
between 500 and 700 W m2 (Oliveira et al. 2014). In equatorial semi-arid regions,
this level of solar radiation is normally recorded after 08:30 h and prior to 16:00 h,
suggesting that cows under pasture based systems should have access to paddocks in
the early hours of day, when the solar radiation is well tolerated, and consequently
cows can devote more time in grazing. Moreover, farmers may implement nocturnal
grazing to complement hours of grazing. Figure 4.5 illustrates how the understand-
ing of shade requirements of domestic animals gives important insights on how to
46 V. F. C. Fonsêca et al.

Fig. 4.4 Shade-seeking behaviour of hair coat sheep during the hottest hours of the day in an
equatorial semi-arid region. (Photo by V.F.C.F)

employ environmental management strategies to improve the thermal comfort and

welfare of grazing dairy cows.

4.4.1 Body Orientation

When shade is not available, changes in body orientation allow animals to manipu-
late the radiant heat gain. This subject is relatively well investigated with animals
living in high latitudes. For instance, in cold days animals may increase absorption
of radiant heat by exposing body long axis perpendicular to the incoming solar
radiation. Oppositely, in hot days animals can reduce surface area exposed to solar
radiation by orienting long axis parallel to the sun’s rays, thus minimizing heat load.
However, studies evaluating changes in body orientation of livestock grazing in low
latitude regions are scarce. When compared to subtropical and temperate latitudes,
the incoming radiation reaches the surfaces of equatorial regions at a highly elevated
angle in relation to horizon over the year. At high solar elevation, we expected that
the benefits in terms of reducing heat load when animals orient their body axis
parallel to incoming radiation may be reduced. Preliminary investigation performed
by our team shows that the body axis orientation seems not to be employed by black
and white hair coat sheep as thermoregulatory strategy when exposed to solar
radiation in an equatorial semi-arid environment. More studies however are needed
to better understand the role of adjustments in body orientation on heat balance of
livestock living in arid and semi-arid equatorial regions.
4 Behavioural Responses of Domestic Animals for Adapting to Thermal Stress 47

Fig. 4.5 Proposed daily routine adaptation for dairy cows raised in equatorial semi-arid regions,
according to the level of solar radiation tolerated by cows (Oliveira et al. 2014). Levels of solar
radiation (RS, W m2) recorded in the study were divided into five classes: Class
1 (RS < 300 W m2), class 2 (300 W m2 < RS  500 W m2), class
3 (500 W m2 < RS  700 W m2), class 4 (RS  900 W m2), class 5 (RS > 900 W m2)

Learning Outcomes
• Behavioural thermoregulatory mechanism is one of the primary means by
which the farm animals cope with thermal stress.
• When compared with large body sized animals, smaller livestock have
better opportunities to seek more buffered microclimates to cope with
extremes in weather.
• Shade-seeking is one of the important behavioural thermoregulatory
mechanisms exhibited by neonates to maintain body temperature during
heat stress exposure.
• Understanding the extent of behavioural plasticity exhibited by livestock is
considered a most promising strategy to identify climate resilient breeds.
48 V. F. C. Fonsêca et al.

4.5 Future Studies: Behavioural Plasticity of Free-Ranging

Livestock

Naturally adapted livestock living in arid and semi-arid regions are facing an
increasing heat load, in addition to food and water shortages, as consequences of
the direct and indirect impacts of climate change. This is occurring at a faster and
more intense rate than the time required for genetic adaptation, leaving only the
expression of phenotypic plasticity of small ruminants to adjust to the new environ-
mental conditions. The compound effects of thermal stress, and lack of food and
water, may impair the ability of dry land ruminants to sustain the high rates of
evaporative water loss and heat production in order to maintain their core body
temperature within a narrow range. Recent findings in wildlife have shown that these
animals employ a plastic behaviour by changing daily activity, grazing patterns, and
microclimate selection to cope with seasonal changes in heat load, food and water
availability. Whether this can be confirmed for farm livestock raised exclusively in
various arid and semi-arid biotopes around the world, still needs to be investigated.
The use of miniaturized animal-implantable devices for logging data recording the
movement and microclimate selection over the extended periods of time will there-
fore elucidate if they use some type of behavioural plasticity in response to heat load,
and lack of food and water. Ultimately, understanding the extent of this behavioural
plasticity will help us to identify the resilient breeds of livestock, and better predict
their potential to adapt to the rapid changes in climate.

References
Bicego KC, Barros RC, Branco LG (2007) Physiology of temperature regulation: comparative
aspects. Comp Biochem Physiol Part A Mol Integrative Physiol 147(3):616–639
Fonsêca VDFC, Maia ASC, Saraiva EP, de Melo Costa CC, da Silva RG, Abdoun KA, Al-Haidary
AA, Samara EM, Fuller A (2019) Bio-thermal responses and heat balance of a hair coat sheep
breed raised under an equatorial semi-arid environment. J Therm Biol 84:83–91
Fuller A, Mitchell D, Maloney SK, Hetem RS, Fonsêca VF, Meyer LC, van de Ven TM, Snelling
EP (2021) How dryland mammals will respond to climate change: the effects of body size, heat
load and a lack of food and water. J Exp Biol 224(Suppl_1):jeb238113
Maia ASC, da Silva RG, Nascimento ST, Nascimento CCN, Pedroza HP, Domingos HGT (2015)
Thermoregulatory responses of goats in hot environments. Int J Biometeorol 59(8):1025–1033
Maia ASC, de Andrade Culhari E, Fonsêca VDFC, Milan HFM, Gebremedhin KG (2020) Photo-
voltaic panels as shading resources for livestock. J Clean Prod 258:120551
Mitchell D, Snelling EP, Hetem RS, Maloney SK, Strauss WM, Fuller A (2018) Revisiting
concepts of thermal physiology: predicting responses of mammals to climate change. J Anim
Ecol 87(4):956–973
Oliveira SEO, de Melo Costa CC, de Souza JBF, de Queiroz JPAF, Maia ASC, de Macedo Costa
LL (2014) Short-wave solar radiation level willingly tolerated by lactating Holstein cows in an
equatorial semi-arid environment. Trop Anim Health Prod 46(8):1413–1417
Oliveira SEO, de Melo Costa CC, Neto MC, Dalla Costa FA, Maia ASC (2019) Effects of shade
location and protection from direct solar radiation on the behavior of Holstein cows. Int J
Biometeorol 63(11):1465–1474
Heat Stress Associated Changes
in the Immune System Related Responses 5
in Sheep

Mariangela Caroprese, Maria Giovanna Ciliberti, Marzia Albenzio,

and Agostino Sevi

Contents
5.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
5.2 Sheep Endocrine Responses to Heat Stress . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
5.3 Sheep Immune Responses Functioning . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
5.4 Sheep Immune Responses to Heat Stress . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
5.5 Future Perspective . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58

Abstract
Very few studies have dealt with sheep under heat stress, with their physiological
mechanisms of adaptation and, above all, with alterations on sheep immunologi-
cal responses. This chapter will focus on the complex network of mechanisms
activated by heat stress as affecting immune responses in sheep. In particular, heat
stress will be discussed as a particular typology of stress response, with the
description of the changes induced by heat stress on both innate and adaptive
immunity, as well as on cellular functioning. Finally, the study of innovative
indicators of cell-to-cell communications will be delineated as future perspectives
for the fully understanding of the changes in sheep immune responses as induced
by heat stress.

Keywords
Cytokines · Heat stress · Immune responses · Sheep · Stress response

M. Caroprese (*) · M. G. Ciliberti · M. Albenzio · A. Sevi

Department of Agriculture, Food, Natural Resources, and Engineering (DAFNE), University of
Foggia, Foggia, Italy
e-mail: [email protected]

# The Author(s), under exclusive license to Springer Nature Singapore Pte 49

Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_5
50 M. Caroprese et al.

Abbreviations

% Percentage

C Degree Celsius
ATP Adenosine triphosphate
B cells B lymphocytes
CD Cluster of differentiation
cDNA Complementary deoxyribonucleic acid
CLR C-type lectin receptor
COX-2 Cyclooygenase-2
DAMP Danger associated molecular pattern
DC Dendritic cell
ECM Extracellular matrix
Fig Figure
h Hour
HMGB1 High mobility group box protein 1
HPA Hypothalamic-pituitary-adrenal
HS Heat stress
HSP Heat shock protein
IFN Interferon
IL Interleukin
iNOS Inducible nitric oxide synthase
miRNA Micro RNA
mL Millilitre
mRNA messenger ribonucleic acid
NF-κB Nuclear factor kappa B
ng Nanogram
NK Natural killer
NLR Nucleotide-binding domain and leucine-rich repeat containing receptor
NO Nitric oxide
PAMP Pathogen-associated molecular pattern
PBMC Peripheral blood mononuclear cell
PGE2 Prostaglandin E2
PHA Phytohemagglutinin
PRR Pattern recognition receptor
RIG-I Retinoic-acid inducible gene I
RLR Retinoic-acid inducible gene I like receptor
ROS Reactive oxygen species
T cells T Lymphocytes
T3 Triiodothyronine
T4 Thyroxine
TCR T cell express cell surface receptor
Th cells T helper cells
TLR Toll-like receptor
5 Heat Stress Associated Changes in the Immune System Related Responses in Sheep 51

TNF Tumour necrosis factor

α Alpha
β Beta
γ Gamma
δ Delta

5.1 Introduction

The alterations of immune responses described during and after heat stress exposure
of livestock are only one typology of the complex system of responses activated to
restore body homeostasis after a stressor. The immune responses are part of the
ancestral and conserved wider response system named stress response, which has the
aim to preserve life and to help coping with aversive situations. Based on this,
defense and adaptation mechanisms activated by livestock define their success to
survive in hostile environments. Heat stress response is a well characterized type of
stress responses which affects even sheep that are considered as one of the most
resistant species to climatic extremes, especially to high ambient temperatures. The
compensatory physiological, endocrine, and behavioral mechanisms activated by
sheep with the intention to survive, and to cope with high environmental conditions
are numerous. After the stimulation of temperature receptors into the hypothalamus,
sheep increase their respiration rate, rectal temperature, and exhibit an alteration of
protein and energy metabolism, of mineral balance, enzymatic reactions, and hor-
monal secretions (Sevi and Caroprese 2012). The first responses including increased
water intake, sweating and respiration rates, are considered homeostatic mechanisms
(Horowitz 2002). If the exposure to thermal extremes is prolonged, heat acclimation
is achieved by an acclimatory homeostasis (Horowitz 2002). It has been proposed
the definition of acclimation as the altered expression of pre-existing mechanisms
driven by the endocrine system in order to sustain animal welfare and health
regardless of environmental challenges. Based on this, acclimation should be con-
sidered a homeorhetic mechanism (Collier et al. 2005) because it alters the set-points
of homeostatic-related systems such as the carbohydrate metabolism rather than
acclimatory homeostasis (Horowitz 2002). However, the control of heat stress
response is characterized by a dynamic orchestration of physiological processes
that simultaneously influences multiple tissues and systems, which results in an
overall coordinated response, and it is mediated by altered responses to homeostatic
signals. The changes in immune responses observed in sheep under heat stress are
therefore the results of alterations in a number of tissues and systems to help the
animals to cope with heat load.
52 M. Caroprese et al.

5.2 Sheep Endocrine Responses to Heat Stress

Dairy animals under heat stress often display a depression of immune system
(Caroprese et al. 2012), which increases their susceptibility to diseases and some-
times lead them to death. Such a depression stems from alterations in particular
endocrine secretions, namely those from the pituitary (prolactin), adrenal (cortisol)
and thyroid glands (Triiodothyronine-T3 and Thyroxine-T4). Prolactin is a multi-
functional hormone involved in a variety of biological functions related mainly to
sustain galactopoiesis and lactogenesis; beside these actions prolactin exerts a
thermoregulatory role in small ruminants. In sheep, a significant association between
heat stress and increased prolactin mRNA expression in sweat glands was found,
thus leading to the hypothesis that prolactin can contribute to increase sweat gland
activity in order to reduce heat load (Choy et al. 1997). Based on previous statements
the increase in prolactin secretions during heat stress seems to improve the ability of
small ruminants to adapt to heat stress. The increase of plasma cortisol secretions in
sheep under heat stress is caused by the activation of hypothalamic-pituitary-adrenal
(HPA) axis with the aim of rising the circulating glucose. The cortisol secretion
during heat stress has the role of stimulating gluconeogenesis by the conversion of
non-carbohydrate molecules into glycogen in the liver to increase blood glucose
levels. The increase in circulating glucose during heat stress is essential for allowing
ruminants to cope with heat stress; therefore, similarly insulin has a crucial role in
post absorptive nutrient partitioning subsequent to heat stress. In dairy cows under
heat stress, basal insulin concentrations gradually increase, stimulating glucose
uptake via glucose transporter type 4 in muscles and adipose tissue, and being
responsible for the heat-induced hypoglycemia frequently reported. Insulin is also
a potent regulator of lipid metabolism as a potent antilipolytic hormone, and this may
explain why heat-stressed cows do not mobilize adipose tissue triglycerides
(Baumgard and Rhoads Jr 2013). On the contrary, recent studies found that both
in dairy ewes (Mehaba et al. 2021), and in dairy goats (Salama et al. 2014) no
increase in the level of insulin during heat stress is observed. Nevertheless, some
metabolic adaptations allowed dairy ewes to spare glucose and to avoid reductions in
milk yield (Mehaba et al. 2021). Ewes under heat stress have higher blood glucose
levels after glucose tolerance test than ewes under thermoneutral conditions imply-
ing the activation of both insulin- and non-insulin–mediated pathways. Further,
adipose tissue of heat stressed ewes became more resistant to the lipolytic signals.
Finally, in sheep under heat stress beside the activation of the pituitary gland and
HPA axis, the thyroid gland shows a reduced functioning, with a consequent
decrease of the secretion of thyroid hormones, triiodothyronine (T3), and thyroxine
(T4) in the attempt to thermoregulate. Such a complex and orchestrated alterations
observed during heat stress in ruminants are associated with and are the cause of
evident changes in immune responses and diseases susceptibility, and relevant
events occurring at cellular level.
5 Heat Stress Associated Changes in the Immune System Related Responses in Sheep 53

5.3 Sheep Immune Responses Functioning

An understanding of immune system structure and function in sheep is of great

importance to provide information in relation to the changes induced by stressful
events on immune functioning. Innate compartment of immune system consists of
physical barriers, soluble factors, and cells among which the most important are
macrophages and neutrophils; cells play an important role in innate immunity by
producing nitric oxide (NO), synthesized by inducible nitric oxide synthase (iNOS),
and prostaglandin E2 (PGE2) synthesized by cyclooxygenase (COX-2), that are
some of the inflammatory mediators. The cells of the innate immune systems, such
as dendritic cells (DCs), macrophages, neutrophils, natural killer (NK) cells and γδ T
lymphocytes, can recognize common structures shared by the vast majority of
microorganisms called pathogen-associated molecular patterns (PAMPs). PAMPs
are recognized through binding to receptors of the innate immune system named the
pattern recognition receptors (PRRs). Several classes of PRRs have been described,
including the Toll-like receptors (TLRs), the retinoic acid-inducible gene I (RIG-I)-
like receptors (RLRs), the nucleotide-binding domain and leucine-rich repeat
containing receptors (NLRs), the C-type lectin receptors (CLRs), and others. Each
TLR is an alert signal for the immune system able to activate the induction of
proinflammatory mediators as a result of the activation of innate immune system,
and to initiate the adaptive immune responses. Full-length cDNA sequences for all
ten TLR genes of the domestic sheep have been found, showing that sheep TLR
sequences share high similarity to cattle, pig, human and mouse genes. The cells of
adaptive immunity are B lymphocytes (B-cells) and T lymphocytes (T-cells). B-cells
are responsible for the secretion of specific antibodies, which enable phagocytes to
recognize microorganisms and destroy them by binding to antigenic sites on both
extracellular microorganisms and toxins. T-cells are the effector of cell-mediated
immune responses of adaptive immunity, are able to recognize and fight against
intracellular viruses and microorganisms. The T-cells express cell surface receptors
(TCRs) consisting of α and β chains in sheep. Among the α/β T-cells there two
different sub-lineages; the first sub-lineage expresses co-receptor molecule CD4
(cluster of differentiation), consists of cells known as T-helper cells (Th cells), and
can neutralize intracellular microorganisms, fungi, and parasites by interacting with
other immune cells, and activating an antigen-specific response; the second
sub-lineage expresses co-receptor molecules T-CD8, consists of cells known as T
cytotoxic cells, and can recognize and neutralize viral infected cells. Th-cells are
further divided into two different subsets: T helper cells type 1 (Th1 cells) and T
helper cells type 2 (Th2 cells). Another group of T-cells is characterized by γ and δ
chains and expresses functions of both innate and adaptive immune cells. Sheep
display a large number of γ/δ T-cells in the blood. In young cattle and sheep, up to
60% of PBMC can be γδ T cells, supporting the notion that they are important for
immunity early in life. TCR genes of both γ and δ chains are well characterized in
cattle, sheep, and in goats. The complex communications between the different cells
of the immune system, as well as the interactions between the immune system and
the stress responses has phylogenetically evolved in a diversified and complicated
54 M. Caroprese et al.

system of cytokines. Cytokines are also considered responsible for the complex
cross-talk between brain and immune system. Both cell-mediated and humoral
responses, as well as innate immunity, are activated and regulated by
proinflammatory cytokines, such as IL-6 and IL-1β. Interleukin-6 stimulates
hepatocytes, B cells, and cytotoxic T cells; IL-1β promotes leukocyte accumulation
in inflamed sites by inducing adhesion receptors on vascular endothelium. The
regulation of transcription of DNA, cytokine production, and cell survival is mainly
attributed to nuclear factor kappa B (NF-κB), a protein complex that controls
intracellular signal transduction pathways. Besides the pivotal role in controlling
the host response to stress and infection, cytokines can also have a negative role in
provoking tissues damages when produced in uncontrolled way. In order to avoid
this event, the cytokine secretion is tightly regulated.

5.4 Sheep Immune Responses to Heat Stress

The effect of stressors on the immune system is either a suppressive or a stimulatory

effect according to a number of factors such as duration and intensity of stressors,
basal animal health status, and also the type of indicators of immune response
measured (Ciliberti et al. 2017). When the animals undergo a stressor, a number of
endogenous danger signals, molecules that are actively secreted by cells in danger,
known as danger-associated molecular patterns or DAMPs, are recognized by the
PRR of the innate immune system. Recognized DAMPs are nucleic acids, ATP, uric
acid, heat-shock proteins (HSP), mitochondrial danger signals, high mobility group
box protein 1 (HMGB1), degradation products of the extracellular matrix (ECM).
The cytokines, such as IL-1β, TNF-α, and the interferons, are considered secondary
endogenous danger signals which, produced by activated immune cells upon PRR
triggering, induce the stimulation of innate and adaptive immunity. The
pro-inflammatory cytokines activate and sustain the signaling pathway and the
effector functions of the classic PRRs, such as the activation of the transcription
factor NFkB, and the induction of proinflammatory processes, respectively. This
type of reaction of the immune system to non-infectious agents is called “sterile
inflammation,” in which the innate immune system is activated by signals that do not
come from microorganisms but can stem from stressors of different nature. The
biology of stress, however, is extremely complex and even more complicated is the
number and type of the interactions between the neuroendocrine and immune
system; the immune system is affected by the stress response and, in turn, it affects
the stress response. In particular, heat stress response is a particular type of stress
response during which the described alterations of endocrine secretions have a
typical suppressive effect on immune responses through the alterations of cytokine
secretions. Conversely, the complex network of cytokines secreted exerts a crucial
regulatory role rules on the stress responses. Prolactin during heat stress can reduce
immune responses in PRL-signaling pathway genes associated with differential
cytokine secretion; particularly, prolactin reduces expression of TNF-α, and lym-
phocyte proliferation (Do Amaral et al. 2010). The cortisol secretion affects the
5 Heat Stress Associated Changes in the Immune System Related Responses in Sheep 55

Fig. 5.1 Percentage increases calculated to stimulated PBMC (CS) of (a) cell proliferation, (b)
IL-6, (c) IL-1beta, and (d) IL-10 levels in stimulated PBMC treated with 100 ng/mL of cortisol
(100) and stimulated PBMC treated with 1000 ng/mL of cortisol (1000) after acute stress (24 h),
chronic stress (96 h), normothermia (39  C) and hyperthermia (43  C) in vitro challenge. Stimulated
PBMC were treated with Phytohemagglutinin (PHA)

expression of a number of genes related to innate immune responses by a strong and

fast activity of down-regulation; especially, it can inhibit cytokine induced transcrip-
tion factors (NF-κB). However, it has been suggested that the reduction in cytokine
production following the down-regulation of NF-κB can lead to a reduction of the
glucocorticoid receptor, thus limiting the cell sensitivity to glucocorticoids and
allowing again the translation of NFkB1 (Sgorlon et al. 2012). The cross-talk
between glucocorticoids and cytokines is bidirectional; via negative feedback,
pro-inflammatory cytokines, such as IL-1, IL-6 and TNF-α, can increase the HPA
axis sensitivity to later stress challenges. However, the precise mechanisms of
glucocorticoids action on cytokine production and cell proliferation during stress,
and in particular during heat stress, are not fully explained. A number of factors can
influence the possible effects of cortisol on immune cell proliferation and cytokine
production, such as the magnitude of the stressor, and its duration (Ciliberti et al.
2017). Physiological levels of cortisol after a stressor can exert both
immunostimulatory and immunosuppressive effects in vitro depending on the dura-
tion of stress (Fig. 5.1). In contrast, levels of cortisol higher than physiological levels
exert immunosuppressive effects after both an acute stress and a chronic stressor
(Ciliberti et al. 2017). During heat stress, physiological levels of cortisol after a
56 M. Caroprese et al.

stressor decrease by 50% cell proliferation of immune cells when in presence of an

antigen (Caroprese et al. 2018). In addition, cortisol is able to regulate cytokine
production by sustaining pro-inflammatory cytokines levels even if during heat
stress cortisol drives the reduction of IL6 levels, as reported in Fig. 5.1 (Caroprese
et al. 2018). The described effects of cortisol on immune cell proliferation and
cytokine production are particularly apparent in the presence of an immune chal-
lenge highlighting that hyperthermia has a relevant inhibitory effect on immune
responses with a crucial reduction of immune competence. Such results from in vitro
study are confirmed by in vivo studies; in sheep under heat stress adaptive responses
are impaired as suggested by a decrease in cellular immune response after the
intradermal injection of mitogens (Sevi and Caroprese 2012; Caroprese et al. 2012).
The suppression of cell-mediated immunity during hyperthermia has been
attributed to a down regulation of Th1 cytokines in favour of the secretion of Th2
cytokines. Environmental conditions can influence the levels of Th1 (IFN-γ, and
IL-12), or Th2 cytokines (IL-10, IL-4, IL-13) secretion. In sheep, a reduction of both
IL-4 and IL-13 was measured during heat stress, also associated to a reduction of
cell-mediated immune response (Caroprese et al. 2014). It is stated that when the
Th2 responses diminish, mainly because of the reduction of IL-4 and IL-13 produc-
tion, concomitantly also the Th1 response appear inappropriate. Beside the reduction
of adaptive responses, heat stress can suppress also innate immune responses, as
demonstrated by the reduced reactivity in terms of phagocytosis and oxidative burst
measured as reactive oxygen species (ROS) production found in polymorphonuclear
cells (Lecchi et al. 2016). During the hot humid season associated with high cortisol
levels, milk neutrophils exhibit a reduction in phagocytic activity. The alterations in
adaptive and innate immune responses are, however, the result of a coordinated
system which modulates gene expression at cellular level in order to minimizing the
negative effects of heat stress on cellular functions and across a variety of cells and
tissues through the changes in endocrine secretions (Collier et al. 2008). The
principal response to heat stress at cellular level is the heat shock cellular response,
with the induction of HSPs, which develops rapidly, and shows the induced
alterations in 24 h. The expression of HSPs has a central role in cytoprotection,
and is the cellular response to the increase of oxidative stress, and of alterations in
protein synthesis, structure and functions. The cellular response to stress is also
modulated by the alterations of hormonal secretions induced by heat stress. Prolac-
tin, insulin, and catecholamines can contribute to the enhancement of HSP expres-
sion and proteins activity (Collier et al. 2008). Glucocorticoids, when enter the
cytoplasm, cause the release of preformed cytoplasmic HSP70 and 90, which are
bound to the glucocorticoid receptor. The hormone-receptor complex then moves
into the nucleus and enhances expression of HSP genes. The preformed HSP70 and
90 represent an instant pool of HSP to protect against protein denaturation and the
first line of defense against HS because the proteins are already in the cytoplasm and
do not require synthesis (Collier et al. 2008). Prolactin also modifies HSP gene
expression although the mechanism is not well known, but it seems to prolactin also
upregulate HSP 70 gene expression unknown (Collier et al. 2008). A crosstalk
between the endocrine secretions, HSPs, and the immune system is established
5 Heat Stress Associated Changes in the Immune System Related Responses in Sheep 57

with HSPs playing both a stimulatory and regulatory role. HSPs are considered
activators of the innate immune system capable of inducing the production of
pro-inflammatory cytokines but also to support immune homeostasis and to have a
dampening effect on immune activation. Intracellular HSPs can activate Treg cells
and inhibit immunity and inflammation. Extracellular HSP70 can exert an anti-
inflammatory response on innate immune responses by binding to cell surface
receptors and inducing IL-10 production.

Learning Outcomes
• Heat stress response causes alterations in endocrine secretions which have a
typical suppressive effect on immune responses through the alterations of
cytokine secretions.
• A crosstalk between the endocrine secretions, HSPs, and the immune
system is established during heat stress with HSPs playing both a stimula-
tory and regulatory role.
• Differed types of miRNA can regulate cellular responses and homeostasis
depending on the types of stressors, suggesting a significant role of miRNA
in thermal stress responses in animals.

5.5 Future Perspective

Heat stress affects immune responses of sheep by reducing the animals’ ability to
promptly cope with immunological challenges, thus being the main cause of
increased diseases susceptibility and even death during hot season. Stress responses
in general, and particularly heat stress responses, are able to perturb multiple
molecular pathways in the organism. At the same time, the molecular responses
try to restore functional and structural homeostasis in stressed cells and tissues. In the
recent years, micro molecules of small non-coding RNA (miRNAs) have been found
to control gene expression at the post-transcriptional level, being considered as novel
regulators in the mammalian stress response. It seems that differed types of miRNA
can regulate cellular responses and homeostasis according to different types of
stressors, suggesting a significant role of miRNA in thermal stress responses in
animals. Recently, it appears that during heat stress the role of several kinds of
miRNA can play regulatory roles by targeting genes that are related to the stress
responses in livestock, therefore controlling the synthesis of heat shock proteins
(HSPs), and the expression of genes responsible for the regulation of immune
responses, apoptosis, and inflammation-related molecules. The study of miRNA
profiling in sheep under heat stress in the next future could contribute to elucidate
the complex network of interactions at cellular level responsible for the fully
understanding of immune response to heat stress.
58 M. Caroprese et al.

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blood mononuclear cell proliferation and cytokine production in sheep as affected by cortisol
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temperature on phagocytosis, reactive oxygen species production and apoptosis in bovine
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Sevi A, Caroprese M (2012) Impact of heat stress on milk production, immunity and udder health in
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Climate Change and Livestock Production:
Significance of Studying Multiple Stressors 6
Impact in Cattle in the Changing Climate
Scenario

C. G. Shashank, M. V. Silpa, P. Ezhil Vadhana, Chinnasamy Devaraj,

M. Tiwari, and Veerasamy Sejian

Contents
6.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
6.2 Concept of Multiple Stresses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
6.3 Effect of Multiple Stressors on Cattle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
6.4 Physiological Changes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
6.5 Metabolic Changes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
6.6 Molecular Changes During Heat and Nutrition Stress . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
6.7 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
6.8 Future Perspective . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72

Abstract
Climate change directly impacts food production system stability, domestic or
through the global food market. This threat necessitates the identification of
various measures to sustain agricultural production to meet the growing
population’s demand. Among agricultural sectors, livestock has the potential to
fortify resilience to climate change, as it tends to be more resilient than crop-based
systems. Therefore, in light of the changing climate, it may benefit from investing

C. G. Shashank
ICAR-National Dairy Research Institute, Karnal, Haryana, India
M. V. Silpa
Institute of Animal Breeding and Genetics, Justus-Liebig-Universität Gießen, Gießen, Germany
P. Ezhil Vadhana · C. Devaraj · V. Sejian (*)
Centre for Climate Resilient Animal Adaptation Studies, ICAR-National Institute of Animal
Nutrition and Physiology, Bangalore, India
M. Tiwari
Animal Biotechnology Division, ICAR-National Dairy Research Institute, Karnal, India

# The Author(s), under exclusive license to Springer Nature Singapore Pte 59

Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_6
60 C. G. Shashank et al.

in the cattle sector to increase its production capacity for the future. Climate
change does not contribute only to the usual suspect, heat stress rather it
contributes to multiple environmental stresses. Unfortunately, predominant
research efforts are oriented towards tackling heat stress alone may not provide
rich dividends in the future as there are several other associated stresses emerge in
the ever changing climate. In tropical countries, especially where the predomi-
nantly extensive rearing system is being practised, it is vital to quantify multiple
stressors’ cumulative impact on animals rather than just concentrating on heat
stress. Milk and meat production will play a huge role in feeding the growing
human population of 9.6 billion by 2050. Therefore sustaining the milk and meat
production from the cattle forms a significant component for future policies to
ensure food security especially in the developing part of the world. This warrens
more research, particularly in quantifying the multiple environmental stressors on
dairy and beef cattle production, which could be the way forward approach to
revisit the amelioration strategies that could play a significant role in sustainable
livestock production in the changing climate.

Keywords
Climate change · Cattle · Food security · Livestock · Multiple stress

Abbreviations

% Percentage


C Degree Celsius
ACOT7 Acyl-CoA thioesterase 7
ATF4 Activating trasncription factor 4
CDC42 Cell division cycle 42
CFTR Cystic fibrosis transmembrane conductance regulator
CO2 Carbon dioxide
CPT1C Carnitine palmitoyltransferase1C
CRYAB Crystallin ALPHA B
DM Dry matter
DMI Dry matter intake
DYRK1A Dual specificity tyrosine phosphorylation regulated kinase 1A
Fig Figure
GH Growth hormone
GPX Glutathione peroxidases
GSTM Glutathione S-transferase mu
HPA Hypothalamic-pituitary-adrenal
HSE Heat shock elements
HSF Heat shock factors
HSP Heat shock proteins
ID2 Inhibitor of DNA binding 2
IGF-1 Insulin-like growth factor-1
6 Climate Change and Livestock Production: Significance of Studying Multiple. . . 61

IGFBP Insulin-like growth factor binding proteins

IPCC Intergovernmental Panel on Climate Change
ITPR2 Inositol 1,4,5-Trisphosphate receptor type 2
MANF Mesencephalic astrocyte derived neurotrophic factor
min Minute
MLPH Melanophilin
NDF Neutral detergent fibre
NEFA Non-esterified fatty acids
pH Potential of hydrogen
PI3-kinase Phosphoinositide 3-kinases
PIK3C2G Phosphatidylinositol-4-phosphate 3-kinase catalytic subunit type
2 gamma
PRKAB2 50 -AMP-activated protein kinase subunit β-2
RAC1 Ras-related C3 botulinum toxin substrate 1
RAP Receptor associated protein
SAM Sympathetic adrenal medullary
SLC23A1 Solute carrier 23 member 1
SOD Superoxide dismutase
STAT-5 Signal tranducer and activator of transcription 5
T3 Triiodothyronine
T4 Thyroxine
THI Temperature humidity index
TNZ Thermo neutral zone
US United States
ZNF516 Zinc finger protein 516

6.1 Introduction

The human population is expected to stretch from 7.2 to 9.6 billion by 2050;
accordingly, agricultural systems worldwide will have to make available extra
food to feed this increasing population. Livestock and crop systems are the central
pillars of an agricultural system. If we peek into livestock systems alone, 17% of
global kilocalorie consumption and 33% of global protein consumption is through
livestock products, and it is considered a vital agricultural commodity for global
food security. Growing demand for livestock products, particularly its rapid growth
in developing countries, have stated it as a “livestock revolution”. Despite this, the
current expansion is insufficient to meet the growing demand for livestock products,
especially in the wake of the diminishing water and land resources caused by the
entrenched rogue known as climate change.
Several stresses other than heat stress constrain livestock production in the current
climate change scenario. Animals, when exposed to single stress at a time, can
counter efficiently without any fluctuations in the productive capabilities due to
62 C. G. Shashank et al.

stored body reserves. However, if animals are exposed to multiple stressors, the
cumulative effects of various stressors might harm the exposed animals. In this case,
the animal’s body reserves are not plentiful enough to efficiently counter multiple
environmental stressors as the overall biological cost is inconsequential to the
animal’s welfare. Resulting in hampered adaptive capabilities, and the animals
struggle to maintain normal homeothermy. With the present climate changing
scenerio, the frequency and intensity of livestock exposure to abiotic and biotic
stressors increased. Abiotic stressors such as heat and nutritional stress significantly
impact livestock productivity (Sejian et al. 2015). According to the IPCC Fifth
Assessment Report, the anticipated intensification in global average surface temper-
ature by 2100 will be between 0.3 and 4.8
C (IPCC 2013). The United States
Livestock industry witnessed an annual economic loss of 1.63–2.36 billion US
dollars because of heat stress, of which the dairy industry bears 50%. Latent impacts
on livestock include fluctuations in production, quality and quantity of feed crop and
forage, water accessibility, animal growth, and milk production. Increased Elevated
temperature and dry conditions lead to discrepancies in water-soluble carbohydrates
and nitrogen levels, affecting the quality of forage and feed crops. On the other hand,
the increased temperature might increase lignin and cell wall components in plants,
which reduces digestibility and degradation rate, leading to a reduction in nutrient
availability for livestock. Hence, it is not only the heat stress that needs to be
counteracted, but nutrition stress is also of great concern.
Livestock has the potential to reinforce climate change resilience, as it is gener-
ally more resilient than crop-based systems. A better understanding of how the
industry is affected is required to augment livestock resilience to climate change.
Multiple stresses affecting livestock proficiency are common due to climate change
in the tropical environment. Most research focuses on a single stress response since
stress responses are multidimensional and comprehensive; balanced multifactorial
experiments are practically difficult to manage, evaluate, and interpret. Reports
assessing the effects of multiple stresses in cattle are limited. Dietary nutrients
wield their roles through numerous nutrient-metabolic and cell signalling pathways;
our current information is still not deep enough to untie the complexities of dietary
nutrients and animal genome expression. Research efforts are required to study
physiological and genotypic traits responsible for growth, improve production
ability, and identify different biological markers used in future breeding programs.

6.2 Concept of Multiple Stresses

Stress continuously moulds evolution on earth, and exposure to stress is the

organism’s daily fate. However, after years of research, understanding the concept
of stress remains a mystery to humankind. Currently, there is an increasing appreci-
ation within livestock stakeholders and industry that stress on animals is real, and
then it must be addressed. With significance for animal health and well-being,
reducing animal stress through better animal management measures and selective
breeding should be our topmost priority. Nevertheless, efficient stress management
6 Climate Change and Livestock Production: Significance of Studying Multiple. . . 63

Fig. 6.1 Pictorial representation of summation effect of multiple stresses on productive functions
in cattle. (Source: Modified from Sejian et al. (2012))

depends on identifying and computing the effects of several stressors and determin-
ing if the individual or combined stressors have different biological effects.
Animals have evolved mechanisms to manage short-term stressors. During the
short-term exposure, the biological cost is minimal because adequate reserves of
biological reserves exist to cope with the stressor and meet the impact of the stress
without any disturbances on biological functions. If the animal gets challenged by
multiple stressors, there will be insufficient biological reserves to satisfy the
biological cost of the stress response; to counteract this, resources will be channelled
from other biological functions. As shown in Fig. 6.1, when resources are side-
tracked from productive functions, it leads to impairment of biological functions. For
example, when multiple stresses deplete body reserves, metabolism shifts away from
growth, the young animal no longer blooms, and growth is restricted. When energy
is shifted from reproduction and its process, reproductive success is reduced. This
metabolic maintenance behaviour of an animal’s body rather than production will
last until the animal restocks its resources/reserves sufficiently to re-establish normal
functions.
Looking at the different climate change predictions, we can envision a future of
immense struggle to adjust and adapt to new environmental challenges both by
humans and livestock. Therefore, to secure food security, recognising animals with
superior genetic traits that are economically beneficial and identifying the
biomarkers to find a solution to animal productivity to climate change should be
of utmost preference to policymakers and researchers, especially when animals get
exposed to multiple stressors.

6.3 Effect of Multiple Stressors on Cattle

Livestock must maintain a steady body temperature to optimise production effi-

ciency and maintain physiological limits within a particular environmental tempera-
ture range, known as the “Thermo neutral zone” (TNZ). Heat stress in cattle is the
primary consequence of rising global temperature. Heat stress is manifested when
64 C. G. Shashank et al.

there is an increase in the core body temperature of animals and the inability to
dissipate the body heat temperature due to high ambient temperature combined with
high humidity and slow air movement. The effects on livestock will vary by region,
animal species, and production type, whether positive or negative. Changes in
rainfall patterns, upsurge in temperatures, more regularity of extreme weather
events, increased heatwave situations, and reduced water availability affects live-
stock productivity and hampers production systems globally, both directly and
indirectly.

6.4 Physiological Changes

Livestock reared under tropical climates are subjected to various challenges. Along
with heat stress, multiple stresses have a negative impact on livestock adaptation and
productivity. Whenever animals are exposed to stress primary sign of identification
by livestock holders is through observing changes in physiological parameters like
respiration rate, rectal temperature, feed intake, water intake and heart/pulse rate.
Multiple stresses negatively affect the dry matter intake (DMI), leading to the
decreased growth rate in cattle. For example, heifers reared in heat stress conditions
(38
C) displayed reduced feed intake; further average daily gain was reduced
compared to the calves in ambient temperature (17
C). Speaking on a percentage
basis, a reduction around 9% of dry matter intake and 22% average daily gain of
prepubertal Holstein heifers maintained at 33
C, while evaporative water loss, such
as sweating, was higher in those heifers, which in turn caused a 23% increase in
water intake compared to heifers maintained at 28
C. Changes in fat-storing
mechanisms (including reduced protein turnover and fat thickness) have also been
noted in prepubertal heifers reared under Heat Stress conditions (33
C). However,
reports suggest that if animals are maintained under proper nutrition balance, the
downfall of body weight due to heat stress can be avoided. Modifying the feeding
time and providing proper shading can be a good strategy to achieve the above
statement. If dissipation of metabolic heat production during digestion can be
achieved during cooler parts of the day by feeding the animal around
mid-afternoon, bodyweight in cattle can be maintained even during heat stress.
Heat stress associated with economic losses is majorly from three factors: declined
performance, increased mortality, and decreased reproduction. Reduced growth
performance and production efficiency could be partially due to sacrificing a fraction
of growth energy for cows heat regulation.
During thermal stress, numerous physiological modifications occur in dairy cows
to reduce the external heat load or metabolic heat load production. Neurons act as
temperature-sensitive sensors, which send information to the hypothalamus in
response to numerous physiological and behavioural responses like fluctuations in
blood flow, endocrine responses, acid-base chemistry, and physiological traits like
increased respiration rate, pulse rate and rectal temperature. Respiration rate is
directly proportional to Temperature Humidity Index in lactating dairy cows. If the
respiration rate is more than 60 breaths/min is considered an indicator of heat stress
6 Climate Change and Livestock Production: Significance of Studying Multiple. . . 65

in cattle. Various studies reported an increase in respiration and panting rate in cows
exposed to heat stress. Increased respiration rate certifies that heat load on animals
beyond the threshold is dissipated through boosted evaporative heat loss, leading to
changes in blood acid-base chemistry. Enhanced respiration rate and panting result
in more exhalation of CO2 via pulmonary ventilation. Lower blood carbonic acid
concentration and an imbalance in carbonic acid to bicarbonate ratio can lead to
respiratory alkalosis in lactating cows. Therefore, the need to counteract a higher
blood pH through increased urinary bicarbonate excretion and a loss of blood
carbonic acid become the utmost priority during heat stress. Another mechanism
through which dairy cattle counteract the heat load is through the thermoregulatory
centre in the hypothalamus, which triggers thermolysis through evaporation,
resulting in increased respiration rate in cattle.
Respiration rate and rectal temperature run parallelly in animals. Rectal tempera-
ture is an efficient indicator to assess stress in animals. Measuring rectal temperature
is generally considered one of the best methods to assess the core body temperature,
and any changes in rectal temperature reflect similar changes in core body tempera-
ture. The average rectal temperature of dairy cattle is around 38.5
C and a threshold
range of 39.4–39.7
C. When cattle get exposed to heat stress and even combined
stresses, the rectal temperature increases due to the reason, once the threshold
temperature crosses, dairy cows initiate to store heat; eventually, rectal temperature
escalates, heat loss through cutaneous evaporative upsurges and variation (in core
body temperature) between cows is much significant than cows exposed to below
35
C. This might be due to differences in sweat glands’ number (and activity) and
hair coat characteristics. Even heart rate increases gradually when cattle are
subjected to heat stress. Increased heart rate helps control blood pressure due to
elevated vasodilatation and increased blood flow due to heat stress. Peripheral
tissues get the blood flow redistributed, ensuring that body heat is transferred from
core body organs to the body’s surface. There is a possibility that heart rate changes
may serve as a cows method of temperature control to keep their heat balance.
Conversely, during nutrition stress, the thermoregulatory mechanism works dif-
ferently. Due to limited feed intake, a drop in body temperature, respiration rate and
pulse rate can be witnessed due to less metabolic rate, as Vant Hoff’s law describes.
Various reports suggest that during hot weather, cows fed with low fibre feeds (NDF,
30% of Dry matter) showed lower body temperature and respiratory rates compared
to animals fed with high fibre feeds (NDF, 42% of DM). This confirms that diet
intake has an applicable effect on metabolic heat production. Physiological traits like
respiration rate, rectal temperature and heart rate increase during initial exposure
then drops progressively with the days; this clearly explains the duration of heat
stress and acclimation capability of cattle affects the intensity of the response. Once
the animals get acclimatised to stimuli, it reduces the endogenous heat production
and helps heat loss. This adaptive mechanism can be observed in zebu cattle. Genetic
adaptation allows zebu cattle to have lesser rectal temperature than Bos taurus when
subjected to comparable heat stress situations. If animal exposure is prolonged (and
not lethal), by the definition of acclimation, changes in endocrine status lead to
gradual homeostasis. However, reaching that equilibrium varies according to breed,
66 C. G. Shashank et al.

age and sex; for example, it ranges from 9 to 14 days for Angus and Polled Hereford
cattle to get acclimated to heat stress (Bernabucci et al. 2010).

6.5 Metabolic Changes

Any changes in the environment are acclimatised by differences in homeostatic

responses, including altered endocrine status. In addition to temperature regulation,
hormones are also implicated in the acclimatory reaction to environmental stresses.
Thyroxine (T4) and Triiodothyronine (T3) hormones play a significant role in
acclimation. Besides producing hormones, the thyroid gland has a significant role
in maintaining the metabolic rate and homeostasis of protein and energy metabolism.
Any changes in the concentration of thyroid hormones mirror the animal body’s
metabolic and nutrient status, and it positively correlates with growth gain. Studies
suggest a decrease in the concentration of T4 and T3 in Holstein cows exposed to
heat stress. Likewise, a reduction in T3 concentration around 18.3, 16.0, 22.0 and
14.4% in cattle species like Alentejana, Frisian, Limousine and Mertolenga and T4
concentrations around 15.3, 15.0, 23.8 and 21.7% in the same cattle species men-
tioned above, when exposed to heat stress were reported. This shows an evident
decrease in T3 and T4 in the blood, which indicates that the pituitary thyroid axis is
depressed during heat stress. Another face of this decrease in thyroid hormones
might be due to increased glucocorticoid hormone concentration. Reports suggest an
interaction between the thyroid gland and adrenaline, noradrenaline released in
response to temperature changes in the environment (Johnson et al. 1988). As
mentioned previously, feed intake and thyroid hormones status go side by side.
When steers are subjected to a restricted diet, T4 concentration was reduced signifi-
cantly after 56 days, and T3 reduced around 83 days. The data infers that feed
restriction in steers regulates both thyroidal secretion and extrathyroidal T3 produc-
tion and even indicates T4 in circulation is more closely linked with energy con-
sumption than T3. T3 is the metabolic active thyroid hormone, whereas T4 is the
thyroid gland’s main product and the most abundant iodothyronine in blood circula-
tion. Hence, maintaining the concentrations of T3, especially during feed restriction,
could be metabolically necessary for growing steers.
During heat stress, the hypothalamic-pituitary-adrenal (HPA) and Sympathetic-
adrenal-medullary (SAM) axis trigger to preserve homeostasis (Sejian et al. 2018). It
results in plasma glucocorticoids’ production, primarily cortisol, the primary stress
hormone in ruminants, which provokes physiological modifications in animals to
tolerate stress. In response to thermoreceptors in the skin, adrenocorticotropin
release in the hypothalamus increases plasma glucocorticoids. Glucocorticoids
help in vasodilatation and stimulate proteolysis and lipolysis by distributing energy
to the animal, even during reduced intake. During acute stress, cortisol levels
increase since glucocorticoid hormones have hyperglycemic action, enhancing glu-
cose formation through the gluconeogenesis process in heat-stressed animals.
On the other hand, the story of chronic heat stress is the opposite of acute stress.
Cortisol is thermogenic in animals; the reduction in the activity of adreno-corticoids
6 Climate Change and Livestock Production: Significance of Studying Multiple. . . 67

during heat stress act as a thermoregulatory protective mechanism to prevent meta-

bolic heat production during the thermal environment. This depicts the relation of the
adrenal-thyroid axis with acute and chronic stress adaptation in animals. Interest-
ingly, when cattle are fed limited or exposed to the restricted feeding regime, the
cortisol concentrations were significantly higher in the fasted group of dairy cattle
than the regular fed group (Chouzouris et al. 2019). Feed restriction is considered
one of the potent stressors; animals adapt to stress by higher glucocorticoid produc-
tion. The following reason can justify the above statement. The concentration of
peripheral cortisol mainly depends less on the energy status than on the intensity of
stress stimulus on animals; in this case, hunger was the significant stress on animals.
Another study confirms that cows were fed only straw displayed higher metabolic
stress and higher cortisol concentrations than cows fed with a silage-based diet,
which suggests more profound physiological stress. Another angle of looking into
the above statement is, Phenylalanine is enzymatically converted to Tyrosine due to
glucagon, and Insulin inhibits it. Catabolism of Tyrosine is brought by cortisol and
other glucocorticoids hormones. During feed restriction, there might be a more
significant degradation of Phenylalanine and Tyrosine, which might be due to a
decrease in Insulin and increase in plasma cortisol concentrations, which deviates
their process of catabolism towards gluconeogenesis from these amino acids and
thus reducing protein synthesis capability.
Non-Esterified fatty acids (NEFA) and glycerol are the hydrolysis products,
unlike triacylglycerols, of stored triglycerides in animals’ adipose tissue by enzyme
hormone-sensitive lipase regulated by Insulin- and catecholamines. NEFA is a vital
marker for evaluating metabolic adaptation in different livestock species. During
heat stress, animal’s Dry Matter Intake reduces to maintain the production of
metabolic heat. As a result, the energy supply through feed reduces; the animal
comes under a negative energy balance. Characteristic response of Negative energy
balance is the reduction in plasma Insulin. Reduced insulin concentrations lead to
adipose lipolysis and mobilisation of NEFA. Many studies reported decreased levels
of NEFA when animals were exposed to heat stress, which might be due to augment
glucose as fuel to counteract the energy deficiency, a novel strategy to reduce
metabolic heat production. Decreased NEFA concentrations might be due to failure
in registering shifts in post-absorptive energetic metabolism (although insufficient
intake of nutrients) may confirm heat stress on animals directly (not mediated by
feed intake) affects energetics. Higher plasma NEFA is a typical glucose-sparing
mechanism that animals on a low nutrient intake implement to increase milk
synthesis. During heat stress, many metabolic and physiological adaptations occur
in animals to maintain homeorhetic hormones. The direct physiological effects of the
growth hormone result from the triggering Growth hormone receptor, resulting in
NEFA mobilisation and gluconeogenesis from adipose tissue by emphasising the
lipolytic stimulus to β-adrenergic signals impeding insulin-mediated lipogenesis and
glucose utilisation correspondingly. Contrastingly, in a study where 50% reduced
feed and fed with straw cow group had the most significant increase in NEFA
concentrations with >fourfold increases the first day of feed restriction and peaking
a sevenfold change on the second day. During feed restriction, insulin levels
68 C. G. Shashank et al.

decrease, final products of lipolysis, NEFA is used as a metabolic energy source to

spare amino acids and glucose.
Growth hormone (GH) or somatotropin, a peptide hormone that triggers cell
growth, reproduction and regeneration. GH is a single chain polypeptide containing
191-amino acids produced, stockpiled and secreted by specified somatotropic cells
in the anterior pituitary gland. Studies reported a decrease in GH production in
chronic heat stress Jersey cows’ group (35
C) compared to thermoneutral zone
Jersey cows (18
C). Few reports suggest when dairy cattle get exposed to THI more
than 70, GH production declined, decrease in GH production might be due to
countermeasure the metabolic heat in dairy cows. Chronic heat stress negatively
impacts GH, followed by a negative energy balance resulting in decreased milk
production and even characterised by a decrease in catecholamines and glucocorti-
coid levels. The change in endocrine status impacts the circulating levels of T4 and
T3, reducing basic metabolic rate and finally, reduction in heat production.
When the animals are subjected to feed restriction, plasma GH concentrations
will increase, linked with GH resistance. During feed restriction, as a consequence of
reduced plasma levels of hormones such as T3, T4 and production of Growth
hormone receptors and GH binding proteins in the plasma are decreased along
with other hormones. The decreased attachment of GH with the target tissue,
especially the liver will be reduced. Few studies reported an increase in serum
growth hormone concentration in Holstein heifers during the restriction phase
compared with the control group results. An increase in GH reflects the animals
attempt to summon energy reserves from adipose tissue and thereby maintain
homeostasis during negative energy balance. Growth hormone has a lipolytic effect
on adipose tissue, especially during the feed restriction due to high GH concentra-
tion, adipose tissue mobilisation and increasing blood Non-esterified fatty acid
concentration. Particularly during feed restriction, growth hormone secretion
changes and even GH inhibiting factors from the hypothalamus, finally resulting
in increased GH.
Insulin-like growth factor-1 (IGF-1), also named Somatomedin C, is a growth-
promoting factor in many body tissues. IGF-1 has a resemblance to insulin molecular
structure. The IGF circulate in the body monocovalently bound to greater molecular
weight Insulin-like growth factor binding proteins (IGFBP). The liver is the primary
source of IGF-1, however it is also produced in other tissues. IGF-1 mediates GH
effects. Generally, GH binds to its specific receptors as a part of the somatotropic
axis pathway on the hepatocyte surface. During summer, the concentration of IGF-1
was found to be decreased in few studies. Circulating IGF-1 in dairy cows helps
regulate milk synthesis, and blood-borne IGF-1 production depends on GH response
on the liver. Heat stress in cattle reduced the hepatic GH receptor abundance, and
along with reduced GH signalling through STAT-5, hepatic IGF-1 mRNA was
significantly lower in heat-stressed animals. The mechanism behind reducing GH
receptors in hepatic tissue during heat stress is still unclear but may help modify the
GH-dependent gluconeogenesis in dairy cows. Because of this, lower levels of
insulin-like growth factor 1 (IGF-1) could also explain the decrease in milk synthesis
6 Climate Change and Livestock Production: Significance of Studying Multiple. . . 69

and increase nutrients utilisation to maintain homeostasis in the liver and mammary
tissues.
Linear decrease in plasma IGF-1 concentrations decreases as a result of lower
GH-receptor availability during feed restriction. Several studies validate the above
statement in growing cattle, under chronic feed restriction phase, IGF-1 concentra-
tion reduced. The number of GH receptors binding sites in the liver which are
available is critically essential for the production of IGF-1. The reason being, during
feed restriction, the liver will have refractory effects to the effects of Growth
Hormone, resulting in lower IGF-1 concentration and conversely higher GH con-
centration production, due to negative feedback regulation of IGF-1 on hypotha-
lamic growth production.

6.6 Molecular Changes During Heat and Nutrition Stress

When the animal gets exposed to high ambient temperature, body temperature
surpasses the average specified for their thermoneutral zone. The total heat load
exceeds the animal’s heat dissipation capacity. An increase in the heat load increases
the health problems and mortality rates, especially for high productive animals and
even for intolerant or less resistant animals. Heat tolerance can be characterised as
animals’ ability to maintain their hereditary functional capacities in hot conditions. A
heat-tolerant animal’s physiological traits are wide skin area to weight ratio, protec-
tive eyes, pigmented skin, and light coloured or whitish body cover. Amelioration of
climate change stress on animals can be addressed by understanding the basic
cellular and molecular responses. Understanding the underlying process that may
provide ways to select stress-tolerant animals and maintain the same animals in these
changing environments is necessary.
Heat stress triggers cytotoxicity by altering the biological molecules, impairs cell
functions, modifying metabolic reactions, inducing cell damage through free
radicals, and triggering necrotic and apoptosis pathways. These abnormalities are
responsible for significant transcription and protein synthesis changes known as the
heat stress response. Heat shock factors (HSF) have been linked as an impactful first
line of defence during cell temperature initiation. These factors coordinate cellular
reactions against thermal stress and influence gene expression in many genes,
including Heat Shock Proteins (HSP). The HSP dissociates from HSF1 monomers
when stimulated by heat stress, which unfolds and binds to other HSF1 monomers to
trimers before their nuclear translocation. In the nucleus, homotrimeric Heat shock
factor1 binds to promoters containing heat shock elements (HSE) to trigger heat
stress specified gene transcription. While HSF1 has predominantly been linked to
HSP regulation, emerging reports now point to its involvement in regulating carbo-
hydrate metabolism, molecules’ transport, cytoskeleton, and ubiquitination, espe-
cially during heat stress. A study on lactating dairy cattle evaluated the hepatic tissue
gene expression profile in response to an extended period of heat stress (Rhoads et al.
2005). The liver’s vital role in whole-body metabolism (integration of both Exo and
endogenous nutrients) prone to be affected by reduced feed intake and metabolism
70 C. G. Shashank et al.

shifts due to heat stress and even suggested that cellular response to heat stress is a
multi-tier process starting with HSF1 activation, preceded by enhanced expression
of HSP, lower fatty acid metabolism, activation of stress response via endocrine
system, and immune system activation all occurring simultaneously.
Many genes contribute to the heat tolerance capability in livestock; Taye et al.
(2017) worked on different cattle breeds of Africa and identified reassuring
signatures of genes for their tolerance capability towards heat stress. Researchers
identified heat stress tolerance is directly associated with IGF-1 and HSF5 genes;
SOD1, GPX7, GSTM2, GSTM4, SLC23A1, and SLC23A1 were positively selected
as the genes related to the oxidative stress response. RAP17 and MLPH genes were
found linked with coat colour, ITPR2 and CFTR genes were responsible for sweat
glands development in native African cattle breeds which helped in more heat
dissipation. Holstein calves’ molecular response to thermal stress involves genes
that may potentially breed Holstein cattle with superior thermotolerance capability.
Increased HSP expression aimed at preventing protein aggregation and disintegra-
tion of misfolded proteins and many differentially expressed genes involved in fatty
acid metabolism like ATF4 (activating transcription factor 4), which ensures fatty
acid synthesis, CPT1C (carnitine palmitoyltransferase 1C) associated in fatty acid
β-oxidation, ACOT7 (acyl-CoA thioesterase 7) engaged in long-chain hydrolation of
fatty acids were up-regulated and CDC42 (cell division cycle 42), RAC1
(Ras-related C3 botulinum toxin substrate 1), and PRKAB2 (50 -AMP-activated
protein kinase subunit β-2) which maintains vital enzymes participating in de-novo
biogenesis of fatty acids were down-regulated.
Another study tried to understand the underlying biology regulating the effects of
dietary restriction and subsequent re-alimentation in the ruminal epithelium papillae
of Holstein Friesian bulls. Researchers found out genes like CRYAB protein which
exhibits chaperon activities and performs to limit protein aggregation in a wide range
of conditions. HSPH1 and HSPB8 genes were down-regulated in skeletal muscle
tissue during dietary restriction. Genes involved in cell proliferation transcription
factor ZNF516 was down-regulated. Genes related to signalling processes involved
in growth and cellular proliferation, such as tyrosine phosphorylation regulated
kinases DYRK1A, DYRK1B, PIK3C2G, which is of vital importance in
PI3-kinase signalling, expressions were reduced. Additionally, up-regulation of
genes like MANF and ID2, associated with cellular proliferation inhibition was
identified at the end of the restricted diet period in hepatic tissue.

6.7 Conclusion

Besides heat stress, several stressors are hindering livestock production in the current
climate change situation. Animals can counterbalance efficiently without
fluctuations in production capacity due to the stored body reserves when exposed
to a single stress. However, the cumulative effects of the various stressors may
damage if animals are exposed to multiple stressors. Reports evaluating the effects of
multiple stresses in cattle are limited. More research efforts are essential to
6 Climate Change and Livestock Production: Significance of Studying Multiple. . . 71

understand the mechanism of how multiple stressors affect the adaptive and produc-
tive capability of cattle, such that policymakers can amplify the endeavours of
researchers to extra more data on the impact of multiple stressors on cattle. On the
other hand, cattle can efficiently counter heat stress, provided nutrition is sufficient to
maintain the normal homeostasis mechanism. Nutrition, particularly during heat
stress, plays a major role in the adaptive capabilities of cattle. Dietary nutrients
wield their roles through numerous nutrient-metabolic and cell signalling pathways;
our current information is still not deep enough to untie the complexities of dietary
nutrients and animal genome expression.

Learning Outcomes
• In the extensive system of rearing, climate change not only causes the usual
suspect heat stress but also leads to nutrition, water and walking stress to
grazing animals.
• The multiple stressors induces very high magnitude of adverse impacts on
the production and adaptation capacity in cattle than heat stress alone.
• Quantification of multiple stressors impact could be the most appropriate
way to support future policies for sustainable livestock production in the
changing climate scenario.

6.8 Future Perspective

Multiple stresses concept is gaining significance among the researchers in the

changing climatic scenario. Presently, various researchers are concentrating primar-
ily on data generation on heat stress and its impact on animals’ productivity and
adaptation. However, other stresses like nutrition, walking, water, and many more
are to be acknowledged and addressed. Through collaborative research (animal,
plant and allied sciences), we can understand the mechanism through which animals
and plants adjust to the multiple stresses. Parallelly, studies on ameliorative
strategies to counteract the multiple stresses, which are economical and practical at
the field level, should be devised and propagated through various extension services.
Research on multiple stresses should be initiated in tropical regions as early as
possible, given that the agricultural land and water resources shrinkage are catching
up speed due to climate change, resulting in increased nomadic behaviour of animals
in search of pasture and water. Considering the emergency to address the multiple
stresses in livestock, especially in the extensive rearing system, might threaten
global food security, policymakers should make rational managerial decisions and
fund more studies on multiple stresses than on single stress.
72 C. G. Shashank et al.

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Comparative Assessment
of Thermo-Tolerance of Crossbred 7
and Indigenous Cattle Breeds

Gyanendra Singh, Hari Abdul Samad, K. Karthiga, K. Priyanka,

Lipika Sarma, Vikrant S. Chouhan, and V. P. Maurya

Contents
7.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
7.2 Assessment of Thermo-Tolerance Between Crossbred and Indigenous Cattle . . . . . . . . . . . 76
7.2.1 Physiological Responses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
7.2.2 Hormonal Responses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
7.2.3 Biochemical Responses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
7.2.4 Heat Shock Responses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
7.2.5 Immune Responses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
7.3 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
7.4 Future Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80

Abstract
One of the important strategies to find solution to the climate change associated
livestock production is to identify the ideal livestock breed to survive in a specific
location. This warrants research efforts to screen the available breeds to identify
climate resilient breeds. Further, comparing the crossbred animals with indige-
nous breeds can help to understand the real differences in livestock adaption in
addition to identifying quantifiable biomarkers covering productive and adaptive
traits. This chapter is an attempt to project to the readers the differences in
adaptation between crossbreds and indigenous cattle. Indigenous cattle possess
various unique characteristics which imparts them the potential to survive in
tropical climate. However, the crossbred cattle which are well known for their
production struggles in harsh climatic condition which ultimately culminates in

G. Singh (*) · H. A. Samad · K. Karthiga · K. Priyanka · L. Sarma · V. S. Chouhan · V. P. Maurya

Division of Physiology and Climatology, ICAR-Indian Veterinary Research Institute, Izatnagar,
Uttar Pradesh, India

# The Author(s), under exclusive license to Springer Nature Singapore Pte 73

Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_7
74 G. Singh et al.

reduced production and welfare. Crossbred cattle shows higher physiological and
biochemical reactions, as well as a lower total antioxidant capacity, as compared
to native breeds, indicating that they were more susceptible to heat stress. Further,
heat shock protein 70 (HSP70), toll-like receptor 2 (TLR2) and TLR4 which is
considered as potential markers for temperature adaptation in livestock was
highly expressed in less adapted crossbred than indigenous breed. More such
efforts are needed to identify more thermo-tolerant breeds to propagate them in
specific agro-ecological zones. Such efforts can ensure the livelihood of poor and
marginal farmers in the changing climate scenario.

Keywords
Adaptation · Breeding · Climate change · Heat stress · HSP70 · Thermo-tolerance

Abbreviations

% Percentage

C Degree Celsius
ALP Alkaline phosphatase
ALT Alanine aminotransferase
APP Acute phase proteins
AST Aspartate aminotransferase
DAHD Department of Animal Husbandry, Dairying
DC Dendritic cells
GDP Gross domestic product
HF Holstein Friesian
HPA Hypothalamic pituitary adrenal axis
HSPs Heat shock proteins
IL Interleukin
MAS Marker assisted selection
ml Millilitre
mRNA messenger ribonucleic acid
ng Nanogram
PBMC Peripheral blood mononuclear cells
RR Respiratory rate
RT Rectal temperature
T3 Triiodothyronine
TLR Toll-like receptors
TNF Tumor necrosis factor
7 Comparative Assessment of Thermo-Tolerance of Crossbred and Indigenous. . . 75

7.1 Introduction

India, a mega diverse country, accounts 8% of all recorded species within 2.4% of
the world’s land area. The overall climate ranges from humid and dry tropical in the
south to temperate alpine in the north with a great diversity of ecosystems. Agricul-
ture, the most climate-dependent economic activity is considered to be the chief
source of livelihoods for about 70% of Indian population. In the past four decades,
the livestock sector’s rapid growth has made a significant contribution to agricultural
growth both nationally and globally. Livestock sector’s annual contribution in total
GDP is nearly 4.9% (DAHD 2016). According to 20th Livestock Census (2019), in
India, the total livestock population is 536.76 million showing an increase of 4.8%,
in which total number of cattle accounts for 193.46 million marking an increase of
1.34% as compared to the last census. In comparison to the previous census total
exotic/crossbred cattle population has been increased by about 29.3% whereas the
indigenous (both descript and non-descript) cattle population declined by about 6%
from the previous census. More over India continues to be the largest milk-
producing nation globally since 1998 (estimated milk production of 187.7 million
tonnes per annum), with an increase of about 6.45% annually. By 2029, world milk
production will increase around 997 million tonnes at the growth rate of 1.6%
p.a. and most importantly, India is likely to contribute more than 30% of world
production. The livestock products demand is expected to double in next three
decades globally. This increase in demand is due to the improved living standard,
growing population, ageing, socio-cultural patterns and also the change in food
preferences across the world from staple food to calorie-dense and taste-based
food items. In order to overcome this growing demand driven by dietary changes
and also the increasing population, 60% of food production has to be increased
by 2050.
Crossbreeding spread like wildfire across the country, with new crossbred cattle
being preferred over native cattle due to their higher milk yield. Most of the genetic
resources used in crossbreeding came from temperate countries, where the climate
and weather, differ greatly from tropical countries like India. Indigenous cattle are
more thermo-tolerant than crossbreds/exotic cattle as they possess various unique
characteristics, which make them well adapted to the tropical climate. Furthermore,
the impact of climate change is predicted to be more severe in tropical countries
where the dairy cattle are primarily reared in an extensive system, unlike that of
chicken and pig which follow a semi-intensive system of rearing. As a result of
crossbreeding, not only has productivity grown, but it has also brought with it issues
of overuse and concerns about crossbred cattle’s long-term viability. In thermal
stress, animals dissipate excess heat which is exhibited by various physiological and
metabolic adaptive mechanisms which are energy consuming, and this is believed to
cause the proportional decline in milk yield in these animals.
Furthermore, it is also reported that heat stress can alter the milk composition by
reducing total protein content and total fat content in milk. In India, crossbreds are
considered to be most susceptible to heat stress in comparison with indigenous cattle
as a result of the long-term intensive breeding in order to improve their milk
76 G. Singh et al.

production. India attained top position as highest milk producing country globally
because of crossbreeding program of dairy cattle. Despite achieving this position,
India still faces many problems in rearing crossbreds compared to those indigenous
breeds of cattle. In the case of crossbreds, primary investment and maintenance
expenditure are high which is not feasible for all farmers who struggle for their
livelihood and also they are highly susceptible to contagious diseases.
With this background, this chapter is primarily targeted to comprehend the
information pertaining to comparative assessment of adaptive capability between
crossbreds and indigenous breeds. This will give us a clue of what are those
biological markers that could be used for future breeding policies to produce well
adapted animals along with production traits. It also tries to elucidate the basic and
molecular mechanism of adaptation, therefore to access the thermo-adaptability of
indigenous and crossbred cattle.

7.2 Assessment of Thermo-Tolerance Between Crossbred

and Indigenous Cattle

7.2.1 Physiological Responses

Generally, a thermal steady state is achieved in animal body through equilibrium

maintained between heat produced in the body by metabolic activities and heat
gained from the environment. Under heat stress, animal undergoes a series of
physiological changes to maintain homeostasis and physiological equilibrium.
These changes are highly dependent on external temperature and also genetic
make-up of an animal. Most of these physiological adjustments are exhibited in
order to dissipate heat from the body to its environment and also to reduce the heat
production through metabolic activities within body.
In a comparative study by Sailo et al. (2017), the respiratory rate (RR) in Sahiwal
were found to be 15.738  0.795, 18.158  0.795 and 29.818  0.795 and in Karan
Fries cattle were 15.779  1.136, 22.979  1.136 and 47.299  1.136 during winter,
spring and summer seasons. In same study, the rectal temperature (RT) also showed
similar results where in Karan Fries RT ( C) showed 37.492  0.115,
38.398  0.115 and 39.186  0.115 while Sahiwal showed 37.300  0.095,
38.178  0.095 and 38.810  0.095 during winter, spring and summer respectively.
When high yielding Holstein Friesian (HF) cows were subjected to high temperature
conditions, their rectal temperature increased significantly from 37.3 to 39.3  C
(Koubkova et al. 2002). According to Khan et al. (2018) the intensity of changes in
physiological parameters were found to be in the order of HF > Crossbreds>
Sahiwal which shows adaptive capability of the breeds. Results from the above
studies reveals that indigenous cattle breed respond better and more tolerant when
compared to that of crossbreds.
7 Comparative Assessment of Thermo-Tolerance of Crossbred and Indigenous. . . 77

7.2.2 Hormonal Responses

Both long and short term environmental stress on animals affects the endocrine
glands through the activation of hypothalamic pituitary adrenal axis (HPA) which is
a key component for heat stress. It is the hallmark response since it reflects the
physiological status of animals responding to heat stress. In the process of acclima-
tion, the secretory patterns of glands changes due to the activation of HPA and there
is consequent change in the plasma level of different hormones along with their
receptors in their target tissue. Endocrine system works to reorient the internal milieu
that is being disturbed during heat load. Hormones responsiveness to heat stress
stimuli primarily includes glucocorticoids, thyroid hormones, growth hormones,
mineralocorticoids, prolactin, catecholamines and anti-diuretic hormones.
Khan et al. (2018) observed the results indicated that level of cortisol was more
prominent in HF and crossbred than Bos indicus. These results were more consistent
with the results of Mc-Manus et al. (2009) showing increased level of cortisol in case
of heat stressed cows that is more severe in Bos Taurus when compared to that of Bos
indicus. Tejaswi et al. (2020), observed that the serum cortisol level and T3 were
significantly higher in crossbred (Hariana  Holstien Friesian/Brown swiss/Jersey)
when compared to Tharparkar and Sahiwal during both the winter and summer
seasons. According to Yousef et al. (1997), the concentration of cortisol increased to
29 ng/ml from 11 ng/ml in Friesian calves and T3 showed a decline in T3 levels from
151 to 126 ng/ml under heat load.

7.2.3 Biochemical Responses

Bhan et al. (2012) reported that the ALT and AST values of growing and adult
Sahiwal was significantly increased during afternoon compared to that of morning.
Georgie et al. (1973) and Shaffer et al. (1981) also reported the significant increase in
AST during exposure to different seasonal changes in crossbred cattle. Studies are
stating that there is seasonal variation that usually influence biochemical parameters
in HF heifers and also reported in the study that there is increase in AST levels in
serum during high ambient temperature. Lower levels of serum AST, ALT and ALP
activities in the native cattle compared to the crossbred cattle both during summer
and winter could be due to their resistance to heat stress which can be attributed to
morphological features like higher density, large sized sweat glands and its proxim-
ity to the skin’s surface, with more layers of cells in the epithelial layer.

7.2.4 Heat Shock Responses

Heat shock proteins (HSPs) are the major cellular proteins which spearhead the heat
shock responses for stress adaptation in animals. In normal cellular physiology, there
are many HSPs that play a very important role. This is highly conserved protein
which is meant to be activated by several environmental stressors. There is an
78 G. Singh et al.

induction of HSPs within minutes after animal exposure, which can go to its peak
expression which might be several hours later. In the stressed cell, there is an
interaction between denatured protein and HSPs which tends to inhibit the aggrega-
tion of cytotoxic proteins and hence maintains the homeostasis of protein in the cell.
The improvement in thermo tolerance is seen in case of prolongation or elevation of
levels of HSPs. Based on the biological functions and molecular weight, HSPs are
divided into HSP10, HSP40, HSP60, HSP70, HSP90, HSP100, HSP 110, and there
are other small families of HSPs. Among this HSP 70 is considered as the most
sensitive heat stress indicator which acts as a cellular thermometer in livestiock
(Dangi et al. 2014; Samad et al. 2019). Different investigators presumes that thermo-
tolerance capacity is associated by assessing intracellular expression of HSP90 in
livestock species (Dangi et al. 2014; Deb et al. 2015; Samad et al. 2019; Slimen et al.
2016). In case of farm animals, during heat stress there is an increase in HSP90 and
HSP70 which is observed in cattle, buffalo, broilers, sheep and goats (Yu et al. 2008;
Slimen et al. 2016; Shilja et al. 2016; Samad et al. 2019; Tejaswi et al. 2020).
Bharati et al. (2017) reported that the HSP90 mRNA is up regulated when
the Tharparkar cattle was exposed to 42  C. Deb et al. (2015) correlates the
thermotolerance with the increase of HSP90 mRNA levels in both Sahiwal and
Frieswal cattle through in-vitro heat stress studies. Kim et al. (2020) also studied the
HSP90 mRNA expression in PBMCs and reported that there is an increased level of
those proteins in response to elevated temperature in beef cattle. Heat stress
responses in different breeds of cattle and buffaloes was studied by Kishore et al.
(2013) and documented a rise in HSP90 2 h post heat stress which was highest in
Murrah buffaloes, followed by Sahiwal and HF. The study also added the ranking
expression between HSPs suggesting that HSP70 was more upregulated and HSP90
was least expressed in thermotolerant animals.

7.2.5 Immune Responses

Immune responses are bodies defensive responses towards infectious and danger
signals associated with cellular damage. Chief mediators of these responses include
body’s physical barrier, chemical barrier, various immune cell and soluble proteins
like cytokines. Heat stress associated cellular damage releases HSPs which have the
tendency to activate various immune cells mainly Dendritic cells (DC) and
Macrophages thereby inducing a cascade of inflammatory responses (Lecchi et al.
2016; Hop et al. 2018). Cytokines are the key secretions of these immune cells that
activate the immune cells as well as transmit downstream signals for ultimate
response. Based on the major response Cytokines are grouped into
pro-inflammatory (IL1 β, TNF α) anti-inflammatory (IL10), which are there to
counter act each other in order to maintain internal cellular homeostasis
(Cannon 2000).
Bharati et al. (2016) reported that there is a significant rise in levels of IL-2 and
IL-6 at various increased temperature of 37  C, 39  C and 42  C in Tharparkar cattle.
The expression of pro-inflammatory cytokines and anti-inflammatory molecules was
7 Comparative Assessment of Thermo-Tolerance of Crossbred and Indigenous. . . 79

higher in crossbred compared to native breeds. The low negative-Acute phase

Proteins (APP) concentrations like lower values of albumin due to elevated levels
of cytokines may cause an increase in disease incidences in crossbreds.

Learning Outcomes
• The indigenous livestock breeds possess unique adaptive characteristics
which imparts them the potential to survive and produce optimally in harsh
climatic condition.
• HSP70 is the most studied molecular chaperon and considered to be the
ideal cellular and molecular marker for heat stress in farm animals.
• As the crossbred animals are predominantly targeted for production traits,
they cannot withstand harsh climate resulting in loosing even the produc-
tive potential when exposed to heat stress for a pronged duration.

7.3 Conclusions

Crossbred cattle shows higher physiological and biochemical reactions, as well as a

lower total antioxidant capacity, as compared to native breeds, indicating that they
were more susceptible to heat stress. Native cattle had much lower HSP70, TLR2,
TLR4, and cytokine expression than crossbred cattle, indicating that they are more
resistant to temperature stress at the cellular level. Indigenous cattle exhibited more
tolerance to heat stress than crossbred during summer, and hence indigenous breeds
are better adaptable to tropical climate.

7.4 Future Perspectives

In the current climate changing scenario, one has to primarily look upon to develop
breeds that are more thermo-tolerant along with production traits. Very limited
studies are conducted in comparing the thermo-tolerance of crossbreds and indige-
nous cattle. The advanced biotechnological tools should be employed in identifying
more traits like adaptation and production. The results mentioned above in compar-
ing crossbreds and indigenous breeds needed to be extrapolated in identifying agro-
ecological zone specific breed. The baseline information is available for only few
indigenous breeds and in case of studying all other breeds will give us clues in
obtaining breed that will have the ability to survive in multiple locations with
optimal milk production. After obtaining the baseline information, different
approach of testing their adaptive capability by shifting them to different agro-
ecological zone and find which breed can survive better. This way those breeds
can be used for marker assisted selection (MAS) breeding with the amalgamation of
productive and adaptive traits. This can only be the solution of producing breeds that
can survive in any environment along production traits. Efforts are much needed not
80 G. Singh et al.

only in ensuring livelihood security of poor and marginal farmers but also makes the
livestock farming more profitable, As world is going to face huge food demand of
growing population by 2050, these efforts of bringing up thermo-tolerant breeds will
be beneficial in ensuring food security as well.

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dynamics of HSP70 during chronic heat stress in Tharparkar cattle. Int J Biometeorol 61(6):
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profile of HSPs in Barbari goats. Int J Biometeorol 58(10):2085–2093
Deb R, Sajjanar B, Pavani KC (2015) Bovine heat shock protein 70 and its application in cellular
thermo tolerance. J Vet Sci Technol 6(6):1000e121
Georgie GC, Chand D, Rardan MN (1973) Seasonal change in plasma cholesterol and serum
alkaline phosphatase and transaminase activity in crossbred cattle. Indian J Exp Biol 11(5):448
Hop HT, Arayan LT, Reyes AWB, Huy TXN, Min WG, Lee HJ, Rhee MH, Chang HH, Kim S
(2018) Heat-stress-modulated induction of NF-κB leads to brucellacidal pro-inflammatory
defense against Brucella abortus infection in murine macrophages and in a mouse model.
BMC Microbiol 18(1):1–12
Khan I, Quershi M, Akhtar S (2018) Crossbred cows respond differently from holstein frisian and
Bos indicus to heat stress under various climatic conditions. J Agric 34:301–310
Kim WS, Ghassemi Nejad J, Roh SG, Lee HG (2020) Heat-shock proteins gene expression in
peripheral blood mononuclear cells as an indicator of heat stress in beef calves. Animals 10(5):
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Kishore A, Sodhi M, Kumari P, Mohanty AK, Sadana DK, Kapila N, Mukesh M (2013) Peripheral
blood mononuclear cells: a potential cellular system to understand differential heat shock
response across native cattle (Bos indicus), exotic cattle (Bos taurus), and riverine buffaloes
(Bubalus bubalis) of India. Cell Stress Chaperones 19(5):613–621
Koubkova M, Knizkova L, Kunc P, Hartlova H, Flusser J, Dolezal O (2002) Influence of high
environmental temperatures and evaporative cooling on some physiological, hematological and
biochemical parameters in high-yielding dairy cows. Czech J Anim Sci 47(8):309–318
Lecchi C, Rota N, Vitali A, Ceciliani F, Lacetera N (2016) In vitro assessment of the effects of
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Sailo L, Gupta ID, Das R, Chaudhari MV (2017) Physiological response to thermal stress in
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Samad HA, Konyak YY, Latheef SK, Kumar A, Khan IA, Verma V, Chouhan VS, Verma MR,
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nutritional) stressors. Int J Biometeorol 60(9):1311–1323
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stress induced alterations in polymorphonuclear (PMN) cell activity in native and crossbred
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 Part III
Methods to Quantify Heat Stress Response
in Livestock
Non-Invasive Methods to Quantify the Heat
Stress Response in Dairy Cattle 8
Gundula Hoffmann, Mullakkalparambil Velayudhan Silpa,
Roman Mylostyvyi, and Veerasamy Sejian

Contents
8.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
8.2 Significance of Non-Invasive Methodologies to Quantify the Heat Stress Response . . . 87
8.3 Measuring Activity and Lying Behavior to Quantify Heat Stress . . . . . . . . . . . . . . . . . . . . . . . 88
8.4 Respiration Rate as a Significant Physiological Variable . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
8.5 Significance of Hair, Urine and Faecal Cortisol Estimation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
8.6 Opportunities to Measure Body Temperatures for Determining Heat Stress in Farm
Animals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
8.7 Infrared Thermal Image Applications in Assessing Thermo-Tolerance in Farm
Animals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92
8.8 Models to Quantify Heat Stress in Farm Animals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
8.9 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
8.10 Future Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97

G. Hoffmann (*)
Department of Engineering for Livestock Management, Leibniz Institute for Agricultural
Engineering and Bioeconomy (ATB), Potsdam, Germany
e-mail: [email protected]
M. V. Silpa
Institute of Animal Breeding and Genetics, Justus Liebig University Giessen, Giessen, Germany
R. Mylostyvyi
Department of Animal Products Processing Technology, Dnipro State Agrarian and Economic
University, Dnipro, Ukraine
V. Sejian
ICAR-National Institute of Animal Nutrition and Physiology, Bangalore, India

# The Author(s), under exclusive license to Springer Nature Singapore Pte 85

Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_8
86 G. Hoffmann et al.

Abstract

The main focus of our research lies on livestock-environment interactions and

how heat load in dairy cows can be measured. Several studies have used animal-
related parameters to measure heat stress in individual cows. In this report, an
overview of different heat stress indicators is given, focusing on non-invasive
indicators that analyze different physiological, ethological and endocrine
responses of livestock animals. However, potentials of digitalization, e.g. by
means of sensor technology, for recording animal-specific adaption reactions of
dairy cows are also described. Sensor information of animals such as milk yield,
vitality (respiration rate, body and skin temperature) and behavior (standing,
lying time) are analyzed taking into account the barn climate. The investigations
contribute to a better understanding of heat stress in farm animals and to the
improvement of animal welfare.

Keywords
Dairy cow · Heat stress · Indicator · Sensors · THI

Abbreviations

% Percentage


C Degree celsius
AT Air temperature
ATB Leibniz Institute for Agricultural Engineering and Bioeconomy
BT Body temperature
DIM Days in milk
ETIC Cattle equivalent temperature index
Fig Figure
GMT Greenwich Mean Time
H Humidity
HR Heart rate
HS Heat stress
IRT Infrared thermography
LR Linear regression
min Minutes
NRC National Research Council
NVB Naturally ventilated buildings
pH Potential of hydrogen
R2 Coefficient of determination
RH Relative humidity
RR Respiration rate
SR Solar radiation
T Temperature
THI Temperature humidity index
8 Non-Invasive Methods to Quantify the Heat Stress Response in Dairy Cattle 87

UK United Kingdom
USA United States of America
USB Universal serial bus
WS Wind speed

8.1 Introduction

Global climate changes are becoming more and more tangible for the world’s
agriculture and the increasing heat load plays a significant role in determining animal
welfare in livestock husbandry. Dairy cows in naturally ventilated barns are particu-
larly affected by the climate change and will suffer under heat stress. As cows are
relatively cold-tolerant but heat-sensitive, the extreme heat periods in the summer
months increasingly lead to restrictions in animal welfare, health and production
performance. Heat-stressed dairy cows dissipate surplus body heat into the environ-
ment and reduce the heat generation rate by adjusting their behavior (body posture,
lying bout, movement, etc.), showing physiological responses (e.g. increased body
temperature and respiration rate, decreased dry matter intake and milk production,
loss of weight, sweating, etc.) and immune responses (changed blood hormone
concentration). In order to keep body functions in a steady state, it is necessary for
homeotherms to maintain body temperature within a narrow range. A deviation from
the set point of the body temperature in a stressful, hot environment leads to an
impairment of the physiological processes and thus has a negative impact on the
productivity of the animals (Naqvi et al. 2012). Literature studies have made it
possible to determine physiological-ethological reactions of dairy cows that correlate
with climatic conditions (Hoffmann et al. 2020). The particular difficulty is to
identify the corresponding meaningful reactions. The objective of this work was to
bring together intensive monitoring of the microclimate and various physiological,
ethological and endocrine reactions of lactating dairy cows. Therefore, several
animal-individual characteristics as well as the microclimatic conditions inside the
barns were taken into account.

8.2 Significance of Non-Invasive Methodologies to Quantify

the Heat Stress Response

The field of science and technology has observed a huge progress over the decades.
These advancements have aided researchers across the globe in several ways like
ensuring quicker and accurate results. However, along with this, there is an increas-
ing rise in animal welfare and ethics especially for the researches involving the use of
animals, be it laboratory animals or even farm animals. All the heat stress
(HS) studies in livestock face a primary obstacle on obtaining ethical clearance as
these experiments subject the animals to severe distress. Having said that, it is
88 G. Hoffmann et al.

inevitable to conduct such live animal studies as they yield the most vital solution to
mitigate the rising concern on livestock associated climate change impact. At this
point, the use of non-invasive technologies to quantify HS is crucial. As these
techniques do not add on to any further stress in animals, adoption of non-invasive
techniques over invasive approaches are more preferred from an ethical point
of view.
Non-invasive methodologies also have an added advantage of being more accu-
rate in estimate the HS impact in animals. Most of invasive technologies require
intense animal handling. These approaches thereby add on to the stress in animals,
which is also an issue from the animal welfare perspective. Since non-invasive
methods require minimal animal handling, they are proposed to depict relatively
accurate stress profile in animals. Furthermore, apart from being accurate, some of
these non-invasive methodologies also have an advantage of being inexpensive and
of high correlation to assess HS both in producing and non-producing animals.
Time invariably, a number of non-invasive methodologies to assess HS impact in
livestock have been adopted. However, with the growing concern on animal welfare
and ethics, several improvisations have also been brought about in these
methodologies. The non-invasive methods also comprise a number of sensor-
based approaches that record series of vital variables in animals. Adoption of such
technologies enables early detection of heat load in animals and thereby quicker
ameliorative measures can be enforced to reduce its impact on livestock production.
Additionally, some of these technologies also aid in predicting the occurrence of
stress, be it HS or any other environmental stress, in livestock. Therefore, this gives
room for the livestock keepers and other stakeholders in this sector to enforce
suitable mitigation measures to combat the predicted adversities. Therefore, the
scope of using non-invasive technologies for HS assessment in livestock is massive
and thus proper emphasis must be given to propagate its usage among both the
livestock farmers and researchers across the globe.

8.3 Measuring Activity and Lying Behavior to Quantify Heat

Stress

As a sensitive indicator of HS, dairy cow activities including the resting behavior
(total lying time, number of lying bouts, lying bout duration) and number of steps per
cow per day were recorded by activity sensors (IceTag3DTM, IceRobotics,
Edinburgh, UK). This trial was carried out in the dairy barn of the Agricultural
Research and Education Center for Animal Breeding and Husbandry (LVAT, Gross
Kreutz, Germany). The effects of HS and individual cow factors (e.g. lactation state,
days in milk) on the total daily lying time are shown in Fig. 8.1. It was found that
cows with heat load had shorter lying times than cows without heat load. When
additional heat load was accumulated three days before the measurement day,
reduced activity responses to heat load (i.e. delayed heat load effects) were observed
(Heinicke et al. 2019).
8 Non-Invasive Methods to Quantify the Heat Stress Response in Dairy Cattle 89

800

Total lying time (min d-1)

699 666
647
568 582 561
600 531
479 472
400

200

0
reference lactation≥4 DIM>150
no heat load contemporaneous heat load delayed heat load

Fig. 8.1 Daily total lying time predicted under different heat load conditions for the reference
group, lactation 4 group, and DIM (days in milk) >150 group. Reference group has normal milk
yield level, lactation state of 1, no pregnancy and estrus (Heinicke et al. 2019)

8.4 Respiration Rate as a Significant Physiological Variable

Another reliable and early indicator of HS in dairy cows is the respiration rate (RR).
This was measured by counting the right thoracoabdominal movements for 30 s and
multiplying by 2. Depending on the measurement series, these measurements were
taken hourly from 0700 h to 1500 h (GMT + 0100 h) or twice daily (morning and
afternoon) in the LVAT (Gross Kreutz, Germany). At the same time it was
documented whether the animal was in a lying or standing position during the
measurement. In addition, the temperature and relative humidity were recorded at
fixed time intervals (10 min) and locations directly in the barn (EasyLog USB 2+
sensor from Lascar Electronics Inc., USA). The THI formula according to NRC
(1971) was applied:

THI ¼ ð1:8  T þ 32Þ  ðð0:55  0:0055  HÞ  ð1:8  T  26ÞÞ

Where, T is the air temperature in
C and H is the relative humidity in %. With the
help of the measuring intervals per 10 min, the heat load duration per heat load
intensity could also be determined. In addition to the average THI per day, the heat
load duration per heat load intensity provide more precise information on how long,
how strongly and when the animals were exposed to heat. It could be shown by Pinto
et al. (2019) that the RR showed high variabilities for the same THI, especially for
higher THI conditions. This illustrates the animal individual influences. The RR
increased with the increase of THI for both cow body postures. In addition, it was
observed that standing cows had a lower RR than lying cows when THI was less
than 80. Since counting flank movements is time-consuming, labor-intensive, and
non-continuous, ATB developed a new differential pressure sensor for monitoring
the RR (Strutzke et al. 2019). The sensor can carry out automatic, continuous and
long-term RR measurements.
90 G. Hoffmann et al.

8.5 Significance of Hair, Urine and Faecal Cortisol Estimation

Heat stress stimulates a number of endocrine responses in animals. Cortisol is one

among the classical HS markers in livestock. Traditionally this glucocorticoid
hormone levels are estimated in blood samples. Though the blood samples serve
as reliable medium for estimating the endocrinal changes occurring during HS, the
methodology is invasive and may add up to the existing stress especially when the
animals are not handled properly. Hence, the use of hair, feces and urine samples, as
an alternative medium is encouraged. Researchers have reported that these biological
samples provide an efficient estimate of cortisol levels in livestock during HS. In
most cases, the cortisol metabolites obtained in some studies also correlated with a
number of other stress indicators. However, prior to incorporation of these
methodologies, one has to be aware of their relevance with HS. Fecal cortisol
estimation provides an overview of the acute stress effect in animals wherein a
cumulative secretion of glucocorticoids may be observed. This secretion occurs for a
few hours coupled with a time delay between plasma glucocorticoid increase and
excretion of fecal cortisol that is primarily due to the gut passage time (between
8 and 16 h). Fecal cortisol estimation during HS was reported in a number of studies
conducted in cattle, sheep and goats. In a study led by Veissier et al. (2018), an
increase in fecal cortisol metabolites were observed with an increase in heat load
index in Holstein cattle.
Hair cortisol, another potential non-invasive methodology having wide applica-
tion in a number of livestock species like cattle, sheep, goat and pig, has been stated
to be a biomarker to detect stress. It may, however, be noted that hair cortisol depicts
the effect of chronic stress in animals. The glucocorticoids are excreted into the hair
shaft and stored within until estimation. The hair samples obtained can be wrapped
in clean sample collection papers or bags and kept at room temperature thereby
ensuring very convenient sample storage options. Additionally, as the estimation
methodology involves extraction of cortisol metabolites stored in the hair shaft,
chances of obtaining over-represented values, as a consequence of sample collection,
is ruled out. Furthermore, a study by Nejad et al. (2019) also proved HS to act as a
strong stressor that can increase hair cortisol levels in Holstein cattle. This method-
ology however needs to be adopted appropriately, as there are reports stating hair
cortisol levels to be influenced by hair color, hair type, body region collected from
and also sex. However, the impact of these factors can be nullified when a uniform
pattern of hair sampling is followed.
The metabolic products of glucocorticoids, released from the adrenal glands
during stress in animals, are excreted in feces and urine. Thus estimating the urine
cortisol metabolite concentrations is also a potential non-invasive approach. The
urine cortisol metabolites reflect the circulating cortisol levels in animals as it is
excreted through the glomerular filtration by kidneys. Similar to the feces and hair
cortisol level, urinary cortisol concentrations are also not influenced by animal
handling and rather provides an integrated index of cortisol production in the animal.
Though the use of urinary cortisol estimation in heat stressed animals is gaining
significance only in the recent past, this technique was extensively used in horses as
8 Non-Invasive Methods to Quantify the Heat Stress Response in Dairy Cattle 91

a doping test. However, with the rising concern of animal welfare, urinary cortisol
estimations have been conducted in cattle, sheep, goats, pigs and other domestic and
wild animals.
Therefore, feces, urine and hair can act as a vital medium to assess the classical
HS marker in livestock. Having known about their significance both from the ethical
point of view and also its increased accuracy to assess HS in animals, researchers can
definitely opt for this non-invasive methodology. However, a parallel study to
improvise the use of these techniques by developing quicker and easier protocols
that can be used directly at the field levels would be of practical high relevance.

8.6 Opportunities to Measure Body Temperatures

for Determining Heat Stress in Farm Animals

Body core temperature is one of the vital variables recorded to assess the impact of
HS in animals. A number of methodologies have been adopted to record body
temperature depending on the site of recording which may include, inter alia, rectum,
vagina, rumen, skin, body surfaces as well as ingestible biosensors and temperature
sensing ear tags with a temperature sensor placed within the ear canal. These
techniques differ in their degree of invasiveness, accuracy and also time lag on HS
response. Body temperature is one vital parameter in farm animals with economic
importance due to its association with health, production, and reproduction. Body
temperature might be elevated in animals due to illness, HS and other health related
issues. Rectal temperature is a routinely used measurement of body temperature.
However, it requires animal handling/restrain, which may cause a stress to the
animals. To overcome this problem associated with measuring the rectal tempera-
ture, a range of subcutaneous microchips and other implantable devices, temperature
sensing ear tags, rumen-reticular boluses, intra rectal and vaginal devices with
remote data transmission ability have been developed for the continuous measure-
ment of body temperature in farm animals.
Temperature sensitive microchips are injected subcutaneously and a handheld
receiver activates the microchip, which transmits a temperature reading to the
receiver (Chen and White 2006). Temperature recorded in this method is almost
instantaneous, and minimal handling of the animal is required. Using subcutaneous
microchips between the shoulder blades, body temperature can be measured compa-
rable to the rectal body temperature.
Rumen boluses, another advanced non-invasive remote sensing technology, are
designed to record rumen pH, rumen temperature, and activity in ruminant animals.
These devices are sensitive in reading body temperature and already transmit the
recordings via reading stations in the barn to a data cloud. Intra-vaginal and intra-
rectal thermo-sensor devices are also extremely sensitive to detect changes in body
temperature (Burdick et al. 2012). Thus, the use of technologies or tools that detect
elevated body temperatures reliably and in time can help to predict and prevent the
negative effects of HS.
92 G. Hoffmann et al.

8.7 Infrared Thermal Image Applications in Assessing

Thermo-Tolerance in Farm Animals

Among the recording of body core temperatures, the infrared thermography (IRT) is
gaining a lot of importance for its advanced features, reliability, non-invasive
approach, wide applicability to assess varied physiological status and diseases in
animals and finally for the minimal human animal interaction required. The infrared
thermal imager records the body temperature of animals by capturing the radiant heat
emitted from them, which involves a combination of body core temperature and
blood flow changes. This technology however differs from an infrared thermometer
that provides only a point based surface temperature recording. While the infrared
thermal imagers, being a field-based temperature measurement system, has a wider
area of detection and provides thermographic images having wide applications.
A number of studies have been conducted to assess the reliability of IRT to
effectively detect heat stressed animals. In a HS study conducted on Holstein (HO),
Girolando (½ Holstein  ½ Gir; considered as ½ HO) and ¾ Holstein  ¼ Gir (¾
HO) dairy cows, Daltro et al. (2017) established IRT to be effective in determining
HS in cows. Based on their study, using the IRT results, Holstein cows were reported
to be more susceptible to HS than ½ and ¾ Holstein cows. Further, based on this
study the authors stated udder region to be the ideal area that can determine the
thermal comfort in animals when compared to the other four regions (eye, right-area,
left-area and foot). In another study by Montanholi et al. (2008), IRT was stated to
have a potential application to detect heat production and methane production along
with detection of skin surface temperature fluctuations as a consequence of heat
increment due to feeding (physiological event) in Canadian dairy cows.
Figure 8.2 shows IRT images of an own experiment in different body parts of a
dairy cow.
Researchers across the globe have employed the use of IRT for HS studies in
other livestock species, too. Barros et al. (2016) stated IRT to be a useful and
accurate non-invasive methodology to detect variations in orbital area, left flank,
right flank and scrotum temperatures in buffaloes, having a positive correlation with
temperature-humidity index (THI). In another study in sheep, Kahwage et al. (2017)
studied the thermal tolerance of Morada Nova and Santa Inês rams with respect to
their ability to maintain body and testicular homeothermy when subjected to thermal
stress. The authors emphasized on the role of IRT as an efficient and alternative
non-invasive method for thermal mapping the animal’s body surface and also
stressed upon their usage as an auxiliary tool to detect thermo-tolerant animals.
Infrared thermography have also been used extensively in other species like goat and
pigs, however, its incorporation in HS studies is minimally documented.
Therefore, IRT can be considered as one of the vital non-invasive methods to
assess thermo-tolerance in farm animals. Having said that, this technology has a
number of confounding factors which need to be looked into so as to ensure
successful outcomes. As this methodology works primarily on the emissions from
the animal, the readings obtained thus are influenced by factors like skin and hair
variables (thickness, reflectivity, color, etc.), emissivity, health status of the animal,
8 Non-Invasive Methods to Quantify the Heat Stress Response in Dairy Cattle 93

Fig. 8.2 Infrared thermal images in different body parts of a dairy cow

feeding time, climatic variables (relative humidity, ambient temperature, wind

speed, sunlight) human factors (user’s distance and angle from animal). Therefore,
utmost care must also be taken to minimize these factors to lowest levels.

8.8 Models to Quantify Heat Stress in Farm Animals

Considering the existing close relationship between the environment and the body,
the direct influence of meteorological factors on the physiological state of productive
animals, the assessment of the animals’ comfort using integral indicators or indices
deserves special attention. The temperature-humidity index (THI), which is based on
air temperature (AT) and relative humidity (RH) measurements, has traditionally
been used to quantify the degree of HS in animals. THI is easy to calculate and it is
94 G. Hoffmann et al.

rather informative. Numerous studies indicate a close relationship between THI and
indicators of body temperature (BT), respiration rate (RR) and heart rate (HR),
widely used to assess the clinical state of animals during HS. A sufficiently high
correlation between THI and milk yield, as well as the content of milk components,
allows this index to be used in predictive models of environmental impact on dairy
cattle. Indices that take into account additional variables (solar radiation (SR), wind
speed (WS), etc.), which can increase or decrease the heat load, in addition to AT and
RH, are also applicable. In particular, Mader et al. (2006) found that the correlations
between THI and the degree of dyspnea ranged from r ¼ 0.47 to 0.87. Therefore,
adjusting the THI equation (for SR and WS) had a higher correlation with the
average dyspnea score (from r ¼ 0.64 to 0.80), which allows for a more accurate
assessment of animal discomfort. Proposed by Wang et al. (2018) Cattle Equivalent
Temperature Index (ETIC), including AT, RH, RS and WS, is more accurate in
predicting changes in physiological responses. The determination coefficient (R2)
for skin temperature, BT and RR was 0.79, 0.40 and 0.49, respectively, so ETIC can
be a useful tool to assess the effect of HS on animal comfort.
HS is an obstacle to the intensive development of animal husbandry, when the
livestock is forced to constantly stay in the buildings, not having free access to a
pasture. The difficulty is the fact that the microclimate of modern naturally ventilated
buildings (NVB) is highly dependent on external conditions that have a direct impact
on livestock welfare. Although the use of data from near located weather stations is a
recognized practice in modeling the response of animals to HS, the design of the
NVB itself (which is associated with indoor air turbulence) can be limiting in
building statistical models using such indices (Wang et al. 2018). In addition, the
error in the predictions themselves can be associated with a small number of tests, as
well as the lack of proper examining of the resulting models in field tests.
A recent study (Mylostyvyi et al. 2020) reported that statistical modeling was
effective in predicting THI values in dairy cowsheds based on multiple AT and RH
records outside and inside the cowshed over a wide temperature range. The use of
linear regression (LR) provided a high forecast accuracy (93–96%) of THI in NVBs
of various designs, depending on the external environment. The coefficient of
determination between the observed and estimated data was in the range
R2 ¼ 0.854–0.921. This approach makes it possible to predict the values of THI in
NVBs without the need for constant measurement of parameters (AT and RH)
indoors, that makes it possible to decrease animals stress. It also pointed out that
one should be careful in predicting the microclimate parameters in NVBs based on
weather station data, since there were significant differences between meteorological
data and measurements of AT and RH (differences of 1–3
C and 8–18%, respec-
tively) outside the cowshed during the warm period.
Thus, the use of thermal indices, as well as statistical models for their prediction,
can be useful in assessing the possible response of productive animals to HS, given
the strong correlation between the value of thermal indices, physiological variables
and the level of productivity. However, many uncertainties remain both in the
methods of monitoring the microclimate, namely the placement of AT and RH
sensors indoors (Hempel et al. 2018).
8 Non-Invasive Methods to Quantify the Heat Stress Response in Dairy Cattle 95

Table 8.1 Overview of different non-invasive methods and their fields of application in
dairy cows
Type of Type of
non-invasive adaptive
method response Field of application
Standing time Behavioural Helps to establish the behavioural reactions of the
Water intake response animals to its microenvironment in a non-invasive way
Feed intake
Drinking frequency
Defecating
frequency
Urinating frequency
Rumination time
Respiration rate Physiological Helps to establish the physiological response oriented
Skin temperature response reactions to the microenvironment
Infrared thermal
images/videos
Hair cortisol Endocrine Helps to identify the influence of heat stress on the
Urine cortisol response ideal stress marker cortisol
Faecal cortisol
Temperature- Barn climate Indices used for quantifying climatic changes in order
humidity index changes to describe the heat load on dairy cattle and its
Heat load index thresholds
Cattle equivalent
temperature index
Subcutaneous Physiological Quantifying changes in the body temperature of dairy
microchips response cattle
Implantable devices Physiological Quantifying changes in the body temperature of dairy
response cattle
Temperature Physiological Quantifying changes in the body temperature of dairy
sensing ear tags response cattle
Rumen-reticular Physiological Quantifying changes in the body temperature of dairy
boluses response cattle
Intra rectal and response Quantifying changes in the body temperature of dairy
vaginal devices cattle

Table 8.1 shows various non-invasive methodologies to quantify HS responses in

dairy cattle.
Technological advancements have also resulted to the invention of several
non-invasive biosensors like neck collars, leg bands, automated tympanic probes,
GPS collars, accelerometers, and so on. All these technologies enable real time
detection of a number of behavioral and physiological responses exhibited by
animals. However, their usage is mostly limited to the developed or well organized
farms. Moreover, these sensors are primarily used to livestock management
purposes, their incorporation into HS associated studies are gaining a slow
momentum.
96 G. Hoffmann et al.

Though such advanced biosensor technologies require substantial investments,

they can effectively detect heat stressed animals at an earlier stage thereby enabling
efficient management intervention to ameliorate its adverse impact that ultimately
prevent economic losses to the farmer and ensure animal welfare.

Learning Outcomes
• Animal welfare and ethical issues enforced to find out suitable non-invasive
techniques for accurate estimation of heat stress response.
• Non-invasive techniques such as activity sensors (behaviour measurement),
urine, hair and faecal cortisol (endocrine responses measurement), are
efficient methods to quantify heat stress response.
• Temperature sensitive subcutaneous microchips, ear tags and other
implantable devices, rumen-reticular boluses, intra rectal and vaginal
devices could be used for the continuous measurement of body temperature
in farm animals.
• Infrared thermography is another vital non-invasive tool gaining impor-
tance to assess the thermal tolerance in farm animals.

8.9 Conclusions

Cows have different possibilities to react under heat stress. It was shown in different
studies that dairy cows under heat stress have higher skin and core body
temperatures, higher respiration rates as well as an increase in standing time. All
parameters have proven to be useful evaluation criterias for the assessment of
physiological-ethological reactions of dairy cows under heat load. The sensors to
measure activity (accelerometers) and devices to measure body temperature (e.g.,
rumen boluses) already used in livestock farming are useful tools for the herd
manager and for the scientist. The respiration rate visually counted on the individual
animal is very sensitive and offers high potential in real-time monitoring for early
detection of heat load. However, visual counting is extremely time-consuming and
thus, difficult to implement in everyday life. Therefore, sensor-based parameters are
the first choice.

8.10 Future Perspectives

Heat stress in dairy cows is often described with a THI above 68, although this index
has some limitations such as the mistaken belief that all animals react in the same
way to thermal factors, and it does not include other parameters, such as solar
radiation, airflow and total hours of exposition. Based on the previously described
findings, the individual animal should be of more interest in the future. Therefore,
the reactions of individual cows to HS and the genetic background of these animals
8 Non-Invasive Methods to Quantify the Heat Stress Response in Dairy Cattle 97

are the subject of an ongoing study, where the climatic parameters find intensive
consideration as well. In this project, data from various digital applications, barn and
animal-specific data, will be merged into a complete system in order to better
manage, analyze, and interpret the individual information of the previous isolated
applications and make them available in the form of a flexible and application-
oriented decision support system. All measured data will be stored on a data platform
and analyzed in order to find thresholds were the cows show a stress reaction.
Furthermore, algorithms should be developed that process physiological and etho-
logical reactions of the cows in a combined way and incorporate them into manage-
ment decisions, so that complex deviations from normal animal behavior act as a
signal.

Acknowledgements The authors gratefully acknowledge the staff of the Department of Engineer-
ing for Livestock Management at the ATB (Potsdam, Germany) and the staff of the dairy farm at the
Agricultural Research and Education Center for Animal Breeding and Husbandry in Gross Kreutz
(Brandenburg, Germany) for their assistance with experimental setup and animal care.

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Applications of Infrared Thermal Imaging
and Rumen Boluses for Quantifying Heat 9
Stress in Cattle

A. M. Lees, A. L. Wallage, L. Labeur, S. L. Sammes, J. C. Lees,

and J. B. Gaughan

Contents
9.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100
9.2 Infrared Thermography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
9.3 Rumen Boluses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
9.4 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
9.5 Future Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109

Abstract
Heat stress is a leading cause of production and financial losses, not to mention
decreased welfare, for animal industries globally. With climate change increasing
the threat of heat stress to livestock enterprises, it is expected that these losses will
become more apparent and significant in future years. Elucidating the impact of
thermal challenges on cattle, and other species, has been a focal research area for
many decades and is likely to continue in future years. Technological
advancements have improved the capacity to collect large scale individualised
datasets, however this then presents challenges for analysis and interpretation of
data. Regardless, the increasing capability and sophistication of technology is

A. M. Lees (*) · L. Labeur · J. C. Lees

University of New England, School of Environmental and Rural Science, Armidale, NSW,
Australia
e-mail: [email protected]
A. L. Wallage · J. B. Gaughan
School of Agriculture and Food Sciences, Animal Science Group, The University of Queensland,
Gatton, QLD, Australia
S. L. Sammes
Feedworks Pty Ltd, Romsey, VIC, Australia

# The Author(s), under exclusive license to Springer Nature Singapore Pte 99

Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_9
100 A. M. Lees et al.

changing the way that thermal challenges are being investigated. Two such
technologies that have been increasingly utilised are infrared thermography
(IRT) and rumen temperature (TRUM) via rumen boluses. Each of these
technologies have been applied in various formats, with IRT in particular having
inconsistent results. This chapter, will focus on the applications of IRT and
boluses for the monitoring of heat stress in cattle and provide a perspective on
the future opportunities and challenges for applying these technologies in the
field.

Keywords
Animal sensors · Beef · Dairy · Heat load · Thermal challenge

Abbreviations


C Degree Celsius
AM Ante meridiem
bpm Beats per minute
BT Body temperature
h Hour
IPCC Intergovernmental Panel on Climate Change
IRT Infrared thermography
L Litre
MID Mid day
min Minutes
PM Post meridiem
RR Respiration rate
SE Standard error
SH AA Shaded Angus cattle
SH BH Shaded Brahman cattle
SH CH Shaded Charolais cattle
THI Temperature humidity index
TRUM Rumen temperature
UNSH AA Unshaded Angus cattle
UNSH BH Unshaded Brahman cattle
UNSH CH Unshaded Charolais cattle

9.1 Introduction

Recent climate change models are predicting an increase in global temperatures by

up to 6
C, before the end of this century (IPCC 2021). Irrespective of the magnitude
of this change, it is evident that our global environment is changing, such that over
the last 40 years each decade has been warmer than the previous one (IPCC 2021).
9 Applications of Infrared Thermal Imaging and Rumen Boluses for Quantifying. . . 101

The major adverse consequences of climate change include an increase in the

number of extreme weather events, and of particular concern is the prediction of
more frequent heat waves in conjunction with longer and hotter summers (IPCC
2021). These events are particularly important as climate change will have profound
impacts on animals, despite the application of mitigation techniques. The interest in
climate change and its impact on animals is not only centred around the fact that
animals will be exposed to more persistent periods of chronic heat stress, but also on
the ability of livestock to cope with climatic extremes. Specifically heat waves that
are longer and hotter in comparison to those events that currently occur. Therefore,
there is a need to investigate alternative non-invasive methods to evaluate how
animals are coping with their thermal environment. Automated monitoring individ-
ual animal responses has the potential to identify animals that are not coping with
heat stress (Islam et al. 2021a), and allow for targeted mitigation techniques to be
applied. The global community is becoming increasingly reliant on technology and
animal agriculture is not excluded from this. The adoption of ‘smart’ technologies in
agriculture, regardless of sector, is increasing to support production efficiencies to
achieve food security. Nevertheless, evaluating body temperature has and will
always remain an important mechanism to quantify of heat stress in animals.
However, collecting body temperature can be challenging under field and commer-
cial conditions (Lees et al. 2018b). Traditional methods of obtaining body tempera-
ture has typically involved relocating animals to handling facilities and/or animal
restraint, which are both associated with increases in body temperature (Mader et al.
2005; Lees et al. 2020a). Infrared thermography (IRT) and rumen temperature
(TRUM) collected by rumen boluses are two current non-invasive technologies that
are available and negate the need to handle and restrain animals to measure body
temperature.

9.2 Infrared Thermography

The use of IRT as a non-invasive measure of body temperature has been generating
an increasing amount of interest. Infrared thermography technology appears to be
useful in highly controlled circumstances, although IRT technology is currently
unpredictable in field conditions and inconsistencies exists between studies. Infrared
thermography measures the infrared radiation emitted from an animal, allowing for
the determination of body surface temperature (McCafferty 2007), which can be
used as a method for measuring and mapping the radiated heat loss at the body
surface (Labeur et al. 2017). Infrared thermography has had extensive diagnostic
applications in veterinary medicine (Schaefer et al. 2004; Metzner et al. 2014),
particularly inflammation (McCafferty 2007) and disease detection (Schaefer et al.
2004; Schaefer et al. 2012). Furthermore, there have been numerous studies
investigating IRT to understand the impact of scrotal heating on male reproductive
function and scrotal thermoregulation (Cruz Júnior et al. 2015; Menegassi et al.
2015, 2016a, b; Wallage et al. 2017). Specifically, in regards to using IRT, eye
temperature is a common focal region of interest and shows a greater potential to be a
102 A. M. Lees et al.

viable alternative non-invasive measure of body temperature although

inconsistencies still exist (Church et al. 2014). This is likely associated with high
blood flow and close proximity to the brain (Church et al. 2014). In addition, animal
restraint is still often required, thus reducing the non-invasive aspect of IRT (George
et al. 2014). Regardless, IRT has a clear advantage in that it is completely
non-invasive and requires little expertise to implement. However, images must be
captured at an angle, between IRT camera and animal, with an ideal angel of 90
,
otherwise there is a rapid decline in radiation measured by the camera, which
subsequently reduces the temperature reading (Speakman and Ward 1998). None-
theless, most IRT studies show that body surface temperatures generated by IRT are
typically generally below internal body temperatures (George et al. 2014; Wallage
et al. 2017; Lees et al. 2018b). If a consistent departure from internal body tempera-
ture could be accureately determined, then it is feasible that a correction factor could
be applied. Regardless, there are still questions concerning the accuracy of data,
remembering that this technology was designed for engineering purposes and
covering broad temperature ranges. When applied for use in animals it must be
considered that small changes in body temperatures have considerable implications
for wellbeing and production.
It is important to consider that the surface temperature of an animal is dependent
on many separate and unrelated factors such as exposure to sunlight, moisture or dirt
in the coat, type of coat and wind speed (McCafferty 2007; Hoffmann et al. 2013;
Church et al. 2014). As discussed by Lees et al. (2018b), some consideration also
needs to be extended to the emissivity value applied during IRT image analysis.
Steketee (1973) describes an emissivity value of 0.98 for biological tissues. How-
ever, it is probable that the emissivity of the coat surface can be influenced by
contaminants (McCafferty 2007), such as dirt (emissivity between 0.93 and 0.96)
and water (emissivity ¼ 0.96; Campbell and Norman (1998)). Therefore, determin-
ing the correct emissivity value where infrared thermography is used in field
conditions and in animals with differing coat characteristics becomes difficult.
However due to the linear relationship between radiative heat transfer emissivity,
McCafferty (2007) suggested that the differences in body surface temperature due to
environmental contaminants would be 0.5
C, for the typical mammalian coat.
Although, all heat stress physiologists would argue that a body temperature differ-
ence of 0.5
C is biologically significant and represents a significant physiological
change in body temperature (Lees et al. 2018b). Furthermore, the determination of
body surface temperature from IRT relies on the body being a 2-dimensional shape
and does not account for the natural curvature of the animal’s body. It remains
unclear what impact this has on the determination of body surface temperature.
Therefore, it is clear that there is a level of assumed errors in the body surface
temperature derived from IRT images.
Another confounding factor is that there is no predefined standard for the
assessment of IRT images, therefore, it becomes difficult to compare results across
studies (Lees et al. 2018b). The analysis, or extraction, of body surface temperatures
from IRT images remains a laborious manual task that has been discussed at length
between the authors of Wallage et al. (2017, Fig. 9.1), Labeur et al. (2017, Fig. 9.2)
9 Applications of Infrared Thermal Imaging and Rumen Boluses for Quantifying. . . 103

Fig. 9.1 An example of infrared thermal (IRT) image using the FLUKE software, showing an
infrared thermal (IRT) image and regions of interest consisting of the flank (a) and scrotum (b)
showing the use of the polygon tool to calculate minimum, maximum and mean temperature of the
selected area as described by Wallage et al. (2017)

Fig. 9.2 An example of infrared thermal image interpretation and analysis using the FLIR
software, displaying the arrangement of four regions of interest across the back of a lamb, being
(1) shoulder, (2) mid loin, (3) hips and (4) rump as described by Labeur et al. (2017)

and Lees et al. (2018b) and was explicitly noted by Cuthbertson et al. (2019).
Progress towards automating this process would be of benefit for the animal
sciences. It is probable, in order to gain a complete understanding of body surface
temperature obtained by IRT, that analysis should occur at the individual pixel level.
104 A. M. Lees et al.

For IRT to become a commercially viable tool that provides accurate real-time
results, an automatic method of data processing is needed (Cuthbertson et al. 2019).

9.3 Rumen Boluses

In many instances technological advancements, specifically rumen boluses, have

considerably improved the volume of individualised data that can be collected (Lees
et al. 2018a). This is an important consideration as previously studies investigating
body temperature have typically been restricted to 10 days. Rumen boluses present
a non-invasive method of obtaining body temperature over long periods of time,
months, and potentially years without compromising animal welfare (Lees et al.
2018a). As such, TRUM is rapidly becoming the most prominent method to obtain
body temperature data. Two obvious challenges with utilising TRUM are (1) heat
generated from digestion (Beatty et al. 2008), and (2) rapid declines in TRUM
associated with water intake (Bewley et al. 2008; Ammer et al. 2016; Cantor et al.
2018). Recently, Cantor et al. (2018) quantified that return to baseline TRUM is
dependent on water temperature and amount of water consumed. The authors were
able to show that 22.7 L of 1.7
C water took 103 min for TRUM to return to its
baseline temperature (Cantor et al. 2018). These findings confirm that water intake
results in a dramatic, but temporary, decrease in rumen temperature that is relatively
quick to return to baseline temperature. Furthermore, the time required to for TRUM
to return to baseline temperature depends on i) the amount of water consumed at
each intake and ii) the temperature of the water ingested (Cantor et al. 2018).
As with the majority of data captured by data loggers, TRUM data requires
extensive data cleaning and management prior to analysis. Specifically, with TRUM
data, decisions regarding the management of water intake need to be considered.
Recently Vázquez-Diosdado et al. (2019) developed a thresholding algorithm to
detect drinking events. Regardless of the management of drinking events, rumen
temperature follows a diurnal rhythm (Lees et al. 2018a, 2019a, 2021) (Fig. 9.3).
Across studies, TRUM generally increases between 08:00 h and 20:00 h, and
decreases, between 20:00 h and 08:00 h. Similar diurnal trends in alternative
measures of body temperature have been reported for TREC (Gaughan et al. 2004),
for abdominal temperature (Lefcourt and Adams 1996; Brown-Brandl et al. 2005;
Gaughan et al. 2010), and for tympanic temperature (Davis et al. 2003; Mader et al.
2010). Ammer et al. (2016) highlighted that TRUM was influenced by THI, in barn
housed lactating cows. Lees et al. (2018a), showed that that breed, ambient
conditions, and availability of shade have an influence on TRUM. This suggests
that TRUM is influenced by ambient conditions and by mitigation techniques.
In our previous, and current studies, rumen boluses capture data at 10 min
intervals, thus capturing 144 data points within each 24-h period (Lees et al.
2018a, b, 2019a, 2020b, 2021). The collection of larger data sets is advantageous
as it allows for more reliable statistical interpretations to occur. However, the
difficulty becomes identifying appropriate statistical modelling techniques for
these large time series data sets. This highlights the importance of consulting with
9 Applications of Infrared Thermal Imaging and Rumen Boluses for Quantifying. . . 105

Fig. 9.3 Diurnal rhythm using mean hourly (h) rumen temperature (TRUM) of (a) shaded Angus
steers over 128 days as adapted from Lees et al. (2019a); (b) un-shaded Angus (UNSH AA),
un-shaded Charolais (UNSH CH), un-shaded Brahmans (UNSH BH), shaded Angus (SH AA),
106 A. M. Lees et al.

biometricians/data scientists that have an intricate knowledge of these types of data

sets and is imperative to the successful evaluation of these data. In addition, the
generation of these large, individualised, data sets are inherently associated with an
increase in computational power required in order to analyse the aforementioned
data sets. The generation of these data sets is valuable for research activities, but may
be of little value to commercial industries. As such those companies that we have
worked within in our previous, and current studies, have established data manage-
ment software, and/or online databases and mobile applications, that have back-
ground algorithms to provide the ‘need to know’ information for commercial
producers, i.e. oestrus detection, high and low body temperature alerts, and insuffi-
cient water and feed intake. Thus, provides a versatile and effective animal health
management tool that extends beyond a heat stress application.

Learning Outcomes
• Infrared thermography (IRT) and rumen temperature (TRUM) measured by
rumen boluses are emerging as viable non-invasive technologies to quan-
tify heat stress response in cattle.
• Both IRT and TRUM are advanced technologies to provide detailed individ-
ual animal based datasets for body temperature, however this then presents
challenges for analysis and interpretation.
• Automated detection of panting score using accelometer based sensors can
also serve as a viable alternative to assess the heat load status in cattle.

9.4 Conclusions

The evaluation of body temperature is a fundamental aspect of quantifying heat

stress and numerous methods to determine body temperature exists. Given the
changing global environment, the negative impacts of heat stress are likely to
increase and expand into more temperate regions. As such non-invasive methods
of obtaining body temperature that are fast, efficient and reliable need to be
investigated. However, to be considered as an alternative method of determining
body temperature, and a viable commercial management tool, new technologies
need to be rapid and reliable. Currently, the inconsistencies and the laborious nature
of IRT evaluations render this technology not yet suitable for application in com-
mercial settings. However, if IRT image evaluation can be automated and used in
conjunction with machine learning techniques as described by Joy et al. (2021), new

Fig. 9.3 (continued) shaded Charolais (SH CH) and shaded Brahman (SH BH) steers over 130 days
adapted from Lees et al. (2018a); and (c) for three 6 day periods (P1, P2 and P3), adapted from Lees
(2016)
9 Applications of Infrared Thermal Imaging and Rumen Boluses for Quantifying. . . 107

opportunities will emerge for IRT to become translatable for commercial industries.
Rumen temperatures appear to be a reliable method to quantify heat stress, however
currently may not be economically viable for commercial industries. Technology
certainly has an important role in quantifying heat stress, the continued development
and sophistication of these technologies will support its extension and adoption in
future years.

9.5 Future Perspectives

In future years, autonomous monitoring for heat stress, and other ailments, are likely
to be based on minimally-invasive technologies such as IRT and rumen boluses, and
likely accelerometer-based technologies. This will enable real-time solutions to
animal responses under various production systems and environmental conditions
(Islam et al. 2021a). The future opportunities for ‘smart’ technologies in animal
science are unlimited. Currently, the greatest limitations are associated with our
ability to extract and analyse data from research and commercial perspective respec-
tively. Machine learning is the most probable method to support the automation of
data analysis moving forward and has been applied to animal behaviour in extensive
pastoral systems (Fogarty et al. 2020; Chang et al. 2022). Machine learning
techniques are likely to improve the effectiveness of both TRUM and IRT as real
time measures of heat stress in ruminants. Recently Joy et al. (2021) showed that
machine learning improved the reliability of IRT to monitor body temperature in
sheep, although individual IRT images appear to have been manually processed.
Thus, automating the processes involved in IRT images and interpretation would be
of great benefit.
Body temperature is going to remain the gold standard for determining an
animal’s thermal status. However, under commercial conditions collecting body
temperature may not be a viable option even with technological advancements.
Changes in respiratory dynamics, i.e. respiration rate and panting score, also
provides a good indication of heat load status in cattle (Mader et al. 2006; Gaughan
and Mader 2014), and sheep (Lees et al. 2019b, 2020b). Specifically, under field
and/or commercial conditions the assessment of panting score (Table 9.1) is a viable
alternative to using body temperature to assess the heat load status of cattle. Gaughan
and Mader (2014) showed that there was a strong relationship between body
temperature, respiration rate and panting score, confirming that panting score is a
good management tool for the assessment of heat stress (Fig. 9.4). Advancements in
accelerometer technologies may support the automated detection of respiration rate
and/or panting score in cattle. Recent heat stress specific studies are showing that
accelerometer-based ear tag sensors can monitor panting scores in cattle under ‘mild
to moderate’ panting (0–2, Table 9.1) (Islam et al. 2020) and ‘high to severe’ panting
(0–4, Table 9.1) (Islam et al. 2021b).
Rumen temperature boluses are likely to be cost prohibitive on a commercial
scale and with the laborious task associated with IRT image interpretation, other
alternative technologies need to be considered. Consideration is also needed
108 A. M. Lees et al.

Table 9.1 Modified assessment of panting score and description of breathing/panting condition
Panting
score Breathing condition
0 No panting
1 Slight panting, mouth closed, no drool, slight chest movement
1.5 Fast panting, mouth closed, no drool, fast easily observed chest movements
2 Fast panting, drool present, no open mouth
2.5 As for 2, but occasional open mouth panting, tongue not extended
3 Open mouth and excessive drooling, neck extended, head
3.5 As for 3, but with tongue out slightly and occasionally fully extended for short
periods
4 Open mouth with tongue fully extended for prolonged periods with excessive
drooling. Neck extended and head up
4.5 As for 4, but head held down. Cattle “breath” from flank. Drooling may cease.
Adapted from Brown-Brandl et al. (2006); Mader et al. (2006), Gaughan et al. (2008) and Lees et al.
(2021)

regarding the implementation of ‘smart’ technologies in extensive environments, for

example the northern pastoral zones in Australia and other regions in the world, that
have vast areas without access to electricity or access to internet. Mechanisms to
overcome these limitations will provide opportunities to investigate how thermal
challenges are impacting animals within grazing pastoral systems. Recently,
Mufford et al. (2021) showed that both behavioural and physiological,
i.e. respiration rate, heat stress responses could be determined by unmanned aerial
vehicles in feedlot and pasture settings. Moving forward, perhaps the combined use
of IRT cameras, accelerometer technologies, unmanned aerial vehicles and TRUM
will combined become the ‘gold standard’ for heat stress monitoring in both research
and commercial environments as we progress into the age of precision livestock
farming.
9 Applications of Infrared Thermal Imaging and Rumen Boluses for Quantifying. . . 109

Fig. 9.4 (a) Relationship between body temperature and panting score at 06:00 h (AM), 12:00 h
(MID), and 16:00 h (PM) and (b) Relationship between body temperature (BT  SE;
C) and
respiration rate (RR  SE; bpm) at panting score values of 0, 1, 2, 2.5, 3 and 3.5. Adapted from
Gaughan and Mader (2014)

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Skin Based Novel Approaches
for Establishing Climate Resilience in Goats 10
M. V. Silpa, Veerasamy Sejian, S. Koenig, Chinnasamy Devaraj,
C. G. Shashank, A. P. Kolte, G. B. Manjunathareddy,
and Raghavendra Bhatta

Contents
10.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115
10.2 Advantage of Goat Over Other Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
10.3 Significance of Assessing Thermo-Tolerance in Indigenous Goats . . . . . . . . . . . . . . . . . . . . 117
10.4 Inter-Relationship Between Skin and Microenvironment in Goats . . . . . . . . . . . . . . . . . . . . . 117
10.5 Skin Based Novel Approaches for Establishing Climate Resilience in Goats . . . . . . . . . 119
10.5.1 Coat Color . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
10.5.2 Hair Based Methodologies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
10.5.3 Significance of Active Sweat Gland and Sweating Mechanism in Goats . . . . 120
10.5.4 Applications of Infrared Thermal Imaging for Assessing Skin Based
Thermo-Tolerance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121
10.5.5 Application of Skin Based Metagenomics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121
10.5.6 Skin Transcriptomics Based Adaptive Assessment in Goats . . . . . . . . . . . . . . . . . 122
10.5.7 Application of Skin Epigenetic Changes as a Novel Tool for Assessing
Climate Resilience in Goats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123
10.6 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124
10.7 Future Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125

M. V. Silpa (*) · S. Koenig

Institute of Animal Breeding and Genetics, Justus-Liebig-Universität Gießen, Gießen, Germany
V. Sejian · C. Devaraj · A. P. Kolte · R. Bhatta
Centre for Climate Resilient Animal Adaptation Studies, ICAR-National Institute of Animal
Nutrition and Physiology, Bangalore, India
C. G. Shashank
ICAR-National Dairy Research Institute, Karnal, Haryana, India
G. B. Manjunathareddy
ICAR-National Institute of Veterinary Epidemiology and Disease Informatics, Bangalore, India

# The Author(s), under exclusive license to Springer Nature Singapore Pte 113
Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_10
114 M. V. Silpa et al.

Abstract

Climate change and food security are the two primary issues that daunt agricul-
tural researchers across the globe. The world is battling to find solution to both
these issues as we need to feed the expected human population of over 9.6 billion
by 2050. Therefore, it is very vital to sustain agricultural production and livestock
sector is particularly considered important for ensuring animal protein. Efforts are
also needed to identify the most heat resilient livestock species which could
produce optimally during the adversities of changing climate. Goats are consid-
ered the ideal climate resilient species. Researchers are currently working on
novel approaches to assess thermo-tolerance in livestock. This chapter is one such
attempt to elucidate the basic adaptive mechanisms in goats based on skin
approaches. The hair and skin are important targets to establish climate resilience
as these are the parts in animals which first get exposed to the microclimate. The
coat color is a vital variable to establish differences in thermo-tolerance in goats.
Further, the hair based variables such as hair characteristics and cortisol could
serve as reliable indicators for climate resilience. The skin based variables such as
sweating rate, active sweat glands distribution, and infrared thermal images may
play vital role in establishing climate resilience in goats. The latest next genera-
tion sequencing services such as skin metagenomics, skin transcriptomics and
skin epigenetic changes could revolutionize pathways for establishing climate
resilience in goats. Thus, this chapter highlights to the readers the above listed
skin based novel approaches to establish climate resilience in goats.

Keywords
DEGs · Goat · Hair · Infrared thermal images · Skin · Skin microbes

Abbreviations

ASIP Agouti signaling protein

CREB3L1 CAMP responsive element binding protein 3-like 1
DCT Dopachrome tautomerase
DEGs Differentially expressed genes
DNA Deoxyribonucleic acid
GDP Gross domestic product
GHG Greenhouse gas
GWAS Genome wide association study
HPA Hypothalamic-pituitary-adrenal
HSP Heat shock protein
mRNA messenger ribonucleic acid
NGS New generation sequencing
PMEL Premelanosome protein
RNA Ribonucleic acid
TRPM 1 Transient receptor potential cation channel subfamily M member 1
10 Skin Based Novel Approaches for Establishing Climate Resilience in Goats 115

TYRP1 Tyrosinase-related protein 1

UV Ultra-violet

10.1 Introduction

Climate change has emerged as the top most environmental challenge that mankind
had to face which is persistently threatening the well-being of future generations.
The alarmingly rising trend in global warming, with its erratic outcomes, has been
imposing a huge risk on the ecosystem, animal diversity and food security. Nearly
45% of the earth’s land surface is occupied by the livestock sector, most of which are
located in regions experiencing harsh and variable environments which are unsuit-
able for other uses. The livestock sector also plays a significant role by contributing
to nearly 40% of the global agricultural gross domestic product (GDP).
In addition to climate change, the exponentially rising human population along
with depletion in agricultural land use due to human invasion is also exerting a
massive pressure on food security. Though the livestock are thought to be least
affected due to climate change, when compared to other agricultural sectors, this also
holds a unique feature. While the livestock sector faces severe negative implications
due to the climatic variability, this sector also contributes to climate change. Live-
stock production across the world contributes directly and indirectly about 18% of
the greenhouse gas (GHG) emissions. Therefore, the livestock sector is of significant
importance from the current climate change perspective for its projected role to
ensure livelihood and also contribution to GHG emissions.
Numerous studies have been reported over the past decades that would assess the
impact of climate change on livestock production, compare this effect across species,
breeds and also at individual population level. Furthermore, a set of researchers have
also opted for a parallel study to establish ameliorative and mitigation strategies to
combat the adversities of climate change. In simpler terms, climate change and
livestock production can be considered as an entirely distinct field of science having
several branches. There are attempts to screen climate resilient livestock breeds with
the intention of identifying agro-ecological zone specific breed. This chapter is
therefore an attempt to collate and synthesis information pertaining to skin based
novel approaches to quantify heat stress response in goats. Such information could
be very valuable for future livestock production as the world is battling to find
solution to ensure food security to meet the requirements of growing human popula-
tion by 2050.
116 M. V. Silpa et al.

10.2 Advantage of Goat Over Other Species

Among all the livestock species, small ruminants are considered to be the go-to-go
species that can relatively adapt well and produce efficiently despite the distressful
climatic conditions. Furthermore, goats are proposed to be the ideal climate animal
model especially in the tropical environment. For the unversed, a major proportion
of the worlds’ livestock population is distributed across the developing countries
which experience most of the harsh environmental conditions. Additionally, these
regions have a prevailing tropical environment wherein livestock farming is mostly
characterized by extensive or semi-intensive farming practices involving small scale
farmers.
Goats are tipped to thrive better in such climatic conditions for their innate
abilities to cope with environmental extremities, sturdy and hardy nature. Goats
possess the ability to survive well even on low pasture and additionally they possess
unique feeding behavior. The narrow muzzle, mobile upper lip and prehensile
tongue in goats gives them an add on benefit to access forage from the soil level
and also for that matter, consume fodder resources like, shrubs, thorns spines, etc.,
that are usually not consumed by other livestock species. Moreover, the bipedal
stance behavior that is uniquely exhibited by goats gives them the superiority to have
access to tree fodder which are inaccessible to other ruminants due to the height. In
addition to these features, goats also possess the ability to travel longer distances in
search of feed and water. Moreover, goats are also stated to have better digestive
efficiency in comparison to other ruminants.
Moving ahead to the crucial climate resilience traits, goats in general are known
for their excellent thermal tolerance abilities. These species possess most of the vital
adaptive feature that enable them to survive efficiently in the extremely hot climatic
conditions. Moreover, goats are also considered to be extremely draught tolerant
species. They possess an inherent potential to conserve its body water reserve
(efficient rumen function), suppress evaporative water loss (modulating sweating
and respiratory mechanism) and also by reducing the excreted water (through urine
concentration and dry feces/pellets). Goats are well known for their better resistance
to diseases when compared to other livestock species. This is also a very crucial
advantageous trait of goats, especially on being aware of the impact of climate
change on health status in animal. Another worthy character possessed by goats is its
relatively lower enteric methane production ability. Therefore, goats stand out to be
the ideal climate resilient species that not only can sustain the climatic adversities
and yield quality products, but can also contribute minimally to GHG emission from
livestock.
10 Skin Based Novel Approaches for Establishing Climate Resilience in Goats 117

10.3 Significance of Assessing Thermo-Tolerance

in Indigenous Goats

The theory of goats being the ideal climate resilient animal species in comparison to
other ruminants has been emphasized and is also gaining concrete evidences for its
establishment. Having said there, the concept of breed variations also prevails. Time
immemorially it has been proved that indigenous germplasm overpowers the exotic
breeds for their immense thermo-tolerance and disease tolerance potential. Majority
of the livestock breeding practices have primarily focused to enhance the productive
potential in animals via incorporation of exotic blood line that are high producers.
Undoubtedly, such approaches did boost the livestock production multifold times,
however over the recent past, livestock breeder especially those located in the tropics
are raising their concern over such breeding practices. With climate change and its
impact on livestock production and food security being inevitable, there is an
increasing pressure to conserve and propagate the use of indigenous livestock
species.
The productive potential and climate resilience ability of indigenous livestock
species need to be studied intensely. There are substantial reports comparing the
productivity, thermo-tolerance and also disease resistant abilities between exotic,
crossbred and native breeds. However it is necessary to understand the variations
among the indigenous germplasm too. Based on the research outputs published from
a series of experiments that were under the leadership of Dr. Sejian and co-workers
(Shaji et al. 2017; Aleena et al. 2018; Pragna et al. 2018), significant differences in
the thermal resilience potential of indigenous goat breeds were observed. The
authors compared the adaptive potential of three indigenous South Indian goat
breeds, Osmanabadi (Karnataka), Malabari (Kerala) and Salem Black (Tamil
Nadu) on exposure to summer induced heat stress in Karnataka state of India.
Based on the cumulative results obtained from this project, it was interesting to
note that the Salem Black goat outperformed Malabari and Osmanabadi goats (that
were native to the experimental location). Thus the Salem Black goats were found to
adapt well for heat stress in addition to producing efficiently. This therefore urges the
need to conduct similar studies, exploring the indigenous goat breeds, so as identify
and promote the use of agro-ecological zone specific goat breeds.
There still lies a lot of hidden intricacies in understanding the thermal resilience
potential of indigenous goat breeds which needs to be addressed in the immediate
future. Thus, this also opens the scope for identifying and assessing novel adaptive
mechanisms that could be adopted by these indigenous goats.

10.4 Inter-Relationship Between Skin and Microenvironment

in Goats

Skin is the primary defense barrier that protects the animal from all the external
adversities. Unfortunately, skin is an underrated tissue for its proposed significant
role in maintaining homeothermy within the individual. Skin is among the very first
118 M. V. Silpa et al.

Fig. 10.1 Overview of skin based approaches to assess climate resilience in goats

body component to be exposed to both macro and micro environmental fluctuations.

For a matter of fact a portion of micro environmental alterations are effectively
tackled at the skin level itself. A number of factors aid to maintain the integrity of
this tissue which in itself hosts a complex niche that gets affected due to the slightest
alterations in the external environment.
The hair, coat and skin characteristics vary among species, breed and also
between individuals. These alterations have a high association with the adaptive
response exhibited by an individual. Therefore, unravelling the complexities at the
skin level could pave way for path-breaking research findings in the area of climate
change and livestock production. There are handful of reports assessing some of the
basic skin and its associated characteristics with respect to climate change, more
specifically heat stress, in livestock. However, like mentioned earlier, these are quite
superficial and needs to be explored further, considering a deeper and holistic
approach. Figure 10.1 illustrated an overview of skin based approaches for assessing
climate resilience in goats.
10 Skin Based Novel Approaches for Establishing Climate Resilience in Goats 119

10.5 Skin Based Novel Approaches for Establishing Climate

Resilience in Goats

10.5.1 Coat Color

Coat color is one of the well established and grossly observable characteristic that
has been associated with thermal stress in all livestock species. It is a proven theory
that darker color, specifically black, absorbs solar radiation while lighter shades,
white, reflect solar radiation. Therefore it is a foregone conclusion that goats with
black or darker coat would be more influenced during heat stress event when
compared to goats with white or lighter coat. However, researchers have also argued
on the protective role of black coats in goats that are found across tropical regions.
As one can reckon most of the indigenous goat breeds have colored coats. Addition-
ally some of the breeds that are well known for its hardy and sturdy nature are black
in color. The darker coat color in such scenario has been associated with the adaptive
response of goats wherein the melanin (responsible for dark pigmentation) produced
shields the animal from the harmful ultra-violet (UV) rays of the sun.
In a recent report from Democratic Republic of Congo by Baenyi et al. (2020), the
effect of coat color and sex on heat stress response was evaluated in goats. The
authors conducted the study in male and female goats with black, grey, brown and
white coat color that were exposed to solar radiation during summer. The impact of
heat stress in goats was observed through significant alterations in a number of
physiological, hematological and biochemical variables. Moreover, female goats
having black coat color were observed to be most affected during heat stress.
Furthermore, in a classic study that was dated back to the year 1995 by Acharya
and co-workers, heat tolerance in Sirohi goats with relevance to their coat
characteristics were assessed. Their study revealed higher alteration in physiological
responses to heat stress (rectal temperature, respiration rate and pulse rate) in goats
with black coat followed by dark brown, light brown and least in white coat goats.
The authors concluded goats with white or light brown coat to possess higher
tolerance to heat stress than those with dark brown or black coat.

10.5.2 Hair Based Methodologies

Hair characteristics are another important variable having high relevance in livestock
from climate change perspective. The hair characteristics like hair length, diameter,
density, and medullation play a crucial role in ensuring thermal insulation/heat
dissipation during the climatic extremities. In a study led by Riberio et al. (2018)
on Brazilian creole goats, the effect of dry and rainy season prevailing in the
Brazilian semi-arid region was assessed on the goats’ endocrine and physiological
profiles. The authors stressed upon the significance of hair variation to maintain
homeostasis in animals especially during seasonal alterations. The authors discussed
that animals having short, thick and well-seated hair are better suited for tropical
120 M. V. Silpa et al.

conditions. Furthermore coat thickness is also an important variable that aids in

homeostasis during environmental alteration in goats.
Hair also serves as a potential biological sample to assess the cellular and
molecular alterations occurring in an animal due to heat stress. Cortisol, considered
as an ideal heat stress marker, is usually estimated from blood samples. With the
rising ethical and animal welfare concern, researchers are encouraged to adopt
non-invasive methodologies during their experimental studies. Hair cortisol estima-
tion is one such approach that is projected to satisfy the animal ethical concern by
being non-invasive and also providing accurate results. Based on the experiment
conducted by Dulude-de Broin et al. (2019) in rocky mountain goats, hair samples
were concluded to be valid biomarkers that can be used to assess the HPA-axis
activity provided the confounding variables are taken into consideration.
Nucleic acids, DNA and RNA, which can be extracted from hair follicles, can
also be used to assess the heat stress induced molecular changes in goats. The DNA
and/or RNA extracted from hair follicles can pave way for molecular studies like,
relative gene expression, GWAS, mRNA and other high end next generation
sequencing approaches. Though the use of this biological sample for such studies
is limited in goats, their adoption is increasing over the years.

10.5.3 Significance of Active Sweat Gland and Sweating Mechanism

in Goats

Animals adopt several mechanisms to maintain their body core temperature when
exposed to harsh environmental conditions. Increased respiration and sweating are
among the many physiological adaptive strategies that aid to dissipate excessive
body heat in goats under heat stress. Goats can efficiently utilize their evaporative
cooling system-respiratory (increased respiration/panting) and cutaneous (sweating);
when exposed to hot climatic conditions. In a study reported decades ago on
dehydrated Black Bedouin goats, Dmi’el (1986) proposed that goats enable brain
cooling via selective sweating and panting modulation.
One of the simplest methods to estimate sweating rate in goats is by using cobalt
chloride impregnated chromatography paper disc (Berman 1957). Though this
methodology includes mild human-handling of the animal, yet it still can be consid-
ered to be a non-invasive approach. Moreover, being easy-to-use, portable and
cheap, this methodology is field friendly and farmer friendly too. Though there are
no much studies reported yet, that assess the sweating rate fluctuation in goats
exposed to heat stress; there are some reports in sheep (Sejian et al. 2012), cattle
(Schleger and Turner 1965; Dikmen et al. 2014) and buffaloes (Debbarma et al.
2020). Therefore, recording both sweating rate and number of activated sweat glands
could add on to the novel methodologies to estimate thermal resilience in goats.
10 Skin Based Novel Approaches for Establishing Climate Resilience in Goats 121

10.5.4 Applications of Infrared Thermal Imaging for Assessing Skin

Based Thermo-Tolerance

Body temperature is among the primary variable that is recorded in every heat stress/
climate change related study in any species. Though there are several possibilities to
record body temperature, recording rectal temperature is the traditionally followed
method. Technological advancements have led to the development of infrared
thermometers that can record the body surface temperatures. Adding on to this is
the development of infrared thermometry, which not only records the surface
temperature but also gives the thermal image of an animal. This technology is
been widely accepted and credited for its vast application in the livestock research.
Not only does it aid in providing an accurate measuring but also is non-invasive.
In an experimental study, the resilience of three indigenous goat breeds,
Osmanabadi, Malabari and Salem Black, to heat stress were assessed based on a
number of physiological traits and HSP70 mRNA expression by Aleena et al.
(2018). In this study, the authors recorded the body surface temperatures using
infrared thermometer. Using this technique, a significantly higher head, should and
flank surface temperature was observed in Osmanabadi and Salem Black goats
during the afternoon exposure to summer induced heat stress when compared to
their control. Furthermore, the heat stressed Malabari goats also exhibited signifi-
cantly higher head and flank surface temperature when compared to their control
however their surface temperature recorded were significantly lower than that of
Osmanabadi and Salem Black goats. The authors attributed this significant differ-
ence to the coat color wherein Malabari goats have pure white coat while Salem
Black and Osmamabadi goats have complete black coat.
In another study by Hooper et al. (2018), the physiological and cellular responses
of Saanen goats on exposure to acute heat stress was studied. The Physiological
variables recorded included respiration frequency, rectal temperature, dorsal and tail
surface temperature (using infrared thermometer); ocular and mammary gland tem-
perature (using infrared thermography). Based on this study, the authors could
understand the heat loss dynamics in goats. Infrared thermography therefore can
be stated as a potential non-invasive methodology having high relevance for
assessing climate resilience in goats.

10.5.5 Application of Skin Based Metagenomics

Metagenomics primarily looks into the isolation of genetic material that is recovered
directly from environmental samples. This next generation approach aids in
elucidating the microbial communities that inhabits in a specific niche and unravels
their population structure, genetic diversity and also their ecological role. Apart from
identification of novel organisms, this methodology also gives a sneak peek into the
functionalities of the associated microbes (Bashir et al. 2014).
In livestock species, metagenomics studies have been mostly used to assess the
rumen, fecal, udder and gastrointestinal microbiome. Narrowing down to studies
122 M. V. Silpa et al.

associated with heat stress, the rumen microbiome is the most explored niche using
metagenomics. Apart from influencing the animal, heat stress also leads to signifi-
cant alteration in the microbial populations that inhabit in an individual. Such
alterations based on the niche considered, would have varied implications on the
productivity and performance of the host. For instance, alteration in rumen
microbiota can impair feed digestion in animals and can also instigate the
methanogenic bacteria that harbors in the rumen.
Skin surface hosts a unique ecosystem comprising of varied micro-organism
which may also get altered due to heat stress. Alteration in the skin microbiota
may affect the skin integrity, skin characteristics and also may impair the primary
immune barrier provided by skin tissue. So far there are no studies reported to assess
the skin microbial diversity in heat stressed goats using metagenomics approach.
This therefore makes it a novel concept to explore and assess thermo-tolerance in
goats.

10.5.6 Skin Transcriptomics Based Adaptive Assessment in Goats

As mentioned in the earlier sections, the skin tissue, being the first tissue to be
exposed to climatic alterations, undergo a number of molecular changes. Such
alterations can be assessed by using the advanced methodology of transcriptomics
approach. This again is very minimally explored area in heat stressed goats, however
is proposed to provide greater inroad to unravel the hidden intricacies. Over the past
few years, there are increasing studies reported that assess the impact of heat stress in
goats through transcriptomics approach. However there is a scarcity of reports on
skin transcriptomics in goats exposed to heat stress or other environmental stress.
So far, most of the reported studies on skin transcriptomics in goats are associated
with either exploring the genes associated with coat color or hair growth. On
comparing the skin transcriptomics profile of Laiwu Black and Lubei White goats,
102 genes were reported to be differentially expressed (Peng et al. 2019). Among
these, six genes predominantly linked with pigmentation were identified; these
included agouti signaling protein (ASIP), dopachrome tautomerase (DCT), CAMP
responsive element binding protein 3-like 1 (CREB3L1), transient receptor potential
cation channel subfamily M member 1 (TRPM1), premelanosome protein (PMEL),
and tyrosinase-related protein 1 (TYRP1). A similar study providing insight into the
transcriptional regulation of black and white coated regions in crossbred goats (with
black head and white body) was studied by Xiong et al. (2020). Among the
165 differentially expressed genes (DEGs) identified on comparing the black with
white coated skin, a number of genes controlling pigmentation of skin and hair
follicles were identified. Furthermore, Agouti, DCT, and TYRP1, were reported as
the key DEGs associated with the melanogenesis pathway in the examined crossbred
goats.
Though such studies provided some vital information pertaining to the DEGs
associated with skin pigmentation, none of these studies assessed the alterations
upon heat stress exposure. So far there are no such reports documented in goats
10 Skin Based Novel Approaches for Establishing Climate Resilience in Goats 123

however there are very minimal studies conducted in cattle. Therefore, this opens up
the urge to incorporate this methodology to assess heat stress impact in goats.

10.5.7 Application of Skin Epigenetic Changes as a Novel Tool

for Assessing Climate Resilience in Goats

Acquiring genomic information has substantially increased the accuracy of selec-

tion/prediction with an appreciable time reduction, to assess the impact of heat stress
on animals and also identify climate resilient individuals. Heat stress induces a
number of cellular and molecular changes in an animal some of which is evident
from altered gene expression profile. Epigenetics is one such phenomenon that
controls gene expression. Epigenetics basically describes the gene-environmental
interaction that results to evident alterations at the phenotypic level. Therefore,
assessing the epigenetic profile in animals under heat stress, which is a classical
host environment interaction, provide a better understanding of the molecular
alterations occurring in an individual.
DNA methlyation and histone modification are the common mechanisms that
describe epigenetic alterations in animals. Among these, DNA methylation is better
studied epigenetic regulatory mechanism in livestock. There are very limited reports
assessing the epigenetics profile of heat stressed livestock species. This should
thereby instigate researchers working on climate change and livestock production
to apply the novel concept of epigenetic profile of animals under environmental
stress. Apart from influencing the gene expression, epigenetic alterations that are
triggered in an individual due to environmental stress can also be passed down to its
future generations. This therefore could explain the epigenetic inheritance acquired
by an individual from its ancestors.
In a heat stress study lead by Del Corvo et al. (2021), the methylome patterns of
Nellore and Angus cattle were explored. Using Reduced Representation Bisulfite
Sequencing, the genome-wide DNA methylation profiles for the heat stress resilient
Nellore and heat stress susceptible Angus cattle were obtained. The authors
identified 819 genes that were significantly methylated during heat stress and
recovery period in cattle. Among these 351 genes were specific to Angus while
366 genes were unique to Nellore. It was observed that heat stress resulted in breed-
specific responses wherein genes associated to stress response and cellular defense
were under methylated/hypo-methylated in Nellore while some of these genes were
hyper-methylated in Angus. This study therefore signifies the impact of heat stress at
the epigenetic level in animals and urges the need for extensive future research.
Epigenetic profile is tissue specific and thereby assessing the skin epigenetic
profile would give a clear understanding on how this potential tissue, that is the first
to be exposed to environmental alterations, responds to the associated stress. So far
there are no published reports assessing the epigenetic profile of caprine skin to heat
stress. Hence this would also be a novel tool to assess heat stress impact and further
screen for thermo-tolerance in goats.
124 M. V. Silpa et al.

Learning Outcomes
• Skin based novel approaches can be considered as reliable non-invasive
tools to identify climate resilient agro-ecological zone specific goat breeds.
• Hair, urine and faecal cortisol and infrared thermal images of skin surface in
animals may provide useful information to assess thermo-tolerance in
livestock.
• Skin based metagenomics, transcriptomics and epigenetic changes analysis
can provide novel insights into goat adaptation to harsh climatic condition.

10.6 Conclusions

This chapter covers in a nutshell the various skin based approaches and variables
which could be considered significant in assessing climate resilience in goats. This
approach could play a vital role as it is easy to assess heat tolerance in goats both
invasively and non-invasively. Hair cortisol and infrared thermal images can be
reliable indicators to quantify heat stress response in goats non-invasively. Further, it
is very easy to take skin biopsy to subject them for advanced NGS analysis for
elucidating the skin microbial diversity, differentially expressed genes and
pathways, as well as to establish skin epigenetic changes associated with climate
resilience. Therefore, this skin based novel approaches could help to concretely
establish thermo-tolerance in goats.

10.7 Future Perspectives

Researchers and policy makers working to sustain livestock production in the

changing climate scenario are looking to establish novel ways and means to quantify
heat stress response. Although there is evidence that light coat color animals may
have the advantage over dark coat color, still this concept has not been fully explored
in all farm animals at mechanistic level. This warrants more such research efforts at
mechanistic level to explore the unexplored aspects of skin based novel approaches
to establish climate resilience in all farm animals. This approach could play a vital
role to ensure food security by 2050 by providing clue as to which are all the species/
breeds those needs to be taken forward for sustainable livestock production. The skin
based approaches may be an important area of research for future to establish the
basic and fundamental aspects of livestock adaptation due to the easy accessibility as
skin being the first body part getting influenced by the microclimate. More intense
research efforts are also needed to compare the adaptive potential of farm animals
based on skin based shotgun metagenomics, whole transcriptomics and bisulphate
sequencing NGS analysis. Such approaches may yield important biomarkers for
climate resilience in farm animals which could be used in future breeding program
10 Skin Based Novel Approaches for Establishing Climate Resilience in Goats 125

using marker assisted selection. This could help to revolutionize the future animal
breeding for climate resilience in farm animals. Therefore, skin based novel
approaches to establish climate resilience in farm animals could be the way forward
approach for policy makers to try and establish permanent solution for climate
change associated livestock production. Further, these novel approaches can help
to identify the best climate resilient breeds specific to agro-ecological zones. By
disseminating these breeds to the resource poor small and marginal farmers can help
to ensure their livelihood in the changing climate scenario.

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 Part IV
Strategies to Ameliorate Heat Stress Impacts
Livestock Shelter Management: Climate
Change Perspective 11
Ravindra Kumar, Ankaj Thakur, Rakesh Thakur, and P. K. Dogra

Contents
11.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
11.2 Shelter Design Considerations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132
11.3 Provision of Shade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132
11.4 Housing System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
11.5 Roofing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134
11.6 Insulation and Flooring . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
11.7 Space Requirements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
11.8 Ventilation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
11.9 Thermal Stress Alleviators . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136
11.10 Thermal Stress Ameliorative Measures Used by the Farmers . . . . . . . . . . . . . . . . . . . . . . . . 136
11.11 Tunnel Ventilation (Fan and Pad System) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136
11.12 Integrated Farming System Shelter Models . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
11.13 Cooling Ponds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
11.14 Manure Management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
11.15 Interventions for Pastoralism Production System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
11.16 Nutritional Amelioration Strategies in Dairy Cattles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
11.17 Future Perspective . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139

Abstract

Increased heat stress of the animals is anticipated to jeopardize food security,

making climate change a serious global concern for livestock production. Live-
stock has an immense role to play in the food security in coming years due to

R. Kumar (*)
Bihar Animal Sciences University, Patna, India
A. Thakur · R. Thakur · P. K. Dogra
COVAS, CSK Himachal Pradesh Agricultural University, Palampur, India

# The Author(s), under exclusive license to Springer Nature Singapore Pte 129
Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_11
130 R. Kumar et al.

rising human population. Therefore, development of appropriate strategies to

sustain livestock production in the changing climate scenario by gains signifi-
cance. Altering the housing design is an immediate and cost-effective approach to
reduce the magnitude of the climatic change on livestock production. Climate
change adaptation through livestock housing can be achieved through appropriate
east-west oriented shelter design, improved ventilation, using fans and other
cooling devices, and using appropriate roof materials to improve the thermal
comfort of the livestock. Naturally ventilated house should always be oriented in
an east-west direction in tropical climate as this direction will minimize the
possibility of direct sunlight entering the house. Properly designed roof overhang
reduces the possibility of direct and indirect sunlight as well as rain water entering
the house during hot and rainy weather respectively. Further the shelter design
should be oriented in such a way to cater the needs of specific agro-ecological
zone. Thus, the chapter provides an overview on appropriate shelter design and
management strategies to reduce the impact of climate change associated envi-
ronmental stresses.

Keywords
Adaptation · Climate change · Cooling devices · Heat stress · Housing · Shelter

Abbreviations

% Percentage

C Degree Celsius

F Degree Fahrenheit
BCS Body condition score
CH4 Methane
FAO Food and Agriculture Organization
Fig Figure
ft Feet
GHG Greenhouse gas
h Hour
IFS Integrated farming system
LCT Lower critical temperature
N 2O Nitrous oxide
NAAS National Academy of Agricultural Sciences
pH Potential of hydrogen
THI Temperature humidity index
11 Livestock Shelter Management: Climate Change Perspective 131

11.1 Introduction

Climate change which includes increase in earth’s near surface temperature, erratic
changes in seasonal patterns, increase in frequency of extreme climatic events like
heat waves, droughts, floods and precipitation has emerged a serious threat to the
environment, economy and welfare of both humans and livestock. As a result, in the
Conference of parties 26th meeting at Glasgow 2021, all countries have taken a
position at key climate related issues. India has promised a massive one-billion-ton
reduction in emission by 2030, a coal phase down, a radical shift to renewable and a
net zero emissions target for 2070. The climate change impacts are visible all over
the world but South-Asia appears to be most vulnerable region. Almost 70% of
livestock in India is owned by small and marginal farmers along with landless
livestock keepers. Further owing to limited financial resources and lack of awareness
cum adoption of climate resilient technologies their animals are even more vulnera-
ble to adverse impacts of climate change (NAAS 2016).
Increased heat stress of the animals is anticipated to jeopardize food security,
making climate change a serious global concern for livestock production. Livestock
are key to food security. Livestock produce contribute 34% of the protein and 14%
of total calorie consumed globally as well as essential micronutrients such as vitamin
B12, A, iron, zinc, calcium and riboflavin. But their contribution to food security and
nutrition goes well beyond that, and includes a range of other goods and services,
such as animal manure and traction. Hundreds of millions of vulnerable people rely
on livestock in a changing climate, because of animals’ ability to adapt to marginal
conditions and withstand climate shocks.
Livestock has an immense role to play in the food security in coming years due to
rising human population; however, its role in contributing to GHG is also being
criticized. Livestock accounts for significant amount of methane (CH4) and nitrous
oxide (N2O) emissions worldwide. Ruminants besides suffering from impact of
climate change are also contributing to it through enteric CH4 emission and manure
management. FAO has proposed the following three ways to substantially reduce
emissions from livestock production: productivity improvements that reduce emis-
sion intensities; carbon sequestration through improved pasture management better
livestock integration in the circular bio economy. Similar alterations like dietary
changes, selection for heat resistant germplasm etc. are being put forth for better
adaptation and mitigation of impact of climate change. In general, livestock systems
under intensive management are less vulnerable to impact of climate change com-
pared to those managed under extensive or semi-intensive management owing to
better shelter and adoption of mitigation strategies. This paper focuses on shelter
management of livestock from climate change perspectives.
Altering the housing design is an immediate and cost-effective approach to
reduce the magnitude of the climatic change. Network project on animal housing
and management, India has also put forth the need to generate more information on
type of most appropriate housing and management for different species in different
climatic zone and different management systems (NAAS 2016). Main goal of
housing is to regulate the microclimate of animals and maximize their productivity
132 R. Kumar et al.

by shielding them from weather extremes. Selection of site is the preliminary step in
livestock housing. Ideal housing can help in providing optimum animal productive
and reproductive performance. Ventilation, cost of construction, proper drainage,
lighting and ease of cleaning need to be considered while designing house for the
livestock.
The structure of housing system (combination of the floor-system, manure col-
lection and removal) impacts the level of emission of gaseous compounds especially
ammonia. At least one or more of the abetment principles are used in housing
systems to reduce gaseous emissions viz. reduction of emitting manure surface,
rapid and complete removal of the liquid manure from the pit to external slurry
storage, applying an additional treatment, such as aeration, to obtain flushing liquid,
cooling the manure surface and changing the chemical/physical properties of the
manure, such as decreasing the pH (Melse et al. 2009). Housing systems that use the
aforementioned principles can minimize gaseous (ammonia) emissions to the atmo-
sphere from 30% to 80%.

11.2 Shelter Design Considerations

• Recommended eave height is 12 ft. for the structures up to 40 ft. width and 16 ft.
for the structure wider than 40 ft.
• Naturally ventilated housed should always be oriented in an east-west direction in
tropical climate as this direction will minimize the possibility of direct sunlight
entering the house. While in temperate areas orientation may be north to south to
get maximum benefit of sunlight.
• Properly designed roof overhang reduces the possibility of direct and indirect
sunlight as well as rain water entering the house during hot and rainy weather
respectively.
• Cows drink up to 50% more water when the temperature humidity index (THI) is
above 80%, hence cool and fresh water should be readily available to the cows
during the summers. Also the water point should be located near to the shade
areas. In close confinement waterer should be available every 50 feet in case of
dairy cows.
• Kumar et al. (2012) proposed different animal shelter designs for different zones
of the Himachal Pradesh, taking in to considerations the climatic variations and
availability of local materials (Fig. 11.1). Similar studies need to be replicated in
different parts of the country so that the impact of the climatic stress can be
reduced through shelter management.

11.3 Provision of Shade

Shade is a cost-effective solution for reducing the heat load on the livestock. Total
heat load of 30–50% can be reduced with a well-designed shade, but it does not
change the air temperature or relative humidity around animals to increase sensible
11 Livestock Shelter Management: Climate Change Perspective 133

Fig. 11.1 Livestock shelter designs for different agroclimatic zones of Himachal Pradesh

heat loss routes. The use of natural shades is not always ideal under grazing
conditions, which is why artificial shades have become popular. Artificial shades
have been shown to improve animal welfare and milk production when used under
grazing situations (Valtorta et al. 1996, 1997). Shade cloth can also be used as an
alternative to solid roofing materials for providing shade to the livestock.

11.4 Housing System

Thermal resistance of the roof negatively radiates heat on animal during night hour.
Roof creates higher indoor temperature during night time which can result in
increased stress to the animals. The outside THI was found to be lower between
17:30 h and 7:43 h, maximum difference was obtained at time point of 6:07 h
(Ambazamkandi et al. 2015). Therefore, the loose housing system in the hot humid
tropical conditions should be practiced as it allow continuous access to the open
paddock. Loose housing system when compared to the tie stall barn system is more
appropriate for the livestock. Bamboo domes for the young livestock are used in the
low cost rearing system of small ruminants during winter which help in preventing
mortality and slow growth rate. This prevents the young one from the cold stress as it
maintains higher temperature during extreme cold. If enclosed livestock housing is
used, the building design must account for a range of ventilation flow rates. During
cold seasons, the minimum ventilation rate should remove the moisture created
while retaining as much sensible heat as possible by using thicker walls
(Fig. 11.2). The maximum ventilation rate should remove enough of the sensible
heat produced to maintain a small temperature difference between inside and
outside, usually 2–4  C.
134 R. Kumar et al.

Fig. 11.2 Inner view of typical animal house (a) and multispecies shelter (b) in colder areas of
Himachal Pradesh

11.5 Roofing

Roof of animal shed should provide sufficient shade, preventing solar radiations
from entering the shed. Insulating capacity should be good in the roof materials.
Aluminum sheets besides being more durable are more reflective to solar radiations.
In small holder production system cheap materials such as palm leaves and dried
grass is used for roofing. Skin temperature of the livestock remains higher in the
asbestos-roofed shed as compared to thatch roofed shed. In tropical climate thatch
and agro-net roof are better when compared to asbestos in preventing the animals
from hot and humid environment (Hatem et al. 2015). During the summer, the roof
inside surface temperature ( C) of a thatch roof was lower than that of an asbestos
roof, and both thatch and agro-net shade material contributed to alleviate thermal
stress in crossbred cattle (Kamal et al. 2014).
Gable roof should have slope of 4:12 and a continuous ridge, overshot ridge or
raised ridge cap to promote natural ventilation. In houses with uninsulated roofs a
steep slope (45 ) is highly desirable as it tend to collect the less radiant heat when
compared to flat roof and the distance between animal and hot celling is also
increased. Painting of the roof (outside or inside with black/white) material can
also reduce the stress on the animals by minimizing the radiations. In colder areas the
roof height should be low with wide overhang and the roof should be painted black.
Roof sprinklers can also be used to reduce the heat gain from the uninsulated roof.
When a metal roof is used that isn’t insulated, sprinkling during hot weather is
particularly effective in reducing heat.
11 Livestock Shelter Management: Climate Change Perspective 135

11.6 Insulation and Flooring

In tropical climate the livestock sheds should be insulated to limit the amount of heat
entering through wall or roof insulation. Roof insulation can also reduce the solar
penetration into the shed, hence should be considered while building some new
cattle shed. Spray polyurethane can also be used for insulation of roof. Thermal
conductivity of the floor influences the thermoregulatory behaviour of the animals.
Straw bedding significantly reduces the lower critical temperature (LCT) as com-
pared to the bare concrete. Gupta et al. (2004) reported that morning rectal tempera-
ture was higher in village type closed barn with kutcha floor than the cemented one.
Rubber mat floor has been reported to have significantly higher temperature than
concrete floor during daytime (Prasad et al. 2013). Though rubber mat flooring offers
a lot of obvious benefits, more research is needed to see how it affects thermolysis,
animal comfort, and physiological relevance in hot, humid environments. The
addition of a thatch ceiling and soft flooring in the form of a sand bed improved
the microclimate of the shed, alleviated stress, and increased milk output in cross-
bred Jersey cows (Sahu et al. 2018).

11.7 Space Requirements

Temperature may rise dangerously if the stocking density is too high for the housing
and ventilation system because more metabolic heat will be contributed to the house
air than was planned for. Space allowance given to the livestock has an effect on the
synchrony of resting and feeding. Floor space availability per animal can be
increased by 10–20% during hot summer i.e. stocking density need to reduce by
10–20%.

11.8 Ventilation

The elimination of contaminants originating from animal excreta and the thermal
exchange between the animal’s surface and the environment are both dependent on
ventilation. Assessment of the airflow can be easily done by using smoke cartridge to
see where physical improvements can be made before switching to the mechanical
ventilation solution. Increasing the airflow above the cows has a dramatic effect on
evaporative heat loss from the skin.
Airflow can be increased by two ways, one by installing fans and other by
opening sides of the barn. Key areas to keep cool and to determine placement of
fans are at feed bunk, holding area and fresh cow pen. Minimum 50% of the floor air
covered with the air movement produced by the circulating fans. If the temperature
of the provided air is lower than the animal’s surface temperature, fans can reduce
body temperature by 0.3–0.4  C. In barns where metal sheet is used for the walls it is
practical to remove the sides and install netting. The netting/curtains can be raised to
increase the airflow during summer and lowered during the winter.
136 R. Kumar et al.

11.9 Thermal Stress Alleviators

In high-humidity situations, every intervention that introduces water into the envi-
ronment should be examined because it may have negative consequences as water
evaporates, increasing humidity and saturating the air. Evaporative systems are
designed to combine forced ventilation with wetness for this reason. Evaporative
cooling systems uses the energy in the air to evaporate water, and the evaporation of
water into warm air lowers the air temperature while raising the relative humidity. If
the area is having higher relative humidity, larger droplets of water with fans are
needed but if lower relative humidity is present, a fog or misting system would
effectively cool the animals. Evaporative cooling should be used if the temperature
remains above 100  F (38  C). Sprinkling increases the amount of heat loss from the
animal via evaporative cooling while lowering water expenditures (Gaughan et al.
2008).
Combination of intermittent wetting and forced ventilation can dramatically
reduce the effects of the heat stress in animals. Spraying should be done in cycles
combined with the air movement. Intermittent wetting and forced ventilation using
automated device had favourable responses on the physiological parameters and
milk production of Murrah buffaloes by alleviating the thermal stress (Sruthi et al.
2019). Fifteen-min intervals have proved suitable for operation, whereby water is
sprayed for about 3 min, controlled by a timer, followed by an evaporation time of
12 min. Approximately 1 L of water per square metre of surface area can be
sprinkled in this process.

11.10 Thermal Stress Ameliorative Measures Used by

the Farmers

Bathing the animals frequently, wetting the body of animals with the gunny bags,
hanging the wet gunny bags against the direction of wind, growing tress or increas-
ing green cover around the animals shed or paddocks are the few thermal stress
ameliorative measures practiced by the farmers. Buffaloes are also taken to village
pond for wallowing during summer to reduce the impact of heat stress in rural areas
of north India, which has been proved very cost effective heat stress ameliorative
measure.

11.11 Tunnel Ventilation (Fan and Pad System)

Tunnel ventilation technology is widely utilized in swine and poultry houses, but it is
new to the dairy industry. It provides air movement and air exchange through fans
placed in one end wall of the barn. A bank of high-powered fans is often installed on
either the long or short end of a rectangular barn to improve wind speed in the barn at
cow level and facilitate cow cooling via conduction and convection (Mondaca and
11 Livestock Shelter Management: Climate Change Perspective 137

Cook 2019). Evaporative cooling with cooling pads, misters, or sprinklers is fre-
quently included in tunnel-ventilation barns.

11.12 Integrated Farming System Shelter Models

Integrated farming system (IFS) or whole farm approach is emerging as a nature

based solution to boost the productivity and income of the farmers by generating a
mix of enterprises viz. fish, pigs, poultry, rice, vegetables etc. in climate stress
scenario. IFS involve use of on-farm resources and reduce dependence on external
inputs such as fertilizers and pesticides. Several design considerations can be used to
integrate to optimize production in the limited space available in the tropics through
vertical height utilization (Ambazamkandi et al. 2015). Roof top cultivation in the
integrated models can also further help in reducing the thermal stress.

11.13 Cooling Ponds

A cooling pond is a dynamic, man-made structure that is maintained in its dynamic

state by adding water to keep a set water level dependent on cow movement and
ambient conditions. Within minutes of entering the pond, the internal temperature of
the cows dropped by 0.5–1  C (Bray et al. 1989). Conduction and coefficient of heat
transmission to water through skin are the primary ways in which animals in ponds
lose heat quickly to cool water. In locations where heat stress is an issue, using a
cooling pond on a farm may help both animal comfort and production without
harming milk quality (Tomaszewski et al. 2005).

11.14 Manure Management

Manure handling related mitigation practices include reducing the exposure of

manure to water (e.g. dry scraping rather than washing into a pond) and changing
management from anaerobic to aerobic conditions. Methane emissions from a
slurry-based manure management system rise as the temperature of stored slurry
rises. The lowering of slurry storage temperature from 20 to 10  C resulted in
30–50% reduction in methane emissions (Hilhorst et al. 1998). Ammonia and
methane emissions from the cow housing can be lowered by more regularly remov-
ing dung to a closed storage facility and scrapping the floor on daily basis. In
advanced poultry housing belt scrapers can be used efficiently to remove litter/
manure continuously, decreasing greenhouse gas emissions (Fournel et al. 2012).
138 R. Kumar et al.

11.15 Interventions for Pastoralism Production System

Grazing systems due to their reliance on climatic conditions and the natural resource
base, as well as their limited adaptation options, are expected to suffer the most
devastating consequences (Aydinalp and Cresser 2008). Pastoralists stay with their
livestock without any shelter for about 4 months in grazing area (in north western
Himalayan region) which is devoid of even trees, however providing basic
nutritional supplements can also help in maintaining them in healthy conditions.
Building the shelters for the accommodation of the livestock along the selected
points of the migratory routes can help in building resiliency of nomadic pastoralism
during extreme climatic conditions.

11.16 Nutritional Amelioration Strategies in Dairy Cattles

The heat stress is the most detrimental constraint to livestock production in lower
gangetic region as the region is affected by both the climatic extremes. Considering
the facts, supplementation of Selisseo and Nutri-ferm on both Indigenous and
crossbred animals at Bihar Animal Sciences University Cattle Farm, Patna, Bihar
as nutritional amelioration strategies for heat stress to sustain milk production in
Dairy Cattles showed very encouraging results. The outcome showed that the
experimental house temperature and humidity varied between 31–33  C and
81–85%, respectively, whereas, it was highest recorded in last fortnight of August
month of 2021. There were non-significant changes in feed intake, body weight and
physiological parameters, however, slight increase in body weight and better feed
efficiency recorded in both the treatment groups. Body condition score (BCS) was
significantly higher in both treatment group in comparison to control. There was
non-significant increase in milk yield and milk composition.

Learning Outcomes
• Scope for better shelter management and amelioration strategies make
intensive system more profitable than extensive and semi-intensive systems
of rearing livestock in the changing climatic condition.
• Providing artificial shades in the extensive system and intermittent wetting
and forced ventilation in intensive system rearing can help to augment
livestock production.
• Climate change calls for multidisciplinary approach in designing innova-
tive integrated and self-sufficient multi-climate shelters to ensure optimum
livestock production.
11 Livestock Shelter Management: Climate Change Perspective 139

11.17 Future Perspective

The vulnerability of livestock to climate change is defined by duration, frequency,

severity and sensitivity, location and related assets. Developed countries, in particu-
lar, are more inclined to implement costly adaptation measures for climatic change,
such as modified housing. New paradigms are required for livestock shelter
buildings to enable low greenhouse gas emissions Climate change adaptation
through livestock housing can be achieved through improved insulation, passive
building designs to improve ventilation, climate control by an appropriate natural
ventilation system, passive and active solar design for heating and cooling, cooling
pads, earth-air heat exchange, geothermal cooling using underground water and by
using smart technologies to improve the thermal comfort of the livestock. Adapta-
tion measures at animal level includes forced ventilation system, cooled lying areas,
thermoregulation with water wallows, cool drinking water, fogging, misting and
sprinkling system. The sensible heat load in the barn can also be reduced by reducing
the stocking density of livestock in summers. Capturing of emissions in the barns
and separation of feces and urine has a great potential to reduce the greenhouse
emissions. Vertical separation of feces and urine employing plates on solid floor
appears promising from animal welfare point of view, as the floor is drier and less
slippery as compared to conventional floors. Artificial floor composed of different
layers can also help in reducing the ammonia emissions due to separation of feces
and urine. Therefore, the climate change’s impact on livestock can be reduced using
a multidisciplinary approach, innovative, integrated, self-sufficient multi climate
shelter designs.

References
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Shelter design for different livestock from a climate change perspective. In: Climate change
impact on livestock: adaptation and mitigation. Springer, New Delhi, pp 399–424
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Agric Environ Sci 3(5):672–676
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Sci 86(1):226–234
Gupta LR, Hariom RS, Grewal SS (2004) Effect of different housing systems on the performance of
Murrah buffalo heifers during winter. Indian J Anim Sci 74(12):1239–1240
Hatem MH, Abdel-Moniem E, Hussein AN (2015) Effect of roofing material type on microclimate
in the sheep buildings. Agric Eng Int: CIGR J 10:52–56
Hilhorst MA, Willers HC, Groenestein CM, Monteny GJ (1998) Effective strategies to reduce
methane emissions from livestock. In: 2001 ASAE annual meeting. American Society of
Agricultural and Biological Engineers, p 1
Kamal R, Dutt T, Patel BHM, Dey A, Chandran PC, Barari SK, Bhusan B (2014) Effect of shade
materials on microclimate of crossbred calves during summer. Vet World 7(10):776–783
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Kumar R, Dogra PK, Thakur YP, Sharma KB (2012) Animal shelter. In: Impact of climate change
on cattle production and mitigations strategies in Himachal Pradesh, p 23
Melse RW, Ogink NWM, Rulkens WH (2009) Air treatment techniques for abatement of emissions
from intensive livestock production. Open Agric J 3:6–12
Mondaca MR, Cook NB (2019) Modeled construction and operating costs of different ventilation
systems for lactating dairy cows. J Dairy Sci 102(1):896–908
NAAS (2016) Policy paper 81 Climate resilient livestock production, National Academy of
Agricultural Sciences, New Delhi, p 26
Prasad A, Savanth VV, Anil KS, Saseendran PC (2013) Impact of rubber mat flooring on the
thermal comfort of dairy cattle in hot humid regions. Tamil Nadu J Vet Anim Sci 9(4):272–278
Sahu D, Mandal DK, Bhakat C, Chatterjee A, Mandal A, Mandal M (2018) Effects of roof ceiling
and flooring on microclimate of shed and physiological indices of crossbred Jersey cows. Int J
Livest Res 8(4):272–280
Sruthi S, Sasidharan M, Anil K, Harikumar S, Simon S (2019) Effect of automated intermittent
wetting and forced ventilation on the physiological parameters and milk production of Murrah
buffaloes in humid tropics. Pharm Innov J 8:315–319
Tomaszewski MA, De Haan MA, Thompson JA, Jordan ER (2005) The impact of cooling ponds in
North Central Texas on dairy farm performance. J Dairy Sci 88(6):2281–2286
Valtorta SE, Gallardo MR, Castro HC, Castelli ME (1996) Artificial shade and supplementation
effects on grazing dairy cows in Argentina. Trans ASAE 39(1):233–236
Valtorta SE, Leva PE, Gallardo MR (1997) Evaluation of different shades to improve dairy cattle
well-being in Argentina. Int J Biometeorol 41(2):65–67
Nutritional Amelioration of Thermal Stress
Impacts in Dairy Cows 12
Frank R. Dunshea, Brian J. Leury, Kristy DiGiacomo,
Jeremy J. Cottrell, and Surinder Singh Chauhan

Contents
12.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142
12.2 Physiological Impact of Heat Stress . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
12.3 Nutritional Strategies to Ameliorate Heat Stress . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
12.3.1 Macronutrient Manipulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
12.3.2 Dietary Betaine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
12.3.3 Dietary Chromium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
12.3.4 Antioxidants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
12.3.5 Other Dietary Strategies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147
12.4 Future Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148

Abstract
Heat stress (HS) has been implicated in several negative impacts on farm animal
welfare and their production. Some of the negative impacts of HS on farm
animals have been alleviated by the advances in management strategies. Despite

F. R. Dunshea (*)
School of Agriculture and Food, Faculty of Veterinary and Agricultural Sciences, The University of
Melbourne, Parkville, VIC, Australia
Faculty of Biological Sciences, The University of Leeds, Leeds, UK
e-mail: [email protected]
B. J. Leury · K. DiGiacomo · J. J. Cottrell
School of Agriculture and Food, Faculty of Veterinary and Agricultural Sciences, The University of
Melbourne, Parkville, VIC, Australia
S. S. Chauhan
Faculty of Veterinary and Agricultural Science, The University of Melbourne, Parkville, Australia

# The Author(s), under exclusive license to Springer Nature Singapore Pte 141
Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_12
142 F. R. Dunshea et al.

all the advances, HS continues to impact farm animal production during summer
heat events, particularly in dairy. Implementation of management strategies, such
as providing shade or additional water troughs, and/or nutritional strategies could
help the producers when a heat event is imminent. Some of the nutritional
strategies that could be utilised include antioxidant supplementation, dietary Cr,
betaine and altering the rate of starch fermentation, which have been
demonstrated to decrease HS under some circumstances. These nutritional
strategies are discussed in this chapter.

Keywords
Antioxidant · Dairy cattle · Heat stress · Nutritional strategies · Starch
fermentation

Abbreviations

% Percentage
Cr Chromium
CrPic Chromium picolinate
d Day
g Gram
GIT Gastro intestinal
HS Heat stress
LWT Live weight
NaOH Sodium hydroxide
NEFA Non-esterified fatty acid
pCO2 Partial pressure of carbon dioxide
pH Potential of hydrogen
pO2 Partial pressure of oxygen
PUN Plasma urea nitrogen
ROS Reactive oxygen species
SCFP Saccharomyces cerevisiae fermentation products
Se Selenium

12.1 Introduction

Heat stress (HS) negatively affects a variety of dairy production parameters

(e.g. milk yield and quality, feed intake) and is already a substantial burden that
will only worsen with climate change. While some of the negative effects of HS have
been mitigated following development and adopting management strategies such as
providing shade, water and cooling systems, but milk production still decreases
during HS. A striking decline in feed intake is a clear indication of HS and is thought
to be primarily responsible for many of the deleterious effects of HS on cow
12 Nutritional Amelioration of Thermal Stress Impacts in Dairy Cows 143

performance. Another major implication of HS is the significant reduction of milk

yield which can decrease by 40–50% in non-cooled dairy during extreme heat
events, and by 10–15% in cooled management systems. Therefore, alternative
measures such as nutritional intervention are needed to augment the other manage-
ment strategies. The nature of the nutritional strategies will be targeted at the
physiological and biochemical adaptations that occur during HS. This chapter will
briefly cover the physiological adaptations to HS and identify targeted dietary
strategies to mitigate HS.

12.2 Physiological Impact of Heat Stress

Lactating dairy cattle are especially vulnerable to HS because of the high metabolic
demand for milk synthesis and hepatic and gastrointestinal (GIT) metabolism
associated with the large feed intakes required for lactation. In dairy cows, sweating
(evaporative heat loss) is the major mode of heat loss during HS while 15% of the
heat loss is via passive evaporation (Dunshea et al. 2013). While the evaporative
efficiency increases with increasing temperature, high humidity may limit evapora-
tive heat loss (Silanikove 2000). HS also increases respiration rate as the dairy cow
attempts to dissipate heat through respiration and panting. Therefore, dietary
strategies aimed at reducing metabolic rate such as betaine or increasing blood
flow to the lungs or skin such as insulin-mimics like chromium offer the potential
to mitigate HS in dairy cows.
Despite the reduction in feed intake during HS in dairy cows, there is no change in
plasma on-esterified fatty acid (NEFA) concentrations indicating that dairy cows
don’t mobilise adipose tissue (Rhoads et al. 2009; Wheelock et al. 2010), which
typically should be the case following feed reduction in feed intake to provide
energy to support milk production. Also, the decrease in milk production and milk
lactose secretion during HS is markedly lower than in pair-fed cows maintained
under thermoneutral conditions. Plasma glucose is decreased, insulin is unchanged
or increased while plasma urea nitrogen (PUN) is increased. Also, glucose utilisation
during a glucose tolerance test increases during HS in dairy cows (Wheelock et al.
2010) while the insulin response to a glucose tolerance test decreases during HS in
non-lactating sheep (Dunshea et al. 2017). Taken together, these findings suggest
that increased insulin sensitivity may be an essential component of the acclimation
mechanism in HS. Increased insulin sensitivity would ensure that cows would
reduce adipose tissue lipid mobilisation and the use NEFA as a preferred energy
substrate despite a reduction in feed intake during HS. Instead, glucose would
become the preferred energy substrate for peripheral tissues increasing the need
for gluconeogenesis, as evidenced by increased PUN, and reducing the amount of
glucose available for lactose synthesis. Heat stress causes a reduction in blood pCO2
and increase in blood pO2 and a resultant decrease in base excess and increase in
blood pH in sheep (Gonzalez-Rivas et al. 2017). These changes in blood gas
parameters are most likely a result of increased respiration rate during HS. Use of
NEFA as a metabolic substrate during HS would result in a low respiratory quotient
144 F. R. Dunshea et al.

and would further decrease the blood base excess resulting in respiratory alkalosis.
An increase in insulin sensitivity with little change in circulating insulin would
inhibit lipolysis and fat mobilization whilst still ensuring hepatic gluconeogenesis
since the effective dose to inhibit plasma NEFA concentrations is within the
physiological range and less than that which inhibits gluconeogenesis. This suggests
that improved insulin sensitivity may reduce the impact of HS on lactating dairy
cows. Dietary supplements that may improve insulin sensitivity include
thiazolidinediones and Cr (antidiabetic compounds) and as well as the osmolyte
betaine.
One of the other implications of HS is the oxidative stress (OS) which if not
controlled can lead to oxidative damage. In dairy cows, HS has been implicated to
cause OS during transition period (Bernabucci et al. 2002) and other lactating
ruminants. Chronic exposure to HS can lead to decreased blood glutathione (reduced
form) and increase oxidised glutathione concentration, resulting in OS (Lakritz et al.
2002). Therefore, antioxidant supplementation is one of the potential nutritional
strategies to prevent oxidative damage during HS.
The redistribution of blood flow away from the body core to the periphery to
increase the radiant heat loss that occurs during HS deprives the viscera of blood
flow, causing ischemia/hypoxia leading to increased production of free radicals or
reactive oxygen species (ROS) and decreased antioxidant status (Bernabucci et al.
2002). Normally, the free radicals produced are scavenged by cellular antioxidant
systems and a balance is maintained. However, when the production of free radicals
is much faster than their neutralization by the antioxidant system, this leads to
damage of macromolecules, disruption of normal metabolism and physiology and
may ultimately lead to loss of cell function. Because of the increased environmental
temperature and paucity of green pastures during summer, the antioxidant status of
dairy cows is often low and not enough to scavenge the excessive free radicals
generated during metabolic challenges (high milk yield) and environmental
challenges such as hot summer conditions. Therefore, dietary antioxidant supple-
mentation would be another nutritional strategy to mitigate some of the negative
impacts of HS in lactating dairy cows.

12.3 Nutritional Strategies to Ameliorate Heat Stress

12.3.1 Macronutrient Manipulation

There are number of potential nutritional strategies that can be utilised to better
manage dairy cows during HS and reduce the losses. Please see review, Beede and
Collier (1986) and Dunshea et al. (2013), for more common and detailed feeding
interventions in addition to those being discussed in this chapter. Many of the dietary
interventions in lactating dairy cows are simply targeted to provide high-energy diets
as HS is known to cause negative energy balance. Thus this energy deficit can be
ameliorated by concentrate feeding or fat supplementation. For example, dietary
supplementation of saturated fatty acids has been reported to reduce rectal
12 Nutritional Amelioration of Thermal Stress Impacts in Dairy Cows 145

temperatures in dairy cattle during the hottest part of the day (Wang et al. 2010). This
reduction in rectal temperature can be explained by reduced metabolic heat produc-
tion associated with these diets and less decline in milk production during HS. Heat
load (total heat contributed by both environmental heat and metabolic heat) on cows
is likely to be increased over summer not just due to high temperatures but also
aggravated by diets which have a high increment of heat of digestion such as fibrous
and dry pasture (Silanikove 2000). Therefore, increasing the energy density of the
diet (concentrate feeding) and decreasing forage contents of the diet to decrease heat
production, are common methods to better manage cows during HS. However, it is
equally important to choose protein type and balance the diet of heat-stressed
lactating cattle, to minimise the increase metabolic heat production involved in
excess nitrogen excretion as urea, following excess protein feeding. During HS,
electrolyte and mineral balances are equally important must be carefully managed,
particularly of those macronutrients lost via sweat such as potassium. However,
cattle are more susceptible to rumen acidosis due to less bicarbonate available for
rumen buffering because of excessive salivation and panting during HS (Renaudeau
et al. 2012). Therefore, careful formulation and properly balanced diets to avoid high
grain content, are required to better manage cattle during HS. Also, as will be
mentioned later, the fermentability of the grain or concentrate can impact responses
to HS.

12.3.2 Dietary Betaine

HS is known to increase the maintenance requirements of animals during HS due to

increased respiration and panting etc. Therefore, feeding methyl donors such as
betaine (tri-methyl glycine), a naturally occurring amino acid derivative, are impor-
tant nutritional interventions to reduce the negative impacts of HS. Betaine has an
important role as osmoregulatory action and therefore can reduce energy required for
ions exchange (sodium/potassium pumping to maintain cellular osmolarity). For
example in pigs, betaine supplementation resulted in improved growth performance
which could be attributed to the reduced maintenance energy requirement of pigs
(Schrama et al. 2003). Additionally, betaine also has antioxidant properties and can
reduce the severity of some enteric infections in poultry by improving the gut
membrane integrity (Klasing et al. 2002; Shakeri et al. 2020). While there are limited
studies of betaine supplementation in ruminants as compared to monogastrics, but
there is evidence that dietary betaine can reduce HS and improve feed intake and
growth performance in beef and dairy cattle (Cronje 2005).
It is well established that milk yield increases with supplemental betaine up to
150 g/d in thermoneutral conditions (Wang et al. 2010; Peterson et al. 2012; Hall
et al. 2016; Dunshea et al. 2019). While the ability of betaine to mitigate HS has been
reported previously in sheep (DiGiacomo et al. 2016), the effects of betaine on milk
yield during summer or HS conditions are more equivocal (Zhang et al. 2014; Hall
et al. 2016). The multi-faceted dose-dependent response to betaine was anticipated
as the underlying factor (Dunshea et al. 2013, 2019). This was evident in a study by
146 F. R. Dunshea et al.

Zhang et al. (2014), where the researchers showed that milk yield increased at a dose
of 15 g/d (ca. 0.125 g/LWT0.75) during summer, beyond which no effect was
observed. A similar quadratic dose-response was observed in sheep during HS
where physiological improvements were seen at 2 g betaine/d (ca. 0.125 g/
LWT0.75) but not 4 g betaine/d (DiGiacomo et al. 2016). Similarly, Dunshea et al.
(2019) found that dietary betaine at 15 g/d increased milk yield in grazing dairy cows
during summer. In contrast, Hall et al. (2016) saw no improvements at 2 and 4 times
this dose in simulated HS conditions. The temperance in response to high doses of
betaine during HS may be that the stimulation of hepatic metabolism and consequent
increase in heat production the liver may offset the reduction in heat production due
to the osmo-protective effects of betaine (Dunshea et al. 2013).

12.3.3 Dietary Chromium

Recall, improving insulin sensitivity may help to reduce HS. Among the essential
trace elements, chromium (Cr) is known to mimic insulin action and plays a key role
in energy and protein metabolism. We (Dunshea et al. 2017; Hung et al. 2021) have
done number of studies on Cr supplementation in ruminants and pigs during HS and
have consistently seen positive effects on the physiological and production
parameters. Similarly, dairy cows supplemented with dietary CrPic during HS
have been reported to increase feed intake and milk yield (Al-Saiady et al. 2004).
More recently, supplemental chromium yeast increased feed intake but not milk
yield in lactating dairy cows exposed to HS (Shan et al. 2020). Importantly, the
physiological responses to HS such as rectal temperature and respiration rate were
reduced in those cows receiving Cr while the antioxidant status and immune function
were improved.
At least some of the action of dietary Cr may be through improving insulin action.
For example, Keshri et al. (2019) found that dietary Cr decreased plasma insulin in
dairy calves undergoing HS. Similarly, both dietary Cr and HS increased insulin
sensitivity in sheep, with the effects being additive (Dunshea et al. 2017). The
mechanism of action of Cr is underpinned by its ability to improve insulin sensitivity
during HS.

12.3.4 Antioxidants

Oxidative stress caused by HS can be managed by dietary antioxidation supplemen-

tation. However supra-nutritional levels of antioxidants are required to prevent the
oxidative damage caused by OS. For example, our studies in sheep have shown that
selenium and Vitamin E supplementation can reduce the oxidative stress and
ameliorate HS in sheep (Dunshea et al. 2017). While many studies have looked at
Se and other antioxidant supplantation in lactating dairy cows, far fewer studies have
been conducted under HS conditions. Similarly, a previous study by Calamari et al.
(2011), reported increased antioxidant systems of heat-stressed lactating dairy cows
12 Nutritional Amelioration of Thermal Stress Impacts in Dairy Cows 147

fed Se yeast. It appears that organic forms of Se are more beneficial than inorganic
forms (Sun et al. 2019). We have found that supplemental Se and Vitamin E reverses
the negative effect of HS on antioxidant status in dairy cows although milk yield was
unchanged (J. Cottrell, unpublished). We have also found that a proprietary antioxi-
dant supplement containing a mixture of vitamins, betaine and Se (OxiCare, DSM
Nutritional Products Pty Ltd) increases milk yield and time spent ruminating in
grazing dairy cows (S. Chauhan, unpublished). Further research is needed in
this area.

12.3.5 Other Dietary Strategies

Given the significant contribution of metabolic heat to the heat load on farm animals,
decreasing the heat of fermentation in ruminants can be one of the simple strategies
to reduce susceptibility to HS and improve feed intake (Russell 2007). Another
strategy to decrease the heat of fermentation could be to reduce the ruminal fermen-
tation of starch. This can be achieved by increasing the amount of starch that escapes
ruminal fermentation (Russell 2007). In Australia, wheat is the most reliable and
cheapest grain available for animal feeding which could be indirectly impacting
dairy industry over the summer because wheat has a very rapid rate of rumen
fermentation, especially when compared to maize (Dunshea et al. 2013). We have
found that maize decreased the magnitude of the increase in respiration rate ( 22%)
and rectal temperature ( 26%) as compared to wheat when sheep were fed 50%
wheat and 50% maize during thermoneutral and HS conditions (Gonzalez-Rivas
et al. 2016, 2017). This response was seen at various feed intakes from 1.3 up to 2.0
times maintenance. We also observed clear differences in flank temperature and the
difference between the right and left flank temperatures which clearly indicate the
difference in heat of fermentation between wheat and corn fed sheep, and these
findings are consistent with observations in dairy cows (Dunshea et al. 2013). Also,
treating wheat with 3% NaOH can reduce simulated rumen fermentation and the
physiological responses to HS in sheep. Alternatively, rate of rumen fermentation
can be reduced by protecting wheat from rumen fermentation. This can be achieved
by treating wheat with starch binding agents (Bioprotect™) which may also reduce
the heat increment of fermentation (Gonzalez-Rivas et al. 2018; Dunshea et al.
2013). In vitro studies have shown that a starch binding agent (Bioprotect™) can
reduce the rate of fermentation of wheat and replacing wheat with wheat treated with
the starch binding agent increased milk yield and milk fat % in dairy cows in summer
(Gonzalez-Rivas et al. 2018).
Saccharomyces cerevisiae fermentation products (SCFP) have been routinely
used to modify rumen fermentation by stimulating cellulolytic and lactate-utilising
bacteria, increasing microbial protein synthesis, and stabilising rumen pH. This in
turn can promote increased feed intake and milk production and has been particu-
larly beneficial for increasing feed intake in early lactation, likely due to a decrease in
inflammation and stress around calving in supplemented animals (Poppy et al.
2012). Optimizing rumen function may mitigate the negative effects associated
148 F. R. Dunshea et al.

with HS and benefit lactation performance by improving feed intake and decreasing
rectal temperatures. However, the impacts of SCFP on milk yield in cattle exposed to
HS are inconsistent, and further research is needed to quantify the effect of SCFP on
lactation performance and persistence (Shwartz et al. 2009).

Learning Outcomes
• The higher metabolic demand for milk synthesis and associated demand for
large feed intake in dairy cattle makes them the most vulnerable livestock
population in the changing climate.
• The essential micronutrients such as chromium, betaine and antioxidant
supplementation or other dietary strategies that alter the rate of ruminal
starch fermentation proved to be beneficial in relieving heat stress in dairy
cattle.
• The additional management strategies (housing, genetic approach, disease
management strategies) when clubbed with nutritional intervention may
provide rich dividends in terms of sustaining production during heat stress
exposure in dairy cattle.

12.4 Future Perspectives

Mitigating HS in lactating dairy cows will require on-farm adoption of multiple

strategies including management interventions such as providing shade (to protect
cows from direct heat) or additional water troughs (to help cows cool down) and
implementing nutritional strategies. Dietary supplementation of antioxidants, beta-
ine and Cr or altering the rate of starch fermentation, depending on conclusive
demonstration of benefit in lactating dairy cows, are some of the potential nutritional
strategies that could be adopted to ameliorate HS. These nutritional strategies should
be targeted towards the physiological and metabolic adaptations that occur during
HS. Further work is needed to understand the role of insulin sensitivity and oxidative
stress in the aetiology of HS. Further work should also target alternative antioxidant
strategies such as plant-derived polyphenols and antioxidants that have beneficial
effects in other species. Finally, given the severe negative consequences that HS can
elicit on dairy cattle production and welfare, research into the use of combinations of
nutritional interventions and other amelioration strategies (such as housing and
genetics) are warranted.

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The Role of Napier Grass (Pennisetum
purpureum Schumach) for Improving 13
Ruminant Production Efficiency and Human
Nutrition in the Tropics

M. R. Islam, S. C. Garcia, N. R. Sarker, B. K. Roy, N. Sultana,

and C. E. F. Clark

Contents
13.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152
13.2 Napier Grass Management, Animal Production and Human Nutrition . . . . . . . . . . . . . . . . 153
13.3 The Opportunity to Address Global Malnutrition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157
13.4 Future Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158

Abstract
Napier grass (Pennisetum purpureum Schumach) comprises up to 80% of small
holder ruminant diets across the (sub)tropics due to its high yield; however,
ruminants offered this feed typically have low productivity. The key reason for
this low productivity is the grass’ low nutritive value under current management.
Consequently, high quality milk and meat consumption in the same regions is low
with many people suffering from malnutrition, impaired cognitive development,
and many children under five are stunted. Therefore, management strategies to
improve the nutritive value of Napier grass are urgently needed to increase
ruminant productivity, increase associated income and subsequently availability
and consumption of animal sourced foods. Currently, well managed Napier grass
offered solely to cattle yield 11 L milk/cow/d and gain 0.5 kg/animal/d and has

M. R. Islam (*) · S. C. Garcia · C. E. F. Clark

Livestock Production and Welfare Group, School of Life and Environmental Sciences, Faculty of
Science, The University of Sydney, Camden, NSW, Australia
e-mail: [email protected]
N. R. Sarker
Krishi Gobeshona Foundation, Dhaka, Bangladesh
B. K. Roy · N. Sultana
Bangladesh Livestock Research Institute, Dhaka, Bangladesh

# The Author(s), under exclusive license to Springer Nature Singapore Pte 151
Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_13
152 M. R. Islam et al.

the further potential to sustain 14 L/cow/d and 1 kg liveweight gain/d in the

tropics and subtropics. This compares against more typical levels of production of
less than 5 L/cow/d from this grass. Thus, there is an immense opportunity to at
least double levels of ruminant food production through simple changes in Napier
grass management from the same land area to improve food security and reduce
malnutrition across vast tropical areas.

Keywords
Animal production · Food security · Napier grass · Nutrition · (Sub)tropical
regions

Abbreviations

$ Dollar
% Percentage
ADF Acid detergent fibre
CP Crude protein
D Day
DESA Department of Economic and Social Affairs
DM/ha Dry matter per hectare
FAO Food and Agriculture Organization
FAOSTAT Food and Agriculture Organization Corporate Statistical Database
FCM Fat corrected milk
Fig. Figure
HI Harvest interval
kg Kilogram
km2 Square kilometre
L Litre
LW Liver weight
ME Metabolizable energy
MJ Megajoule
NDF Neutral detergent fibre
UNICEF United Nations Children’s Fund
WHO World Health Organisation
yr Year

13.1 Introduction

More than half of the worlds humans and cattle are situated in the (sub)tropics. The
UNICEF, WHO, and The World Bank (World Health Organization 2018) reported
more than 815 million people in the world were hungry and chronically undernour-
ished in 2018 which equates to ~11% of the world’s 7.6 billion people. About 23%
13 The Role of Napier Grass (Pennisetum purpureum Schumach) for. . . 153

of our undernourished people live in sub-Saharan Africa, 12% in South Asia and 6%
in Latin America of the sub (tropics) (Concern Worldwide 2014). In addition,
150–151 million children (over 22% of total children) under 5 years old are stunted
(low height for age), with half of these children living in Asia and over a third living
in Africa. Also, ~51 million children (8% of total children) are wasted (low weight
for height) and two-thirds of all these children live in Asia (10% of all children), a
quarter in Africa and 1.3% in Latin America. Furthermore, ~38 million babies are
born with iodine deficiency and ~250 million preschool children are vitamin A
deficient (FAO et al. 2017; FAO 2018a). The highest prevalence of undernourish-
ment in the world occurs in the sub-Saharan region (23%) followed by South Asia
(includes India, Pakistan and Bangladesh; 11.5%) and South America (5.6%) (FAO
2017). Geographically, these countries are all located in the tropics and subtropics.
Those who live in these regions are typically low to middle income earners, with
associated poor food and nutrition security and many live on less than $2/person/day
(Table 13.1; FAO 2017). Consumption of animal protein in these regions is as low as
10 kg meat/person/year, ten times less than the 110 kg/person/year consumed in
Australia, European and North American countries (Ritchie and Roser 2019; https://
ourworldindata.org/meat-production). Thus, almost all stunting, wasting, vitamin A
and iodine deficiencies in children under five are due to the lack of animal sourced
foods (UNICEF, WHO, and The World Bank 2018; Table 13.1). Importantly, the
ruminants in these regions that supply such animal sourced food primarily consume
Napier grass. Here, we investigate the current situation of production of milk and
meat using Napier grass and its best management practice to improve yield and
nutritive quality aiding nutrition security, ending malnutrition, stunting and the
wasting of children in these regions.

13.2 Napier Grass Management, Animal Production and Human

Nutrition

Napier grass is the main forage source in tropical and subtropical regions,
compromising up to 80% of the forage base used by the smallholder dairy farmers
(Kabirizi et al. 2017). Thus, Napier grass is also known as ‘small holders grass’ or
‘poor man’s grass’ as it is widely grown and used by those with low incomes. Napier
grass is typically harvested when the plant grows to between 2 and 3.5 m high
(Zhang et al. 2010; Rengsirikul et al. 2013) to maximise forage yield (10–86 t DM/
ha; Cook et al. 2005; Vicente-Chandler et al. 1959) and feed livestock from typically
limited (small) areas of land. This robust growth to a late maturity ensures high fibre,
and low protein and energy content (Gwayumba et al. 2002; Muinga et al. 1992)
with cows offered Napier grass alone producing 8.1 L/cow/d (7.4 L at 4% FCM) but
losing ~500 g liveweight/cow/d (Islam et al. 2021; unpublished results). In contrast,
average CP and ME content of ryegrass, which is the main grass in the temperate
regions are >18% and >10 MJ ME/kg DM respectively which enables the produc-
tion of 20 L milk from 600 kg dairy cows (Fulkerson 2007). Thus, differences in
management are causing large disparities between areas where cattle are located and
154 M. R. Islam et al.

Table 13.1 Characteristics of tropical and subtropical countries in the world

Tropical and
subtropical Rest of the world
Area (km2)a 46,457,647 (36.5%)b 80,885,573
(63.5%)c
Human (in billion)d 4.0 (52%) 3.7 (48%)
Cattle (in million)e 900 (61%) 568 (39%)
Milk production share (%)f 49% 51%
Income ($) 996–3895g 3896 to >12,056h
Poverty (million)i 767 n/a
Hunger (million)j 815–821k >672 (obese)l
Undernourished (million)j 815m n/a
Stunted (million, children with low height for 150–151n n/a
age)j
Wasted (million, children with low weight for 50o 38p
height)j
Iodine deficiency (million)j 38q n/a
a
Land area, FAO (2018b)
b
Areas include South Asia, Australia, Latin America and Caribbean
c
Temperate, Mediterranean, Middle East, Saharan, cold, equators
d
WHO (2017)
e
World Cattle Inventory (2018)
f
Include Asia (29%; India and Pakistan’s share 23% of the), Africa (5%), South America (10%),
Oceania (5%) for tropical and subtropical. Include EU & USA 35%, China & Japan 8% of the
temperate share. The World Dairy Situation (2016)
g
Low to lower middle income ($996–3895) (upper middle in south American & Caribbean; $3896–
12,055)
h
Upper middle to high income
i
9767 million people living below poverty line <$1.90/person/day
j
World Hunger and Poverty Facts and Statistics (2018)
j, k
815–821 (10.7%; almost all hungry people live in lower to middle income countries)
j, l
>672 million adults are obese mostly in these countries
j, m
815 million people (10.7% of world 7.6 billion people) 23% people in sub-Saharan Africa;
11.5% in South Asia; 5.6% in Latin America
j, n
150–151 million children under 5 globally (over 22%); 1/2 of all in Asia and 1/3rd in Africa
j, o
50 million children globally (7.5% of total children); 2/3rd of all in Asia (9.7%) and 1/4th in
Africa and 1.3% in Latin America
j, p
38 million under five are overweight mostly in these countries
j, q
38 million babies born with Iodine deficiency especially in Asia and Africa

their production (Table 13.2). In this regard, FAOSTAT (2018) reported 77% of the
word’s dairy cattle to be located in Africa and Asia but produce only 33% of the
world’s milk production. Therefore, improvements in feed quality are critical to the
success of the smallholder dairy sector.
Data on animal production performance from a sole Napier grass diets are limited.
Available data indicate harvest interval (HI) plays an important role on nutritive
value of this grass and subsequently milk yield of cows with Napier grass harvested
from 42 to 91 days HI containing 6–8% CP and 7.0–7.6 MJ ME/kg DM and yielding
13 The Role of Napier Grass (Pennisetum purpureum Schumach) for. . . 155

Table 13.2 Production of cattle meat and milk by region in 2017 (Source FAOSTAT 2018) in
relation to world population
% cattle meat % whole fresh cow milk % people in the worlda
Asia 23 30 60
Africa 10 5 16
Oceania 4 5 1
Europe 16 33 10
Americas 47 27 14b
Tropical and subtropical regions are mainly in Asia, Africa and in part of Americas
a
World population by continent in 2019 (https://en.wikipedia.org/wiki/List_of_continents_by_
population)
b
South America, 6%

4.0–7.2 L milk/cow/d (Table 13.3). However, when this grass was harvested
between 28 and 30 days, it contained 7–14% CP and 7.1–7.9 MJ ME/kg DM and
yielded 7.8–11.4 L milk/cow/d (Table 13.3). Aroeira et al. (1999) in an experiment
conducted over 2 years in Brazil demonstrated Napier grass containing 11–14% CP
and 7.9 MJ ME/kg DM, grazed by Holstein Friesian x Zebu cows at 30 days HI can
yield on average 11.4 L/cow/d. However, this yield was achieved through the
supplementation of 0.9–1.4 kg concentrate during winter and early spring, although
no supplementation was required during summer and autumn. The available data,
although limited, further indicate for each per cent increase in Napier grass CP (%)
from 6.7 to 12.5 and each mega joule increase in ME (MJ/kg DM) from 7.0 to 7.9
corresponds to a milk yield increase of 0.83 and 3.5 L/cow/d, respectively
(Fig. 13.1). In line with milk yield, the growth of cattle increases alongside Napier
grass quality. Kariuki et al. (1998, 1999) offered Napier grass containing 12% CP
and 8.6 MJ ME/kg DM to steer or heifers and reported 390–500 g liveweight gain
per cattle/day (Table 13.3). Further, Napier grass quality (Sileshi et al. 1996;
Goorahoo et al. 2005) can be maintained at similar levels to that of Kikuyu grass
(Fulkerson et al. 2006; Fariña et al. 2011; García et al. 2014) or ryegrass (Fulkerson
2007). Goorahoo et al. (2005) in California for a high input production system
showed Napier grass to contain high protein (17–25% CP) and relatively low fibre
(49–53% NDF, neutral detergent fibre; 30–35% ADF; acid detergent fibre) as a HI
between 14 and 49 days. Similarly, Sileshi et al. (1996) in Ethiopia with three
cultivars showed Napier grass to contain high protein (22% CP), low fibre (53%
NDF, 29% ADF) and high energy (ME 11.6 MJ/kg DM) when harvested at 28 days
(i.e. 13 harvests/yr). In this regard, a sole Napier grass diet could support 24 L milk/
cow/d (Islam et al. 2021; unpublished results) provided intake is maintained at
>20 kg DM/cow/d. However, such intake levels would be limited by the high
NDF levels of Napier grass, but the supplementation of ~2 kg concentrate could
enable such levels to be achieved Sileshi et al. (1996) and Goorahoo et al. (2005).
Nonetheless, research using Kikuyu and Rye grass suggest that it may be possible to
harvest 14–20 L milk/cow/d when Napier grass is managed following principles of
Sileshi et al. (1996) and Goorahoo et al. (2005).
 156

Table 13.3 Current cattle production from Napier grass and potential to increase cattle production following examples of Kikuyu grass
Grass ME LW loss
Trial harvest MJ/kg Live wt DMI Milk yield or gain
days interval CP (%) DM Cow type (LW) (kg) (kg/cow/d) (L/d/cow) (g/cow/d)
Dairy
Muinga et al. (1992) 50 6.4–6.7 4–6
4 to
23
Muinga et al. (1992) 98 42 5.6 7.0 Ayr/Brown 430 6.8 6.9
490
SxShahiwal
Gwayumba et al. 37 70–91 6.1–7.2 7.6 Friesian 400 8.4–9.1 6.1–7.2
530 to
(2002)
890
Shem et al. (2003) 56 Flowera 7.8 7.1 AyrxFriesian 323 7.8
87
Muinga et al. (1992) 98 28 7.2 7.6 Ayr/Brown 430 9.3 8.6
165
SxShahiwal
Anindo and Potter 28 8.6 (7.5– Friesian 424–441 10.5
(1986) 10.2)
Aroeira et al. (1999) 730 30 11.2–13.7 7.9 HFxZebu 483 12.9 11.4 + 450 to +500
Fulkerson et al. 20.0 10 Holstein 13.0 14.2
(2007)b
Fariña et al. (2011)c 730 20–24 19.0–20.5 10.2 Holstein 613 20.4 25.1
Beef
Kariuki et al. (1999) 120 56 11.7 Friesian or 163–181 5.6–6.0 +390 to +420
Shahiwal
Kariuki et al. (1998) 104 11.8 8.6 Holstein heifers 144 5.0 +500
a
Harvested at flowering stage
b
With kikuyu grass (Pennisetum clandestinum)
c
With kikuyu in spring-summer, ryegrass (Lolium multiflorum) pasture in autumn-winter in 65% of land and an annual rotation of maize (Zea mays), brassica
(Brassica napus) and a legume either Persian clover, Trifolium resupinatum L or maple pea (Pisum sativum) in 35% land to support pasture during scarcity. The
system supported 26 t DM/ha/yr to produce 27,835 L milk/ha/yr with forages and 35,000 L/ha/yr when supported with 3 kg grain per cow/d
M. R. Islam et al.
13 The Role of Napier Grass (Pennisetum purpureum Schumach) for. . . 157

12 12
Milk yield, L/d/cow 10 10

Milk yield, L/d/cow

8

6 6

4 4

2 2

0 0
0 5 10 15 6.5 7.0 7.5 8.0
Crude protein, CP% Metabolisable energy (ME,
(MJ/kg DM)

Fig. 13.1 Impact of increasing protein (Milk yield, L/d/cow ¼ 0.83  CP% of grass + 1.63,
R2 ¼ 0.69) and energy (Milk yield, L/d/cow ¼ 3.49  ME of grass + 17.69, R2 ¼ 0.48) on milk
yield of cows using Napier grass (Source: Table 13.3)

Therefore, simple changes in Napier grass defoliation interval are key to improv-
ing the quality of Napier grass and in so doing, improve the production of animal
protein and the nutrition of people in the (sub) tropics. The dedicated demonstration
of such Napier grass management practices in a systems context as part of an
adoption strategy for such regions is thus urgently required. The FAO (2019)
reported that global livestock product demand was expected to increase by 70%
by 2050, and Adesogan et al. (2020) reported that livestock and animal sourced
foods play a critical role in improving livelihood, reducing poverty, increasing food
security, improving health and nutrition and gender equity and are vital to
sustainability.

13.3 The Opportunity to Address Global Malnutrition

The World Bank targeted (Target 2) to end all forms of malnutrition by 2030
including stunting and wasting in children under 5 by 2025. Nutrition is central to
these agendas as good nutrition can lay the foundation to achieve these targets. This
report also stressed the importance of good nutrition to ending malnutrition, stunting
and wasting whilst also enabling healthy lives, a quality education, gender equality,
economic growth and for ending poverty. In addition, good nutrition lays the
foundation for sustainable development and drives the changes needed for a more
sustainable and prosperous future. Adesogan et al. (2020) reported livestock and
animal sourced food are vital to sustainability for their critical role in improving
nutrition, livelihoods, health and gender equity, reducing poverty and increasing
food security. WHO (2014) described animal source foods as the best source of high
quality nutrient rich food for children aged 6–23 months and these livestock can play
a central role in supplying highly nutritious food such as meat and milk. Thus, we
propose that the target to end malnutrition, stunting and wasting can be tackled by
158 M. R. Islam et al.

increasing the efficiency of livestock production through simple changes in the

management of Napier grass as provided here.

13.4 Future Perspectives

The ‘Green revolution’ prioritised plant calories over protein to save millions of lives
in Asia and other (sub) tropical continents but in so doing, marginalised ruminant
production. ‘Grain’ was viewed as food, but meat or milk were used as buffers for
times of shortage. However, with increased income per capita, the consumption of
animal protein has increased with ruminants now a key food resource, supported by
tropical feeds such as Napier grass. This trend towards ruminant protein production
has generally doubled farmer income compared to grain production on the same area
of land. Despite this, the management of tropical feeds such as Napier grass could be
markedly improved to enable vastly greater levels of animal production. Such
research and adoption programmes on improved management of Napier grass are
urgently required to support the World Bank program vision to end malnutrition and
associated stunting and wasting by 2025–2030.

Learning Outcomes
• Animal sourced foods, milk and meat plays a pivotal role in providing best
nutrition to the children and because of which nutrition of ruminant live-
stock finds its importance.
• Napier grass, occupies 80% of the small holder farmer’s forage in (sub)
tropics and due to its low nutritive value, it causes low productivity in
ruminants.
• Imperative management strategies such as changes in Napier grass’s defo-
liation interval is of key importance to improve its quality and thereby
increasing the animal’s protein and in turn the nutrition of the (sub) tropics
people.

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Strategies to Ameliorate Heat Stress
Impacts in Sheep 14
Kristy DiGiacomo, Surinder Singh Chauhan, Frank R. Dunshea,
and Brian J. Leury

Contents
14.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162
14.2 Management and Housing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 164
14.3 Nutrition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 164
14.3.1 Water . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165
14.3.2 Nutritional Supplements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165
14.4 Breeding/Selection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 170
14.5 Precision Feeding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 170
14.6 Future Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 172

Abstract
Sheep are continually exposed to fluctuating environmental temperatures and
employ multiple behavioural, physiological and cellular mechanisms to maintain
a stable core temperature during temperature extremes. Heat stress events during
gestation can also impact the health and physiology of the offspring via prenatal

K. DiGiacomo (*) · B. J. Leury

School of Agriculture and Food, Faculty of Veterinary and Agricultural Sciences, The University of
Melbourne, Parkville, VIC, Australia
e-mail: [email protected]
S. S. Chauhan
Faculty of Veterinary and Agricultural Science, The University of Melbourne, Parkville, Australia
F. R. Dunshea
School of Agriculture and Food, Faculty of Veterinary and Agricultural Sciences, The University of
Melbourne, Parkville, VIC, Australia
Faculty of Biological Sciences, The University of Leeds, Leeds, UK

# The Author(s), under exclusive license to Springer Nature Singapore Pte 161
Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_14
162 K. DiGiacomo et al.

programming. The clinical responses to heat stress in ruminants include, to a

degree, a manifestation of the general stress response. Heat stress can also
increase energy requirements due to the processes of sweating, respiration and
hormone and cellular protection employed to cope with increased core
temperatures in response to heat stress. While there are many studies exploring
the physiological and productivity effects of heat stress, less is known about the
underlying metabolic and cellular responses and adaptations to heat. Previous
research in ruminants, including sheep, has shown that heat stress alters intake as
well as nutrient and energy metabolism. Although ruminants often respond to
heat exposure by decreasing voluntary feed intake, heat stress increases mainte-
nance energy requirements (due to panting, sweating and the production of
hormones), which can in turn lead to production losses and animal health issues.
This chapter will examine some of the key nutritional strategies that can be
employed to reduce the negative impacts of heat stress on health and productivity
in sheep.

Keywords
Climate change · Heat stress · Feed additives · Nutritional supplements · Sheep

Abbreviations

% Percentage
DMI Dry matter intake
g Gram
GPS Global positioning system
HSP Heat shock protein
kg Kilogram
NaOH Sodium hydroxide
NEFA Non-esterified fatty acid
NRC National Research Council
pH Potential of hydrogen
RFID Radio frequency identification
ROS Reactive oxygen species
SCFP Saccharomyces cerevisiae fermentation products
VFA Volatile fatty acid

14.1 Introduction

The impact of climate change on small ruminants is multifaceted, but includes

changes to water availability and quality, land access and soil quality, forage (and
feed) growth and quality, changes to weather events, increased ambient temperature
14 Strategies to Ameliorate Heat Stress Impacts in Sheep 163

extremes and number of extreme heat events; all of which will have direct effects on
animal physiology, health and productivity.
Heat stress is caused by a combination of environmental factors including
ambient temperature, solar radiation, humidity and wind properties and can impact
upon animals in a direct manner by reducing feed intake, altering production traits,
negatively affecting reproduction, decreasing disease resistance and consequently
affecting the overall efficiency and health of the animal. While sheep are more
tolerant to temperature variations than other ruminants, the negative impact of heat
stress on sheep productivity are well documented.
The effects of climate change on environments (and hence animals) will differ by
region, and it is anticipated that along with an overall rise in global temperatures,
there will also be a shift in rainfall patterns and thus water availability for plant and
animal growth. This will impact all aspects of ruminant production, from the regions
in which they can be grown, the species and breeds that are utilized, the types of feed
that can be grown and accessed and the housing requirements for the animals. The
combination of changes to the available forage type and quality and the reduction in
intake and changes in nutrient utilization and partitioning changes that occur in
response to heat stress in sheep highlight the important role that nutrition and
supplementation will play in ensuring that sheep production can continue in
response to climate change. It is also clear that nutritional interventions that can
improve water balance in ruminants will be highly valued in response to climate
change.
Ruminant livestock are essential for the provision of high-quality protein and
other essential nutrients (meat and milk) and fibre for human consumption and
manufacturing use. By converting ligno-cellulose plant matter to nutrients that can
be digested by humans, ruminants occupy a valuable niche and it is unlikely humans
would cease to produce ruminant products in response to climate change, although
production practices may require adaptation and change. Small ruminants provide
food security for lower socio-economic populations, particularly in underdeveloped
nations. Thus, producers are seeking methods that can be utilized to maintain, and
perhaps even improve, the productivity of sheep in warmer climates. Numerous
reviews, including those from authors of this chapter (Dunshea et al. 2017; Joy et al.
2020; Zhang et al. 2020), have outlined some nutritional supplements that have been
successfully employed to improve production in heat stressed ruminants. This
chapter will briefly outline some of these key supplements and interventions. As
other chapters in this book have covered the effect of heat stress on sheep behaviour,
welfare, meat quality, immune system, monitoring of heat stress, breed differences
and genetic selection, shelter management, rumen microbial adaptation and impacts
on reproduction these topics will not be covered in depth here.
164 K. DiGiacomo et al.

14.2 Management and Housing

While modifying the environment around animals is a faster alternative to

employing breeding or selection strategies, environmental alterations are expensive
and thus not always economically viable for producers. A successful method to
alleviate heat stress is the provision of shade. Shaded dairy cattle have lower rectal
temperatures and respiration rates and increased milk yields than those not given
access to shade (Collier et al. 2006). However, the provision of shade does not
always lead to reduced core temperatures as the usage of the shade will vary amongst
individuals. Sheep that use shade have similar rectal temperatures to that of animals
that remain out of shade (Johnson 1991) and shaded and unshaded sheep spend
comparable amounts of time grazing, drinking and ruminating (Johnson and Strack
1992). The construction of shade must be carefully designed to account for the
environmental conditions likely to be experienced by the animals, as well as the
breed of animal being cooled. For example, shade must be purposely orientated,
adequately insulated and must be large enough to allow sufficient space per head to
not increase the radiative heat load upon the animal. As trees can both protect from
sunlight and act as a heat sink by removing heat from the air, shade provided by tress
are the most effective form but requires forward planning and is not always possible
due to paddock or enclosure designs. In more humid regions the provision of shade
is less important as the effects of wind speed and humidity are more damaging than
that of solar radiation. Although shade provision may not always lead to improved
production and decreased core temperature, shade does reduce the mortality rate of
animals in higher environmental temperatures. The cost of construction use and
maintenance of these modifications must be accounted for to ensure that its use is
economically viable. A simpler and potentially more cost-effective method of
improving animal production during thermal stress is the modification of the diet
and/or the provision of nutritional supplements during stress events.

14.3 Nutrition

The heat increment of feeding describes the heat produced internally when an animal
digests feed and is influenced by the type and quality of feed consumed, as well as
the specific rumen environment of the animal. This form of heat gain is of particular
importance in ruminants due to the fermentation processes of the gut. Often summer
pastures are fibrous and dry and thus have a high heat increment of feeding, adding
to the heat load on the animal (Silanikove 2000). Excess protein in the diet of
ruminants can further exasperate the heat increment of feeding, as the energy
required to excrete excess nitrogen as urea increases the internal heat load (metabolic
heat) (NRC 2007). Fat and protein supplementation was shown to increase the
respiration rate, rectal temperature and heart rate of sheep in a meta-analysis, while
concentrate and non-fermentable protein supplementation reduced rectal
temperatures (Slimen et al. 2019). Additionally, concentrate supplementation will
increase the ratio of propionate to acetate in the rumen, and propionate produces a
14 Strategies to Ameliorate Heat Stress Impacts in Sheep 165

lesser heat increment of feeding (accounting for fermentation and nutrient metabo-
lism) than acetate. The mixture of feeds and ration balance will likely need to be
adjusted to suit what is available in a climate-altered world. Research is required to
examine novel feeds and feed combinations to ensure that animal production
efficiency can be maintained in a changing climate. For example, producers may
need to adapt to grow crops native to their specific region to ensure fodder can thrive
in the climate and soil type etc. Native non-traditional feeds have received limited
research to date, but certainly warrant further attention globally.

14.3.1 Water

Water is a vital component for health and often called the ‘forgotten nutrient’.
Despite the clear importance of animals maintaining adequate hydration, especially
during heat stress, producers often do not consider water access or quality when
designing animal rations. Although some breeds of sheep, particularly those native
to tropical regions, can be well adapted to water deprivation and as ruminant animals
can quickly rehydrate after a bout of dehydration as the rumen can store water and
release it slowly, water remains an essential ingredient for maintaining adequate
production. Moreover, water requirements are increased in growing, pregnant and
lactating animals. Increased ambient temperatures will impact an animal’s water use
by decreasing DMI and thus decreasing water gained from feed (preformed water),
and by increasing water use via the evaporative cooling mechanisms employed by
the animal to dissipate excess heat. Breed differences exist in water intake in sheep.
For example, sheep from mountain and desert conditions have increased water
turnover than that of other breeds (Marai et al. 2007). As reviewed by Naqvi et al.
(2017), the effects of climate change on water availability and quality will impact
sheep physiology from metabolism to endocrine function and reproduction. Inter-
mittent water supplies and fluctuations in ambient temperature are likely to lead to a
plethora of impacts on water balance in sheep, including increased consumption of
saline water, consumption of contaminated or toxic water (via contamination caused
by flooding and other natural events) and decreased overall access to water. It is also
likely that changes to weather and precipitation patterns will increase animals’
exposure to mycotoxins and contaminated feed as wet/warm climates can promote
toxin contamination and growth.

14.3.2 Nutritional Supplements

The provision of nutritional supplements to ameliorate the effects of heat stress is

potentially the fastest and easiest method for producers to adopt, particularly for
feedlots and dairy animals. Nutritional supplements include additives such as amino
acids, vitamins, minerals, osmolytes, buffers and electrolytes. Supplements can
improve both animal performance and also animal health, for example by improving
the immune system and reducing oxidative stress (Chauhan et al. 2021). Additives
166 K. DiGiacomo et al.

such as chromium, glycine and betaine have been examined with varying degrees of
success. If successful and cost effective, supplementation provides a management
strategy that has the potential to be widely adopted by producers, and hence has great
potential to improve production during times of heat exposure. Nevertheless, sup-
plementation must be met with careful consideration as some negative responses to
the dietary modification may occur. A recent meta-analysis of the nutritional
supplements used to alleviate heat stress in sheep (66 studies examining 38 different
heat stress alleviating strategies) found that some supplements actually increased the
rectal temperature of sheep (such as concentrates and Saccharomyces Cerevisiae)
whereas others (such as seaweed and a combination of antioxidants and minerals)
successfully reduced rectal temperature to that of or below that seen in thermoneutral
sheep (Slimen et al. 2019). Thus, the determination of any naturally occurring and
organic compounds that can ameliorate the effects of heat stress is highly desirable.
There are a variety of nutritional strategies and supplements that can be utilized to
maintain or improve production in sheep. While this chapter cannot adequately
describe all published data on currently available interventions, some of the com-
monly utilized nutritional interventions used for sheep will be briefly described here.

14.3.2.1 Betaine
Betaine, also known as trimethylglycine and glycine betaine, is a non-toxic amino
acid found widely in nature. Mammals utilize betaine for two major functions, as a
methyl donor participating in protein and lipid metabolism, and when not
catabolised, betaine is used as an organic osmoprotectant. Betaine helps to maintain
cellular integrity during times of osmotic stress thus sparing energy by reducing the
need for cellular ion-pumps. During sustained periods of water loss organic
osmolytes, such as betaine, are employed to maintain cellular water balance. Inor-
ganic ions like urea cannot be tolerated for long periods by the cell as they disrupt
protein function, while organic osmolytes elicit no such effects, and thus organic
osmolytes are employed when long-term osmolytes are required. Moreover, protein
synthesis is reduced in cells during times of osmotic stress, and the accumulation of
organic ions can permit cells to recover and improve protein synthesis. Additionally,
betaine is shown to have anti-apoptotic properties that can promote cell proliferation,
which is of particular importance for the maintenance of gut tissue integrity in heat
stressed animals (Eklund et al. 2005). In heat stressed sheep betaine (2 g/head/day)
reduced rectal and skin temperatures, respiration rates and plasma NEFA
concentrations, although a higher dose (4 g/head/day) either had no effect or an
additive effect such that responses to heat exposure were increased (DiGiacomo
et al. 2016). In lactating cattle, betaine supplementation improved milk yield and
quality, feed intake and osmotic balance (Dunshea et al. 2019; Shah et al. 2020).
While studies in heat stressed meat-sheep have not been published, betaine supple-
mentation increased feed intake and average daily gain and improved some meat
quality measures with particularly favourable responses observed in response to
rumen protected betaine (Dong et al. 2020). However, Fernández et al. (1998)
demonstrated no improvement in sheep liveweight or muscle area but did show a
reduction in fat thickness when supplemented with 2 g/kg betaine.
14 Strategies to Ameliorate Heat Stress Impacts in Sheep 167

14.3.2.2 Amino Acids

Heat stress can lead to a negative nitrogen balance due to changes in metabolism and
reductions to feed intake. Excess dietary protein can exacerbate the impact of heat
stress and further decrease DMI and production (such as milk yield) in ruminants
(West 1999). Supplementations of specific amino acids are being increasingly
investigated in ruminant models. While the published data to date is in low animal
numbers, there is evidence that supplementation of rumen protected choline can
improve milk yield in dairy cattle (Holdorf and White 2021). In goats, supplementa-
tion of rumen protected choline during the summer period increased feed intake,
liveweight gain and feed conversion efficiency (Habeeb et al. 2017). Research into
feeding amino acids and combinations of amino acids to heat stressed sheep is
required to determine if such responses are also observed in other small ruminants.

14.3.2.3 Fibre and Concentrates

As acetate is used as by peripheral tissues for the de novo synthesis of long chain
fatty acids and cholesterol (NRC 2007), acetate metabolism produces more heat than
propionate. Thus, reducing the fibre intake of sheep can reduce the heat increment of
feeding (West 1999) and decrease the animals’ heat load. Although in some cases
(depending on the forage type) high roughage diets can decrease body temperature
as the overall level of intake, and thus rate of digestion, is decreased. Furthermore,
increased water intake during heat stress can dilute feed volume and further impact
intake and digestibility. Evidence also shows that VFA production is decreased
during heat stress, even when animals are force fed via a cannula, which highlights
that changes other than just level of intake are contributing to the change in VFA
production and likely relate to rumen microbiome changes. As heat (and other)
stressors can reduce intake, more energy dense diets are often provided to meet
energy requirements. This can include the addition of short chain fatty acids or fats,
which have a low metabolic heat production. However, excess fat supplementation
can potentially disrupt the rumen microbiome which can further decrease feed
intake. The timing of concentrate feeding may also be important and feeding
concentrates at night when ambient temperatures are lower may improve feed intake
and energy balance.

14.3.2.4 Grain Feeding

Feed fermentation and the heat increment of feeding are major contributors towards
the heat load of sheep. High carbohydrate grain feeds (high in rapidly fermentable
starch) have a reduced heat increment from fermentation compared to forages and
roughages, but the rapid rate at which they are fermented can increase heat loads.
Thus, starch sources must be carefully considered before being fed to heat stressed
ruminants. For example, wheat is rapidly fermented and produces a greater physio-
logical response to heat (greater heat load) in sheep compared to corn (Gonzalez-
Rivas et al. 2016). Modification of starch to slow the rate of fermentation may
therefore reduce heat loads and improve production in heat stressed sheep. A starch
binding agent (Bioprotect®) was able to reduce the heat increment of feeding wheat
grain to match that produced when feeding corn in sheep (Prathap et al. 2021). In
168 K. DiGiacomo et al.

addition, feeding NaOH treated wheat to sheep reduced respiration rates compared to
untreated wheat diets with no negative impact on overall starch digestibility
(Gonzalez-Rivas et al. 2017, 2021). Research into novel starch sources and starch
modification techniques that slow the rate of digestion without negatively impacting
digestibility or energy content is therefore warranted.

14.3.2.5 Electrolytes
Reduced intake can lead to decreased mineral consumption, while increased water
consumption can alter mineral excretion and changes to metabolism associated with
heat stress can alter mineral metabolism. For example, absorption of calcium and
potassium are reduced by heat. Thus, mineral supplementation is likely to assist in
the maintenance of health and production in heat stressed animals. In sheep, mineral
supplementation combined with antioxidants (zinc, cobalt, chromium, selenium and
vitamin E) ameliorated the negative impact of heat stress on feed intake and reduced
respiration rate and rectal temperatures (Sejian et al. 2014).

14.3.2.6 Essential Oils

While experimental studies examining the feeding of essential oils to heat stressed
ruminants are scarce, as essential oils commonly have antimicrobial and antiparasitic
properties it is hypothesised that they will improve animal performance and health
during times of heat stress. Further, there is potential that essential oils can reduce
methane production although the emissions associated with oil extraction techniques
might negate any such gains. In lactating sheep in Greece, a combination of cornus 
thyme oil supplementation improved milk yield and milk fat % (Kalaitsidis et al.
2021). Plantain, which contains bioactive components that have free radical scav-
enging properties, reduced feed intake, rectal temperatures and plasma glucose
utilization in response to exogenous insulin infusion in heat stressed sheep
(Al-Mamun et al. 2017). Research into novel supplements and native feeds are
predicted to elucidate further essential oils that may be useful for heat stress
abatement and even rumen modification to reduce methane production. However,
current industrial extraction methods used to isolate the oils are harmful to the
environment and technological advancements in extraction methods are required if
extracted oils are to be fed. Future research might therefore also examine the feeding
of whole fresh or dried herbs, rather than extracts.

14.3.2.7 Antioxidants
Heat stress can induce oxidative stress which contributes to the associated produc-
tion losses. Antioxidants such as selenium and vitamin E are able to act as
scavengers for reactive oxygen species (ROS) and can maintain redox balance,
protect membrane integrity, modulate the immune response, alleviate leaky gut
and improve heat shock protein (HSP) expression in stressed animals (Chauhan
et al. 2016). In heat stressed sheep, supranutritional doses of selenium and vitamin E
reduced respiration rates, rectal temperatures, plasma ROS and the oxidative stress
index which resulted in supplemented sheep maintaining feed intake under heat
stress compared to non-supplemented sheep (Chauhan et al. 2014). According to the
14 Strategies to Ameliorate Heat Stress Impacts in Sheep 169

recent meta-analysis, the feeding of a combination of antioxidants and minerals

produced the greatest reduction to rectal temperature in heat stressed sheep (Slimen
et al. 2019).

14.3.2.8 Minerals
Chromium plays an important role in the metabolism of fat, carbohydrate and protein
and is also involved in glucose metabolism. Chromium has demonstrated beneficial
physiological and production responses in sheep exposed to various stressors,
including heat. Chromium can improve energy utilization, augment the actions of
insulin (Gardner et al. 1998), reduce the hyperinsulinemic effects of heat and may
enhance skin micro-circulation and vasodilation to enhance heat loss from the skin.
Nano-chromium picolinate reduced rectal temperatures and respiration rates and
improved average daily feed intake (Hung et al. 2021) and improved insulin
sensitivity (Dunshea et al. 2013) in heat stressed sheep. While further research is
required to confirm appropriate forms and dose rates of chromium, and to compare
responses in different sheep breeds and physiological states (i.e. lactating and
growing animals), currently available evidence suggests chromium supplementation
has beneficial impacts on thermal tolerance and production in sheep.
Zinc supplementation can improve intestinal barrier function in animals, and as
heat stress can damage gastric and intestinal tissues in response to blood flow
redistribution to the extremities, zinc supplementation may improve animal health
and performance under heat stress conditions. In steers, a zinc and amino acid
complex supplement reduced rectal temperature and improved some blood gas
markers (Opgenorth et al. 2021).
A combination of supplements may be beneficial in combating the negative
effects of heat stress by targeting the different metabolic and physiological pathways
impacted. For example, Sejian et al. (2014) observed an improvement to feed intake
and a reduced respiration rate and rectal temperature in heat stressed sheep fed a
combination of minerals and antioxidants (zinc, cobalt, chromium, selenium,
vitamin E).

14.3.2.9 Saccharomyces Cerevisiae Fermentation Products (SCFP)

Saccharomyces cerevisiae fermentation products (SCFP, yeast) supplements have
been utilised in cattle to improve rumen stability and maintain milk production in
lactating animals. While there are no published experiments examining SCFP in heat
stressed sheep, one experiment in dairy cattle demonstrated improved milk yield and
feed efficiency (Zhu et al. 2016). In a meta-analysis of the impacts of SCFP in
ruminants (cattle and sheep), SCFP overall increased DMI, increased rumen pH and
VFA concentrations, tended to decrease rumen lactic acid concentration, increased
milk yield and tended to increase milk fat (with no impact on milk protein
concentrations) (Desnoyers et al. 2009). As heat stress can disrupt rumen stability,
it is likely that SCFP supplementation can improve health and production in
ruminants and thus warrants further research in small and large ruminant species.
170 K. DiGiacomo et al.

14.4 Breeding/Selection

Breeding animals to improve production and efficiency is the aim of most producers,
as animals that are better able to convert feed to product are more cost effective.
Additionally, animals that are better able to cope with the external environment will
have improved production output as they dissipate less energy maintaining thermal
homeostasis. Nevertheless, it is generally concluded that animals bred to embody
better thermoregulatory control and are thus better able to cope with extreme
environmental temperatures will offset this coping mechanism with a decline in
overall production. Research into breeding and selection programs, particularly
those that might employ the use of native and adapted breeds of sheep, are
continuing globally. In the future researchers and producers will need to work
together across regions and countries to ensure that breeding programs are utilizing
the most appropriate technology and genetic material to maintain production in a
climate changed world.

14.5 Precision Feeding

Technology developments in agriculture are required to ensure that production can

continue in a changing climate, particularly those that can assist in resource man-
agement (feed, land and water). Advances in technology in agriculture has driven the
development of an emerging industry and body of research called precision livestock
farming (or precision agriculture). Precision livestock farming is the application of
technology such as real-time sensors on farm that can be used by the producer to help
make decisions and/or directly manage animals or groups of animals on farms. As
the cost of feeding animals is estimated to be 60–80% of the cost of production,
technological tools that can limit feed wastage and maximise nutrient usage are
highly sought after by producers. However, adoption of technologies on farm
remains relatively low as producer’s acceptance towards these new technologies is
limited. Innovations in animal feeding (such as automatic feeders that can be
programmed to tailor a diet to each specific animal or group) and virtual fencing
are becoming increasingly important in a resource limited modern farming world.
While the greatest uptake of such technologies has been in dairy cattle, other
ruminant systems (including pasture-based systems) are gradually increasing tech-
nology into their daily animal management, although use remains limited by costs. A
combination of tools such as RFID ear tags, GPS, walkover scales, automatic
feeders, automatic drafters and virtual fencing are key technologies that can be
utilized on farm to manage feed (and supplements) and increase feed efficiency in
small ruminants (Odintsov Vaintrub et al. 2021).
14 Strategies to Ameliorate Heat Stress Impacts in Sheep 171

Learning Outcomes
• Heat stress is one of the primary factors which negatively influences the
health and performances of sheep.
• Managemental (shade and shelter, breeding programme) strategies could
help to relieve heat stress in sheep.
• Nutritional strategies (grain feeding, precision feeding, amino acids,
antioxidants, vitamins, electrolytes and minerals) could be employed to
alleviate the negative impacts of heat stress on health and productivity in
sheep.

14.6 Future Perspectives

While much has been learned regarding the impacts and thus management of heat
stress in sheep, much remains unknown and research is required to fill the gaps and
ensure production of sheep meat, milk and fibre can occur in a world adjusting to
climate change induced increases in heat events. There remains a knowledge gap
regarding the metabolic responses to heat, particularly in non-lactating ruminants.
While the actions of insulin play an important role in the metabolic adaptations to
heat stress, the exact mode of action of these responses remains to be elucidated.
Further studies examining the metabolic responses to heat stress are required to fully
discern these processes and advancing our knowledge in this area may lead to
improved feeding and breeding practices to improve production during heat stress.
While no one intervention in isolation is likely to ameliorate all the negative
responses to heat stress, to date there is a lack of published data examining a
combination of interventions in sheep. Research and commercialisation of new
feed supplements and technologies are required to allow production to continue,
and perhaps improve in efficiency, during higher temperature environments. This
might include things like precision feeding that would allow for timed delivery of
feed and/or supplements to sheep. As sheep are generally housed in extensively
managed/grazing systems, the delivery of feed and feed supplements is a major
constraint and any advancements in this space have the potential to rapidly improve
the ability to manage heat stress in sheep.
While this review briefly described some commonly utilized supplements that can
mitigate the impacts of heat stress in sheep, there are others that were not covered in
detail and it is likely novel supplements will be discovered in the future. Further,
there is a lack of research into potential to combine supplements to have an additive
response. This has generally been prevented by the cost required to provide multiple
supplements ensuring that producers would be unlikely to adopt such measures; but
as the impacts of climate change become more prevalent and damaging there might
be an increased cost benefit to providing a combination of feed supplements. The
rumen microbiome and its potential manipulation will be described elsewhere in this
172 K. DiGiacomo et al.

book and remains another area of rapid discovery and advancement. Given the large
contribution that the rumen activity and heat increment of feeding contributes to heat
loads in ruminants, methods that can improve fermentation efficiency (and reduce
heat production) can likely reduce heat production and improve productive
efficiency.

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Strategies to Ameliorate Heat Stress Effects
on Sheep Reproduction 15
Susan Robertson and Michael Friend

Contents
15.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176
15.2 Impacts of Heat Stress on Reproduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176
15.3 Strategies to Optimise Reproduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177
15.3.1 Heat-Tolerant Genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178
15.3.2 Time of Mating and Risks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178
15.3.3 Shearing and Shade to Protect from Solar Radiation . . . . . . . . . . . . . . . . . . . . . . . . . . 179
15.3.4 Nutritional Interventions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180
15.4 Targeted Nutrition to Increase Fertility and Fecundity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180
15.5 Targeted Management and Nutrition to Improve Lamb Survival . . . . . . . . . . . . . . . . . . . . . . 181
15.6 Future Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183

Abstract

The reproductive performance of sheep will be challenged by increasing

temperatures and more extreme weather events due to climate change. Heat stress
potentially reduces the mating behaviour and fertility of ewes and rams, while
increasing embryonic mortality, impairing fetal development and reducing lamb
birth weight, and increasing perinatal lamb mortality. Heat stress can occur above
32  C, but the degree, duration, and timing of heat stress determines whether
reproductive performance is reduced, and to what extent. This chapter considers

S. Robertson
Gulbali Institute and School of Agricultural, Environmental and Veterinary Sciences, Charles Sturt
University, Wagga Wagga, NSW, Australia
M. Friend (*)
Office of the Pro Vice-Chancellor (Research & Innovation), Charles Sturt University, Wagga
Wagga, NSW, Australia
e-mail: [email protected]

# The Author(s), under exclusive license to Springer Nature Singapore Pte 175
Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_15
176 S. Robertson and M. Friend

practical husbandry and nutritional interventions to optimise sheep reproduction

in extensive grazing systems. Strategies include genetics, choice of month of
mating and lambing, shearing date, provision of shade, nutritional supplements
and targeted nutrition.

Keywords

Fertility · Fecundity · Lamb mortality · Nutrition · Ovulation

Abbreviations

% Percentage

C Degree celsius
cm Centimeter
e.g. Exempli gratia (for example)

15.1 Introduction

Maintaining or increasing the reproductive rate of sheep is required to meet future

demand for animal protein and to retain profitability for sheep breeders. An increase
in reproductive efficiency is desirable, rather than a simple increase in the number of
ewes, to minimise methane emissions/kg meat produced, offset reductions in the
availability of agricultural land, and minimise capital outlay and risk for producers.
The predicted global increases in temperatures can be expected to negatively
impact on sheep reproduction, although temperatures in some regions are already
sufficient to impair performance. Reduced rates of reproduction decrease the profit-
ability of sheep farming, the rate of genetic gain, and can be an animal welfare
concern, which in certain cases will increase mortality. Identification and implemen-
tation of suitable management strategies to minimise any adverse effects of heat
stress on reproduction across the range of global production systems and
environments is required. This chapter will briefly outline the possible impacts of
heat stress on ewes and rams, then consider key on-farm strategies to increase
reproductive rates in extensive grazing systems, including time of mating and
shearing, provision of shade and nutritional management.

15.2 Impacts of Heat Stress on Reproduction

Heat stress occurs when sheep need to use energy to dissipate heat from the body
into the environment. The thermoneutral zone for sheep has been cited in a review by
Al-Dawood (2017) as 12–32  C, and heat stress is a function of temperature,
humidity, but also solar radiation. Clearly, grazing sheep can be exposed to
prolonged periods of heat stress in many locations.
15 Strategies to Ameliorate Heat Stress Effects on Sheep Reproduction 177

The impacts of heat stress on reproduction in ewes and rams have been reviewed
and described in detail elsewhere (Al-Dawood 2017; Krishnan et al. 2017; van
Wettere et al. 2021), and are summarised here. Temporary ram infertility or
sub-normal sperm production or quality can result from heat stress, and may persist
for approximately 50 days, which is the duration of spermatogenesis. Ambient
temperatures above 32  C applied to rams have been reported as reducing
fertilisation or pregnancy rates, with the reduction in ram fertility increasing with
the severity and duration of heat stress. Both fertilisation rate and embryo mortality
can be adversely affected, evident in increased returns to service if further mating
opportunities are available, or reduced pregnancy rates. Under field conditions the
impact of heat stress on ram fertility is less clear, since diurnal variation in tempera-
ture provides relief, and extended mating periods to multiple rams can mask short-
term impacts on the more susceptible individuals. It is clear however, that
minimising the risk of heat stress in rams for around 8 weeks prior to and throughout
the mating period will optimise pregnancy rates.
Heat stress in ewes can impair cyclicity, oestrous behaviour, alter follicular
development and reduce oocyte quality, ovulation rate, fertility and increase embryo
mortality. Pregnancy rates have been reduced by up to 100% when ewes are exposed
to continuous 32  C conditions. The impact appears to be most severe where heat
stress occurs in the period from 5 days before to 5 days after oestrus, as the
susceptibility of the embryo declines with maturity. However, sufficient diurnal
variation in temperature, as may occur under field conditions, may prevent most of
this impact. In naturally cycling flocks in environments where heat stress is not a
daily event, only a portion of ewes may be exposed to heat stress in the days around
oestrus. Re-mating of non-pregnant ewes in the subsequent oestrus cycle may mask
any impact on pregnancy rates. However, where oestrus is synchronised, or where
heat stress is frequent, avoiding or mitigating heat in the days around insemination is
important to achieve high rates of pregnancy.
Prolonged periods (approximately 50 days) of heat stress in pregnant ewes can
reduce placental and fetal growth, and while feed intake is also reduced by heat
stress, the reduction in growth appears not to be due to reduced intake. Reductions in
the birthweight of lambs of 16–74% have been reported. The extent of reduction
depends on the severity, timing and duration of heat stress. Lower birthweights
increase the risk of lamb mortality, and more than a 25% increase in lamb mortality
has been reported for ewes heat stressed during the last two thirds of pregnancy (van
Wettere et al. 2021). Heat stress can also reduce immunity levels, increasing the risk
of disease, and reduce milk production, largely associated with the reduction in
intake. Both of these factors contribute to an increased risk of post-natal lamb
mortality.

15.3 Strategies to Optimise Reproduction

Extensive grazing systems dominate the large sheep-producing nations such as

Australia and New Zealand. The extensive nature of these enterprises and the large
scale of operations means any intervention to ameliorate heat stress requires minimal
178 S. Robertson and M. Friend

infrastructure and sheep handling or labour, in addition to being cost-effective. A

number of practical methods are currently available for implementation by sheep
breeders.

15.3.1 Heat-Tolerant Genetics

The consequences of heat stress are optimally avoided by using sheep well adapted
to their environment. While an increased risk of more extreme and longer duration of
hot temperatures in summer months may occur with climate change, in some
locations more extreme chill events may also occur. Sheep in these scenarios need
to be resilient to both heat and cold, whereas sheep in warmer environments may
only require greater tolerance to heat to maintain welfare and productivity. As such,
care is needed to select sheep with the appropriate traits for their production
environment.
Breeds derived from tropical regions have higher resistance to heat stress than
those from cooler zones (Slimen et al. 2019). However, heat resilient breeds may be
less productive or lack particular traits for target markets e.g. hair sheep are not
alternatives for fine wool production. There is potential for within-breed selection if
appropriate traits are identified for each breed. For example, in the Australian
Merino, rams with higher levels of skin wrinkle are more susceptible to heat stress,
displayed by a 20% lower conception rate from summer matings (Fowler and Dun
1966). The identification and use in breeding programs of traits or genetic markers
for heat resilience must balance any benefit from resilience against other production
consequences relevant to each breed.

15.3.2 Time of Mating and Risks

One complication of managing sheep under extensive conditions is optimising

weather conditions for the duration of the reproductive cycle, and this difficulty
will vary with the climatic location. The potentially adverse effect of heat stress from
50 days pre-mating for semen viability, throughout 150 days of gestation, and
through a lambing period of several weeks means sheep will be at a susceptible
reproductive stage for approximately 8 months per year.
In some locations, heat stress around the mating period may be easily avoided by
not mating during the summer months. However, if the consequence of this is to
incur heat stress during later pregnancy or for ewes to lamb during inclement (hot or
cold) weather, resulting in poor lamb survival, this may not be advantageous.
Mortality of new-born lambs due to hypothermia can exceed 60% in extreme
conditions, with lambs born as twins or multiples to Merino ewes more susceptible
than singles and crossbred ewes (Donnelly 1984). While hyperthermia during the
lambing period does contribute to perinatal lamb mortality, there is insufficient data
to inform actual levels of loss under field conditions, since extensive grazing flocks
typically are not mated to lamb in summer months and estimates are derived from
housed studies. Van Wettere et al. (2021) have predicted a 26% reduction in lamb
15 Strategies to Ameliorate Heat Stress Effects on Sheep Reproduction 179

survival associated with lower birthweights when ewes were continuously exposed
to 32  C for various stages during pregnancy. In practice, the impact of heat stress
during pregnancy on lamb survival will depend on the timing, severity and duration
of heat, and diurnal variation will reduce the effect. The seasonality of the ewe
breeding cycle and potential ovulation rate are other factors which influence choice
of month of mating. A decision is therefore required as to what month of mating the
flock will optimise the overall reproductive rate. Location-specific assessments of
the monthly climatic risks are needed to enable this, along with consideration on the
feed supply to meet demands, and in some cases market demand at the point of sale,
as discussed later.
Extending the period of mating may allow non-pregnant ewes another mating
opportunity, although extended joining and subsequent lambing can complicate
flock management. An alternative strategy to reduce climatic risk is to split mating
over more than one period. Simulation studies, based on a field experiment in
southern Australia, have shown a February split with April mating, compared with
mating all ewes in April, resulted in similar lamb production but reduced production
risk (Robertson and Friend 2020). In addition to spreading the risk due to month of
mating, such a system allows re-mating of ewes failing to conceive or losing
embryos from the earlier mating, and as such may increase the number of lambs
born, without disrupting the farm management schedule. However, multiple mating
periods do require a higher managerial skill, and pregnancy scanning is
recommended to identify the ewes to re-join.
Month of mating will also alter the potential stocking rate, weaner growth rates
and risk of mortality, the need to supplementary feed ewes or lambs, and ability to
meet particular market targets, all of which impact on risk, cost and potential income
(Robertson and Friend 2020). Minimising the risk of heat stress is therefore only one
of several key factors which must be considered in choice of month of mating.
Use of higher ram percentages than normal (1–2%) may reduce the risk of using
individual rams with heat-impaired fertility or performance. However, this may not
be cost-efficient or effective if the heat stress is too severe, or the timing of heat stress
is inopportune relative to the mating period. The use of several rams with a mob of
ewes reduces the risk that sub-fertile rams will reduce conception rates, since ewes
may mate with more than one ram.

15.3.3 Shearing and Shade to Protect from Solar Radiation

Time of shearing wool sheep such as the Merino can be used to reduce heat stress
resulting from solar radiation. Increasing fleece length up to 4 cm reduces heat stress
due to radiant heat, by providing insulation, although the optimum wool length may
vary with breed and the density of fleece (see review by Al-Dawood 2017). How-
ever, under hot but humid conditions (33  C dry bulb temperature; 55% relative
humidity) Merino sheep with a long fleece length (>8 cm) were less able to regulate
body temperatures than recently shorn sheep (Beatty et al. 2008). Therefore, very
short wool during hot months is desirable in humid, but not hot-dry environments.
Many factors contribute to the choice of shearing date, including disease risk
180 S. Robertson and M. Friend

associated with fleece length (e.g. flystrike, fleece rot, grass-seeds, sunburn), con-
tamination with vegetable matter, shearer availability, and the timing of other farm
management activities, and as such will also need to be considered.
Provision of shade also reduces heat resulting from solar radiation, and has been
reported to reduce heat load by up to 50% (Al-Dawood 2017). Adequate air
movement and ventilation is necessary to allow heated air to escape if structures
are used. Sheep readily seek shade in warm to hot weather, modifying behaviour to
seek water and graze during the cooler periods of the day while idling in shade
during the heat. In the extensive grazing situation, trees or shrubs may be the only
cost-effective form of shade.

15.3.4 Nutritional Interventions

The reduction in feed intake resulting from heat stress is accompanied by an increase
in maintenance energy requirements reported as between 7% and 30%, meaning
sheep are likely in negative energy balance (Al-Dawood 2017; Krishnan et al. 2017).
Nutritional restriction during heat stress therefore exacerbates potential impacts on
productivity (Slimen et al. 2019). As such, improving nutritional status can be an
effective means of reducing the impact of heat stress on reproduction.
Numerous nutritional interventions have been reviewed and proposed to mitigate
the production impacts of heat stress, including feeding of energy-dense
concentrates, non-fermentable protein, fat (Slimen et al. 2019), feeding during cooler
parts of the day and feeding in smaller feeds more frequently, use of antioxidants
including vitamin C and E, adequate supply of water (Al-Dawood 2017), higher
quality forages, minerals including sodium and potassium, and ascorbic acid
(Krishnan et al. 2017). In the extensive grazing systems, not all of these strategies
are practical, as sheep are usually grazing pastures, rather than being fed complete
rations. Additionally, economic viability must be considered. There may be oppor-
tunity however, to supplement sheep at specific stages of the reproductive cycle, or
to alter the forage base to improve nutrition.

15.4 Targeted Nutrition to Increase Fertility and Fecundity

Feeding of a grain supplement, such as lupins (Lupinus angustifolius), to rams for

8 weeks prior to the mating period has long been recommended to promote sper-
matogenesis. This strategy has not been evaluated for mediation of heat stress and
may not be effective if this period coincides with ample high-quality pasture, or if
feeding occurs outside the natural breeding season in unresponsive breeds. However,
it will assist in overcoming the reduced intake due to heat stress. Well-nourished
rams in good but not over-fat condition are required to optimise both semen quantity
and ram behaviour.
Similarly, it is well known that undernutrition impairs the ability of ewes to cycle
and conceive. Ewes in higher condition score also have a higher ovulation rate.
However, there is opportunity to target nutrition around mating to increase ovulation
15 Strategies to Ameliorate Heat Stress Effects on Sheep Reproduction 181

rate even when ewes are in optimal condition. Ovulation rate sets the maximum
potential number of offspring. Our group has shown that grazing high quality green
feed for 1 week before and after the start of mating during the natural breeding
season improves fecundity by 10% or more, through targeting the critical period of
days 10–14 of the oestrous cycle for the majority of ewes mated (Robertson et al.
2014). While feeding an energy supplement is an alternative strategy, grazing quality
pasture reduces labour and feed costs, and allows sheep to modify behaviour to
compensate for heat (e.g. grazing at night). The short duration of increased nutrition
improves the efficiency of resource use, but could be extended under hot conditions.
This would improve the nutrition of ewes during the period to 5 days after oestrus
when embryos are most susceptible to heat stress (van Wettere et al. 2021). The
benefit of improved nutrition under heat stress may mean producers reconsider the
forage base available at the time of mating, although soil and environmental
constraints may limit the type and quality of forage able to be produced.

15.5 Targeted Management and Nutrition to Improve Lamb

Survival

Peri-natal mortality has historically been a significant concern for the global sheep
industry, even prior to the added risk of reductions due to heat stress. Mortality
causes substantial reductions in potential income for producers, and is increasingly
being perceived as an animal welfare issue, with future risks in the social licence of
husbandry and market access. Preventing increases in mortality due to rising global
temperatures is therefore critical.
The lower birth weight of lambs born to heat-stressed ewes poses a risk of
reduced perinatal survival (van Wettere et al. 2021), particularly if lambs are born
during adverse weather. Changes to climate, resulting in milder temperatures during
lambing, may mean that lamb survival may not be reduced to the extent indicated by
birthweight alone, since hypothermia is also a significant contributor to mortality in
some locations. Warmer weather may also improve pasture growth and ewe nutrition
at some locations during the lambing period, potentially improving colostrum
production and maternal behaviour.
Hyperthermia can cause mortality in new-born lambs, regardless of whether ewes
were heat-stressed during pregnancy. Strategies which reduce mortality during hot
weather involve limiting heat stress through provision of shade, but also limiting the
potential for excess movement of ewes to access feed or water. The need to move large
distances or in difficult terrain in heat exposes new-born lambs to the risk of heat
exhaustion and mismothering whilst attempting to follow their mothers. Congregation
of large numbers of ewes and lambs at feeding sites or water points at the same time, as
occurs when a mob seeks water at dawn and dusk, may also lead to mismothering and
mortality. Paddock selection, smaller mob size, and feeding strategies are factors
which may reduce the risk of mortality in new-born lambs born during hot weather.
Adequate nutritional supply of energy, protein, minerals and vitamins and main-
tenance of a suitable condition score of ewes during pregnancy and lactation are
required to optimise perinatal lamb survival. Where fetal growth and development
182 S. Robertson and M. Friend

has been compromised due to heat stress during pregnancy, or heat stress is
occurring during the lambing period, adequate nutrition increases in importance to
promote appropriate ewe-lamb behaviours, bonding, and colostrum and milk
production.
The reduced intake and higher maintenance requirements associated with heat
stress also increase the risk of metabolic disorders in ewes. Recent research has
shown that the energy balance of ewes and the immune response in lambs is
increased by feeding above-maintenance levels of calcium and magnesium
(Ataollahi et al. 2018). Although not measured, these changes could potentially
improve lamb survival, particularly under heat stress since heat stress reduces
immune function (Al-Dawood 2017). The study also indicates that the current
estimates for mineral requirements may be too low to maximise ewe and lamb
survival.

Learning Outcomes
• Extreme weather events greatly challenge the reproductive performances of
sheep.
• Heat stress has the potential to significantly impair the reproductive
behaviour and fertility of ewes and rams, increase embryonic mortality,
and impair fetal development, growth and perinatal lamb survival.
• Husbandry strategies (month of mating, lambing and shearing, provision of
shade) and nutritional intervention including targeted nutrition could opti-
mize the sheep reproduction.

15.6 Future Perspectives

Excessive heat stress is undeniably a threat to maintaining the reproductive perfor-

mance of sheep at many locations around the world. The susceptibility of sheep
throughout the reproductive process means adaptive practices are needed for each
stage. Breeding sheep which are resilient to heat is a viable and attractive goal,
although long-term. For selection to be a successful strategy, resilience needs to be
gained without significant negative consequences on other traits relevant to the
particular production system.
A clearer definition of the temperature regimens which are detrimental under
extensive grazing conditions is required to allow managers to know when and what
level of intervention is warranted. Provision of shade is required to maintain animal
welfare, but additional targeted nutritional modifications have the potential to reduce
the impacts of heat stress and improve reproductive performance. While a number of
supplements have been identified, the effectiveness, cost and delivery of these has to
be considered. Improving nutritional levels at key times through strategic allocation
of the existing forage base and feeds, and timing mating and lambing to minimise the
15 Strategies to Ameliorate Heat Stress Effects on Sheep Reproduction 183

risk of extreme weather, may be sufficient in some locations. In locations with

prolonged, severe heat stress, further strategies may be required.

References
Al-Dawood A (2017) Towards heat stress management in small ruminants - a review. Ann Anim
Sci 17:59–88
Ataollahi F, Friend M, McGrath S, Dutton G, Peters A, Bhanugopan M (2018) Effect of calcium
and magnesium supplementation on minerals profile, immune responses, and energy profile of
ewes and their lambs. Livest Sci 217:167–173
Beatty DT, Barnes A, Fleming PA, Taylor E, Maloney SK (2008) The effect of fleece on core and
rumen temperature in sheep. J Therm Biol 33(8):437–443
Donnelly JR (1984) The productivity of breeding ewes grazing on lucerne or grass and clover
pastures on the tablelands of Southern Australia. III. Lamb mortality and weaning percentage.
Aust J Agric Res 35(5):709–721
Fowler DG, Dun RB (1966) Skin folds and Merino breeding. 4. The susceptibility of rams selected
for a high degree of skin wrinkle to heat induced infertility. Aust J Exp Agric 6(21):121–127
Krishnan G, Bagath M, Pragna P, Vidya MK, Aleena J, Archana PR, Sejian V, Bhatta R (2017)
Mitigation of the heat stress impact in livestock reproduction. Theriogenology 8:8–9
Robertson SM, Friend MA (2020) Performance of sheep systems grazing perennial pastures.
4. Simulated seasonal variation and long-term production. Anim Prod Sci 60(3):423–435
Robertson SM, King BJ, Allworth MB, Rummery J, Friend MA (2014) The effect of peri-
conceptual grazing of live pasture on fetal numbers in unsynchronised ewes. Anim Prod Sci
54(8):1008–1015
Slimen IB, Chniter M, Najar T, Ghram A (2019) Meta-analysis of some physiologic, metabolic and
oxidative responses of sheep exposed to environmental heat stress. Livest Sci 229:179–187
van Wettere WH, Kind KL, Gatford KL, Swinbourne AM, Leu ST, Hayman PT, Kelly JM, Weaver
AC, Kleemann DO, Walker SK (2021) Review of the impact of heat stress on reproductive
performance of sheep. J Anim Sci Biotechnol 12(1):1–18
Applications of Genetic Selection
in Breeding for Thermo-Tolerance 16
in Livestock

Surinder Singh Chauhan, Richard Osei-Amponsah,

Veerasamy Sejian, and Frank R. Dunshea

Contents
16.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187
16.2 Identification of Thermo-Tolerant Phenotypes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187
16.3 Breeding for Thermo-Tolerance in Livestock . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190
16.4 Genomic Selection for Thermo-Tolerance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191
16.5 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192
16.6 Future Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193

Abstract

Climate change and rising global temperature are expected to exacerbate further
the problem of heat stress which is very well known to affect livestock produc-
tion, reproduction, and welfare negatively. Despite the tremendous advances in

S. S. Chauhan (*)
Faculty of Veterinary and Agricultural Science, The University of Melbourne, Parkville, Australia
e-mail: [email protected]
R. Osei-Amponsah
Faculty of Veterinary and Agricultural Science, The University of Melbourne, Parkville, VIC,
Australia
Department of Animal Science, University of Ghana, Legon, Accra, Ghana
V. Sejian
ICAR-National Institute of Animal Nutrition and Physiology, Bangalore, India
F. R. Dunshea
Faculty of Veterinary and Agricultural Science, The University of Melbourne, Parkville, VIC,
Australia
Faculty of Biological Sciences, The University of Leeds, Leeds, UK

# The Author(s), under exclusive license to Springer Nature Singapore Pte 185
Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_16
186 S. S. Chauhan et al.

strategies to alleviate the impact of heat stress on livestock production, heat stress
remains a substantial challenge for sustainability of global livestock production
causing substantial economic losses to livestock industries in the warmer parts of
the world. Physical modification of the environment, nutritional interventions,
and genetic selection for heat tolerance are strategies that could be used to
attenuate some of the negative impacts of heat stress. While genetic selection
for heat tolerance would be the best long-term strategy to produce heat resilient
future animals, identification of heat-tolerant animals is a big challenge due to the
negative association between production traits and heat tolerance. Due to this
unfavorable correlation between production traits and heat tolerance, a sustained
reduction in heat tolerance has occurred due to continuous selection for increased
production over the last 50 years. More work is thus needed to find phenotypes
that are more convenient and economically viable to measure and identify heat-
tolerant animals under farm conditions. Similarly, more advancements and cost
reduction in data collection and genotyping are needed to develop genomic
breeding values for heat tolerance. Genomic selection for heat tolerance in
high-producing dairy cattle breeds such as Holsteins and Jerseys may be useful
for the developed nations with more advanced data collection, efficient manage-
ment systems, and genomic technologies. However, less developed countries
with adapted local breeds may have better crossbreeding options between local
stock and high producing exotic breeds. Nevertheless, selection for heat tolerance
will improve the heat resilience of future animals and play an important role in
supporting sustainable livestock production under changing climates.

Keywords
Breeding · Genetic selection · Global warming · Heat stress · Livestock

Abbreviations

C Degree celsius
ADG Average daily gain
AUD Australian Dollar
BCS Body condition score
BV Breeding value
CO2 Carbon dioxide
GEBV Genomic breeding value
GWAS Genome Wide Association Study
H+ Hydrogen ion
H2CO3 Carbonic acid
HCO3 Bicarbonate
HS Heat stress
HSP Heat shock protein
16 Applications of Genetic Selection in Breeding for Thermo-Tolerance in Livestock 187

NEFA Non-esterified fatty acids

pH Potential of hydrogen
SNP Single nucleotide polymorphism
THI Temperature humidity index

16.1 Introduction

Heat stress (HS) caused by increased environmental temperature is one of the

greatest challenges livestock production faces in the warmer parts of the world.
Increased environmental temperature compromises an animal’s ability to dissipate
heat from the body, and core body temperature exceeds the physiological limits for
the species causing HS (West 2003). HS has negative effects on animal physiology,
welfare and production, which will be exacerbated by global warming (Osei-
Amponsah et al. 2020). Significant progress has been achieved in understanding
the impacts of HS on various livestock species, economic losses associated with HS,
and the potential strategies for amelioration of HS (Dunshea et al. 2013) and have
been thoroughly discussed by other authors in this book. However, HS remains a
significant challenge for sustainable livestock production and continues to cause a
substantial financial burden to global livestock production, especially in warmer
climates. There are three types of strategies: physical modification of the environ-
ment, nutritional interventions, and genetic selection for heat tolerance, which could
be adopted to ameliorate HS in livestock species. Significant progress has been
achieved in the physical modification of the animal’s environment (providing shade
and shelter, cooling down during hot times) and nutritional interventions (feed
supplements) which we and other authors have discussed in separate chapters in
this book. This chapter summarizes the genetic manipulations to improve heat
tolerance, particularly recent advances in genomic technologies and their application
for selecting heat-tolerant animals.

16.2 Identification of Thermo-Tolerant Phenotypes

One of the biggest challenges of selecting animals for thermo-tolerance is the lack of
convenient, efficient, and economically viable methods to identify heat tolerant
versus heat-sensitive animals under farm conditions (Carabaño et al. 2019). Differ-
ent methods of measuring heat tolerance are used, which are mainly based on
animal’s physiological and behavioral responses or ability to maintain homeothermy
and production performance when exposed to hot conditions. One of the common
behavioral mechanisms exhibited by most livestock species on exposure to high
ambient temperature is reduced feed intake. For example, in sheep, various adaptive
responses, including reduced feed intake, rumination time and frequency, and
increased water intake, are invoked to cope with the increased environmental
188 S. S. Chauhan et al.

temperature (Joy et al. 2020a). Such responses are invoked to reduce metabolic heat
production, one of the important contributors to heat load on animals, the other being
ambient temperature. Research involving various livestock species has consistently
reported a significant decline in feed intake (West 2003; Dunshea et al. 2013;
Chauhan et al. 2014; Joy et al. 2020b; Zhang et al. 2021). In addition, animals
exposed to HS tend to increase their total water intake as a compensatory mechanism
to restore the elevated fluid loss through respiratory and cutaneous routes to cool the
body. Apart from maintaining the body water status, the increased ingestion of water
also helps the heat-stressed animals cope with the adverse conditions by immediate
rumen-reticular cooling, which effectively reduces their core body temperature.
However, genetic differences among different breeds show a variable decline in
feed intake. For example, our recent work on HS in sheep in Australia, involving
Dorper and second cross lambs [Poll Dorset  (Border Leicester  Merino)],
showed reduced feed intake in second-cross Merino lambs, but not in Dorper
lambs when both breeds were subjected to HS in the climatic chambers (Joy et al.
2020b). Thus, reduced feed intake in second cross lambs under HS could be
attributed to them being more heat sensitive and on the other hand, Dopers can be
identified as heat tolerant, given less reduction in feed intake and less increase in
water intake, only 27% in Dorper lambs compared to 63% in second cross Merino
lambs in this study.
Physiological and metabolic responses of the animals following exposure to
higher environmental temperatures are also used to measure heat tolerance. Physio-
logical measures of heat tolerance include respiration rate, rectal temperature, and
pulse rate (Chauhan et al. 2014). The magnitude of increase in these standard
physiological measures can be used to evaluate HS in livestock. Increased rectal
temperature on exposure to heat indicates that the thermoregulatory responses are
insufficient to maintain homeothermy. Generally, in hot environments, increased
respiratory rate is the first thermoregulatory mechanism recruited by the animals to
lose heat and to maintain their body temperature. Panting is another response
recruited by cattle and sheep in response to severe heat, which marks a substantial
increase in the respiratory rate (Joy et al. 2020b). Our recent summer study (Osei-
Amponsah et al. 2019) involving 120 lactating dairy cows in Northeast Victoria,
Australia, showed that the cows experienced significant increase in average body
temperature (11%), respiration rate (67%) and panting scores (64%) on exposure to
summer conditions with temperature humidity index (THI) ranging 72–84. Simi-
larly, sheep studies have demonstrated an increase in respiration rate, rectal temper-
ature, and panting scores over the summer season. However, variation in
physiological changes on exposure to heat can in different genotypes and, therefore,
compare heat tolerance. For example, Dorper sheep exhibited lower rectal tempera-
ture, respiratory rate and skin temperature than Merino cross lambs which indicated
better heat tolerance of Dorper lambs (Joy et al. 2020b).
Production traits are also used to measure thermo-tolerance, as HS influences the
animal’s performance negatively. For example, in dairy cattle, reduction in milk
production is one of the most significant implications of HS, and up to 30% decline
in milk production has been recorded (West 2003). Cows that show a greater
16 Applications of Genetic Selection in Breeding for Thermo-Tolerance in Livestock 189

reduction in milk production on exposure to heat can be termed as heat-sensitive,

while the cows with smaller decline in milk production are termed as heat tolerant
(Nguyen et al. 2016). This is a cumulative effect of reduced grazing in extensive
system and declined feed intake in the intensive system. Our recent study (Osei-
Amponsah et al. 2020) in dairy cattle in Australia, showed a 14% decline in milk
yield despite the provision of the shade and cooling system in the milk parlor, which
clearly showed that multiple HS mitigation strategies are required to ameliorate the
adverse effects of HS. On an average, on a hot day with an ambient temperature
above 32  C, 95% of the cows sought shade and were reluctant to go grazing, and
60% of the cows spent most of the day near the water bodies to cool down.
Similarly, a reduction in production performance occurs in sheep as an effect of
harsh conditions and reduced feed intake. This then leads to a drop in body weight
and average daily gain (ADG) (Zhang et al. 2021). Loss of body weight during HS
might also be attributed to the increase in energy expended for heat dissipation
through respiratory evaporation and, subsequently, the reduction in the amount of
water available for storage. Additionally, the warm environment also causes a
decline in body condition score (BCS) due to insufficient fat storage and
compromised energy availability. The reduction in feed intake for an extended
period may have implications for carcase and meat quality characteristics. Activation
of the adrenergic system, initiated by HS, leads to peripheral vasodilation and
muscle glycogenolysis, resulting in higher ultimate pH and darker meat from heat-
stressed ruminants (Zhang et al. 2021). The meat from heat-stressed goats exhibited
increased cooking loss, reduced water holding capacity, and increased toughness
(Archana et al. 2018). In contrast, a recent study conducted in Dorpers and second
cross Merinos, did not show any effect of higher temperature (2 weeks exposure) on
meat color, water holding capacity and texture (Zhang et al. 2021). It is worth
mentioning that the variation in these results could be attributed to the duration of
HS to which those animals were exposed to. It appears that the impact of HS on these
parameters is quite limited when the heat exposure time is shorter than 3 weeks.
The biochemical profile of ruminants is also modified under hot environments,
with changes in hematocrit values, hemoglobin and lowered blood glucose, protein,
cholesterol, and non-esterified fatty acids (NEFA) concentrations (Joy et al. 2020a).
Increased respiration rate during severe HS may result in respiratory alkalosis in
sheep (Chauhan et al. 2014), greater heat dissipation, and increased alveolar ventila-
tion. An associated elevation in CO2 excretion shifts the bicarbonate equilibrium to
H2CO3 from H+ and HCO3 . At the molecular level, adaptive responses to heat
exposure are driven by increased expression of heat shock proteins (HSPs) which
function as intra-cellular chaperones and prevent protein and cell damage in heat-
stressed animals, along with many other genes (Collier et al. 2008). Research in
various livestock species (See in review Chauhan et al. 2021) have also
demonstrated compromised immune responses during summer.
While a number of physiological, biochemical, and molecular markers and
production responses to HS have been identified across species, better understanding
of the genetic basis of thermo-tolerance is needed for accurately phenotyping heat
tolerance. Potential tools capturing the changes in biomarkers of HS on farms are
190 S. S. Chauhan et al.

required for the successful application of genetic selection for thermo-tolerance in

various livestock species.

16.3 Breeding for Thermo-Tolerance in Livestock

Genetic improvement for thermo-tolerance in dairy cattle is more profitable as it

produces a permanent and cumulative change in the animal population. Genetic
selection for thermo-tolerance may provide a sustainable means of augmenting other
HS mitigation strategies such as feeding and/or housing modification to reduce
losses. There are considerable genetic resources and local breeds of cattle in tropical
and subtropical countries that are well adapted to hot environmental conditions.
However, the lower production of these animals is a big challenge due to the
negative association between production traits and heat tolerance. Due to this
unfavorable correlation between the production traits and heat tolerance, a signifi-
cant reduction in heat tolerance has occurred in the dairy animals where continuous
selection for productivity has been practiced (Pryce and Haile-Mariam 2020).
Identifying the stress-adaptive genes in heat-tolerant breeds could provide the
selection targets for selection in new breeding populations of high-producing
animals. Furthermore, the selection signature for thermo-tolerance can be identified
through functional genomics. The heat resilience of high-producing breeds could be
improved by introgression of HS tolerant genes or through crossbreeding with
resilient genotypes. For example, heat-tolerant Holstein genetics have already been
developed through the introgression of SLICK haplotype, initially identified in
Senepol cattle. SLICK haplotype Holstein cows have a better ability to regulate
the body temperature and experience a less-pronounced reduction in milk yield
when exposed to hot conditions (Dikmen et al. 2014). While introgression can be
quite useful to incorporate favorable alleles from different breeds, it is usually a very
long process involving several generations of backcrossing to restore production
traits.
Another option for improving the thermo-tolerance while maintaining or improv-
ing production could be crossbreeding involving local breeds and high-producing
selected exotic breeds. This could be more appropriate for the tropical regions
characterized by relatively high HS environments than the temperate regions.
Local breeds, particularly in Asia and Africa, are known for their better thermo-
tolerance and ability to thrive under harsh conditions. In general, tropical and
subtropical breeds of cattle have greater adaptive capacity to stressful environments
than exotic cattle. For example, Zebu cattle (Bos indicus) can better regulate body
temperature under hot conditions than Bos taurus breeds of cattle. Bos indicus
breeds have been naturally selected for survivability rather than production traits.
For example, the majority of the breeds from warm climates prioritize fattening
rather than milk production under good management and feeding conditions, clearly
indicating that lower production is one of their constitutional characteristics. Addi-
tionally, the introduction of high-producing germplasm and genomic selection
16 Applications of Genetic Selection in Breeding for Thermo-Tolerance in Livestock 191

(described below) may not be best suited to the regions facing scarcity of quality
feed and fodder, poor housing management, and greater incidence of diseases.

16.4 Genomic Selection for Thermo-Tolerance

Recent advances in genomics and transcriptomics may help to develop genomic

breeding values for heat tolerance across breeds. Genomic selection uses genome-
wide DNA markers to capture the effects of many mutations that influence variation
in a complex trait like heat tolerance (Nguyen et al. 2016). Information from gene
expression or genome-wide association studies (GWAS) can be used to improve the
accuracy of selection further. GWAS has often been used to identify regions on the
genome that have a specific effect on a trait of economic importance in ruminant
livestock. Genomic selection presents the advantage of accelerated genetic gain, as
livestock can be selected from a young age using their estimated genomic breeding
values (GEBVs), rather than waiting to test their progeny. Australia was the first
country to release breeding values (BVs) for heat tolerance in 2017, with an accuracy
of 0.42–0.61 using high-density SNP genotypes (Nguyen et al. 2016; Garner et al.
2016). GEBVs for thermo-tolerance of Australian Holsteins and Jerseys are relevant
because profitability and animal welfare can be improved by identifying the animals
that can adapt to current and future climate challenges. When exposed to simulated
heatwaves, genomically predicted heat-susceptible and -tolerant animals showed
significant differences in their decline in milk production and rectal and intravaginal
temperatures (Garner et al. 2016). Heat tolerance ABV allows farmers to identify the
animals with greater ability to tolerate hot, humid conditions with less impact on
milk production. It is expressed as a percentage with a base of 100. An animal with
BVs of 105 is 5% more tolerant to hot, humid conditions than average, and its drop
in production will be 5% less than the average. On the other hand, an BVs of
95 means the animal is 5% less tolerant to hot, humid conditions than the average
and its drop in production under heat stress is 5% more than the average (DataGene).
Given the negative association between milk production and heat tolerance, the
association of GEBVs for thermo-tolerance with BVs for other production traits will
help implement genomic predictions for thermotolerance. Preliminary data
(Fig. 16.1) from our recent study conducted in a herd of Holstein Friesian cows
exposed to HS over Australian summer showed a positive association between heat
tolerance GEBVs and Australian Feed Saved breeding values indicating improving
feed efficiency can improve thermo-tolerance.
However, because thermo-tolerance is a genomic trait identified only recently, the
GEBVs for HT are less reliable than those of already established phenotype-based
production traits such as feed efficiency, fertility, and mastitis resistance. The
accuracy of GEBVs for thermo-tolerance is further limited using variation in test-
day milk production to measure heat stress, as the variation between days is not
wholly dependent on ambient conditions alone. Data on core body temperature with
increasing temperature and humidity and altered fatty acid profiles in milk using
mid-infrared spectroscopy may improve the accuracy of GEBVs. However, the cost
192 S. S. Chauhan et al.

200

150

100
Mean

50

0
93 95 97 99 100 101 102 103 104 105 106 107 108 110 112
-50

-100 Heat Tolerance

Temperament Mastitis resistance Feed saved Fertility

Fig. 16.1 Variation of temperament, mastitis resistance, feed saved, and fertility GEBVs with heat
tolerance GEBVs of Holstein dairy cows

of genomic selection is an important issue; if this is too high, farmers may not wish to
participate in a breeding scheme involving genomic selection. In 2011, the cost of
genotyping was approximately 100 AUD per heifer which was proven to be not
economically viable as a selection tool. It is encouraging that the cost of genotyping
of cattle has shown a steady reduction, making genome-wide selection more afford-
able in the future. If the price of genotyping is further reduced in developing
countries (perhaps by a subsidy), this would have a major impact on farm profitabil-
ity by selecting heat-tolerant animals.

Learning Outcomes
• Genetic selection is an very effective tool to identify climate resilient
animals.
• Crossbreeding between local breeds and high producing exotic breeds may
be better strategy for producing thermo-tolerant animals in developing
countries.
• Selection for heat tolerance will improve the climate resilience of future
animals and play an important role in supporting sustainable livestock
production.

16.5 Conclusion

Despite the significant advances in strategies to alleviate the impacts of heat stress on
livestock production, heat stress remains a significant challenge for sustainability of
global livestock production, especially in the warmer parts of the world. Physical
16 Applications of Genetic Selection in Breeding for Thermo-Tolerance in Livestock 193

modification of the environment, nutritional interventions and genetic selection for

heat tolerance are some of the strategies that could be used to attenuate some of the
negative impact of HS. While genetic selection for heat tolerance can provide a long-
term strategy to produce heat resilient animals in future, identification of heat
tolerant animals itself is a big challenge due to lack of accurate and commercially
viable tools for identifying heat tolerant phenotypes, which is further exacerbated by
the negative association between the production traits and heat tolerance. Therefore,
more research is required and should be supported by stakeholders to improve the
phenotyping for thermo-tolerance and improve the understanding of the genetic
basis of thermo-tolerance.

16.6 Future Perspectives

Global warming and HS will continue to challenge our livestock industries,

compromising animal welfare and farm productivity. HS is and will remain a
complex issue eliciting multiple responses and triggering various physiological
pathways in the animal body, negatively impacting their production. Genetic
improvement of animal populations and physical modification of their environment
and feeding strategies to reduce the negative impacts of HS will play an important
role in ameliorating HS in animals. Existence of considerable variation in heat
tolerance between and within the breeds of cattle provides an excellent opportunity
for the selection of thermo-tolerant livestock and will be an important step towards
breeding resilient animals. While the genetic selection for thermo-tolerance will help
produce heat tolerant animals for the future, more research should be aimed at
accurately and conveniently identifying the heat-tolerant phenotypes. In this regard,
global strategic collaborations would be critical to generate and share quality data on
thermo-tolerant phenotypes that could be used for genomic predictions. While
genomic selection will play a key role in future breeding programs, its application
and adoption by different countries will depend upon the type of resources available.
Genomic selection for thermo-tolerance within high producing animal populations
will be an ideal strategy for producing heat tolerant animals in resource-rich
countries. On the other hand, resource poor nations will need to focus on better
identifying and characterizing breeds that are best suited for their production
environments where nutritional deficiency and disease burden might be bigger
challenges than HS. In those countries, introgression of the slick gene for instance
and crossbreeding between local and high producing exotic breeds, will play an
important role in improving thermo-tolerance and production of more resilient
genotypes for climate change adaptation.

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 Part V
Enteric Methane Emission and Amelioration
Enteric Methane Mitigation in Livestock: An
Indian Perspective 17
Raghavendra Bhatta, Pradeep Kumar Malik, and Veerasamy Sejian

Contents
17.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 199
17.2 GHG Emissions from Livestock . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200
17.3 Enteric Methane: A Necessity for Ruminants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201
17.4 Enteric Methane Emission: Indian Scenario . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203
17.5 Enteric Methane Amelioration: Challenges and Opportunities . . . . . . . . . . . . . . . . . . . . . . . . . 203
17.6 GHG Emissions from Manure Management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 204
17.7 Measures for Reducing GHG Emissions from Manure Management . . . . . . . . . . . . . . . . . . 208
17.8 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 208
17.9 Future Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210

Abstract
An interdependent relationship exists between livestock and climate change,
where one adverse event can have a ripple effect on the other. Methane (CH4)
emissions from livestock can occur due to enteric fermentation and manure
management; nevertheless, nitrous oxide (N2O) emissions are entirely due to
animal manure management systems. In general, Indian livestock emits 8–10 Tg
enteric CH4 per year, with cattle and buffaloes generating more than 90% of the
total enteric CH4 emission from livestock in India, while small ruminants emit
7.7%. Reducing cattle numbers, feeding quality concentrates, fodders, supple-
mentation of Ionophores, balancing ration, Defaunation of protozoa, reducing
acetogenesis, use of plant secondary metabolites, bio-hydrogenation, and active
use immunisation and disabling of surface proteins are the essential ameliorative
techniques for reducing enteric methane emissions. The primary greenhouse gas

R. Bhatta (*) · P. K. Malik · V. Sejian

ICAR-National Institute of Animal Nutrition and Physiology, Bangalore, India

# The Author(s), under exclusive license to Springer Nature Singapore Pte 197
Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_17
198 R. Bhatta et al.

emitted by manure is CH4, which is emitted during the decomposition of organic

matter anaerobically, while N2O during storage and soil application. Standard
management procedures could lower both CH4 and N2O emissions from manure.
Considering that India has the world’s most enormous livestock population,
reducing livestock-related GHG emissions has to be the key focus. This might
improve farm profitability by diverting all energy into productive use, signifi-
cantly minimising livestock-induced climate change.

Keywords
Climate change · Enteric methane · GHG · Manure · Nitrous oxide

Abbreviations

% Percentage
Bo(T) Maximum methane producing capacity for manure produced by
livestock category T
CH4 Methane
CO2 Carbon dioxide
EF(T) Emission factor for livestock category T
EPA Environmental Protection Agency
FAO Food and Agriculture Organization
Fig. Figure
GHGs Greenhouse gases
H2 Hydrogen
ha Hectare
kg Kilogram
MCF Methane conversion factors
MS (T,S,k) Fraction of livestock category T’s manure handled using manure
management system S in climate region k, dimensionless
N Number of head of livestock species
N2 Dinitrogen
N 2O Nitrous oxide
N2OD Direct N2O emissions from manure management
Nex Annual average N excretion
NH3 Ammonia
NOX Nitric oxide
S Manure management system
T Species/category of livestock
Tg Teragram
VS Volatile solids
WRI World Resources Institute
yr Year
17 Enteric Methane Mitigation in Livestock: An Indian Perspective 199

17.1 Introduction

During 2005, the global greenhouse gas (GHG) emission was around 49 Gigatonnes
(WRI 2011). China ranks first in methane emission, followed by the United States of
America and the European Union (WRI 2011). India accounts for only 4.25% of
global GHG emissions (Fig. 17.1). Livestock is considered a crucial component of
agriculture and provides subsistence for over a billion people. This sector alone
provides 13% of calories and 28% of protein requirements word-wide. Global
demand for milk, beef and eggs is predicted to increase by 30%, 60%, and 80%,
respectively, by 2050, compared to 1990, due to rapidly changing food patterns. This
additional demand can be met by either expanding the number of animals or
strengthening current livestock stock productivity. FAO estimates that 1526 million
bovines and 1777 million bovines are maintained, while these populations are
anticipated to expand around 2.6 and 2.7 billion by the end of 2050, respectively.
Livestock and climate change are inextricably linked via a complex process in
which one’s adversity impacts the other. Climate change is a constant threat to the
livestock sector, but the severity of adversity varies according to the prevailing agro-
climatic conditions. Climate change, directly and indirectly, impacts livestock pro-
duction through changes in the atmosphere (stress), nutritional and volumetric
changes in fodder crops, health, and other factors. In other words, we can consider
livestock as both culprit and victim as far as climate change is debated. Climate
change’s deleterious effect on livestock productivity, either directly or indirectly, is
outside the scope of this chapter; so, only the involvement of livestock in global
greenhouse gas (GHG) emissions and mitigation methods will be discussed.

Fig. 17.1 Nation wise greenhouse gas emissions (WRI 2011)

200 R. Bhatta et al.

17.2 GHG Emissions from Livestock

Livestock produces a significant amount of carbon dioxide (CO2), methane (CH4),

and nitrous oxide (N2O). Emissions of CO2 from livestock are considered part of the
perpetual cycling biological system, in which plant absorbed CO2 is metabolised by
livestock through digestion and subsequently released into the atmosphere via
respiration to be reabsorbed by plants (FAO 2006). The emitted and absorbed CO2
is considered equivalent, and therefore, the release of CO2 from livestock on a net
basis is almost zero. Due to the net-zero contribution, CO2 as a byproduct of
respiration is excluded from the calculation of livestock GHG emissions (Holtkamp
et al. 2006). Agriculture, including livestock, is the third-largest sector after power
and land-use change to emit greenhouse gases in the atmosphere (Fig. 17.2).
On the whole, a total of 14% of global GHG emissions are attributed to agricul-
ture (Fig. 17.2). Soil alone is accountable for 38% of the GHG emissions (as N2O)
from the agriculture sector; however, enteric fermentation is also not far behind and
contributes 32% of the total GHG emission from the agriculture sector (Fig. 17.3).
Rice cultivation, biomass burning and manure management constitute the rest 30%
of the sector.
Enteric fermentation and manure management contribute to CH4 emissions from
livestock production, but N2O emissions are solely the result of livestock manure
management systems. Large populations and high emissions of CH4 from ruminants
make them a crucial CH4 source in many countries (Bhatta et al. 2009). CH4
emissions from manure management are comparatively very less than enteric
emissions. Confined animal management operations deal with liquid-based manure
management systems, where large amounts of CH4 are released into the atmosphere.
The amount of N2O emitted by manure management systems varies significantly
according to different management systems and may also result in indirect emissions
due to various kinds of nitrogen loss from the system. Globally, livestock only
accounts for 9% CO2 emission, but this sector contributes 35 and 65% of CH4 and
N2O from anthropogenic sources, respectively.

Fig. 17.2 GHG emissions according to different sectors

17 Enteric Methane Mitigation in Livestock: An Indian Perspective 201

Fig. 17.3 Global agricultural

GHG emission

Fig. 17.4 Region wise enteric methane emission (modified from O’Mara 2011)

According to Fig. 17.4, livestock in Latin America release the most enteric CH4,
trailed by Africa and China. With an annual emission of 8–10 Tg, India contributes
significantly to world enteric CH4 emissions. Projections indicate a substantial
increase in enteric CH4 emission from 2010 to 2030 (Fig. 17.5). However, the
increase in CH4 from manure management is not much. Likewise, an increase in
N2O emission is also predicted (Fig. 17.6) by the end of 2030. China’s N2O
emissions from excrement will also increase significantly. The increase in enteric
CH4 emissions in the next 15 years is likely due to the increasing livestock numbers
and their continuous low productive performance.

17.3 Enteric Methane: A Necessity for Ruminants

Rumen serves as a home for a diverse range of microbes that perform various
functions, ranging from complex carbohydrate degradation to the removal of fer-
mentation metabolites syntrophically under a strict anaerobic environment. Enteric
fermentation produces a considerable volume of H2 that is effectively disposed of
202 R. Bhatta et al.

Fig. 17.5 Projections for CH4 emission (2010–2030)

Fig. 17.6 Projections for N2O emission (2010–2030)

from the anaerobic vat to carry out various duties and maintain the favourable rumen
conditions for the dwelling bacteria and the host animal. In a healthy rumen,
metabolic H2 reduces CO2 to CH4, which is then vented into the atmosphere by
the mouth and nostrils. The bacteria referred to as ‘archaea’ are the rumen’s
methane-producing machinery. Although these archaea or methanogens were previ-
ously classified as bacteria (prokaryotes), recent classification indicated that they
belong to archaea’s domain, which is markedly distinct from the bacterial domain.
Among the numerous end products of rumen fermentation, H2 is an essential
metabolite since its partial pressure in the rumen controls the extent to which
methanogenesis occurs and the extent to which feedstuffs can be oxidised (Hegarty
and Gerdes 1999). H2 is commonly referred to as the currency of fermentation in the
rumen (Hegarty and Gerdes 1998). A ruminant can produce around 6–12% of its
17 Enteric Methane Mitigation in Livestock: An Indian Perspective 203

total energy intake in the form of methane through methanogenesis, which is

necessary yet inefficient.

17.4 Enteric Methane Emission: Indian Scenario

India is home to 512 million animals (19th Livestock census, Govt. of India). Cattle
and buffaloes account for 60% of the livestock population and produce the most
methane (CH4) than any other livestock species. Estimates from various
organisations indicate that Indian livestock produces 8–10 Tg of enteric CH4
annually. More than 90% of the enteric CH4 emissions from livestock in India
come from cattle and buffaloes (Fig. 17.7); Small ruminants, such as sheep and
goats, make up only 7.7% of total methane production. Species such as yaks,
mithuns, and others found only in a few places provide the balance of the emissions.
Crossbred cattle produce far more enteric methane than indigenous cattle (46 kg/ha/
yr vs. 25 kg/ha/yr).

17.5 Enteric Methane Amelioration: Challenges

and Opportunities

As far as enteric CH4 emission and its amelioration is concerned, these are the
challenges that should be addressed in a planned manner for achieving long term
CH4 reduction from livestock

• Uncertainty over the validity of the country-specific data

• Improper methodologies for methane prediction
• Shortage of feed and fodder
• Quality of feed and fodder

Fig. 17.7 Species wise % enteric methane emission (modified from Kamra 2014)
204 R. Bhatta et al.

• Low functional productivity

• Twinkling methane reduction attributes
• Complicated rumen microbial system
• Unexplored community of rumen archaea

In order to reduce CH4 emissions from the rumen successfully, any technique under
evaluation must fulfil the criteria of shrinking H2 production (Bhatta et al. 2008,
2013a). A brief overview of the potential opportunities to minimise enteric methane
emission and ameliorative measures (Table 17.1) is presented below.

• Cutting down on the number of unproductive or under-productive animals.

• Dipping of H2 production in the rumen to obstruct the supply
• Diverting H2 utilisation from methanogenesis process
• Targeting H2 utilising microbes for indirect inhibition of methanogenesis
• Targeting rumen archaea for direct inhibition

17.6 GHG Emissions from Manure Management

Soil quality, tilth, and productivity can all be improved by using livestock manure as
a supplement to the soil. CH4 and N2O are the major GHGs produced by manure
during the anaerobic decomposition of organic matter during storage and soil
application, respectively. Indirect sources of nitrous oxide include the gases NH3
and NOX generated by manure, which also contribute to the odour. Estimated and
projected CH4 and N2O emissions from manure management are presented in
Table 17.1. It is clear from the table that the CH4 emission from manure management
in the country is about 9.6% of the world’s emission and projected to be remaining
9.5–10.2% in the next 40 years. N2O emission from livestock waste was estimated at
3.9% of the world’s total (Table 17.2).
Manure management is essential in minimising GHG emissions caused by
microbial activities during manure decomposition. The decomposition of dung
under anaerobic conditions during storage produces CH4. These conditions occur
most readily when manure is disposed of in liquid-based systems. The bulk of dung
produced and the portion of dung that decomposes anaerobically are the main factors
that influence CH4 emissions. When manure is stored or treated as a liquid in
lagoons, ponds, tanks or pits, it decomposes anaerobically and can produce a
significant quantity of CH4. The temperature and the retention time of the storage
unit significantly affect the amount of CH4 produced. Handling dung as a solid (e.g.,
in stacks or piles) or when it is deposited on pastures and rangelands, it tends to
decompose under aerobic conditions and hence, produce very little CH4. The
anaerobic decomposition of the faeces produces CH4 due to its volatile solids
(VS). There are both biodegradable and non-biodegradable fractions of VS in animal
manure. The VS excretion rates can be derived from publicly available sources for
country-specific estimations. Feed intake levels can be used to determine country-
specific VS excretion rates if daily average VS excretion rates are not readily
17 Enteric Methane Mitigation in Livestock: An Indian Perspective 205

Table 17.1 Ameliorative measures for enteric CH4 mitigation

Measures Opportunities/limitation Remarks
Reducing the CH4 emissions per kilogramme of Animals that are not productive
livestock livestock product are high because of the should be graded up through
numbers large number of low- or non-producing rigorous, selective breeding to
ruminants. It is impossible to slaughter increase their productivity and
these animals because of a ban on cow produce less methane
slaughter in India
Feeding of To reduce methane emissions, feed With an ever-increasing human
quality fodders, interventions are the most effective population and food-feed-fuel
concentrate method. The uninterrupted availability is competitiveness, it appears
a question mark. Since the last three impossible to increase the
decades, the area under pasture and availability of excellent fodders
permanent fodder production has
decreased or remained flat. About 7–8%
of the arable land in the country is used
to feed livestock
Ionophore Selective inhibition of microbes and Rotation and combination may be
failure to achieve long-term reduction used to sustain the long-term
are major issues. After a short period of decrease
time, animals return to their normal
emission levels. It is prohibited in
several European nations to use them
Ration Ration balancing with feed resources Ration balance is critical for
balancing available at the farmer’s doorstep will farmers, and they must be made
improve the productivity with aware of the financial benefits
concurrent methane reduction at a low that accrue from it
input level
Removal of Lower levels of CH4 are produced when However, even after complete
protozoa the rumen is freed of ciliate protozoa. removal, enteric methane
Fibre digestibility may be reduced. reduction can be achieved
However, maintaining rumen free of partially through defaunation
protozoa is nearly impossible without compromising fibre
digestion
Reductive The rumen’s methanogenesis is By simultaneously targeting
acetogenesis favoured by thermodynamics. When it rumen archaea, a reduction in
comes to H2 substrate affinity, acetogens acetogenesis can be encouraged.
have a far lower affinity than As a result, there will be less
methanogens. It will not function until methane and more acetate for the
the rumen’s methanogens are targeted host animal
Use of plant The use of PSM as methane mitigation Enteric methane amelioration
secondary agents is a viable option in the event of a may be possible if the degree of
metabolites lack of quality fodder. Dose inclusion is safe and does not
optimisation and validation of methane adversely influence feed
migration potential in vivo on a large fermentability—the combined
scale should be compulsory before effect of PSM on in vivo methane
recommendation emission warrants further
investigation
Nitrate/Sulfate To a more considerable extent, nitrate Slow-release sources for these
and sulphate can reduce methane chemicals may lessen the risk of
emissions. Thermodynamically, these toxicity from intermediate
(continued)
206 R. Bhatta et al.

Table 17.1 (continued)

Measures Opportunities/limitation Remarks
reductive mechanisms outperform metabolites. A safe inclusion
methanogenesis. The outcome of this level must be determined and
production process will have no benefit tested on many animals, taking
to the animal in terms of energy. into account all of the species
Intermediary products are toxic to the accountable for methane
host animal production
Active This strategy has the potential to reduce For consideration of this technique
immunisation methane emissions significantly if the for the amelioration of enteric
methanogenic archaea of the rumen are methane, it is necessary to gather
thoroughly studied to identify the best information on the diversity of
candidate vaccination target methanogenic archaea
Disabling of Identifying and disabling the In order to investigate the
surface proteins methanogens which adhere to other possibilities, significant
microbes surfaces for H2 transfer fundamental and advanced study
through surface proteins might reduce is required in this relatively
enteric methane emission unknown region
Bio- Restricting the H2 supply to This approach is not practical due
hydrogenation methanogens through alternate use in to the high cost of fat/lipids and
bio-hydrogenation decrease enteric fibre depression at a high level of
methane amelioration. The use of use
fat/lipids at a high level depresses fibre
digestion. Of the total, about 5–7% of H2
is only utilised in this process
Source: Bhatta et al. (2000, 2001, 2005, 2012a, b, 2013b, c, 2015) and Malik et al. (2014, 2015)

Table 17.2 Estimate and projected emissions of methane and nitrous oxide from manure
management
Manure methane (kg
105)
Estimated Projected
Methane 2010 2025 2050
World 11,414 12,849 15,046
India 1096 1221 1543
% of total 9.6 9.5 10.2
Manure nitrous oxide (kg
105)
Nitrous Estimated Projected
oxide 2010 2025 2050
World 383 445 516
India 15.3 17.5 21.4
% of total 3.9 3.9 4.1
Source: Patra (2014)

available. The ‘Enhanced’ characterisation approach can estimate cow and buffalo
feed intake. The VS content of dung is the percentage of the animal’s food that isn’t
digested and is excreted as faecal matter. The following equation can be used to
figure out the rate of excretion of VS.
17 Enteric Methane Mitigation in Livestock: An Indian Perspective 207

Volatile solids excretion rate


 h i
DE% 1  ASH
VS ¼ GE ∙ 1  þ ðUE ∙ GE ∙
100 18:45

Using the VS excretion rate, the CH4 emission factor from dung can be calculated
using the equation presented below.
" #
  X MCF S,k
EF ðT Þ ¼ VSðT Þ • 365 • BoðT Þ • 0:67kg=m3 • • MSðT,S,kÞ
S, k
100

Where:
EF(T) ¼ Annual CH4 emission factor for livestock category T, kg CH4
animal1 yr1
VS (T) ¼ Daily volatile solid for livestock category T, kg dry matter
animal1 day1
Bo(T) ¼ Maximum methane producing capacity for manure produced by livestock
category T, m3 CH4 kg1 of VS excreted
0.67 ¼ Conversion factor
MCF (S,k) ¼ Methane conversion factors for each manure management system
S by climate region k, %
MS (T,S,k) ¼ Fraction of livestock category T’s manure handled using manure
management system S in climate region k, dimensionless
Nitrification and denitrification processes work together to produce N2O
emissions directly from manure management processes. During the storage and
treatment of manure, the amount of N2O released depends on the nitrogen and
carbon content of the dung, as well as the storage period. Nitrification (the oxidation
of ammonia nitrogen to nitrate nitrogen) is a necessary prerequisite for the emission
of N2O from stored animal manures. Nitrification is likely to occur when stored dung
has a sufficient supply of oxygen. In order to release N2O from stored animal dung,
ammonia must be oxidised to nitrate nitrogen. When dung is appropriately stored,
nitrification is more likely to occur as long as enough oxygen is available. Anaerobic
denitrification produces N2O and dinitrogen from nitrite and nitrate. Direct N2O
emission from manure management may be estimated using the following equation.
Direct N2O emission from manure management
" " # #
X X  44
N 2 ODðmmÞ ¼ N ðT Þ • NexðT Þ • MSðT,SÞ • EF 3ðSÞ •
S T
28

Where:
N2OD (mm) ¼ direct N2O emissions from Manure Management, kg N2O yr1
N (T) ¼ number of head of livestock species/category T in the country
Nex (T) ¼ annual average N excretion kg animal1 yr1
208 R. Bhatta et al.

MS (T,S) ¼ fraction of total annual nitrogen excretion for each livestock species/
category T that is managed in manure management system S in the country,
dimension less
EF3 (S) ¼ emission factor for direct N2O emissions from manure management
system S in the country, kg N2O-N/kg N in manure management system S
S ¼ manure management system
T ¼ species/category of livestock
44/28 ¼ conversion of (N2O-N) (mm) emissions to N2O (mm) emissions

17.7 Measures for Reducing GHG Emissions from Manure

Management

Table 17.3 lists preventive and ameliorative techniques for reducing manure-related
emissions of CH4 and N2O.

17.8 Conclusions

Livestock produces methane and nitrous oxide due to enteric fermentation and
manure management, which are considered the primary sources of anthropogenic
GHG emissions. These GHGs have a far higher global warming potential than
carbon dioxide. Enteric methane emissions from livestock contribute to global
warming and rob the host animal of a significant portion of its energy supply.
India cannot afford this energy loss since it will require extra feed supplies to
compensate for the shortfall. The implementation of mitigation measures to reduce
enteric methane should be based on the viability of the intervention(s) in the specific
region. Our attention should be on measures that have the potential to endure over
time and result in a 20–25% reduction in enteric methane emission. Different storage
conditions are necessary for manure management’s methane and nitrous oxide
emissions. The methane emission from manure in the country is not very alarming,
so our focus should be on reducing nitrous oxide emissions from manure manage-
ment by developing interventions that ensure that methane emission has not gone up
while trying to mitigate nitrous oxide emission from manure management. Many
strategies have been developed and evaluated in the country. However, their con-
finement to the laboratory is a crucial issue and needs the attention of the extension
workers and line departments for undertaking front line demonstration and
popularising the farmers’ mitigation strategies. Large enteric methane emissions in
absolute quantity from Indian livestock warrants the urgent attention of the
researcher and policymakers.
17 Enteric Methane Mitigation in Livestock: An Indian Perspective 209

Table 17.3 Precautionary/ameliorative measures for reducing GHG emissions from manure
management
GHG Measures
CH4 • It might be a viable option rather than storing manure in a lagoon (this may increase
N2O emission), handling manure as a solid or putting it on pasture.
• Using manure decomposition as a source of CH4 to generate renewable energy
• Avoid incorporating straw with manure, which acts as a substrate for anaerobic
bacteria.
• Prolong storage of manure stimulates anaerobic decomposition leading to increased
CH4; it is better to add manure to the soil as soon as possible.
• Avoiding the process of manure application when is soil is too damp, which might lead
to increased methane production.
• Enhance animal feed conversion efficiency by giving high-quality feeds to reduce GHG
emissions
• Covering the tanks or lagoons with impermeable covers like plastic to capture GHGs.
N2O • Apply manure immediately before crop growth to allow the crop to utilise the
maximum amount of available nitrogen.
• To avoid excessive nitrogen loss during spring and N2O emissions, do not apply
manure in the late fall or winter.
• Applying manure during hot and windy or before a storm should be avoided, as these
conditions may increase N2O emissions.
• Few soil and water management practices like increasing soil aeration, avoiding soil
compaction, improving drainage and using nitrification inhibitors for nitrogen gas
production instead of nitrogen oxide.
• Distribute manure evenly around the pasture.
• Maintaining healthy pastures using effective grazing management strategies contributes
to fodder quality improvement.
• Including the proper amount of amino acid levels in the feeding regime minimises
nitrogen excretion, primarily in the urine. Use phase feeding to match diet to growth and
development
• Reduce the amount of N2O emitted by moving fresh manure into a covered storage
facility.
• Storage at lower temperatures inhibits microbial activity

Learning Outcomes
• Cattle and buffaloes account for more than 90% of the enteric CH4
emissions from livestock in India, while small ruminants account for just
7.7% of the total enteric CH4 emissions from livestock in India.
• Methane emissions can be reduced at a low input level if rations are
balanced with the feed supplies available at the farmer’s doorstep.
• If the methanogenic archaea of the rumen is studied to the full extent, the
active immunisation strategy has the potential to reduce methane emissions
significantly.
210 R. Bhatta et al.

17.9 Future Perspectives

Agricultural GHG emissions come primarily from livestock, producing methane

through enteric fermentation and manure management. GHGs have much greater
global warming potential than carbon dioxide. Enteric methane emissions from
livestock contribute to global warming and divert a significant amount of energy
from the host animal. India cannot afford the energy loss in the form of methane
since it will require extra feed supplies to compensate for the shortfall. Mitigation
options for enteric methane reduction should be chosen based on the feasibility of
the intervention(s) in a particular region. Our attention should be on strategies that
have the potential to endure over time and result in a 20–25% reduction in enteric
methane emission. Different storage conditions are required for methane and meth-
ane emissions from manure management to the fact that methane emission from
manure is not particularly alarming in the country; our focus should be on reducing
methane emission from manure management by developing interventions that
ensure that methane emission does not increase while attempting to mitigate meth-
ane emission from manure management.
Enteric methane amelioration from Indian livestock is required from monetary
benefits to stakeholders and environmental points of view. To achieve the substantial
reduction under the present scenario of feed and fodders deficit, a set of likely
interventions alone and in combinations need to be explored under different feeding
systems and species. The target of the researchers should be long term methane
reduction rather than short term. Basic information on methanogenic archaea should
be explored to effectively control methane generating machinery in the rumen. Even
though the absolute methane emission corresponding to the total livestock popula-
tion in the country is less, due to high enteric methane emission per unit of product,
Indian livestock always remains a matter of discussion. Large enteric methane
emissions in absolute quantity from Indian livestock warrants the urgent attention
of the researcher and policymakers. Given the acute feed shortage and seasonal
variability of the feed resources, the researcher should focus on exploring the
prospects of biological interventions for reducing enteric methane emission from
the livestock, and efforts should be taken up for the genetic up-gradation of the
livestock for low methane emission. Many strategies have been developed and
evaluated in the country. However, their confinement to the laboratory is a crucial
issue and needs the attention of the extension workers and line departments for
undertaking front line demonstration and popularising the farmers’ mitigation
strategies. Large enteric methane emissions in absolute quantity from Indian live-
stock warrant the urgent attention of the researcher and policymakers.

References
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seed husk as a source of tannin on dry matter intake, digestibility of nutrients and production
performance of crossbred dairy cows in mid-lactation. Anim Feed Sci Technol 83(1):67–74
17 Enteric Methane Mitigation in Livestock: An Indian Perspective 211

Bhatta R, Krishnamoorthy U, Mohammed F (2001) Effect of tamarind (Tamarindus indica) seed

husk tannins on in vitro rumen fermentation. Anim Feed Sci Technol 90(3–4):143–152
Bhatta R, Vaithiyanathan S, Shinde AK, Jakhmola RC (2005) Effect of feeding complete feed block
containing Prosopis cineraria leaves and polyethylene glycol (PEG)-6000 on nutrient intake, its
utilisation, rumen fermentation pattern and rumen enzyme profile in kids. J Sci Food Agric
85(11):1788–1794
Bhatta R, Enishi O, Takusari N, Higuchi K, Nonaka I, Kurihara M (2008) Diet effects on methane
production by goats and a comparison between measurement methodologies. J Agric Sci
146(6):705–715
Bhatta R, Uyeno Y, Tajima K, Takenaka A, Yabumoto Y, Nonaka I, Enishi O, Kurihara M (2009)
Difference in the nature of tannins on in vitro ruminal methane and volatile fatty acid production
and on methanogenic archaea and protozoal populations. J Dairy Sci 92(11):5512–5522
Bhatta R, Saravanan M, Baruah L, Sampath KT (2012a) Nutrient content, in vitro ruminal
fermentation characteristics and methane reduction potential of tropical tannin-containing
leaves. J Sci Food Agric 92(15):2929–2935
Bhatta R, Mani S, Baruah L, Sampath KT (2012b) Phenolic composition, fermentation profile,
protozoa population and methane production from sheanut (Butryospermum Parkii) byproducts
in vitro. Asian Australas J Anim Sci 25(10):1389–1394
Bhatta R, Enishi O, Yabumoto Y, Nonaka I, Takusari N, Higuchi K, Tajima K, Takenaka A,
Kurihara M (2013a) Methane reduction and energy partitioning in goats fed two concentrations
of tannin from Mimosa spp. J Agric Sci 151(1):119–128
Bhatta R, Baruah L, Saravanan M, Suresh KP, Sampath KT (2013b) Effect of medicinal and
aromatic plants on rumen fermentation, protozoa population and methanogenesis in vitro. J
Anim Physiol Anim Nutr 97(3):446–456
Bhatta R, Saravanan M, Baruah L, Sampath KT, Prasad CS (2013c) In vitro fermentation profile
and methane reduction in ruminal cultures containing secondary plant compounds. J Appl
Microbiol 115(2):455–465
Bhatta R, Saravanan M, Baruah L, Prasad CS (2015) Effects of graded levels of tannin-containing
tropical tree leaves on in vitro rumen fermentation, total protozoa and methane production. J
Appl Microbiol 118(3):557–564
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Hegarty RS, Gerdes R (1999) Hydrogen production and transfer in the rumen. Recent Adv Anim
Nutr Aust 12:37–44
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approaches in enteric methane amelioration. Livestock production and climate change. In:
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Heat Stress on the Rumen Fermentation
and Its Consequence 18
Yutaka Uyeno

Contents
18.1 Rumen Microbial Community Overview . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214
18.2 Heat Stress and Rumen Acidosis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
18.3 Enteric Methane Emission and Mitigation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
18.4 Future Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221

Abstract
Ruminants are considered as one among the most sensitive animals to heat stress.
Very little information is available regarding the ruminal microbiota, playing
significant role in ruminant digestion, in connection with the physiological
changes associated with heat stress. A study was conducted by our team for
assessing the effects of temperature and relative humidity on the ruminal
microbiome composition using RNA-based method and the results were used
to effectively describe the predominant bacterial colony colonizing the rumen.
Four female Holstein animals (Heifers) were used in the study, with 2 weeks each
animal maintained in a climatic chamber under three different temperatures
(20–33  C). Ruminal fluid samples are collected from each animal on the
termination day. In this experiment it was observed that at elevated environmental
temperatures and humidity, body weight gain and dry matter intake were
decreased and feed digestibility was increased with increasing temperature.
Increase in the comparative populations of the genus Streptococcus and the
Blautia coccoides-Eubacterium rectale group and decrease in the genus
Fibrobacter’s population was observed as a response to rising temperature. It

Y. Uyeno (*)
Shinshu University, Nagano, Japan
e-mail: [email protected]

# The Author(s), under exclusive license to Springer Nature Singapore Pte 213
Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_18
214 Y. Uyeno

was also established that the ruminal microbiota was affected by various factors
such as age, diet and stress. These findings suggests that the ruminal eco-system
and its microbial community remain in a relatively unperturbed and matured state
throughout a longer duration, with typical daily fermentation patterns remaining
largely unchanged across the different stages.

Keywords
Heat stress · Holstein heifers · Metagenomics · Rumen · Rumen microbiota

Abbreviations

% Percentage

C Degree celsius
16S rRNA 16 Sedimentation coefficient rRNA
CO2 Carbon dioxide
CP Crude protein
DMI Dry matter intake
Fig. Figure
H Hydrogen ion
H2 Hydrogen
LPS Lipopolysaccharides
pH Potential of hydrogen
PS Persimmon skin
PSM Plant secondary metabolite
PSS Persimmon skin silage
RH Relative humidity
RNA Ribonucleic acid
rRNA Ribosomal RNA
SARA Sub-acute rumen acidosis
SCFA Short chain fatty acid
VFA Volatile fatty acid

18.1 Rumen Microbial Community Overview

The complex microbial ecosystem of rumen efficiently converts the carbohydrates

from plant source to organic acids. Any disturbances caused to this ecosystem will
negatively influence the host by causing diseases or impairing its productivity.
Factors such as farm management, feeding practices followed in the farm, and
importantly, composition of diet will severely affect the rumen’s ecosystem of
animal, producing negative effects on animal’s health and production. There is a
renewed interest in this complex system due to the development of various molecu-
lar strategies, with particular mention to the techniques that rely on 16S rRNA genes.
18 Heat Stress on the Rumen Fermentation and Its Consequence 215

These technologies do not only provide a phylogenetic framework of the

microbiome, but also in addition, they describe the impacts caused by environment
and host factors on the dynamics and structures of rumen microbiome. Eventually
the extreme molecular diversity exhibited by the bacterial community shall be
understood. Phylum Firmicutes and phylum Bacteroidetes are the ones under
which majority of the ruminal bacteria are affiliated (Uyeno et al. 2007). All
the fibrolytic bacteria that contributes to the characteristics of the organ, belong to
the lower taxa, such as Ruminococcus flavefaciens and Ruminococcus albus and the
genus Fibrobacter, and they also possess niches in the ruminal community.
Methanogens are exclusively represented by the archael community of the ruminal
microbiota, which is claimed for hydrogen removal. Methane generation, being the
crucial component involved in maintenance of the rumen ecosystem’s stability, is at
its peak of attaining attention worldwide, due to its major part played in contribution
to greenhouse effect and global warming. Mitigation strategies implemented to
reduce the enteric methane production have increased the energy deposition in the
ruminant’s body which made ruminal archaea to acquire an indispensable segment
of rumen’s ecosystem.
Over the years, innumerable numbers of attempts were made by the researchers
worldwide to manipulate the composition and functions of ruminal ecosystem for
improving the production of animals. Microbial colonization in the rumen that is
inherently harmful for the rumen microbiota is usually unsuccessful, because of the
robust nature of ruminal ecosystem. However, some common microbial colonization
also becomes harmful under certain conditions. For example, pH reduction caused
by the organic acids accumulation in the rumen that are absorbed little via the
ruminal epithelium. This condition is very commonly observed among the cattle
undergoing heat stress and the ones fed with excess amount of concentrates.

18.2 Heat Stress and Rumen Acidosis

The combination of excessively high temperature above the upper critical limit and
the higher relative humidity always causes heat stress in animals. Ruminants are
considered as one among the most sensitive animals to heat stress that is primarily
caused by various environmental factors like relative humidity, temperature, air
movement and solar radiation in combination (Collier et al. 2006). Metabolic heat
produced in the rumen during rigorous fermentation is the main reason for heat load
in case of cattle. The increased temperature attained from this process is nullified by
the means of heat loss. (Bohmanova et al. 2007). To get over heat stress, cattle
respond physiologically by reducing the exercise and feed intake. Since, mass
energy is spent to tackle heat stress effects rather than energy consumption, this
process in turn leads to deleterious effects on reproduction, milk yield and animal
health. In these occasions, ruminal ecosystem probably exhibits response to heat
stress. Changes produced at the group level of ruminal bacteria was monitored by
using the ruminal fluid. Total RNA was extracted from the collected ruminal fluid to
investigate the changes in bacterial community.
216 Y. Uyeno

Fig. 18.1 Effect of heat stress [ambient temperature and relative humidity (RH)] on the population
of specific bacterial groups in the rumen of Holstein heifers

Fig. 18.2 Effect of heat

stress [ambient temperature
and relative humidity (RH)]
on the rumen VFA level and
the composition of Holstein
heifers

Our study conducted using rumen-fistulated, four healthy female Holstein

animals of age between 9 and 15 months that are exposed to different temperatures,
showed reduction in body weight gain and dry matter intake (DMI) particularly with
consumption of forages and with increase in temperature (20–33  C) (Uyeno et al.
2010; Tajima et al. 2007). Profiling of rumen’s bacterial colony structure was
investigated. Increase in the genus Streptococcus, a major saccharolytic bacteria
(2.5–10 times) and decrease in the genus Fibrobacter, a major fibrolytic bacteria
(50–80%) was observed as a response to increment in environmental temperature
(Fig. 18.1).
Under heat stress, one can observe the consistent changes in the population level
of particular group of bacteria. It is the preferential increase in concentrates con-
sumption justifies the increase in activity of saccharolytic bacterial population and
reduction in activity of fibrolytic bacterial population. These changes in turn lead to
the notifiable reduction in concentration of SCFA (Fig. 18.2). This reduction in
concentration of SCFA could be an attributing factor for impairment observed in
growth performance of animal suffering under extreme conditions. Heat stress effect
on the fermentation can be indirectly explained by the reduction in pH caused by
increment in the activity of bacteria responsible for starch degradation. It is also said
18 Heat Stress on the Rumen Fermentation and Its Consequence 217

Fig. 18.3 A schematic flow

about how SARA occurs due
to changing feed

that, may be these bacteria undergoing changes may be small in population, still their
comparative population in the rumen holds an important position in deciding the
rumen’s adaptive response to heat stress. It is very vital to note that the diversity of
bacterial community as a whole at both species and genus level, remained intact even
under heat stress. If one group gets reduced under certain situation, meanwhile the
nullification for this reduction of certain group is taken care under different situation
and hence the diversity of ruminal bacterial community is maintained.
From the current understanding of ruminal physiology, it can be noted that the
combination of different strategies are required to optimize the animal’s production
under this hot and humid conditions. Various studies demonstrated the effects of
different diets on the composition and diversity of ruminal bacterial community.
Knowledge in this perspective is now good enough to have nutritional intervention
as a simple approach to tackle the heat stress related challenges. Reducing forage in
the ruminant’s diet can be a very simple and better way to increase the nutrient
density and decrease the adverse effects on productivity. In addition, this forage
reduction in the diet, also contributes for increase in the milk production of dairy
cattle undergoing heat stress in comparison with forage fed cattle, which can be
explained by decrement in the production of metabolic heat. But it should also be
understood that at times this feeding strategy may lead to increase in the population
of lactate producers (S. bovis) and henceforth drop in the pH of rumen will be
observed, because of the fact that the absorption of lactate is very much less than that
of VFAs by the ruminal epithelium.
There are few other conditions which is associated with ruminal pH reduction but
not linked to increase in lactic acid accumulation. Sub-acute rumen acidosis (SARA)
is one such condition, which often occurs with accompanying laminitis and liver
abscesses during the early and mid phase of lactation and is characterized by the
reduction in ruminal pH. Etiology of this condition is related to the factors like shift
in ruminal microbe population and immunological responses but not with lactic acid
accumulation in the rumen (Fig. 18.3).
218 Y. Uyeno

It is postulated that the moderate reduction in ruminal pH can increase the

lipopolysaccharides production of ruminal bacteria and also the permeation capacity
of LPS which then acts as endotoxins for triggering the systemic infection. Under
extreme conditions, to maintain the animal’s health and productivity, nutritional
intervention can be a key option to cater the negative effects on the rumen’s function.
Application of radio transmission pH sensing system was demonstrated to monitor
and assess the changes in pH of cow’s rumen in the transition period (Hasunuma
et al. 2016). Cows suffering from SARA in transition, early and mid phase of
lactation periods were successfully identified through consecutive monitoring of
changes in ruminal pH. For easy and better diagnosis of SARA in the on-farm basis,
this technique can be considered as a promising strategy to track changes in ruminal
pH. Acquisition of vast amount of data along with the study of relationship of
ruminal pH changes with various other factors can give better understanding regard-
ing the factors causing susceptibility and tolerance of animals to SARA in near
future.

18.3 Enteric Methane Emission and Mitigation

Various forms of carbohydrates are converted into VFAs by the complex ecosystem
of rumen to meet out the energy requirement of host through categorical disposal of
H by reducing carbon dioxide to CH4. It is in this step of reducing H, lies the primary
concept of manipulating methane generation, which is aimed at diverting this H
away from CH4 production. For instance, alternative pathways of metabolism to
divert this hydrogen like pathways generating propionate from succinate. Another
vital way is to reduce the H available for methanogenesis which can be done by
utilizing the alternate electron sinks. But equivocal results attained from the trails
conducted with this concept demands more information regarding the function,
composition and interactions taking place within this complex ecosystem of rumen.
In ruminants, methane production principally depends on the dry matter intake
(DMI), particularly fiber. The more carbohydrate digestion in the rumen leads to
more disposal of reducing power in the form of methane. An inverse relationship has
been described long back between the rumen’s methane and propionate production,
reflecting the competitiveness exhibited by both the processes in consuming the
reducing power. Three groups of ruminal bacteria were classified on the basis of its
H2 utilizing/producing ability: (1) bacteria producing lactate, butyrate, propionate
and/or ethanol, (2) bacteria producing hydrogen and acetate and, (3) archae produc-
ing methane (methanogenic and hydrogen users) (Fig. 18.4). The first classified
bacterial community theoretically undergoes fermentation completely without
requiring any disposal of hydrogen, which means that they don’t rely on methane
production and the second classification shows a marked dependency on third
classified bacterial community for effective fermentation. Some bacteria are potent
in utilizing molecular hydrogen for reductive acetogenesis, and even threshold for
this reaction is much higher than the methanogenesis resulting in very poor oppor-
tunity to alter methanogenesis with acetogenesis.
18 Heat Stress on the Rumen Fermentation and Its Consequence 219

Fig. 18.4 Schematic representation of fermentation of carbohydrates and categorization of bacteria

according to their demand for H2 utilization/production

Direct ruminal interventions aim to manipulate any of the steps, where possibility
appears for the mitigation of methane emission in ruminants. Several efforts have
focused on decreasing enteric methane emission through the addition of lipids, plant
compounds, monensin, and other organic compounds, or by otherwise controlling
diet composition. There is a blooming interest in the field of utilizing plant extracts
and plants itself for mitigating enteric methane emissions. Interest is gaining in
tropical parts of the world to use the plant secondary metabolites (PSM) for mitiga-
tion of livestock’s methane emission and to improve production performance of
livestock. Antimicrobial property of the PSM, makes it a better substitute for
artificial feed additives. Major classes of PSM are constituted by saponins and
tannins, which are under major area of research among multiple laboratories around
the globe (Bhatta et al. 2009). Tannins can bind with microbial proteins and feed and
hence it can be used to limit the substrate required by methanogens for methane
production and therefore as a better supplement to inhibit enteric methane produc-
tion without producing any undesirable effects on ruminal function (Patra and
Saxena 2011). Much more investigations are required to confirm the feasibility
and potential nature of tannins as additives in the feed of ruminants. Effects of
tannins are not limited to carbohydrate metabolism alone. Sufficient documentation
is available for the effect produced by tannins in the metabolism of nitrogen in
rumen. Tannins (phenolic hydroxyl group) react with proteins predominantly
through hydrogen bonding. Tannin-protein complex prevents the breakdown of
proteins by proteases. This bonding occurs at the ruminal pH which in turn affects
220 Y. Uyeno

the metabolism by ruminal microbiota. This characteristic of tannins can be utilized

in the production of rumen escape CP. This complex in turn can be reversed back in
abomasum for animal’s utilization and this strategy can play a significant role in
improving the availability of protein for the host. For more details, please refer to
other articles in this book.
A widely distributed fruit in the Asian continent, Persimmon scientifically named
as Diospyros kaki, occupies majorly the food industries of South Korea and Japan
and is produced more than 200 kilotonnes per year. Hoshigaki, dried form of
persimmon is consumed as such sometimes. The process of preparing Hoshigaki
involves peeling its skin, which is associated with the production of 20% of
persimmon skin (PS). This skin of persimmon is very rich in polysaccharides like
pectin and various soluble forms of carbohydrates, which makes this as a best
suitable source of energy for animals. To understand its effects on CH4 mitigation,
in vitro experiment (incubation study) was conducted using the unfermented and
ensiled PS (Mousa et al. 2019). Outcome of this experiment proved that the in vitro
production of methane was reduced by PS silage (PSS) and it was also found that this
reduction in methane generation is attributed to the carbohydrate composition and
changes in the composition of PSS. It can be concluded that the changes occurring
during this ensiling procedure is the sole responsible factor for the effect absorbed on
fermentation and methane production. Hence, by utilizing different dry absorbents
for adjusting the moisture in PSS, reduction in the effluent loss shall be achieved to a
trivial amount (Abdelazeem et al. 2020). Furthermore, sheep after consuming a
partially (upto 20%) substituted ration with PSS, showed no undesirable effects on
palatability and feed intake. Henceforth, detailed investigations along with intro
experiments (incubation study) are needed for determining the effects of PS on
animal’s feed efficiency and its impacts on enteric methane production and emission.

18.4 Future Perspectives

Many years of research were aimed at manipulation of composition and function of

rumen microbiota for improving its production. Even the minor changes in the
rumen’s microbiome composition have a potential impact on food intake, satiety
and energy expenditure of the animal. Therefore, it can be understood that the
animal’s health and its productivity is determined by its ability to prolong the
balance within rumen’s microbial ecosystem. Balance maintained in the rumen
ecosystem chiefly determines the animal’s health and its productivity. Hence, the
fundamental aspect of future research should aim at warranting the interactions
between the microbial community and its interaction with the host. The ruminal
ecosystem and its microbial community remain in a relatively unperturbed and
matured state throughout a longer duration. In order to link the ruminal microbiome
composition to the changing rumen metabolites, understanding the effects of differ-
ent factors, including diet, individual animal variability, and post-feed time, on the
rumen microbiome is warranted.
18 Heat Stress on the Rumen Fermentation and Its Consequence 221

Learning Outcomes
• Factors such as age, diet, and heat stress influence the microbial composi-
tion of rumen.
• Heat stress significantly increases the relative populations of the genus
Streptococcus, and the Blautia coccoides-Eubacterium rectale group
while decreases the genus Fibrobacter in cattle.
• 16S rRNA gene sequencing technology apart from providing the phyloge-
netic framework of the microbiota, can also provide an insight into the
impact of environmental and host factors on the dynamics and structure of
ruminal microbial community.
• Phytochemicals like tannin and saponins serves as best feed additives and
their antimicrobial property can be used to inhibit enteric methane produc-
tion in ruminants.

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analysis for evaluating the effect of heat stress on the rumen microbial composition of Holstein
heifers. Anaerobe 16(1):27–33
Comparative Assessment of Rumen
Microbial Diversity in Cattle and Buffaloes 19
Pradeep Kumar Malik, S. Trivedi, A. P. Kolte, Veerasamy Sejian,
A. Mech, S. Biswas, A. V. K. Bhattar, A. Mohapatra,
and Raghavendra Bhatta

Contents
19.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 224
19.2 Rumen Microbial System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
19.2.1 Bacteria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226
19.2.2 Archaea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227
19.2.3 Protozoa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227
19.2.4 Fungi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 228
19.2.5 Virome . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 228
19.3 Comparative Microbial Diversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229
19.4 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 234
19.5 Future Directions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235

Abstract

Cattle and buffalo are essential components of the livestock industry, which
contribute significantly to the economy. Globally, the rumen microbiota in
general, and methanogen community in particular in buffaloes, have received
less attention than cattle. The rumen microbiota has a strong correlation with the
digestion and metabolism of host diet. Furthermore, buffalo rumen exhibits
higher bacterial, fungal and protozoal populations compared to cows, indicating
different metagenome profiles. Firmicutes and Bacteroidetes are the most preva-
lent ruminal bacterial phyla in both buffaloes and cows, indicating the existence
of a core microbiome in ruminants. Diversity of microbes is much higher in
buffaloes compared to cattle. Between buffalo and Jersey cows, the methanogen
community was comparable, with Methanobrevibacter genus accounting for

P. K. Malik (*) · S. Trivedi · A. P. Kolte · V. Sejian · A. Mech · S. Biswas · A. V. K. Bhattar ·

A. Mohapatra · R. Bhatta
ICAR-National Institute of Animal Nutrition and Physiology, Bangalore, India

# The Author(s), under exclusive license to Springer Nature Singapore Pte 223
Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_19
224 P. K. Malik et al.

more than 90% of the total archaea. The protozoal community are more variable
than bacteria and archaea; Metadinium being higher in buffaloes, while the
Entodinium genus predominated in Jersey cows. Fungal diversity has been
found to be influenced more by the host rather than effect of diet. Genera
Caeocomyces, Cyllamyces, and Orpinomycesi have been principally found in
cattle while Aspergillus, Candida, or Kluyveromyces found to be highly abundant
in buffaloes. It is critical to characterize the ruminal microbiota differences
between buffalo and Jersey cows on a worldwide geographic scale for developing
innovative methods to their productivity while minimising environmental impact.

Keywords
Buffalo · Cattle · Metagenome · Methanogen · Microbiome · Rumen microbiota

Abbreviations

% Percentage
> Greater than
< Lesser than
CH4 Methane
CO2 Carbon dioxide
FAO Food and Agriculture Organization
GOI Government of India
MCP Microbial crude protein
mL Millilitre
pH Potential of hydrogen
RNA Ribonucleic acid
Tg Teragram
VFA Volatile fatty acid

19.1 Introduction

Cattle and buffalo are a vital part of the livestock sector, which contribute signifi-
cantly to the economy. Cattle are raised across the world, whereas buffalo, which
have evolved to tropical and subtropical temperatures, are confined in low-latitude
regions (Asai et al. 2021). India is endowed with a vast variety of 43 indigenous cow
breeds and 13 buffalo breeds that have endured for hundreds of years due to their
adaptability for certain functions in the local environment (GOI 2021a, b). Indian
livestock inventory now total 536.76 million head of livestock, including 193.46
million cattle and 109.85 million buffaloes (GOI 2019). The cattle and buffaloes
contribute 48% and 49% to the total annual milk production (198 million metric
tonnes) in India (GOI 2021a, b).
19 Comparative Assessment of Rumen Microbial Diversity in Cattle and Buffaloes 225

With the livestock sector’s fast growth, various challenges have arisen, including
low feed conversion efficiency, nitrogen use efficiency, product quality, and signifi-
cant methane emissions (Kumar et al. 2013). Ruminants face more of the above
challenges than monogastric animals do due to their unique digestive system, which
must be better understood in order to handle these concerns. Attempts to understand
and manipulate the rumen microbiome for the benefit of global agricultural
challenges have been ongoing for decades with limited success, owing primarily
to a lack of a detailed understanding of this microbiome and our inability to culture
the majority of these microbes outside the rumen.
There have been several reports claiming that buffaloes are better at digesting
fibrous foods than cattle, particularly diets rich in cellulose (Norton et al. 1979).
Buffaloes are more efficient at using high-roughage feeds (Lapitan et al. 2008); they
digest fibrous fractions, crude protein, and crude fibre more efficiently than cattle
(Robles 1971). Dry matter, organic matter, crude protein, neutral detergent fibre, and
acid detergent fibre digestibility were shown to be higher in buffalo than in cattle
(Chanthakhoun et al. 2012), showing clear variations in the rumen microbiome
composition or function between buffalo and cattle. Globally, the rumen microbiota
in general and methanogen community in buffaloes is particular less characterized as
compared to cattle (Malik et al. 2020). Thus, it is critical to characterise the ruminal
microbiota variations between buffalo and Jersey cows in order to formulate appro-
priate feed for them.

19.2 Rumen Microbial System

Rumen is one of the most diverse and complicated microbial habitats, containing
both prokaryotes and eukaryotes. Ruminants’ ability to utilise a wide variety of feeds
is a result of their highly diverse rumen microbial ecosystem, which includes
bacteria (1010–1011 cells/mL, representing more than 50 genera), ciliate protozoa
(104–106 cells/mL, representing 25 genera), anaerobic fungi (103–105 zoospores/
mL, representing five genera), and bacteriophages (108–109 cells/mL). These figures
might be substantially higher, considering the majority of them are unculturable
(Kamra 2005). Due to the enormous variety and functions assigned to rumen
bacteria, the rumen microbiota is recognised as a unique organ consisting of billions
of microbes with gene content hundreds of times that of host cells (Human
Microbiome Project Consortium 2012).
The rumen microbiota generates enzymes that efficiently break down complex
lignocellulosic plant components into simpler molecules that the host can easily
absorb. Vertebrates evolved without the ability to manufacture enzymes capable of
degrading cellulose and other complex polysaccharides. The primary benefit of
rumen bacteria is their capacity to absorb non-protein-nitrogen and plant
polysaccharides in a diet that cannot be digested and utilised by the host animal.
Several microbes in the rumen can secrete cellulase, which is a key enzyme to use
plant cell wall materials by ruminants. Rumen fermentation is carried out by bacteria
(two main phyla are Firmicutes and Bacteroidetes), fungi, archaea
226 P. K. Malik et al.

(Euryarchaeaota), and protozoa, which produce volatile fatty acids, vitamins,

microbial proteins (Hobson and Stewart 2012), and amino acids. Certain rumen
microorganisms also synthesise their own proteins (referred to as microbial crude
protein, MCP) for growth, employing the energy and nitrogen obtained from the
diet. The MCP are digested and absorbed by the host in the small intestine, therefore
contributing to the host’s nutrition also. Additionally, several bacteria are capable of
producing vitamins, such as vitamins B and K. Rumen microorganisms have several
enzymes involved in the de novo production of vitamin B12, which are absent from
the human gastrointestinal microbiome (Seshadri et al. 2018).
Carbon dioxide (CO2) and methane (CH4) are the end products of anaerobic
fermentation in the rumen, which are excreted into the environment (Morgavi et al.
2012). Enteric fermentation, which emits between 87 and 97 Tg of methane per year,
continues to be one of the primary sources of methane in agriculture (Chang et al.
2019). Globally, cattle and buffaloes contribute 77% and 13%, respectively, to
yearly enteric methane emissions (FAO 2021). The cattle and buffaloes contribute
48% and 49% to the total annual milk production (198 million metric tonnes) in
India (GOI 2021a, b). India is the world’s largest milk producer, owns 13% of the
worldwide cow population and 53% of the global buffalo population (GOI 2019),
which together account for 4.92 and 2.91 Tg of yearly global enteric methane
emission from the respective species (Bhatta et al. 2019). These two largest bovine
species together accounts for more than 85% of India’s total enteric methane output
(Malik et al. 2021). Methane is generated entirely by methanogenic archaea during
ruminal fermentation.
The rumen microbiota has a strong correlation with the digestion and metabolism
of host diet. Numerous prior researches have shown that one or more groups of
ruminal microbiota have an effect on feed efficiency, nitrogen digestibility, and
methane generation in ruminants (Schären et al. 2018). Rumen methanogenic
archaea produce methane mostly from the products of fermentation processes. It
becomes critical to understanding the roles played by the constituent microbes for
devising any strategies to manipulate the rumen microbiome to improve ruminant
production whilst reducing environmental impact.

19.2.1 Bacteria

Despite considerable advances in microbiology, the function of rumen bacteria and

their interactions with other members of the rumen microbiome remain poorly
understood, and as a result, there are only a few cases of direct modification of
this community’s composition yielding favourable results. Gram negative bacteria
prevail when animals are given a high forage diet, while Gram positive bacteria, such
as Lactobacilli, predominate when animals are fed a high grain diet (Hungate 1966).
Prevotella, Butyrivibrio, and Ruminococcus are the most abundant bacteria in the
rumen, and their community structure is influenced by the host’s diet (Henderson
et al. 2015; Iqbal et al. 2018). Fibrobacter succinogenes and Ruminococcus albus
are primarily responsible for cellulose breakdown, while Streptococcus bovis,
19 Comparative Assessment of Rumen Microbial Diversity in Cattle and Buffaloes 227

bovine amylolytic bacteria, is responsible for starch digestion (Matthews et al.

2019). According to studies, the shape of the microbial community may be deter-
mined by diet rather than animal type.
The feature of milk production is closely associated with the rumen microbial
composition, particularly with bacteria (Xue et al. 2020). Firmicutes:Bacteroidetes
ratio is substantially connected with milk-fat yield (Jami et al. 2014), but milk yield,
protein percentage, fat yield, and bacterial communities all have modest associations
(Bainbridge et al. 2016). Prevotella sp. and Succinimonas amilolytica were shown to
have favourable correlations with VFA concentrations in the rumen of high milk
producing cows.

19.2.2 Archaea

The archaea found in the rumen are classified as Euryarchaeota. These methanogens
produce methane in the rumen, which is subsequently eructed and released into the
environment, acting as a significant greenhouse gas. Methane is mostly created
through the hydrogenotrophic pathway, which involves the reduction of CO2, and
less often via the methylotrophic pathway, which involves the use of methyl groups,
or even less frequently from acetate (acetoclastic pathway) (Morgavi et al. 2010).
Though ruminal methanogens are not as diverse as bacteria, their necessity for a
different substrate makes this community complex (Malik et al. 2021). The primary
phylogenetic groupings of methanogens in the rumen include Methanobrevibacter
(>62% of methanogens), Methanosphaera (>15% of methanogens), and the newly
described Methanoplasmatales (>16% of methanogens). The remainder are classi-
fied as minor genera, including Methanimicrococcus, Methanosarcina, and
Methanobacterium (St-Pierre and Wright 2013). The Methanobrevibacter
gottschalkii and Methanobrevibacter ruminantium clades, as well as
Methanosphaera sp. and two Methanomassiliicoccaceae-related groupings with
variable abundance, dominate the archaeal community in all animal species world-
wide (Henderson et al. 2015).

19.2.3 Protozoa

Rumen protozoa account for a significant proportion of the rumen’s microbial

biomass (about 20–50% in certain circumstances); their involvement in ruminal
fermentation, metabolism, and nutrition of the animal remains a source of much
debate (Williams and Coleman 1992). They are, together with fungi, the eukaryotic
members of the rumen microbiota. Both flagellated and ciliated protozoa have been
described in the rumen, with ciliate protozoa (e.g. genera Diplodinium, Dasytricha,
Isostricyha, Eremoplastron, Entodinium, or Trichostomatia) being far more abun-
dant and belonging to two distinct orders: the Entodiniomorphida and the
Vestibuliferida or Holotricha (Williams and Coleman 1992).
228 P. K. Malik et al.

The rumen protozoa engulf organic materials, typically bacteria, and transport it
to digestive vacuoles where it undergoes hydrolysis and fermentation. Protozoa have
also been shown to boost the rumen pH faster than bacteria do, hence stabilising
rumen fermentation (Williams and Coleman 1992). As with other rumen bacteria,
certain ciliate protozoa are capable of digesting plant structural polysaccharides such
as cellulose and hemicellulose, whereas all ciliate protozoa are capable of engulfing
particulate matter. They can engulf and utilise starch grains, insoluble protein
and even microbial cells. Ciliate protozoa are capable of fermenting carbohydrates
and generating end products such as formate, acetate, propionate, butyrate, CO2, and
hydrogen. Defaunation, a process of protozoa elimination, lowers methane produc-
tion by around 11% (Morgavi et al. 2010). Rumen protozoa show variability in their
contributions to plant degradation and methane production. For example, Epidinium
spp. play a significant role in plant degradation (Huws et al. 2009), and holotrichs in
general encourage methanogens and methanogenesis (Belanche et al. 2015).

19.2.4 Fungi

Anaerobic fungi comprise 10–20% of the rumen microbiome and are critical in the
breakdown of low-quality forages (Krause et al. 2013). Fungi are one of the most
powerful fiber-degrading organisms known to science, owing to their large and
effective array of enzymes for the destruction of plant structural polymers (Solomon
et al. 2016). Additionally, their rhizoids are capable of physically penetrating plant
structural barriers (Orpin 1977), benefiting other rumen bacteria by expanding the
plant cell surface area available for colonisation. Six genera of fungi are found in the
rumen: the monocentric Neocallimastix, Caecomyces, and Piromyces, and the poly-
centric Anaeromyces, Orpinomyces, and Cyllamyces, each with 21 species, and the
newly discovered genera Oontomyces and Buwchfawromyces. As with the protozoal
population, rumen fungi’s close interaction with methanogenic archaea promotes
fungal activity and contributes to methane generation.

19.2.5 Virome

Virome is comprised of the bacteriophage and viral population. Bacteriophage are

obligatory pathogens of bacteria that exist in the rumen at a rate of around 107–
109 particles per gram of digesta (Klieve et al. 1996). Although the high prevalence
of bacteriophage indicates that they contribute significantly to the rumen system’s
equilibrium, they remain by far the least well-characterized component of the rumen
microbiome. The complete genome sequences of lytic phages, of the order
Caudovirales was obtained (Gilbert et al. 2017), that can infect Bacteroides,
Ruminococcus and Streptococcus. In the rumen, the majority of viruses are linked
with Firmicutes, Bacteroidetes, and Proteobacteria (Berg Miller et al. 2012). Recent
research has shown the occurrence of RNA-based viruses that infect fungus
(mycoviruses). However, there is a lack of sufficient molecular and physiological
19 Comparative Assessment of Rumen Microbial Diversity in Cattle and Buffaloes 229

information for this class of rumen microbe. The precise function of phages in the
rumen, their effect on fungal populations in the rumen, and fibre breakdown all need
more investigation to acquire a better understanding of their role and function in the
rumen.

19.3 Comparative Microbial Diversity

In India, the buffalo population is growing at a higher pace than cattle (Paul et al.
2018). Thus, it is critical to comprehend and compare their fermentation patterns and
microbial community composition in order to fully understand the variations and
effects of animal species on their gut microbial community. The rumen and reticu-
lum of grazing buffaloes contained a high quantity of bacteria and a low concentra-
tion of archaea (bacteria, methanogenic archaea, and ciliate protozoa). Buffalo had a
larger bacterial population and a higher Shannon diversity index than Jersey cows,
with the highest abundance of Bacteroidetes and Firmicutes (Iqbal et al. 2018).
Firmicutes population in the rumen is proportional to carbohydrate fermentation
(Fernando et al. 2010); hence, Firmicutes are always more abundant in the guts of
efficient feed consuming animals. Buffalo are more efficient in breaking down feed
nutrients than Jersey cows, owing to their larger Firmicutes population. Similar to
cows, buffalo rumen had a greater number of ruminal cellulolytic bacteria and
fungus (Chanthakhoun et al. 2012). This conclusion is bolstered further by the fact
that buffaloes have higher ruminal VFA concentrations (Iqbal et al. 2018).
Firmicutes and Bacteroidetes were the major ruminal bacterial phyla in both
buffalo and Jersey cow, confirming the existence of a core microbiome in the rumen
of ruminants (Henderson et al. 2015). Bacteroidetes were more abundant in the
rumen of cattle, which may account for their tolerance to high roughage feeding.
Prevotella was substantially less abundant in buffalo than in Jersey cows; however
Ruminococcus was significantly higher in buffalo. Prevotella is engaged in protein
breakdown and polysaccharide digestion, but Ruminococcus is highly effective at
cellulose and hemicellulose degradation (Flint et al. 2012). Bacterial diversity and
Shannon index were much higher in buffalo than in Jersey cows (Iqbal et al. 2018),
demonstrating that buffaloes have a greater rumen microbial diversity than cattle.
Ruminicoccus albus and Ruminicoccus flavefaciens were found in greater numbers
in the rumen of swamp buffaloes than in cattle (Chanthakhoun et al. 2012). R. albus
levels in rumen fluid were considerably greater in buffalo than in cattle, whereas
Ruminicoccus flavefaciens and Fibrobacter succinogenes levels tended to be higher
in cattle than in buffalo. The dissimilar microbiome profiles in cattle and buffalo
became more closely related after 16 weeks of similar dietary regime (Asai et al.
2021). The population of Fibrobacter succinogens was increased significantly in the
tannin and tannin + saponin supplemented groups as compared to control. In
contrast, the population of Ruminococcus albus and Ruminococcus flavifaciens
was decreased significantly ( p < 0.05) in the tannin + saponin (Ruminococcus
albus) and tannin (Ruminococcus flavifaciens) groups respectively, as compared to
230 P. K. Malik et al.

control; reporting differential response of fibrolytic microbes to tannins and saponins

supplementation.
The rumen archaeal population is profoundly influenced by the host physiology.
The geographical region and host have an effect on the structure of the archaeal
communities. Between buffalo and Jersey cows, the methanogen community was
comparable, with Methanobrevibacter genus accounting for more than 90% of the
total archaea (Iqbal et al. 2018). Six archaeal orders have been recognised in cattle,
whereas seven orders have been identified in buffaloes (Malik et al. 2021).
Methanogens related with the Methanocellales have been found solely in buffaloes.
Methanogens belonging to the Methanobacteriales were predominant in both hosts,
accounting for 72–73% of total ruminal archaea (Fig. 19.1a). Methanobrevibacter
was the most prevalent genus (55–62%) in both cattle and buffaloes, but
Methanobrevibacter gottschalkii constituted the largest proportion of the archaeal
community at the species level (Trivedi et al. 2020). However, other Indian studies
discovered that Methanomicrobium dominated the methanogen community in
buffaloes (Singh et al. 2011).
Methanotrophic Methanomassiliicoccales comprised 10% of the entire archaeal
community and had the greatest variation between cattle and buffaloes (hosts) and
geographical locations. Methanomassiliicoccales, including Methanosphaera sp.,
Group 4 sp., Group 8 sp., Group 12 sp., and Group 9 sp., were substantially more
abundant in cattle than in buffaloes. Thus, except for the presence of
Methanobrevibacter gottschalkii, the host species appears to have minimal effect
on the organisation of the rumen methanogens community. A meta-analysis (Paul
et al. 2018) demonstrated that the community structure of methanogens differs at the
genus and species level between cattle and buffaloes. Buffaloes had a larger abun-
dance of Methanomassiliicoccales, Group 10, and Methanobacterium alkaliphilum,
whereas Jersey cows had a lower abundance of Methanobrevibacter, demonstrating
that host species do have an effect on the rumen archaeal community (Iqbal et al.
2018).
The archaeal diversity between cattle and buffaloes fed on the same diet
(Fig. 19.2) demonstrated that Methanobrevibacter dominated the archaea commu-
nity (66–68%) and that its abundance was comparable across cattle and buffaloes
(Sirohi et al. 2013; Parmar et al. 2017; Malik et al. 2021). Additionally, Sulfolobus
thuringiensis was detected in Indian cattle and buffaloes, making this the world’s
second report and the first from India (Malik et al. 2021).
The Shannon (alpha diversity) and Bray-Curtis (beta diversity) indices at the
archaeal genus level in cattle and buffaloes are displayed in Fig. 19.3 and demon-
strate that the total diversity of ruminal methanogens was considerably different
between cattle and buffaloes. Additionally, the examination of the core methanogens
revealed a minor variation in the core microbiome of cattle and buffaloes.
Methanomicrobium was found only as a component of the cow archaeal core
microbiome. Methanogens such as Methanomicrobium mobile and
Methanobrevibacter wolinii were found to be present only in the cow archaeal
core microbiome (Fig. 19.4c). In contrast, Methanobrevibacter smithii was shown
 19
Comparative Assessment of Rumen Microbial Diversity in Cattle and Buffaloes

Fig. 19.1 Ruminal archaea community composition in cattle and buffaloes both at (a) order level and (b) family level (Malik et al. 2021). The inner-circle
represents the methanogens distribution in cattle; while the outer circle represents the distribution in buffaloes
231
232 P. K. Malik et al.

Fig. 19.2 Ruminal archaea community composition at genus and species levels in cattle and
buffaloes (Malik et al. 2021). Each genus is represented by larger bars that are underlaid on the
smaller bars representing the abundance of all the species affiliated to the corresponding genus

Fig. 19.3 (a) Alpha diversity and (b) beta diversity of the ruminal methanogens in cattle and
buffaloes (Malik et al. 2021)

to be the sole representative of the methanogens’ core microbiome in buffaloes

(Fig. 19.4d).
The protozoal community was more variable than bacteria and archaea
(Henderson et al. 2015), with more than 90% of samples including Entodinium
and Epidinium genera. Buffaloes have a substantially larger protozoal population
than cattle (Franzolin and Dehority 2010; Iqbal et al. 2018). The prevalence of
Metadinium was greater in buffaloes, but the Entodinium genus predominated in
Jersey cows. Epidinium and Dasytricha are the dominant genera in Zebu cattle,
19 Comparative Assessment of Rumen Microbial Diversity in Cattle and Buffaloes 233

Fig. 19.4 Ruminal archaea representing the core microbiome at 50% minimum prevalence in (a)
cattle at genus level (b) buffalo at genus level (c) cattle at species level (d) buffaloes at species level
(Malik et al. 2021). The colour gradient indicates variability in prevalence

whereas the Diplodiniinae subfamily is prevalent in the rumen fluid of buffaloes

(González et al. 2007). Entodinium are prevalent in cattle fed a variety of diets.
Numerous earlier investigations (Kittelmann et al. 2013) established Entodinium as
the dominating protozoa in cow and water buffalo rumen. Water buffaloes, on the
other hand, exhibit a high prevalence of Metadinium (73.2%) in their rumen (Iqbal
et al. 2018) Strong positive associations between Entodinium and M. gottschalkii
and Methanomassiliicoccales suggested a symbiotic relationship within the rumen
(Iqbal et al. 2018). Although the Epidinium genera (E. cuadatum and E. ecuadatom)
and D. crystagali are not found in cow rumens, they have been observed in buffalo
234 P. K. Malik et al.

rumens. Entodinium, Epidinium, Ophryoscolex, and Isotricha were found in greater

abundance in cows than in buffaloes (Jabari et al. 2014), suggesting that water
buffaloes have a greater rumen metabolic capacity than Holstein cows.
In the case of anaerobic gut fungi, it has been demonstrated that the host species is
critical in defining community composition, with the influence of the host being
greater than the effect of diet (Janssen and Kirs 2008). Lewia, Neocallimastix, and
Phoma were found to be more abundant in calves on a high grain diet, but
Alternaria, Candida Orpinomyces, and Piromyces spp. were found to be less
abundant (Ishaq et al. 2017). At the family level, Neocallimastigaceae was identified
with three major genera (Caeocomyces, Cyllamyces, and Orpinomyces).
Caecomyces and Cyllamyces collectively accounted for 25% of all anaerobic fungus
reported in cows (Kumar et al. 2015). Neocallimastix has the capacity to physically
or enzymatically degrade lignin plant cells, whereas Orpinomyces has the ability to
dissolve lignocellulose and produce an effective enzyme for hydrolyzing cellulose
and hemicellulose.
Cow anaerobic fungal communities were more variable than bacteria (73.2%),
archaea (77.9%), and even ciliate protozoa (41.4%) (Kittelmann et al. 2013). About
60% of Aspergillus, Candida, or Kluyveromyces were found in the rumens of all
buffaloes. Numerous investigations have detected Engyodontium and Sarocladium
in the rumen of buffaloes; certain members of Engyodontium have been reported to
possess protease capacity (Zhou et al. 2021). Sarocladium, on the other hand, is a
well-known phytopathogen that is specialised in secreting enzymes that destroy the
cell wall and other tough plant cellular components. The anaerobic fungi isolated
from buffalo rumen fluid have the potential to accelerate the decomposition of
lignocellulose in diet (Paul et al. 2004).

Learning Outcomes
• Bacterial diversity and Shannon index were much higher in buffalo than
cattle demonstrating that buffaloes have a greater rumen microbial diversity
than cattle.
• Both cattle and buffaloes have a comparable rumen methanogen commu-
nity, with Methanobrevibacter genus accounting for more than 90% of the
total archaeal population.
• Higher populations of bacteria, fungi and protozoa are found in buffalo
rumen when compared with cattle which indicates different metagenome
profiles exhibited by both the species.

19.4 Conclusions

There are few comparative studies on the rumen microbial ecology of buffaloes and
cattle. Numerous earlier researches comparing the microbiome compositions of
cattle and buffaloes found that buffaloes are more effective at using food due to
19 Comparative Assessment of Rumen Microbial Diversity in Cattle and Buffaloes 235

their higher rumen fermentation efficiency. Furthermore, buffalo rumen exhibits

higher bacterial, fungal and protozoal populations compared to cows, indicating
different metagenome profiles. However, studies indicate that the host has a minor
effect on the ruminal archaea population in cattle and buffaloes. These studies
provided novel and fascinating insights into different aspects of the structure,
function and activity of the rumen microbiome. The relationships between the host
genotype and the bacterial, fungal, protozoal, and viral populations inside the rumen
may be similar or variable.
Future research at molecular level on the rumen microbiome may offer informa-
tion on the role of host genetics in affecting these microbial communities. Funda-
mental knowledge of the rumen microbiome on a worldwide geographic scale is
critical for developing innovative methods for increasing animal productivity while
minimising environmental impact.

19.5 Future Directions

Over the last 10 years, a lot of effort has gone into figuring out what makes up the
rumen microbiome and how it influences the host’s growth and health. The study of
the rumen microbiome has primarily focused on the comprehensive study of rumen
bacteria alone, ignoring the other eukaryotic counterparts, which limits our ability to
understand the rumen microbiome. Future research focusing on other members of
the rumen microbiome might give information on how host genetics shape these
microbial communities. Furthermore, studies need to be focused on the influence of
species and breed genomics on microbiome composition and its implications on
breed specific physiological differences (nutrient digestibility, innate immunity,
stress adaptability) as well as product quality (milk meat etc.). Our knowledge of
how a stable rumen microbial community forms and what variables influence the
makeup and function of rumen microbial communities is limited. With research
showing that host genetics influences host immunity, it’s tempting to explore how
host genetics may alter rumen microbial community composition directly or indi-
rectly in ruminants.

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Metagenomics Application
in Understanding Rumen Functions 20
S. Trivedi, A. P. Kolte, Pradeep Kumar Malik, Veerasamy Sejian,
A. Mech, and Raghavendra Bhatta

Contents
20.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 241
20.2 Anatomy and Physiology of Rumen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 241
20.3 Rumen Microbiome . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 242
20.4 Why Metagenome? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 243
20.5 Understanding Complex Rumen System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 244
20.6 Rumen Metagenomics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245
20.7 Methods for Understanding Rumen Microbiome . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245
20.8 Culture Based Methods to Study Rumen Microbes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247
20.9 Culture Independent Approaches for Studying the Rumen Microbes . . . . . . . . . . . . . . . . 247
20.10 Phylogenetic Marker Genes in Microbes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 248
20.11 The 16S rRNA Gene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 248
20.12 Molecular Characterization Assays Based on 16s rRNA Gene for Community
Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249
20.13 Sequencing Technologies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 250
20.13.1 First Generation Sequencing Approach . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 250
20.13.2 Second Generation Sequencing Platforms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251
20.13.3 Third Generation Sequencing Platforms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 252
20.14 The Hungate1000 Collection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 253
20.15 Metagenomics in Ruminant Research . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 253
20.16 Whole Metagenome Analysis of Rumen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 254
20.17 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 255
20.18 New Frontiers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 256
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 256

S. Trivedi (*) · A. P. Kolte · P. K. Malik · V. Sejian · A. Mech · R. Bhatta

ICAR-National Institute of Animal Nutrition and Physiology, Bangalore, India

# The Author(s), under exclusive license to Springer Nature Singapore Pte 239
Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_20
240 S. Trivedi et al.

Abstract

Cattle and other ruminant food production is critical to the health of both humans
and the environment. A unique organ in the digestive system known as the
“rumen” allows ruminants to transform low-nutrient rich diet into food for
humans. As part of the digestion process, the rumen microflora is key to the
generation of essential nutrients for the animal. Rumen bacteria are the most
researched, however only 15% of the species in the rumen microbiota have been
cultivated in a laboratory. Metagenomics offers the ability to address fundamental
questions concerning the ecology of microorganisms. In the past, genetic finger-
printing techniques were used to study the structure and dynamics of microbial
communities. The advent of whole-genome metagenomics has opened new
avenues for studying the makeup of the microbiome and its relationship to the
metabolic processes involved in the digestion, absorption, and use of nutrients.
The rumen’s enormous diversity of metabolic capabilities may hold the key to
boosting the efficiency of animal production and industrial fermentation. A better
understanding of the animals’ microbial communities would allow for the modi-
fication of growth and other desired qualities, such as minimizing methane
emissions and extracting the most nutrients from their diets.

Keywords

Methane emissions · Metagenomics · Microbiota · Nutrients · Ruminants · Rumen

Abbreviations

16s rRNA 16 sedimentation coefficient rRNA

ARISA Automated ribosomal intergenic spacer analysis
Bp Base pair
CH4 Methane
DGGE denaturing gradient gel electrophoresis
DNA Deoxyribonucleic acid
FAOSTAT Food and Agriculture Organization Corporate Statistical Database
Gbp Giga base pairs
GC Guanine and cytosine
ITS Internal transcribed spacer
Kb KiloByte
KEGG Kyoto encyclopedia of genes and genomes
Ml Milliliter
mRNA Messenger ribonucleic acid
NGS Next generation sequencing
OUT Operational taxonomic unit
PCR Polymerase chain reaction
pH Potential of hydrogen
RdRP Ribonucleic acid polymerase
20 Metagenomics Application in Understanding Rumen Functions 241

RFLP Restriction fragment length polymorphisms

RNA Ribonucleic acid
rRNA Ribosomal RNA
SCFA Short chain fatty acid
SMRT Single molecule real time
Tbp Tera base pairs
TGGE Temperature gradient gel electrophoresis
T-RFLP Terminal restriction fragment length polymorphism
VFA Volatile fatty acids
ZMW Zero-mode waveguides

20.1 Introduction

Hosts and microbes can be thought of as a holobiont organism, with the microbiome
occupying various niches and interacting with the host in a symbiotic manner (Hall
et al. 2017). Complex features such as body mass index in humans or feed efficiency
in farm animals may be accurately predicted by the microbiome (Ross et al. 2013;
Camarinha-Silva et al. 2017). There are also associations between host genotype and
microbial composition in the gastrointestinal tract, confirming that particular diges-
tive niches are not only influenced by the environment and nutrition, but are also
impacted by the host’s genetic makeup. Food production from cattle and other
ruminants is crucial for human and environmental well-being. Over 296 million
beef and 273 million dairy cattle are included in the FAOSTAT database from 2013,
as well as an additional 468 million milk-producing ruminants (Stewart et al. 2018).
Though it has been studied extensively by both industry and the scientific commu-
nity, most of the rumen’s microbiota is still to be cultured. As a result, research into
how ruminants transform their food into energy and, ultimately, milk and muscle
protein, is gaining traction among livestock species. We may be able to produce
more food with less resources if we can enhance ruminant digestive efficiency,
which is an important objective in addressing global food security.

20.2 Anatomy and Physiology of Rumen

The digestion of tough plant tissues, which are particularly high in lignin and
cellulose, is a barrier that all herbivores must overcome. Relative to their gut
physiology, animals can be divided into three categories: ruminants, monogastrics,
and hind gut fermenters. The ruminant stomach is divided into four compartments:
the rumen and reticulum (which account for 84% of the total stomach volume), the
omasum (12%), and the abomasum (4%), each of which performs a specific role.
The stomach of a ruminant takes up nearly 75% of the abdominal cavity.
242 S. Trivedi et al.

Rumen Microbial fermentation takes place in the rumen, which acts as a fermenta-
tion vat. In the rumen, 50–65% of the starch and soluble sugar eaten is digested.
Rumen bacteria absorb cellulose from plant cell walls, digest complex starch, create
protein from nonprotein nitrogen, and produce B vitamins and vitamin K. The pH of
the rumen is usually between 6.5 and 6.8. The rumen is an anaerobic environment
(without oxygen). Carbon dioxide, methane, and hydrogen sulphide are among the
gases produced in the rumen.

Abomasum Also known as the “real stomach,” the abomasum is a part of the
digestive system. This compartment serves the same purpose as the stomachs of
other nonruminants, such as humans. The pH of the abomasum is usually between
3.5 and 4.0. The abomasum is where microbial and dietary proteins are digested.

Omasum The omasum is a spherical tissue fold that looks like a leaf and is attached
to the reticulum by a small tunnel. The many folds or leaves that mimic pages of a
book are referred to as the “many piles” or “butcher’s bible.” These folds enhance
surface area, which improves absorption of nutrients from feed and water. It also
serves as a filtration system, allowing only little amounts of fluid and fine particles
entering the abomasum.

Reticulum The reticulum gets its name from the honeycomb-like appearance of its
lining. Ingesta pass freely between the rumen and the reticulum because the reticu-
lum is not physically separated from the rumen. The reticulum’s primary role is to
gather smaller digesta particles and transport them to the omasum, while bigger
particles are regurgitated and rechewed with more saliva (a process known as
rumination).

20.3 Rumen Microbiome

Rumen is a fascinating complex microenvironment that includes a multi-kingdom

community of 1011 bacteria cells/ml, 104–106 protozoa cells/ml, 103–106 fungi cells/
ml, 106 cells/ml methanogens, and 107–1010 particles/ml bacteriophages (Morgavi
et al. 2013), all of which benefit each other in a symbiotic relationship. These
microbes that thrive in a strict anaerobic environment perform an important and
necessary activity called anaerobic fermentation, which involves the destruction of
complex plant polysaccharides such as cellulose, hemicellulose, lignin, and pectin,
which are indigestible to humans. There is a sequential syntropic connection of
microflora, comprising bacteria, protozoa, fungi, archaea, and bacteriophages, as
well as host, for completing complete feed digestion. Cellulolytic bacteria break-
down polysaccharic plant fibres, anaerobic fungi degrade lignocellulosic
components of feed particles, and the resulting substrate is acted upon by other
bacteria. As a result of these microbial action, volatile fatty acids (VFAs) such as
acetate (used in fat synthesis), propionate (used in glucose production), and butyrate
are produced. These VFAs are later used as a source of energy by the animal.
20 Metagenomics Application in Understanding Rumen Functions 243

Fig. 20.1 Functions of rumen microbiome

Methane (CH4), one of the end product gases, is a powerful greenhouse gas. The
health of the local communal flora determines overall energy production as well as
digestion. Rumen microbiome perform a variety of functions (Fig. 20.1) that benefit
the host.

20.4 Why Metagenome?

It was long widely believed that organisms isolated in pure culture from their natural
environment represented numerically and functionally important species. The
organisms obtained using traditional culture techniques, on the other hand, were
seldom dominant. Due to their capacity to thrive on nutrient-rich medium at a
moderate temperature and under aerobic circumstances, these organisms have been
isolated (Hugenholtz 2002). The culturable species make up less than 1% of the
overall diversity in some environments. This phenomenon was previously known as
the “great plate count anomaly,” and the unculturability could not be confirmed until
molecular technologies arrived. To discover uncultured bacteria in the environment,
ribosomal RNA and rRNA genes were sequenced. This method was primarily used
to reconstruct phylogenies, compare microbial distribution in samples using
sequencing or RFLP, and quantify relative abundance of each taxonomic group
using hybridization with probes unique to that taxonomic group.
Carl Woese proved in the late 1980s that the 16S ribosomal ribonucleic acid
(rRNA) gene could be used to establish prokaryote phylogeny (Woese 1987). A
similar version of this gene was found in all species. Woese used the gene sequence
to classify living things into three categories: Eukaryote (multicellular), Archaea and
244 S. Trivedi et al.

Bacteria (singe celled). Norman Pace demonstrated that the polymerase chain
reaction (PCR) may be used to isolate 16S rRNA genes from the environment.
Most taxa that underwent culture-independent study were new, indicating that there
are thousands of species yet to be identified. However, PCR analysis can only
amplify the genes of interest in organisms, leaving the remainder of the genome
unaccounted for. Microbiologists need a new technique to study genetics and
physiology, which prompted the development of the science of metagenomics
(Handelsman 2004). Scientists may use metagenomics to examine the genetic
material of whole microbial populations. This area detects and classifies genetic
material in environment by combining genetics and molecular biology. The micro-
bial research in many ecological niches have been fueled by the sequencing technol-
ogy and infrastructure created during the human genome sequencing project.

20.5 Understanding Complex Rumen System

Recent research on rumen/gut microorganisms has demonstrated that the rumen

microflora is vital in the degradation of ingested feed as well as the production of
key nutrients. The rumen microflora is dynamic, changing with the animal’s age and
species, responding to feeds and environmental conditions, and the imbalance/
dysbiosis has an impact on the animal’s nutrition and health. Robert Hungate, the
“Father of Rumen Microbiology,” pioneered the creation of anaerobic bacterial
growth techniques that are still the “gold standard” method across the world
(Hungate 1966). However, due to the rumen microorganisms’ complicated substrate
requirements, adequate cultures could not be grown. As a result, for a long period, a
significant portion of the rumen microbial diversity was unknown (Krause et al.
1999). The rumen microbial communities have never been thoroughly explored due
to the inherent difficulty in culturing and research of these microbes. Despite the fact
that rumen bacteria are the most studied, only 15% of the species in the microbiota of
the rumen have been cultured in the lab (Morgavi et al. 2013; Creevey et al. 2014).
There are now at least 70 rumen bacterial species accessible in pure culture from
public sources (Creevey et al. 2014).
Despite the development of culture-based approaches, cloning and sequencing of
the 16S rRNA gene-based identification of rumen bacteria is a more potent tool for
identifying and characterising community members in greater depth (Sundset et al.
2007). It is possible to study the rumen microflora at several taxonomic levels thanks
to the advent of high-throughput, culture-independent technologies. Understanding
the rumen microbiome composition will offer new possibilities for improving
production, disease resistance, novel enzyme discoveries, sustaining health, and
reducing methane emissions from ruminal fermentation in ruminants. The rumen
microbial community’s composition and evolution in response to a variety of
selection forces have been widely researched (Huws et al. 2018).
20 Metagenomics Application in Understanding Rumen Functions 245

20.6 Rumen Metagenomics

Even if steps have been taken to cultivate various ruminal microbiota members, there
are still a large number that have not yet been characterised (Seshadri et al. 2018).
The rumen microbiome holds approximately 100 times more gene pool as compared
to the host (McSweeney and Mackie 2012). Therefore, the total gene pool of the
microbiome is also regarded as “Second Genome of the animal” (Bath et al. 2013).
In order to get a thorough understanding of the genetic structure, population dynam-
ics, and metabolic processes of these uncultured bacteria in the environment,
including the rumen, sophisticated culture-independent approaches must be devel-
oped. Because of metagenomics, the rumen microbiota’s whole genetic repertoire
may be accessed without the time-consuming culture procedure (Glendinning et al.
2021). The metagenomics has the potential to provide answers to basic concerns
about microbial ecology. Bacterial and archaea genome sequencing has transformed
our understanding of microorganisms’ diverse roles.

20.7 Methods for Understanding Rumen Microbiome

Bacteria, archaea, protozoa, fungus, yeast, mycoplasma, and bacteriophages are all
part of the rumen’s microbial community. Anaerobic fermentation of ingested feeds
takes place because of their interaction and participation in the detoxification of
many anti-nutritional components as well as some toxic substances. As a result,
ruminant nutrition relies heavily on microorganisms. Light and electron microscopy
were used to study the microbial members, although only 11% of the overall variety
of microbes present in the rumen was known to have been studied (Wolin et al. 1997;
Edwards et al. 2007). For the identification and quantification of rumen microbiota,
molecular approaches employing the 16/18s rRNA gene and it’s associated ITS
region were developed. A culture-free approach to microbial community structure
and function is provided by metagenomics. A variety of PCR techniques, including
clone libraries, terminal restriction fragment length polymorphism (T-RFLP),
denaturing gradient gel electrophoresis (DGGE), ARISA (Welkie et al. 2010),
amplicon sequencing, and shotgun metagenome sequencing, have been used to
study the diversity. The final two strategies rely significantly on genetic sequencing
and computational technologies. To better understand the structure and composition
of microbial communities, 16s rRNA gene amplicon sequencing can be used. Whole
metagenome shotgun sequencing, metatranscriptomics, and metaproteomics can be
used to discover the functional capacities of the metagenome. Different approaches
for studying metagenomics have been represented in Fig. 20.2.
 246

Fig. 20.2 Different approached used for study of rumen microbiome

S. Trivedi et al.
20 Metagenomics Application in Understanding Rumen Functions 247

20.8 Culture Based Methods to Study Rumen Microbes

The creation of the anaerobic glove box for decreasing the exposure of the medium
and bacteria during inoculation and subculturing improved the early attempts for
cultured the rumen microbes (Aranki et al. 1969). The roll tube method of anaerobic
microbe culture was shown to be preferable for researching anaerobic bacteria and
was used to examine rumen microbes (Hungate 1960). Using traditional approaches,
10–11% of the entire microbial species might be cultured (Jami et al. 2013; Millen
and Romero 2016). Around 200 species of bacteria, 100 species of protozoa, and
100 species of fungus have been identified from the rumen using traditional culture
methods (Williams and Coleman 1997; White et al. 1999). Due to the unavailability
of universal media that could sustain the growth of the majority of species, these
approaches were time consuming and difficult (Kamra 2005). Although genetic
identification of rumen bacteria has been widely used to research rumen microbial
composition, culture-based investigations will be critical for isolating new species
and understanding the physiological and genomic characteristics of specific
microbiome members. To extract new rumen bacteria, a liquid culture based on
the dilution strategy (also known as extinction culturing method) was devised and
successfully proved to grow a high fraction of hitherto unculturable rumen
microbiota members (Kenters et al. 2011).

20.9 Culture Independent Approaches for Studying the Rumen

Microbes

Due to severe constraints in cultivating microorganisms in anaerobic niches such as

the rumen and other biological sites, techniques that do not need growing microbes
to identify them were necessary. In many ecosystems, DNA identification of
microorganisms has revealed that these bacteria are seldom dominant (Hugenholtz
2002). In some ecological niches, the fraction of cultivable organisms is less than
1–10% of overall microbial diversity (Prakash et al. 2013). The causes for environ-
mental species unculturability are several. The first is a lack of understanding of the
exact growth requirements, or they may grow at a much slower rate in the laboratory
than we anticipate. The laboratory media could be toxic for the microbes, or they
simply cannot exist alone, and may grow only in communities. The uncultivability
could not be established until the development of molecular technologies for micro-
bial identification, which was previously known as “the great plate count anomaly.”
To detect uncultured microorganisms in the environment, sequencing of phyloge-
netic marker genes was developed. The invention of the polymerase chain reaction
(PCR), DNA sequencing technology, and the use of the 16S rRNA gene as a
taxonomic identifier (Woese 1987) have made gene-based identification of
prokaryotes in various contexts much easier.
Because of the evolutionary conservation of 16S ribosomal RNA, molecular
approaches based on 16S rRNA may be used to identify and quantify the abundances
of bacteria without the need to culture them. The 16S rRNA approach had revealed
248 S. Trivedi et al.

the third microbial kingdom ‘Archaebacteria’ (Woese et al. 1990). Total DNA
extracted from ambient samples, as well as amplification, cloning, and sequencing
of the 16S rRNA gene using universal degenerate primers (Rappé and Giovannoni
2003), gave a window into the uncultured world. Several reviews and technical
guides have published methods for understanding the rumen microbial population
(Zoetendal et al. 2003, 2004, 2008; Makkar and McSweeney 2005; McSweeney
et al. 2006; Morgavi et al. 2012).

20.10 Phylogenetic Marker Genes in Microbes

In the past, housekeeping genes were employed to detect and categorize

microorganisms. However, most of these genes were effective in identifying previ-
ously discovered bacteria. The research on the yet-to-be-cultured bacteria
necessitated the use of a highly conserved gene or genes that could be targeted
using PCR amplification and sequencing. Universality, conserved function across
organisms, adequate gene length for discriminating, absence of horizontal transfer,
and availability of high-quality sequencing databases are all desired characteristics
of the gene. The 16S rRNA gene, out of all the known genes, contains all the
necessary characteristics and has been widely employed in microbial molecular
phylogeny. Today, different genes are used to identify different classes of microbes,
such as 16S rRNA for bacteria and archaea (Rampini et al. 2011; Salipante et al.
2013), 18S rRNA for eukaryotes, ITS regions for fungi (Wagner et al. 2018), and
RNA dependent RNA Polymerase (RdRP) for RNA viruses (Culley et al. 2010).

20.11 The 16S rRNA Gene

Ribosomes, which are made up of RNA and proteins, are essential components of all
living forms. The ribosome includes the 5S, 16S, and 23S RNA molecules in
prokaryotes and archaea. However, 16S rRNA was widely utilized in microbial
phylogeny owing to its size advantage (Konstantinidis and Tiedje 2007). The 16S
rRNA gene transcript is a component of the small ribosomal subunit, which is
responsible for translating messenger RNA (mRNA)-coded information into
proteins. Variable regions inside the gene were produced because of the accumula-
tion of evolutionary mutations in regions that did not impact ribosome function. In
the late 1980s (Lane et al. 1985), the 16S rRNA gene was originally proposed for
phylogenetic study of bacteria, and it has since been the method of choice for
molecular identification of isolates as well as characterizing microbial communities
(Tringe and Hugenholtz 2008). The 16S rRNA gene is 1500 bases long and has nine
hypervariable regions (Fig. 20.3) separated by conserved sections (Baker et al.
2003).
Because of the conserved sections of the gene, universal primers may be designed
for PCR amplification of the gene across all bacterial members of the community.
Several works have reviewed the design and assessment of primers for different
20 Metagenomics Application in Understanding Rumen Functions 249

Fig. 20.3 Schematic diagram of 16S rRNA gene showing the distribution of the conserved and
hypervariable regions within the gene

variable areas, as well as the comparison of the region for resolving the taxonomy
(Klindworth et al. 2013).

20.12 Molecular Characterization Assays Based on 16s rRNA

Gene for Community Analysis

Due to the diversity of ruminal microorganisms, genetic fingerprinting techniques

have been utilized in the past to examine population structures and dynamics.
Typically, the fingerprinting process begins with the collection of community
DNA, followed by PCR amplification of the assay’s target gene and gel electropho-
resis examination of the PCR results. Comparing almost full 16S rRNA gene
sequences has been extensively utilised to identify Bacteria and Archaea and to
establish taxonomic relationships among prokaryotic strains (Kim and Chun 2014).
The Terminal Restriction Fragment Length Polymorphism (T-RFLP) approach is
used to investigate bacterial diversity. It is based on variation in the 16S rRNA gene,
which may be exploited by restriction enzymes to create terminal restriction
fragments of varying length. The 16S rRNA gene is amplified using primers that
are fluorescently labelled. Tetra cutter restriction enzyme is used to digest the PCR
products. Capillary electrophoresis is used to separate the fragments, and the frag-
ment lengths are analyzed to determine the phylotypes. Based on the length of the
terminal fragment, a tentative identification can be made by comparing the fragment
length to a database of virtual digests of reference 16S rRNA sequences. The 16S
rRNA gene was utilized to detect microbial communities in the reticulo-rumen
(Fernando 2008; Li et al. 2009). The rumen microorganisms of dairy sheep were
examined using TRFLP during sunflower and fish oil supplementation. The
approach demonstrated a varied response of Lachnospiraceae members to lipid
feeding. TRFLP demonstrated that these bacteria do not play a dominating role in
the process, although uncultured members may be more significant (Belenguer et al.
2010). Tannin supplementation in lambs resulted in distinct clustering of TRFLP
patterns (Vasta et al. 2010). The approach was effectively utilized to investigate the
influence of breed on the microbial profiles of sheep guts, revealing that the bacterial
populations in the foregut, midgut, and hindgut were significantly different (Douglas
et al. 2016).
Denaturing Gradient Gel Electrophoresis (DGGE) and Temperature Gradient Gel
Electrophoresis (TGGE) are two types of electrophoresis that employ a temperature
or chemical gradient to separate denatured nucleic acids. DGGE/TGGE are primarily
based on the principle of amplifying rRNA or functional gene PCR products isolated
250 S. Trivedi et al.

from community DNA using primers that contain a 50-bp GC-clamp and then
separating them on polyacrylamide gels (Rastogi and Sani 2011). DGGE separates
identical gene sequences based on their differential denaturing ability, which is
dictated by their base pair sequence, whereas TGGE separates nucleic acids based
on temperature-dependent changes in structure. Additionally, Denaturing Gradient
Gel Electrophoresis (DGGE) and Automated Ribosomal Intergenic Spacer Analysis
(ARISA) were utilized to characterize the rumen bacteria (Petri et al. 2012).
According to the DGGE profiles, animal to animal variability in the ciliate popula-
tion was highest in sheep compared to deer and cattle (Kittelmann and Janssen
2011). The microbiome of the sheep mammary gland found that 15 DGGE bands
were strongly linked with changes in somatic cell count (Monaghan 2013).
While full-length amplification and sequencing of the 16S rRNA gene is ideal for
molecular taxonomic identification of bacteria, restrictions imposed by sequencing
methods and universal primers limit its application in community characterization. It
is established that the various variable regions have diverse taxonomic resolutions,
and the proper variable region selection is dependent on the assay and the source of
the samples. Different primers produce distinct fingerprints on DGGE gels (Yu and
Morrison 2004). The studies based on amplicon sequencing and primer selection do
skew the evaluation of community composition, resulting in under or over represen-
tation of specific taxonomic groupings (Yang et al. 2016). Nonetheless, given the
cheap cost per sample and the requirement of low template DNA concentrations, 16S
rDNA sequencing continues to be one of the most popular high-throughput sequenc-
ing technologies (Cao et al. 2017).

20.13 Sequencing Technologies

20.13.1 First Generation Sequencing Approach

The sanger sequencing approach, which is based on capillary electrophoresis, can

sequence the whole 16S rRNA gene (1500 bp), but not in a single run and requires
the use of two or more primer pairs. While the approach is quite precise, it falls short
of providing the depth of coverage necessary for studying microbial communities
(Mitreva 2017). This led in difficulty recognizing rare species and insufficient
coverage of microbial diversity. Additionally, examining microbial communities
with Sanger sequencing is time consuming and expensive in comparison to next
generation sequencing platforms (NGS) (Sorek et al. 2007). While next-generation
sequencing systems are extremely scalable, cost effective, and capable of concur-
rently sequencing millions of copies of a gene, they suffer from sequencing length
constraints, which limit the identification of bacteria to lower taxonomic ranks. To
increase taxonomic resolution, several variable regions of the 16S rRNA gene were
sequenced using next-generation sequencing technology, and various primers were
described and assessed (Soergel et al. 2012).
20 Metagenomics Application in Understanding Rumen Functions 251

20.13.2 Second Generation Sequencing Platforms

In 2005, the 454/Roche sequencing platform was launched. The platform has been
extensively utilised in the field of microbial ecology (Tringe and Hugenholtz 2008).
The 454 platform use emulsion polymerase chain reaction to clonally amplify DNA
fragments that were sequenced in parallel using pyrosequencing technology.
Although the platform had limitations in terms of the introduction of artificial
replicate sequences and homopolymer sequencing errors that resulted in indels, it
provided 10–100 fold greater diversity coverage at a 10–100 fold lower cost than
previous methods of cloning and sequencing metagenomic libraries (Tamaki et al.
2011). The FLX+ platform yielded an average sequence length of 600–800 bp,
which was adequate to cover more than one variable region in the 16S rRNA gene.
Multiplexing of samples was introduced utilizing barcoded primers to increase
throughput above the requirement for a single ecological sample. The platform
was used to conduct microbiological surveys in various habitats (Dinsdale et al.
2008).
The Illumina/Solexa technology is based on sequencing by synthesis and was
released in 2007 with a data output of 1Gbp (Giga base pairs), which was later
increased to 1.8 Tbp (Tera base pairs) in the Illumina HiSeq systems. Initially, the
platform yielded shorter sequences than the 454/Roche platform, but with better
chemistry, the cost of sequencing has decreased significantly (Di Bella et al. 2013),
allowing for faster throughput and longer read lengths of up to 250–300 bp. The
Illumina platform’s primary strength was paired end sequencing, which involved
sequencing a bit of DNA from both ends, allowing the platform to cover 600 bases
for each piece of DNA. The Illumina MiSeq benchtop sequencer is used to perform
the 300 bp paired end chemistry.
With shorter read lengths acquired from Illumina platforms, it is not possible to
correctly classify bacteria beyond the genus level using 16S rDNA amplicon
sequencing (Claesson et al. 2010). Another drawback of this strategy is the selection
of nine hypervariable regions (V1–V9) inside the 16S rDNA gene. Numerous
previous investigations have employed various hypervariable areas, including
V1/V2/V4 (Sundquist et al. 2007), V2/V3/V4 (Liu et al. 2008) V2/V4 (Wang
et al. 2007) and V2/V3 (Chakravorty et al. 2007). PCR amplicon fragments as
short as 82 bp targeting the 16S rDNA V5 variable region have been shown to be
sufficient for phylum-level bacterial classification (Lazarevic et al. 2009), and a
longer amplicon fragment of 100 bp combined with appropriate primer design and
downstream analysis could display the same clustering information as a longer 16S
rDNA sequence. Even with longer variable regions, pyrosequencing, and increased
coverage, amplification of various polymorphic regions resulted in a bias in
estimating the microbiome. Despite this, 16S rDNA sequencing remains one of the
most common high-throughput sequencing technologies due to its low cost per
sample and necessity for low input template DNA concentrations.
252 S. Trivedi et al.

20.13.3 Third Generation Sequencing Platforms

One of the most widely utilised third-generation sequencing methods is PacBio’s

SMRT (Single Molecule Real Time) sequencing. PacBio long-read sequencing,
enabled by SMRT Sequencing technology, requires no PCR amplification and has
a read length 100 times longer than NGS. A unique flow cell with hundreds of tiny
picolitre wells with transparent bottoms—zero-mode waveguides—is used for
SMRT sequencing (ZMW). The polymerase is attached to the well’s bottom and
permits the DNA strand to pass through the ZMW. Real-time imaging of fluores-
cently tagged nucleotides produced alongside individual DNA template molecules is
possible with SMRT sequencing. When the template and polymerase separate, the
sequencing process is complete. The PacBio instrument’s average read length is
about 2 kb, with some readings exceeding 20 kb. Longer reads are very advanta-
geous for de novo genome assembly.
The PacBio method is being used to unravel the puzzle of the complex rumen
microbiome. Myer et al. (2016) compared full-length 16S rRNA reads (V1–V8)
sequenced with PacBio to shorter V1–V3 reads sequenced using Illumina Mi-Seq to
categorize rumen community members in greater depth. While the two platforms
revealed similar microbial OTUs, species richness, Good’s coverage, and Shannon
diversity metrics, the Pac-Bio platform improved taxonomic depth. Another study
used the Pacific Biosciences single-molecule real-time (PacBio SMRT) sequencing
technique to assess the intestinal microbiota of nine dairy cows during lactation
period (Li et al. 2018). Firmicutes (83%) was the most common phylum, while
Bacteroides (6.16%) was the most common genus. All the samples included a lot of
microbial diversity, including a lot of short chain fatty acid (SCFA) producers. The
research shed new light on the relationship between dairy cow gut microbes and milk
production.
Nanopore sequencing is a scalable, yet another technique for sequencing long
DNA or RNA segments in real time. It functions by tracking how an electrical
current changes when nucleic acids flow through a protein nanopore. The resultant
signal is decoded to reveal the DNA or RNA sequence in concern. Flow cells, which
include an array of microscopic holes—nanopores—embedded in an electro-
resistant membrane, are used in all Oxford Nanopore sequencing equipment. Each
nanopore has its own electrode, which is connected to a channel and sensor chip that
detects the electric current flowing through it. The current is disturbed when a
molecule travels through a nanopore, resulting in a distinctive’ squiggle.’ The
DNA or RNA sequence is then determined in real time by decoding the squiggle
using base calling algorithms. MinIONTM, GridIONTM, and PromethIONTM are
nanopore systems; MinIONTM is a tiny and portable device introduced in 2014.
Nanopore sequencing has the potential to create ultra-long reads, with read lengths
exceeding 2 megabytes, allowing for a more comprehensive assessment of genetic
variants and the reconstruction of complicated genomes.
A comprehensive assessment of over 6.5 terabytes of sequencing data from
283 cattle’s rumen was used to create the rumen genome catalogue, which contains
5845 assembled rumen microbial genomes (Stewart et al. 2019). The study
20 Metagenomics Application in Understanding Rumen Functions 253

unravelled a metagenomic assembly that included at least three entire bacterial

chromosomes as separate contigs demonstrating the ability of long reads for assem-
bling full, entire chromosomes from complicated metagenomes. The nanopore
assembly was 268 kb, which was more than 56 times longer than the typical Illumina
assembly (4.7 kb). Delgado et al. (2019) used the MinION (Nanopore) and Illumina
MiSeq to sequence the rumen content of 12 cows selected for high feed efficiency
phenotype. Although Nanopore sequencing produced similar findings as Illumina
sequencing, it was able to categorise a greater number of observations at the species
level. The findings also suggest that, due to the size of the reads, the same amount of
information might be obtained in a shorter duration of time.

20.14 The Hungate1000 Collection

Seshadri et al. (2018) compiled the most comprehensive reference genome library
till date, consisting of 410 cultured bacteria and archaea, as well as their reference
genomes, representing major rumen-associated archaeal and bacterial families.
Illumina or PacBio technology was used to sequence all the Hungate1000 genomes.
They investigated polysaccharide breakdown, short-chain fatty acid synthesis, and
methanogenesis pathways, as well as attributing functions to specific taxa. 336 spe-
cies were found in rumen metagenomic data sets, whereas 134 were found in human
gut microbiome data sets. 75% of the rumen’s genus-level bacterial and archaeal
species are represented in the Hungate genome collection. The Hungate Collection
serves as the basis for designing rumens to reduce CH4 emissions, boost productiv-
ity, and sustainability (Mcallister et al. 2015), offering insight on what has been
termed “the world’s biggest commercial fermentation process” (Weimer 1992).

20.15 Metagenomics in Ruminant Research

Pyrosequencing was utilized to examine the bacterial community composition in the

rumen of dairy cows and their relationship to physiological factors. According to the
study, several physiological indicators, such as milk production and composition,
are strongly connected with the prevalence of specific bacteria found in the rumen
microbiome. A notable discovery was a substantial link between the proportion of
Firmicutes to Bacteroidetes and milk fat output (Jami and Mizrahi 2012). The
combined use of amplicon, shotgun, and metatranscriptome sequencing revealed
an enhanced population of sharpea in sheep that produce less methane. The
microbiome of high methane emitters was dominated by members of the
Ruminococcaceae and Lachnospiraceae families. The study revealed that the low
methane emitter sheep microbiome will produce roughly 24% less methane. The
rumen size was lower in low emitters than in high emitters (Kamke et al. 2016). In
both high and low methane emitters, deep metagenomic and metatranscriptomic
sequencing revealed a similar quantity of methanogens and methanogenesis pathway
genes. However, transcription of genes involved in the methanogenesis pathway was
254 S. Trivedi et al.

significantly elevated in sheep producing large amounts of methane (Shi et al. 2014).
The global rumen census employed 16S rRNA sequencing of rumen samples from
742 animals from eight different ruminant species all around the world to try to
figure out the core rumen microbiome (Henderson et al. 2015). The predominant
rumen bacteria were Prevotella, Butyrivibrio, and Ruminococcus, as well as unclas-
sified Lachnospiraceae, Ruminococcaceae, Bacteroidales, and Clostridiales, which
may represent a core bacterial rumen microbiome. Large collections of rumen
microbial genomes have been revealed in two prior investigations. Seshadri et al.
(2018) reported 410 reference archaeal and bacterial genomes from the Hungate
collection, while Stewart et al. (2018) compiled 913 draft MAGs from the rumens of
43 cattle bred in Scotland. The Hungate genomes are typically of superior value as it
contains genomes of isolated rumen microbes those exist in culture collection and
thus can be investigated in the lab. Recently a metagenomic investigation of the
ruminal contents of cows (Bos taurus), sheep (Ovis aries), reindeer (Rangifer
tarandus), and red deer (Cervus elaphus) was performed by Glendinning et al.
(2021). They constructed 391 metagenome-assembled genomes from 16 microbial
phyla and discovered 279 new species by comparing rumen genomes to other
publicly accessible microbial genomes. The study also reported substantial changes
in the number of microbial taxonomies, carbohydrate-active enzyme genes, and
KEGG orthologs in the microbiota of various ruminant species.

20.16 Whole Metagenome Analysis of Rumen

Most of the prior research have used 16S rRNA sequencing to characterize the
microbiota. Because readings must be matched against incomplete databases that
lack some specific rumen microorganisms, this technique yields minimal informa-
tion. In addition to culturing and 16S rRNA sequencing, whole genome sequencing
and assembly enable a more accurate taxonomic categorization based on several
marker genes and avoids PCR biases. Additionally, various taxonomic groupings
may perform comparable activities, masking genuine relationship at the gene func-
tion level when just the taxonomic makeup is considered. When a DNA molecule is
too lengthy to sequence in a single run, it must be sliced into several more manage-
able “bite-sized” pieces, each of which is copied and sequenced separately with
overlapping regions and then reassembled to generate the overall genetic picture.
While each DNA cutting, replication, and sequencing step is achievable manually,
they are the equivalent of a whole first-generation research using the Sanger or
Maxam-Gilbert methods. The research is crucial to create more efficient techniques
for perturbing the microbiome.
The era of whole-genome metagenomics has created new opportunities to analyze
microbiome composition to assess an individual’s feed consumption and its link to
metabolic processes involved in nutrient digestion, absorption, and utilization. The
ruminant gut microbiota contributes to the heterogeneity in feed consumption
efficiency. Delgado et al. (2019) used whole metagenome sequencing to demonstrate
20 Metagenomics Application in Understanding Rumen Functions 255

a link between the microbiome and feed utilization and consumption levels in a
Holstein cow herd. Their investigation discovered that the most effective cows have
a greater quantity of Bacteroidetes and Prevotella but lacked Firmicutes. Addition-
ally, Methanobacteria and Methanobrevibacter were less prevalent in high-
efficiency cows, indicating decreased methane emission. Their findings suggested
that there were variations in the microbiota compositions of efficient and inefficient
animals at both the taxonomic and gene levels.

Learning Outcomes
• Cloning and sequencing of 16s rRNA gene-based identification of rumen
bacteria is a more potent tool for identifying and characterizing rumen
microbiome.
• Understanding the rumen microbiome composition offer new possibilities
for improving production, disease resistance, novel enzyme discoveries,
and reducing enteric methane emissions in ruminants.
• Whole metagenome shotgun sequencing, metatranscriptomics, and
metaproteomics can be used to discover the functional capacities of the
metagenome.

20.17 Conclusions

The rumen microbiome plays a critical role in ensuring food security and mitigating
the effects of climate change. Recent research has resulted in the publication of over
1300 draught and full rumen genomes (Stewart et al. 2019). The vast array of
metabolic capabilities seen in the rumen microbiome may hold the key to increasing
the efficiency of animal production and industrial fermentation. Nucleic acid-based
technology advancements have reshaped our capacity to characterize the rumen
microbiome and opened new avenues for investigating the intricate interactions
and niches found within the rumen microbial population. The rumen microbiome
has been characterized in novel ways using next generation sequencing and other
molecular techniques. Pyrosequencing of the 16S ribosomal RNA gene has been
used for the taxonomic identification of bacteria and archaea at genus level. Whole
genome shotgun sequencing yields genuine metagenomic sequences that can be
used to forecast a microbiome’s functional capabilities and can also be utilised to
synthesized isolated species’ genomes. Integrating high-throughput data defining the
rumen microbiome with traditional fermentation and animal performance indicators
has resulted in significant breakthroughs and opened new research avenues.
To further understand the roles of the rumen microbiome, additional rumen
bacteria and archaea must be cultured. Understanding the microbiome and the
substrates used by bacteria, as well as how they interact with one another and with
their ruminant host to devise reasonable treatments to alter rumen feed conversion or
methane emissions, would be necessary. Improved rumen microbiome culture
256 S. Trivedi et al.

collections and future modification of the rumen microbiome can be better achieved
by genome sequencing and microbial genome assemblies.

20.18 New Frontiers

While genome assemblies provide insight into microorganisms’ metabolic

capabilities, next study will focus on the genes expressed in the microbiome
(metatranscriptome) utilizing RNA-seq. Increased data creation rates via high-
throughput technologies will necessitate the incorporation of bioinformatics tools
for data processing and meaningful (functional) outputs. Considering that diet
supplies substrates for the microbiome, fundamental nutrition principles will con-
tinue to be crucial for the design of experiments and their interpretation. Further
explicating the relationship between the host and microbiome might aid in
elucidating a variety of variables impacting microbial community alterations.
Knowledge of the animals’ microbial communities would allow for the manipulation
of growth and other desirable traits, such as minimal methane emissions and
maximum nutrition extraction from their diets. Furthermore, the high-throughput
microbiome investigations will help us better understand the development of animal
systems, immunology, tolerance to biotic and abiotic stress, resistance to infections,
metabolic diseases, etc. Using these methods, we want to gain a better understanding
of how methane lowering additives affect the rumen microbial population, as well as
how methanogenesis works. Safety evaluation of feed additives and rumen manipu-
lation effects could be studied through the profiling of the microbial community
analysis. Dysbiosis of the gut microbiome affect the metabolism as well as favor the
opportunistic infections to establish, the microbiome dynamics would facilitate the
study of such illnesses and remedies. Systems biology approaches to increase animal
productivity and welfare might potentially benefit from investigations of the
microbiome of the lung, reproductive tract, skin, udder, and genitalia. Understanding
the rumen microbiome could help in improving feed efficiency and reducing the
carbon footprint of animal products.

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 Part VI
Adaptation Strategies and Future Perspectives
Genetic Adaptation of Livestock to Heat
Stress Challenges 21
Concepta McManus, Andrea Queiroz Maranhão, Daniel Pimentel,
Felipe Pimentel, and Marcelo de Macedo Brigido

Contents
21.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 264
21.2 Methods for Identifying Specific Genes or Alleles for Adaptation . . . . . . . . . . . . . . . . . . . . . 266
21.3 Genes Linked to Heat Stress . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267
21.4 Future Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 273
Appendix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 277

Abstract
Genetic/genomic selection between and within species and breeds can aid in
maintaining production levels in farm animal species under climatic stress. This
chapter looks at how genes and animals can be identified and used for this
purpose. We also look at over 19,600 genes reported from studies on adaptation
cited in the scientific literature for cattle, sheep, goats and horses. Functional
analysis revealed pathways involved in developmental and growth processes,
regulation (positive and negative) of biological process, regulation of response to
stimulus and stress, immune system regulation, function and development, leu-
kocyte activation, oxidoreductase activity, metabolism and behaviour. Future
works will look at how we can select for increased tolerance to heat stress and
its related traits while maintaining productivity. Solutions may include landscape
genomics, genome editing and multi-omics studies. Overall, there is a need to
integrate different stakeholders with the development of statistical methodologies

C. McManus (*) · A. Q. Maranhão · D. Pimentel · M. de Macedo Brigido

Universidade de Brasília, Instituto de Ciências Biológicas, Brasilia, DF, Brazil
e-mail: [email protected]
F. Pimentel
CEUB, Brasília, DF, Brazil

# The Author(s), under exclusive license to Springer Nature Singapore Pte 263
Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_21
264 C. McManus et al.

(including artificial intelligence and machine learning) and a regulatory frame-

work to ensure animal welfare and consumer safety.

Keywords

Gene ontology · GWAS · Landscape · Selection · Sequencing

Abbreviations

B4GALT6 Beta-1,4-galactosyltransferase 6
Cas9 CRISPR associated protein 9
CRISPR Clustered regularly interspaced short palindromic repeats
DNA Deoxyribonucleic acid
DSC Desmocollins
DSG Desmogleins
EGFR Epidermal growth factor receptor
FDR False discovery rate
Fig Figure
FST Fixation index
GEBV Genomic breeding value
GO Gene ontology
GWAS Genowe-wide association study
HS Heat stress
KEGG Kyoto Encyclopedia of genes and genomes
LISA Local indicators of spatial association
MAPK Mitogen-activated protein kinase
mTOR mammalian target of rapamycin
PI3k/AKT phosphatidylinositol 3-kinase/protein kinase B
QTL Quantitative trait loci
RNA Ribonucleic acid
SNPs Single nucleotide polymorphisms
TTR Transthyretin

21.1 Introduction

There is an increased risk of extreme heat stress (HS) in domestic animals due to
climate change, especially in the tropics. Animals undergo stress (internal or external
stimuli that disrupt homeostasis) in all types of production systems. Still, how the
organism responds to this determines if it remains healthy and productive. This
response can be at several levels, including physiological, cellular, molecular or
behavioural. Nevertheless, to understand how stress affects an organism, it is
necessary to comprehend how response can be affected by several complex and
overlapping factors.
21 Genetic Adaptation of Livestock to Heat Stress Challenges 265

Heat stress does not typically occur in isolation. It can be accompanied by a lack
of water and feed (Sejian et al. 2013), the need to walk long distances, increased or
decreased humidity, and disease/vector prevalence. Input constraints such as forage
availability and quality, feed digestion and absorption occur in sequence. Therefore,
effects may be directly (thermoregulation, effects on endocrine, metabolism, pro-
duction, and reproduction systems) or indirectly (environmental effects on food and
water supply, pest and pathogen prevalence, and immune system resistance immu-
nological pressures) affect the animal. Under HS, as described above, animals reduce
food intake and, thereby, digestive capacity. Consequently, fewer nutrients are
absorbed. Reproduction is compromised, followed by production and growth, with
energy conserved for vital maintenance functions.
In animal production, selective breeding has improved productivity using modern
animal breeding techniques and reproductive technologies. The response is clear
with marked increases in efficiency and production levels but with a reduction in
fitness, resistance to disease and tolerance to increasing stresses in the environment
such as higher temperatures or lower water availability. Pryce and Haile-Mariam
(2020) show that heat tolerance has worsened in dairy cattle due to a lack of selection
for these traits. Climatic variables determine the choice of livestock species after
soils, geography, household characteristics, and country fixed effects have been
considered. An individual’s susceptibility to heat stress depends on both intrinsic
(mainly of genetic origin) and extrinsic (primarily environmental) factors (Gaughan
et al. 2002). Also, traits that were not previously deemed relevant, such as methane
production, have become important for the livestock industry (Hayes et al. 2013), as
well as the need to meet consumer demands for healthier animal products.
Genetic selection is incorporated into breeding programs based on linkage-
disequilibrium between polymorphisms in relevant characteristics and markers
(usually SNPs – Single Nucleotide Polymorphisms). Genomic breeding values
(GEBV) are calculated as the sum of each genetic marker’s effects or haplotypes.
As these are spread across the entire genome, potentially all quantitative trait loci
(QTL) that contribute to variation in the trait of interest are captured. For this, a
reference population is used whereby animals are both phenotyped and genotyped
for the characteristics of interest. Hayes et al. (2009) showed that linkage disequilib-
rium varied between breeds. When the prediction equations are to be used across
multiple breeds, the definition of the optimal composition of reference populations is
necessary.
Genomic selection can aid in selection schemes, as traits related to the response to
HS may be challenging to measure or have low heritability. This depends on
sufficient phenotyping on reference populations to determine linkages between
genotypes and phenotypes. Another option is the identification of genes of large
effects, such as the slick hair gene in locally adapted cattle breeds (Olson et al. 2003).
Dikmen et al. (2014) showed that introgression of this gene into Holstein cattle
improved thermoregulatory ability, with positive effects on milk yield even under
conditions whereby housing was modified to reduce HS. More recently, genome
editing (especially using CRISPR-Cas9 technology) has been used to introduce
beneficial alleles (e.g., heat tolerance, disease resistance) and haplotypes from native
266 C. McManus et al.

locally adapted to commercial populations to improve their productivity under

climate change (Singh and Ali 2021).
While mitigation of the effects of climate change on livestock through modifica-
tion of management conditions may represent a short-term solution to heat stress
challenges, genetic changes are more long term. Studies with locally adapted breeds
can help identify genes and metabolic pathways of interest in adaptation studies, as
they have had sufficient time to become genetically adapted to the environment.
These adaptations arise through equilibrium between evolutionary (crossbreeding,
artificial selection, genetic drift) and local environment sources.
Collier et al. (2019) identified three responses of the animal to HS: acclimation
(phenotypic response to a specific stressor), acclimatisation (coordinated response to
more than one stressor) and adaptation (genetic modification as adverse environment
persists over a few generations). Therefore, coping involves (McManus et al. 2020)
the ability to limit the heat load (resistance) and the ability to limit the harm caused
by a specific stressor (tolerance).
The animal’s response tries to minimise the effects of HS on cellular functions. It
does this through a coordinated gene network involving various cells, tissues and
systems. These genes are involved in (McManus et al. 2020) energy production/
metabolism, classical heat shock protein genes/chaperones, protein degradation/
turnover/DNA repair, genes reducing the impact of oxidative stress/cellular repair,
and transcriptional regulation. Gene expression is a crucial component of the cellular
response to thermal stress.

21.2 Methods for Identifying Specific Genes or Alleles

for Adaptation

Adaptation traits are complex and controlled by many genes control. These genes are
commonly grouped based on function or common expression profiles. This grouping
attempts to summarise a complex response network into fewer categories. These
response categories can then be interpreted due to their perceived roles in
re-establishing cellular homeostasis.
Molecular genetic approaches (such as microarray analyses, whole transcriptome
analysis, genome-wide association studies and next-generation sequencing) have
been used to identify adaptation-related genes. These, in turn, have been used to
group the genes associated with thermo-tolerance into different categories. Genome-
wide association studies (GWAS) use genetic variations in the genotype and pheno-
type by scanning the genomes of many different animals and looking for those
statistically associated with a specific trait or disease. This method aims to identify
common single nucleotide polymorphisms (SNPs) in the animal genome and deter-
mine how these polymorphisms are distributed across different populations. Other
methods analyse the patterns of genomic diversity within and between populations,
as well as the level of admixture in specific genomic regions, to identify adaptive
selection signatures. Landscape Genomics use environmental variables as proxies
21 Genetic Adaptation of Livestock to Heat Stress Challenges 267

for phenotypes. These approaches require genomic tools to look at individual loci
via whole genomic sequence analyses.
HS has been shown to stimulate signal transduction pathways. This stimulus
alters gene expression of immune cell mediators, thereby activating the heat shock
response, promoting cytokine activity. In sequence, this impairs the cellular immune
response by increasing cortisol concentrations, which binds to a specific Transcrip-
tion Factor that control gene function. Physiological responses are linked to these
molecular pathways and processes. Adaptive studies quantify responses through
mechanisms such as gene/protein expression, enzyme activity and genome-wide
analyses. For these studies to be feasible, there must be sufficient genetic variation
within a population.

21.3 Genes Linked to Heat Stress

Vertebrate genomes have millions of single nucleotide polymorphisms (SNPs).

Genetic variation can be due to deletions, duplications, copy-number variants,
insertions, inversions and translocations, mobile genetic elements, splicing junction
heterogeneity, regulatory elements and different sorts of ploidies. On the other hand,
crossing over, independent assortment and sexual reproduction are principal
mechanisms that maintain genetic diversity within populations. Genotype-
environment interactions cause phenotypic variation, which provides the substrate
for adaptive mechanisms.
We used three online tools (http://bioinformatics.sdstate.edu/go/; https://tools.
dice-database.org/GOnet/ and https://david.ncifcrf.gov/tools.jsp) to analyse over
19,906 genes (Fig. 21.1) found in published papers on the adaptation of cattle,
sheep, goats and horses worldwide.
As can be seen, most genes were cited in articles on cattle, and all species showed
significant overlaps. The genes found in at least three of the four species were then
used in further analyses (1703 genes used for enrichment, KEGG pathways, gene
ontology analyses).
Functional analysis revealed pathways involved in developmental and growth
processes, regulation (positive and negative) of biological process, response to
stimulus and stress, immune system regulation, process and development, leukocyte
activation, oxidoreductase activity, metabolism and behaviour (Fig. 21.2). As can be
seen, the response to HS involves several response levels. As the papers from which
these genes were identified are globally distributed and responses may be region or
breed-specific, further studies are needed to improve our understanding of these
processes.
Enrichment analysis (Fig. 21.3) shows 19 enriched regions (Bos taurus used as
the reference genome for the four species) on chromosomes 3, 7, 8, 13, 18, 19 and
24. For example, the enriched region on chromosome 24 shows genes linked to
epidermal, epithelial and mucosal structures (DSC1, DSC2, DSC3, DSG1, DSG2,
DSG3, DSG4), thyroxine transport (TTR), glycolipid biosynthesis (B4GALT6), and
protein modification (GALNT1), among others.
 268

Fig. 21.1 Upset Plot (https://gehlenborglab.shinyapps.io/upsetr/) of the number of genes linked to climatic adaptation cited in scientific papers per species
(based on 19,609 genes and 127 scientific papers)
C. McManus et al.
21 Genetic Adaptation of Livestock to Heat Stress Challenges 269

Fig. 21.2 Functional analysis network from 1703 genes related to heat response in cattle, sheep,
goats and horses (https://tools.dice-database.org/GOnet/)

B
A

Fig. 21.3 Chromosomal localization of 1703 genes found in published papers on the adaptation of
cattle, sheep, goats and horses worldwide (a) and a closeup (b) of one of the enriched regions based
on cattle genomic organization (individual genes in red, enriched regions underlined in purple -
these regions were defined by chi-square tests using all 19,906 genes found in the literature. http://
bioinformatics.sdstate.edu/go/)
270 C. McManus et al.

Signalling networks, such as MAPK (immune response, gene expression, cell

proliferation, differentiation, apoptosis, and cell motility), mTOR (growth, energy
metabolism, ageing), ErbB (cell growth, development and survival, lipid metabo-
lism), and PI3k/AKT (metabolism, proliferation, cell survival, growth and angio-
genesis) are also involved in this response (Fig. 21.4 and Table 21.1).
In the reactome (https://reactome.org/) analysis, the importance of the immune
response can be seen due to the prevalence of interleukin and MAPK signalling.
Here too, the response to heat stress is highlighted. AKT (serine/threonine kinase) is
an upstream positive regulator of the mammalian target of rapamycin (mTOR)
which controls cell growth. The KEGG analyses (https://www.genome.jp/kegg/
pathway.html) highlights some of the same reactions, but also includes melanogen-
esis and epidermal growth factor receptor (EGFR) tyrosine kinase inhibitor resis-
tance possibly involved in primary defence of the organisms to ultraviolet light.
Rap1 regulates the cell’s response to external stimuli, and is also involved in cell
survival and proliferation. In the Gene Ontology (http://geneontology.org/) for
biological processes, development and metabolic processes are emphasised.
Directly annotated Gene Ontology mappings and Tissue expression locations
(Fig. 21.5) further enhance the complexity of the heat stress response, as noted in
Fig. 21.4 above. The response involves KEGG metabolic pathways in the liver,
during foetal development, the digestive system, brain, heart, blood, and several
other organs. The liver is an important regulator of growth and metabolism. It
controls many physiological processes that are impacted by heat stress (Hubbard
et al. 2019). These authors also show that changes in the cell membrane exert
important downstream effects on heat stress response genes and metabolites
(Fig. 21.5b). Other organs and structures are also important. For example,
interleukins (Fig. 21.4a) are expressed in white blood cells and endothelium, while
melanocytes (Fig. 21.4b) in the skin, eye, meninges and bones. The thymus
(Fig. 21.5b) makes up part of the immune system and is responsible for T cell
differentiation and development. Heat stress also affects the digestive and renal
systems, heart and blood flow, as well as parts of the brain such as the hippocampus,
cerebellum and hypothalamus.
An increase or decrease in the frequency of genetic variants in a population can be
caused by natural or artificial selection pressures. Selection signatures are regions of
the genome that hold functionally important sequence variants. They, therefore, are
or have been under selection (natural or artificial), creating specific patterns of DNA.
The reach of such signatures in the genome, up- and downstream of this functional
variant, is a consequence of the so-called hitchhiking effect. Therefore, the increase
in the prevalence of advantageous alleles under positive directional selection will
leave distinctive signatures (patterns of genetic variation) in the DNA sequence.
Local adaptation selection signatures can be studied using whole-genome data.
Spatial association with these molecular markers is also possible as candidate loci for
adaptation traits can be identified through genome-environment associations (Stucki
et al. 2017). Measuring Local Indicators of Spatial Association (LISA) for these
candidate loci enables means that we can evaluate how similar genotypes associate
spatially (Cesconeto et al. 2017). Paim et al. (2018) described the main statistical
 21
Genetic Adaptation of Livestock to Heat Stress Challenges

Fig. 21.4 A hierarchical clustering tree summarising the correlation among significant pathways from the Enrichment analysis using Reactome (a), KEGG (b)
and Gene Ontology (GO) analyses (c). Pathways with many shared genes are clustered together (http://bioinformatics.sdstate.edu/go/). Numbers are False
Discovery Rate (FDR). Bigger blue dots indicate more significant P-values
271
272 C. McManus et al.

Fig. 21.5 Word clouds for Gene Ontology mappings directly annotated by the source database (a)
and Tissue expression locations (b) (https://david.ncifcrf.gov/tools.jsp)

methods and software used to analyse genomic data and the identification of
selection signatures.
The spatial genetic differentiation of loci can vary due to the demographic history
of a species (Hoban et al. 2017). Even without selection, all have the same influence
of genetic drift and migration. Possible neutral differentiation patterns need to be
separated from loci under local selection. Focusing on departures from neutrality for
detecting selection is difficult. Random processes affect each locus in different
manners. The population structure and demography of the species in question also
affect differentiation distribution. This can be problematic when there is a high
average level of differentiation. The variance in FST values (Fixation index)
among loci increases with average FST, even with selective neutrality, making
detecting outlier loci difficult for highly differentiated populations.
The genetic bases of species adaptation to geographic conditions or climate
change can be carried out using Geographical Information Systems. Manel et al.
(2003) described how geographical and environmental features could facilitate
genetic variation structure at population and individual levels. This method focuses
on fine spatial and temporal scales, such as those with recent farm livestock
migration. It has been used to identify genes responsible for adaptive evolution of
species at a population level, such as quantifying the influence of spatial environ-
ment on genomic divergence and uncovering environmental factors that shape
adaptive genetic variation and the genetic basis of adaptive change (Li et al.
2017). The focus in these studies needs to be on phenotypic and genetic variation
so as to validate the function and adaptive generality of the detected loci. Therefore,
genes involved in the regulation of metabolic pathways, as well as the adaptive
phenotypes controlled by these genes, need to be identified. At the same time, Storfer
et al. (2018) show that understanding the underlying demographic structure of study
populations is essential when selecting genome scan methods. Species adaptability
should be used to determine its distribution range, as responding to climate change
depends on their landscape adaptability. This is usually determined by the genome
adaptive differentiation potential as well as the species/breed’s gene dispersal ability.
21 Genetic Adaptation of Livestock to Heat Stress Challenges 273

Learning Outcomes
• Heat stress, when combined with other stresses affects drastically the
productivity of farm animals.
• Traits related to the heat stress response have low heritability values for
genomic selection.
• Advanced molecular and genetic tools provide greater opportunities to
identify potential biomarkers for heat stress.
• Multi-omics approaches involving genome, proteome, transcriptome,
epigenome, metabolome, and microbiome could revolutionise genomic
selection for heat tolerance in farm animals.

21.4 Future Perspectives

Under controlled management conditions (sufficient feeding, heat mitigation, and

controlled parasite and pathogenic environment), selection for heat tolerance within
highly productive breeds is likely to offer more opportunity than improving local
breeds. On the other hand, crossing local and selected breeds and monitoring of heat
tolerance may improve production system productivity where these conditions are
not present. Research in functional genomics provides new information on HS
impacts on livestock production. Genetically superior animals can be selected by
identifying genes that are up- or down-regulated during HS.
Possible candidate genes have been identified in this work that could be
associated with adaptation to HS. Stress causes a series of response mechanisms in
animal physiology that harm the whole production chain. The identification of major
genes positively correlated to heat tolerance can be used as markers in
marker-assisted selection, Genomic selection, and gene editing programs. Also,
other potential biomarkers on DNA and RNA, including the non-coding ones,
have been proposed recently. However, there is still a gap between transforming
these markers into tangible tools for breeding programs. Much research is still
necessary.
Difficulties in breeding animals for resistance/tolerance to HS (Romero et al.
2015) include the lack of (1) an unambiguous definition of the stress phenotype; (2) a
robust (or group of) biomarker to characterise the proposed phenotype; (3) reliable
models (theoretical and quantitative) for prediction of how animals react to stressors;
(4) a clear understanding of individual variability in responses to stress and
transitions between acute and chronic stress. As a whole, these problems limit the
ability to assess an individual’s physiological status and develop techniques to
reverse/control the effects of chronic stress before they become pathological. Essen-
tial for the success of breeding programs for improved HS tolerance is the availabil-
ity of High-throughput phenotyping. Developing low-cost sensors and automated
data collection and storage for genotyping reference populations is necessary for the
efficient use of genomic breeding values and understanding the biology of the
underpinning traits.
274 C. McManus et al.

Understanding the physiological basis for adaptation requires increased

characterisation with high-throughput single nucleotide polymorphism (SNP) assays
or genome sequencing (Boettcher et al. 2015). Species-wide HapMap studies and
multi-species studies are the first step in understanding the link between the genome
and adaptation. Nevertheless, information on more breeds, geographical areas and
production system environments are required to acquire a complete picture.
Metagenomics of associated microbiome can also help understand the
co-adaptation of Animal Genetic Resources with other organisms in production
environments. This can add information on novel biocatalysts, enzymes, genetic
linkage, as well as phylogenetic and evolutionary profiles of microorganism’s
community function and structure and their effects on host phenotypes.
The Multi-omics approach to addressing heat tolerance and methane emissions
will become important over the next decade. Examples of a multi-omics data set
include some of the following: direct and indirect selection criteria, the metagenome
(e.g., rumen, reproductive, and so on), mid-infrared spectral data, in addition to
information from the proteome/metabolome (protein/metabolite structure and func-
tion) and functional genomic assays (e.g., methylation, transcriptomics). Causal
variants can be identified when these approaches are used together with whole-
genome sequencing, leading to improved responses to selection. If significant effect
variants are found, these could become candidates for gene editing. In a collabora-
tive environment, a multi-omics approach (including genome, proteome,
transcriptome, epigenome, metabolome, and microbiome) could revolutionise geno-
mic selection for traits of interest under climate change. This requires the integration
of multi-omics data analysis and machine learning. Combined with the fact that
mixed models have been applied to GWAS, and can reduce the number of false-
positive associations, it is clear that statistical knowledge will play a significant
factor in future applications of these technologies.
Genomic selection and advanced biotechnologies such as gene editing can help
improve pure- and crossbreeding programmes to include traits linked to adaptation,
assuming that phenotypes are available. Therefore, programs for performance
recording animals in regions suffering climate change and stressful environments
are needed. Integration of geographical and genetic information is also essential and
adequate referencing, organisation, storage, and dissemination to stakeholders. This
requires new and improved databases and information systems and the capability to
link different geographical and production scales in formats that can be analysed.
Regulatory frameworks for the use of advanced technologies such as gene editing
will also determine the future of their application as animal health and consumer
safety has to be considered. Therefore, cooperation between all stakeholders will be
needed to optimise the genetic and genomic resources to adapt livestock to climate
change.

Acknowledgements Thanks are due to the Coordenação de Aperfeiçoamento de Pessoal de Nível

Superior (CAPES) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)
for financing.

Conflict of Interest The authors declare no conflict of interest.

21 Genetic Adaptation of Livestock to Heat Stress Challenges 275

Appendix

Table 21.1 Pathway enrichment analysis of 1703 genes (from a total of 19,606) related to heat
stress in at least three species of livestock (cattle, sheep, goats or horses)
Enrichment Number Pathway Fold
FDRa of genes Genes enrichment Pathways
b
GO biological processes
3.10E-07 273 1747 1.4 Response to organic substance
6.70E-08 313 2019 1.4 Regulation of multicellular organismal
process
1.20E-07 331 2180 1.4 Phosphate-containing compound
metabolic process
2.80E-07 331 2198 1.4 Phosphorus metabolic process
1.20E-07 381 2579 1.3 Positive regulation of metabolic process
2.00E-09 471 3208 1.3 Multicellular organism development
7.10E-08 428 2944 1.3 System development
4.80E-11 558 3841 1.3 Developmental process
6.80E-09 512 3577 1.3 Anatomical structure development
3.10E-07 556 4056 1.2 Positive regulation of biological process
GO molecular function
1.20E-08 190 1093 1.6 ATP binding
1.20E-08 197 1142 1.6 Adenyl nucleotide binding
1.30E-08 195 1132 1.6 Adenyl ribonucleotide binding
1.30E-08 224 1348 1.5 Purine ribonucleoside triphosphate
binding
1.10E-08 234 1413 1.5 Purine nucleotide binding
1.30E-08 232 1410 1.5 Ribonucleotide binding
1.10E-08 261 1609 1.5 Nucleotide binding
1.10E-08 261 1609 1.5 Nucleoside phosphate binding
1.10E-08 293 1862 1.4 Small molecule binding
1.10E-08 320 2065 1.4 Anion binding
KEGGc
3.20E-05 25 76 3 EGFR tyrosine kinase inhibitor
resistance
1.90E-04 26 91 2.6 Melanogenesis
8.00E-05 35 136 2.3 Gastric cancer
8.20E-08 65 263 2.2 MAPK signalling pathway
4.10E-04 33 137 2.2 Cellular senescence
2.30E-04 39 168 2.1 Lipid and atherosclerosis
3.00E-04 41 183 2 Proteoglycans in cancer
6.90E-04 40 185 2 Rap1 signalling pathway
6.50E-05 63 307 1.9 PI3K-Akt signalling pathway
4.90E-06 92 471 1.8 Pathways in cancer
(continued)
276 C. McManus et al.

Table 21.1 (continued)

Enrichment Number Pathway Fold
FDRa of genes Genes enrichment Pathways
Pantherd
4.30E-02 11 39 2.6 Interleukin signalling pathway
4.20E-02 16 64 2.3 Cadherin signalling pathway
4.30E-02 16 68 2.1 PDGF signalling pathway
2.50E-02 25 113 2 Inflammation mediated by chemokine
and cytokine signalling pathway
4.30E-02 20 91 2 Angiogenesis
2.00E-02 33 157 1.9 Wnt signalling pathway
Reactomee
3.00E-03 10 19 4.8 HSF1-dependent transactivation
3.00E-03 21 69 2.8 Cellular response to heat stress
6.80E-03 20 68 2.7 Negative regulation of the PI3K/AKT
network
1.50E-02 19 67 2.6 PI5P PP2A and IER3 regulate PI3K/
AKT signalling
2.10E-02 28 125 2 PIP3 activates AKT signalling
2.10E-02 33 156 1.9 Other interleukin signalling
2.10E-02 34 161 1.9 MAPK family signalling cascades
3.00E-03 57 285 1.8 Signalling by interleukins
1.40E-03 71 364 1.8 Cytokine signalling in immune system
2.10E-02 170 1188 1.3 Immune system
Wikipathwaysf
5.30E-04 18 51 3.2 Thyroid Stimulating Hormone (TSH)
signalling pathway
1.10E-02 14 45 2.8 Kit receptor signalling pathway
1.30E-02 14 48 2.6 Endochondral ossification
1.30E-02 14 48 2.6 Oncostatin M signalling pathway
1.20E-02 17 63 2.4 Leptin signalling pathway
9.50E-04 30 121 2.2 Integrated breast cancer pathway
9.50E-04 31 127 2.2 EGF/EGFR signalling pathway
2.10E-02 19 80 2.1 WP3231 senescence and autophagy in
cancer
2.40E-04 43 182 2.1 MAPK signalling pathway
1.20E-02 25 109 2.1 BDNF signalling pathway
Cellular component
3.50E-03 9 17 4.8 Desmosome
1.10E-07 346 2302 1.4 Cytosol
3.50E-03 373 2802 1.2 Intracellular non-membrane-bounded
organelle
3.50E-03 373 2803 1.2 Non-membrane-bounded organelle
1.60E-03 465 3526 1.2 Protein-containing complex
1.50E-02 338 2571 1.2 Nucleoplasm
3.50E-03 404 3076 1.2 Membrane-enclosed lumen
(continued)
21 Genetic Adaptation of Livestock to Heat Stress Challenges 277

Table 21.1 (continued)

Enrichment Number Pathway Fold
FDRa of genes Genes enrichment Pathways
3.50E-03 404 3076 1.2 Organelle lumen
3.50E-03 404 3076 1.2 Intracellular organelle lumen
8.90E-03 374 2856 1.2 Nuclear lumen
All available gene sets
4.90E-08 234 1413 1.5 Purine nucleotide binding
4.30E-08 261 1609 1.5 Nucleotide binding
4.30E-08 261 1609 1.5 Nucleoside phosphate binding
4.90E-08 293 1862 1.4 Small molecule binding
4.90E-08 313 2019 1.4 Regulation of multicellular organismal
process
4.30E-08 320 2065 1.4 Anion binding
3.20E-09 471 3208 1.3 Multicellular organism development
5.30E-08 428 2944 1.3 System development
7.90E-11 558 3841 1.3 Developmental process
1.10E-08 512 3577 1.3 Anatomical structure development
a
FDR false discovery rate
b
GO gene ontology
c
KEGG Kyoto encyclopaedia of genes and genomes—https://www.genome.jp/kegg/pathway.html
d
Panther protein analysis through evolutionary relationships—http://www.pantherdb.org/pathway/
e
Reactome-database of reactions, pathways and biological processes—https://reactome.org/
f
https://www.wikipathways.org/index.php/WikiPathways

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Goat as the Ideal Future Climate Resilient
Animal Model 22
Veerasamy Sejian, M. V. Silpa, M. R. Reshma Nair,
Chinnasamy Devaraj, A. Devapriya, N. Ramachandran,
V. B. Awachat, C. G. Shashank, Vinícius F. C. Fonsêca,
and Raghavendra Bhatta

Contents
22.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281
22.2 Goat as the Future Animal from Food Security Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . 282
22.3 Advantages of Goat Farming in Tropical Countries . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 283
22.4 Climate Resilient Goat Production . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 284
22.5 Concepts Associated with Climate Resilient Goat Production . . . . . . . . . . . . . . . . . . . . . . . . . 285
22.6 Biomarkers for Heat Stress in Goat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 287
22.6.1 Phenotypic Markers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 287
22.6.2 Genotypic Markers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 287
22.7 Adaptation Strategies to Sustain Goat Production Under Changing Climate . . . . . . . . . . 288
22.8 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 290
22.9 Future Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 290
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 291

V. Sejian (*) · M. R. Reshma Nair · C. Devaraj · A. Devapriya · V. B. Awachat · R. Bhatta

Centre for Climate Resilient Animal Adaptation Studies, ICAR-National Institute of Animal
Nutrition and Physiology, Bangalore, India
M. V. Silpa
Institute of Animal Breeding and Genetics, Justus-Liebig-Universität Gießen, Gießen, Germany
N. Ramachandran
Bioenergetics and Environmental Sciences Division, ICAR-National Institute of Animal Nutrition
and Physiology, Bangalore, India
C. G. Shashank
ICAR-National Dairy Research Institute, Karnal, India
V. F. C. Fonsêca
Animal Biometeorology and Ethology Group (BIOET), Department of Animal Science, Federal
University of Paraiba, Areia, Brazil

# The Author(s), under exclusive license to Springer Nature Singapore Pte 279
Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_22
280 V. Sejian et al.

Abstract

The negative effects of climate change on livestock production was mediated

predominantly through the indirect impacts such as sudden disease outbreaks,
less feed and water available as well as shrinking pasture lands. Therefore, it is
very essential for the scientific community to identify the best species among the
livestock which could cope with the adversities associated climate change. The
very few recent research efforts based on modeling approach clearly
demonstrated that goat is the only species among the livestock to face very less
number of extremely severe heat stress days both by 2050 as well as 2100. Thus it
is very important to concentrate on this otherwise neglected species so far because
rearing goats could help the livestock farmers to optimize their economy in the
changing climate scenario. Goats are predominantly distributed in the tropical
countries and they possess the ability to survive in harsh locality due to their high
thermo-tolerance, drought tolerance, unique feeding behaviour as well as disease
resistance. Most goat breeds recorded are indigenous in nature. Differences in
climate resilience were established even among the indigenous breeds. This
warrants more research efforts on the indigenous breeds to screen for their climate
resilience probably with the intention to identify the best breed which could
survive and produce optimally in specific agro-ecological zones. The chapter
also highlights the various adaptive strategies that need to be implemented to
sustain goat production in the changing climate scenario.

Keywords
Adaptation · Climate resilience · Goat · Heat stress · Thermo-tolerance

Abbreviations

γ Gamma
% Abbreviation
DAGT Diacylglycerol acyltransferase
Fig Figure
FSHR Follicle stimulating hormone receptor
GH Growth hormone
GHR Growth hormone receptor
HCT Hematocrit
HGB Hemoglobin
HSF Heat shock factor
HSP Heat shock protein
ICAR Indian Council of Agricultural Research
IFN Interferon
IGF-1 Insulin-like growth factor-1
LEP Leptin
LHR Luteinizing hormone receptor
22 Goat as the Ideal Future Climate Resilient Animal Model 281

MSTN Myostatin
NIANP National Institute of Animal Nutrition and Physiology
NOS Nitrous oxide synthase
NRAMP1 Natural resistance-associated macrophage protein 1
PLR Prolactin receptor
SOD Super oxide dismutase
THR Thyroid hormone receptor
TLR Toll-like receptor
TNF Tumour necrosis factor

22.1 Introduction

Livestock farming plays a vital role in ensuring livelihood security especially

towards the marginal or economically weaker segments of the society, who lack
even the basic resources required to create favorable microclimate for managing the
farming system. Global demand for livestock products are predicted to double
during the first half of this century as a consequence of the growing human popula-
tion and their affluence. On the other hand, drastic changes are expected to occur
from the global climate change point of view. In the present scenario, climate change
is among the most serious challenges faced by farmers and livestock owners around
the globe, having a long term effect. The indirect impacts of climate change are
proved to significantly impair livestock production. This is even more pronounced in
tropical countries which inhabit the major proportion of the livestock population,
especially with their indigenous germplasm. Additionally, severe reduction in pas-
ture availability as along with shrinking grazing lands also lead to marked reduction
in livestock production. Furthermore, the addition of climate change impact on
pasture availability, to the ongoing reduction, would aggravate the issue severely
(Sejian et al. 2018). Hence, scientific communities are under enormous pressure to
sustain livestock production amidst the chaos and ensure food security from the
future perspectives. Therefore, it is very vital to channelize the research efforts to
identify potential livestock species that can effectively cope with the adversities
associated with climate change and also produce optimally.
With the projected alarming impacts of climate change on the pasture availability,
small ruminants and especially goat production gain significance because of their
ability to survive on limited pastures. The farming communities are looking to invest
in the small ruminant livestock production systems because of their lower feed
requirement, lower input cost and better climate resilience than the large ruminants.
Among the small ruminants, goats are considered ideal climate animal model due to
their better thermo-tolerance, drought tolerance, ability to survive on limited pastures
as well as the disease resistance capacity (Reshma Nair et al. 2021). In a recent
research effort by Carvajal et al. (2021) and Rahimi et al. (2021) it was clearly
established that among the livestock species, goats will experience least number of
282 V. Sejian et al.

Fig. 22.1 Model projections of extremely severe heat stress to be experienced by different
livestock species by 2050 and 2100 in East Africa (source Rahimi et al. 2021; Reshma Nair et al.
2021)

extremely severe stress days both by 2050 as well as by 2100 in the globe as a whole
as well as East Africa respectively. Further, rearing goat is considered to be more
economical when compared to large ruminants, due to their comparatively lower
feed requirement, monetary investment and management (Reshma Nair et al. 2021).
Figure 22.1 describes the modeling approach to project the extremely severe heat
stress days that major livestock species are going to experience by both by 2050 and
2100 scenario. This compilation is therefore an attempt to collate and synthesis
information on this line to project the various advantages associated with goat
production to prove them as ideal animal model in the changing climate scenario.

22.2 Goat as the Future Animal from Food Security Perspectives

In order to sustain livestock production under the alarmingly changing climatic

conditions, researchers are now channelizing their work towards identification of
ideal species that can cater the needs of the growing human population. Several
studies have identified goats to be the go-to species, for possessing the ability to
efficiently sustain animal agriculture amidst the changing environmental conditions.
Pioneers in livestock research had identified goats to be more potential among the
other small ruminants, for their better adaptation to a wide range of environmental
conditions. Goats being opportunistic feeders, are stated to be least affected due to
the ongoing and projected depletion of pasture. Apart from this, the selective feeding
behavior exhibited by goats are proved to be advantageous; as these species can
consume even the poor quality forages, the nutrients obtained from which are then
converted into high quality products. Furthermore, the bipedal stance behavior
22 Goat as the Ideal Future Climate Resilient Animal Model 283

exhibited by goats gives them an advantage over other livestock species as this gives
them access to the tree fodders that are usually inaccessible to other species. Further,
the feed efficiency of goats are considered to be better than other ruminant species.
Another benefit of goat rearing, especially from economics perspective is their
minimal housing requirement. Goats can survive well in any location even with
minimum protection from the weather. Additionally, labor costs associated with goat
production is minimal as they can be managed efficiently even by using family
members, which is the usual practice followed especially by the poor farmers.
The human population across the globe is projected to touch an alarming count of
9.6 billion by 2050. This is a major concern from the food security aspect as animal
proteins are the most preferred protein source by majority of the individuals,
especially in the developing world. Goats are considered to be the ideal climate
animal model that has the ability to perform better than other species amidst the
projected climatic adversities and also feed and fodder shortage. As per the model by
Ngambi et al. (2013), dairy goats produce approximately 15.2 million tons of milk
that constitute 2% of total livestock milk production. Moreover, goat meat and milk
are considered to have several health benefits and therapeutic values that further
boosts up their demand. In harmony with this, recent reports also suggest that the
wide acceptance of goat products across the globe makes the goat enterprises to be of
more commercial value. This highlights their pivotal role in meeting the growing
humanitarian animal protein needs by the end of this century along with their most
important role to financially support poor and marginal farmers around the world.
Therefore, it is the need of the hour to gather researchers and policymakers who can
efficiently set priorities to design appropriate programs to ensure sustainable live-
stock production and also satisfy the food demands of the growing human popula-
tion by 2050.

22.3 Advantages of Goat Farming in Tropical Countries

Goat farming is one of the largest subsectors of agriculture in arid, semi-arid regions
of tropical and subtropical countries. It plays a vital role in livelihood and economic
wellbeing of marginal, small farmers and landless labourers in both rural and peri-
urban areas in developing countries. Goat farming is advantageous over the other
livestock farming system since it maintains their production potential under extreme
climate conditions of the tropical and subtropical regions. The anatomical and
physiological characteristics of goats and economics of goat farming make them
more beneficial comparing to the other farm animals.
Goats have been considered as ideal animal to be reared in tropics and subtropical
regions. Goats have a capacity to thrive and perform better under harsh environmen-
tal conditions because of their superior thermoregulation capacity and ability to
withstand during feed and water scarcity. Goats better utilises low quality fodder and
shrubs or benefit from feed resources which are not consumed by the other farm
animals and convert them into meat and milk. Goats have ability to walk long
distance for search of feeds. Comparing to the other domesticated farm animals,
284 V. Sejian et al.

goat emits less methane. In addition to their great adaptability to extreme climatic
conditions and to different feeds, it require less space (smaller body size), less
housing requirement, less maintenance cost, easy to handle, faster growth rate,
higher resistance to disease as well as heat stress which make them goats are much
better than other ruminant species.
Goat forming requires low investment but gives high rates of return and always
available for sales of live goats to generate revenues to meet their immediate needs.
Therefore, goats are considered as a poor man’s cow since it is an important source
of income for rural poor farmers. Thus, goat farming in tropics and subtropical
countries stabilize the rural economy.
Goat meat (chevon) is a delicious food for most people across the globe since it is
lean and has low cholesterol content. Similarly, goat milk and products demand are
increasing and it is most consumed dairy product in the world in recent times. Goat
milk are rich in vitamin and mineral content, easily digestible and better taste
compared to cow milk. Goat products are widely accepted, no religious taboo and
having higher demand across the world. Goat manure are good source natural
fertilizers for rural farmers which are good source of nitrogen, phosphorus and
potassium that improves the soil fertility and ensure healthy crop growth and
productivity.

22.4 Climate Resilient Goat Production

Identification of indigenous breeds that can survive efficiently across varied agro-
ecological zones is the key to ensure sustained livestock production. In series of
studies conducted at Indian Council of Agricultural Research- National Institute of
Animal Nutrition and Physiology (ICAR-NIANP), three different indigenous goat
breeds Osmanabadi, Malabari and Salem Black goats from three South Indian states;
Karnataka, Kerala and Tamil Nadu, respectively, were compared for their climate
resilience capacity. Innumerable phenotypic and genotypic traits associated with
heat stress were studied in these goats which thereby led to the conclusive report that
Salem Black breed was able to adapt and produce better when compared to
Osmanabadi and Malabari breeds upon heat stress exposure (Archana et al. 2018;
Angel et al. 2018; Amitha et al. 2019; Rashamol et al. 2019). The significantly lower
respiration rate, rectal temperature and heat shock protein 70 (HSP70) gene expres-
sion in the Salem Black in comparison to the Osmanabadi and Malabari goats during
heat stress indicate better thermal resilience of the Salem Black goats (Aleena et al.
2018). Therefore, promoting the Salem Black breed among the local farmers could
benefit them and improve their livelihood security. Figure 22.2 depicts the three
indigenous goat breeds, Osmanabadi, Malabari and Salem Black.
22 Goat as the Ideal Future Climate Resilient Animal Model 285

Fig. 22.2 Three indigenous goat breeds of South India

22.5 Concepts Associated with Climate Resilient Goat

Production

The livestock sector in general holds a uniqueness for both being sensitive to climate
change and also a contributor to the phenomenon. Though goats are excellent
climate resilient species, they too face the deleterious impact of climate change.
Amongst the multiple climatic stresses faced by goat, heat stress is of major concern
as it destabilizes the production efficiency of the animals. The deleterious effects of
heat stress include decline in growth, meat and milk production in goat. Further,
climate change leads to the increase and rapid spread of several vector borne diseases
in goat which further compromises the immune status of the animals. Nevertheless
animals employ some vital adaptive mechanisms through behavioural, physiologi-
cal, neuro-endocrine, cellular and molecular responses that aid them to efficiently
combat the changing climatic condition. Further, climate change can lead to alter-
ation in the rumen function and thereby digestibility in goats. This could aggravate
the issue of enteric methane emission apart from the dietary losses incurred. Hence
suitable ameliorative strategies need to be enforced to counter the adverse impact of
climate change on goat production and reduce its contribution towards global
warming. The management strategies can be broadly categorized as housing man-
agement, animal management and climate monitoring. Nutritional interventions
considering season specific feeding and micronutrient supplementation could also
play a efficient role both to ameliorate the climate change impact on goats as well as
sustain their production. Adoption of goat specific body condition scoring system
may help to optimize economic return by minimising the unnecessary feeding costs.
Finally, proper emphasis must be given to develop and enforce appropriate adapta-
tion strategies by active involvement of policy makers. These strategies may include
development thermo-tolerant breeds, ensuring water availability, promoting women
empowerment, establishing accurate early warning system and effective capacity
building programmes for all the stakeholders. These efforts would be vital to ensure
sustainable goat production in the changing climate scenario. Figure 22.3 describes
the various concepts associated with climate resilient goat production.
 286

Fig. 22.3 Various concepts associated with climate resilient goat production
V. Sejian et al.
22 Goat as the Ideal Future Climate Resilient Animal Model 287

22.6 Biomarkers for Heat Stress in Goat

22.6.1 Phenotypic Markers

Drinking frequency, respiration rate, and rectal temperature have been reported to be
the ideal biological markers for assessing the impact of both heat and nutritional
stress in goats. In another study conducted in Osmanabadi goats, Insulin-like growth
factor 1 (IGF-1), leptin (LEP), growth hormone (GH) and plasma HSP70 were
established as biomarkers for assessing nutritional stress (Pragna et al. 2018;
Abhijith et al. 2021). Based on the study conducted to comparatively assess the
climate resilience of three indigenous goat breeds, Osmanabadi, Malabari and Salem
Black breeds, drinking frequency, water intake, haemoglobin, packed cell volume,
plasma cortisol, aldosterone, tri-iodo-thyronine and thyroxin were established to be
reliable biomarkers for heat stress response in goats cutting across breeds (Shilja
et al. 2016). These findings may also be useful for assessing the implications of heat
stress from the perspective of animal welfare.

22.6.2 Genotypic Markers

The classical heat stress associated molecular chaperone, HSP70 gene was identified
to act as an ideal biological marker for assessing the impact of combined stress (heat
and nutritional stress) in Osmanabadi goats. In the same study, the significantly
higher expression of TLR8 and TLR10 in heat stressed goats indicated the probable
role of these genes as immunological markers of heat stress in goats. In another study
conducted in Osmanabadi goats, growth hormone receptor (GHR), HSP70 and
HSP90 were identified as efficient markers for assessing nutritional stress in goats.
Further, based on the results obtained from a mega study at ICAR-NIANP a number
of biomarkers were reported that were significantly associated with heat stress.
These included some of the well established heat stress associated genes like heat
shock factor 1 (HSF1), HSP27, HSP60, HSP70, HSP90, HSP110; growth and
metabolism associated genes like GH, GHR, IGF-1, LEP, leptin receptor (LEPR),
thyroid hormone receptor (THR); genes associated with reproduction, prolactin
receptor (PLR), follicle stimulating hormone receptor (FSHR), luteinizing hormone
receptor (LHR) (Sejian et al. 2019). It also included a number of immune response
linked genes like interleukin 10 (IL10), IL18, interferon β (IFNβ), IFNγ, tumour
necrosis factor α (TNFα), toll-like receptor-1 (TLR1), TLR4, TLR5, Natural
resistance-associated macrophage protein 1 (NRAMP1), nitrous oxide synthase
(NOS), superoxide dismutase (SOD) (Sophia et al. 2016; Rashamol et al. 2019).
Figure 22.4 describes the different biomarkers for quantifying heat stress response in
goats.
288 V. Sejian et al.

Association Analysis between THI and Phenotypic and Genotypic Traits

Behavioural Traits Immune System

Physiological Traits
Drinking Frequency, Feed Related
Respiration Rate
Intake, Water Intake, TLR2, TLR4, TLR8, IL-
Rectal Temperature
rumination time 2, IL-6, IL-10, IL-18,
Skin Temperature
TNF-α, IFN-β, and IFN-
Panting Score
γ genes
Sweating rate

Blood & Endocrine

Traits
Growth Related Traits
HGB
GH, GHR, IGF-1,
HCT
Adiponectin genes
Cortisol
Aldosterone
T3 & T4
HSP70

Thermo-tolerant Genes Non Invasive Methods

Meat Quality Genes HSF-1, HSP27, HSP40, Hair Cortisol
MSTN, LEP, CAPN1, HSP60, HSP70, HSP90, Saliva Cortisol
CAPN2, CAST, HSP110 , PRLR, SOD, NOS , Faeces Cortisol
CRYA, DAGT1 genes THR genes Urine Cortisol

Fig. 22.4 Different biomarkers for quantifying heat stress response in goats

22.7 Adaptation Strategies to Sustain Goat Production Under

Changing Climate

Adapting to climate change comprises of the inclusion of accurate measures to

reduce the adverse effects of climate change along with enforcing suitable strategies
to reduce the contribution of livestock sector to climate change. Development of
thermo-tolerant breeds, that not only adapt well but also produce efficiently, using
various genomics approach could be one of the most efficient and long term solution.
Additionally, reduced methane emission should also be considered as a vital trait
while considering the revised livestock breeding program for climate resilient.
Further, encouraging the active participation of women is also crucial as they possess
rich knowledge and efficient management skills especially for maximizing the use of
natural resources. Hence, organizing occasional trainings and participatory research
approach into the roles of women can assists to tackle climate change impact from
the rural areas point of view. In addition, well-organized and accurate early warning
systems can play a pivotal role by avoiding the severe damages that especially occur
22 Goat as the Ideal Future Climate Resilient Animal Model 289

during unexpected disasters, by providing sufficient time for the policy maker and
people to prepare effective response. Another crucial strategy involves development
of skilled disease surveillances supported with effective health services. This is often
neglected despite the fact that they can effectively control the spread of the diseases
especially those that are climate change induced. Furthermore, so as to meet with the
water requirements for goat production, especially in tropical regions, improved
water resource management should be developed. Another efficient adaptive strat-
egy involves the cultivation of drought tolerant fodder varieties in hot and dry areas
as this can ensure feed supply during scarcity period. Finally, the role of extension
services to build awareness through capacity building programs among the livestock
keepers is vital to improve their adaptive capacities against climate change. Hence,
there is an urgent need to improvise the policies and practices so as to ensure cost
effective adaptive strategies to combat climate change. Figure 22.5 describes the
different adaptation strategies that are vital to sustain goat production in the changing
climate scenario.

Fig. 22.5 Different adaptation strategies to sustain goat production in the changing climate
scenario
290 V. Sejian et al.

22.8 Conclusions

The limited available literature clearly project goat as the ideal climate resilient
animal model. As the scientific community are looking forward to identify ways and
means to ensure food security by 2050 to feed the expected human population of 9.6
billion, it is very essential to invest on goat promotion. Such approaches can ensure
economic security of the weaker sections of poor and marginal farmers. However,
there are lot of basic information are lacking in this neglected species till date. Hence
research efforts needed to generate baseline information on the indigenous breeds of
goats. Such efforts would help the farming community to recommend ideal species
specific to specific agro-ecological zone. Generating the baseline information might
help to identify the targeting point for intervention in terms of sustaining goat
population in the changing climate scenario. Further, appropriate extension system
should be in place to transfer these information generated to the ultimate target
groups of farmers. Capacity building program should be developed to cater the need
of different stakeholders involved in goat production. Research efforts are also
needed to develop standard package of practices comprising shelter management,
nutritional interventions and disease management for goats. Additionally, genetic
approach should be focussed to identify more advanced thermo-tolerant markers
with high heritable values to incorporate them in marker assisted selection program
to develop more climate resilient goat breeds to disseminate to poor and marginal
farmers to ensure their livelihood.

Learning Outcomes
• Goats are considered ideal climate resilient species because of its better
thermo-tolerance, drought tolerance, ability to survive on limited pastures
as well as the disease resistance capacity in tropical region.
• Differences in climate resilience were achieved even among the indigenous
goat breeds. This warrants more such efforts in screening all indigenous
goat breeds to identify the best breed for the specific agro-ecological zone.
• Genetic approach to develop more climate resilient goat breeds should
focus on identification of more advanced biomarkers with high heritable
values for incorporation in marker assisted selection programs.

22.9 Future Perspectives

Substantial evidences are now available to project goat as the ideal climate resilient
species and therefore future research must focus on generating baseline information
pertaining to adaptive variables in all unexplored indigenous breeds. Such an
approach would help the scientific fraternity to identify the best climate resilient
indigenous goat breeds which could survive and produce optimally in a given agro-
ecological zone. Such identified breed could be disseminated to the poor and
22 Goat as the Ideal Future Climate Resilient Animal Model 291

marginal farmers for ensuring their livelihoods. Further such identified best climate
resilient breeds has to be subjected for testing their survival and production ability in
multiple locations. This way it is possible to identify a region specific breeds which
has the ability to survive in multiple locations and produce optimally. Such breeds
should be subjected for biomarkers identification and such markers could be used in
future breeding programs to produce new goat breeds with potential to survive and
produce optimally in multiple agro-ecological zones. The policy makers must
propagate such breeds for ensuring sustainable goat production in the changing
climate scenario by designing appropriate capacity building programs for different
stakeholders and providing additional incentives to farmers for rearing such breeds.

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Future Vision for Climate Change
Associated Livestock Production 23
Veerasamy Sejian, Surinder Singh Chauhan, Chinnasamy Devaraj,
Pradeep Kumar Malik, E. Vadhana, M. V. Silpa, C. G. Shashank,
and Raghavendra Bhatta

Contents
23.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 295
23.2 Thermal Indices with More Applicability . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 295
23.3 Predictive Livestock Early Warning Systems (PLEWS) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 295
23.4 Applications of Advanced Molecular Tools for Inducing Thermo-Tolerance . . . . . . . . 297
23.5 More Focus on Goat as Future Food Animals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 297
23.6 Interrelationship Between Soil, Climate and Livestock . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 298
23.7 Climate Smart Livestock with A Farmer Centric Approach . . . . . . . . . . . . . . . . . . . . . . . . . . 298
23.8 Improved Balanced Animal Nutrition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 298
23.9 Genetic Selection and Breeding Management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 299
23.10 Low Carbon Livestock . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 299
23.11 Artificial Meat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 299
23.12 Vaccine Technology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 300
23.13 Innovations in Methane Reduction: Methane-Oxidizing Nose Ring . . . . . . . . . . . . . . . . . 300
23.14 Technologies to Assess Animal Welfare: Precision Livestock Farming . . . . . . . . . . . . . . 301
23.15 Nanotechnology in Animal Production . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 301
23.16 Climate-Smart Animal Agriculture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 302
23.17 Exploiting the Genetic Potential of Local/Indigenous Breeds . . . . . . . . . . . . . . . . . . . . . . . . 302

V. Sejian (*) · C. Devaraj · E. Vadhana · R. Bhatta

Centre for Climate Resilient Animal Adaptation Studies, ICAR-National Institute of Animal
Nutrition and Physiology, Bangalore, India
S. S. Chauhan
Faculty of Veterinary and Agricultural Sciences, The University of Melbourne, Parkville, Australia
P. K. Malik
ICAR-National Institute of Animal Nutrition and Physiology, Bangalore, India
M. V. Silpa
Institute of Animal Breeding and Genetics, Justus-Liebig-Universität Gießen, Gießen, Germany
C. G. Shashank
ICAR-National Dairy Research Institute, Karnal, India

# The Author(s), under exclusive license to Springer Nature Singapore Pte 293
Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1_23
294 V. Sejian et al.

23.18 Exploiting the Unconventional Feed Resources . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 303

23.19 Identification of Agro-ecological Zone Specific Breeds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 303
23.20 Water Resources Management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 304
23.21 Bridging the Knowledge Gap Between Climate Change and Livestock Diseases . . . 304
23.22 Concluding Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 305
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 306

Abstract

Climate change is inevitable and thus there is an immediate need to plan for the
future amelioration and mitigation strategies to sustain livestock production. This
chapter is therefore aimed to shed some light on few supreme strategies that need
to be considered while planning to tackle future climate change effects in
livestock sector. To begin with, improvisation of thermal indices which reveal
the impact of stress is necessary. Furthermore, adoption of predicted livestock
early warning systems, advanced molecular tools to assess thermo-tolerance and
also assess animal welfare, incorporation of nanotechnology in animal production
and so on are some of the proposed strategies. Additionally, a lot more consider-
ation should also be given to animal management and livestock farming practices
like, improvising the nutritional charts in animals, encouraging climate smart
livestock farming keeping a regular check on the soil, climate and livestock
harmony. Along with these, approaches to reduce the livestock further contribu-
tion to climate change should be considered. Animal breeding is another major
division that can play a promising role in future livestock farming in the face of
global warming threat. Screening large livestock populations, breeding for
climate-resilient livestock, actively incorporating the local indigenous germplasm
and further substantiating the target to identify agro-ecological zone specific
breeds are some of the vital aspects to be considered. The proposed approaches
to effectively combat climate change are of great relevance and hence these
concepts should be disseminated across the globe to ensure future sustainability
in livestock farming.

Keywords
Food security · Livestock · Policy · Sustainability · Technologies

Abbreviations

% Percentage
CO2 Carbon dioxide
CSA Climate smart agriculture
FAO Food and Agriculture Organization
GHG Greenhouse gas
GS Genomic selection
GWP Global warming potential
23 Future Vision for Climate Change Associated Livestock Production 295

IgG Immunoglobulin gamma

MAS Marker assisted selection
NCFR Non-conventional feed resources
PLEWS Predictive livestock early warning systems
PLF Precision livestock farming

23.1 Introduction

The future livestock production in the changing climate scenario requires paradigm
shift in the existing policies to develop and propagate new strategies and
technologies which could cater the need of growing human population both by
2050 and 2100 (Fig. 23.1). The livestock adaptation strategies developed should
be focused and teller made to suit the projected future climatic condition. A wide
range of strategies comprising early warning system, advanced thermal indices;
identifying more appropriate livestock species; climate smart livestock production;
applications of molecular tools; balanced nutrition; low carbon livestock; artificial
meat; breeding management; vaccine technologies; advancements to assess animal
welfare; vaccine and nanotechnologies applications; promoting indigenous animals;
exploiting unconventional feed resources; water resource management and under-
standing in depth climate change associated livestock disease occurrences. These
strategies are discussed in brief in this chapter probably to give an overview to the
readers on the possible future technologies which could play a significant role in
sustaining livestock production in the future climate change perspectives.

23.2 Thermal Indices with More Applicability

Although there are several thermal indices available to quantify heat stress response,
still there are lots of scopes for improvement. Advanced thermal indices
incorporating all cardinal weather variables such as temperature, humidity, wind
velocity and solar radiation needs to be developed. Such indices should be developed
separately for specific agro-ecological zones. Such approach could help to generate
useful basic information which may play vital role in determining the appropriate
amelioration strategies to sustain livestock production in the changing climate
scenario.

23.3 Predictive Livestock Early Warning Systems (PLEWS)

The PLEWS involves a combination of advanced technological and predictive

modelling functions that amalgamates the impact of climatic extremities, weather,
and can also improve food security concerns especially among the most vulnerable
296 V. Sejian et al.

Fig. 23.1 Future strategies to sustain livestock production in the changing climate scenario
23 Future Vision for Climate Change Associated Livestock Production 297

populations. Such predictive models eliminates the time gap between a warning and
its required intervention thereby providing an anticipatory action rather than the
usual combative action. Furthermore, as a consequence of this feature, the PLEWS
are also thought to drastically cut down the costs associated with early warning.
Additionally, PLEWS has an advantage of strengthening the resilience of
populations at-risk, ensure better disaster mitigation strategies and also provide
more time to the communities and global actors for preparation, planning and
mitigation rather than only response. However, though such advancements promise
great possibilities to combat the adversities of climate change, it can be effective
only when adopted satisfactorily by the end users. It is equally necessary to educate
the potential users about the functions, notifications and all other relevant details
associated with such facilities and also their application towards decision making
process (Matere et al. 2020).

23.4 Applications of Advanced Molecular Tools for Inducing

Thermo-Tolerance

The latest molecular biological tools offer huge scope for improving thermo-
tolerance in domestic animals. The latest technologies such as metagenomics,
transcriptomics, bisulfite sequencing, genome wide association studies and selection
signature may play a greater inroad to understand in depth or may help to elucidate
further the hidden intricacies of molecular mechanisms governing livestock adapta-
tion to climate change. Such information may prove very vital in identifying more
reliable biomarkers for climate resilience in farm animals. This could revolutionize
future animal breeding to evolve more climate resilience breeds which has the
potential to survive and produce optimally in a given location.

23.5 More Focus on Goat as Future Food Animals

Recent research findings using modeling approach clearly projects goat as the ideal
climate resilient animals (Nair et al. 2021; Carvajal et al. 2021). Their drought
tolerance, thermo-tolerance potential, anatomical advantage for browsing, ability
to survive on poor quality and limited pastures, and disease resistance potential in
tropical regions imparts them the potential to survive and produce optimally in
multiple agro-ecological zones. With the projected climate change associated reduc-
tion in both green and dry fodders as well as reduction in water resources makes
large animal farming very difficult. Based on the above advantages associated with
goat and the very low initial investments required for goat farming makes it a go to
go species for ensuring food security by 2050 scenario. Hence the focus should shift
to investing on goat farming which otherwise remains a neglected species in many
parts of the world. Such approach could help to sustain goat production to ensure
animal protein for the expected 9.6 billion peoples by 2050.
298 V. Sejian et al.

23.6 Interrelationship Between Soil, Climate and Livestock

Strategies need to be developed for efficient grassland management aimed at

optimizing its capacity to serve as a carbon sink. More supports needs to be provided
by the government to encourage grazing. Similarly, the land-use change should be
brought under strict control, including that related to imported animal feed. Further-
more, the livestock production should have a stronger link to the regional feed data
base (Idel et al. 2013).

23.7 Climate Smart Livestock with A Farmer Centric Approach

There are opportunities to transform livestock sector in low and middle income
countries through farmer centric approach to make the sector more profitable and
sustainable both socially and ecologically. Creating a global livestock climate-smart
livelihoods fund could be the way forward approach for creating employment
opportunities and building resilient livelihoods of farming households. The crucial
factor which could determine this is the global partnership involving the farmers on
the ground to the global organisations looking to protect our environment for the
next generation. This approach has to be amalgamated with public-private
partnerships and this becomes increasingly important for testing innovations, speed-
ing up change and taking success to scale. In this we can build a more sustainable
global livestock sector.

23.8 Improved Balanced Animal Nutrition

Ration balancing is a very effective means to sustain livestock production in addition

to achieving low emission. This involves (1) formulating balanced feed rations with
high-quality feed ingredients; (2) improving feed conversion rates and forage digest-
ibility, through effective silage and hay preservation; (3) better grassland manage-
ment, e.g. by introducing legumes into grass pasture; (4) selecting better feed
resources, including alternative feed products and feed additives; (5) supplementa-
tion with small amounts of concentrate feed and mineral/urea/molasses n; and
(6) low emission feed. Reducing the amount of feed required per unit of output
(e.g. beef, milk) has the potential to reduce greenhouse gas emissions and increase
farm profits. Feed efficiency can be increased by developing breeds that grow faster,
are hardier, gain weight more quickly, or produce more milk. Rotational grazing is
another approach to effectively manage grasslands. This would stop land degrada-
tion and increase soil carbon sequestration. Further this will improve the quality of
the diet as the animals will be consuming forages in a relatively earlier growth stage.
Low emission feed resources is achieved by feeding by-products from agriculture
and the agri-food industry or feed that has been produced by using conservation
agriculture. Other low emission feed products include insect-based feeds, and
specific feed additives (FAO 2021).
23 Future Vision for Climate Change Associated Livestock Production 299

23.9 Genetic Selection and Breeding Management

Breeding management and genetic selection has the potential to form the formidable
climate smart livestock production comprising both mitigation and adaptation
strategies. The future breeding strategies must concentrate on marker assisted selec-
tion comprising traits pertaining to adaptation, production and low methane emis-
sion. Conservation of local animal genetic resources may play an important role in
making available elite animals for the farmers. Involving local breeds in breeding
program offers scope for maintaining traits related to thermo-tolerance, coping
ability to poor nutrition and disease resistance (FAO 2021).

23.10 Low Carbon Livestock

Low-carbon livestock can help countries achieve a balance whereby animal-source

foods feed the hungry and malnourished, yet are produced in a way that is
minimizing the overall output of greenhouse gases. FAO (2019) suggests that this
could be achieved by five ways: (1) Boosting efficiency of livestock production and
resource use; (2) Intensifying recycling efforts and minimizing losses for a circular
bioeconomy; (3) Capitalizing on nature-based solutions to ramp up carbon offsets;
(4) Striving for healthy, sustainable diets and accounting for protein alternatives;
(5) Developing policy measures to drive change.

23.11 Artificial Meat

Cultured meat (also known as artificial, in vitro or lab grown meat) promises to be
the best alternative to satisfy the growing demand of highly increasing population. It
might drastically reduce the need of rearing animals for meat production, though
fewer animals have to be grown for the purpose of harvesting cells for production of
in vitro meat. Muscle cells will be cultured in a culture medium consisting of
nutrients, growth factors and hormones, such that the muscle cells proliferates and
in turn produces lumpsum amount of meat from minimum number of cells (Baca-
González et al. 2020). In spite of owning advantages from animal welfare and human
health point of view by producing pathogen free meat, it bears its own disadvantages
like usage of hormone and growth factors, palatability issues, etc. Being a new
technology, it faces its own critical points such as struggle to fetch public accep-
tance, huge price, religious debate and so on. Henceforth, with all its pros and cons,
still there is long way of research needed in multiple dimensions to overcome all the
questions raised and to overcome all the drawbacks.
300 V. Sejian et al.

23.12 Vaccine Technology

Enteric methane production, being the burning topic of interest, its contribution
towards GHG pool is quite alarming and the efforts are repeatedly posted towards
reducing it. There are various methods suggested in pipeline to mitigate enteric
methane emission, among which vaccination against methanogens attracted the
attention of many scientists around the globe. Major advantage is that its ability to
leave no traces of residues in the product and its potential of universal applicability to
all ruminant farm animal species. In a nut shell, it is all about eliciting immune
response in rumen against methanogens. Since, rumen lacks any lymphoid tissue,
the idea is to use saliva as a immunoglobulin delivering vector into the rumen (Baca-
González et al. 2020). This mitigation strategy is at its initial stage of research, with
minimal literature available to compare and get a conclusive idea. There are lots of
bottleneck hampering the evolution and utilization of this strategy in each step of its
development like choice of antigen, selection of appropriate adjuvant, limited IgG
transfer from blood to saliva, survival of immunoglobulins in the rumen, determina-
tion of booster frequency and finally an exact evaluation method to analyze the
evoked immune response. In addition, scarce literature availability majorly holds
back the technique from successful implementation in the field. Henceforth, there is
vast area of science need to be studied before successful implementation of anti-
methanogens vaccination in practice. Therefore, more intensified research efforts are
needed to standardise the protocol for a possible universal vaccine against rumen
methanogens and such a breakthrough in near future can help to reduce the livestock
associated climate change.

23.13 Innovations in Methane Reduction: Methane-Oxidizing

Nose Ring

There are few emerging innovative technologies to combat the effect of enteric
methane emission (Norris 2018). This technology uses a device in the form of a
simple nose ring, to reduce the methane exhaled by ruminants which was produced
through the process of enteric methane fermentation. A smart cattle nose-ring is
attached to the cattle’s nose. Whenever the animal exhales, this device detects the
methane in the exhaled air, which activates the micro-oxidation chamber in the
device. This is turn leads to the conversion of methane into CO2 and water vapour.
This device is expected to convert upto 80% of exhaled methane. In addition, it also
transmits data which shall be stored for the animal’s lifetime. Carbon dioxide, being
a green house gas with very low global warming potential (GWP) than methane, if
studied in detail, this technology could emerge as a promising alternative to reduce
the impact of enteric methane emission caused by animals.
23 Future Vision for Climate Change Associated Livestock Production 301

23.14 Technologies to Assess Animal Welfare: Precision

Livestock Farming

To meet out the blooming demand for meat and milk production, increasing live-
stock production becomes inevitable. This demand for increase in livestock produc-
tion should not compromise animal welfare. Henceforth, the need for some
innovative technologies which could help to ensure livestock production without
compromising animal health and welfare rises. Precision livestock farming (PLF), is
one of its kind designed to monitor and control animal productivity, along with
animal health and welfare parameters in an automated manner (Schillings et al.
2021). Various technologies such as cameras, sensors or microphones are used to
caution the farmers then and there, such that farmers will remain informed about any
undesirable conditions of animals at the earliest. PLF technologies have highly
encouraging potential to reduce the occurrence of disease and injuries. Any technol-
ogy will have its own drawbacks. Likewise, PLF might reduce the human animal
relationship by reducing the frequency of interaction between animals and farmers,
which likely to reduce the opportunity of animals to become acclimatize with people.
In addition, the extent to which the PLF technologies could help to improve the
animal welfare seems to be limited. Literature and evidences are lacking to prove
PLF technologies as welfare indicators. Having said that, integrating such PLF
technologies with climate change prediction tools can also aid to assess and predict
heat stress or environmental stress events in livestock thereby giving the livestock
keepers sufficient time to adopt suitable ameliorative and mitigation strategies.

23.15 Nanotechnology in Animal Production

Nanotechnology, a very novel field of research, which has already been applied in
certain areas of animal production, is a scientific approach to study the matter in its
atomic and molecular scale, usually in nanometer (nanoparticles). These are very
small entities which have large surface-to-volume ratio, which in turn provide them
various unique properties. Nanoparticles helps in improving the absorbability,
solubility, half-life and bioavailability of various products. Nanomaterials,
Nanosensors and microfluidics are few nanotechnology devices which helps in
improving animal health, reproduction, production, diagnosis and treatment of
diseases. Nanotechnology has the promising potential to improve livestock produc-
tion system. It is an eco-friendly and economical way to control pathogenic micro
organisms (Fesseha et al. 2020). But the major challenge is all about the environ-
mental contamination that might be caused by this very technology. The major risk
being, if released into the environment, the threat that might be faced by workers and
consumers, pose a serious concern from safety risk perspective. In addition, it is
readily accepted that few nanoparticles can evoke toxic and dangerous side effects
upon ingestion. Even though, it is one of the major novelties in the current era, it is
still in its prime stage of development. There are much more hampering factors from
animal and people welfare perspectives which need to be studied in depth. Wide
302 V. Sejian et al.

hazard analysis and assessments need to be performed before implementing nano-

technology for livestock production.

23.16 Climate-Smart Animal Agriculture

The FAO has identified Climate Smart Agriculture (CSA) as one among the
approaches to address the interlinked challenges of climate change and food secu-
rity. The FAO has bridged the knowledge gap associated with the terminology of
CSA by defining it as ‘the integrated approach to enhance the capacity of the
agricultural systems to support food security, incorporating the need for adaptation
and the potential for mitigation into sustainable agriculture development strategies.
The CSA work with the aim of achieving increased productivity, enhanced resilience
and reduced emission from agricultural sector. It involves various components like
crops, livestock and trees, arranged in an effective way such that the available
resources are effectively utilized and recycled for generating maximum productivity.
Although substantial progress has been made in successfully implementing CSA,
still such efforts should be more intensified to take up in mass scale throughout the
world in the future. Such efforts require development of more innovative
technologies to achieve the basic principle of CSA. Implementation of such
technologies at farmers’ field requires innovative policies and therefore the policy
makers must engage all stakeholders to develop a common policy compromising all
aspects of environmental influence on livestock production. The government also
must develop reliable financial mechanisms which could help to implement such
innovative technologies by the ultimate targeted beneficiaries of farmers.

23.17 Exploiting the Genetic Potential of Local/Indigenous

Breeds

It is an unequivocal fact that indigenous germplasm have a higher significance in the

current changing climatic scenario. These indigenous livestock however lag behind
the established exotic breeds for their lower production ability. Hence it is the need
of the hour to conduct extensive studies to assess and improvise the productive
performance of indigenous germplasm. Furthermore, with the concept of
establishing agro-ecological zone specific livestock breeds (Sejian et al. 2021); it
would also be advisable to conduct studies that assess the performance of livestock
breeds in a given local environment. The advancements in the field of molecular
biotechnology have made it possible to identify several biomarkers for climate
resilience. Stimulation studies conducted in climate controlled chambers have
made is easier to subject animals to varied environmental conditions across the
global by programming the desired temperature and humidity variables. Such
experiments thereby enable researcher to subject animals to a diverse combination
of environmental stress and with the use of the advanced biotechnological tools can
also identify potential markers for climate-resilience. Identification of such markers
23 Future Vision for Climate Change Associated Livestock Production 303

can pave way towards Marker Assisted Selection (MAS) or Genomic Selection
(GS) which can ultimately aid towards breeding of agro-ecological zone specific
germplasm.

23.18 Exploiting the Unconventional Feed Resources

In the current scenario, in order to meet out the imperative demand for alternative
feed resources, utilization of non-conventional feed resources (NCFR), popularly
known as ‘new feeds’, could be exploited for livestock production (Amata 2014).
But this valuable source of feed supply is facing challenges in every step of its
development like collection, storage and importantly during detoxification
procedures. Major constraint faced by NCFR is its anti-nutritional factors, which
can be tackled by utilizing the emerging processing methods which includes physi-
cal, chemical and biological processing. Basically, NCFR involves variety of feed-
stuff such as shrub fodder, tree fodder, agro-industrial by-products, animal
by-products, single cell proteins and so on. Among them, agro-industrial
by-products, browse foliage and crop residues are gaining importance in the farming
community. Furthermore, considering the rapid depletion of fresh water resources
and increase in the saline agriculture, use of haplotype plants as NCFR may satisfy
major critical points such as food and water demand. Haplotype such as Acacia
species, are highly saline tolerant and these plants should be fed with energy rich
feed supplements like molasses, corn, etc., in order to overcome the effect of its anti-
nutritional factors (Abd El-Hack et al. 2018). Practically, from the view of landless
and marginal farmers, Cactus (thornless) and Azolla can also be considered as
foremost NCFR (Keerthi et al. 2019). Azolla, being easily cultivable and economi-
cal, also it serves as the best protein substitute. Cactus is the best plantation in highly
water deficit and drought prone areas and it shall be an ideal NCFR when mixed with
other fibrous feedstuffs. Apart from the above stated feed/fodder sources, a variety of
other NCFR have been fed to livestock like, sugarcane tops, Moringa oleifera,
mango seed kernel, mango pulp, jackfruit waste, cassava, pineapple waste, tamarind
seed hulls and so on (Singh 2018; Onte et al. 2019). NCFR can serve as best feed
substitutes, when considerable attention is given along with the involvement of
farmers and local extension agencies.

23.19 Identification of Agro-ecological Zone Specific Breeds

Indigenous animals developed the traits to survive in a specific location and they
acquire such potential from generation after generation. However, even among
indigenous breeds, differences were established in their responses to climatic
stresses. Therefore, it is very essential to channelize new research initiatives to
identify the most suited indigenous breeds to specific agro-ecological zones. Such
efforts can help the farming community to reap maximum by using the most
appropriate breeds for rearing. Efforts are also needed to test the potential of
304 V. Sejian et al.

indigenous breeds to survive in different location and those breeds which exhibit the
potential to survive and produce optimally in different agro-ecological zones must be
identified and disseminated apart from using them in breeding programs. These
efforts can secure the livelihoods of poor and marginal farmers by helping them to
rear the most appropriate breed to generate income.

23.20 Water Resources Management

Water is another vital resource that needs great attention especially in the prevailing
climate change scenario. Understanding the water footprint associated with the
livestock sector is therefore imperative. Additionally, appropriate methodologies
should also be adopted to conserve water resources and sustain fodder production.
Broadly, the practical measures to conserve fodder production during water scarcity
are categorized into land management, irrigation management, in-situ water
harvesting and ex-situ rainwater harvesting. Among all these, the management of
irrigation water is considered to be the most harmonizing methodology for water
conservation. Land management measures primarily target the reduction of water
erosion and loss of water from soil surface. Crop rotation is yet another effective and
feasible practice that aids in the reduction of water shortage risk. In-situ land
management, a self-explanatory terminology, comprises of strategies that involves
conversion of landscapes into permanent and/or temporary structure that conserve
water, like terraces, bunds to cover rainwater, thus reducing soil erosion. While on
the other hand, ex-situ water conservation practices consists of construction of
permanent structures; dams and artificial ponds; that store excess rainwater and
also support irrigation during water scarcity. With the projected impact of climate
change on the global perspective and also the predicted rise in the global tempera-
ture, it is therefore necessary to implement multiple water conservation strategies.
Such strategies not only will address the water scarcity concern but can also enhance
effective forage production which is also necessary to meet the rising human and
livestock demand.

23.21 Bridging the Knowledge Gap Between Climate Change

and Livestock Diseases

Interrelating climate change and sudden disease outbreaks is one of the most crucial
aspect to reduce animal production. However, not much information on this line is
currently available. More intensified research efforts are required to interlink climate
change associated disease outbreak in livestock. Particularly the vector-borne
diseases are very important as the weather variables directly influence the multipli-
cation of these vectors which acts as the causative agent for the disease. To find
solution to the problem such sudden disease outbreaks have to be linked with
geographical information system to establish its connection with changing climatic
conditions. This has to be followed with more field and laboratory oriented
23 Future Vision for Climate Change Associated Livestock Production 305

researches. Such attempts should be agro-ecological zone or country specific.

Tackling climate change associated disease outbreak in livestock and developing
solution to the problem through laboratory research may go in a long way to sustain
livestock production. As it is one of the important impact which causes drastic
reduction in animal production even among the indigenous breeds.

Learning Outcomes
• The latest technologies such as metagenomics, transcriptomics, bisulfite
sequencing, genome-wide association studies, and selection signature may
play a greater inroad in understanding the hidden intricacies of molecular
mechanisms governing livestock adaptation.
• The future breeding strategies to evolve new climate-resilient breeds must
concentrate on marker-assisted selection comprising traits pertaining to
adaptation, production, and low methane emission.
• The developed adaptation strategies and designed technologies should be
agro-ecological zone specific to cater the needs of the poor and marginal
farmers to secure their livelihood.

23.22 Concluding Remarks

Investing in livestock sector in future becomes inevitable probably due to their better
climate resilience than other sectors in agriculture. Thus, the livestock sector could
play a vital role in ensuring food security to feed the growing human population by
2100. Therefore, strategies needs to be worked out by policy makers to identify most
suitable strategies which are cost effective and may play a significant role to ensure
sustainable livestock production in the future. All the identified strategies must be
concentrated around ensuring optimum production in livestock in the changing
climate scenario. A systematic road map needs to be developed keeping in view
the anticipated increase in average earth temperature to realistically implement the
suitable strategies for augmenting livestock production. A multidisciplinary
approach should be in place to effectively utilize the strategies developed to achieve
the goal towards sustainable livestock production. Further, the developed strategies
and designed technologies should be agro-ecological zone specific to cater the need
of the poor and marginal farmers of the location. From the available strategies, only
those which are promising should be narrowed down for up scaling by the policy
makers for propagating them under field condition. Such systematic approach could
help the farming communities to sustain livestock production in the changing
climate scenario.
306 V. Sejian et al.

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Index

A Agricultural sector, 5, 115, 302

Abdominal temperature, 104 Agriculture, 13, 87, 101, 170, 199, 200, 226,
Abiotic stress, 62, 256 282, 283, 298, 302, 303, 305
Abomasum, 220, 241, 242 Agroclimatic zone, 133
Abscess, 217 Agro-ecological zone, 79, 115, 117, 124, 125,
Absorption, 46, 168, 217, 242, 254, 265 284, 290, 291, 295, 297, 302–305
Accelerometer, 13, 95, 96, 107, 108 Agro-net roof, 134
Accelerometer-based technologies, 107, 108 Air exchange, 136, 139
Acclimation, 51, 65, 66, 143, 266 Airflow, 96, 135
Acclimatisation, 266 Air movement, 64, 135, 136, 180, 215
Acetate, 164, 165, 167, 205, 218, 227, 228, 242 Air temperature (AT), 6, 43, 89, 93, 94, 132,
Acetogenesis, 205, 218 136
Acid detergent fibre (ADF), 155, 225 Algorithm, 34, 97, 104, 106, 252
Acute heat stress, 7, 17, 121 Alveolar ventilation, 189
Acute stress, 55, 66, 90 Ambient conditions, 104, 137, 191
Adaptability, 17, 22, 224, 235, 272, 284 Ambient temperature, 7, 18, 51, 64, 69, 77, 93,
Adaptations, viii, 8, 13, 22, 30–37, 47, 48, 51, 162, 163, 165, 167, 177, 187–189, 216
52, 64, 65, 67, 71, 76, 77, 79, 124, 131, Ameliorate, viii, 6, 96, 144–148, 162–172,
138, 139, 143, 148, 163, 171, 193, 176–183, 187, 189, 285
264–276, 282, 288–289, 295, 299, 302, Amelioration, viii, 69, 138, 142–148, 187,
305 203–206, 210, 295
Adaptive immunity, 53, 54 Ammonia, 132, 137, 139, 207
Adaptive mechanism, 18, 65, 75, 267, 285 Amplicon sequencing, 245, 250, 251
Adaptive response, vii, 9, 56, 189, 217 Amplification, 248–252
Additive effect, 34, 146, 166 Amylolytic, 227
Additive response, 171 Anaerobic, 18, 137, 201, 202, 204, 207, 209,
Additives, 34, 146, 165, 166, 171, 219, 221, 225, 226, 228, 234, 242, 244, 245, 247
256, 298 Anatomy, 241–242
Adjuvant, 300 Angiogenesis, 276
Adrenal-thyroid axis, 67 Animal, 5, 16, 30, 41, 51, 61, 75, 87, 101, 115,
Adrenergic stress response, 18 131, 145, 153, 163, 176, 187, 199, 214,
Adults, 42, 77, 154 225, 241, 264, 281, 295
Aeration, 132, 209 Animal agriculture, 87, 101, 282, 302
Aerobic, 137, 204, 243 Animal behaviour, 20, 41–48, 63, 71, 107, 135
Age, 7, 9, 35, 36, 66, 108, 153, 154, 191, 216, Animal factor, 21
221, 244 Animal feed, 147, 170, 209, 298
Agricultural land, 71, 115, 176 Animal husbandry, 87, 88, 94
Agricultural products, vii, 5, 115 Animal manure, 131, 204, 207

# The Author(s), under exclusive license to Springer Nature Singapore Pte 307
Ltd. 2021
V. Sejian et al. (eds.), Climate Change and Livestock Production: Recent Advances
and Future Perspectives, https://doi.org/10.1007/978-981-16-9836-1
308 Index

Animal production, 5–13, 41, 43, 153–158, Baseline temperature, 104

164, 165, 255, 265, 301–302, 304, 305 Beef, 6, 7, 16–22, 30–37, 78, 145, 156, 199,
Animal productivity, 12, 17, 63, 71, 131, 193, 241, 298
218, 220, 235, 256, 265, 273, 301 Beef cattle, 7, 18–21, 30–37, 78, 145
Animal protein, 5, 30, 153, 158, 176, 283, 297 Beef production, 22, 31, 36
Animal response, 32, 101, 107 Behavior, 8–9, 12, 13, 42–43, 87–89, 97, 116,
Animal restraint, 101, 102 182, 282
Animals health, 13, 17, 54, 62, 106, 165, 169, Behavioural mechanism, 47, 161, 285
218, 220, 274, 301 Behavioural responses, 8, 9, 41–48, 64, 95, 187
Animal source, 153, 157, 158, 299 Biocatalyst, 274
Animal-source foods, 153, 157, 158, 299 Biochemical, 10, 77, 79, 119, 143, 189
Animal welfare, 5–13, 17, 42, 51, 87, 88, 91, Biochemical adaptation, 143
96, 120, 133, 139, 176, 181, 191, 193, Biodegradable, 204
256, 287, 299, 301, 401 Biogenesis, 70
Antagonism, 32 Bio-hydrogenation, 206
Anthropogenic, 200, 208 Bioinformatics, 256
Antibodies, 53 Biological intervention, 210, 287
Antigen, 53, 56, 300 Biological markers, 62, 76
Anti-inflammatory, 57, 78 Biological reserve, 63
Anti-methanogens vaccination, 300 Biomarkers, 63, 90, 120, 124, 189, 273,
Anti-nutritional factors, 303 287–288, 290, 291, 297, 302
Antioxidants, 11, 19, 79, 144–148, 166, Biotechnology, 32, 274, 302
168–169, 171, 180 Biotic stress, 62, 256
Apoptosis, 57, 69, 270 Biotopes, 48
Archaea, 202, 204–206, 209, 210, 215, Birthweight, 177, 179, 181
225–235, 242, 243, 245, 248, 249, 253, Bisulfite sequencing, 123, 297, 305
255 Blood pressure, 65
Arid, 42–46, 48, 283 Blooms, 63
Artificial insemination, 34 Body condition score (BCS), 6, 7, 138, 189,
Artificial meat, 295, 299 285
Artificial selection, 30, 266, 270 Body core temperature, 91, 92, 120
Ascorbic acids, 180 Body mass, 42, 241
Atmosphere, 199, 200, 202 Body mass index, 241
Atomic, 301 Body reserve, 62, 63, 70, 116
Australia, 17, 19, 21–22, 108, 147, 153, 154, Body surface temperature, 42, 91, 101–103,
177, 179, 188, 189, 191 121
Australian Merino, 178 Body temperature (BT), 18, 20, 41, 43, 47,
Automated tympanic probes, 95 63–65, 69, 87, 91–96, 101, 102, 104,
Autonomic nervous system (ANS), 18 106, 107, 109, 121, 135, 179, 187, 188,
Autonomic response, 41, 44 190, 191
Average daily gain (ADG), 64, 166, 189 Body weight, 6, 7, 13, 64, 138, 189, 216
Azolla, 303 Bos indicus, 21, 22, 77, 190
Bos taurus, 33, 65, 77, 190, 254, 267
Brahman, 22, 106
B Breed, 7, 22, 31, 45, 65, 76, 104, 115, 163, 178,
Backcrossing, 190 188, 224, 249, 265, 284, 297
Bacteria, 122, 147, 202, 209, 215–219, Breeding, 22, 32–36, 62, 75, 76, 79, 88, 117,
225–229, 232, 234, 235, 242–245, 124, 125, 164, 170, 171, 178–182,
247–251, 253–255 187–193, 205, 265, 273, 288, 291, 295,
Bacteriophage, 225, 228, 242, 245 297, 299, 303–305
Bacteroidales, 254 Breeding programs, 22, 33–36, 62, 124, 170,
Bacteroides, 215, 225, 227–229, 252, 253, 255 171, 178, 193, 265, 273, 288, 291, 299,
Bacteroidetes, 215, 225, 228, 229, 253, 255 304
Index 309

Breeding season, 180, 181 Chronic stress, 55, 67, 90, 273
Breeding values (BVs), 34–36, 191, 265, 273 Ciliates, 205, 225, 227–229, 250
Buffaloes, 7, 9, 10, 78, 92, 120, 136, 203, 206, Circular bioeconomy, 299
209, 224–235 Climate, 5, 17, 31, 42, 61, 75, 87, 100, 115,
Bushfires, 16–22 131, 142, 162, 187, 199, 255, 264, 281,
Butyrate, 218, 228, 242 295
Butyrivibrio, 226, 254 adaptation, viii, 131, 139, 163, 193,
288–289, 299
adoption, 131
C animal model, 116, 281
Cactus, 303 change, vii, viii, 3–13, 15–22, 33, 35, 36,
Calcium, 11, 131, 168, 182 42, 48, 59–71, 75, 87, 88, 95, 100, 101,
Candidate gene, 273 115, 117–119, 121, 123, 129–139, 142,
Capacity building program, 289–291 162, 163, 165, 170, 171, 178, 193, 199,
Capital outlay, 176 255, 264, 266, 272, 274, 281, 285, 288,
Carbon dioxide (CO2), 65, 189, 200, 202, 208, 289, 293–305
210, 218, 226–228, 242, 300 controlled chambers, 302
Carbon sequestration, 131, 298 extreme, 283
Carcase, 16, 17, 20, 21, 34, 36 resilience, 9, 10, 36, 113–125, 192, 281,
quality, 17, 189 287, 290, 297, 302, 305
weight, 7, 16, 20, 22 smart livelihood, 298
yield, 20 smart livestock production, 295, 299
Cascade of inflammation, 78 stress, 137
Cascade reaction, 18 variability, 6, 31, 115
Catabolism, 10, 67 Climate smart agriculture (CSA), 302
Cattle, 6, 17, 30, 53, 62, 75, 90, 106, 120, 134, Climatic extremes, 51, 101, 138
143, 152, 164, 188, 203, 215, 224, 241, Climatic extremities, 119, 295
265, 300 Climatic stress, 132, 285
Cell, 11, 19, 33, 53, 62, 77, 144, 166, 189, 225, Cloning, 34, 244, 248, 251, 255
242, 266, 287, 299 Clostridiales, 254
damage, 69, 78, 189 Coat characteristics, 65, 102
function, 144 Cold, 19, 20, 43, 46, 133, 154, 178
growth, 270 Cold stress, 31, 41, 133
mediators, 267 Cold-tolerant, 87
motility, 270 Colonization, 215
proliferation, 55, 56, 70, 166, 303 Colostrum, 43, 181, 182
signalling, 62, 71 Combined stress, 65, 287
Cell-mediated immunity, 56 Concentrate, 144, 145, 155, 164, 166, 167, 180,
Cellular damage, 78 205, 215, 216, 298, 299, 305
Cellular osmolarity, 145 Conception rate, 178, 179
Cellular response, 56, 57, 70, 121, 266, 276 Conduction, 136, 137
Cellulolytic, 147 Contagious, 76
Cellulolytic bacteria, 229, 242 Contaminants, 102, 135
Cellulose, 225, 226, 228, 229, 234, 241, 242 Contigs, 253
Census, 75, 203, 254 Convection, 42, 43, 136
Central nervous system, 41 Cooking loss, 20, 189
Cervus elaphus, 254 Cooler zone, 178
Cheese production, 7 Cooling pond, 137
Chemical barrier, 78 Cow, 10, 21, 33, 42, 52, 64, 76, 87, 104, 132,
Chromatography, 120 142, 153, 188, 205, 218, 224, 252, 284
Chromosomal localization, 269 Critical temperature, 41, 135
Chromosomes, 253, 267 Crop rotation, 304
Chronic heat stress, 8, 17, 66, 68, 101 Crossbred, 73–80, 117, 122, 134, 138, 178, 203
310 Index

Crossbreed, 36 Digestion, 64, 104, 122, 145, 167, 168, 200,

Crossbreeding, 75, 76, 190, 192, 193, 266, 274 205, 206, 218, 226, 227, 229, 241–243,
Crossing over, 267 254, 265
Cultivation, 5, 137, 200, 289 Digestive system, 225, 242, 270
Cultured meat, 299 Dilution, 247
Cyclicity, 177 Discomfort, 12, 94
Cytoprotection, 56 Disease, 6, 11–13, 31, 35, 43, 52, 57, 76, 79, 92,
Cytotoxicity, 19, 69 101, 116, 117, 148, 177, 179, 191, 193,
214, 256, 265, 266, 284, 285, 289, 290,
295, 301, 304–305
D outbreaks, vii, 5, 13, 304, 305
Dairy, 6, 32, 42, 52, 62, 75, 87, 132, 153, 165, resistance, 34, 163, 244, 255, 265, 281, 290,
188, 217, 241, 265, 283 297, 299
Dairy cattle, 6, 7, 11, 35, 65, 67–69, 75, 76, tolerance, 117
85–97, 138, 143, 145, 148, 154, 164, vulnerability, 11
167, 169, 170, 188–190, 241, 265 Dissipation, 41, 42, 64, 69, 70, 119, 189
Danger-associated molecular pattern (DAMP), Diurnal rhythm, 104, 105
54 Diversity, 35, 75, 115, 121, 122, 124, 206, 215,
Dark colour meat, 19 217, 223–235, 243–245, 247, 249–252,
Dark-cutting meat, 17–22 266, 267
Dark, firm and dry (DFD) meat, 8 DNA methylation, 123
Data analysis, 107, 274 Domestic animals, 39–48, 264, 297
Data loggers, 104 Doping test, 91
Data management, 106 Dose-dependent response, 145
Decomposition, 204, 209, 234 Dose response, 146
Deer, 250, 254 Down-regulation, 55, 56
Defecation, 9, 95 Downstream regulation, 270
Defense, 11, 51, 56, 123 Downstream signals, 78
Defense barrier, 117 Drip loss, 20
Defensive response, 78 Drooling, 108
Deficiency, 67, 153, 154, 193 Drought, vii, 31, 131, 303
Defoliation interval, 157 Drought master, 22
Dehydration, 18, 43, 165 Drought tolerance, 281, 290, 297
Deletion, 267 Dry matter intake (DMI), 64, 67, 87, 156, 165,
Demand, 5, 13, 30, 41, 61, 75, 80, 143, 148, 167, 169, 216, 218
157, 176, 179, 199, 218, 219, 265, 281, Duplication, 267
283, 284, 299, 301, 303, 304 Dysbiosis, 244, 256
Denaturation, 56 Dyspnea, 94
Denitrification, 207
De novo synthesis, 167
Deoxymyoglobin, 19 E
Deoxyribonucleic acid (DNA), 34, 36, 54, 120, Early warning, 297
123, 191, 247–252, 254, 266, 270, 273 Early warning system, 285, 288, 295–297
Depletion, 22, 115, 282, 303 Ear tag sensors, 107
Detoxification, 245, 303 Earth, 43, 62, 115, 131, 139, 305
Diagnosis, 218, 301 East Africa, 7, 282
Differentially expressed genes (DEGs), 70, 122, Ecology, 234, 245, 251
124 Economic loss, vii, 6, 12, 16, 62, 64, 96, 187
Differential pressure, 89 Economic returns, 7, 285
Differentiation, 53, 270, 272 Economic trait, 6, 7, 13
Digesta, 228, 242 Economic value, 7
Digestibility, 62, 167, 168, 205, 225, 226, 235, Economy, 16, 17, 131, 224, 284
285
Index 311

Ecosystem, 6, 75, 115, 122, 214, 215, 218, 220, Eukaryote, 225, 243, 248
225, 247 Evaporation, 18, 43, 45, 65, 136, 143, 189
Egg production, 6, 7, 13 Evaporative cooling, 41, 45, 136, 137, 165
Electric current, 252 Evaporative heat loss, 65, 135, 143
Electrode, 252 Evaporative water loss, 43, 45, 48, 64, 116
Electron microscopy, 245 Evolution, 62, 244, 272, 300
Elite animals, 299 Ewes, 52, 176–182
Embryo, 35, 36, 177, 179, 181 Excretion, 18, 65, 145, 168, 189, 206–209
mortality, 177 Excretion rate, 204, 207
transfer, 34 Exotic, 7, 75, 117, 190, 192, 193, 302
Emission, vii, viii, 92, 115, 116, 131, 132, 137, Extension agencies, 303
139, 168, 176, 199–210, 218–220, 225, Extensive, 8, 71, 101, 104, 108, 116, 123, 131,
226, 244, 253, 255, 256, 285, 288, 298, 138, 177, 178, 302
300, 302, 305 grazing systems, 176, 177, 180
Emissivity value, 102 system, 71, 75, 138, 189
Endocrine, 9–10, 51, 52, 54, 56, 57, 65, 66, 68, Extrinsic factors, 265
70, 77, 87, 119, 165, 265 Eye temperature, 101
mechanism, 51
response, 52, 65, 66, 90, 95, 96
Endogenous, 54, 65, 69 F
Endotoxin, 218 Farm animals, 8, 13, 47, 78, 79, 87, 91–96, 124,
Energy, 18, 19, 22, 41, 42, 45, 51, 63, 64, 125, 147, 241, 273, 283, 297, 300
66–68, 75, 136, 143–146, 153, 155, 157, Farmers, 13, 42, 45, 76, 80, 88, 96, 116, 120,
164, 166–170, 176, 180, 181, 189, 206, 125, 131, 136, 137, 153, 158, 191, 192,
208–210, 215, 218, 220, 226, 241–243, 205, 208–210, 281, 283, 284, 290, 291,
265, 266, 270, 303 298, 299, 301–305
balance, 67, 68, 144, 167, 180, 182 Farming, 96, 108, 116, 137, 170, 176, 281,
density, 145 283–284, 290, 297, 298, 301, 303, 305
reserve, 18, 68 Farm management, 179, 214
Ensiling, 220 Fasting, 18
Enteric fermentation, 200, 208, 210, 226 Fat, 7, 17, 20, 22, 64, 75, 144, 147, 166–169,
Enteric methane, viii, 116, 197–210, 215, 180, 189, 206, 242, 253
218–221, 226, 255, 285, 300 score, 17
Enteric methane emission, viii, 201, 204, 206, stores, 64
208, 210, 218–220, 226, 285, 300 yield, 7, 227
Enterprises, 137, 177, 283 Fecal cortisol estimation, 90
Environment, 17, 30–35, 41–46, 51, 62, 66, 67, Fecal cortisol metabolites, 90
69, 76, 80, 87, 93, 94, 100, 101, 106, Fecundity, 180–181
108, 115, 116, 118, 123, 131, 134, 135, Feed additives, 219, 221, 256, 298
164, 168, 170, 171, 176–179, 187–190, Feed conversion efficiency, 167, 209, 225
193, 201, 215, 224, 226, 227, 241–245, Feed conversion rate, 298
247, 265–267, 273, 274, 298, 301, 302 Feed efficiency, 35, 169, 170, 191, 220, 226,
Environmental stress, 66, 77, 88, 122, 123, 301, 241, 253, 256, 283, 298
302 Feed intake, 6–8, 10, 18, 19, 33, 64–67, 69, 95,
Environmental stressors, 6, 13, 62, 77 105, 138, 142, 143, 145–148, 163,
Environmental temperature, 164, 170, 187, 188, 166–169, 177, 180, 187–189, 204, 206,
216 215, 220
Enzyme, 10, 67, 70, 225, 226, 228, 234, 244, Feed intervention, 205
249, 254, 255, 267, 274 Feedlot cattle, 20
Epigenetics, 32, 123–124 Feed resources, 205, 210, 283, 295, 298, 303
Epigenome, 273, 274 Fermentable grains, 167
Equilibrium, 43, 45, 65, 76, 189, 228, 265, 266 Fermentation, 17, 147, 148, 164, 165, 167, 169,
Ethological response, 87 172, 200–202, 208, 210, 213–221,
312 Index

225–229, 235, 242, 244, 245, 253, 255, Gene, 8, 33–36, 53–57, 69, 70, 122–124, 189,
300 190, 193, 214, 221, 225, 243–245,
Fertilisation, 177 247–251, 253–256, 265–273, 275–277,
Fertilisation rate, 177 287
Fertility, 16, 31, 32, 34, 36, 177, 179–181, 192, Gene editing, 273, 274
284 Gene-environmental interaction, 123
Fetal growth, 177, 181 Gene expression, 32, 56, 57, 69, 120, 123, 266,
Fibre, 65, 153, 155, 163, 167, 171, 206, 225, 267, 270
229, 242 Generation interval, 33
Fibre digestion, 205, 206 Genetic, 21, 31, 48, 63, 75, 96, 121, 148, 163,
Fibrolytic, 215, 216, 230 176, 187, 210, 235, 241, 265, 290, 299
Filtration, 90, 242 adaptation, 48, 263–277
Fingerprints, 250 change, 33–36, 266
Firmicutes, 15, 225, 227–229, 252, 253, 255 difference, 188
Fixation index, 272 diversity, 35, 121, 267
Flagellated, 227 drift, 266, 272
Flank temperature, 147 evaluation, 35
Fleece, 179 fingerprinting, 249
Fleece rot, 179 gain, 176, 191
Flocks, 177–179 improvement, 32, 34, 36, 190
Flora, 243 makeup, 35, 241
Fluid, 43, 188, 215, 229, 233, 234, 242, 301 markers, 178, 265
Fluorescent, 249 progress, 34
Flystrike, 180 resources, 190, 274, 299
Fodder, 5, 6, 116, 165, 191, 203, 205, 209, 210, selection, 163, 185–193, 265, 299
283, 289, 297, 303, 304 strategies, 22
Fodder crops, 199 traits, 63
Foetus, 33 trend, 35
Follicular development, 177 up-gradation, 210
Food and agriculture organization (FAO), 131, Genome, 31, 35, 123, 191, 192, 244, 245,
153, 154, 157, 199, 200, 226, 298, 299, 252–256, 265–267, 270, 272–274, 277,
302 297, 305
Food safety, 8 editing, 265
Food security, vii, 5, 7, 61, 63, 71, 80, 101, 115, estimated breeding value, 36, 191
117, 124, 131, 157, 163, 241, 255, expression, 62, 71
281–283, 290, 295, 297, 302, 305 sequence, 35, 36, 228
Forages, 30, 62, 116, 145, 153, 156, 158, 162, Genome wide association study (GWAS), 120,
163, 167, 180–182, 216, 217, 226, 228, 191, 266, 274
265, 282, 298, 304 Genomic, 35, 36, 123, 187, 190, 191, 193, 235,
Forced ventilation, 136, 138, 139 247, 265–267, 269, 272–274, 288
Foregut, 249 assay, 274
Free radicals, 19, 144, 168 diversity, 266
Free-ranging, 42, 48 selection, 35, 36, 190–193, 265, 273, 274,
Freis wal, 78 303
Fungi, 53, 225, 227, 228, 234, 242, 248 Genotype, 188, 190, 191, 193, 235, 241,
265–267, 270
Genotypic markers, 287–288
G Genus, 215–217, 221, 230, 232–234, 251–253,
Galactopoiesis, 52 255
Gases, vii, 115, 137, 139, 143, 169, 199, 200, Geographical information system, 272, 304
204, 209, 227, 242, 243, 299 Germplasm, 34, 117, 131, 190, 281, 302, 303
Gastrointestinal tract, 241 Gestation, 32, 178
Gel electrophoresis, 245, 249, 250 Global food security, 71, 241
Index 313

Global surface temperature, 7 Heat, 6, 16, 30, 41, 51, 61, 75, 87, 101, 115,
Global temperature, 63, 100, 163 131, 143, 163, 176, 187, 215, 264, 282,
Global warming, 22, 115, 187, 193, 210, 215 295
Global warming potential (GWP), 208, 210, dissipation, 41, 42, 69, 70, 119, 189
300 exchange, 41–43, 139
Gluconeogenesis, 52, 66–68, 143, 144 gain, 42–46, 76, 134, 164
Glycogenesis, 18 increment, 92, 147, 164, 165, 167, 172
Glycogenolysis, 8, 17, 18, 189 load, 12, 42–44, 46, 48, 51, 52, 64, 65, 69,
Glycogen reserve, 17–19, 22 77, 87–90, 94–96, 106, 107, 132, 139,
Goat, 7, 42, 52, 78, 90, 115, 167, 189, 203, 267, 147, 164, 167, 180, 188, 215, 266
281, 297 load index, 95
Goat manure, 284 load influx, 44
Grain supplement, 180 loss, 42, 43, 65, 101, 121, 133, 135, 136,
Gram negative, 226 143, 144, 169, 215
Gram positive, 226 period, 87
Grass-seeds, 179 regulation, 64
Grazing, 13, 30, 42, 45–48, 71, 108, 133, 138, resilience, 178, 190
146, 147, 164, 171, 176–178, 180–182, resilient breeds, 178
189, 209, 229, 281, 298 resistance, 131
Green feed, 181 sensitive, 87, 187–189
Greenhouse gas (GHG), 131, 200, 204, 208, Heat shock element (HSE), 69
209, 227, 243, 299, 300 Heat shock factor (HSF), 69
Greenhouse gas (GHG) emission, vii, 115, 116, Heat shock proteins (HSP), 8, 33, 54, 57, 69,
137, 139, 199–201, 204–210, 298 77, 168, 189, 266, 280
Green pasture, 144 Heat stress, vii, viii, 6, 16, 30, 41, 51, 61, 75, 87,
Green revolution, 158 101, 115, 131, 142, 163, 176, 187, 215,
Gross domestic product (GDP), vii, 115 264, 295
Growth, 16, 30–32, 36, 61–64, 66, 68–70, 75, indicator, 78
122, 145, 153, 155, 157, 162, 163, 165, response, 51, 54, 57, 69, 85–97, 106, 108,
177, 181, 182, 209, 216, 225, 226, 235, 115, 119, 124, 270, 273, 287, 288, 295
244, 247, 256, 265, 267, 270, 284, 285, Heat tolerance, 22, 31–33, 35–37, 69, 70, 119,
287, 298 124, 178, 187–193, 265, 273, 274
factors, 9, 68, 270, 299 Heat transfer, 42, 102
hormone, 9, 67, 68, 77, 287 Heat transmission, 137
rate, 7, 17, 64, 75, 133, 179, 284 Heat wave, 101, 131
Gut, 90, 145, 164, 166, 168, 229, 234, 241, 244, Heifers, 20, 21, 35, 36, 64, 68, 77, 155, 156,
249, 252–254, 256 192, 216
Hematocrit (HCT), 189
Hematology, 11, 119
H Hemicellulose, 228, 229, 234, 242
Habitat, 33, 225, 251 Hemoglobin (HGB), 189
Hair cortisol, 90, 95, 120, 124 Herbivore, 241
Hair sheep, 178 Heritability, 34, 36, 265, 273
Haplotypes, 190, 265, 303 Heterogeneity, 254, 267
Harvesting cells, 299 High-roughage, 167, 225, 229
Harvest interval (HI), 154–156 Hindgut, 249
Hay preservation, 298 Hind gut fermenters, 241
Hazard analysis, 302 Histone modification, 123
Health, 8, 11–13, 16, 17, 19, 33, 51, 54, 62, 69, Hitchhiking effect, 270
87, 91, 92, 106, 116, 138, 157, 163, 165, Holobiont organism, 241
168, 169, 171, 199, 202, 209, 214–216, Holstein heifers, 64, 68, 156, 216
218, 220, 235, 243, 244, 264, 274, 283, Homeostasis, 19, 41, 51, 57, 65, 66, 68, 69, 71,
284, 289, 299, 301 76, 78, 119, 120, 170, 264, 266
Heart rate (HR), 18, 65, 94, 164 Homeotherms, 87
314 Index

Homeothermy, 62, 92, 117, 187, 188 Implantable devices, 48, 91, 95, 96
Homopolymer, 251 Incubation, 220
Honeycomb, 242 Independent assortment, 267
Hormonal growth promotants (HGP), 21, 22 Indigenous, 8, 10, 73–80, 117, 119, 121, 138,
Hormones, 9, 10, 13, 52, 56, 66–69, 77, 90, 299 203, 224, 281, 284, 285, 287, 290, 295,
Host, 12, 54, 118, 122, 123, 202, 205, 206, 208, 302–304
210, 214, 215, 218, 220, 221, 225, 226, Indigenous breed, 76, 79, 284, 290, 303–305
230, 234, 235, 241–243, 245, 255, 256, Infertility, 176
274 Inflammation, 54, 57, 101, 147, 276
Host response, 54 Infrared radiation, 101
Hot carcase weight, 20 Infrared thermal imaging, 92–93, 95, 99–109,
Hot temperature, 178 121, 124
Housing system, 132–134 Infrared thermography (IRT), 92, 96, 101–104,
Human, vii, 5, 8, 12, 13, 32, 33, 53, 61, 63, 92, 106, 107, 121
93, 115, 120, 131, 151–158, 163, 205, Infrared thermometer, 92, 121
226, 241, 242, 244, 253, 281–283, 290, Infrastructure, 36, 177, 244
295, 299, 301, 304, 305 Ingesta, 242
Human health, 299 Innate immunity, 53–57, 235
Human population, 5, 12, 13, 61, 115, 131, 205, Inoculation, 247
282, 283, 290, 295, 305 Insect-based feeds, 298
Humidity, 6, 12, 21, 64, 136, 138, 143, 163, Insemination, 34, 177
164, 176, 179, 191, 265, 295, 302 Integrated farming system (IFS), 137
Hunger poverty, 154 Intergovernmnetal panel on climate change
Husbandry, 87, 88, 94, 181, 182 (IPCC), 62, 100, 101
Hydrogen, 215, 218, 219, 228, 242 Intermittent wetting, 136, 138
Hydrogen disposal, 218 Intracellular, 53, 54, 57, 78, 276, 277
Hydrogenotrophic, 227 Intra rectal device, 91, 95, 96
Hydrogen sulphide, 242 Intra vaginal device, 91, 95, 96
Hyperthermia, 55, 56, 178, 188 Intrinsic factors, 265
Hypervariable regions, 248, 249, 251 Introgression, 190, 193, 265
Hypoglycemia, 52 Inversion, 267
Hypothalamo-pituitary-adrenal (HPA) axis, 9, In vitro meat, 299
11, 52, 55, 66, 77, 120 Isoelectric point, 19
Hypothalamus, 41, 51, 64–66, 68, 270 Isolation, 31, 121, 171, 265
Hypothermia, 178, 181

J
I Jersey, 10, 11, 35, 68, 77, 135, 191, 225, 229,
IgG transfer, 300 230, 232
Illumina technology, 251, 253
Immune activation, 57
Immune barrier, 122 K
Immune cell, 11, 53–56, 78, 267 Kanni Aadu, 8
Immune competence, 56 Karan fries, 76
Immune function, 33, 53, 146, 182 Kikuyu grass, 155, 156
Immune response, 12, 51–57, 78–79, 87, 168,
182, 189, 270, 287, 300
Immune system, 13, 49–57, 70, 163, 165, 265, L
267, 270, 276 Lab grown meat, 299
Immunity, 11, 33, 53, 54, 56, 57, 177, 235 Lachnospiraceae, 249, 253, 254
Immunoglobulin, 300 Lactation, 32, 88, 89, 143, 147, 148, 181, 217,
Immunological markers, 287 218, 252
Immunology, 256 Lactogenesis, 52
Immunostimulatory, 55 Lactose, 7, 143
Immunosuppression, 55 Lactose synthesis, 143
Index 315

Lamb, 42–45, 103, 177–179, 181–182, 188, Magnesium, 11, 182

249 Maintenance energy, 145, 180
Lambing, 178, 179, 181, 182 Malabari, 10, 117, 121, 284, 287
Lambing period, 178, 181, 182 Malnutrition, 153, 157, 158
Lamb mortality, 177, 178, 181 Management strategies, 42, 46, 142, 148, 158,
Laminitis, 217 176, 209, 285
Layers, 7, 77, 139 Manure, 131, 132, 137, 200, 201, 204–210, 284
Legume, 156, 298 Manure management system, 137, 200, 207,
Lignin, 62, 234, 241, 242 208
Lignocellulose, 234 Mapping, 35, 92, 101, 270, 272
Linkage, 265, 274 Marbling, 17, 20
Linkage-disequilibrium, 265 Marbling score, 20
Lipogenesis, 67 Marker assisted selection (MAS), 79, 125, 273,
Lipolysis, 67–68, 144 290, 299, 303, 305
Lipolytic, 52, 67, 68 Maternal behaviour, 181
Liquid culture, 247 Mating, 176–182
Livelihood, vii, 75, 76, 80, 115, 125, 157, 281, Mating period, 177–180
283, 284, 290, 291, 298, 304, 305 Maturity score, 20
Livelihood security, vii, 80, 281, 284 Meat, 6–8, 13, 16–22, 30, 32, 34, 36, 153, 155,
Livestock, 5, 18, 36, 42, 51, 61, 75, 87, 101, 157, 158, 163, 166, 171, 176, 189, 235,
115, 131, 148, 153, 163, 187, 199, 219, 283–285, 295, 299, 301
224, 241, 265, 281, 295 Meat production, 7, 22, 32, 36, 285, 299, 301
Livestock census, 75, 203 Meat quality, 7, 8, 16–22, 34, 163, 189, 235
Livestock farming, 96, 108, 116, 170, 281, 283, Meat quality traits, 7
301 Meat yield, 6, 13
Livestock husbandry, 87, 94 Membrane integrity, 145, 168
Livestock production, vii, viii, 5, 6, 8, 12, 13, Merino, 178, 179, 188, 189
61–71, 88, 115, 117, 118, 123–125, 131, Messenger ribonucleic acid (mRNA), 52, 68,
138, 158, 187, 192, 200, 273, 281, 282, 78, 120, 121, 248
284, 295–305 Meta-analysis, 164, 166, 169, 230
Livestock products, 5, 7, 61, 157, 205, 281 Metabolic acidosis, 18
Livestock sector, vii, 5–7, 13, 75, 115, 199, Metabolic activity, 76
200, 224, 225, 285, 288, 298, 304, 305 Metabolic capacity, 234
Livestock species, 6, 7, 10, 11, 45, 67, 78, 90, Metabolic changes, 33, 66–69
92, 116, 119, 121, 123, 187–190, 203, Metabolic diseases, 256
207, 208, 241, 265, 281–283, 295 Metabolic disorders, 182
Liveweight (LW), 22, 153, 155, 156, 166, 167 Metabolic heat, 9, 10, 17, 18, 43, 64, 65, 67, 68,
Local breeds, 190, 192, 193, 273, 284, 299, 135, 145, 147, 164, 167, 188, 215, 217
302–303 Metabolic heat production, 9, 18, 43, 64, 65, 67,
Loci, 34, 265, 267, 270, 272 145, 167, 188
Loin yield, 7 Metabolic markers, 10
Longevity, 31, 34 Metabolic processes, 245, 254, 270, 275
Longissimus muscle area, 20 Metabolic profile, 11
Loose housing system, 133 Metabolic rate, 17, 65, 66, 68, 143
Low carbon livestock, 295, 299 Metabolic responses, 9, 10, 18, 22, 171, 188
Lupins, 180 Metabolism, 10, 11, 43, 45, 51, 52, 63, 66, 67,
Lupinus angustifolius, 180 69, 70, 143, 144, 146, 165–169,
Lymphoid tissue, 300 218–220, 226, 227, 256, 265–267, 270,
Lytic phages, 228 287
Metabolites, 10, 11, 13, 90, 201, 202, 205, 206,
219, 220, 270, 274
M Metabolome, 273, 274
Machine learning, 106, 107, 274 Metagenome profile, 235
Macro environment, 118 Metagenomics, 121–122, 124, 241–256, 274,
Macronutrient, 144–145 297, 305
316 Index

Metaproteomics, 245, 255 188–191, 217, 224, 226, 227, 253, 283,
Metatranscriptomics, 245, 253, 255 285, 301
Meteorological data, 94 Milk protein, 169
Methane, viii, 92, 116, 131, 137, 168, 176, Milk quality, 7, 137
199–210, 215, 218–221, 225–228, Milk yield, 6, 7, 13, 34, 52, 75, 89, 94, 138, 142,
242–244, 253–256, 265, 274, 284, 285, 144–148, 154–157, 164, 166–169, 215,
288, 299, 300, 305 227, 265
Methane emission, viii, 137, 176, 199, 201, Mineral balance, 51, 145
203, 205, 206, 208–210, 218–220, 225, Minerals, 7, 10, 51, 145, 165, 166, 168, 169,
226, 244, 255, 256, 274, 285, 288, 300, 171, 180–182, 284, 298
305 Minimally-invasive technologies, 107
Methane-oxidizing nose ring, 300 Mithuns, 203
Methanobacteria, 255 Mitigate, 6, 20, 88, 143–145, 147, 171, 180,
Methanobrevibacter, 227, 230, 234, 255 208, 210, 300
Methanogen, 225, 230, 234 Mitochondria, 19
Methanogenesis, 202–206, 218, 228, 253, 256 Mitogens, 56
Methanogenic archae, 218 Mixed model, 274
Methanotrophic, 230 Molasses, 298, 303
Methylotrophic, 227 Molecular biology, 34, 244
Microarray, 266 Molecular biotechnology, 32, 302
Microbes, 12, 121, 201, 204–206, 217, 225, Molecular marker, 79, 189, 270
226, 229, 230, 241, 242, 244, 245, Molecular mechanism, 8, 76, 297, 305
247–248, 252, 254 Molecular response, 57, 69, 70, 285
Microbial biomass, 227 Monensin, 219
Microbial community, 121, 214–215, 220, 221, Monogastrics, 145, 225, 241
227, 229, 235, 244, 245, 248–250, 256 Morbidity, 12
Microbial diversity, 122, 124, 224–235, 244, Mortality, 11, 12, 43, 64, 69, 133, 164,
247, 250, 252 176–179, 181, 182
Microbial protein, 147, 219, 226 Mortality rate, 69, 133, 164
Microbiologists, 244 Mouse, 53
Microbiology, 226, 244 Multiple climatic stresses, 285
Microbiome, 121, 122, 167, 171, 215, 220, 225, Multiple stresses, 9, 62–64, 70, 71
226, 228–230, 232–235, 241–247, Multiplexing, 251
250–256, 273, 274 Muscle protein, 19, 241
Microbiota, 122, 215, 220, 221, 225–227, 241, Muscles, 8, 17–22, 52, 70, 166, 189, 241, 299
244, 245, 247, 252, 254, 255 Mutations, 191, 248
Micro-circulation, 169 Mycoplasma, 245
Microclimate, 43, 48, 87, 94, 124, 131, 135, Myofibril, 19
171, 281 Myostatin (MSTN), 8
Microenvironment, 95, 117–118, 242
Microflora, 242, 244
Microfluidics, 301 N
Micronutrients, 131, 148, 285 Nanomaterials, 301
Microorganisms, 53, 54, 226, 244, 245, Nanoparticles, 301
247–249, 254, 256, 274 Nanopores, 252, 253
Micro-oxidation chamber, 300 Nanosensors, 301
Micro ribonucleic acid (miRNA), 57 Nanotechnology, 295, 301–302
Midgut, 249 Napier grass, 152–158
Milk composition, 7, 13, 75, 138 National research council (NRC), 89, 164, 167
Milk fat, 7, 75, 147, 168, 169, 227, 253 Native breeds, 79, 117, 170
Milk-fat yield, 7, 227 Natural selection, 31, 33
Milk production, 6, 31, 62, 75, 79, 87, 133, 136, Negative energy balance, 67, 68, 144, 180
138, 142, 143, 145, 147, 154, 169, 177, Negative feedback regulation, 69
Index 317

Neonates, 42–45, 47 Operative temperature, 42, 43

Neuro-endocrine response, 285 Organic, 147, 166, 204, 214, 215, 219, 225,
Neutral detergent fibre (NDF), 155, 225 228, 275
New generation sequencing (NGS), 124 Organic matter, 204, 225
New Zealand, 177 Osmanabadi, 117, 121, 284, 287
Next-generation sequencing (NGS), 250, 252, Osmoprotectant, 166
266 Osmotic balance, 166
Nitrate, 205, 207 Osmotic stress, 166
Nitrification, 207, 209 Ovis aries, 254
Nitrous oxide (N2O), 131, 200–202, 204, Ovulation rate, 177, 179, 180
206–209 Oxidation, 11, 17, 19, 207
Nocturnal, 44, 45 Oxidative burst, 56
Non-biodegradable, 204 Oxidative stress (OS), 19, 22, 56, 70, 144, 146,
Non-conventional feed resources (NCFR), 303 148, 165, 168
Non-esterified fatty acids (NEFA), 10, 67, 68, Oxidative stress index, 168
143, 144, 166, 189 Oxygen, 19, 56, 144, 168, 207, 242
Non-invasive methods, 87–97, 101, 104, 106,
120, 121
Non-pregnant, 177, 179 P
Non-protein nitrogen, 225, 242 Paddock, 42, 45, 133, 136, 164, 181
Non ruminants, 171 Pale, soft and exudative (PSE) meat, 8, 17
Nuclear transfer, 34 Panting, 18, 41, 44, 65, 106–109, 120, 143,
Nucleic acids, 54, 120, 249, 250, 252 145, 188
Nutrients, 17, 52, 62, 66, 67, 69, 71, 157, 163, Panting rate, 65
165, 170, 217, 229, 235, 242, 244, 254, Panting score, 106–109, 188
265, 282, 299 Parasites, 12, 30, 31, 53, 273
Nutrition, 18, 31, 62, 64, 65, 69–71, 131, Parsimonious model, 21
152–158, 163–169, 180–182, 226, 227, Passive immunity, 33
241, 244, 245, 256, 284, 295, 298, 299 Pastoralism, 138
Nutritional interventions, 143, 145, 148, 163, Pasture, 5, 17, 19, 21, 45, 71, 94, 108, 116, 131,
166, 180, 182, 187, 193, 217, 218, 285, 144, 145, 156, 164, 180, 181, 204, 205,
290 209, 281, 282, 290, 297, 298
Nutritional management, 176 Pasture quality, 19, 180, 181
Nutritional strategies, 22, 138, 143–148, 166, Pathogen, 228, 265, 299
171 Pathogen associated molecular patterns
Nutritional stress, 9, 62, 142–148, 287 (PAMPs), 53
Nutritional supplements, 138, 163–179 Pathogen free meat, 299
Nutrition balance, 64 Pectin, 220, 242
Nutrition security, 153 Pedigree, 35, 37
Nutrition stress, 62, 65, 69–70 Peri-natal, 181
Nutritive value, 154, 158 Phagocytosis, 56
Phenotype, 33–36, 187–191, 193, 253, 265,
267, 272–274
O Photovaltaic panels, 42
Oestrous behaviour, 177 Phylogenetic, 215, 227, 248, 274
Oestrus, 106, 177, 181 Phylogenetic marker genes, 248
Offspring, 33, 181 Phylogenies, 243
Omasum, 241, 242 Phylotypes, 249
Omental fat, 7 Phylum, 215, 251, 252
Omics, 273, 274 Physical barrier, 53, 78
On-farm strategies, 176 Physiological adaptation, 67, 120, 142
Ontology, 267, 270–272, 277 Physiological effect, 67
Oocyte quality, 177 Physiological mechanism, 30
318 Index

Physiological parameters, 64, 136, 138 252, 266, 267, 274, 276, 277, 283, 297,
Physiological response, 76, 87, 94, 95, 146, 299, 303
147, 267 aggregation, 70
Physiological stress, 67 disintegration, 70
Physiological traits, 64, 65, 69, 121 nanopore, 252
Physiology, 9–11, 77, 144, 163, 165, 187, 217, Proteome, 273, 274
230, 241–242, 244, 273 Protozoa, 205, 225–229, 232–235, 242, 245,
Phytopathogen, 234 247
Picolitre, 252 Pulse rate, 64, 65, 119, 188
Pigs, 7, 10, 11, 17, 91, 92, 137, 145, 146 Pyrosequencing technology, 251
Placenta, 177
Plant polysaccharides, 242
Ploidy, 267 Q
Polyacrylamide gels, 250 Quantitative trait loci (QTL), 34, 265
Polymorphisms, 34, 35, 265, 266 Quantitative traits, 34
Polysaccharide, 220, 225, 228, 229, 242, 253
Poor man’s grass, 153
Population, 6, 11, 21, 34, 61, 75, 80, 115, 116, R
121, 122, 148, 155, 163, 190, 193, 199, Radiant heat, 17, 42, 43, 45, 46, 92, 134, 144,
200, 203, 205, 216, 217, 221, 226, 179
228–230, 232, 234, 235, 244, 245, 248, Radiant temperature, 44, 45
249, 253, 255, 256, 265–267, 270, 272, Radiation, 6, 9, 12, 41–47, 94, 96, 101, 102,
273, 281–283, 290, 295, 297, 299, 305 119, 134, 163, 164, 176, 179–180, 215,
Population mean, 34 295
Post-mortem, 19, 20 Radiative heat transfer, 102
Post-natal, 177 Rams, 92, 176–180, 182
Post-slaughter, 20 Rangelands, 204
Potassium, 11, 145, 168, 180, 284 Rangifer tarandus, 254
Poultry, 6, 7, 10, 13, 17, 136, 137, 145 Rations, 165, 180, 205, 209, 220, 298
Poultry meat, 7 Reactive oxygen species (ROS), 19, 54, 144,
Precision livestock farming (PLF), 108, 170, 168
301 Receptor, 51, 53, 54, 57, 68, 77
Precocial, 43–45 Rechewed, 242
Predictive livestock early warning systems Rectal temperature (RT), 51, 64, 65, 76, 91,
(PLEWS), 295–297 119, 121, 145–148, 164, 166, 168, 169,
Pregnancy, 89, 177, 178, 181, 182 188, 284, 287
Pregnancy rates, 177 Regression, 21
Pregnancy scanning, 179 Regurgitation, 242
Pre-mating, 178 Relative humidity, 12, 18, 89, 93, 94, 132, 136,
Pre-slaughter stress, 19 179, 215, 216
Prevotella, 226, 227, 229, 254, 255 Re-mating, 177, 179
Primers, 248–251 Renewable, 131, 209
Production, vii, viii, 5, 17, 31, 41, 54, 61, 75, Reproduction, 13, 31, 34, 63, 64, 68, 91, 163,
87, 101, 115, 131, 142, 153, 163, 176, 165, 175–183, 215, 265, 267, 287, 301
187, 200, 214, 224, 241, 264, 281, 295 Reproductive cycle, 178, 180
Pro-inflammatory, 54–57, 78, 79 Reproductive efficiency, 35, 176
Prokaryote, 202, 225, 243, 247, 248 Reproductive performance, 31, 132, 182
Propionate, 164, 167, 218, 228, 242 Reproductive rate, 176, 179
Protein, 5, 7, 10, 17, 19, 22, 30, 33, 35, 51, 54, Reproductive stage, 178
56, 57, 61, 64, 66–70, 75, 77–79, 131, Resistance, 31, 34, 68, 77, 116, 133, 163, 178,
145–147, 153, 155, 157, 158, 163, 164, 191, 192, 244, 255, 256, 265, 266, 273,
166, 167, 169, 176, 180, 181, 189, 199, 275, 281, 284, 290, 297, 299
206, 219, 220, 225–229, 241, 242, 248, Respiration, 120, 143, 145, 200
Index 319

Respiration frequency, 121 Saline agriculture, 303

Respiration rate, 51, 64, 65, 87, 89, 94–96, Saliva, 242, 288, 300
107–109, 119, 143, 146, 147, 164, 166, Sanger sequencing, 250
168, 169, 188, 189, 284, 287 Saponin, 219, 221, 229, 230
Respiratory alkalosis, 18, 65, 144, 189 Scrotal, 101
Respiratory dynamics, 107 Seasonal pattern, 131
Respiratory evaporation, 45, 189 Seasonal variation, 77
Respiratory quotient, 143 Selection, 17, 22, 30–37, 48, 80, 125, 131, 132,
Response, vii, viii, 6, 8–10, 12, 18, 19, 22, 32, 163, 164, 170, 178, 181, 182, 185–193,
33, 39–57, 62–70, 76–79, 85–97, 101, 244, 250, 251, 265, 266, 270, 272–274,
106–108, 115, 118, 119, 121, 123, 124, 290, 297, 299, 300, 303, 305
136, 143, 145–147, 163, 166–169, 171, Semen, 178, 180
182, 187–189, 193, 215–217, 230, 244, Semen sexing, 34
249, 264–267, 269, 270, 273–276, Semi-arid, 42, 43, 45–48, 119, 283
285–289, 295, 297, 300, 303 Semi-intensive system, 75, 138
Restraining, 91, 101 Sensor-based parameters, 96
Restriction enzymes, 249 Sensor chip, 252
Reticulum, 241, 242 Service, 71, 131, 177, 289
Reticulum-rumen, 242 Sex, 7, 20, 34, 36, 66, 90, 119
Rhizoids, 228 Sex-limited traits, 36
Ribeye area, 20 Shade, 8, 20, 42, 44–47, 104, 132–134, 142,
Ribonucleic acid (RNA), 120, 215, 228, 243, 148, 164, 171, 176, 179–182, 187, 189
247, 248, 252, 255, 256, 273 Shearer, 179
Ribosomal RNA (rRNA), 243, 247, 249, 255 Shearing, 176, 179–180, 182
Rigour mortis, 20 Sheep, 7, 9, 10, 19, 42, 43, 45, 46, 49–57, 78,
RNA polymerase, 248 90–92, 107, 120, 143, 145–147,
Roof sprinklers, 134 161–172, 175–183, 187–189, 203, 220,
Roof top cultivation, 137 249, 250, 253, 254, 267, 269, 275
Rotational grazing, 298 Sheep breeders, 176
Roughage, 165, 225, 229 Sheep farming, 176
Rumen, 18, 30, 91, 101, 116, 145, 163, 188, Shelf life, 8, 17, 19
201, 214, 225, 241, 274, 285, 300 Shelter, 42, 43, 129–139, 163, 171, 187, 290
acidosis, 145, 215–218 Shelter-seeking behaviour, 42
bolus, 91, 96, 99–109 Short-term stressors, 63
epithelium, 70, 215, 217 Shrubs, 116, 180, 283, 303
fluid, 229, 233, 234 Signalling pathway, 62, 71, 275, 276
pH, 91, 147, 169, 228 Signal transduction, 54, 267
temperature, 91, 101, 104–107 Silage, 67, 220, 298
Rumen-reticular boluses, 91, 95, 96 Single nucleotide polymorphism (SNP), 191,
Ruminants, 7, 17, 19, 20, 31, 48, 52, 66, 91, 265–267, 274
107, 116, 117, 131, 133, 144–147, Skin, 18, 33, 41, 45, 66, 69, 77, 91, 92, 96,
151–158, 162–165, 167–172, 189, 191, 113–125, 135, 137, 143, 169, 178, 220,
200–203, 205, 209, 215, 217–219, 221, 256, 270
225, 226, 229, 235, 241, 244, 245, Skin temperature, 94, 95, 134, 166, 188, 288
253–255, 281–284, 300 Slaughter, 7, 18–21, 36, 205
Rumination, 9, 95, 187, 242 Slick Hair Gene, 33, 265
Ruminococcaceae, 253, 254 Slurry storage, 137
Ruminococcus, 215, 226, 228, 229, 253, 254 Small ruminants, 7, 48, 52, 116, 133, 162, 163,
Rye grass, 155 167, 170, 203, 209, 281, 282
Smart technologies, 101, 107, 108, 139
Sodium, 11, 145, 180
S Soil aeration, 209
Saccharolytic, 216 Soil carbon sequestration, 298
Sahiwal, 76–78 Soil compaction, 209
Salem black, 10, 117, 121, 284 Soil quality, 162, 204
320 Index

Solar radiation (SR), 9, 12, 41–45, 47, 94, 119, T

134, 163, 164, 176, 179–180, 215, 295 Tannin, 210, 219–220, 229, 230, 249
Spatial association, 270 Tanzania, 6
Species, 6, 19, 32, 45, 51, 64, 75, 90, 115, 131, Targeted nutrition, 180–182
144, 163, 187, 203, 217, 226, 241, 265, Taxa, 215, 244, 253
281, 295 Taxonomy, 243, 244, 247, 249, 250, 254, 255
Spermatogenesis, 177 Temperate, 8, 30, 33, 34, 46, 75, 106, 132, 153,
Sperm production, 177 154, 190
Starch, 147, 148, 167, 168, 216, 227, 228, 242 Temperature, 6, 18, 41, 51, 62, 76, 87, 100, 119,
Statistical methodologies, 34 131, 143, 161, 176, 187, 204, 215, 224,
Statistical models, 94, 104 243, 265, 284, 295
Steer, 18, 20, 21, 66, 105, 106, 155, 169 Temperature humidity index (THI), 6, 10, 64,
Stimulus, 67, 267 68, 89, 92–95, 104, 132, 133, 188, 288
Stocking density, 135, 139 Temperature sensing ear tags, 91, 95
Stocking rate, 179 Temperature sensitive microchips, 91, 96
Stress, 6, 16, 30, 41, 51, 61, 75, 87, 101, 115, Temperature-sensitive sensors, 64
131, 143, 163, 176, 190, 199, 215, 235, Tetra cutter restriction enzyme, 249
256, 264, 282, 295 Tharparkar, 77, 78
indicators, 78, 90 Thatch roof, 134
response, 9, 19, 51, 53, 54, 57, 62, 63, 69, Thermal challenges, 42, 43, 101, 104, 108
70, 85–97, 106, 108, 115, 119, 123, 124, Thermal energy, 45
162, 270, 273, 287, 288, 295 Thermal equilibrium, 43, 45
tolerance, 70 Thermal gradient, 44
Structural barrier, 228 Thermal imprinting, 32
Stunted growth, 153, 154 Thermal indices, 94, 295
Sub-acute rumen acidosis (SARA), 217, 218 Thermal insulation, 119, 135, 179
Subcutaneous microchips, 91, 95, 96 Thermal load, 41, 44
Sub-fertile, 179 Thermal mapping, 92
Sub-Saharan African climate, 6, 153, 154 Thermal resilience, 117, 120, 284
Substrate, 70, 143, 205, 209, 219, 227, 242, Thermal resistance, 133
244, 255, 256, 267 Thermal stress, 39–48, 57, 64, 69, 70, 119, 136,
Subtropical, 46, 153–155, 158, 190, 224, 283, 137, 141–148, 164, 266
284 Thermal tolerance, 32, 33, 78, 79, 92, 96, 116,
Sugars, 242, 303 121, 169
Sulphate, 205 Thermal zone, 41
Summer, 8, 10, 19–21, 68, 76, 77, 79, 87, 101, Thermo-adaptability, 76
117, 119, 121, 132, 134–136, 139, Thermodynamics, 205
144–147, 155, 156, 164, 167, 178, 188, Thermographic images, 92
189, 191 Thermolysis, 65, 135
Summer matings, 178 Thermometer, 78, 92, 121
Sunburn, 180 Thermoneutral condition, 10, 52, 143, 145
Supplementary feed, 179 Thermoneutral zone (TNZ), 41, 63, 68, 69, 176
Surface area, 42, 43, 46, 47, 136, 228, 242 Thermoreceptor, 66
Surface temperature, vii, 7, 42, 44, 92, 101, 102, Thermoregulation, 9, 17, 18, 42–47, 101, 139,
121, 131, 134, 135 265, 283
Susceptibility, 52, 57, 147, 177, 182 Thermo-sensor devices, 91
Sweating, 18, 41, 44, 51, 64, 87, 116, 120, 143, Thermo tolerance, 7, 17, 22, 70, 73–80, 92, 117,
288 121–124, 185–193, 266, 281, 290, 297,
Sweating rate, 120, 288 299
Swine, 6, 7, 136 Tissue, 10, 19, 43, 51, 52, 54, 56, 57, 65, 67–70,
Symbiotic, 233, 241, 242 77, 102, 117, 118, 122, 123, 143, 166,
Sympathetic-adrenal-medullary (SAM) axis, 167, 169, 241, 242, 266, 270, 272, 300
11, 66 Tissue acidosis, 19
Index 321

Tolerance, 7, 17, 31, 52, 69, 76, 92, 116, 143, Vitamins, 131, 146, 147
169, 178, 186, 218, 229, 256, 265, 281, Volatile fatty acids (VFAs), 10, 19, 217, 218,
297 226, 241, 242
Traits, 6, 17, 31, 62, 76, 116, 163, 178, 186, Volatile solids (VS), 204, 206, 207
256, 265, 284, 299
Transcription, 54, 57, 69, 70, 122, 253, 266
Transcriptome, 266, 273, 274 W
Transcriptomics, 122–124, 191, 245, 253, 274, Water, 5, 17, 32, 41, 51, 61, 95, 102, 116, 132,
297, 305 142, 162, 180, 187, 209, 233, 242, 265,
Transition period, 144, 218 283, 295
Tropical, 6, 8, 30, 32, 36, 44, 45, 62, 64, 71, 75, intake, 9, 64, 95, 104, 165, 167, 187, 188,
79, 116, 119, 132–135, 153–155, 158, 287, 288
165, 178, 190, 219, 224, 281, 283–284, scarcity, 5, 283, 304
289, 290, 297 trough, 148
Tropical areas, 36 vapour, 300
Tropical countries, 75, 281, 283–284 Water holding capacity (WHC), 8, 17, 189
Tunnel ventilation technology, 136–137 Weaning, 32
Tympanic temperature, 104 Welfare, 3–13, 17, 42, 46, 51, 62, 87, 88, 91,
94, 96, 104, 120, 131, 133, 139, 148,
163, 176, 178, 181, 182, 186, 187, 191,
U 193, 256, 287, 295, 299, 301
Ultimate meat pH, 8, 17, 20, 21, 189 Whole-genome metagenomics, 245, 254
Undernutrition, 180 Wind speed, 102, 136, 164
United States, 6, 62, 89, 154, 199 Wind velocity, 6, 12, 295
Wool production, 178, 179
World bank, 148, 152, 153, 157
V World health organization (WHO), 152–154,
Vaccine, 295, 300 157
Vasodilatation, 65, 66
Vector, 12, 265, 300, 304
Vector-borne diseases, 12, 285, 304 Y
Vegetable matter, 180 Yak, 203
Ventilation, 20, 65, 132–136, 138, 139, 180 Yeast, 146, 147, 169, 245
Ventilation rate, 133 Yield, 6–7, 13, 20, 34, 52, 75, 76, 88, 89, 94,
Vertebrate, 225, 267 116, 138, 142, 144–148, 153–157, 164,
Veterinary, 17, 101 167–169, 189, 190, 215, 226, 227, 251,
Viability, 75, 178, 180, 208 254, 255, 265
Virome, 228–229
Virus, 53, 228, 248
Vitamin C, 180 Z
Vitamin E, 146, 147, 165, 168, 169 Zebu, 22, 65, 190, 232