Feed Additives and Supplements for Ruminants
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Editors
M. S. Mahesh Vinod Kumar Yata
Faculty of Veterinary and Animal Sciences Animal Biotechnology Centre
Banaras Hindu University National Dairy Research Institute
Mirzapur, Uttar Pradesh, India Karnal, Haryana, India
ISBN 978-981-97-0793-5 ISBN 978-981-97-0794-2 (eBook)
https://doi.org/10.1007/978-981-97-0794-2
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Foreword
Livestock is an integral part of the agrarian economy in most parts of the world,
providing quality animal protein and thus contributing to human food and nutrition
security. Owing to the greater demand for milk and meat products driven by the
change in dietary preferences and increased per capita income of consumers, rumi-
nant production has been poised to increase by >50% by the year 2050 compared to
the current level. The livestock industry, therefore, aims to produce high-quality,
safe animal products while duly considering factors such as consumer awareness,
public health, ethical standards, and environment.
‘Balanced feeding’ is a key to harnessing the production potential of ruminants,
and feed represents a standalone cost of about 65–70% of recurring farm expenses.
While energy, protein, fat and micronutrients constitute the core components of any
typical balanced diet, the feed additives and supplements occupy niche among the
list of scientific interventions that has guided the enhancement of livestock produc-
tion for augmenting the growing demands for animal products by the global popula-
tion. However, application of some of those feed additives such as antibiotics raised
grave concerns to both animal industry and public health. This has emphasised the
need for developing alternate strategy to enhance the nutrient utilisation towards
achieving higher productivity. Evidently, feed additives seem to be the front runner
to cater to the new and emerging challenges of livestock sector. In addition to regu-
lating rumen fermentation, feed additives help to enhance nutrient utilisation,
improve metabolic functions and nutrient-use efficiency as well as the overall health
status of animals. With the present emphasis on precision feeding to avoid wastage
of feed, it becomes all the more relevant to provide critical nutrients and supple-
ments as per the animal needs to have more efficient and sustainable animal produc-
tion. The use of feed additives is much greater in poultry and pig production than in
ruminants. However, because of its beneficial effects in improving gut health and
providing proper rumen environment for augmenting better performance that trans-
lates into higher yields of saleable farm produce (milk and meat), its use in rumi-
nants is gaining significance. In addition, usage of feed additives may become
increasingly relevant in a changing climatic scenario, where livestock production is
seen alongside ‘efficiency’ as well as ‘environmental sustainability’. Besides, in
practical terms, additives and supplements assist nutritionists in formulating cost-
optimised diets at the farm level.
v
vi Foreword
This book entitled Feed Additives and Supplements for Ruminants brings about
in detail the applications of various additives and supplements for enhancing the
productivity and profitability of ruminants. The editors have covered all the latest
information on various aspects of feed additives and their relevance in present-day
feeding systems. This book is very comprehensive and well-articulated. The updated
information provided in this book would serve as a very useful resource material for
all those interested in studying ruminant production science including researchers,
nutrition advisors, veterinarians, progressive farmers, agripreneurs and feed indus-
try personnel as well as students of animal, dairy, veterinary and agricultural sci-
ences to enrich their knowledge in the subject and provide insights in developing
economical feeding practices for farm animals.
I appreciate the editors and contributors for bringing out this valuable book per-
taining to ruminant nutrition, which is very timely and need of the hour.
Maharashtra Animal and Fishery Sciences University C.S. Prasad
Nagpur, India
ICAR–National Institute of Animal Nutrition and Physiology
Bengaluru, India
Indian Council of Agricultural Research, New Delhi, India
Animal Nutrition Society of India, Karnal, India
Preface
‘Mother cow expects from us nothing but grass and grain’
—M.K. Gandhi, Father of Nation, India
Although the above statement broadly holds good for most of the ruminants in gen-
eral, the diets of modern cows with increased dairy merit demand even more than
mere grass (forage/roughage) and grain (concentrates). That is how science-based
feeding systems have evolved to meeting nutritional requirements in commensurate
with the production performance of today’s ruminant livestock.
Globally, the demand for livestock-source foods are in rise manifolds in the near
future, and hence, the livestock systems—including ruminants—are transforming
from traditional low-input model to advanced commercial farming. Given the
genetic progress-led increased productivity of ruminants, maintaining equilibrium
between the key performance indicators like production, reproduction, health and
welfare has become a new challenge to ruminant producers worldwide. And, it is
well-known that feeds and feeding regimen, among other factors, directly reflect the
balance sheet of farms on a day-to-day basis by acting as 0.7 of standalone farm
expense and thus influence sustainability on a long-run. In this context, apart from
balancing the rations at farm levels for major and micronutrients, there is an increas-
ing necessity/scope to incorporate various feed additives and supplements, which
potentially help navigate with the above challenges associated with intensive
farming.
The Association of American Feed Control Officials defines “additive” as ‘an
ingredient or combination of ingredients added to the basic feed mix or parts thereof
to fulfil a specific need, and usually used in micro quantities and requires careful
handling and mixing’. Unlike feed additives that are generally ‘non-nutritive’, the
supplements are ‘nutritive’ in nature. Although, typically, feed additives and supple-
ments constitute not greater than 10% of the total diet expense, their significance
cannot be overlooked, owing to their myriads of health and production benefits,
ultimately enabling sustainable farming. In other words, while ‘supplements’ may
correct the underlying nutritional deficiencies of diet, ‘additives’ modify nutrient
metabolism such that it benefits the measurable farm outcomes.
This book is a unique collection of feed additives and supplements that are sys-
tematically dealt in 23 different chapters addressing both well-known and new-gen-
eration intriguing compounds. Broadly, this book envisages production enhancement
vii
viii Preface
through various additives and supplements by means of rumen fermentation manip-
ulation, immunomodulation, nutrient utilisation, cellular metabolism, etc. that ulti-
mately dictate milk production, body weight gain, feed efficiency and reproduction.
Besides, this book takes a brief discourse on improving environmental stewardship
of ruminant production by minimising carbon footprint associated with greenhouse
gas emissions, enhancing ruminant-derived food safety through mycotoxin binders,
probiotics, exogenous enzymes, ionophores, rumen buffers, flavours and natural
phytogenic (herbal) feed additives with an emphasis on plant secondary metabolites
(tannins, saponins and essential oils). In addition, recent advances in nutritional
supplements such as trace minerals, amino acids, B vitamins, slow-release nitrogen
and functional nutrients have also been dealt in detail. Moreover, insights on feed
additives for calves, transition cows as well as heat stress amelioration are covered.
Furthermore, some of the lesser-known or emerging additives comprising of newer
trace elements, biochar, rare-earth elements, seaweeds and silage additives consti-
tute an additional highlight of this book. The conclusions from meta-analyses and
the overview on commercial products of select additives along with cost implica-
tions of their usage further help users to get a practical outlook on the benefits of
each supplement, which may also lead to new product development by the feed
industry. Lastly, it may be noted that feed additives and supplements per se are not
magic bullets to boost productivity; rather, when these are used judiciously in con-
junction with proper ration balancing and a good feeding management, these could
catalyse performance, health and profitability of ruminants.
This book holds a wide scope, and hence, it is strongly believed that it would
enable worldwide readers—including researchers, animal producers, nutrition spe-
cialists, veterinarians, feed industry personnel—to appreciate the significance of
proven additives and supplements in bolstering the production of global ruminant
agriculture.
The authors invited to contribute the chapters are active researchers and well-
qualified scientists from both academia and feed industry, who are specialists in
their own specific subject domain as reflected by their quality publications in the
recent past. The editors are grateful to all the authors for their quality contributions,
and both the editors sincerely acknowledge the support of their respective institu-
tions during editing of this book. Finally, we sincerely thank the publishers (Springer
Nature) for their diligent efforts in bringing out the book in this shape.
Mirzapur, Uttar Pradesh, India M.S. Mahesh
Karnal, Haryana, India Vinod Kumar Yata
About the Book
This book is an extensive compilation of self-contained chapters on various feed
additives and supplements employed in ruminant production. Broadly, this book
envisages production enhancement through various additives and supplements by
means of rumen fermentation manipulation, immunomodulation, nutrient utilisa-
tion, cellular metabolism, etc. that ultimately dictate performance outcomes like
milk production, body weght gain, feed efficiency and reproduction. Besides, this
book takes a brief discourse on improving environmental stewardship of ruminant
production by minimising carbon footprint associated with greenhouse gas emis-
sions, enhancing ruminant-derived food safety through mycotoxin binders, micro-
bial feed additives, exogenous enzymes, ionophores, rumen buffers, flavours and
natural phytogenic (herbal) feed additives with an emphasis on plant secondary
metabolites (tannins, saponins and essential oils). In addition, recent advances in
nutritional supplements such as trace minerals, amino acids, B vitamins, slow-
release nitrogen and functional nutrients have also been dealt in detail. Moreover,
insights on feed additives for calves, transition cows as well as heat stress ameliora-
tion are covered. Furthermore, some of the lesser-known or emerging additives
comprising of newer trace elements, biochar, rare-earth elements, seaweeds and
silage additives constitute an additional highlight of this book. The conclusions
from meta-analyses and the details on commercial products and cost implications of
using additives further help users to get a practical outlook on the rationale to use
each additive and supplement. Overall, this book holds a wide scope and would
enable worldwide readers—including researchers, nutrition specialists, veterinari-
ans, feed industry personnel and animal producers—to appreciate the significance
of proven additives and supplements in bolstering the production of global ruminant
agriculture.
Features
• A comprehensive collection encompassing conventional as well as new-genera-
tion feed additives used in ruminant production
• Presents both nutritive supplements (minerals, amino acids, B vitamins, etc.) and
non-nutritive additives (phytogenics, direct-fed microbials, enzymes, flavours,
toxin binders, etc.)
• Holistic coverage of each additive/supplement along with their mode of action
and scientifc rationale to use
ix
x About the Book
• Explores specialty topics like role of feed additives in mitigating enteric methane
emission, heat stress amelioration, controlling gastrointestinal parasitism and
natural phytogenic additives
• Examines the role of additives and supplements in critical life stages of rumi-
nants such as successful calf rearing and transition management
• Uses simple and lucid language, acting as a ready reference material for animal
science and veterinary professionals
• Showcases practical benefits to dairy producers to enhance bottom-line profit-
ability in ruminant farming
Contents
1 Dietary Applications of Exogenous Enzymes
to Improve Nutrient Utilization and Performance
in Ruminants���������������������������������������������������������������������������������������������� 1
Ahmed E. Kholif and Amlan K. Patra
2 cope of Microbial Feed Additives in Ruminant Nutrition�������������������� 29
S
Vinay Venkatesh Varada and Sachin Kumar
3 Nano-Mineral Technology in Ruminant Feeding: Progress
and Prospects���������������������������������������������������������������������������������������������� 51
Partha Sarathi Swain, D. Rajendran, S. B. N. Rao,
and Sonali Prusty
4 Organic Trace Minerals in Ruminant Nutrition:
Production, Reproduction, Health, Economics,
and Environmental Implications�������������������������������������������������������������� 69
N. K. S. Gowda, D. T. Pal, and Debpriyo Kumar Dey
5 ewer Trace Elements for Ruminants ���������������������������������������������������� 87
N
Ravi Prakash Pal, Veena Mani, Hujaz Tariq,
Srobana Sarkar, Amit Sharma, and Deepanshu Gupta
6 Advancements in Methane-Mitigating Feed Additives
in Ruminants���������������������������������������������������������������������������������������������� 119
Y. A. Soltan and A. K. Patra
7 umen-Protected Amino Acids for Ruminants�������������������������������������� 143
R
Jasmine Kaur, Ravneet Kaur, M. S. Mahesh,
and S. S. Thakur
8 hytochemicals as Natural Feed Additives for Ruminants�������������������� 167
P
Abhishek K. Singh, Lamella Ojha, Punita Kumari,
Mahipal Choubey, and Sandeep K. Chaudhary
xi
xii Contents
9 Herbal Feed Additives and Supplements
for a Sustainable Ruminant Production�������������������������������������������������� 197
Thulasiraman Parkunan, Mukesh Kumar Bharti,
Thirumalaisamy Govindasamy, Manish Kumar,
Dhinesh Kumar Ramasamy, and M. S. Mahesh
10 Exploring Condensed Tannin to Control Gastro-Intestinal
Parasitism in Small Ruminants���������������������������������������������������������������� 235
Sandeep Uniyal, Amit Kumar Chaurasiya, Pramod Chaudhary,
and Udeybir Singh Chahal
11 Mycotoxin Binders for a Safe Ruminant-Derived
Food Production���������������������������������������������������������������������������������������� 249
Prateek Shukla, R. Rajendra Moorthy, and M. S. Mahesh
12 ole of Flavoured Additives in Ruminant Production �������������������������� 265
R
Vishal Gaikwad, Ankita Rautela, Gopal Kedare,
Srobana Sarkar, Ravi Prakash Pal, and Amit Sharma
13 otential of Slow-Release Nitrogen in Ruminant Feeding �������������������� 281
P
K. M. Rashmi, T. M. Prabhu, and M. S. Mahesh
14 Feed Additives for Calves: A Brief Insight
on Their Classification and Applications ������������������������������������������������ 301
P. Ravikanth Reddy, D. Yasaswini, P. Pandu Ranga Reddy,
I. Hyder, and D. Srinivasa Kumar
15 dditives and Nutritional Supplements for Transition Cows���������������� 337
A
Dilshad Kour, Divya Sharma, Vijay Kumar Sharma,
and M. S. Mahesh
16 Heat Stress in Ruminants and Its Mitigation Through
Nutritional Additives��������������������������������������������������������������������������������� 367
V. Sejian, C. Devaraj, C. G. Shashank, A. P. Madhusoodan,
M. V. Silpa, A. Devapriya, M. R. Reshma Nair, R. U. Suganthi,
N. Ramachandran, G. B. Manjunathareddy, and R. Bhatta
17 ecent Developments in B-Vitamin Nutrition of Dairy Cattle�������������� 399
R
A. K. Singh, Shilpi Kerketta, Punita Kumari, M. S. Mahesh,
S. K. Rajak, and Ravindra Kumar
18 iochar as a Novel Feed Additive for Ruminants ���������������������������������� 423
B
Sudhir Kumar Rajpoot, Ph. Romen Sharma, Jayesh Singh,
Adarsh Kumar, S. Vijayakumar, Richa Chaudhary,
and Dinesh Kumar
19 rospects of Rare-Earth Elements in Ruminant Production���������������� 437
P
Hujaz Tariq, Amit Sharma, Neha Sharma, Srobana Sarkar,
and Ravi Prakash Pal
Contents xiii
20 Silage Additives������������������������������������������������������������������������������������������ 449
Deepesh Bharat Mishra and Nitin Tyagi
21 cope of Seaweeds to Boost Ruminant Productivity������������������������������ 459
S
Muneendra Kumar, Vinod Kumar, and Raju Kushwaha
22 Ionophoric Additives to Augment Performance
Outcomes and Economics in Ruminants ������������������������������������������������ 479
Sonali Prusty, M. S. Mahesh, Rakesh Sheel, Partha Sarathi Swain,
A. P. Madhusoodan, Neeraj Thakur, and G. E. Chethan
23 Rumen Buffers to Harness Nutrition, Health
and Productivity of Ruminants���������������������������������������������������������������� 495
B. S. Bharath Kumar, Hujaz Tariq, Ranjan K. Mohanta,
Muhammad Umar Yaqoob, Vinu M. Nampoothiri, M. S. Mahesh,
Dinesh Kumar, Brishketu Kumar, and Chander Datt
About the Editors
M.S. Mahesh is an Assistant Professor (Animal
Nutrition) of Livestock Farm Complex at the Faculty
of Veterinary and Animal Sciences of Banaras Hindu
University, Mirzapur, India. He has graduated in
Veterinary Science and Animal Husbandry from
Veterinary College, Bengaluru, and earned master’s and
Ph.D. in Animal Nutrition from ICAR-National Dairy
Research Institute, Karnal, India. He was awarded with
university gold medals for academic excellence during
undergraduation and afterwards with Junior and Senior
Research Fellowships. His areas of specialisation
include use of novel feed ingredients in ruminants, feed
characterisation, on-farm diet formulation and
optimisation, designing species-specific micronutrient
premixes, troubleshooting nutritional issues and
rationalising feed additive applications for an augmented
performance of dairy cattle and buffaloes. His previous
association with animal feed industry helped him gain a
broad-spectrum of hands-on experience on the diverse
facets of practical dairy nutrition. He has published over
30 research/review papers in journals of national and
international repute with a few other technical articles.
He served as the former member of editorial board of
Indian Journal of Animal Nutrition and is currently
associated as a reviewer for many international journals
on animal science and multidisciplinary areas. He is a
life member of professional societies such as Animal
Nutrition Society of India, Animal Nutrition
Association, Indian Poultry Science Association and
Karnataka Veterinary Council. In his present role, he
teaches students and researchers on various aspects of
animal nutrition.
xv
xvi About the Editors
Vinod Kumar Yata is an interdisciplinary researcher
working at Center of Excellence for Aging and Brain
Repair, University of South Florida (USF), Tampa,
Florida, USA. Previously, he worked as a Research
Associate at National Dairy Research Institute, Karnal,
India. He served as an Assistant Professor at the
Department of Biotechnology, Dr. B.R. Ambedkar
National Institute of Technology, Jalandhar, Punjab,
India. He received his Ph.D. in Biotechnology from
Indian Institute of Technology, Guwahati. He specialises
in interdisciplinary research which includes
nanotechnology, microfluidics, animal biotechnology,
cancer biology and bioinformatics. He has developed a
microfluidic device for the separation of live and motile
spermatozoa from cattle semen samples. His research
interests have been focused on the development of
nanocarriers, understanding prodrug enzyme therapy
and targeted drug delivery. He elucidated the structural
features and binding interactions of several biomolecules
by in silico methods. He has published four books as an
editor and one book as an author with Springer Nature
publisher. He has published several research papers in
peer-reviewed international journals and presented
papers in several international conferences.
Dietary Applications of Exogenous
Enzymes to Improve Nutrient Utilization 1
and Performance in Ruminants
Ahmed E. Kholif and Amlan K. Patra
Abstract
Competition among animal feeds, human foods, and demands for biofuel pro-
duction forces livestock producers to feed their animals on the available low-cost
raw materials. Ruminant farmers provide their animals forage-based diets to
reduce the cost of feeding and to keep normal rumen function. However, only for-
age feeding does not sometimes provide animals enough available energy and
protein required for milk production and growth. The major problem with feed-
ing low-quality forages or raw materials is their content of rigid and less-
digestible cell wall components. However, ruminants have a digestive system
with a complex and dynamic microbiota and microbial activities that allow them
to utilize plant cell walls and degrade them to be utilized as energy sources by
host. The ability of ruminants to degrade structural polysaccharides (e.g., cellu-
lose, hemicellulose, and lignin) has a limit, and under perfect conditions, is less
than 65%. Different approaches have been developed to improve nutrient digest-
ibility and production responses in ruminants including chemical, physical, and
biological strategies. Exogenous enzymes have gained an increasing interest to
improve the nutritive value of ruminant feeds. The aim of administration of exog-
enous enzymes is to improve fiber degradation and efficiency of feed utilization
A. E. Kholif (*)
Dairy Science Department, National Research Centre, Giza, Egypt
Department of Animal Sciences, North Carolina Agricultural and Technical State University,
Greensboro, NC, USA
e-mail: [email protected]; [email protected]
A. K. Patra
Department of Animal Nutrition, West Bengal University of Animal and Fishery Sciences,
Kolkata, India
American Institute for Goat Research, Langston University, Langston, USA
© The Author(s), under exclusive license to Springer Nature Singapore Pte 1
Ltd. 2024
M. S. Mahesh, V. K. Yata (eds.), Feed Additives and Supplements for Ruminants,
https://doi.org/10.1007/978-981-97-0794-2_1
2 A. E. Kholif and A. K. Patra
by enhanced nutrient utilization and reduce the wastage. Exogenous enzymes are
capable of breaking down specific bonds in carbohydrates, starch, protein, fats,
cellulose, hemicellulose, pectin, glycoproteins, and lignin. Lactobacillus aci-
dophilus, L. plantarum, Streptococcus faecium, and Bacillus subtilis are the main
bacterial species that produce a wide array of exogenous enzymes, while
Trichoderma reesei, Aspergillus oryzae, and Saccharomyces cerevisiae are the
main fungal enzyme producers. The ability of exogenous enzymes to improve
growth performance in feedlot cattle and small ruminants as well as milk produc-
tion in dairy cattle has been demonstrated, but with inconsistent effects. Enzyme
type and preparation, application method, level of enzyme supplementation,
diets, animals and interaction between enzymes and feeds are the factors that
determine the response to enzyme supplementation and are responsible for varia-
tions among experiments. This chapter reviews information about sources of
enzymes, enzyme activity, modes of action, optimal inclusion, and response of
ruminant to dietary applications of various enzymes.
Keywords
Animal performance · Exogenous enzymes · Feed additives · Forage fiber ·
Ruminants · Structural carbohydrates
1.1 Introduction
Many countries have limited resources for good-quality fodder crops. Forages are
the cheapest available ruminant feeds; however, feeding animals on poor quality
forages limits their performance. Poor-quality fodders do not provide enough
metabolizable energy and protein intakes for optimum milk production and daily
gain in ruminants due to their high contents of structural fibrous carbohydrate com-
ponents, e.g., cellulose, hemicellulose, and lignin (Taghizadeh and Nobari 2012).
Furthermore, slow and/or incomplete digestion of fiber in diets limits their total
tract digestion, especially when forages or fibrous feeds are the major component in
the diets.
Animal nutritionists have explored new techniques to increase the digestibility of
low-quality fibrous feeds (Elghandour et al. 2014; Azzaz et al. 2020; Simon et al.
2024). Different approaches including physical, chemical, and biological treatments
of forages were employed to achieve these goals. The supplementation with exog-
enous enzymes showed mixed results with milk and feedlot performance; however,
improved feed utilization and performance were observed in most studies. The rea-
sons for the administration of exogenous enzymes in ruminant diets include: (1) to
increase the availability of nutrients enclosed within fiber-rich cell walls; (2)
improving the capability of ruminal endogenous enzymes to breakdown specific
chemical bonds in raw materials; and (3) to break down antinutritional factors in
feeds. The improvements in performance and nutrient utilization with feeding exog-
enous enzymes observed in many studies with cattle (Tewoldebrhan et al. 2017),
1 Dietary Applications of Exogenous Enzymes to Improve Nutrient Utilization… 3
goats (Azzaz et al. 2020), sheep (Salem et al. 2015a), and buffaloes (Morsy et al.
2016) encouraged their commercial applications. Exogenous feed enzymes increase
the digestibility of nutrients, feed efficiency, minimize the environmental impact in
animal industry by allowing better use of natural resources, and reducing wastage
(Kholif et al. 2017b).
The first investigations on enzyme administration in animal feeds were per-
formed in early 1920s; however, the first commercial production of enzymes using
submerged fermentation systems in large fermenters was started in the 1950s. At
that period, responses were variable and modes of action were not clear. The enor-
mous development in fermentation technology allowed for the economic produc-
tion of large quantities of enzymes with high biological activities to be used as
livestock feed additives to manipulate the digestive processes in ruminants. Most
enzyme applications in ruminants have focused on the use of polysaccharide-
digesting enzymes to improve the ruminal degradation of the relatively slowly
digested fiber. Cellulases and xylanases are the two major types of enzymes com-
monly considered to improve fiber digestion. Cellulases are complex mixtures of
proteins that act to hydrolyze β-(1-4)-glucosidic linkages, which connect the indi-
vidual glucose molecules in the cellulose found in forages. Xylanases degrade
xylans in the hemicellulose fraction in forages. The main methods of application of
exogenous enzymes in animal feeding are the pretreatment of feeds before feeding
or the direct feeding enzyme preparations to improve the degradation activities in
the gastrointestinal tract of animals. Thus, microbiologists and nutritionists recom-
mend the addition of active enzyme preparations to ruminant diets prior to feeding.
This chapter provides some recent information demonstrating the important effects
of specific enzyme preparations on microbial activities in the rumen and discusses
their impact on the performance and nutrient utilization in ruminants.
1.2 Cell Wall Components
The maturity of the forages or the amount of lignification can influence the compo-
sition and structure of plant polysaccharides and cell wall components. Each plant
cell contains a tough and rigid layer consisting mainly of cellulose, hemicellulose,
lignin, and pectin, which provides the cell with physical and structural support and
protection. The cell wall primarily contains three major polymers including: cellu-
lose at 35–50%, xylan (hemicellulose) at 20–35%, and lignin at 10–15% (Mendoza
et al. 2014). The cellulose microfibrils are linked via hemicellulosic tethers to form
the cellulose-hemicellulose network, which is embedded in the pectin matrix.
Xyloglucan is the most common hemicellulose in the primary cell wall (Mendoza
et al. 2014). Cellulose is a linear molecule of insoluble β-glucans, comprising of
D-anhydro-glucopyranose residues linked by β(1-4) bonds (Paloheimo et al. 2010).
Cellulose is present in the cell walls of plants, trees, tunicates, algae, and several
species of bacteria as exopolysaccharide membranes (Lynd et al. 2002; Khandelwal
and Windle 2013). The cellulose composition differs between microorganisms and
4 A. E. Kholif and A. K. Patra
plants (Lin et al. 2013). With hemicellulose and lignin, cellulose forms a cellulosic
biomass complex.
Hemicellulose has a substantial role in maintaining cell wall structure in plants.
It accounts for about a quarter of total plant biomass (Scheller and Ulvskov 2010).
Hemicellulose is a heterogeneous group of polysaccharides characterized by β-(1-4)
linkages, which includes xyloglucans, glucuronoxylans, glucuronoarabinoxylans,
glucomannans, and galactoglucomannan (Scheller and Ulvskov 2010). The main
component in hemicellulose is xylan.
Lignin is a macromolecule composed of highly cross-linked phenolic molecules
(Cosgrove 2001). There are different forms of lignin including guaiacyl,
5-hydroxygualacyl, p-hydroxyphenyl, and syringyl lignin (Moore and Jung 2001).
The concentration of lignin varies depending on the nature of the plant species,
maturity, soil and environmental conditions, organs, and cell wall layers (Mendoza
et al. 2014). Lignin has a high molecular weight that limits the availability of the
structural carbohydrates to rumen microorganisms (Van Soest 2019), which, in turn,
limits the digestibility and overall forage nutrient availability to ruminants.
Additionally, cell wall contains structural proteins at about 1–5%, which are
classified as hydroxyproline-rich glycoproteins and arabinogalactan proteins,
glycine-rich proteins, and proline-rich proteins (Moire et al. 1999). The concentra-
tions of all components in the cell wall differ between plants, cell type, and cell age
(Taghizadeh and Nobari 2012).
1.2.1 Ruminal Degradation of Cell Wall Components
Ruminants have their digestive system for efficient utilization of plant cell walls.
Under optimal conditions, cell wall digestibility in the total gastrointestinal tract is
less than 65% (Van Soest 2019). Ruminants have a compartment known as ‘rumen’,
which is a very efficient fermentation system that houses a vast array of different
microbes. The rumen microbial ecosystem shows high resilience and inertia proper-
ties due to core microbiota composition and also has great plasticity owing to other
parts of the microbiota (Patra 2020). The consortial activities of the ruminal micro-
biota render ruminants to utilize complex forages including browse forages/foli-
ages, grasses, and other plant materials due to the ability to degrade and ferment
carbohydrates in plant cell walls and provide volatile fatty acids and protein to the
host animals. Taghizadeh and Nobari (2012) determined four factors controlling
fiber degradation in the rumen: (1) the structure and composition of ingested plants;
(2) densities of the predominant fiber digesters in the rumen; (3) microbial factors
regulating microbial attachment to feed particles and hydrolysis; and (4) animal
factors that increase the availability of nutrients through mastication, salivation and
digesta kinetics.
Ruminants digest cellulosic feeds mainly in the rumen by the microbiota. The
environment in the rumen is anaerobic and contains dense populations of several
species of bacteria (10–50 × 109/mL), protozoa (105–106/mL), archaea (107–108/
mL), bacteriophages, fungi (104–105/mL), and sometimes yeasts (Russell and
