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 Fish Vaccines
This book is a timely reference text that highlights the role of vaccination in the fast-­growing
­aquaculture industry. It discusses topics such as vaccine formulation, vaccine delivery, and
­enhancing the immune response of fish using nanoparticles. Information related to vaccine safety,
ethical approval, and regulations is also discussed, together with dissemination of vaccines to
fish farms across the globe. This cutting-edge book presents novel strategies to meet the growing
demand for vaccines in finfish aquaculture. This book is useful to students, academics, clinicians,
and professionals in the field of fisheries sciences, aquaculture, and veterinary sciences.
 Fish Vaccines
Health Management for Sustainable Aquaculture

Edited by
Preetham Elumalai
Kim Thompson
Sreeja Lakshmi
First edition published 2023
by CRC Press
2385 Executive Center Drive, Suite 320, Boca Raton, Florida 33431-8530

and by CRC Press

4 Park Square, Milton Park, Abingdon, Oxon, OX14 4RN

CRC Press is an imprint of Taylor & Francis Group, LLC

© 2023 selection and editorial matter, Preetham Elumalai, Kim Thompson, Sreeja Lakshmi; individual chapters, the
contributors

Reasonable efforts have been made to publish reliable data and information, but the author and publisher cannot
assume responsibility for the validity of all materials or the consequences of their use. The authors and p ­ ublishers
have attempted to trace the copyright holders of all material reproduced in this publication and apologize to
­copyright ­holders if permission to publish in this form has not been obtained. If any copyright material has not been
­acknowledged please write and let us know so we may rectify in any future reprint.

Except as permitted under U.S. Copyright Law, no part of this book may be reprinted, reproduced, transmitted, or
utilized in any form by any electronic, mechanical, or other means, now known or hereafter invented, ­including
­photocopying, microfilming, and recording, or in any information storage or retrieval system, without written
­permission from the publishers.

For permission to photocopy or use material electronically from this work, access www.copyright.com or contact the
Copyright Clearance Center, Inc. (CCC), 222 Rosewood Drive, Danvers, MA 01923, 978-750-8400. For works that are
not available on CCC please contact [email protected]

Trademark notice: Product or corporate names may be trademarks or registered trademarks and are used only for
identification and explanation without intent to infringe.

ISBN: 978-1-032-48341-2 (hbk)

ISBN: 978-1-032-48344-3 (pbk)
ISBN: 978-1-003-38854-8 (ebk)

DOI: 10.1201/9781003388548

Typeset in Times
by codeMantra
Contents
Acknowledgements...........................................................................................................................vii
Foreword by Shri. Pinarayi Vijayan...................................................................................................ix
Foreword by Prof Chris Secombes....................................................................................................xi
Foreword by Dr. Tim Connelley..................................................................................................... xiii
Editors............................................................................................................................................... xv
Contributors....................................................................................................................................xvii

Section I Vaccines in Aquaculture: A Preface

Chapter 1 Understanding Vaccine Development in Aquaculture..................................................3

Nayomi John, Anu Joseph, Aparna Sasi, Bismi Mujeeb, Joyal Elisabeth Baiju,
Eleeswa Celin Syrus, and Nivya Mariam Paul

Chapter 2 Antigens and Immune Responses in Fishes: An Overview........................................ 17

Nivya Mariam Paul, Aruna Babu, Amrutha Rajesh, Neema, Alanta Maria Binu,
Anu Jose, Aneena Mariya Abraham, and Nayomi John

Chapter 3 Aquaculture: Facing an Array of Pathogens............................................................... 33

Wanna Sirimanapong, Prawporn Thaijongrak, Chiranan Sudpraseart, and
Boonsirm Withyachumnarnkul

Section II Vaccination in Fishes: Types and Methods

Chapter 4 Concepts and Types of Vaccines................................................................................. 49

Subramaniam Sivakumar, Panneerselvam Punniyakotti, Kuppuswamy Kavitha,
and Panneerselvam Saravanan

Chapter 5 Antigen Discovery....................................................................................................... 65

Nhat HM Truong, Nam Tri Vo, Phuong Hong Vo, and Hoang Duc Nguyen

Chapter 6 Novel Advancements in Vaccine Delivery and Methods of Vaccination:

Experimental Strategies.............................................................................................. 75
Abdullah Bin Abdul Nazar, Muhammed Salih Salim, Aswathi Ashok,
Anuj Sharma, Sanchu Prakash, Malavika Biju Reni, Meril Mary Mathew,
Muzammil Siddiqui, Subramaniam Sivakumar, Sreeja Lakshmi, and
Preetham Elumalai

v
vi Contents

Section III Adjuvants in Vaccination: An Underpinning

Chapter 7 Role of Adjuvants in Vaccination Studies................................................................. 101

Md Yasin Ina-Salwany, Tilusha Manchanayake, Aslah Mohamad,
Md. Shirajum Monir, Mohammad Noor Amal Azmai, Salleh Annas, and
Mohd Zamri-Saad

Chapter 8 Development of Mucosal Adjutants for Fish Vaccination......................................... 113

Ina Salwany Md Yasin, Aslah Mohamad, Mohammad Noor Amal Azmai,
Annas Salleh, and Mohd Zamri Saad

Section IV T
 echnological Advancements and
Efficacy of Fish Vaccination

Chapter 9 Biotechnological Approaches to Vaccines................................................................ 127

Thavasimuthu Citarasu, Ganapathi Uma, Ramamoorthy Sathish Kumar,
Sugumar Vimal, and Mariavincent Michael Babu

Chapter 10 Safety and Efficacy of Vaccines in Aquaculture....................................................... 155

Eakapol Wangkahart, Po-Tsang Lee, Chou-Min Chong, and
Fernando Yamamoto

Chapter 11 Mass Vaccination in Aquaculture: Possibilities and Limitations.............................. 169

Hethesh Chellapandian, Sivakamavalli Jeyachandran, Joseph Selvin,
Kiyun Park, and Ihn-sil Kwak

Section V V
 accines in Aquaculture:
Challenges and Perspectives

Chapter 12 Fish Vaccination: How Close Are the Fish Farmers?............................................... 189
Saurabh Dubey, Kizito K. Mugimba, Jitendra Kumar, and
Hetron M. Munang’andu

Chapter 13 Development of Fish Vaccines: Challenges and Future Perspectives.......................209

A. P. Desbois and S. J. Monaghan

Chapter 14 Fish Vaccine: From Field to Development................................................................ 229

R. Cascione, Adrian Astier, Philippe Mahl, and Øystein Evensen

Index............................................................................................................................................... 237
Acknowledgements
The editors cordially acknowledge ‘International Veterinary Vaccinology Network’ (IVVN) and
‘BactiVac – The Bacterial Vaccines Network’ for their excellent support for the successful produc-
tion of the book. The authors are sincerely appreciated and acknowledged for their ­skillfull presen-
tations of their respective chapters in articulating this book. I would like to thank Dr. Subramaniam
Sivakumar, Associate Professor, Sri Sankara Arts and Science College, Enathur, Kancheepuram
for offering his handful assistance and for technical support throughout the publishing process. The
editors also greatfully acknowledge CRC Press/Taylor & Francis Group, especially Madhurima
Kahali, Neha Bhatt and Vijay Shanker, for their patience and timely support towards the commend-
able task of organizing and integrating this compendium.

vii
Foreword by Shri. Pinarayi Vijayan
As part of the Cochin University of Science and Technology’s
international project with the UK and Norway, on aquatic vac-
cination, this book has been jointly edited by Dr. Punnadath
Preetham Elumalai, Dr. Kim Thomson, and Dr Sreeja
Lakshmi. Fish Vaccines: Health Management for Sustainable
Aquaculture, published by the CRC Press, USA, incorporates
various chapters authored by experts from countries in which
aquaculture is a prominent economic activity. Therefore, it will
not merely be an academic document, but will also serve as a
useful reference material for Kerala, especially as we are under-
taking several measures to modernize and improve our fisheries
sector.
Kerala’s international collaborations in the fisheries sector
date back to almost seven decades. Back in the early 1950s, we
had collaborated with Norway to develop the capture fisheries
sector in the State. Subsequently, the Neendakara Fishing Harbour was established under the
Indo Norwegian Fisheries project, in 1953. Modern fishing techniques like trawling were learnt
by our traditional fishing communities from the Norwegians. It is to further strengthen such col-
laboration, that an official delegation from Kerala recently visited Finland, Norway, and the UK.
Throughout the discussions and deliberations held by Kerala’s delegation in these countries, the
fisheries and aquaculture sector was featured prominently. Agreements were also made, with regard
to co-operation in the sector. The Norwegian Institute of Marine Research has assured their support
for research in fish nutrition, fish feed, and fish health management.
The NORD University that focusses on blue and green growth, and has a prominent Faculty
of Biosciences and Aquaculture, has expressed its willingness to provide research orientation to
students and teachers in Kerala. We have also decided to ensure more cooperation in student and
faculty exchanges and research aimed at employment generation through offshore aquaculture, cage
farming, and sustainable fish farming.
Fisheries and aquaculture play a significant role in the overall development of the world in terms
of poverty alleviation, employment creation, and protein security. Unfortunately, owing to various
reasons like climate change, over exploitation, marine pollution etc. our marine fish stocks are
getting depleted and the production from the capture sector is showing dwindling trends for a few
decades now. So, aquaculture has emerged as a sunrise sector to compensate this gap between pro-
duction and demand.
The world has advanced significantly in aquaculture with varying production protocols and strate-
gies, making this health food available in all the markets at a nominal price. Kerala’s aquaculture
production is low compared to countries like Norway and Vietnam and a few Indian States too. We
aim to correct this situation through joint action and collaboration. In fact, both Finland and Norway
have expressed interest in collaborating with the Maritime Cluster with a Sustainable Maritime
Technology Hub, which is coming up in Kochi. It is in this larger backdrop of Kerala’s focussed inter-
ventions toward emerging as a prominent maritime hub that this book assumes greater significance.
Apart from providing an introduction to vaccines in aquaculture, this book discusses various
aspects of fish vaccines including their types and methods, technological advancements and chal-
lenges in the field, and so on. Certainly, it can serve as a much useful reference material for those
who are engaged in improving the productivity of fisheries in general, and aquaculture in par-
ticular. Its contents would also go a long way in promoting cutting edge scientific and sustainable

ix
x Foreword by Shri. Pinarayi Vijayan

aquaculture. In that regard, this book benefits not only academia but also the general public who
are keen to learn about and earn from aquaculture.
Kerala has the potential to emerge from a follower of aquaculture techniques developed around the
world, to a creator of cutting edge technologies and techniques that can be followed around the world.
In order to ensure that academic works such as this that balance research with experience are essential.
I congratulate the authors and wish the readers an enriching experience.

Shri. Pinarayi Vijayan

The Hon’ble Chief Minister of Kerala
Foreword by Prof Chris Secombes
Over the last couple of years, the importance of
vaccination as a means of infectious disease con-
trol has been apparent to everyone. During the
Covid pandemic, modern vaccination approaches
were adopted and proven to be highly efficient in
the fight against this disease. Such advances are
likely to enable novel vaccine development in many
areas of clinical and veterinary science, on a scale
not previously seen. The need for vaccines to con-
trol diseases of fish in aquaculture is also readily
apparent and crucial for the expansion of aquacul-
ture in many developing countries, where fish are an
important source of animal protein. Vaccination of
fish is already highly successful in controlling many
of the diseases of concern in the salmonid fish farm-
ing industry. Indeed, in Atlantic salmon culture,
it is common for all fish to be vaccinated before
transfer to seawater. This book, edited by Preetham
Elumalai, Kim Thompson, and Sreeja Lakshmi, contains a number of review articles pertaining to
the demand and ways in which vaccination can be used in aquaculture to improve fish health. There
is an initial overview of the need for vaccination of fish, and the pathogens commonly encountered.
It then describes the approaches taken to develop new and improved vaccines for aquaculture, using
modern approaches and methodologies, including the requirement to ensure the safety as well as
efficacy of those that are commercialized. The use of adjuvants to maximize (fish) vaccine efficacy
is well known. For example, the benefit of incorporating adjuvants into salmonid vaccines against
furunculosis (caused by the bacterium Aeromonas salmonicida) was realized at an early stage of
their development. However, better adjuvants are needed, especially those that aid mucosal vaccina-
tion of fish, and their role and the advances on the horizon are covered in some of the later chapters.
Such progress bodes well for the future. The challenges of vaccine commercialization, in terms of
taking successful vaccines to market and increasing their use by fish farmers, is the final topic cov-
ered in this book. That there is an economic benefit as well as improved fish welfare is key to their
uptake and should not be overlooked during the enthusiastic hunt for novel, highly effective vaccines
to fish pathogens. Overall, this book highlights the vital role that vaccination can play in fish health
management and the advances being made currently, which will help to ensure a robust and sustain-
able future for aquaculture.

Prof Chris Secombes

University of Aberdeen, Aberdeen, Scotland, UK

xi
Foreword by Dr. Tim Connelley
The rapid growth of aquaculture as a primary source of providing
the increasing needs for animal-derived protein in large parts of
the world has been matched by a highly dynamic and progressive
development in the science and application of vaccines. This has
been achieved, in large part, by scientists from around the globe
working collaboratively together. It has been a privilege to be
able to support small-scale collaborations between some of these
scientists as part of the portfolio of activities conducted by the
International Veterinary Vaccinology Network (IVVN).
This book covers the broad spectrum of perspectives of vaccin-
ology—from the foundational sciences such as immunology and
microbiology, through the translational aspects of vaccine produc-
tion and manufacture and onto the delivery of vaccines to farmers. As such it represents a useful
text which considers and summarizes the current successes as well as the future challenges and
opportunities in a comprehensive and holistic manner.
In India, aquaculture continues to increase as a source of both income and food security - ­growing
at an annual rate of 8%, generating >USD6 billion in revenue and supporting the livelihoods of
~28 million fishermen and others involved in the aquaculture economy. With the expansion of the
aquaculture sector comes increasing risks of disease outbreaks from a number of viral, bacterial,
parasitic and fungal pathogens with the potential to lead to substantial morbidity and mortality for
the farmed fish and economic loss to farmers. Vaccination is, and will continue to be, a principal
method to mitigate and minimize the losses incurred from such diseases. As such it gives me great
pleasure to provide a foreword to Fish Vaccines: Health Management for Sustainable Aquaculture
edited by Dr. Preetham Elumalai, Dr. Kim D Thompson and Dr. Sreeja Lakshmi. I am certain the
readers will find this book a useful reference by which a better understanding of the current status
of fish vaccine development and deployment can be gained.

Dr. Timothy Connelley

Director of the International Veterinary Vaccinology Network,
The Roslin Institute, University of Edinburgh, United Kingdom

xiii
Editors
Preetham Elumalai, PhD, is an Associate Professor at the Department of Marine Biology, Cochin
University of Science and Technology, Kochi, Kerala, India. He earned a Master’s degree from the
University of Madras and a PhD in biochemistry and molecular immunology from the Institute for
Immunology, University of Regensburg, Germany. His research practice includes bioassay-guided
identification of novel marine compounds, unveiling fish lectins in innate immune defense, aquatic
vaccine development, evaluation of cost-effective feed additives and nutrigenomics, and effects of
environmental pollutants on marine ecosystems. He has been a partner in numerous EU-, Indian-,
and UK-funded projects (e.g., IVVN, BactiVac). He has written more than 70 peer-reviewed articles
and has 2 patents in his name apart from editing 5 books (Springer, CRC Press) and has presented
his work at more than 60 national and international conferences. He has been awarded the presti-
gious INSA fellowship (2018); MASTS, Fellowship (2019); IVVN award, UK (2020); FRSB, UK
(2021); and BactiVac award, UK (2022).

Kim Thompson, PhD, is a Principal Investigator at Moredun Research Institute, where she heads
the Aquaculture Research Group. She earned a BSc in biochemistry from the University of Stirling,
an MSc in immunology from the University of Aberdeen, and a PhD in fish immunology from the
University of Stirling. She has published over 160 articles in international journals relating to her work
on fish health. Her research interests include vaccine development for fish pathogens (viral, bacterial,
and parasitic), examining the immune response of fish to these pathogens and understanding the factors
that influence this response. She is interested in developing alternative therapies for disease control in
fish (e.g., functional feeds, immunostimulants, and probiotics).

Sreeja Lakshmi, PhD, is a Postdoctoral Research Scientist in collaboration with Moredun Research
Institute (MRI), UK. She graduated from Calicut University and earned a PhD in biochemistry and
functional genomics from the Institute for Molecular Biology, University of Regensburg, Germany.
She has published research articles in peer-reviewed international journals and authored books
and book chapters. She has been awarded prestigious research grants from the Bavarian Research
Foundation (Bayerische Forschungsstiftung), Government of Bavaria, Germany; an HRD-Fellowship
for Women Scientists from the Department of Health Research, Government of India; and a MASTS
(Marine Alliance Science and Technology, Scotland) Award for Postdoctoral and Early Career
Research Exchanges (PECRE), visited the University of Aberdeen, Scotland, and received IVVN
Fellowship grant from the International Veterinary Vaccinology Network (IVVN), UK.

xv
Contributors
Aneena Mariya Abraham R. Cascione
Department of Microbiology Virbac Asia Pacific Co. Ltd.
Mar Athanasius College (Autonomous) Bangkok, Thailand
Kerala, India
Hethesh Chellapandian
Department of Biotechnology and
Mohammad Noor Amal Azmai
Microbiology
Department of Biology
National College (Autonomous)
Faculty of Science, Universiti Putra Malaysia
Tiruchirappalli, Tamil Nadu, India
Serdang, Selangor, Malaysia
Chou-Min Chong
Salleh Annas Aquatic Animal Health and Therapeutics
Department of Veterinary Pathology and Laboratory (AquaHealth)
Microbiology Institute of Bioscience
Faculty of Veterinary Medicine Universiti Putra Malaysia
Universiti Putra Malaysia Serdang, Selangor, Malaysia
Serdang, Selangor, Malaysia
Saurabh Dubey
Department of Production Medicine
Aswathi Ashok
Norwegian University of Life Sciences
Department of Marine Biology, Microbiology
Ås, Norway
and Biochemistry
School of Marine Sciences A. P. Desbois
Cochin University of Science and Technology Institute of Aquaculture, University of Stirling
(CUSAT) Stirling, Scotland, United Kingdom
Kochi, India
Preetham Elumalai
Adrian Astier Faculty of Fisheries
Aquaculture division Kerala University of Fisheries and Ocean
Virbac Asia Pacific Co. Ltd. Studies
Bangkok, Thailand and
Department of Marine Biology, Microbiology
and Biochemistry
Aruna Babu
School of Marine Sciences
Department of Microbiology
Cochin University of Science and Technology
Mar Athanasius College (Autonomous)
Kochi, India
Kerala, India
Øystein Evensen
Joyal Elisabeth Baiju Norwegian University of Life Sciences
Department of Microbiology Ås, Norway
Mar Athanasius College (Autonomous)
Kerala, India Uma Ganapathi
Aquatic Animal Health Laboratory, Centre for
Alanta Maria Binu Marine Science and Technology (CMST)
Department of Microbiology Manonmaniam Sundaranar University
Mar Athanasius College (Autonomous) Tirunelveli, Tamil Nadu, India
Kerala, India

xvii
xviii Contributors

Md Yasin Ina-Salwany Philippe Mahl

Aquatic Animal Health and Therapeutics Virbac Asia Pacific Co. Ltd.
Laboratory Bangkok, Thailand
Institute of Bioscience
Universiti Putra Malaysia Tilusha Manchanayake
Serdang, Selangor, Malaysia Department of Veterinary Pathology and
Microbiology
Sivakamavalli Jeyachandran Faculty of Veterinary Medicine
Department of Biotechnology and Universiti Putra Malaysia
Microbiology Serdang, Selangor, Malaysia
National College (Autonomous)
Tiruchirappalli, Tamil Nadu, India Meril Mary Mathew
Faculty of Fisheries
Nayomi John Kerala University of Fisheries and Ocean
Department of Microbiology Studies
Mar Athanasius College (Autonomous) Kochi, India
Kerala, India
Mariavincent Michael Babu
Anu Joseph Aquatic Animal Health Laboratory
Department of Microbiology Centre for Marine Science and Technology
Mar Athanasius College (Autonomous) (CMST)
Kerala, India Manonmaniam Sundaranar University
Tirunelveli, Tamil Nadu, India
Jitendra Kumar
College of Fisheries Aslah Mohamad
Acharya Narendra Deva University of Aquatic Animal Health and Therapeutics
Agriculture and Technology Laboratory
Uttar, Pradesh, India Institute of Bioscience
Universiti Putra Malaysia
Kavitha Kuppuswamy Serdang, Selangor, Malaysia
Department of Microbiology
Sri Sankara Arts and Science College S. J. Monaghan
Kanchipuram, Tamil Nadu, India Institute of Aquaculture
University of Stirling
Ihn-sil Kwak Stirling, Scotland, United Kingdom
Department of Fisheries
Chonnam National University Md. Shirajum Monir
Yesou, South Korea Bangladesh Fisheries Research Institute
Freshwater Station
Sreeja Lakshmi Mymensingh, Bangladesh
Aquaculture Research Group
Moredun Research Institute Kizito K. Mugimba
Penicuik, United Kingdom Department of Biotechnical and Diagnostic
Sciences
Po-Tsang Lee College of Veterinary Medicine
Department of Aquaculture Animal Resources and Biosecurity
National Taiwan Ocean University Makerere University
Keelung, Taiwan Kampala, Uganda
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