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Francesc Piferrer is Research Professor and Head of the Reproductive Physiology and Environmental Epigenetics Group at the Institute of Marine Sciences, Spanish National Research Council, Barcelona, Spain.
Hanping Wang is Principal Scientist, Research Professor, and Director of the Ohio Center for Aquaculture Research and Development at The Ohio State University, Piketon, Ohio, USA.
About the Editors xvii
List of Contributors xix
Preface xxiii
Acknowledgments xxv
Part I Theoretical and Practical Bases of Epigenetics in Aquaculture 1
1 The Potential Role of Epigenetics in Aquaculture: Insights from Different Taxa to Diverse Teleosts 3Han-Ping Wang and Zhi-Gang Shen
1.1 Introduction 3
1.2 Key Players of Epigenetics 4
1.3 Divergent Epigenetic Mechanisms from Different Taxa to Diverse Teleosts 10
1.4 The Roles and Applications of Epigenetics 11
1.5 Conclusion and Perspectives 25
2 Transcriptional Epigenetic Mechanisms in Aquatic Species 45Laia Navarro-Martín, Jan A. Mennigen, and Jana Asselman
2.1 Epigenetic Mechanisms as Modulators of Transcription 45
2.2 Transcriptional Epigenetic Mechanisms in Aquatic Species 51
2.3 Modulation of Biological Functions by Transcriptional Epigenetic Mechanisms in Aquaculture Species of Interest 54
2.4 Conclusions and Perspectives 57
3 Epigenetic Regulation of Gene Expression by Noncoding RNAs 65Elena Sarropoulou and Ignacio Fernández
3.1 General Introduction 65
3.2 Major Types of ncRNAs 65
3.3 Roles of ncRNA in Key Processes of Teleosts 76
3.4 ncRNAs as Biomarkers and Future Perspectives 84
4 Epigenetic Inheritance in Aquatic Organisms 95Ramji K. Bhandari
4.1 Introduction 95
4.2 Epigenetic Reprogramming of Embryo and Germline Cells 101
4.3 Heritable Effects of Environmental Stress 104
4.4 Past Exposure and Future Phenotypic Consequences in Aquatic Species 108
4.5 Conclusions and Perspectives 114
5 Environmental Epigenetics in Fish: Response to Climate Change Stressors 127Zhi-Gang Shen, Yue Yu, and Han-Ping Wang
5.1 Overview of Climate Change and Environmental Stressors 127
5.2 Epigenetic Response to Climate Change 129
5.3 Conclusions and Future Perspectives 137
6 Analytical Methods and Tools to Study the Epigenome 149Oscar Ortega-Recalde and Timothy A. Hore
6.1 Introduction 149
6.2 Recommendations for Choosing a Method to Study the Epigenome 150
6.3 Methods and Tools to Analyze Epigenetic Modifications 150
6.4 Bioinformatics Analysis 165
6.5 Databases and Other Public Resources 166
6.6 Conclusions and Outlook 166
Part II Epigenetics Insights from Major Aquatic Groups 175
7 Epigenetics in Sexual Maturation and Gametes of Fish 177Marta Lombó Alonso, Audrey Laurent, María Paz Herráez, and Catherine Labbé
7.1 Introduction 177
7.2 Epigenetics During Spermatogenesis and Oogenesis 177
7.3 Epigenetic Changes in the Gametes Triggered by Environmental Constraints 181
7.4 Conclusion 186
8 Epigenetics in Sex Determination and Differentiation of Fish 193Qian Wang, Qian Liu, Xiaona Zhao, Wenxiu Ma, Lili Tang, Bo Feng, and Changwei Shao
8.1 Introduction 193
8.2 Epigenetics and Sex Chromosome Evolution 195
8.3 Epigenetics and Sex Determination 198
8.4 Epigenetic Regulation of Sex Differentiation in Gonochoristic Species and Sex Change in Hermaphrodites 199
8.5 Transgenerational Epigenetic Sex Reversal 201
8.6 Conclusions and Future Perspectives 203
9 Epigenetics in Fish Growth 209Jorge M.O. Fernandes, Artem V. Nedoluzhko, Ioannis Konstantinidis, and Paulo Gavaia
9.1 Myogenesis in Teleosts 209
9.2 Skeletogenesis in Teleosts 213
9.3 Epigenetic Regulation of Sexually Dimorphic Growth 215
9.4 Epigenetic Control of the Skeleton in Teleosts 218
9.5 Mitochondrial Epigenetics 219
9.6 Conclusion 221
10 Epigenetics in Fish Nutritional Programming 231Kaja H. Skjærven, Anne-Catrin Adam, Takaya Saito, Rune Waagbø, and Marit Espe
10.1 Epigenetic Basis of Nutritional Programming 231
10.2 Nutritional Programming 233
10.3 Key Nutrients and Metabolites for Epigenetic Mechanisms 235
10.4 Case Examples 237
10.5 Conclusions and Perspectives for Nutritional Programming in Aquaculture 239
11 Microbiome, Epigenetics, and Fish Health Interactions in Aquaculture 245Sofia Consuegra, Tamsyn Uren Webster, and Ishrat Anka
11.1 Introduction 245
11.2 The Fish Microbiome in Aquaculture 245
11.3 Microbiome-Epigenome Interactions 252
11.4 Gaps in Knowledge and Future Research Avenues 255
11.5 Conclusions 255
12 Epigenetics of Stress in Farmed Fish: An Appraisal 263Bruno Guinand and Athanasios Samaras
12.1 Introduction 263
12.2 Stress and Stress Response 264
12.3 Is There an Epigenetics of Stress in Cultured Fish? 267
12.4 The Neuroepigenetics of Stress: Fishing with Mammalian Models 269
12.5 Epigenetic Biomonitoring of Stress 273
12.6 Conclusions 274
13 Epigenetics in Hybridization and Polyploidization of Aquatic Animals 287Li Zhou and Jian-Fang Gui
13.1 Hybridizing and Hybridization 287
13.2 Polyploidy and Polyploidization 287
13.3 Epigenetic Changes and Effects During Hybridization and Polyploidization in Aquatic Animals 289
13.4 Association of Epigenetic Changes with Heterosis 292
13.5 Conclusions and Future Perspectives 293
14 Epigenetics in Aquatic Toxicology 301Sara J. Hutton and Susanne M. Brander
14.1 Introduction 301
14.2 Epigenetic Endpoints in Aquatic Toxicology Studies 303
14.3 Epigenetics During Early Development Related to Toxicology 310
14.4 Multigenerational and Transgenerational Toxicology 311
14.5 Epigenetics in Ecological Risk Assessment 313
14.6 Rapid Evolution 314
14.7 Epigenetics in Aquaculture 315
14.8 Conclusion and Perspectives 316
15 Epigenetics in Mollusks 325Manon Fallet
15.1 Introduction 325
15.2 DNA Modifications in Mollusk Species 328
15.3 Chromatin Conformation and Histone Modifications/Variants in Mollusks 330
15.4 Noncoding RNAs in Mollusks 331
15.5 Epigenetic Responses to Environmental Fluctuations in Mollusks 336
15.6 Mechanisms of Meiotic Epigenetic Inheritance in Mollusks and Their Impact in Evolution 340
15.7 Perspectives 345
15.8 General Conclusions 346
16 Epigenetics in Crustaceans 355Günter Vogt
16.1 Introduction 355
16.2 Epigenetics Research with Brine Shrimps and Copepods 356
16.3 Epigenetics Research with Water Fleas 359
16.4 Epigenetics Research with Amphipods 363
16.5 Epigenetics Research with Freshwater Crayfish 363
16.6 Epigenetics Research with Shrimps and Crabs 371
16.7 State of the Art of Epigenetics in Crustaceans 373
16.8 Potential Application of Epigenetics in Crustacean Aquaculture 374
17 Epigenetics in Algae 383Christina R. Steadman
17.1 Introduction: What Are Algae 383
17.2 Algae Epigenetics 388
17.3 Environmental Stress Alters Microalgae Epigenomes 404
17.4 Conclusions and Perspectives 405
Part III Implementation of Epigenetics in Aquaculture 413
18 Development of Epigenetic Biomarkers in Aquatic Organisms 415Dafni Anastasiadi and Anne Beemelmanns
18.1 Biomarkers 415
18.2 Epigenetic Biomarkers 415
18.3 Development of Epigenetic Biomarkers 417
18.4 Epigenetic Biomarkers in Aquatic Organisms and their Applications in Aquaculture 425
18.5 Future Perspectives 431
18.6 Concluding Remarks 432
19 Genetics and Epigenetics in Aquaculture Breeding 439Shokouoh Makvandi-Nejad and Hooman Moghadam
19.1 Overview 439
19.2 Breeding in Aquaculture and Evolution of Genetic Markers 440
19.3 Epigenetics and Missing Heritability 442
19.4 Transgenerational Inheritance of Epigenetic Marks 444
19.5 Epigenetic Marks -- Possible Biomarkers to Improve Breeding 444
19.6 Association Analysis and Search for Epigenetic Biomarkers 445
19.7 Concluding Remarks 446
20 Epigenetics in Aquaculture: Knowledge Gaps, Challenges, and Future Prospects 451Francesc Piferrer
20.1 Introduction 451
20.2 Knowledge Gaps 452
20.3 Challenges 456
20.4 Prospects 458
Acknowledgments 461
References 461
Index-Species 465
Index-Subjects 469
Aquaculture is the fastest food production sector in the world and prospects are that this position will be maintained for years to come. According to data from the Food and Agriculture Organization (FAO) of the United Nations, in 2020 global aquaculture production reached a record of 122.6 million tons worth USD 281.5 billion. Animals accounted for 87.5 million tons while algae comprised 35.1 million tons. However, aquaculture must become more sustainable to meet the growing demand for aquatic foods of an ever-increasing human population. Thus, improved aquaculture production requires further technical innovations, including more focus on breeding programs, feed utilization, well-being, and disease control. Similar to any other food production system, aquaculture is about producing the phenotypes with superior value. In this endeavor, new advances on our understanding of the epigenetic regulation of the phenotype have the potential to play an increasing role in achieving aquaculture production sustainability.
The term "epigenetics" was coined by Conrad Waddington in the 1940s, but with a meaning different from how it is understood today. Initially, it was essentially related to what today is understood as the field of developmental biology and how the phenotype comes into being. However, the modern concept of epigenetics, i.e., "the study of mitotically and/or meiotically heritable changes in gene function that cannot be explained by changes in DNA sequence," arose in the mid-1990s and around the turn of this century. The field has largely benefited especially from the advancements made after the sequence of the human genome, the characterization of the regulatory elements, and all emerging technologies to interrogate different aspects of the genome and epigenome.
Epigenetics is now considered one of the "hot topics" in biology. Epigenetic modifications or "marks" can be easily identified, and they constitute therapeutic approaches for the treatment of an increasing number of diseases. Thus, there is a lot of research ongoing in the epigenetics of cancer, for example.
There are three very important aspects to take into account when dealing with epigenetics. First, epigenetics integrates genomic and environmental influences to bring about the phenotype. Second, there is a fraction of the phenotypic variance that cannot be explained solely on genetic variation, but that can be explained by taking into account epigenetic variation. Third, epigenetic changes can be inherited and thus passed from parents to offspring into the following generations. Combined, this has prompted the implementation of epigenetic research, not only in ecology and evolution for its contribution to adaptation to new environments, but also into agriculture and livestock for improved food production. Consequently, recently there has been both a clear interest in marine epigenetics and in the application of epigenetics in aquaculture. One of the main reasons is that aquatic organisms are quite susceptible to environmental cues since, for example, temperature in a cold-blooded animal influences growth rates more strongly than in a warm-blooded animal. Further, in contrast to mammals, fishes seem to have less reprogramming and erasing of epigenetic marks after fertilization, thus facilitating epigenetic transmission of environmental influences to the next generation. Thus, there is a lot of interest for the application of epigenetics in aquaculture. However, and to the best of our knowledge, there are currently no books that address this need.
"Epigenetics in Aquaculture" consists of 20 chapters and is arranged into three parts: Part I: Theoretical and practical bases of epigenetics in aquaculture; Part II: Epigenetics insights from major aquatic groups; and Part III: Implementation of epigenetics in aquaculture. All chapters are written by top specialists with ample experience and at the forefront in their respective research fields.
Part I contains six chapters (Chapters 1-6) and provides the necessary background to understand what epigenetics is about and what are the major mechanisms and phenomena. The first chapter covers the overall roles and the diversity of epigenetic mechanisms across major taxa and provides insights into their potential applications in aquaculture and aquatic animals. The following two chapters are devoted to the three main epigenetic mechanisms regulating gene expression, namely, DNA methylation, histone modifications, and non-coding RNAs. The next two chapters are devoted to two key aspects of epigenetics. One explains how epigenetic modifications can be inherited across different generations, a hot topic in different areas of biology, and the other elucidates the role of epigenetics in integrating environmental cues as a powerful mechanism in the adaptation and the basis of organismal plastic responses to rapid environmental change. The last chapter of Part I presents the currently available methods to analyze the epigenetic modifications including the latest developments, as well as some basic resources for the bioinformatics analysis of the data. It also explains how to choose among the different approaches based on the type of question that one aims to answer.
Part II contains 11 chapters (Chapters 7-17) and constitutes the bulk of the book. These chapters explore the roles of epigenetic regulatory mechanisms in key biological process and their relevance for aquatic production. The first two chapters deal with epigenetic sex determination and differentiation as well as the dynamics of epigenetic marks during gametogenesis and early development. The following two chapters are devoted to growth, with one focusing on skeletal muscle and the other emphasizing nutritional programming. The next two chapters of Part II are devoted to the epigenetics of stress response, immune response, and the emerging topic of the role of the microbiome in shaping epigenetic responses of the host. Additionally, one chapter is focused on epigenetics in hybridization and polyploidy and another on how epigenetics can contribute to explain organismal responses to toxins present in the aquatic environment. Many of the above-cited chapters of Part II focus on fish, where considerable work has been carried out so far. Thus, this part ends with three chapters dedicated to the epigenetics of other taxa that are also very important for aquaculture production, namely mollusks, crustaceans, and algae, where interesting discoveries related to similarities and differences with the situation in vertebrates are being made.
Finally, Part III includes the final three chapters (Chapters 18-20), dealing with the actual integration of epigenetics into aquaculture practice. For this, the development of biomarkers and their applications in aquaculture is discussed. Particular attention is then paid on the integration of epigenetic selection into current genetic breeding programs. The final chapter identifies knowledge gaps, discusses challenges that must be overcome, and outlines future prospects on the application of epigenetics in aquaculture. In addition to tables, figures, and abundant bibliography, each chapter contains a glossary of terms used with pertinent definitions.
Thus, this book provides an update on the state-of-the-art on the knowledge of epigenetic regulatory mechanisms in major taxa of aquatic organisms including algae, crustaceans, mollusks, and fish and how this new knowledge can be applied to increase aquaculture production. It covers both basic and applied aspects of epigenetics related to reproduction, development, growth, nutrition, and disease of aquatic species, which we hope will benefit the aquatic scientific community and the aquaculture sector.
This book will be appealing to anyone interested in knowing all major aspects related to epigenetics, including mechanisms, inheritance, methodology, etc. Information contained within will be particularly useful to researchers working on epigenetics in aquatic animals and aquaculture, including basic aspects of fish and shellfish epigenetics, reproductive endocrinology, genetics, and evolutionary and environmental biology. It will also appeal to PhD and MSc students and biologists working in hatcheries or in breeding companies, who will all benefit from reading about epigenetics and the opportunities it can provide. More broadly, aquatic biologists, including fisheries managers and conservation biologists, will also benefit from clear and practical information in epigenetics. The epigenetic insights from fish, shellfish, and aquatic model species will attract readers from other disciplines as well, who might find inspiration in findings made on epigenetics of aquatic organisms.
Our previous book, "Sex Control in Aquaculture," was based on knowledge accumulated after decades of applying sex control techniques to improve aquatic productivity. In contrast, the present book is based on research that is just a few years old because the study of epigenetics in aquatic organisms and its application in aquaculture is still in its infancy. Thus, a lot remains to be done. We hope that our efforts in providing a comprehensive picture of the current situation will be of much help and foster future research. The well-known British zoologist D'Arcy Thompson (1860-1948) wrote in the preface of his most famous book "On Growth and Form" that "this book of mine has little need of preface, for indeed it is 'all preface' from beginning to end." Given...
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