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A Primer of Ecological Aquaculture
Endorsements
“This important new volume on aquaculture presents a remarkable synthesis of all the key issues involved in the com-
mercial culturing of freshwater and marine species. It provides a comprehensive overview of the basic biological elements
of aquaculture, including new information on the use of genomic and genetic tools, and incorporates these analyses into
a broader ecological, environmental, and sociological context. The emphasis on strategies for protecting water quality—
a vital part of the aquaculture enterprise—is especially well-presented. The volume deserves wide readership among
students and scientists who not only have interests in aquaculture, but also in sustaining robust aquatic ecosystems in
freshwater and marine environments.”
George N. Somero, David and Lucile Packard Emeritus
Professor of Marine Science, Hopkins Marine Station, Stanford University, USA
“Dietmar Kültz has distilled many years of teaching an undergraduate course in aquaculture into an engaging and acces-
sible text. The book is a huge endeavour and presents an honest and rigorous assessment of aquaculture practices with
clear, detailed descriptions of invertebrate and vertebrate aquatic culture, diseases, and water management. The ‘Key
Conclusions’ at the end of every chapter are particularly useful for students.”
Patricia A. Wright, Professor, Department of Integrative Biology,
University of Guelph, Canada
“A must-read primer for anyone considering aquaculture either as undergraduate and postgraduate students or aquacul-
ture farmers. Dietmar Kültz takes readers on a fascinating journey into the world of ecological aquaculture to stimulate
awareness of the need to reach a more sustainable future.”
Silvia Gomez-Jimenez, Director of Accredited Aquaculture Diagnosis Laboratory,
Research Centre for Food and Development, Mexico
“A Primer of Ecological Aquaculture by Dietmar Kültz is the latest addition to the growing list of entry-level books on the
topic of aquaculture. It differs from other books, however, on its emphasis on ecological aspects of aquaculture. As such,
it is like a new tool that seems impossible to have lived without before it was acquired. The book succeeds by providing
a holistic assessment of various aquaculture production systems including inputs, outputs and how they contrast and
compare among various species groups. Most importantly, the book presents both problems and opportunities associ-
ated with aquaculture in a concise and balanced manner. Aquaculture production must increase to meet future seafood
demand and this book provides solid information and guidance on how society can meet seafood demand by employing
sustainable aquaculture practices.”
Ronald Hardy, Professor Emeritus, Department of Animal,
Veterinary, and Food Sciences, University of Idaho, USA
“As the demand for wild caught seafood is continuing to increase while stock volumes are rapidly decreasing, our future
food security will rely more heavily on ecologically sustainable aquaculture practices. Kültz presents a detailed exploration
of its limitations and novel paths to diminish its ecological footprint, enriched with comprehensive analysis and clear
graphics. This accessible primer systematically dissects the rationales behind current practices, from monocultures and
polycultures to microcosms and mesocosms. It also provides clear signposts to sustainable practices aimed at minimizing
the exploitation of natural ecosystems, while maximizing production efficiency and food security.”
Gillian Renshaw, Professor, School of Allied Health Sciences,
Griffith University, Australia
A Primer of Ecological
Aquaculture
Dietmar Kültz
Professor of Physiological Genomics, College of Agricultural and Environmental
Sciences, University of California Davis, USA
Great Clarendon Street, Oxford, OX2 6DP,
United Kingdom
Oxford University Press is a department of the University of Oxford.
It furthers the University’s objective of excellence in research, scholarship,
and education by publishing worldwide. Oxford is a registered trade mark of
Oxford University Press in the UK and in certain other countries
© 2022 Dietmar Kültz
The moral rights of the author have been asserted
Impression: 1
All rights reserved. No part of this publication may be reproduced, stored in
a retrieval system, or transmitted, in any form or by any means, without the
prior permission in writing of Oxford University Press, or as expressly permitted
by law, by licence or under terms agreed with the appropriate reprographics
rights organization. Enquiries concerning reproduction outside the scope of the
above should be sent to the Rights Department, Oxford University Press, at the
address above
You must not circulate this work in any other form
and you must impose this same condition on any acquirer
Published in the United States of America by Oxford University Press
198 Madison Avenue, New York, NY 10016, United States of America
British Library Cataloguing in Publication Data
Data available
Library of Congress Control Number: 2022930150
ISBN 978–0–19–885022–9 (hbk)
ISBN 978–0–19–885023–6 (pbk)
DOI: 10.1093/oso/9780198850229.001.0001
Printed and bound by
CPI Group (UK) Ltd, Croydon, CR0 4YY
Cover image: High-density aquaculture of Bigeye scad (Selar crumenophthalmus)
is supported by their natural schooling behaviour. Photo by author.
Links to third-party websites are provided by Oxford in good faith and
for information only. Oxford disclaims any responsibility for the materials
contained in any third party website referenced in this work.
About the Author
Aquaculture exemplifies the ongoing global strug- is necessary. The book carries out such an assess-
gle to strike a sustainable balance between the ment from the standpoint of how best to align the
conflicting needs of a large human world popula- two major (and often conflicting) goals of future
tion, human health, ecosystem health, the welfare aquaculture development: minimizing reliance on
of wild and domesticated animals, and the eco- ecosystem services while maximizing productivity.
nomic principles of globalized economies. On the This book represents a broad snapshot assess-
one hand, aquaculture has great potential for pro- ment of the current state, past developments, and
viding us with a healthy and nutritious food sup- prospects of applying knowledge in aquatic animal
ply while alleviating pressure on capture fisheries biology and engineering to the captive culture of
and reducing fisheries-induced habitat destruction, aquatic animals. My intent was to transcend tra-
overfishing, genetic modification of wild popula- ditional disciplinary boundaries whenever possi-
tions, and wholesale waste of bycatch. On the other ble and present contrasting perspectives whenever
hand, aquaculture relies heavily on clean water, an applicable to stimulate critical assessment of aqua-
increasingly precious (and dwindling) resource that culture practices by students. There are many differ-
is subject to intense pressure through being used ent opinions about the best way to further develop
for many competing purposes. The multifaceted aquaculture. The perspective presented here should
nature of aquaculture is most evident in the divi- not be taken dogmatically, but rather should stimu-
sion of approaches between those that emphasize late critical thinking in readers. To make the book
minimal reliance on ecosystem services (e.g. recir- more accessible, e.g. to students taking a semester-
culating aquaculture systems, aquaponics) versus or quarter-long course on this subject, it presents
those aimed at maximizing profitability (e.g. open the material concisely and provides select exam-
pond, cage, and raceway systems). Ecological con- ples to illustrate pertinent points and key concepts,
siderations have received increasing attention in and it explains scientific and aquaculture-specific
aquaculture since the turn of the millennium and terminology when first used. It is not my intent
their proper implementation promises to greatly to comprehensively review specific commodities or
accelerate the rate at which the field moves forward. go into undue detail regarding specific aspects of
This book aims to introduce students to the basic aquaculture if such detail is unnecessary to support
concepts, opportunities, and challenges of aquacul- salient facts and arguments.
ture with an emphasis on ecological considerations. The motivation for writing this book arose from
I contrast the many values and promises of aquacul- feedback provided by my students and colleagues.
ture (including specific approaches and commodi- I have been teaching a general education (GE) class
ties) with their pitfalls, drawbacks, and challenges. on aquaculture (ANS 18) at the University of Cali-
My goal is to provide students with a broad under- fornia, Davis for more than a decade but a suitable
standing of the general state of aquaculture to equip text accompanying the class material was missing.
them with the knowledge they need to contribute Most students I have taught prefer single-author
to future advances in the field. To achieve this texts that require little prior knowledge of a subject
goal, a critical assessment of current aquaculture and are written in a consistent style, rather than
practices from a broad, interdisciplinary perspective more specialized volumes comprised of chapters by
P R E FA C E vii
different authors. Although there are many excel- suggestions for improving individual chapters of
lent books focusing on aquaculture, some of which this book, including Professors Steve McCormick
are referenced in this volume, an integrated discus- (University of Massachusetts), Ron Hardy (Univer-
sion of the ecological cornerstones of aquaculture in sity of Idaho), Colin Brauner (University of British
the form of a concise ‘primer’ was needed to help Columbia), Patricia Wright (University of Guelph),
students learn pertinent concepts and principles. Brian Sardella (California State University Stanis-
This book thus attempts to serve as such a ‘primer’ laus), Wes Dowd (Washington State University),
in a format that is accessible to readers from a broad Jonathon Stillman (San Francisco State Universi-
range of academic backgrounds and experience. It ty), Silvia Gomez-Jimenez (CIAD and University of
is meant to consolidate and expand on my lecture Sonora), Alison Gardell (University of Washington),
notes to provide a study guide for students inter- Peter Allen (Mississippi State University), Frederick
ested in both the natural biology and the culture of Silvestre (University of Namur), Jason Podrabsky
aquatic organisms, both marine and freshwater. (Portland State University), Katie Gilmour (Uni-
Although I majored in marine ecology when at versity of Ottawa), Tyler Evans (California State
university, my own research now focuses heavi- University East Bay), Xiaodan Wang (East China
ly on biochemical mechanisms of environmental Normal University Shanghai), and Esteban Soto
tolerance and stress-induced evolution in fish and and Jackson Gross (both University of California
marine invertebrates using molecular, cellular, and Davis). Many thanks also go to my graduate stu-
systems biology approaches. It seems natural to dents Elizabeth Mojica, Meranda Corona, Larken
investigate molecular phenomena at an increasing Root, and Jens Hamar, who read chapter drafts
level of depth to explain how organisms function and provided photos and valuable feedback that
within their environment and how life works. How- helped improve the book. I also thank my col-
ever, at the same time it is important to integrate leagues including Professors Avner Cnaani (Agri-
reductionist approaches with holistic approaches cultural Research Organization, Beth Dagan), Delin
that explain the function of systems, e.g. ecosystems Duan (Institute of Oceanology Chinese Academy
or aquaculture systems, that are more than just the of Sciences), Steve McCormick (University of Mas-
sum of their parts. This book emphasizes the holistic sachusetts), Olivier DeClerck (Ghent University)
view throughout. and Esteban Soto (University of California, Davis),
This book could not have been completed without and Miguel Sepulveda (Rio de Janeiro, Brazil),
the support and encouragement from my family, Christian Veterlaus (marinecultures.org), and Mark
friends, and colleagues. Many thanks go to my fam- Zivojnovich (Hydromentia Technologies) who have
ily, especially my wife, son, and parents for always kindly shared illustrations for the book. Their con-
supporting my pursuit of scientific research and tributions are acknowledged in the corresponding
enduring my spending long hours in the laborato- figure legends.
ry and at the desk pondering scientific questions. Finally, I thank my editorial contacts at Oxford
Their patience, encouragement, and feedback were University Press, Ian Sherman, Charlie Bath and
invaluable for completing this book project, which Sharmila Radha, who kept my writing on track with
took more effort and time than I had originally the planned schedule for publication and compat-
anticipated. ible with the projected size of this book. Ian and
I am also particularly grateful to Professor Charlie helped with all aspects of the book produc-
George Somero (Stanford University), my post- tion process and in keeping the page number from
doc mentor and long-term friend, who has read increasing beyond a limit that we felt would dimin-
and commented on early drafts of all chapters. ish accessibility. The collective feedback and great
His knowledgeable feedback and constructive com- advice I have received from many people helped
ments helped greatly in shaping the final draft of substantially improve this book although, of course,
the book and pushing it over the finish line on I am solely responsible for any omission or errors
time. I am also very grateful to other colleagues that may remain.
and friends who have read and made valuable
Dietmar Kültz,
Davis, CA, November 10, 2021
Contents
12.6 The impact of public media and perception on ornamental fishes 161
12.7 Invasiveness of ornamental fishes 162
Key conclusions 164
References 165
Index 277
PART 1
‘Science and engineering is meant to be questioned, tested and re-tested, but the road of true
progress can be long, convoluted and tiresome.’
Adey and Loveland, Dynamic aquaria: building living ecosystems (2007)
This book introduces the basic concepts, approaches focuses on their potential for meet-
opportunities, and challenges of aquaculture ing future demand in an economically feasible
with an emphasis on ecological considerations. manner while minimizing the exploitation of com-
The road to progress in aquaculture has been mon property resources and reliance on ecosystem
followed for several millennia leading to a dramatic services.
increase in its global productivity. We have arrived Although socio-economic aspects are critical for
at a major, challenging junction on this road and practical implementation of aquaculture solutions,
future advancement of aquaculture depends on a they are not considered here in the interest of brevi-
thorough understanding of the ecological value of ty and to maintain focus. They have been discussed
clean water as its main resource. extensively elsewhere and the reader is referred
Therefore, this book outlines links between aqua- to recent works that highlight this important topic
culture, trophic cascades, nutrient cycles, and (Bunting, 2013; FAO, 2020).
ecosystem services that determine water quality Specific aquaculture practices are illustrated for a
and ecosystem stability. A basic understanding of selection of economically important aquatic species
such links reveals opportunities for increasing the in Part 2 of this book. These examples illustrate key
ecological sustainability of aquaculture and makes concepts and contrast the values and promises of
the diverse but interwoven facets of ecological aqua- aquaculture with their pitfalls, drawbacks, and chal-
culture accessible to students entering this field. lenges to instil a broad understanding of the current
I assess the prospects of various aquaculture prac- state of aquaculture and prospects for increasing its
tices for key species groups based on the premise ecological sustainability.
of aligning two major (and often conflicting) devel- It was not my intent to comprehensively review
opment goals: minimizing reliance on ecosystem specific aquaculture commodities or practices and
services while maximizing productivity. omissions in this regard are unavoidable in this
The first part of the book builds a general under- introductory overview. The reader is referred to
standing of terminology, the history, objectives, excellent recent volumes that focus on particular
approaches, and ecological implications of aqua- aspects of aquaculture more comprehensively (e.g.
culture. It also emphasizes prospects for further Pillay and Kutty 2005; Bert, 2007; Davenport et al.,
development of ecologically sustainable aquacul- 2008; Costa-Pierce, 2014; Boyd and McNevin, 2015;
ture approaches. The discussion of aquaculture Stickney, 2017; Lucas, Southgate, and Tucker, 2019).
A Primer of Ecological Aquaculture. Dietmar Kültz, Oxford University Press. © Dietmar Kültz (2022). DOI: 10.1093/oso/9780198850229.003.0001
4 A P R I M E R O F E C O L O G I C A L A Q UA C U LT U R E
In addition, there are many volumes that cover are overexploited, and 8% are depleted (with 1%
aquaculture practices for a particular species or recovering), while 20% are moderately exploited
species group in great detail, e.g. oysters (Hanes, and only 3% are underexploited by capture fisheries
2019), shrimp (Leung and Engle, 2010), and tilapia (FAO, 2011).
(El-Sayed, 2019). Aquaculture and capture fisheries play an essen-
Part 3 provides an overview of abiotic parameters tial role in ensuring food supply and security in the
and biotic factors that are critical for water qual- context of climate change, in particular in develop-
ity management in aquaculture. Clean water is a ing countries. These tasks are recognized in the 2030
common property resource that affects all. Aqua- UN Agenda for Sustainable Development that com-
culture competes for this resource with other uses, prises seventeen sustainable development goals
including as drinking water, for sanitary purposes, (SDGs) (United Nations, 2015). One of these goals
for ecosystem health and species conservation, for (SDG 14) emphasizes the conservation and respon-
agricultural irrigation, and for human recreational sible use of the oceans and marine resources for
and cultural activities. sustainable development, which includes aquacul-
Most importantly, clean freshwater is limited and ture development governed by proper ecological
its value as a resource is increasing in parallel to stewardship.
the rising world population. Nevertheless, freshwa- Aquaculture exemplifies the ongoing global
ter (FW, also referred to as land-based) aquaculture struggle to strike a sustainable balance between
has greater prospects for future sustainable devel- the many (often conflicting) needs of a rapidly
opment than mariculture. Throughout this book increasing world population. Such needs pertain
I provide examples supporting this argument. to human health and nutrition, ecosystem health,
Clean water, whether fresh or marine, can no longer the welfare of wild and domesticated animals,
be considered a free natural resource but must be the cultural history of peoples, and the economic
valued by properly incentivizing its conservation. principles of globalized economies.
On the one hand, aquaculture has great poten-
tial for securing a healthy and nutritious food sup-
1.1 What is aquaculture?
ply while alleviating pressure on capture fisheries
According to the Food and Agriculture Organiza- and reducing fisheries-induced habitat destruction,
tion (FAO) of the United Nations (UN), ‘Aquacul- overfishing, genetic modification of wild popula-
ture is the farming of aquatic organisms in both tions by fisheries-induced evolution, and wholesale
coastal and inland areas involving interventions in waste of bycatch.
the rearing process to enhance production.’ (FAO, On the other hand, aquaculture relies heavi-
2019). Farming is the rearing of organisms under ly on clean water, an increasingly precious (and
(semi-)controlled conditions and implies owner- dwindling) resource that is subject to intense pres-
ship (by an individual or corporation) over the sure of being used for many competing purposes.
organisms that are being reared. In contrast to Although transitions between solid and liquid state
aquaculture, capture fisheries collect seafood from water occur more rapidly because of climate change,
wild populations, i.e. from natural, unmanaged the total amount of water on earth has not changed
resources. much.
The organisms that are subject to capture fish- The amount of available clean water is decreas-
eries are considered common property resources ing because water pollution has increased great-
that can be exploited and are not owned by any- ly over the past few centuries, which affects the
one, although fishing rights often apply. Cap- utility of available water for many human needs.
ture fisheries yield has plateaued at approximately The rate of pollution of the world’s inland water
90 million metric tons annually since the turn of bodies mirrors the exponential growth of the
the millennium while aquaculture yield continues human population, although environmental pro-
to increase steadily (Figure 1.1). In fact, 52% of world tection efforts have improved this trend in recent
marine fish stocks are already fully exploited, 17% decades. Even the seemingly vast reservoir of ocean
A Q UA C U LT U R E T E R M I N O L O G Y A N D B A S I C C O N C E P T S 5
Marine capture fisheries - All other countries Marine capture fisheries - China
Freshwater capture fisheries - All other countries Freshwater capture fisheries - China
Aquaculture - All other countries Aquaculture - China
180
Millions
160
140
Production yield (tons)
120
100
80
60
40
20
0
2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018
Year
Figure 1.1 Yields of seafood production by aquaculture and capture fisheries from 2005 to 2018. Global yields excluding China and yields
produced in China are shown separately. Data source: OECD.stat.
water is increasingly impacted by anthropogenic rivalries as this resource became scarcer and its
pollution. usage more prolific.
A rising world population requires more FW for Although crucial for assessment of the ecological
drinking, sanitary, agricultural, and other purposes. impact of aquaculture practices, ‘comparatively lit-
FW is also essential for maintaining the balance of tle effort has been expended on documenting and
inland ecosystems. Clean seawater (SW) is required analysing the range of methods used in exploita-
to maintain ecosystem health and species diversi- tion of aquatic environments’ (Beveridge and Little,
ty in the oceans, which is particularly critical in 2014). Nonetheless, consumer awareness and inter-
near-shore areas exploited by aquaculture, includ- est in the sources of seafood is increasing.
ing coral reefs, kelp forests, and mangrove lagoons. Educational programmes, such as the SeaFood
Watch programme of the Monterey Bay Aquari-
um, offer relevant knowledge and many markets
now provide detailed information about seafood
sources and methods of harvest such that this
1.2 Sustainable aquaculture
knowledge can be factored into consumer choices
development (Figure 1.2).
Ecological considerations have received increas- There is a growing public resentment against
ing attention in aquaculture during the past few semi-intensive farming systems that rely heavily
decades and their proper implementation promis- on common property resources and ecosystem ser-
es to greatly accelerate the rate at which the field vice capacity (open systems). Another main concern
moves forward. As a result of the industrial revo- with semi-intensive farming is animal welfare. To
lution, clean water, in particular FW, became pro- meet high seafood demand and address such con-
gressively precious as a resource and competition cerns, aquaculture development goals should pri-
over water rights and access have resulted in fiercer oritize practices that maximize yields while also
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Pessimist, The, Ben King, 727
Peter Simple (extracts), Frederick Marryat, 474
Petronius, 101
Ingenious Cook, An (from Trimalchio’s Banquet), 102
Philippides, Epigrams, 84
Philips, John,
Splendid Shilling, The, 423
Phillis’ Age, Matthew Prior, 389
Philosopher, A, Sam Walter Foss, 718
Philosopher, The (from The Fables), Ivan Chemnitzer, 631
Phoenix, John. See Derby, George Horatio
Phoenixiana (extract), George Horatio Derby, 678
Phyllis Lee, Oliver Herford, 744
Pictorial humor, 27, 46, 47, 48
Pigtail, The, Adelbert von Chamisso, 605
Pike County Ballads (extract), John Hay, 690
Pilpay (or Bidpai), Fables, 120;
(Selections), 164–170
Pitrá,
The Peasant of Larcarà, 218
Plato,
idea of humor, 4
Miser and the Mouse, The, 190
Thief and the Suicide, The, 189
Plato Comicus, fragments, 66
Plautus, 87
Military Swagger (from The Braggart Captain), 88
Suspicious Miser, The (from The Pot of Gold), 91
Playfulness of animals, 18
Play’s the Thing, Martial, 107
Pleasant Conceits of Old Hobson, The (extract), Richard
Johnson, 265, 267
Pleasantries of Cogia Nasr Eddin Effendi, The (extracts), 199
Pleasure of Fishes, The (from Autumn Floods), Chwang Tze,
157
Poems in Prose, Ivan Turgenieff, 638
Poetic Fame (from Satires), Persius, 104
Poets, Samuel Butler, 377
Poets at Tea, The, Barry Pain, 551
Poggio, Italian stories, 182
Polish humor, 639–641
Pontalais, Jehan du,
Money, 322
Pope, Alexander, 17
Epigram on Mrs. Tofts, 421
Lines by a Person of Quality, 419
Worms, 420
Pope and Sultan (German Student Song), 613
Pope and the Net, The, Robert Browning, 502
Popularity, Sung Yu, 158
Praed, Winthrop Mackworth,
Song of Impossibilities, A, 484
Praise of Folly, The (extracts), Desiderius Erasmus, 337
Prayer, Ivan Turgenieff, 638
Prior, Matthew, 386
Epitaph, An, 387
Phillis’ Age, 389
Reasonable Affliction, A, 389
Simile, A, 388
Prodigal Egg, The, Oliver Herford, 747
Professional entertainers of the Middle Ages, 231–236
Professor with a Small Class, A, Lucilius, 77
Professor’s Malady, The (from Water Babies), Charles Kingsley,
498
Proverbial Wisdom, Anton Chekov, 639
Provinciales, Les (extract), Blaise Pascal, 400
Psycholophon, Frank Gelett Burgess, 749
Puffing, Samuel Butler, 377
“Punning” (from Speeches), Sydney Smith, 446
Purple Cow, The, Frank Gelett Burgess, 748
Python, The, Hilaire Belloc, 555