Table Of ContentAquaculture and Fisheries Biotechnology:
Genetic Approaches, 2nd Edition
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Aquaculture and Fisheries
Biotechnology: Genetic Approaches,
2nd Edition
Rex A. Dunham
Department of Fisheries and Allied Aquacultures
Auburn University
Alabama
USA
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A catalogue record for this book is available from the British Library, London,
UK.
Library of Congress Cataloging-in-Publication Data
Dunham, Rex A.
Aquaculture and fi sheries biotechnology: genetic approaches / Rex A. Dunham.
– 2nd ed.
p. cm.
Includes bibliographical references and index.
ISBN 978-1-84593-651-8 (alk. paper)
1. Fishes--Breeding. 2. Shellfi sh--Breeding. 3. Fishes--Molecular genetics.
4. Shellfi sh--Molecular genetics. I. Title.
SH155.5.D86 2011
639.3--dc22
2011003482
ISBN-13: 978 1 84593 651 8
CABI South Asia Edition ISBN: 978 1 84593 868 0
Commissioning Editor: Rachel Cutts
Editorial Assistant: Gwenan Spearing
Production Editor: Simon Hill
Typeset by AMA DataSet, Preston, UK.
Printed and bound in the UK by MPG Books Group.
Dedicated to
Petrcyle (Azset) – akong usa ka lawasnon, gihigugma ko ikaw hangtod sa hangtod
Aubrey, Nick and Julian; Christian, Nicole and Aaliyah; Amy; Gabrielle; Sean; and
Jeef – proof that genes work
Carol Jean – our Saint
Mom – enduring and loving
Scott, Earl and Dixie – educators all
Terry Abella – a true friend
The Dunham and Gealon families
In memory of my Dad
Richard Vincent (Dick) Dunham
December 1, 1919–August 11, 1971
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Contents
1. History of Biotechnology, Genetics and Selective Breeding in
Aquaculture and Fisheries 1
2. Phenotypic Variation and Environmental Effects 8
3. Basic Genetics, Qualitative Traits and Selection for Qualitative Traits 23
4. Strain Evaluation, Domestication and Strain Selection 33
5. Population Size, Inbreeding, Random Genetic Drift and Maintenance of
Genetic Quality 37
6. Gynogenesis, Androgenesis, Cloned Populations and Nuclear Transplantation 47
7. Intraspecifi c Crossbreeding 58
8. Interspecifi c Hybridization 65
9. Selection and Correlated Responses to Selection 73
10. Polyploidy and Xenogenesis 93
11. Sex Reversal and Breeding 128
12. B iochemical and Molecular Markers 150
13. Population Genetics and Interactions of Hatchery and Wild Fish 170
14. G enomics, Gene Mapping, Quantitative Trait Locus Mapping and
Marker-assisted Selection 190
15. Gene Expression 215
vii
viii Contents
16. Gene-transfer Technology 240
17. Combining Genetic Enhancement Programmes 278
18. Genotype–Environment Interactions 283
19. Commercial Application of Fish Biotechnology 288
20. Environmental Risk of Aquatic Organisms from Genetic Biotechnology 293
21. Food Safety of Transgenic Aquatic Organisms 311
22. A Case Example: Safety of Consumption of Transgenic Salmon
Potentially Containing Elevated Levels of Growth Hormone and
Insulin-like Growth Factor 315
23. Government Regulation of Transgenic Fish and Biotechnology Products 329
24. S trategies for Genetic Conservation, Gene Banking and
Maintaining Genetic Quality 339
25. Ethics 341
26. Constraints and Limitations of Genetic Biotechnology 344
Glossary 347
References and Further Reading 387
Index 481
1
History of Biotechnology, Genetics and
Selective Breeding in Aquaculture and Fisheries
Aquaculture is an ancient form of farming importance. Genetic variation is one key
dating back 2000 years or more in China variable in the survival of various species.
and to the Roman Empire (Balon, 1995; Also, natural populations are perhaps the
Dunham et al., 2001). However, only in the best gene banks, a critical resource for
last few decades has aquaculture grown into genetic variation for current and future
a global practice resulting in tremendous application in genetic improvement for
worldwide production. Aquaculture pro- farmed species and specialized sport-fi sh
duction has enlarged dramatically since the applications.
early 1980s, and will become increasingly Recreational fi shing is also of great
important as demand for fi sh products importance in many countries. When the
increases, world harvest by capture fi sher- revenue from the fi sh, licences, fi shing
ies reaches a plateau or declines and human equip ment, boats, travel, food and lodging
population numbers expand. Aquaculture is considered, recreational fi shing is proba-
now accounts for 50% of global fi sh con- bly more than tenfold more important eco-
sumption (FAO, 2009). The biomass of fi sh nomically than aquaculture in the USA.
that can be prod uced per surface area is Biotechnology is permanently linked not
much greater than that for terrestrial ani- only to aquacul ture, but also to commercial
mals, indicating that aquaculture could be and recreational fi sheries because of its
the key for providing global food security. potential positive and negative impacts on
Humans were hunter–gatherers prior to these resources.
being farmers and fi shermen before they Currently, the quantity of animal protein
were aquaculturalists. Although aquacul- harvested from global aquatic sources via the
ture is growing in importance and must capture of natural fi sh populations is at maxi-
expand to meet future demand for fi sh prod- mum sustainable yield. Many major fi sh
ucts, commercial harvest of natural popula- stocks are showing precipitous declines in
tions has traditionally been of higher productivity due to overfi shing and further
economic value than aquaculture and will increases are not anticipated under the cur-
continue to be of great importance. Even as rent global conditions and environment.
aquaculture closes the gap or surpasses the Wild fi sh stocks have been heavily fi shed or
value of commercial fi sheries, the genetic overfi shed, which has resulted in a noticeable
management and conservation of natural levelling of fi sh landings at around 60 million
fi sh stocks and gene pools will be of great t, with harvest from oceans unlikely to
© R.A. Dunham 2011. Aquaculture and Fisheries Biotechnology: Genetic Approaches,
2nd Edition (R.A. Dunham) 1
2 Chapter 1
expand (Hardy, 1999). Almost two-thirds of to the point that it is now having an impact
marine stocks in the Pacifi c and Atlantic on aquaculture worldwide, but potential
Oceans are being fully exploited or have maximum improvement in overall perfor-
already been overfi shed (Pauly et al., 1998). mance is not close to being achieved. As
The Food and Agriculture Organization of space for aquaculture becomes more
the United Nations (FAO) predicts a 36% limiting, the necessity for more effi cient
increase in the world population, with only a production or increased production within
30% increase in production from aquacul- the same amount of space will further
ture and fi sheries. Hardy (1999) predicts a increase the importance of genetic improve-
55 million t shortage in demanded seafood ment of aquaculture species. Genetic
products by 2025 resulting from levelled research and its application have had a sig-
wild catch and increasing demand. By 2025, nifi cant role in the development of aquacul-
aquaculture will have to increase by 350% to ture, and this role and impact will become
cover the impending shortage (Hardy, 1999). increasingly important as aquaculture
In 1993, approximately 16 million t of aqua- develops further.
cultured animal protein were produced, rep- Aquaculture genetics actually had its
resenting some 13% of the total aquatic origin with the beginning of aquaculture in
animal protein harvested or produced China and the Roman Empire more than
(Tacon, 1996). This grew to 59.4 million t by 2000 years ago. Without realizing it, the fi rst
2004 (FAO, 2006) and was approaching fi sh culturists changed gene frequencies
90 million t for 2009 (FAO, 2009). The growth and altered performance of the wild-caught
of aquacultured animal protein increased at a fi sh, actually genetically enhancing the fi sh
rate of over 11.8% annually from 1985 to for fi sh-farming application by closing the
1994, but slowed to 7.1% from 1995 to 2004, life cycles and domesticating species such
compared with the more modest growth of as the common carp, Cyprinus carpio. When
terrestrial meat production, which ranges the Chinese, Europeans and others observed
from 0.7% (beef) to 5.2% (poultry). mutations and phenotypic variation for
With increased demand for aqua- colour, body conformation and fi nnage, and
cultured foods has come a need for then selected for these phenotypes as well
more effi cient pro duction systems. Major as for body size, genetics and selective
improvements have been achieved through breeding of fi sh and shellfi sh was born.
enhanced husbandry procedures, improved Additionally, fi sh culturists and scientists
nutrition, enhanced disease diagnosis and who compared and evaluated closely
therapies and the application of genetics to related species for their suitability for aqua-
production traits. Although several aqua- culture application over the past two mil-
culture species have been greatly improved lennia were also unknowingly conducting
through the application of genetics, much some of the fi rst fi sh genetics research.
greater improvements can be accomplished Closely related species are reproductively
(Dunham et al., 2001; Dunham, 2004). isolated and have species status because of
Genetics can greatly contribute to produc- their genetic distance from one another;
tion effi ciency, enhancing production and therefore, the comparison of different spe-
increasing sustainability. Resource utiliza- cies is a genetic comparison (Dunham et al.,
tion can be greatly improved and impedi- 2001). However, directed breeding and
ments to sustainability, such as slow growth genetics programmes were probably not
of fi sh, ineffi cient feed conversion, heavy intense and strongly focused until the Japa-
mortality from disease and the associated nese bred koi in the 1800s and the Chinese
use of chemicals, loss of fi sh from low oxy- developed fancy goldfi sh.
gen levels, ineffi cient harvest, poor repro- Of course, fi sh biotechnology and
duction, ineffi cient use of land space and molecular genetics research and develop-
processing loss, can all be diminished by ment share the same beginnings as biotech-
utilizing genetically improved fi sh. Genetic nology and molecular genetics applied to
enhancement of farmed fi sh has advanced other organisms when in 1665 Robert Hooke