Genetica 111: 155173, 2001.

2001 Kluwer Academic Publishers. Printed in the Netherlands.

155
Genetic manipulations in aquaculture: a review of stock improvement by
classical and modern technologies
G. Hulata
Department of Aquaculture, Agricultural Research Organization, The Volcani Center, PO Box 6, Bet Dagan 50250,
Israel (Phone: +972 3 9683388; Fax: +972 3 9605667; E-mail: vlaqua@volcani.agri.gov.il)
Key words: aquaculture genetics, chromosome-set manipulations, crossbreeding, DNA markers, hybridization,
selection, sex manipulation, transgenesis
Abstract
The aim of this review was to highlight the extent to which the genetic technologies are implemented by the
aquaculture industry. The review shows that some of the modern genetic technologies are already extensively
applied by the diverse aquaculture industries, though not to the same extent for all important aquacultured species
(according to FAO 1998 gures). Some species (common carp, Atlantic salmon, rainbow trout, channel catsh,
Nile tilapia, and the Pacic oyster) received concentrated breeding efforts, while other major cultured species
(Chinese and Indian carps and the giant tiger shrimp) received, so far, relatively limited attention, and a few species
(Yesso scallop, blue mussel, white Amur bream, and milksh) have, apparently, not been genetically improved at
all. Most of the genetically improved strains reaching the aquaculture industry were developed through traditional
selective breeding (selection, crossbreeding, and hybridization). Emerging, more modern technologies for genetic
manipulation seem to take 1020 years frombeing established experimentally until applications affect the industry.
Thus, chromosome-set and sex manipulations started to affect the industry during the 1980s and 1990s. DNA
marker technology and gene manipulations have yet hardly affected the industry. The former have not matured
yet, but hold much promise. The latter could have affected the industry already had it not been restricted by public
concern.
Introduction
Progress in genetic improvement of cultured sh and
shellsh has been reviewed or discussed on several
occasions during the last decade (e.g., Gjedrem, 1997;
Benzie, 1998b; Knibb, Gorshkova &Gorshkov, 1998).
The eld is regularly visited at the triennial Genetics
in Aquaculture symposia (Gall & Chen, 1993; Doyle
et al., 1996; McAndrew, Penman & Hulata, 1999;
Proceedings of Genetics in Aquaculture VII (Towns-
ville 2000), in preparation). It was recently the topic
of an Australian symposium on the application of ge-
netics to the aquaculture industry (see Lymberly et al.,
2000) and some key presentations therein (Gjedrem,
2000; Jones, McPhee & Ruscoe, 2000; Knibb, 2000;
Lawrence & Morrissy, 2000; Lymberly, 2000; Nell,
Smith & McPhee, 2000; Ward et al., 2000). The
status of aquaculture genetics at the turn of the mil-
lennium has also been recently reviewed by Dunham
et al. (2001). Thus, the aim of the current paper is not
to comprehensively review the eld again, but rather
to highlight the extent to which the genetic and ad-
vanced reproductive technologies are implemented by
the aquaculture industry.
For intraspecic genetic improvement aquacultur-
ists commonly use two major conventional breeding
methods: crossbreeding and selection. Both are fre-
quently integrated into comprehensive breeding pro-
grams. Interspecic hybridization is also used in some
aquaculture species. Crossbreeding has the advantage
of simplicity and the often immediate and visible im-
pact on animal performance. However, it is a static
procedure in the sense that no further genetic pro-
gress beyond that achieved in the rst generation can
be expected if crossbreeding is the only component
of the breeding program (Gjedrem, 1985). Selection
schemes, on the other hand, can realize higher gains
and progress at accelerated rates as long as additive
156
genetic variance exists for the traits of interest, as
has been shown for farm animals as well as Atlantic
salmon (Gjedrem, 1997). When a considerable com-
ponent of the genetic variance in the population is
non-additive, the breeding program should combine
both selection and crossbreeding (Gjedrem, 1985).
Until the early 1990s, a relatively small number of
aquacultural breeding programs existed worldwide.
However, it is hoped that improving and optimizing
the design of such programs (e.g., Bentsen & Gjerde,
1994; Gjerde, Gjen & Vilanueva, 1996; Gjerde &
Rye, 1997) will enable further gains in these as well
as in new breeding programs.
During the last decade or two, modern techno-
logies (including sex and chromosome-set manipula-
tions, gamete cryopreservation, transgenesis, genome
mapping and use of DNA markers) have matured from
the experimental phase and reached a stage where
they can be incorporated in aquacultural breeding pro-
grams. The extent to which each of those methods
and technologies are actually advancing the world
aquaculture industry is reviewed below. The review is
based on over 30 responses received to a questionnaire
distributed among ca. 200 aquaculture geneticists, as
well as search of databases covering the scientic lit-
erature. The data are presented in Table 1, which sum-
marizes the information known and available to me,
and does not necessarily list all cases of application
of genetic technologies in the worldwide aquaculture
industry.
Selection programs
As early as the 1920s, Embody and Hyford (1925)
selected surviving brook trout (Salvelinus fontinalis
Mitchill) from a population with endemic furuncul-
osis and increased the survival rate from 2% in the
initial population to 69% after three generations of
selection. Another historical report of a selection pro-
gram is the production of the Donaldson strain of
rainbow trout (Oncorhynchus mykiss Walbaum) (Don-
aldson & Olson, 1955). Donaldson began breeding
work with rainbow trout in 1932 and reported signic-
ant increase in growth and fecundity. After more than
35 years of directed individual selection, this strain
of rainbow trout was appreciated by trout culturists
in the USA and throughout the world and has been
widely distributed (Parsons, 1998). In spite of these
pioneering successes, very little selective breeding re-
search was conducted in aquacultured species prior to
1970.
Breeding work with common carp (Cyprinus car-
pio Linnaeus), traditionally carried out in European
closed populations, was based on phenotypic selec-
tion of the largest individuals with some consideration
of appearance (Wohlfarth, 1995). In the mid 1960s,
two large-scale directed selection efforts were initiated
in the common carp. Moav and Wohlfarths (1976)
mass selection program, initiated in 1965, did not
result in any improvement in growth rate. The reas-
ons for this might have been one, or a combination
of a few of the following reasons: the use of mass
selection, which is inefcient in estimating breeding
values for growth; the environmentally-based skewed
size distribution of common carp, which means that
largest individuals are not necessarily genetically su-
perior; the accumulation of high level of inbreeding,
which counteracted the selection performed; or the
occurrence of limited additive genetic variance in the
base population. Consequently, their efforts were di-
verted to the initiation of a crossbreeding program
(Hulata, 1995). At about the same time, a long-
term program for selecting common carp resistant to
dropsy, a serious infectious disease, for farming in
the cold zones of North Caucasus (Krasnodar) was
initiated (Kirpichnikov, 1993). The breeding program
consisted of selection within the local and the Siberian
wild carps from the river Amur, and crossing between
them. Mass selection for resistance to dropsy and for
high growth rate was carried out. The selection pro-
gram was successful and resistance to dropsy has been
improved. As a result of this successful breeding pro-
gram, three stocks of Krasnodar common carp were
developed, and heterotic crossbreds among them are
used for commercial production in that region and
other parts of the former Soviet Union (Kirpichnikov,
1993).
Several large scale selection experiments and
breeding programs, aiming at increasing growth rate,
were conducted more recently, resulting in 1020%
gain per generation, in channel catsh, Ictalurus
punctatus Ranesque (Smitherman & Dunham, 1985;
Dunham & Brummet, 1999), rainbow trout (Gjerde,
1986), Atlantic salmon, Salmo salar L. (Gjerde,
1986; Gjerde & Korsvoll, 1999; Gjedrem, 2000),
coho salmon, Oncorhynchus kisutch Walbaum (Her-
shberger et al., 1990; Myers et al., 2001), Nile
tilapia, Oreochromis niloticus L. (Eknath et al., 1993;
Bentsen et al., 1998; Basiao & Doyle, 1999; Lon-
galong, Eknath & Bentsen, 1999; Rye & Eknath,
1999), and other tilapias (Brzeski & Doyle, 1995).
Although the Norwegian trout and salmon, and the
157
Table 1. Aquacultured species (countries) to which different genetic manipulation technologies are applied in breeding programs or
practical culture operations
Technology Species involved in breeding programs Species involved in practical aquaculture
Selection Atlantic salmon, Salmo salar (Chile, Iceland, Atlantic salmon, Salmo salar (Chile, Iceland,
Ireland, Norway, Scotland); coho salmon, Ireland, Norway, Scotland); Pacic salmon,
Oncorhynchus kisutch (Chile, USA); rainbow Oncorhynchus sp. (Canada); coho salmon,
trout, Oncorhynchus mykiss (Chile, France, Oncorhynchus kisutch (Chile, USA); rainbow
Norway, USA); brown trout, Salmo trutta trout, Oncorhynchus mykiss (Czech Republic,
(France); channel catsh, Ictalurus punctatus France, Norway, USA,); silver bream
(USA); Gilthead sea bream, Sparus auratus and (Bangladesh, Thailand, Vietnam); Gilthead
Mediterranean sea bass, Dicentrarchus labrax sea bream, Sparus aurata and Mediterranean
(Israel); turbot, Scophthalmus maximus (Spain, sea bass, Dicentrarchus labrax (Israel);
Norway); rohu, Labeo rohita (India); Nile tilapia, common carp, Cyprinus carpio (China,
Oreochromis niloticus (China, Egypt, Indonesia, Czech Republic, Estonia, Indonesia, Russia,
Philippines Thailand); Oreochromis shiranus Vietnam); blunt snout brean, Megalobrama
(Malawi); common carp, Cyprinus carpio (China, amblycephala (China); crucian carp,
Czech Republic, Estonia, Indonesia, Russia, Carassius carassius (China); tench, Tinca
Vietnam); blunt snout brean, Megalobrama tinca (Czech Republic); Nile tilapia,
amblycephala (China); crucian carp, Carassius Oreochromis niloticus (China, Egypt,
carassius (China); tench, Tinca tinca (Czech Indonesia, Malaysia, Philippines, Vietnam);
Republic); Pacic oyster, Crassostrea gigas Pacic oyster, Crassostrea gigas (USA);
(Australia, USA); Sydney rock oyster, Saccostrea channel catsh, Ictalurus punctatus (USA);
glomerata (Australia); white Pacic shrimp, white Pacic shrimp, Litopenaeus vannamei
Litopenaeus vannamei (Brazil, Colombia, Mexico, (USA); blue shrimp, Penaeus stylirostris
USA); giant tiger shrimp, Penaeus monodon (Mexico); Kuruma shrimp, Penaeus japonicus,
(Australia, Thailand, USA); Kuruma parwn, brown tiger prawn, P. esculentus and banana
Penaeus japonicus (Australia) prawn, P. merguiensis (Australia)
Crossbreeding Gilthead sea bream (Israel); rainbow trout (Ireland); Common carp (China, Czech Republic, Estonia,
channel catsh (USA); common carp (China, Hungary, Israel, Russia, Thailand, Vietnam); Nile
Czech Republic, Hungary); Nile tilapia (Malaysia); tilapia (Malaysia); channel catsh (USA);
Pacic oyster (USA) Gilthead sea bream (Israel); rainbow trout (Ireland);
Pacic oyster (USA)
Hybridization Rainbow trout x Oncorhynchus clarkii Tilapia (China, Israel, Taiwan); channel and
(France); channel and blue catsh (USA); Asian x blue catsh (USA); Asian x African catshes,
African catshes, Clarias macrocephalus Clarias macrocephalus x C. gariepinus (Vietnam);
x C. gariepinus (Vietnam) hybrid striped bass (USA, Israel, Taiwan)
Sex manipulation Rainbow and brown trout (France)
a
; Nile tilapia (China, Fiji, Philippines,
channel catsh (USA); Nile tilapia (Philippines); Thailand, USA, Vietnam); Jordan tilapia (Israel);
silver barb (Thailand) Atlantic salmon (Canada) and coho salmon (Canada,
Japan); rainbow trout (France, Japan); amago and
masu salmon, ayu and hirame (Japan)
Gynogenesis/clonal Rainbow and brown trout (France)
a
; Rainbow and brown trout (France)
a
;
lines Atlantic, amago, coho and masu salmon (Japan); common carp (China, Japan); hirame (Japan)
ayu (Japan); common carp (China, Israel,
Japan); goldsh and crucian carp (Japan)
silver barb (Vietnam); red sea bream (Japan)
Androgenesis Rainbow trout (Japan, USA); amago salmon
(Japan); common carp (Japan, Netherlands)
158
Table 1. (continued)
Technology Species involved in breeding programs Species involved in practical aquaculture
Polyploidy Rainbow trout (France, Japan); brown trout Rainbow trout (Canada, France, Japan);
(France)
a
; Pacic oyster (USA); Catarina scallop, brown trout (France); Pacic oyster (Australia,
Argopecten ventricosus, lion paw scallop, USA, Japan); Catarina scallop, Argopecten
Nodipectenn subnodusus, red abalone, Haliotis rufescens ventricosus (Mexico); Atlantic salmon (Canada);
(Mexico); loach, Misgurnus anguillicauda (Japan) amago and masu salmon, ayu and hirame
(Japan); grass carp (USA)
Gene transfer Pacic salmon (USA, Canada, New Zealand
b
)
DNA markers Atlantic salmon (Norway); coho salmon (Chile) Pacic salmon (Canada)
Gamete/embryo Atlantic salmon (Canada, Norway, USA); Atlantic and Pacic salmon (Canada)
cryopreservation Pacic salmon (Canada); rainbow trout
(Norway); carps (China, Vietnam); Pacic oyster (USA)
a
Combination of gynogenesis and triploidy to produce all-female sterile trout.
b
Technology ready for application, but not implemented due to public concern and lack of marketing permit.
Summary of data extracted from responses to a questionnaire, therefore not necessarily a complete list.
GIFT Nile tilapia breeding programs have yet to be
comprehensively and critically described in full de-
tails, the products from these breeding programs are
commercially used by the industry in the respective
countries. An evaluation of the performance of the
GIFT Nile tilapia in comparison to local stocks has
been performed in several Asian countries, resulting
in variable degree of superiority of the selected sh
(Eknath et al., 1998). Several of these breeding pro-
grams, which started as experimental projects, have
been expanded to include the industry (e.g., Atlantic
salmon), or continue by private companies (e.g., At-
lantic salmon, rainbow trout, channel catsh, Nile
tilapia).
Domestication and selective breeding of the Kur-
uma shrimp Penaeus japonicus Bate in Australia com-
menced in 1993 (pilot research trials in collaboration
with a commercial farm). Following the success of
these trials (Hetzel et al., 2000), there has been a pro-
gressive increase in the use of selectively bred stocks.
Participating farms now stock up to 75% of their farm
ponds with genetically improved stocks. A commer-
cial prawn farming company pioneered commercial
scale selective breeding of the banana shrimp P. mer-
guiensis De Man in Australia. The use of domesticated
selectively bred stocks now accounts for the majority
of farmed P. merguiensis production in Australia. The
brown tiger prawn P. esculentus Haswell (endemic
to Australia) has also been successfully domesticated
and selectively bred stocks are now being used in
pilot-scale farm trials (Preston & Crocos, 2001, pers.
comm.).
The last decade has seen the initiation of many new
local and/or national breeding programs in a range of
countries and species, some in cooperation with the
industry or private companies. Among those are the
coho salmon breeding programs in Chile; Atlantic sal-
mon breeding programs in Chile, Iceland (Jnasson,
1996; Jnasson, Gjerde & Gjedrem, 1997), Ireland
and Scotland; rainbow trout in Chile, France and the
USA; brown trout (Salmo trutta L.) in France; Oreo-
chromis shiranus Boulenger in Malawi; multi-species
tilapia hybrid in Israel (Hulata et al., 1999); channel
catsh in the USA; Gilthead sea bream (Sparus aurata
L.) and Mediterranean sea bass (Dicentrarchus lab-
rax L.) in Israel; turbot (Scophthalmus maximus L.)
in Norway and Spain; rohu (Labeo rohita Hamilton)
in India; common carp in China, the Czech Repub-
lic, Estonia, Russia and Vietnam; blunt snout bream
(Megalobrama amblycephala Yih) in China (Li &
Yang, 1996; Li & Cai, 2000); tench (Tinca tinca L.)
in the Czech Republic; Pacic oyster (Crassostrea
gigas Thunberg) in Australia (Ward et al., 2000)
and Oregon, USA (Hedgecock et al., 1996); Sydney
rock oyster (Saccostrea glomerata Gould) in Australia
(Nell, Smith & McPhee, 2000); white Pacic shrimp
(Litopenaeus vannamei Boone) in Brazil, Colombia,
Mexico and the USA; Penaeus stylirostris in Mex-
ico; P. japonicus, P. esculentus Haswell and P. mer-
guiensis in Australia; and P. monodon Fabricius in
Australia, Thailand and the USA. The products from
these breeding programs will signicantly increase
the use of genetically improved stocks in commercial
aquaculture.
159
Crossbreeding and hybridization
Intraspecic crossbreeding
Intraspecic crossbreeding often increases growth rate
of aquacultured species, although specic heterotic
combinations need to be experimentally identied.
Only a few commercially aquacultured species have
been improved by crossbreeding. Variable propor-
tions of crossbreds showing heterosis for growth rate
have been obtained in the channel catsh; rainbow
trout; common carp and the Pacic oyster. Heterosis
was also found in survival, disease resistance and
reproductive traits.
Common carp crossbreds exhibiting heterosis are
the basis for the carp culture industries in Israel, Vi-
etnam, China and Hungary (Wohlfarth, 1993; Bakos
& Gorda, 1995; Hulata, 1995). Based on experimental
results, several crossbreds (two in Israel, three in Hun-
gary) were chosen for commercial application, whose
growth rate and other qualitative features gave about
20% better performance than the parental lines and
other control strains. The majority (80%) of Hun-
garian carp production was made of broodstocks of
parental lines developed for crossbreeding and dissem-
inated from Szarvas (Bakos & Gorda, 1995). Almost
all Israeli carp production was made of broodstocks
of parental lines developed for crossbreeding and dis-
seminated from Dor (Wohlfarth, 1993; Hulata, 1995).
In Indonesia, a breeding programbased on crossbreed-
ing among 10 strains of common carp, some of which
were developed using articial gynogenesis and sex-
reversal, is being carried out (Sumantadinata, 1995).
Signicant heterosis was observed in F
1
generations
of crossbreeds among local Vietnamese varieties of
common carp and foreign strains introduced from
Hungary, Ukraine, Indonesia, and Czech Republic.
Commercial crossbreeding between Vietnamese white
carp and Hungarian carps, and between Hungarian and
Indonesian yellow carp is being carried out (Thien &
Trong, 1995).
Crossbred channel catsh are widely cultured in
the southeastern states of the USA. Farmers realized
the superiority of crossbred catsh over pure strains
and highly appreciate it (N. Chatakondi, Gold Kist
Inc., personal communication).
Crossbreeding to improve yield has been exper-
imentally shown to be useful in the Pacic oyster
(Hedgecock, McGoldrick & Bayne, 1995; Hedge-
cock et al., 1996). The use of crossbreeding is being
extended to the commercial level in the west coast
of the USA, as a joint venture with private industry
(Hedgecock, 2001, pers. comm.).
Interspecic hybridization
Interspecic hybridization was successfully obtained
in many sh and shellsh genera and/or families
(carps, catshes, cichlids, moronids, salmonids, spar-
ids, sunshes, oysters, craysh, and others; e.g.,
Bakos, Krasznai & Marian, 1978; Chevassus, 1983;
Refstie, 1983; Hulata, 1995; Harrell, 1997; Lawrence
et al., 2000b; Dunham et al., 2001) as a means of
improving production traits (e.g., growth rate, sur-
vival, disease resistance) as well as to manipulate
sex ratios. Sometimes an interspecic hybrid does not
exhibit heterosis for any trait, but is still quite im-
portant for aquaculture application as it expresses a
good combination of benecial traits from both par-
ent species. Most of the hybrids evaluated did not
prove benecial for aquaculture. However, a few valu-
able hybrids are commercially exploited. The sunshine
bass, a hybrid between white bass (Morone chrysops
Ranesque) female and the striped bass (M. saxatilis
Walbaum) males grows faster and has better over-
all culture characteristics (good osmoregulation, high
thermal tolerance, resistance to stress and disease,
high survival in culture and modied water bodies,
and ability to utilize soy beans as a protein source)
than either parent species (Harrell, 1997, 1998). This
hybrid (and sometimes also the reciprocal, palmetto
bass hybrid) is commercially cultured mainly in the
USA (Van Olst & Carlberg, 1990), and recently also
in Israel and Taiwan.
Of the many hybrids among cyprinids tested,
most suitable for aquaculture were the two reciprocal
crosses between silver carp (Hypophthalmichthys mo-
litrix Valenciennes) and bighead carp (Aristichthys
nobilis Val.) which show higher survival and yield
than the parental species (Issa et al., 1986). This hy-
brid was commercially cultured in Israel during the
1970s to 1980s (and preferred over the pure silver
carp due to its tameness) before it was abandoned due
to the limited market for silver carp and species alike.
Another cyprinid hybrid of interest is the grass carp
(Ctenopharyngodon idella Val.) x bighead carp hy-
brid, originally produced in Hungary (Bakos, Krasznai
& Marian, 1978). The reported triploidy of this hybrid
(Marian & Krasznai, 1978, 1979; Beck, Biggers &
Dupree, 1980) attracted US sheries biologists look-
ing for an efcient herbivorous sh to control aquatic
vegetation (Sutton, Stanley & Miley, 1981; Shireman
160
& Smith, 1983) that is unable to naturalize in water
systems. The incomplete triploidy of the hybrid (Allen
& Stanley, 1983) has led to the development of large-
scale induction techniques for sterile, triploid grass
carp (Shelton, 1986, 1995).
The intergeneric hybrids between common carp
(Cyprinus carpio communis L.) and the Indian ma-
jor carps catla (Catla catla Ham.), rohu (Labeo rohita
Ham.) and mrigal (Cirrhinus mrigala Ham.) were also
found to be triploid and sterile (Reddy et al., 1990).
All three exhibit faster growth rate under monoculture,
possess higher esh content and show lower seine es-
capability than the maternal parent, but the former is
the most viable (Khan et al., 1990). The rohu x catla
hybrid grows almost as fast as pure catla, but has the
small head of the rohu and is therefore useful in In-
dian aquaculture. Catla x L. mbriatus Bloch hybrids
were reported to have small heads of mbriatus and
deep body and nearly equal growth rate to the catla.
Dressing percentage was also improved in this hybrid
(Basavaraju, Devaraj & Ayyar, 1995).
The hybrid between the channel catsh female and
the blue catsh (I. furcatus Val.) male exhibits het-
erosis for several traits, and is the only one of 28 cat-
sh hybrids evaluated exhibiting overdominance for
economic traits (Smitherman & Dunham, 1985). The
channel x blue hybrid has increased growth, growth
uniformity, disease resistance, tolerance of low oxy-
gen, dressing percentage and harvestability. Just one
producer sold over 3 million hybrid sh to commercial
catsh farmers during 19962000. Impressive produc-
tion performance in terms of growth, survival, feed
conversion and processing yield has been documented.
It is the best catsh to raise, however, technology
to produce it in commercial quantities is still a chal-
lenge due to mating blocks between the two species
(Chatakondi, 2000, pers. comm.).
The main catsh cultured in Thailand is a cross
between the African (Clarias gariepinus Burchell) and
the Thai (C. macrocephalus Gnther) catsh. This
cross combines fast growth rate of the African cat-
sh with the desirable esh characters of the Thai
catsh, although it does not grow as fast as the
pure African catsh. The intergeneric hybrid between
the two African catshes, Heterobranchus longilis
(Val.) x Clarias gariepinus also show culture potential
(Nwadukwe, 1995), though it is not clear whether this
has been realized yet.
Hybridization between tilapia species, to produce
all-male progeny, was considered as a promising
method to prevent uncontrolled reproduction, lead-
ing to over-population of culture ponds. Hybridiza-
tion between some species of tilapias such as Nile
tilapia and the blue tilapia, O. aureus Steindachner,
result in the production of predominantly male off-
spring. This hybrid combines well the advantageous
characteristics of both species, being more cold toler-
ant than O. niloticus and less borrowing in the mud
than O. aureus. It also has good salinity tolerance.
Other tilapia crosses producing predominately male
offspring include Nile tilapia x O. urolepis hornorum
Trewavas or O. macrochir Boulenger, and O. mos-
sambicus Peters x O. urolepis hornorum (Wohlfarth &
Hulata, 1983; Penman & McAndrew, 2000).
The most widespread culture of nearly all-male
hybrid tilapias is in Israel. The potential of tilapia
hybrids for culture is under exploited, due mainly to
management problems (Wohlfarth, 1994). The major
reason for this failure is the instability in produc-
tion of all-male hybrids. All-male progeny are only
produced from crossing pure species. Without care-
ful broodstock management the system breaks down,
and females start to appear in what was all-male hy-
brid before. It appears that this breakdown is largely
due to the inltration of parental broodstock by indi-
viduals of a different genotype, predominantly hybrids
between the two species involved, which are difcult
to distinguish from their parents (Wohlfarth, 1994).
Maintaining the purity of broodstock for hybridiza-
tion is a difcult and tedious task for a well-managed
tilapia hatchery, let alone ngerling production facil-
ities in the less developed countries. Hybridization of
tilapia has been largely suppressed by hormonal sex
inversion, and more recently by breeding sex-inversed
neo-males (see below), as alternative methods of pro-
ducing all-male tilapias. It should be noted, however,
that maintaining purity of broodstocks for the latter
suffers from exactly the same problems.
Female green sunsh (Lepomis cyanellus Rane-
sque) x male bluegill (L. macrochirus Ranesque)
hybrids also have improved overall performance for
aquaculture systems and can be an alternative for
aquaculture in regions of North America where the
growing season is unsuitable for production of tra-
ditional coldwater and warm-water species (Tidwell
et al., 1994; Webster, Tiu & Tidwell, 1997).
Salmonid hybrids do not express heterosis for
growth rate (Chevassus, 1979). Some diploid sal-
monid hybrids are potentially valuable because of
disease resistance inherited from the parent species
that is usually not cultured, but these hybrids have low
viability. The induction of triploidy can increase the
161
hatchability and viability of these potentially import-
ant hybrids (Parsons et al., 1986; Dorson, Chevassus
& Torhy, 1991). The rainbow trout (Oncorhynchus
mykiss) x char (Salvelinus spp.) triploid hybrids had
general resistance to several pathogenic salmonid vir-
uses (Dorson, Chevassus & Torhy, 1991), but these
triploid hybrids grew slower than diploids. Triploid
Pacic salmon hybrids can be seawater-acclimated
earlier than diploids (Seeb et al., 1993). Triploidiza-
tion of Atlantic salmon (Salmo salar) x brown trout (S.
trutta) hybrids increased their survival and growth rate
to a level comparable to Atlantic salmon (Galbreath
& Thorgaard, 1996). Nevertheless, hybrid salmonids
are not cultured commercially in signicant quantities.
The widely used triploidization (see below) in the sal-
mon industry may interfere with genetic improvement
programs for salmonids (Blanc & Vallee, 1999).
The recent nding that all-male progeny are ob-
tained from hybridization among Australian fresh-
water crayshes (Lawrence et al., 2000b) may soon
be implemented by craysh growers who currently
use manual sexing to establish all-male populations
(Lawrence et al., 2000a).
Chromosome-set and sex manipulations
Worldwide expansion of aquaculture requires spe-
cies or stocks more adapted to localized conditions
and less likely to affect genetic diversity of natural
populations in the event of escape. Sex control tech-
nologies (including direct endocrine treatments (sex
reversal) and indirect, combined endocrine and genetic
manipulations (gynogenesis, androgenesis, induced
polyploidization)
*
) to prevent uncontrolled reproduc-
tion, as well as to capitalize on differential culture
performances of the sexes, were recently reviewed
by Dunham (1990), Donaldson (1996), Pandian and
Koteeswaran (1998), Piferrer (2001) and Beardmore,
Mair and Lewis (2001). The indirect technique res-
ults in monosex gametes (usually sperm) that enable
*
Gynogenesis is a reproductive manipulation resulting in all-
maternal inheritance. Induction of gynogenesis involves egg activa-
tion by genetically inactivated homologous or heterologous sperm,
and diploidization by retention of the second polar body (meiotic
gynogenesis), or suppression of the rst mitotic cleavage (mitotic
gynogenesis). Androgenesis is a reproductive manipulation result-
ing in all-paternal inheritance, involving genetic inactivation of the
eggs genome and fertilization with haploid sperm (followed by dip-
loidization) or diploid sperm (Pandian & Koteeswaran, 1998). Poly-
loidy is the induction of individuals with extra (triploids, tetraploids)
sets of chromosomes (Thorgaard, 1986).
the production of monosex diploid or triploid popu-
lations. Monosex female salmonids are produced by
indirect feminization (Donaldson & Devlin, 1996)
while sterile (monosex female triploid) salmonids are
produced by indirect feminization followed by the
use of pressure or temperature shock to induce trip-
loidy (e.g., Johnstone, Mclay & Walsingham, 1991;
Johnstone, 1993). It should be noted that these tech-
nologies for production of chromosomally manipu-
lated sh/shellsh, are often used in combination,
or sequentially, for example, sex reversal of gyno-
genetic offspring to produce neomales or neofemales
for monosexing, or triploidization of gynogenetic fe-
males to produce XXX females (see examples below).
These technologies, which were intensively invest-
igated for several decades have reached commercial
application in many aquacultured species and coun-
tries. The most remarkable examples are mentioned
below.
Already in the mid 1980s, all-female stocks ac-
counted for over half of the production of UK trout
farms, and the use of sterile trout, particularly for sport
sheries and saltwater culture, was increasing (Bye
& Lincoln, 1986). Gynogenesis and triploidy to pro-
duce all-female sterile populations (Chourrout et al.,
1986; Chourrout & Nakamura, 1987; Quillet et al.,
1991) are commonly practiced in private trout farms
throughout France (Guyomard, 2000, pers. comm.)
and Japan (Arai, 2001). All-female and/or triploid sal-
mon are widely cultured in Canada (Benfey, 1996)
and on a smaller scale also in Japan (Arai, 2001).
These are produced either by direct triploidization in-
duction or by breeding tetraploid (XXXX) females
with diploid (XX) males. The reasons for the use of
all-female monosex seed in chinook salmon (Onco-
rhynchus tshawystscha Walbaum) and coho salmon
(O. kisutch) are to prevent losses arising from early
maturation of males, and because the roe has recently
become valuable and thus all-female populations are
desirable. For Atlantic salmon in British Columbia,
the reasons for monosex are for genetic containment
purposes, since they have been shown to breed with
Pacic salmon (Devlin, 2000, pers. comm.). Monosex
(all-female) strains of Oncorhynchus species are being
produced by gynogenesis and by the use of Y chro-
mosomal DNA markers (Devlin et al., 1991, 1994).
Triploidy is being explored by the Canadian salmon
industry for containment purposes. Triploidy is re-
quired by some Canadian Provinces for the culture
of non-native salmonid species, for example, rain-
bow trout in New Brunswick (Benfey, 1999, 2001).
162
However, the performance of the triploids has un-
fortunately been not as good as that of diploids, so
the industry is reluctant to adopt this technology un-
less they have to (Devlin, 2000, pers. comm.). In
Japan, culture of chromosomally manipulated sh
and shellsh is regulated by the governmental Fish-
eries Agency. Authorization has been granted for the
commercial culture of all-female diploid or triploid
salmonids (rainbow trout, coho, masu and yamame
salmons), ayu (Plecoglossus altivelis Temminck and
Schlegel) and oyster, as well as cloned hirame (Japan-
ese ounder, Paralichthys olivaceus Temminck and
Schlegel) (Yamamoto, 1999), in various prefectures
(Arai, 2001).
Triploidization seems to interfere with genetic im-
provement programs for salmonids. Genetic analyses
were carried out within two species (brown trout,
Salmo trutta and rainbow trout), by recording several
survival and growth traits in diploid and triploid pro-
genies from the same sires. The survival and growth
of triploid interspecic hybrids (O. mykiss female x S.
trutta male or Salvelinus fontinalis male) displayed a
special family variation hardly correlated with that of
maternal or paternal monospecic controls. Therefore,
the genetic improvement of such allo-triploids, as well
as other (diploid) hybrids, would require the evalu-
ation of breeders through progeny testing of the hybrid
offspring. This might be a serious restriction to the
development of interspecic hybridization in salmonid
farming (Blanc & Valle, 1999). Similarly, Bonnet et
al. (1999) indicate that due to interactions between
ploidy and family parameters in rainbow and brown
trout, selection of diploid male breeders for growth
performance may not be effective for their triploid
progeny.
Triploid grass carp production for control of nuis-
ance aquatic plants (Cassani, 1996) has been com-
mercialized (multi-million $ business) in the USA
(Shelton, 2001, pers. comm.). Early work on
monosexing as a means of preventing naturalization
(Shelton, 1986) was never applied because of the
development of large-scale induction techniques for
sterile triploids (Shelton, 1995). Combined techno-
logy of gynogenesis and hormonal sex-reversal is be-
ing developed for the production of all-female XXX-
triploid grass carp and black carp (Mylopharyngodon
piceus Richardson) that are completely sterile, unlike
the XXY- or XYY-triploids that possess fragments of
testes, and are thus able to produce some active sperm
(Rothbard et al., 1997). Their potential has yet to be
realized.
YY male genotypes of Nile tilapia proved to be
as viable and fertile as normal XY males, and to sire
progeny that are nearly 100% males (Mair, 2000, pers.
comm.). Their all-male progeny, XY, known as ge-
netically male tilapia (GMT), are mass produced on a
commercial scale, providing a reliable solution to the
problem of early sexual maturation and uncontrolled
reproduction leading to overpopulation in Nile tilapia
culture (Mair et al., 1995, 1997; Tuan, Little & Mair,
1998, 1999; Abucay et al., 1999; Beardmore, Mair &
Lewis, 2001). Results from on-station trials indicate
that GMT have considerable benets under culture,
signicantly increasing yields compared to mixed sex
tilapia of the same strain or sex reversed male tilapia
(Mair et al., 1995). The YY male line has been further
selected for growth rate and for combining ability for
GMT sex ratio, using intensive within-family selection
in a synthetic female line (Abucay &Mair, 2000). Cul-
ture of all-male Nile tilapia, based on the YY-GMT has
been widely practiced in the Philippines since 1995
and in Thailand since 1997, and to a lesser extent in
a number of other countries including China, Fiji, Vi-
etnam, Central American countries, and USA (Mair,
2000, pers. comm.). A similar approach was also ap-
plied to O. aureus. By breeding sex-inverted O. aureus
males (which became ZZ neo-females) with normal
(ZZ) males, all-male progeny are produced (Lahav,
1993; Rosenstein & Hulata, 1994). All-female com-
mon carp populations (Cherfas et al., 1996) have been
established in Israel, by sex reversing XX gynogenetic
females to males (Gomelsky et al., 1994), and using
these XX males for breeding. All-female seed released
to commercial farms resulted in 1015% yield im-
provement over existing commercial stocks. However,
dissemination was halted due to a severe viral disease
outbreak affecting the Israeli carp industry a few years
ago.
Polyploid technology has been commercially ap-
plied in the Pacic oyster culture industry on the US
West Coast since the mid-1980s, based on earlier work
summarized by Allen, Downing and Chew (1989).
The current push is making tetraploids, in order to
make triploids by a natural 4N x 2N cross (Guo,
DeBrosse & Allen, 1996; Ward et al., 2000), as
already done in nsh (Arai, 2001). One US com-
pany is already doing this. The research is beginning
to put the crossbreeding technology together with the
polyploid technology, but this is still experimental
(Hedgecock, 2000, pers. comm.).
Recent and on-going research will likely lead to
commercial application of chromosome-set manipu-
163
lations for monosexing (coupled with sex reversal)
or polyploidy to more species, including the Medi-
terranean sea bass, Dicentrarchus labrax (Colombo
et al., 1995; Chatain, 1998; Felip et al., 1998,
1999); European sea bream, Sparus aurata (Gor-
shkov et al., 1998); red sea bream, Pagrus major
Temminck and Schlegel (Arakawa, 1999); hirame,
Paralichthys olivaceus (Tabata & Mizuta, 1997;
Yamamoto, 1999); Thai silver barb, Puntius go-
nionotus (Bleeker) (Pongthana et al., 1999; Koedprang
& Na-Nakorn, 2000); African catsh, Clarias gar-
iepinus (Volckaert et al., 1997; Vardi et al., 1999;
Galbusera, Valckaert & Ollevier, 2000); Asian cat-
sh, Clarias macrocephalus (Na-Nakorn, Rangsin
& Witchasunkul, 1993); channel catsh, Ictalurus
punctatus (Goudie et al., 1995); black carp, Mylopha-
ryngodon piceus (Rothbard et al., 1997); paddlesh,
Polyodon spathula Walbaum (Mims et al., 1997); hy-
brid Lepomis (Wills, Paret & Sheehan, 1994); brook
trout, Salvelinus fontinalis (Galbreath & Samples,
2000); Atlatic halibut, Hippoglossus hippoglossus L.
(Hendry, Martin-Robichaud & Benfey, 1999); and
Manila clam, Ruditapes philippinarium Adams and
Reeve (Utting, Millican & Laing, 1996).
Gamete cryopreservation
The technology for gamete cryopreservation of
aquatic species (nsh as well as shellsh) has pro-
gressed signicantly during the last two decades, and
reviewed recently by Diwan (2000), Gwo (2000),
Lahnsteiner (2000), Suquet et al. (2000), Tiersch
and Mazik (2000), and Chao and Liao (2001). For
mollusks, the technology enables also cryopreserva-
tion of late embryos and early larvae (Chao et al.,
1997; Lin & Caho, 2000; Paniagura-Chavez et al.,
2000). Protocols developed experimentally have been
tested under production conditions (e.g., Fauvel et al.,
1998) and evaluated economically (Caffey & Tier-
sch, 2000a,b). While not a strictly genetic technology,
gamete cryopreservation has a potential role in genetic
conservation of wild resources and endangered wild
or cultured stocks (e.g., McAndrew, Rana & Penman,
1993; Pullin et al., 1998; Gwo et al., 1999; Harvey,
2000), and can be incorporated into breeding pro-
grams for storage of milt of specic genotypes (e.g.,
for monosexing, as discussed above) or for preserva-
tion and dissemination of improved breeds (Van der
Lende, Komen & Brascamp, 1998). Retrieval of the
whole genome of endangered or extinct stock from
cryopreserved milt can be done through androgenesis
(e.g., Bercsenyi et al., 1998; Rothbard et al., 1999).
Efcient use of the technology for all these applica-
tions require the establishment of sperm (cryo)banks
(Cross & Rogan, 1992; Lubzens et al., 1997), which
already exist for some species (e.g., salmonids Cloud
et al., 2000). Assistance is provided to aquaculture
producers wishing to cryopreserve sperm from special
salmon strains (Devlin, 2001, pers. comm.).
The aquaculture industry in general, and those of
us running large-scale breeding programs in partic-
ular, seem to underestimate the genetic potential of
establishing such spermbanks. Compared with cryo-
preservation of dairy bull and other livestock sperm
(Johnson, 2000) which is a mature, commercialized
technology, commercialization of sh sperm conser-
vation is somewhere between conception and infancy
of the technology (Caffey & Tiersch, 2000a). In-
creased commercialization is expected to occur when
research protocols for sh sperm cryopreservation are
applied in the private sector and markets for cryo-
preserved sperm are established (Caffey & Tiersch,
2000b).
Transgenesis
The rst successful gene transfer into sh was reported
fromChina in the mid 1980s. Since then, gene transfer
has been accomplished in many aquacultured species
and countries, in most cases involving the growth hor-
mone gene. Signicant breakthrough occurred when
promoters (Moav et al., 1992) and sh gene sequences
(Houdebine & Chourrout, 1991; Devlin et al., 1994;
Devlin et al., 1995a; Donaldson & Devlin, 1996) be-
came available, since mammalian genes used in the
early works were poorly expressed. The technology is
still being experimentally rened, and applied to more
species (e.g., Rahman, Iyengar & Maclean, 1997;
Venugopal et al., 1997; Rahman et al., 1998; Hin-
its & Moav, 1999; Kang et al., 1999; Rahman &
Maclean, 1999; Li & Tsai, 2000; Sin et al., 2000; Tsai,
2000). The aquacultured species transformed include
Atlantic salmon (Salmo salar), rainbow trout (On-
corhynchus mykiss), tilapia (Oreochromis niloticus),
carp (Cyprinus carpio), channel catsh (Ictalurus
punctatus), African catsh (Clarias gariepinus) and
northern pike (Esox lucius L.). In addition to these
commercially orientated species, certain small model
species have also been successfully exploited gold-
sh (Carassius auratus L.), loach (Misgurnus fossilis
164
L.), medaka (Oryzias latipes Temminck and Schlegel)
and zebrash (Danio rerio Hamilton) (see Maclean
(1998) for a recent review).
The scientic progress achieved in applying bio-
technological manipulations to aquacultured species,
and indeed to other edible crops and livestock, res-
ulted in the production of genetically modied or-
ganisms (GMOs). The scientic community did not
neglect concerns about their possible escape to the
environment if held outside experimental laborator-
ies, as well as the safety of transgenic sh as food,
and the possible unexpected consequences of gene
transfer (pleiotropic effects). Unanticipated effects of
gene insertion are likely to be reected (positively
or negatively) in the shape, function or health of the
transgenic shes, as was indeed observed (Chatakondi
et al., 1994, 1995; Devlin et al., 1995b; Ostenfeld,
Devlin & McLean, 1998; Stevens & Devlin, 2000a,b).
The practical precautions which must be observed if
this novel technology is to safely complement conven-
tional sh production strategies were repeatedly dis-
cussed (e.g., Kapuscinski & Hallerman, 1990, 1991;
McEvoy, Gannon & Sreenan, 1992; Bartley & Haller-
man, 1995; Kapuscinski & Hallerman, 1995; Hew &
Fletcher, 1997; Knibb, 1997; Devlin, 1998; Volck-
aert & Ollevier, 1998; Maclean & Laight, 2000; Hew
& Fletcher, 2001). Berkowitz and Kryspin-Sorensen
(1994) concluded that food safety evaluation of trans-
genic shes has requirements similar to evaluations of
the safety of added substances such as chemicals and
drugs used orally or by injection.
After about a little over a decade of very intens-
ive research, the technology has reached the stage of
commercial application (Entis, 1997). Unlike other
genetic applications in aquaculture, which are taken
up by a wide range of producers, this one is mainly
led by large, corporate operations (e.g., in the USA,
Canada, New Zealand) that have already produced
fast-growing transgenic Atlantic and Pacic salmon,
and will soon have similarly produced trout, tilapia
and ounder. However, there is a growing public con-
cern about the potential hazards of raising such sh,
as exemplied by articles in Times (Thompson, 2000)
and the New York Times (Yoon, 2000). This has led
all producer groups in Canada to clearly indicate they
do not want to utilize transgenic sh, and they do not
support the research in any way. Consumer attitudes
are such that it is difcult to market transgenic sal-
mon, at least in the near future (Devlin, 2000, pers.
comm.). Applications have also been led with the
FDA for marketing in the USA, but no permits have
been granted yet. Fearing of the consumer backlash, a
major New Zealand producer of Chinook salmon de-
cided to suspend their gene- modication experiments
(Thompson, 2000), and the EC is maintaining a sim-
ilar policy. The Canadian research group is conducting
research only on risk assessment for the environment,
in case the public and producers do change their mind
in the future, and to compare the technology to tra-
ditional approaches such as selection (Devlin, 2000,
pers. comm.). While Entis (1997) believes that under-
standing and acceptance will grow gradually, and the
benets will become clear to most only in hindsight,
it remains to be seen if the public will accept such
products on the global marketplace any time soon.
DNA markers, gene mapping, QTLs and
marker-assisted selection
DNA markers replaced allozymes (Utter, 1991) as
the major analytical tool in genetic research in the
mid 1980s, following Jeffreys, Wilson and Theins
(1985) nding of the usefulness of the hypervariab-
ility of the genome at the DNA level. Aquaculture
and sheries geneticists quickly adopted them as well
(e.g., Wright, 1990; Ward & Grewe, 1995; Wright
& Bentzen, 1995). Genomic research and gene map-
ping of aquatic organisms developed fast during the
late 1990s. Novel genetic technologies involving the
use of DNA-based tools are under development for a
range of aquaculture species and applications, includ-
ing species/strain identication and pedigree tracing
(e.g., Heath et al., 1995; Thongpan et al., 1997; Perez-
Enriquez, Takagi & Taniguchi, 1999; Tan et al., 1999;
Norris, Bradley & Cunningham, 2000), conrmation
of isogenicity in products of chromosome-set manip-
ulations and clones (e.g., Young et al., 1996; Jen-
neckens et al., 1999), linkage mapping (e.g., Alcivar-
Warren et al., 1997; Kocher et al., 1998; Young et al.,
1998; Liu et al., 1999a,b; Agresti et al., 2000; Li
et al., 2000; McConnell et al., 2000; Sakamoto et al.,
2000; Nichols et al., 2001), identication of QTLs
(e.g., Jackson et al., 1998; Danzmann, Jackson &
Ferguson, 1999; Moore et al., 1999; Palti, Parsons
& Thorgaard 1999; Sakamoto et al., 1999), and
other uses in breeding programs (Davis & Hetzel,
2000). Knowledge of linkages between marker and
QTL alleles may be used for marker-assisted selec-
tion (MAS), increasing the rate of genetic progress
above that for selective breeding alone (Poompuang
& Hallerman, 1997). While holding great promise for
1
6
5
Table 2. Genetic manipulation technologies that are applied in breeding programs (B) or practical culture (P) of the worlds 19 principal aquacultured species (FAO, 1998). Also marked
are cases of experimental work (E) when not yet B or P
Species World Selection Cross- Hybridi- Sex Gyno- Andro- Ployploidy Gene MAS Gamete
production breeding zation manipulation genesis genesis transfer or cryopreser-
(MT) QTL vation
Pacic oyster 3,438,824 B, P B, P B, P E B
Silver carp 3,308,419
a
E
b
E B
Grass carp 2,894,017
a
E E B, P
c
E B
Common carp 2,465,283 B, P B, P B, P B, P E, B E E E B
Bighead carp 1,584,289
a
E
b
B
Manila clam 1,426,501 B E E
Crucian carp 1,036,164 B, P E E E, B B, P E E
Yesso scallop 856,217
Nile tilapia 793,931 B, P B, P P B, P E E E
Rohu 754,677 B E E E
Atlantic salmon 687,906 B, P E B, P E, B B, P E
d
E, B B
Catla 628,757 E
Giant tiger shrimp 577,990 B
Mrigal 560,556 E E
Blue mussel M. edulis 499,891
e
E
White Amur bream 449,282
Rainbow trout 438,013 B, P B, P B B, P B, P E, B B, P E B
Milksh 369,003
Channel catsh 256,129 B, P B, P B, P B B E E E
a
Several Chinese population genetics studies (e.g., Wu & Wang, 1997a,b; Li, Lu & Bernatchez, 1998; Yang & Gui, 1999), but no record found for any breeding program.
b
The hybrid between these two species was commercially cultured in Israel in the 1970 1980s. Experimentally cultured in Vietnam.
c
In the USA. In China, where the bulk of production occurs, it is still experimental (Li, 2001).
d
Technology ready for application for Pacic salmon, but not implemented due to public concern and lack of marketing permit.
e
Numerous population genetics studies (e.g., Hilbish, Bayne & Day, 1994; Tremblay, Myrand & Sevigny, 1998; Daguin & Borsa, 1999; Penny & Hart, 1999), but no record found for any
breeding program.
166
advancing breeding programs, these genomic meth-
ods had, so far, only marginal commercial application.
The notable exception is the identication of YY
genotypes in salmonids using a DNA probe (Devlin
et al., 1991, 1994), as mentioned previously under the
monosexing section.
Summary and conclusions
This review shows that while we still have a long
way to go in applying the existing scientic informa-
tion and modern genetic technologies into the diverse
aquaculture industries, some of them have already
found their application. However, this is not true for all
important aquacultured species, as can be seen from
Table 2. While some species (common carp, Atlantic
salmon, rainbow trout, channel catsh, Nile tilapia,
and the Pacic oyster) received concentrated breeding
efforts, other major cultured species (Chinese and In-
dian carps and the giant tiger shrimp) received, so far,
relatively limited attention, and a few species (Yesso
scallop, blue mussel, white Amur bream, and milksh)
have, apparently, not been genetically improved at all.
The number of breeding programs, both in private
sector and national levels, is increasing, and the
use of stocks genetically improved through selective
breeding, monosexing and polyploidy manipulations
is gaining popularity. This trend seems to be stronger
for nsh, and weaker for crustaceans (Benzie, 1998a)
and mollusks. For the latter, however, commercial
applications become evident, not only for the Pa-
cic oyster (the number one farmed aquatic species
in the world, according to FAOs latest statistics),
for which Allens (1998) assessment was that pro-
gress in commercial shellsh improvement programs
are rudimentary, but for other molluscan species as
well. Most of the genetically improved strains reach-
ing the aquaculture industry were developed through
traditional selective breeding (selection, crossbreeding
and hybridization). Emerging, more modern techno-
logies for genetic manipulation seem to take 1020
years from being established experimentally until ap-
plications affect the industry. Chromosome-set and
sex manipulations, which were intensively investig-
ated in the 1970s and 1980s in several species, and
are still being rened for others, started to affect
the industry during the 1980s. Molecular techniques
(DNA markers and gene manipulations), intensively
investigated since the mid 1980s, reached rst applic-
ation in the mid 1990s. In spite of attracting huge
amounts of research funds, DNA marker technology
has yet hardly affected the industry. This is also true
for transgenic sh, which could have affected the in-
dustry already had it not been restricted by public
concern.
Acknowledgements
The author thanks the following colleagues who
kindly supplied information by responding to a ques-
tionnaire: C. Araneda (Chile), B. Argue (USA),
T. Benfey (Canada), H.B. Bentsen (Norway), N.
Chatakondi (USA), P. Crocos (Australia), T. Cross
(Ireland), R.H. Devlin (Canada), P.F. Galbreath
(USA), G.A.E. Gall (USA), A.J. Gharrett (USA), B.
Gjerde (Norway), B. Gomelsky (USA), S. Gorshkov
(Israel), R. Gross (Estonia), R. Guyomard (France),
A. Hardjamulia (Indonesia), D. Hedgecock (USA),
G. Hrstgen-Schwark/A. Mller-Belecke (Germany),
A.M. Ibarra (Mexico), S. Li (China), O. Linhart
(Czech Republic), E. Lubzens (Israel), G.C. Mair
(Thailand), S.D. Mims (USA), N. Preston (Australia),
W.L. Shelton (USA), J. Thodesen (ICLARM, Malay-
sia), P.A. Tuan (Vietnam), F. Volckaert (Belgium), R.
Ward (Australia), and W.R. Wolters (USA). Thanks
are also due to Elisha Gootwine (ARO, Israel) and Ro-
ger Doyle (Canada) for their comments on this review,
and for useful discussions.
References
Abucay, J.S. & G.C. Mair, 2000. Divergent selection for growth
in the development of a female line for the production of im-
proved genetically male tilapia (GMT) in Oreochromis niloticus
L., pp. 7172 in Tilapia Aquaculture, Proc. of the Fifth Inter-
national Symposium on Tilapia in Aquaculture, edited by K.
Fitzsimmons & J.C. Filho. Rio De Janeiro, Brazil.
Abucay, J.S., G.C. Mair, D.O.F. Skibinski & J.A. Beardmore, 1999.
Environmental sex determination: the effect of temperature and
salinity on sex ratio in Oreochromis niloticus L. Aquaculture
173: 219234.
Agresti, J.J., S. Seki, A. Cnaani, S. Poompuang, E.M. Hallerman,
N. Umiel, G. Hulata, G.A.E. Gall & B. May, 2000. Breeding new
strains of tilapia: development of an articial center of origin and
linkage map based on AFLP and microsatellite loci. Aquaculture
185: 4356.
Alcivar-Warren, A., R.A. Dunham, P. Gaffney, T.D. Kocher & G.H.
Thorgaard, 1997. First aquaculture species genome mapping
workshop. Anim. Gene. 28: 451452.
Allen, S.K., Jr., 1998. Commercial application of bivalve genetics:
not a solo effort. World Aquacult. 29: 3843.
Allen, S.K., Jr. & J.G. Stanley, 1983. Triploid grass carp: status and
management implications. Fisheries 12(4): 2024.
167
Allen, S.K., Jr., S.L. Downing & K.K. Chew, 1989. Hatchery
Manual for Producing Triploid Oysters. University of Washing-
ton Press, Seattle, WA, USA.
Arakawa, T., 1999. Present state of red sea bream Pagrus major cul-
ture and breeding experiments in Nagasaki prefecture, in Goals
and Strategies for Breeding in Fisheries, edited by M. Kato, K.
Fukusho, K. Ikuta, J.P. McVey, P.K. Park & B.J. Keller. Proceed-
ings of the 27th UJNR Aquaculture Panel Symposium, Ise, Mie,
Japan, 1112 November, 1998. Bull. Nati. Res. Inst. Aquacult.
Suppl. 1: 5561.
Arai, K., 2001. Genetic improvement of aquaculture nsh species
by chromosome manipulation techniques in Japan. Aquaculture
197: 205228.
Bakos, J. & S. Gorda, 1995. Genetic improvement of common carp
strains using intraspecic hybridization. Aquaculture 129: 183
186.
Bakos, J., Z. Krasznai & T. Marian, 1978. Cross-breeding ex-
periments with carp, tench and Asian phytophagous cyprinids.
Aquacult. Hungarica 1: 5157.
Bartley, D.M. & E.M. Hallerman, 1995. A global perspective on the
utilization of genetically modied organisms in aquaculture and
sheries. Aquaculture 137: 17.
Basavaraju, Y., K.V. Devaraj & S.P. Ayyar, 1995. Comparative
growth of reciprocal carp hybrids between Catla catla and Labeo
mbriatus. Aquaculture 129: 187191.
Basiao, Z.U. & R.W. Doyle, 1999. Test of size specic mass selec-
tion for Nile tilapia, Oreochromis niloticus L., cage farming in
the Philippines. Aquacult. Res. 10: 373378.
Beardmore, J.A., G.C. Mair & R.I. Lewis, 2001. Monosex male
production in nsh as exemplied by tilapia: applications,
problems, and prospects. Aquaculture 197: 283301.
Beck, M.L., C.J. Biggers & H.K. Dupree, 1980. Karyological ana-
lysis of Ctenopharyngodon idella, Aristichthys nobilis and their
F
1
hybrid. Transac. Am. Fish. Soc. 109: 433438.
Benfey, T.J., 1996. Use of all-female and triploid salmonids for
aquaculture in Canada. Bull. Aquacult. Assoc. Can. 1996 (2):
68.
Benfey, T.J., 1999. Use of triploid Atlantic salmon (Salmo salar) for
aquaculture. Can. Dept. Fish. Oceans Res. Doc. 98/166: 11.
Benfey, T.J., 2001. Use of sterile triploid Atlantic salmon (Salmo
salar L.) for aquaculture in New Brunswick. ICES J. Marine Sci.
(in press).
Bentsen, H.B. & B. Gjerde, 1994. Designs of sh breeding pro-
grams, pp. 353359 in Proceedings of the 5th World Congress
on Genetics Applied to Livestock Production, Vol. 19, Guelph,
Canada.
Bentsen, H.B., A.E. Eknath, M.S. Palada-de Vera, J.C. Danting,
H.L. Bolivar, R.A. Reyes, E.E. Dionisio, F.M. Longalong, A.V.
Circa, M.M. Tayamen & B. Gjerde, 1998. Genetic improvement
of farmed tilapias: growth performance in a complete diallel
cross experiment with eight strains of Oreochromis niloticus.
Aquaculture 160: 145173.
Benzie, J.A.H., 1998a. Genetic improvement of prawns, pp. 103
110 in Proceedings of the 6th World Congress on Genetics
Applied to Livestock Production, Vol. 27, Armidale, Australia.
Benzie, J.A.H., 1998b. Penaeid genetics and biotechnology. Aqu-
aculture 164: 2347.
Bercsenyi, M., I. Magyary, B. Urbanyi, L. Orban & L. Horvath,
1998. Hatching out goldsh from common carp eggs: interspe-
cic androgenesis between two cyprinid species. Genome 41:
573579.
Berkowitz, D.B. & I. Kryspin-Sorensen, 1994. Transgenic sh:
safe to eat? A look at the safety considerations regarding food
transgenics. BioTechnology 12: 247252.
Blanc, J.M. & F. Valle, 1999. Genetic variability of farming per-
formances in some salmonid species and hybrids modied by
triploidization. Cybium 23(Suppl. 1): 7788.
Bonnet, S., P. Haffray, J.M. Blanc, F. Valle, C. Vauchez, A. Faur
& B. Fauconneau, 1999. Genetic variation in growth parameters
until commercial size in diploid and triploid freshwater rainbow
trout (Oncorhynchus mykiss) and seawater brown trout (Salmo
trutta). Aquaculture 173: 359375.
Brzeski, V.J. & R.W. Doyle, 1995. A test of an on-farm selection
procedure for tilapia growth in Indonesia. Aquaculture 137: 219
230.
Bye, V.J. & R.F. Lincoln, 1986. Commercial methods for the con-
trol of sexual maturation in rainbow trout (Salmo gairdneri).
Aquaculture 57: 299309.
Caffey, R.H. & T.R. Tiersch, 2000a. Economics and marketing of
cryopreserved sh sperm, pp. 388408 in Cryopreservation in
Aquatic Species, edited by T.R. Tiersch & P.M. Mazik. World
Aquaculture Society, Baton Rouge, Louisiana, USA.
Caffey, R.H. & T.R. Tiersch, 2000b. Cost analysis for integrat-
ing cryopreservation into an existing sh hatchery. J. World
Aquacult. Soc. 31: 5158.
Cassani, J.R. (ed.), 1996. Managing Aquatic Vegetation With
Grass Carp: A Guide for Water Resource Managers. American
Fisheries Society, Bethesda, MD, USA.
Chao, N.-H. & I.C. Liao, 2001. Cryopreservation of nsh and
shellsh gametes and embryos. Aquaculture 197: 161189.
Chao, N.-H., T.T. Lin, Y.J. Chen, H.W. Hsu & I.C. Liao, 1997.
Cryopreservation of late embryos and early larvae in the oyster
and hard clam. Aquaculture 155: 3144.
Chatain, B., 1998. Triploidie et gynogenese chez le bar: un moyen
dobtenir des populations monosexes femelles steriles. Piscicult.
Francaise 134: 1116.
Chatakondi, N., A.C. Ramboux, A. Nichols, M. Hayat, P.L. Duncan,
T.T. Chen, D.A. Powers & R.A. Dunham, 1994. The effect of
rainbow trout growth hormone gene on the morphology, dressing
percentage and condition factor in the common carp, Cyprinus
carpio, pp. 481484 in Proceedings of the 5th Congress on
Genetics Applied to Livestock Production, Vol. 17.
Chatakondi, M., R. Lovell, P. Duncan, M. Hayat, T.T. Chen, D.A.
Powers, T. Weete, K. Cummins & R.A. Dunham, 1995. Body
composition of transgenic common carp, Cyprinus carpio, con-
training rainbow trout growth hormone gene. Aquaculture 138:
99109.
Chen, T.T., J.-K. Lu & F. Richard, 1998. Transgenic sh technology
and its application in sh production, pp. 527547 in Agricul-
tural Biotechnology, edited by A. Altman. Marcel Dekker, N.Y.,
USA.
Cherfas, N.B., B. Gomelsky, N. Ben-Dom, D. Joseph, S. Co-
hen, I. Israel, M. Kabessa, G. Zohar, Y. Peretz, D. Mires &
G. Hulata, 1996. Assessment of all-female common carp pro-
genies for sh culture. Israeli J. Aquacult. Bamidgeh 48: 149
157.
Chevassus, B., 1979. Hybridization in salmonids: results and per-
spectives. Aquaculture 17: 113128.
Chevassus, B., 1983. Hybridization in sh. Aquaculture 33: 245
262.
Chourrout, D. & I. Nakayama, 1997. Chromosome studies of pro-
genies of tetraploid female rainbow trout. Theoret. Appl. Genet.
74: 687692.
Chourrout, D., B. Chevassus, F. Krieg, A. Happe, G. Burger &
P. Renard, 1986. Production of second generation triploid and
tetraploid rainbow trout by mating tetraploid males and diploid
females. Potentials of tetraploid sh. Theoret. Appl. Genet. 72:
193206.
168
Cloud, J.G., R. Armstrong, P. Wheeler, P.A. Kucera & G.H. Thor-
gaard, 2000. The Northwest salmonid germplasm repository,
pp. 338342 in Cryopreservation in Aquatic Species, edited by
T.R. Tiersch & P.M. Mazik. World Aquaculture Society, Baton
Rouge, Louisiana, USA.
Colombo, L., A. Barbaro, A. Libertini, P. Benedetti, A. Francescon
& I. Lombardo, 1995. Articial fertilization and induction
of triploidy and meiogynogenesis in the European sea bass,
Dicentrarchus labrax L. J. Appl. Ichthyol. 11: 118125.
Cross, T.F. & E. Rogan, 1992. The feasibility of developing and
utilising gene banks for sea trout (Salmo trutta) conservation.
Fisheries Technical Report, No. 4. National Rivers Authority,
Bristol, UK, 33 p.
Daguin, C. & P. Borsa, 1999. Genetic characterisation of Mytilus
galloprovincialis Lmk. in North West Africa using nuclear DNA
markers. J. Exp. Marine Biol. Ecol. 235: 5565.
Danzmann, R.G., T.R. Jackson &M.M. Ferguson, 1999. Epistasis in
allelic expression at upper temperature tolerance QTL in rainbow
trout. Aquaculture 173: 4558.
Davis, G.P. & D.J.S. Hetzel, 2000. Integrating molecular genetic
technology with traditional approaches for genetic improvement
in aquaculture species. Aquacult. Res. 31: 310.
Devlin, R.H., 1998. Production and evaluation of transgenic sh for
aquaculture. Austral. Biotechnol. 8: 222227.
Devlin, R.H., B.K. McNeil, T.D.D. Groves & E.M. Donald-
son, 1991. Isolation of Y-chromosomal DNA probe cap-
able of determining genetic sex in chinook salmon (Onco-
rhynchus tshawytscha). Can. J. Fish. Aqua. Sci. 48: 1606
1612.
Devlin, R.H., B.K. McNeil, I.I. Solar & E.M. Donaldson, 1994.
A rapid PCR-based test for Y-chromosomal DNA allows simple
production of all-female strains of chinook salmon. Aquaculture
128: 211220.
Devlin, R.H., T.Y. Yesaki, C.A. Biagi, E.M. Donaldson, P. Swanson
& W.-K. Chan, 1994. Extraordinary salmon growth. Nature 371:
209210.
Devlin, R.H., T.Y. Yesaki, E.M. Donaldson, S.J. S.J. Du & C.L.
Hew, 1995a. Production of germline transgenic Pacic salmonids
with dramatically increased growth performance. Canad. J. Fish.
Aqua. Sci. 52: 13761384.
Devlin, R.H., T.Y. Yesaki, E.M. Donaldson & C.L. Hew, 1995b.
Transmission and phenotypic effects of an antifreeze/GH gene
construct in coho salmon (Oncorhynchus kisutch). Aquaculture
137: 161169.
Diwan, A.D., 2000. Cryopreservation of crustacean gametes
and embryos: potential tool in biotechnological applications,
pp. 249271 in Recent Advances in Marine Biotechnology;
Aquaculture. Part A: Seaweed and invertebrates, Vol. 4, edited
by M. Fingerman & R. Nagabhushanam. Science Publishers,
Eneld, USA.
Donaldson, E.M., 1996. Manipulation of reproduction in farmed
sh. Anim. Reprod. Sci. 42: 381392.
Donaldson, E.M. & R.H. Devlin, 1996. Uses of biotechnology to
enhance production, pp. 9691020 in Principles of Salmonid
Culture. Developments in Aquaculture and Fisheries Science,
Vol. 29, edited by W. Pennell & B. Barton. Elsevier Science
Publishers, Amsterdam, The Netherlands.
Donaldson, L.R. & P.R. Olson, 1955. Development of rainbow trout
broodstock by selective breeding. Transact. Am. Fish. Soc. 85:
93101.
Dorson, M., B. Chevassus & C. Torhy, 1991. Comparative suscept-
ibility of three species of char and of rainbow trout x char triploid
hybrids to several pathogenic salmonid viruses. Diseases Aqua.
Organ. 11: 217224.
Doyle, R.W., C.M. Herbinger, M. Ball & G.A.E. Gall (eds), 1996.
Genetics in Aquaculture V. Aquaculture 137: 1358.
Dunham, R.A., 1990. Production and use of monosex or sterile
shes in aquaculture. Rev. Aquat. Sci. 2: 117.
Dunham, R.A. & R.E. Brummett, 1999. Response of two genera-
tions of selection to increased body weight in channel catsh,
Ictalurus punctatus, compared to hybridization with blue catsh,
I. furcatus, males. J. Appl. Aquacult. 9: 3745.
Dunham, R.A., K. Majumdar, E. Hallerman, D. Bartley, G. Mair, G.
Hulata, Z. Liu, N. Pongthana, J. Bakos, D. Penman, M. Gupta, P.
Rothlisberg & G. Hrstgen-Schwark, 2001. Review of the status
of aquaculture genetics, pp. 129157 in Aquaculture in the Third
Millenium, edited by R.P. Subasinghe, P. Bueno, M.J. Phillips,
C. Hough, S.E. McGladdery & J.R. Arthur. Technical Proceed-
ings of the Conference on Aquaculture in the Third Millennium,
Bangkok, Thailand, 2025 February 2000. NACA, Bangkok,
Thailand and FAO, Rome, Italy.
Eknath, A.E., M.M. Tayamen M.S. Palada-de Vera, J.C. Danting,
R.A. Reyes, E.E. Dionisio, J.B. Capili, H.L. Bolivar, T.A. Abella,
A.V. Circa, H.B. Bentsen, B. Gjerde, T. Gjedrem & R.S.V. Pul-
lin, 1993. Genetic improvement of farmed tilapias: the growth
performance of eight strains of Oreochromis niloticus tested in
different farm envuronments. Aquaculture 111: 171188.
Eknath, A.E., M.M. Dey, M. Rye, B. Gjerde, T.A. Abella, R.
Sevilleja, M.M. Tayamen, R.A. Reyes & H.B. Bentsen, 1998.
Selective breeding of Nile tilapia for Asia, pp. 8996 in Proceed-
ings of the 6th World Congress on Genetics Applied to Livestock
Production, Vol. 27.
Embody, G.C. & C.D. Hyford, 1925. The advantage of rearing
brook trout ngerlings from selected breeders. Transact. Am.
Fisher. Soc. 55: 135138.
Entis, E., 1997. Aquabiotech: a blue revolution? World Aquacult.
28(1): 1215.
FAO yearbook, 1998. Fishery statistics. Aquacult. Prod. 86(2): 47.
Fauvel, C., M. Suquet, C. Dreanno, V. Zonno & B. Menu, 1998.
Cryopreservation of sea bass (Dicentrarchus labrax) sperma-
tozoa in experimental and production simulating conditions.
Aquat. Living Resour. 11: 387394.
Felip, A., S. Zanuy., M. Carrillo & F. Piferrer, 1998. Study of the
treatment conditions leading to the mass-production of triploid
and gynogenetic sea bass. Options Mediterra. 34: 123129.
Felip, A., S. Zanuy., M. Carrillo & F. Piferrer, 1999. Growth and
gonadal development in triploid sea bass (Dicentrarchus labrax
L.) during the rst two years of age. Aquaculture 173: 389399.
Galbreath, P.F. & G.H. Thorgaard, 1996. Sexual maturation and
fertility of diploid and triploid Atlantic salmon x brown trout
hybrids. Aquaculture 137: 299311.
Galbreath, P.F. & B.L. Samples, 2000. Optimization of thermal
shock protocols for induction of triploidy in brook trout. North
Am. J. Aquacult. 62: 249259.
Galbusera, P., F.A.M. Volckaert & F. Ollevier, 2000. Gynogenesis
in the African catsh Clarias gariepinus (Burchell, 1822). III.
Induction of endomitosis and the presence of residual genetic
variation. Aquaculture 185: 2542.
Gall, G.A.E. & H. Chen (editors), 1993. Genetics in Aquaculture
IV. Aquaculture 111: 1331.
Gjedrem, T., 1985. Improvement of productivity through breeding
schemes. GeoJournal 10: 233241.
Gjedrem, T., 1997. Selective breeding to improve aquaculture
production. World Aquacult. 28: 3345.
Gjedrem, T., 2000. Genetic improvement of cold-water sh species.
Aquacult. Res. 31: 2533.
Gjerde, B., 1986. Growth and reproduction in sh and shellsh.
Aquaculture 57: 3755.
169
Gjerde, B. & M. Rye, 1997. Design of breeding programs in
aquaculture species possibilities and constraints, in Proc. of
the Genetics and Breeding of Mediterranean Aquaculture Spe-
cies, Seminar of the CIHEAM Network on Technology of
Aquaculture in the Mediterranean (TECAM), Jointly Organized
by CIHEAM and FAO, Zaragoza, Spain. Options Mediterr. 34:
181192.
Gjerde, B. & S.A. Korsvoll, 1999. Realized selection differentials
for growth rate and early sexual maturity in Atlantic salmon,
pp. 7374 in Towards Predictable Quality, Abstracts of Con-
tributions Presented at the Aquaculture Europe 99. European
Aquaculture Society Special Publication 27, Oostende, Belgium.
Gjerde, B., H.M. Gjen & B. Villanueva, 1996. Optimum designs
for sh breeding programmes with constrained inbreeding: mass
selection for a normally distributed trait. Livestock Prod. Sci. 47:
5972.
Gomelsky, B., N.B. Cherfas, Y. Peretz, N. Ben-Dom & G. Hulata,
1994. Hormonal sex inversion in the common carp (Cyprinus
carpio L.). Aquaculture 126: 265270.
Gorshkov, S., G. Gorshkova, A. Hadani, H. Gordin & W. Knibb,
1998. Chromosome set manipulations and hybridization exper-
iments in gilthead seabream (Sparus aurata). I. Induced gyn-
ogenesis and intergeneric hybridization using males of the red
seabream (Pagrus major). Israeli J. Aquacult. Bamidgeh 50:
99110.
Goudie, C.A., B.A. Simco, K.B. Davis & Q. Liu, 1995. Produc-
tion of gynogenetic and polyploid catsh by pressure-induced
chromosome set manipulation. Aquaculture 133: 185198.
Guo, X.M., G.A. DeBrosse & S.K. Allen, 1996. All-triploid Pacic
oyster (Crassostrea gigas) produced by mating tetraploids and
diploids. Aquaculture 142: 149161.
Gwo, J.C., 2000. Cryopreservation of aquatic invertebrate semen: a
review; in special issue: cryopreservation of gametes in aquatic
species. Aquacult. Res. 31: 259271.
Gwo, J.C., H. Ohta, K. Okuzawa & H.C. Wu, 1999. Cryop-
reservation of sperm from the endangered Formosan landlocked
salmon (Oncorhynchus masou formosanus). Theriogenology 51:
569582.
Hallerman, E.M. & A.R. Kapuscinski, 1995. Incorporating risk as-
sessment and risk management into public policies on genetically
modied nsh and shellsh. Aquaculture 137: 917.
Harrell, R.M., 1997. Hybridization and genetics, pp. 217234, in
Striped Bass and Other Morone Culture, edited by R.M. Harrell.
Elsevier Science Publishers, Amsterdam, The Netherlands.
Harrell, R.M., 1998. Genetics of stripped bass. World Aquacult. 29:
5659.
Harvey, B., 2000. The application of cryopreservation in sh genetic
conservation in North and South America, pp. 331337 in Cryo-
preservation in Aquatic Species, edited by T.R. Tiersch & P.M.
Mazik. World Aquaculture Society, Baton Rouge, Louisiana,
USA.
Heath, D.D., R.H. Devlin, T.J. Hilbish & G.K. Iwama, 1995. Multi-
locus DNA ngerprints in seven species of salmonids. Can. J.
Zool. 73: 600606.
Hedgecock, D., D.J. McGoldrick & B.L. Bayne, 1995. Hybrid vigor
in Pacic oysters: an experimental approach using crosses among
inbred lines. Aquaculture 137: 285298.
Hedgecock, D., C. Langdon, M. Blouin & S.K. Allen, Jr. 1996.
Genetic improvement of cultured pacic oysters by selection.
Coastal Marine Experiment Station Annual Report. Hateld
Marine Science Center and Agricultural Experiment Station,
Oregon State University, Oregon, USA, 40 p.
Hedgecock, D., D.J. McGoldrick, D.T. Manahan, J. Vavra, N. Ap-
pelmans &B.L. Bayne, 1996. Quantitative and molecular genetic
analyses of heterosis in bivalve mollusks. J. Exp. Marine Biol.
Ecol. 203: 4959.
Hendry, C.I., D.J. Martin-Robichaud & T.J. Benfey, 1999. Hor-
monal production of all-female Atlantic halibut (Hippoglos-
sus hippoglossus): the timing of sexual differentiation. Bull.
Aquacult. Assoc. Canada 99(4): 1315.
Hershberger, W.K., J.M. Meyers, W.C. McAuley & A.M. Saxton,
1990. Genetic changes in growth of coho salmon (Oncorhynchus
kisutch) in marine netpens, produced by 10 years of selection.
Aquaculture 85: 187197.
Hetzel, D.J.S., P.J. Crocos, G.P. Davis, S.S. Moore & N.C.
Preston, 2000. Response to selection and heritability for growth
in the Kuruma prawn, Penaeus japonicus. Aquaculture 181: 215
223.
Hew, C.L. & G. Fletcher, 1997. Transgenic sh for aquaculture.
Chem. Industry (London) 8: 311314.
Hew, C.L. & G. Fletcher, 2001. The role of aquatic biotechnology
in aquaculture. Aquaculture 197: 191204.
Hilbish, T.J., B.L. Bayne & A. Day, 1994. Genetics of physiolo-
gical differentiation within the marine mussel genus Mytilus.
Evolution 48: 267286.
Hinits, Y. & B. Moav, 1999. Growth performance studies in
transgenic Cyprinus carpio. Aquaculture 173: 285296.
Hulata, G., 1995. A review of genetic improvement of the common
carp (Cyprinus carpio L.) and other hybrids by crossbreeding,
hybridization and selection. Aquaculture 129: 143155.
Hulata, G., A. Cnaani, N. Umiel, J.J. Agresti, S. Seki, B. May,
S. Poompuang, E.M. Hallerman & G.A.E. Gall, 1999. Develop-
ment of a tilapia articial center of origin and genetic linkage
map based on AFLP and microsatellite loci, pp. 9394 in To-
wards Predictable Quality, Abstracts of Contributions Presented
at the Aquaculture Europe 99. European Aquaculture Society
Special Publication 27, Oostende, Belgium.
Issa, M.A., L. Horvath, M.A. Kosba & M. Sharrabi, 1986. A note
on the survival, growth, feed conversion and some morphological
characters of the reciprocal hybrids of silver carp (Hypophthal-
michthys molitrix Val.) and bighead carp (Aristichthys nobilis
Rich.) raised in polyculture. Aquacult. Hungarica 5: 714.
Jackson, T.R., M.M. Ferguson, R.G. Danzmann, A.G. Fishback,
P.E. Ihssen, M. OConnell & T.J. Crease, 1998. Identication of
two QTL-inuencing upper temperature tolerance in three rain-
bow trout (Oncorhynchus mykiss) half-sib families. Heredity 80:
143151.
Jeffreys, A.J., V. Wilson & S.L. Thein, 1985. Individual-specic
ngerprints of human DNA. Nature (London) 316: 6773.
Jenneckens, I., A. Mller-Belecke, G. Hrstgen-Schwark & J.-N.
Meyer, 1999. Proof of the successful development of Tilapia (Or-
eochromis niloticus) clones by DNA-ngerprinting. Aquaculture
173: 377388.
Johnson, L.A., 2000. Lessons from the cryopreservation of livestock
sperm, pp. 383387 in Cryopreservation in Aquatic Species, ed-
ited by T.R. Tiersch & P.M. Mazik. World Aquaculture Society,
Baton Rouge, Louisiana, USA.
Johnstone, R., 1993. Maturation control in Atlantic salmon, pp. 99
105 in Recent Advances in Aquaculture, Vol. 4, edited by J.F.
Muir & R.J. Roberts. Blackwell, Oxford, UK.
Johnstone, R., H.A. McLay & M.V. Walsingham, 1991. Production
and performance of triploid Atlantic salmon in Scotland. Cana-
dian Technical Report Fisheries and Aquatic Sciences 1789, pp.
1536.
Jnasson, J., 1996. Selection experiment in Atlantic salmon ranch-
ing. II. Variation among release sites and strains for return rate,
body weight and ratio of grilse to total return. Aquaculture 144:
277294.
170
Jnasson, J., B. Gjerde & T. Gjedrem, 1997. Genetic parameters
for return rate and body weight of sea-ranched Atlantic salmon.
Aquaculture 154: 219231.
Jones, C.M., C.P. McPhee & I.M. Ruscoe, 2000. Areview of genetic
improvement in growth rate in redclaw craysh Cherax quad-
ricarinatus (von Martens)(Decapoda: Parastacidae). Aquacult.
Res. 31: 6167.
Kang, J.-H., G. Yoshizaki, O. Homma, C.A. Strssmaan & F.
Takashima, 1999. Effect of an osmotic differential on the ef-
ciency of gene transfer by electroporation of sh spermatozoa.
Aquaculture 173: 297307.
Kapuscinski, A.R. & E.M. Hallerman, 1990. Transgenic sh and
public policy: anticipating environmental impacts of transgenic
sh. Fisheries 15: 211.
Kapuscinski, A.R. & E.M. Hallerman, 1991. Implications of in-
troduction of transgenic sh into natural ecosystems. J. Fisher.
Aquat. Sci. 48: 99107.
Khan, H.A., S.D. Gupta, P.V.G.K. Reddy, M.S. Tantia & G.V.
Kowtal, 1990. Production of sterile intergeneric hybrids and their
utility in aquaculture and reservoir stocking, pp. 4148 in Carp
Seed Production Technology, edited by P. Keshavanath & K.V.
Radhakrishnan. Special Publication Asian Fisheries Society, In-
dian Branch No. 2.
Kirpichnikov, V.S., Ju.I. Ilyasov, L.A. Shart, A.A. Vikhman, M.V.
Ganchenko, L.A. Ostashevsky, V.M. Simonov, G.F. Tikhonov &
V.V. Tjurin, 1993. Selection of Krasnodar common carp (Cyp-
rinus carpio L.) for resistance to dropsy: principal results and
prospects. Aquaculture 111: 720.
Knibb, W., 1997. Risk from genetically engineered and modied
marine sh. Transgenic Res. 6: 5967.
Knibb, W., 2000. Genetic improvement of marine sh which
method for industry? Aquacult. Res. 31: 1123.
Knibb, W., G. Gorshkova & S. Gorshkov, 1998. Genetic improve-
ment of cultured marine nsh: case studies, pp. 111149 in
Tropical Mariculture, edited by S. De Silva. Academic Press,
Sydney, Australia.
Kocher, T.D., W.-J. Lee, H. Sobolewska, D. Penman & B.
McAndrew, 1998. A genetic linkage map of a cichlid sh, the
tilapia (Oreochromis niloticus). Genetics 148: 12251232.
Koedprang, W. & U. Na-Nakorn, 2000. Preliminary study on
performance of triploid Thai silver barb, Puntius gonionotus.
Aquaculture 190: 211221.
Lahav, E., 1993. Use of sex-reversed females to produce all-male
tilapia (Oreochromis aureus) fry. Israeli J. Aquacult. Bamidgeh
45: 131136.
Lahnsteiner, F., 2000. Semen cryopreservation in the Salmonidae
and in the northern pike; in special issue: cryopreservation of
gametes in aquatic species. Aquacult. Res. 31: 245258.
Lawrence, C.S. & N.M. Morrissy, 2000. Genetic improvement of
marron Cherax tenuimanus Smith and yabbies Cherax spp. in
Western Australia. Aquacult. Res. 31: 6982.
Lawrence, C.S., Y.W. Cheng, N.M. Morrissy & I.H. Williams,
2000a. A comparison of mixed sex verses monosex growout and
diets on the growth rate of freshwater craysh (Cherax albidus).
Aquaculture 185: 281289.
Lawrence, C.S., N.M. Morrissy, P.E. Vercoe & I.H. Williams,
2000b. Hybridization in Australian freshwater craysh produc-
tion of all-male progeny. J. World Aquacult. Soc. 31: 651658.
Li, S., 2001. Aquaculture genetic research in China, pp. 1524 in
Fish Genetic Research in Member Countries and Institutions of
the International Network on Genetics in Aquaculture, edited by
M.V. Gupta & B.O. Acosta. ICLARM Conf. Proc. 64, 179 p.
Li, S., G.-Q. Lu & L. Bernatchez, 1998. Diversity of mitochondrial
DNA in the populations of silver carp, bighead carp, grass carp
and black carp in the middle- and lower reaches of the Yangtze
River. Acta Zool. Sinica 44: 8293.
Li, S. &X.M. Yang, 1996. Effects of two-way selection on biochem-
ical genetics of blunt snout bream (Megalobrama amblycephala).
J. Fish. Sci. China 3: 15.
Li, S. & W. Cai, 2000. Two-way selective response of Megalobrama
amblycephala. J. Fisher. China 24: 201205.
Li, S.S. & H.J. Tsai, 2000. Transfer of foreign gene to giant fresh-
water prawn (Macrobrachium rosenbergii) by spermatophore-
microinjection. Mol. Reprod. Develop. 56: 149154.
Li, Y., K.J. Wilson, K. Byrne, V. Whan, D. Iglesis, S.A. Lehnert, J.
Swan, B. Ballment, Z. Fayazi, M. Kenway, J. Benzie, S. Pong-
somboon, A. Tassanakajon & S.S. Moore, 2000. International
collaboration on genetic mapping of the black tiger shrimp, Pen-
aeus monodon: progress Update, in Proceedings of the Plant &
Animal Genome VIII Conference. San Diego, CA, USA.
Lin, T.-T. & N.-H. Caho, 2000. Cryopreservation of eggs and em-
bryos of shellsh, pp. 240-250 in Cryopreservation in Aquatic
Species, edited by T.R. Tiersch & P.M. Mazik. World Aquacul-
ture Society, Baton Rouge, Louisiana, USA.
Liu, Z.J., P. Li, H. Kucuktas, A. Nichols, G. Tan, X. Zheng,
B.J. Argue & R.A. Dunham, 1999a. Development of amplied
fragment length polymorphism (AFLP) markers suitable for ge-
netic linkage mapping of catsh. Transact. Am. Fisher. Soc. 128:
317327.
Liu, Z.J., P. Li, B.J. Argue & R.A. Dunham, 1999b. Random ampli-
ed polymorphic DNA markers: usefulness for gene mapping
and analysis of genetic variation of catsh. Aquaculture 174:
5968.
Longalong, F.M., AA.E. Eknath & H.B. Bentsen, 1999. Response
to bi-directional selection for frequency of early maturing fe-
males in Nile tilapia (Oreochromis niloticus). Aquaculture 178:
1325.
Lubzens, E., N. Daube, I. Pekarsky, Y. Magnus, A. Cohen, F. Yuse-
fovich & P. Feigin, 1997. Carp (Cyprinus carpio L.) spermatozoa
cryobanks-strategies in research and application. Aquaculture
155: 1330.
Lymberly, A.J., 2000. Genetic improvement in the Australian
aquaculture industry. Workshop summary. Aquacult. Res. 31:
145149.
Lymberly, A.J., R.G. Doup, G. Jenkins & T. Thorne, 2000.
Genetics in the aquaculture industry. Aquacult. Res. 31: 12.
Maclean, N., 1998. Regulation and exploitation of transgenes in
sh. Mutat. Res. 399: 255266.
Maclean, N. & R.J. Laight, 2000. Transgenic sh: an evaluation of
benets and risks. Fish Fisher. 1: 146172.
Mair, G.C., J.S. Abucay, J.A. Beardmore & D.O.F. Skibinski, 1995.
Growth performance of genetically male tilapia (GMT) derived
from YY-males in Oreochromis niloticus L.: on-station com-
parisons with mixed sex and sex reversed male populations.
Aquaculture 137: 313322.
Mair, G.C., J.S. Abucay, D.O.F. Skibinski, T.A. Abella & J.A.
Beardmore, 1997. Genetic manipulation of sex ratio for the large
scale production of all-male tilapia Oreochromis niloticus L.
Canad. J. Fisher. Aquat. Sci. 54: 396404.
Marian, T. & Z. Krasznai, 1978. Karyological investigations on
Ctenopharyngodon idella and Hypophthalmichthys molitrix and
their cross-breeding. Aquacult. Hungarica 1: 4450.
Marian, T. & Z. Krasznai, 1979. Comparative karyological studies
on Chinese carps. Aquaculture 18: 325336.
McAndrew, B., D. Penman & G. Hulata (eds), 1999. Genetics in
aquaculture VI. Aquaculture 173: 1497.
McAndrew, B.J., K.J. Rana & D.J. Penman, 1993. Conservation and
preservation of genetic variation in aquatic organisms, pp. 295
171
336 in Recent Advances in Aquaculture, Vol. 4, edited by J.F.
Muir & R.J. Roberts. Blackwell, Oxford, UK.
McConnell, S.K.J., C. Beynon, J. Leamon & D.O.F. Skibinski,
2000. Microsatellite marker based genetic linkage maps of Or-
eochromis aureus and O. niloticus (Cichlidae): extensive linkage
group segment homologies revealed. Anim. Genet. 31: 214218.
McEvoy, T.G., F. Gannon & J.M. Sreenan, 1992. Gene transfer in
sh: potential and practice. Anim. Biotechnol. 3: 221243.
Mims, S.D., W.L. Shelton, O. Linhart & C.Z. Wang, 1997. Induced
meiotic gynogenesis of paddlesh Polyodon spathula. J. World
Aquacult. Soc. 28: 334343.
Moav, B., Z.J. Liu, Y. Groll & P.B. Hackett, 1992. Selection of
promoters for gene transfer into sh. Molec. Mar. Biol. and
Biotechnol. 1: 338345.
Moav, R. & G.W. Wohlfarth, 1976. Two-way selection for growth
rate in the common carp (Cyprinus carpio L.). Genetics 82: 83
101.
Moore, S.S., V. Whan, G.P. Davis, K. Byrne, D.J.S. Hetzel & N.
Preston, 1999. The development and application of genetic mark-
ers for the Kuruma prawn Penaeus japonicus. Aquaculture 173:
1932.
Myers, J.M., P.O. Heggelund, G. Hudson & R.N. Iwamoto, 2001.
Genetics and broodstock management of coho salmon. Aquacul-
ture 197: 4362.
Na-Nakorn, U., W. Rangsin & S. Witchasunkul, 1993. Suitable
conditions for induction of gynogenesis in the catsh, Clarias
macrocephalus, using sperm of Pangasius sutchi. Aquaculture
118: 5362.
Nell, J.A., I.R. Smith & C.C. McPhee, 2000. The Sydney rock
oyster Saccostrea glomerata (Gould 1850) breeding programme:
progress and goals. Aquacult. Res. 31: 4549.
Nichols, K.M., W.P. Young, B.D. Robison, R. Danzmann, R.
Phillips, K.H. Brown, P. Bentzen, J. Brunelli, S. Ristow &
G.H. Thorgaard, 2001. An updated linkage map of rainbow trout
(Oncorhynchus mykiss) using doubled haploids. Proceedings of
the Plant and Animal Genome IX Conference, San Diego, CA,
USA.
Norris, A.T., D.G. Bradley & E.P. Cunningham, 2000. Parentage
and relatedness determination in farmed Atlantic salmon (Salmo
salar) using microsatellite markers. Aquaculture 182: 7383.
Nwadukwe, F.O., 1995. Analysis of production, early growth and
survival of Clarias gariepinus (Burchell), Heterobranchus longi-
lis (Val.) (Pisces: Clariidae) and their F
1
hybrids in ponds.
Netherlands J. Aquat. Ecol. 29: 177182.
Ostenfeld, T.H., R.H. Devlin & E. McLean, 1998. Transgenesis
changes body and head shape in Pacic salmon. J. Fish Biol. 52:
850854.
Palti, Y., J.E. Parsons & G.H. Thorgaard, 1999. Identication of
candidate DNA markers associated with IHN virus resistance
in backcrosses of rainbow and cutthroat trout. Aquaculture 173:
8194.
Pandian, T.J. & R. Koteeswaran, 1998. Ploidy induction and sex
control in sh. Hydrobiologia 384: 167243.
Paniagura-Chavez, C.G., J.T. Buchanan, J.E. Supan & T.R. Tiersch,
2000. Cryopreservation of sperm and larvae of the eastern oyster,
pp. 230239 in Cryopreservation in Aquatic Species, edited by
T.R. Tiersch & P.M. Mazik. World Aquaculture Society, Baton
Rouge, Louisiana, USA.
Parsons, J., 1998. Status of genetic improvement in commercially
reared stocks of rainbow trout. World Aquacult. 29: 4447.
Parsons, J., R. Busch, G. Thorgaard & P. Scheerer, 1986. Resistance
of diploid and triploid rainbow trout, coho salmon and reciprocal
hybrids to infectious hematopoietic necrosis (IHN). Aquaculture
57: 337343.
Penman, D.J. & B.J. McAndrew, 2000. Genetics for the man-
agement and improvement of cultured tilapias, pp. 227266 in
Tilapias: Biology and Exploitation, edited by M.C.M. Beveridge
&B.J. McAndrew. Kluwer Academic Publishers, Dordrecht, The
Netherlands.
Penney, R.W. & M.J. Hart, 1999. Distribution, genetic structure, and
morphometry of Mytilus edulis and M. trossulus within a mixed
species zone. J. Shellsh Res. 18: 367374.
Perez-Enriquez, R., M. Takagi & N. Taniguchi, 1999. Genetic
variability and pedigree tracing of a hatchery-reared stock of
red sea bream (Pagrus major) used for stock enhancement,
based on microsatellite DNA markers. Aquaculture 173: 413
423.
Piferrer, F., 2001. Endocrine sex control strategies for the feminiza-
tion of teleost sh. Aquaculture 197: 229281.
Pongthana, N., D.J. Penman, P. Baoprasertkul, M.G. Hussain,
M.S. Islam, S.F. Powell & B.J. McAndrew, 1999. Monosex fe-
male production in the silver barb (Puntius gonionotus Bleeker).
Aquaculture 173: 247256.
Poompuang, S. & E.M. Hallerman, 1997. Toward detection of
quantitative trait loci and marker-assisted selection in sh. Rev.
Fisher. Sci. 5: 253277.
Pullin, R.S.V., J. Bell, J.C. Danting & F. Longalong, 1998. Gene
banking for sh and other aquatic organisms: ICLARMSs per-
spectives and experience, pp. 3145 in Action Before Extinction:
Proceedings of an International Conference on Conservation of
Fish Genetics Diversity, edited by B. Harvey, C. Ross, D. Greer
& J. Carolsfeld. World Fisheries Trust, Victoria, BC, Canada.
Quillet, E., L. Foisil, B. Chevassus, D. Chourrout & F.G. Liu,
1991. Production of all-triploid and all-female brown trout for
aquaculture. Aquat. Living Resour. 4: 2732.
Rahman, M.A. & N. Maclean, 1999. Growth performance of trans-
genic tilapia containing an exogenous piscine growth hormone
gene. Aquaculture 173: 333346.
Rahman, M.A., A. Iyengar & N. Maclean, 1997. Co-injection
strategy improves integration efciency of a growth hormone
gene construct, resulting in lines of transgenic tilapia (Oreo-
chromis niloticus) expressing an exogenous growth hormone
gene. Transgenic Res. 6: 369378.
Rahman, M.A., R. Mak, H. Ayad, A. Smith & N. Maclean,
1998. Expression of a novel piscine growth hormone gene res-
ults in growth enhancement in transgenic tilapia (Oreochromis
niloticus). Transgenic Res. 7: 357369.
Reddy, P.V.G.K., H.A. Khan, S.D. Gupta, M.S. Tantia & G.V
Kowtal, 1990. On the ploidy of three intergeneric hybrids
between common carp (Cyprinus carpio communis L.) and
Indian major carps. Aquacult. Hungarica 6: 511.
Refstie, T., 1983. Hybrids between salmonids species. Growth and
survival in seawater. Aquaculture 33: 281285.
Rosenstein, S. & G. Hulata, 1994. Sex reversal in the genus
Oreochromis. Optimization of feminization protocol. Aquacult.
Fisher. Manag. 25: 329393.
Rothbard, S., W.L. Shelton, Z. Kulikovsky, I. Rubinshtein, Y.
Hagani & B. Moav, 1997. Chromosome set manipulations in the
black carp. Aquacult. Int. 5: 5164.
Rothbard, S., I. Rubinstein, L. David & W.L. Shelton, 1999.
Ploidy manipulations aimed to produce androgenetic Japanese
ornamental (koi) carp, Cyprinus carpio L. Israeli J. Aquacult.
Bamidgeh 51: 2639.
Rye, M. & A.E. Eknath, 1999. Genetic improvement of tilapia
through selective breeding experience from Asia, pp. 207
208, in Towards Predictable Quality, Abstracts of Contributions
Presented at the Aquaculture Europe 99. European Aquaculture
Society Special Publication 27, Oostende, Belgium.
172
Sakamoto, T., R.G. Danzmann, N. Okamoto, M.M. Ferguson &
P.E. Ihssen, 1999. Linkage analysis of quantitative trait loci as-
sociated with spawning time in rainbow trout (Oncorhynchus
mykiss). Aquaculture 173: 3343.
Sakamoto, T., R.G. Danzmann, K. Gharbi, P. Howard, A. Ozaki,
S.K. Khoo, R.A. Woram, N. Okamoto, M.M. Ferguson, L.-E.
Holm, R. Guyomard & B. Hoyheim, 2000. A microsatellite link-
age map of rainbow trout (Oncorhynchus mykiss) characterized
by large sex-specic differences in recombination rates. Genetics
155: 13311345.
Seeb, J.E., G.H. Thorgaard & T. Tynan, 1993. Triploid hybrids
between chum salmon female x chinook salmon male have early
sea-water tolerance. Aquaculture 117: 3745.
Shelton, W.L., 1986. Broodstock development for monosex produc-
tion of grass carp. Aquaculture 57: 311319.
Shelton, W.L., 1995. Genetic manipulation to produce sterile shes,
pp. 98102 in Herbivorous Fishes: Culture and Use for Weed
Management, edited by K. Opuszynski & J.V. Shireman. CRC
press, Boca Raton, FL.
Shireman, J.V. & C.R. Smith, 1983. Synopsis of biological data
on the grass carp, Ctenopharyngodon idella (Cuvier and Valen-
ciennes, 1844). FAO Fisheries Synopsis 135, 86 p.
Sin, F.Y.T., S.P. Walker, J.E. Symonds, U.K. Mukherjee, J.G.I. Khoo
& I.L. Sin, 2000. Electroporation of salmon sperm for gene trans-
fer: efciency, reliability, and fate of transgene. Mol. Reprod.
Develop. 56: 285288.
Smitherman, R.O. & R.A. Dunham, 1985. Genetics and breeding,
pp. 283316 in Channel Catsh Culture, edited by C.S. Tucker.
Elsevier Scientic Publishing, Amsterdam, The Netherlands.
Stevens, E.D. & R.H. Devlin, 2000a. Gill morphometry in growth
hormone transgenic Pacic coho salmon, Oncorhynchus kisutch.
Environ. Biol. Fish. 58: 113117.
Stevens, E.D. & R.H. Devlin, 2000b. Intestinal morphology in
growth hormone transgenic Pacic coho salmon, Oncorhynchus
kisutch Walbaum. J. Fish Biol. 56: 191195.
Sumantadinata, K., 1995. Present state of common carp (Cyp-
rinus carpio L.) stocks in Indonesia. Aquaculture 129: 205
209.
Suquet, M., C. Dreanno, C. Fauvel, J. Cosson & R. Billard, 2000.
Cryopreservation of sperm in marine sh, in special issue: cryo-
preservation of gametes in aquatic species. Aquacult. Res. 31:
231243.
Sutton, D.L., J.G. Stanley & W.W. Miley, 1981. Grass carp hybrid-
ization and observations of a grass carp x bighead carp hybrid. J.
Aquat. Plant Mana. 19: 3739.
Tan, G., A. Karsi, P. Li, S. Kim, X. Zheng, H. Kucuktas, B.J. Ar-
gue, R.A Dunham & Z.J. Liu, 1999. Polymorphic microsatellite
markers in Ictalurus punctatus and related catsh species. Mol.
Ecol. 8: 17581760.
Thien, T.M. & T.D. Trong, 1995. Genetic resources of common carp
in Vietnam. Aquaculture 129: 216.
Thompson, D., 2000. Make way for Frnakensh. Time, 13 June
issue.
Thongpan, A., M. Mingmuang, S. Thinchant, R. Cooper, T. Tiersch
& K. Mongkonpunya, 1997. Genomic identication of catsh
species by polymerase chain reaction and restriction enzyme ana-
lysis of the gene encoding the immunoglobulin M heavy chain
constant region. Aquaculture 156: 129137.
Thorgaard, G.H., 1986. Ploidy manipulation and performance.
Aquaculture 57: 5764.
Tidwell, J.H., C.D. Webster, J.A. Clark & M.W. Brunson, 1994.
Pond culture of female green sunsh (Lepomis cyanellus) x
male bluegill (L. macrochirus) hybrids stocked at two sizes and
densities. Aquaculture 126: 305313.
Tiersch, T.R. & P.M. Mazik (eds), 2000. Cryopreservation in
Aquatic Species. World Aquaculture Society, Baton Rouge,
Louisiana, USA.
Tremblay, R., B. Myrand & J.M. Sevigny, 1998. Genetic charac-
terization of wild and suspension-cultured blue mussels (Mytilus
edulis Linneaus, 1758) in the Magdalen Islands (southern Gulf
of St. Lawrence, Canada). J. Shellsh Res. 17: 11911202.
Tsai, H.J., 2000. Electroporated sperm mediation of a gene transfer
system for nsh and shellsh. Mol. Reprod. Develop. 56: 281
284.
Tuan, P.A., D.C. Little & G.C. Mair, 1998. Genotypic effects on
comparative growth performance of all-male tilapia Oreochromis
niloticus (L.). Aquaculture 159: 293302.
Tuan, P.A., G.C. Mair, D.C. Little & J.A. Beardmore, 1999. Sex
determination and the feasibility of genetically male tilapia pro-
duction in the Thai- Chitralada strain of Oreochromis niloticus
(L.). Aquaculture 173: 257269.
Utter, F.M., 1991. Biochemical genetics and shery management:
an historical perspective. J. Fish Biol. 39 (Supp. A): 120.
Utting, S.D., P.F. Millican & I. Laing, 1996. The breeding potential
and biochemical composition of triploid Manila clams, Tapes
philippinarum Adams and Reeve. Aquacult. Res. 27: 573580.
Van der Lende, T., J. Komen & E.W. Brascamp, 1998. The role
of reproductive techniques in genetic improvement and dissem-
ination of genetic gain in livestock production and aquaculture,
in Impact of Reproductive Technology on Animal Breeding and
Genetic Conservation, Proceedings of the Bertebos Prize Sym-
posium, edited by S. Einarsson & J. Rendel. Acta Agricult.
Scand. (Sec. A, Anim. Sci.) 29: 9097.
Van Olst, J.C. & J.M. Carlberg, 1990. Commercial culture of hybrid
striped bass: status and potential. Aquacult. Mag. 16(1): 4959.
Vardi, L., I. Benk, J. Varga & L. Horvth, 1999. Induction of
diploid gynogenesis using interspecic sperm and production
of tetraploids in African catsh, Clarias gariepinus Burchell
(1822). Aquaculture 173: 401411.
Venugopal, T., T.J. Pandian, S. Mathavan & N. Sarangi, 1997.
Gene transfer in Indian major carps by electroporation. Curr. Sci.
(Bangalore) 74: 636638.
Volckaert, F. & F. Ollevier, 1998. Transgenic sh. The future of
sh with novel genes, pp. 3358 in Genetics and Aquaculture
in Africa, edited by J.F. Agnese. Actes du colloque Genetique
et aquaculture en Afrique, Abidjan. Editions ORSTOM, Bondy,
France.
Volckaert, F.A., C. Van den Haute, P.H. Galbusera & F. Ollevier,
1997. Gynogenesis in the African catsh, Clarias gariepinus
(Burchell, 1822). Optimizing the induction of polar-body gyn-
ogenesis with combined pressure and temperature shocks.
Aquacult. Res. 28: 329334.
Ward, R.D. & P.M. Grewe, 1995. Appraisal of molecular genetic
techniques in sheries, pp. 2954 in Molecular Genetics in Fish-
eries, edited by G.R. Carvalho & T.J. Pitcher. Chapman & Hall,
London, UK.
Ward, R.D., L.J. English, D.J. McGoldrick, G.B. Maguire, J.A. Nell
& P.A. Thompson, 2000. Genetic improvement of the Pacic
oyster Crassostrea gigas (Thunberg) in Australia. Aquacult. Res.
31: 3544.
Webster, C.D., L.G. Tiu & J.H. Tidwell, 1997. Growth and body
composition of juvenile hybrid bluegill Lepomis cyanellus x L.
macrochirus fed practical diets containing various percentages
of protein. J. World Aquacult. Soc. 28: 230240.
Wills, P.S., J.M. Paret & R.J. Sheehan, 1994. Pressure induced trip-
loidy in hybrid Lepomis. J. World Aquacult. Soc. 25: 507511.
Wohlfarth, G.W., 1993. Heterosis for growth rate in common carp.
Aquaculture 113: 3146.
173
Wohlfarth, G.W., 1994. The unexploited potential of tilapia hybrids
in aquaculture. Aquacult. Fisher. Manag. 25: 781788.
Wohlfarth, G.W., 1995. The common carp and Chinese carps, pp.
138176 in Conservation of Fish and Shellsh Resources: Man-
aging Diversity, edited by J.E. Thorpe, G.A.E. Gall, J.E. Lanan
& C.E. Nash. Academic Press.
Wohlfarth, G.W. & G. Hulata, 1983. Applied Genetic of Tilapias.
ICLARM Studies and Reviews, 2nd edn. Vol. 6, 26 p.
Wright, J.M., 1990. DNA ngerprinting in shes, pp. 5791 in
Biochemistry and Molecular Biology of Fishes, Vol. 2, edited
by P. Hochachka & T. Mommsen. Elsevier, Amsterdam, The
Netherlands.
Wright, J.M. & P. Bentzen, 1995. Microsatellites: genetic markers
for the future, pp. 117121 in Molecular Genetics in Fisher-
ies, edited by G.R. Carvalho and T.J. Pitcher. Chapman & Hall,
London, UK.
Wu, L. & Z. Wang, 1997a. Studies on the expressions and reg-
ulations of isozymic genes in silver carp (Hypophthalmichthys
molitrix) during ontogenesis. Acta Hydrobiol. Sinica 21: 4958.
Wu, L. & Z. Wang, 1997b. Biochemical genetic structure and vari-
ation in a natural population of silver carp from the middle
reaches of the Yangtze River. Acta Hydrobiolo. Sinica 21:
157162.
Yamamoto, E., 1999. Studies on sex-manipulation and production of
cloned populations in hirame, Paralichthys olivaceus (Temminck
et Schlegel). Aquaculture 173: 235246.
Yang, S. & J.F. Gui, 1999. Isozyme analysis and preliminary con-
rmation of the genetic markers in two articial gynogenetic
populations of silver carp, Hypophthalmichthys molitrix. Acta
Hydrobiol. Sinica 23: 264268.
Young, W.P., P.A. Wheeler, R.D. Fields & G.H. Thorgaard, 1996.
DNA ngerprinting conrms isogenicity of androgenetically
derived rainbow trout lines. J. Heredity 87: 7781.
Young, W.P., P.A. Wheeler, V.H. Coryell, P. Keim & G.H. Thor-
gaard, 1998. A detailed linkage map of rainbow trout produced
using doubled haploids. Genetics 148: 839850.
Yoon, C.K., 2000. Altered salmon leading way to dinner plates, but
rules lag. The New York Times, May 1.