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The biology of triploid fish

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DOI: 10.1007/s11160-004-8361-8

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Reviews in Fish Biology and Fisheries (2004) 14: 391–402
Springer 2005
DOI 10.1007/s11160-004-8361-8

The biology of triploid fish

Basant K. Tiwary1, R. Kirubagaran2 & Arun K. Ray3

1
Department of Life Science, Assam (Central) University, Silchar 788 011, India (Phone: +91-3842-270823;
fax: +91-3842-270802; E-mail: basant68@email.com; 2National Institute of Ocean Technology, Chennai 601
302, India; 3Department of Animal Physiology, Bose Institute, Kolkata 700 054, India

Accepted 8 December 2004

Contents
Abstract page 391
Introduction 391
Induction of triploidy 392
Detection of triploids 392
Morpho-anatomical changes in triploids 393
Growth performances in triploids 393
Haematology of triploids 394
Food conversion efficiency in triploids 394
Behavior of triploids 395
Endocrinology of triploids 395
Gonadal development in triploids 396
Future prospects in triploid research 397
Conclusion 398
References 398

Key words: detection, growth, induction, morpho-anatomy, physiology, triploidy

Abstract

This review deals with major areas of triploidy research in fish. It includes not only methods for induction
and detection of triploidy but also the impact of triploidy on morphology, anatomy, growth, haematology,
energetics, behaviour, endocrinology and gonads in various species of fish, studied so far. The future
prospects of research on triploid fish are discussed inviting researchers with diverse areas of interest in fish
biology.

Introduction may be directed towards improving flesh quality

Induction of triploidy in fish has been found to be
useful for the control of overpopulation, for
increasing growth in juveniles, and for extending
survival and improving growth in mature fish.
Triploids have three sets of chromosomes instead
of two sets in diploids. Triploids are generally
sterile due to irregular meiotic division of chro-
mosomes resulting in reduced gonadal develop-
ment and aneuploid gametes. Therefore, in such
fish, the energy consumption for sexual matura-
tion may be avoided and more biological effort
and somatic growth. Induction of triploidy has
been achieved in many species of fish by treatment
designed to block the second meiotic division or to
prevent extrusion of the second polar body of the
egg. Retention of second polar body of the fertil-
ized egg may happen without special treatment in
some cases or may result from temperature, pres-
sure or chemical treatment of the fertilized egg.
The natural occurrence of triploids in fish popu-
lations is due to occasional failure of extrusion of
the second polar body of the fertilized egg
(Thorgaard and Gall, 1979).
392

Induction of triploidy Table 1. Methods for inducing triploidy in fish

Method Species References

Temperature treatments (both heat and cold
shock) of fertilized eggs have widely been used to Cold shock Oncorhynchus Thorgaard and
suppress second polar body extrusion for induc- mykiss Gall (1979)
tion of triploidy in fish. Temperature shock treat- Heat shock Oreochromis Varadaraj and
ments are inexpensive to apply and can be mossambicus Pandian (1988)
successfully adapted for mass production by fish Pressure shock Danio rerio Streisinger et al.
farms. Cold shock in general has been most suc- (1981)
cessful in warm water fishes (Thorgaard and Gall, Nitric oxide Salmo salar Johnstone et al.
1979). However, heat shock was 100% effective in (1989)
tilapia (Oreochromis mossambicus) (Varadaraj and Freon Salmo salar Johnstone et al.
Pandian, 1988). (1989)
Hydrostatic pressure shock has also been suc- High pH & Ca++ Oncorhynchus Ueda et al. (1988)
mykiss
cessfully used to block second polar body extrusion in
Cytochalasin B Salmo salar Refstie et al. (1977)
rainbow trout (Onchorhynchus mykiss) (Lou and
Colchicine Salvelinus Smith and Lemoine
Purdom, 1984), zebrafish (Danio rerio) (Streisinger
fontinalis (1979)
et al., 1981), common carp (Cyprinus carpio) (Linhart
et al., 1991), Nile tilapia (Oreochromis niloticus)
(Hussain et al., 1991), yellow perch (Perca flavescens) cellular sizes of erythrocytes in triploids are
(Malison et al., 1993), coho salmon (Oncorhynchus increased to accommodate the increased amount of
kisutch) (Piferrer et al., 1994a), and hybrid channel nuclear DNA while maintaining a normal ratio of
catfish (Ictalurus punctatus) · blue catfish (Ictalurus nuclear to cytoplasmic volume (Small and Benfey,
furcatus) (Goudie et al., 1995). 1987). Unfortunately, this method does not always
Chemicals were also used to block polar body accurately reflect ploidy (Thorgaard and Gall, 1979;
extrusion in fertilized eggs of fish. Refstie et al. Wolters et al., 1982a). Therefore, it is generally
(1977) produced mosaic polyploid–diploid Atlantic calibrated using direct ploidy measurement tech-
salmon (Salmo salar) after exposing fertilized eggs to niques. However, erythrocyte volume measurement
cytochalasin B. Smith and Lemoine (1979) reported using a coulter counter with channelizer was
similar production in brook trout (Salvelinus fonti- extremely accurate and rapid method of determin-
nalis) after exposing fertilized eggs to colchicine. ing ploidy in grass carp (Ctenopharyngodon idella)
Among anaesthetics, nitrous oxide induced high (Wattendorf, 1986). A fluorescent nuclear stain,
triploid yield (79.7%) followed by freon induced Ethidium bromide has been used for detection of
lower yield in Atlantic salmon (Johnstone et al., triploidy in Asian catfish (Clarias macrocephalus)
1989). Either sperm or freshly fertilized eggs of (Thititananukij et al., 1996).
rainbow trout when treated with high pH and cal- Triploid cell contains three nucleoli in their
cium resulted in high incidence of triploidy (Ueda nucleus instead of the two in a diploid cell. Nucleolar
et al., 1988) (Table 1). counts have been used to identify triploid hybrids of
crucian carp (Carassius carassius) and common carp
(Cherfas and Ilyasova, 1980), triploid European
Detection of triploids catfish (Silurus glanis) (Linhart and Flajshans, 1995)
and rainbow trout (Al-Sabti, 1995).
Triploid fish have been identified by various indi- Protein electrophoresis has also been used to
rect methods such as measurement of nuclear and identify triploid fish due to differences in relative
cellular size, counting of nucleoli, electrophoresis of dosage of alleles between diploids and triploids.
proteins and examination of morphology. Direct Liu et al.(1978) used muscle myogen and creatine
methods include chromosome counting and DNA kinase pattern differences to detect triploid and
content determination (Thorgaard, 1983). Nuclear diploid ginbuna (Carassius auratus langsdorfi).
and cell size measurements of erythrocyte are most Allozyme markers at three loci (ADH, EST &
convenient and widely used methods when identi- 6PGD) were used for proving triploidy in red sea
fying a triploid fish (Purdom, 1993). Nuclear and bream (Pagrus major) (Sugama et al., 1992).
 393

Chromosome counting is a highly precise and segregation based on condition factor in the grass
direct method to identify triploidy in a fish carp and bighead carp hybrid. Diploids and
(Thorgaard, 1983). It is a tedious and time-con- autotriploids, which have three sets of conspecific
suming method that generally requires the sacrifice chromosomes, had few morphological differences.
of experimental fish. However, chromosomes can Remarkable morphological similarities between
be prepared from regenerating fin (Streisinger triploids and diploids have been observed in rainbow
et al., 1981; Tiwary et al., 1997) and lymphocyte trout (Leary et al., 1985). However, triploid stickle-
culture (Thorgaard and Gall, 1979) of adult fish back (Gasterosteus aculeatus) had a shorter trunk
without any sacrifice. and larger tail than diploids (Swarup, 1959). Trip-
The DNA measurement of individual cells loid male channel catfish had smaller heads than
with flow cytometry is another accurate and rapid diploid males (Wolters et al., 1982b). Minor differ-
method for triploidy determination. It has been used ences in body depth, predorsal length, width of gape,
to detect triploidy in rainbow trout (Thorgaard fin ray and scale counts have been observed between
et al., 1982), grass carp · bighead carp (Aristichthys diploids and autotriploid grass carp (Bonar et al.,
nobilis) hybrids (Allen and Stanley, 1983), walleye 1988). Morphological differences between diploids
(Sander vitreus) (Ewing et al., 1991), silver carp and triploids have also been observed in common
(Chao et al., 1993) and Amazon molly (Poecilia carp (Gomelsky et al., 1992) and bighead carp (Tave,
formosa) (Lamatsch et al., 2000). Feulgen positive 1993). Tiwary et al. (1999) observed significant
blood cell nuclei were analyzed through microden- differences in the ratio of nine different morpho-
sitometry to detect triploid silver carp (Gervai et al., metric characters between diploid and triploid
1980). The relative frequencies of triploids in a mixed catfish (Heteropneustes fossilis). The ratio between
population of diploids and triploids have been standard length and body depth (SL/BD) was
estimated with iterative mathematical analysis of effective in separating triploids from diploids.
microsatellite DNA data (Krieger and Keller, 1998) Lower jaw deformity in triploid Atlantic
(Table 2). salmon (Sutterlin et al., 1987; Jungalwalla, 1991;
Sutterlin and Collier, 1991; McGeachy et al.,
1996) is the most frequently reported skeletal
Morpho-anatomical changes in triploids
abnormality associated with triploidy in fish. Tave
(1993) observed deformities in the head of triploid
Morphological differences between diploid and
bighead carp and grass carp. In contrast, Fast
autotriploid fish are not apparent in most species
et al. (1995) reported a reduction in frequency of
of fish except grass carp, common carp, bighead carp
deformities in triploid Asian catfish. Only two
and Indian catfish (Bonar et al., 1988; Gomelsky
anatomical differences were reported between
et al., 1992; Tave, 1993; Tiwary et al., 1999). Cassani
triploid and diploid fish including impaired
et al. (1984) reported 95% accuracy in visual
gonads. The first difference was the divided spleen
in some triploid rainbow trout (Okada, 1985).
Table 2. Methods for detection of triploidy in fish
Additionally, muscle growth and development of
Method Species References Atlantic salmon displayed a lower density of
satellite cells, reduced rates of fibre recruitment,
Nuclear and Pleuronectes Purdom (1972) hypertrophy of muscle fibres, advanced develop-
Cellular size platessa ment of myotubes, myofibrils and acetylcholines-
Coulter counter Ctenopharyngodon Wattendorf (1986) terase staining at the myosepta (Johnston et al.,
idella
1999). The profound changes in size of the air sac
Nucleoli count Carassius carassius Cherfas and
and spinal deformities were observed in triploid
· Cyprinus carpio Ilyasova (1980)
Indian catfish (Tiwary and Ray, 2004).
Protein C. auratus Liu et al. (1978)
electrophoresis langsdorfi
Chromosome Danio rerio Streisinger et al.
number (1981)
Growth performances in triploids
DNA content Oncorhynchus Thorgaard et al.
mykiss (1982) Growth rate is an important characteristic for
the genetic improvement in fish for commercial
394
production (Purdom, 1993). Triploids are expected
to have a higher growth potential due to sterility and
reduced gonadal development. Growth perfor-
mances between diploids and triploids vary between
species (Table 3). Growth rates in triploids are not
significantly different compared to their diploid
controls in mud loach (Misgurnus mizolepis), Atlan-
tic salmon and two species of tilapia (Oreochromis
niloticus and O. aureus) (Chang et al., 1993; Kim
et al., 1994; Hussain et al., 1995; McGeachy et al.,
1995). Triploids in species such as rainbow trout,
sunshine bass (a hybrid), coho salmon, Atlantic
salmon tend to grow poorly when compared to their
diploid counterparts (Solar et al., 1984; Galbreath
et al., 1995; Kerby et al., 1995; Withler et al., 1995;
McCarthy et al., 1996). However, triploids have a
better growth rate than diploids at maturation in
catfish and tilapia (Wolters et al., 1982b; Bramick
et al.,1995; Tiwary et al., 1997). Better growth has
been reported in triploids of tilapia and European
catfish at the age of 14 weeks (Valenti, 1976;
Krasznai and Marian, 1986). No weight gain was
observed in female triploid brook trout even after
long-term treatment with anabolic steroid, estradiol-
17b (Schafhauser-Smith and Benfey, 2003a).
Haematology of triploids
The cellular and nuclear dimensions of erythrocytes
are increased in various species of triploid fish
(Benfey, 1999). However, this increase in cell size is
Table 3. Growth performances of triploid fish
Species Growth References
Ictalurus punctatus + Wolters et al. (1982b)
Oreochromis niloticus + Bramick et al. (1995)
Heteropneustes fossilis + Tiwary et al. (1997)
Oreochromis aureus + Valenti (1976)
Silurus glanis + Krasznai and Marian
(1986)
Oncorhynchus mykiss ) Solar et al. (1984)
Lepomis gibbosus ) Kerby et al. (1995)
Oncorhynchus kisutch ) Withler et al. (1995)
Salmo salar ) Galbreath et al. (1995)
Misgurnus mizolepis NC Kim et al. (1994)
Salmo salar NC McGeachy et al. (1995)
Oreochromis niloticus NC Hussain et al. (1995)
Oreochromis aureus NC Chang et al. (1993)
(+) positive, ()) negative, (NC) no change.
compensated by a proportional decrease in number
of erythrocytes, thereby maintaining a constant
haematocrit value for a particular species (Barker
et al., 1983; Sezaki et al., 1983; Benfey and Sutterlin,
1984b; Ueno, 1984; Graham et al., 1985; Small and
Randall, 1989; Aliah et al., 1991; Sezaki et al., 1991,
Parsons, 1993; Biron and Benfey, 1994; Yamamoto
and Iida, 1994). The one exception was no decrease
in erythrocyte numbers in rainbow trout O. mykiss
(Virtanen et al., 1990). Increases in the size of ery-
throcytes in triploid fish were followed by higher
haemoglobin content per cell (Barker et al., 1983;
Sezaki et al., 1983; Benfey and Sutterlin, 1984; Ueno,
1984; Graham et al., 1985; Aliah et al., 1991; Sezaki
et al., 1991, Parsons, 1993; Yamamoto and Iida,
1994), but total haemoglobin levels in triploid fish
showed variable results in different species. Triploids
of Atlantic salmon (Benfey and Sutterlin, 1984b;
Graham et al., 1985), coho salmon (Small and
Randall, 1989), white crappies (Pomoxis annularis)
(Parsons, 1993) and rainbow trout (Yamamoto and
Iida, 1994) exhibited lower haemoglobin levels than
diploids, whereas fish like hybrids of grass carp and
bighead carp (Barker et al., 1983), ginbuna (Sezaki
et al., 1983, 1991), ayu (Plecoglossus altivelis) (Aliah
et al., 1991), brook trout (Stillwell and Benfey, 1994,
1996) had haemoglobin levels similar to their diploid
counterparts. There was a significant decrease in the
haematocrit value and total blood haemoglobin
after long term treatment with estradiol-17b
(Schafhauser-Smith and Benfey, 2003a). Increased
enzyme activity was assayed in triploid ginbuna and
spinous loach (Cobitis biwae) (Sezaki et al., 1983,
1988). However, this enzyme activity was balanced
by fewer erythrocytes in triploid fish, keeping the
activity level similar to diploids in per unit volume
(Barker et al., 1983; Sezaki et al., 1983, 1988).
Physiological parameters of blood such as oxygen
carrying capacity, pH and nucleotide triphosphate
(NTP) levels in the blood of triploid Atlantic salmon
(Graham et al., 1985) and size of the heart in triploid
white crappies (Parsons, 1993) were not significantly
different from diploids. Plasma osmolality and oxy-
gen consumption recovered earlier in triploid brook
trout after exhaustive exercise when compared to
diploids (Hyndman et al., 2003).
Food conversion efficiency in triploids
The feed conversion efficiency has been well stud-
ied in triploid grass carp (Wiley and Wike, 1986),

Plasma hormone levels (ng/ml)
8 12
7 2n
3n
6
5
4 *
*
3
* 2
2
* 0
1
0 Figure 2. Plasma levels of gonadotropin (GtH-II), testosterone
GtH-II T E2
Figure 1. Plasma levels of gonadotropin (GtH-II), testosterone
(T) and Estradiol-17b (E2) during annual reproductive cycle of
male catfish (H. fossilis) (* indicates P < 0.001, 2n–3n).
Redrawn after Tiwary et al. (2001).
African catfish (Clarias gariepinus) (Henken et al.,
1987) and rainbow trout (Oliva-Teles and Kaushik,
1990b). No significant differences were observed in
protein efficiency ratio, net protein utililization,
efficiency of energy gain, total energy intake
and nitrogen balance in these species. Triploid
females, however, showed higher fat deposition in
viscera (Chevassus et al., 1984; Lincoln and Scott,
1984; Johnson et al., 1986; Henken et al., 1987;
Fauconneau et al., 1990) and better muscle
pigmentation (Choubert and Blanc, 1989) than
maturing diploid females. During development,
embryo, larvae and juveniles of triploid rainbow
trout do not differ from diploids with respect to
ammonia excretion rate, energy expenditure, oxi-
dation of glucose and amino acids (Fauconneau
et al., 1986, 1989; Oliva-Teles and Kaushik, 1987a,
1990a). Still, utilization of protein for energy pro-
duction was significantly lower in juvenile triploids
(Oliva-Teles and Kaushik, 1987b, 1990b).
Behavior of triploids
The central nervous system (CNS) and hormones
regulate the initiation, coordination and execution
of all kinds of behavior. The reduction in cell
numbers in CNS and reduced levels of hormones
395
Plasma hormone levels (ng/ml)
2n
10 3n
8
*
6
*
4
GtH-II T E2
(T) and Estradiol-17b (E2) during annual reproductive cycle of
female catfish (H. fossilis). (* indicates P < 0.001, 2n–3n).
Redrawn after Tiwary et al. (2001).
(Benfey, 1999) in triploid fish suggest lower
responsiveness to various environmental stimuli
leading to expression of abnormal behavior in
triploid fish. Triploids were found to be less
aggressive in rainbow trout (Boulanger, 1987), ayu
(Aliah et al., 1990; Inada and Taniguchi, 1992),
fighting fish (Beta splendens) (Kavumpurath and
Pandian, 1992a, b), Atlantic salmon (Carter et al.,
1994) than diploids. Triploid ayu were less
responsive to sound and light stimuli than diploid
fish (Aliah et al., 1990). Indicators, such as mini-
mum voltage required for eliciting a response and
number of days spent in learning to avoid electric
shock, in triploid brook trout (Deeley and Benfey,
1995), and lower jumping efficiency in triploid ayu
when crossing a barrier (Inada and Taniguchi,
1992), did not differ from diploids.
Endocrinology of triploids
Gonadotropin releasing homone (GnRH) has
central role in vertebrates by regulating the release
of gonadal steroids mediated through gonadotro-
pin (GtH). A positive correlation has been
observed between brain GnRH and gonadal
maturity in teleosts (Gentile et al., 1986; Amano
et al., 1992). However, there are only two reports
available on the nature of GnRH in triploid fish.
The first is reduced level of GnRH in triploid
rainbow trout (Breton and Sambroni, 1996) and
396
the second finding is decrease in size and number
of immunoreactive GnRH cells in preoptic area
(POA) of Indian catfish (Tiwary et al., 2002) and
low immunoreactivity in pituitary. The low activ-
ity of GnRH in triploid fish may be due to absence
of positive feedback stimulation by sex steroids
and/or decreased responsiveness of sensory cells to
environmental cues required for gonadal matura-
tion in teleosts.
Female triploid fish exhibited low levels of
gonadotropin and sex steroids in various species
(Lincoln and Scott, 1984; Sumpter et al., 1984;
Nakamura et al., 1987a; Benfey et al., 1989b;
Kobayashi et al., 1998; Tiwary et al., 2001, Fig-
ure 1). Triploid females have a low biosynthesis of
vitellogenin due to low levels of sex steroids.
However, exogenous estradiol-17b (E2) can induce
production of vitellogenin in coho salmon (Benfey
et al., 1989a) and rainbow trout (Atteke et al.,
2003; Kobayashi et al., 1998). Schafhauser-Smith
and Benfey (2003b) observed similar ovarian ste-
roidogenesis and gonadotropin induced E2 pro-
duction in both diploid and triploid female brook
trout. Either progesterone or E2 was able to
upregulate the mRNA levels of estradiol receptor
(ER) in the brain of triploid female rainbow trout
but they have no co-operative effect (Atteke et al.,
2003). This suggests occurrence of ER in the brain
of triploid female as ER-mRNA and immunore-
active ER were also previously detected in triploid
rainbow trout (Anglade et al., 1994). Female
triploids showed reduced levels of E2 and T when
compared to diploids (Hussain et al., 1995).
Female triploid brook trout showed low plasma E2
and testosterone levels after long-term treatment
with E2 in the diet (Schafhauser-Smith and Benfey,
2003a). However, the plasma levels of sex steroids
were positively correlated with advancing devel-
opmental stages of ovaries (Freund et al., 1995).
Plaice (Pleuronectes platessa) and flounder (Pla-
tichthys flesus) hybrid females had comparable
levels of sex steroids in both diploids and triploids
(Lincoln, 1981a).
In contrast to females, male triploid fish have
shown normal levels of sex hormones in most spe-
cies except Indian catfish. A key enzyme in steroid
biosynthesis, steroid dehydrogenase (3b-HSD) and
immunoreactive testosterone was localized in ste-
roidogenic cells of male tilapia (Koteeswaran et al.,
1995). Male tilapia also had normal levels of tes-
tosterone similar to diploids (Hussain et al., 1995).
Triploid males of Indian catfish showed reduced
levels of maturational gonadotropin (GtH-II) and
sex steroids in plasma (Tiwary et al., 2001, Figure 2).
11-ketotestosterone is a prominent male sex steroid
in fish. Further study on this male specific hormone
in triploid fish may show the impact of triploidy on
reproductive endocrinology in male fish.
Gonadal development in triploids
Triploid males generally show more gonadal devel-
opment than females. This may be due to fact that
triploidy does not interfere with many mitotic divi-
sions required in testes (Thorgaard, 1983). Triploid
males in various species have normal development of
testes resulting in large testes with fully active ste-
roidogenic cells (Benfey, 1999). However, impaired
gonadal development was reported in triploid male
carp (Gervai et al., 1980) and Indian catfish (Tiwary
et al., 2000). There were significant increases in
size of secondary spermatocytes and spermatids in
triploid rainbow trout (Lincoln and Scott, 1984)
and spermatogonia in coho salmon (Piferrer et al.,
1994b) when compared to diploid fish. Mature
spermatozoa were not observed in triploid channel
catfish (Wolters et al., 1982b), Atlantic salmon
(Benfey and Sutterlin, 1984a) and loach (Misgurnus
anguillicaudatus) (Suzuki et al., 1985) despite a
well-developed testes. Most frequently, the testes of
rainbow trout had spermatocytes and morphologi-
cally abnormal spermatozoa during final stages of
spermatogenesis (Carrasco et al., 1998).
The impaired development of oocytes and
corresponding decrease in ovaries may be due to
meiotic failure in triploid females (Figure 3). The
impact of triploidy on ovarian development is well
studied in coho salmon (Piferrer et al., 1994b),
rainbow trout (Lincoln and Scott, 1984; Nakamura
et al., 1987a, b), masu salmon (Oncorhynchus
masou) (Nakamura et al., 1993), plaice (Purdom,
1972; Lincoln, 1981b), channel catfish (Wolters
et al., 1982b), European catfish (Krasznai and
Marian, 1986), and Indian catfish (Tiwary et al.,
2000). Salmonid studies have shown lack of oocyte
maturation in female triploids at different stages of
development (Thorgaard and Gall, 1979; Lincoln
and Scott, 1983, 1984; Yamazaki, 1983; Chevassus
et al., 1984; Solar et al., 1984; Solar and Donaldson,
1985; Piferrer et al., 1994b). However, some oocyte
development has been reported in rainbow trout,
 397

25 when compared to diploids during pre-spawning

2n
20
phase of the reproductive cycle (Figure 4). Impaired
3n
oocyte development in triploid females may be due
15 to inadequate release of vitellogenin (Benfey et al.,
GSI

10 * 1989a, b). Thus, triploid fish have a unique brain-

* pituitary-gonad axis, which is affected at each level
*
5
of endocrine hierarchy (Figure 5).
0
Resting Preparatory Prespawning Spawning

Figure 3. Gonadosomatic index (GSI) in diploid (2n) and

Future prospects in triploid research
triploid (3n) female catfish (H. fossilis) during different phases
of reproductive cycle. (* indicates P < 0.001, 2n–3n). Triploid fish have a great potential for commercial
exploitation in aquaculture and as a model for
plaice and coho salmon (Purdom, 1972; Johnson
et al., 1986). Gonadal development was also signif-
icantly reduced in triploid female carp (Gervai et al., Environmental Cues
1980) and tilapia (Chang et al., 1993). Impaired
oocyte development has also been reported in trip-
loid female channel catfish, European catfish and ?
Indian catfish (Wolters et al.,1982b; Krasznai and
Marian, 1986; Tiwary et al., 2000, Figure 4) but not (-)
in African catfish (Richter et al.,1994). Triploid
BRAIN
Indian catfish showed numerous atretic follicles

Chromatin
nucleolar stage Low GnRH
Perinucleolar (-)
stage (-)
Yolk vesicle PITUITARY
stage
Vitellogenic
stage
Low GtH
Ripe stage
(-)
Atretic stage
LIVER
2n
Chromatin (Vitellogenesis)
nucleolar
stage
Perinucleolar
Low Vitellogenin
stage

Yolk vesicle OVARY

stage
(Impaired Oogenesis)
Vitellogenic
stge

Ripe stage
3n
Figure 4. Proportion (%) of different stages of oocyte in ovary
Low Estradiol 17
of triploid (3n) and diploid (2n) catfish (H. fossilis) during Figure 5. Schematic diagram showing the effect of triploidy on
spawning phase of reproductive cycle. reproductive endocrinology in female fish.
398
basic research. New methods for inducing trip-
loidy in fish may be forthcoming. This may help in
alleviating the problem of morpho-anatomical
changes encountered in some species. Triploid
detection methods should have a combination of
precision and cost-effectiveness. Research should
be conducted to determine a simple but accurate
method for detection of triploidy in fish.
Growth performance in triploid fish is a var-
iable factor in different species. Other culture
species for cultivation may be utilized for the
induction of triploidy and show benefits of
enhanced growth performance. This may open up
new avenues for commercial exploitation of
triploid fish in aquaculture. The public health
aspects of triploid food fish production must be
evaluated before their introduction into com-
mercial aquaculture. Studies on growth factors
such as growth hormones, insulin and insulin-like
growth factors (IGFs) may help further in eluci-
dating the mechanism of variable growth in trip-
loid fish.
Since the triploid fish have abnormal physio-
logical features, they provide a unique potential
for discovery of new biochemical and molecular
mechanisms in fish physiology. Fish have a vital
step in evolution of higher vertebrates. Physio-
logical studies using triploid fish may be able to
help in focus more light on biochemical and
molecular evolutionary processes. Triploid fish
show some behavioral alterations when compared
to diploids. The structure of neurones and nature
of neuropeptides and neurotransmitters in their
brain may give new clues to workings of neural
control which results in the altered behavior in
triploid fish. The relationship between shock and
morpho-anatomical changes in some species may
be clarified by studies involving expression of
homeotic genes.
Conclusion
There are many research opportunities investigat-
ing the biology of triploid fish. Researchers having
backgrounds in fish physiology, biochemistry,
molecular biology, structural biology, neurobiol-
ogy and behavioral science have an opportunity to
contribute significantly to development of triploid
fish research.
References
Aliah, R.S., Yamaoka, K., Inada, Y. and Taniguchi, N. (1990)
Effects of triploidy on tissue structure of some organs of ayu.
Nippon Suisan Gakkaishi 56, 569–575.
Aliah, R.S., Inada, Y., Yamaoka, K. and Taniguchi, N. (1991)
Effects of triploidy on haematological characterstics and
oxygen consumption of ayu. Nippon Suisan Gakkaishi 57,
833–836.
Allen, S.K., Jr. and Stanley, J.G. (1983) Ploidy of hybrid grass
carp · bighead carp determined by flow cytometry. Trans.
Am. Fish. Soc. 112, 431–435.
Al-Sabti, K. (1995) Detection of triploidy in fish using the
cytokinesis-blocked method for erythrocyte and hepatic cells.
Cytobios 82, 181–187.
Amano, M., Aida, K., Okumoto, N. and Hasegawa, Y. (1992)
Changes in salmon GnRH and chicken GnRH-II contents in
the brain and pituitary, and GtH contents in the pituitary in
female masu salmon, Oncorhynchus masou, from hatching
through ovulation. Zool. Sci. 9, 375–386.
Anglade, I., Pakdel, F., Bailhache, T., Petit, F., Salbert, G.,
Jego, P., Valotaire, Y. and Kah, O. (1994) Distribution of
estrogen receptor-immunoreactive cells in the brain of the
rainbow trout (Oncorhynchus mykiss). J. Neuroendocrinol. 6,
573–583.
Atteke, C., Vetillard, A., Fostier, A., Garnier, D.H., Jego, P.
and Bailhache, T. (2003) Effects of progesterone and estradiol
on the reproductive axis in immature diploid and triploid
rainbow trout. Comp. Biochem. Physiol. 134A, 693–705.
Barker, C.J., Beck, M.L. and Biggers, C.J. (1983) Haematologic
and enzymatic analysis of Ctenopharyngodon idel-
la · Hypophthalmichthys nobilis F1 hybrids. Comp. Biochem.
Physiol. 74A, 915–918.
Benfey, T.J. (1999) The physiology and behaviour of triploid
fishes. Rev. Fish. Sci. 7, 39–67.
Benfey, T.J. and Sutterlin, A.M. (1984a). Growth and gonadal
development in triploid landlocked Atlantic salmon (Salmo
salar). Can. J. Fish. Aquat. Sci. 41, 1387–1392.
Benfey, T.J. and Sutterlin, A.M. (1984b) The haematology of
triploid and landlocked Atlantic salmon, Salmo salar L. J.
Fish Biol. 24, 333–338.
Benfey, T.J., Dye, H.M., Solar, I.I. and Donaldson, E.M.
(1989a) The growth and reproductive endocrinology of
adult triploid Pacific salmonids. Fish Physiol. Biochem. 6,
113–120.
Benfey, T.J., Dye, H.M. and Donaldson, E.M. (1989b) Estro-
gen induced vitellogenin production by triploid coho salmon
(Oncorhynhus kisutch), and its effect on plasma and pituitary
gonadotropin. Gen. Comp. Endocrinol. 75, 83–87.
Biron, M. and Benfey, T.J. (1994) Cortisol, glucose and
hematocrit changes during acute stress, cohort sampling, and
the diet cycle in diploid and triploid brook trout (Salvelinus
fontinalis Mitchill). Fish Physiol. Biochem. 13, 153–160.
Bonar, S.A., Thomas, G.L. and Pauley, G.B. (1988) Evaluation
of the separation of triploid and diploid grass carp, Cteno-
pharyngodon idella (Valanciennes), by external morphology.
J. Fish Biol. 33, 895–898.
Boulanger, Y. (1987) Preliminary results on the growth of sterile
rainbow trout (triploidy induced by a pressure shock) in fish
culture. Proc. Ann. Meet. Aquacult. Assoc. Can. 1, 55–58.

Bramick, U., Puckhaber, B., Langholz, H.J. and Horstgen-
Schwark, G. (1995) Testing of triploid tilapia (Oreochromis
niloticus) under tropical pond conditions. Aquaculture 137,
343–353.
Breton, B. and Sambroni, E. (1996) Steroid activation of the
brain-pituitary complex gonadotropic function in the triploid
rainbow trout, Oncorhynchus mykiss. Gen. Comp. Endocrinol.
101, 155–164.
Carrasco, L.A., Doroshov, S., Penman, D.J. and Bromage, N.
(1998) Long term, quantitative analysis of gametogenesis in
autotriploid rainbow trout, Oncorhynchus mykiss. J. Reprod.
Fert. 113, 197–210.
Carter, C.G., McCarthy, I.D., Houlihan, D.F., Johnstone, R.,
Walsingham, M.V. and Mitchell, A.I. (1994) Food consump-
tion, feeding behaviour, and growth of triploid and diploid
Atlantic salmon, Salmo salar L., parr. Can. J. Zool. 72, 609–617.
Cassani, J.R., Caton, W.E. and Clark, B. (1984) Morphological
comparisons of diploid and triploid hybrid grass carp,
Ctenopharyngodon idella female · Hypophthalmichthys nobi-
lis male. J. Fish Biol. 25, 269–278.
Chang, S.L., Chang, C.F. and Liao, I.C. (1993) Comparative
study on the growth and gonadal development of diploid and
triploid tilapia, Oreochromis aureus. J. Taiwan Fish. Res. 1(1),
43–49.
Chao, N.H., Li, F.G., Huang, C.F., Yang, C.P., and Liao,
I.C. (1993). Investigation on artificially induced triploidy in
common carp using flow cytometer. J. Taiwan Fish. Res. 1 (2),
39–54.
Cherfas, N.B. and Ilyasova, V.A. (1980) Induced gynogenesis in
silver crucian carp and carp hybrids. Genetika 16, 1260–1269.
Chevassus, B., Quillet, E. and Chourrout, D. (1984) La
production de truites steriles par voie genetique. Piscic. Fr.
78, 10–19.
Choubert, G. and Blanc, J.M. (1989) Dynamics of dietary
canthaxanthin utilization in sexually maturing female rain-
bow trout (Salmo gairdneri Rich.) compared to triploids.
Aquaculture 83, 359–366.
Deeley, M.A. and Benfey, T.J. (1995) Learning ability of
triploid brook trout. J. Fish Biol. 46, 905–907.
Ewing, R.R., Scalet, C.G. and Evension, D.P. (1991) Flow
cytometric identification of larval triploid walleyes. Prog.
Fish-Cult. 53, 177–180.
Fast, A.W., Pewnim, T., Keawtabtim, R., Saijit, R., Te, F.T.
and Vejaratpimol, R. (1995) Comparative growth of diploid
and triploid Asian catfish Clarias macrocephalus in Thailand.
J. World Aquacult. Soc. 26, 390–395.
Fauconneau, B., Kaushik, S.J. and Blanc, J.M. (1986) Utilisa-
tion de differents substrates energetiques chez la truite:
influence de la ploide. Diab. Metabol. 12, 111–112.
Fauconneau, B., Kaushik, S.J. and Blanc, J.M. (1989) Uptake
and metabolization of dissolved compounds in rainbow
trout (Salmo gairdneri R.) fry. Comp. Biochem. Physiol.
93A, 839–843.
Fauconneau, B., Aguirre, P. and Blanc, J.M. (1990). Protein
synthesis in different tissues of mature rainbow trout (Salmo
gairdneri R.). Influence of triploidy. Comp. Biochem. Physiol.
97C, 345–352.
Freund, F., Hoerstgen-Schwark, G. and Holtz, W. (1995)
Plasma steroid hormones in adult triploid tilapia (Oreochr-
omis niloticus). In: Goetz, F.W. and Thomas, P. (eds.),
Proceedings of the Fifth International Symposium on the
399
Reproductive Physiology of Fish. The University of Texas,
Austin, TX, p. 116.
Galbreath, P.F. and Thorgaard, G.H. (1995) Salt-water per-
formance of all female triploid Atlantic salmon. Aquaculture
138, 77–85.
Gentile, F., Lira, O. and Marcano-de cotte, D. (1986)
Relationship between brain gonadotropin-releasing hormone
(GnRH) and seasonal reproductive cycle of the caribe Colorado
Pygocentrus notatus. Gen. Comp. Endocrinol. 64, 239–245.
Gervai, J., Peter, S., Nagy, A., Horvath, L. and Csanyi, V.
(1980) Induced triploidy in carp, Cyprinus carpio L. J. Fish
Biol. 17, 667–671.
Gomelsky, B.I., Emelyanova, O.V. and Recoubratsky, A.V.
(1992) Application of the scale cover gene (N) to identifica-
tion of type of gynogenesis and determination of ploidy in
common carp. Aquaculture 106, 233–237.
Goudie C.A., Simco, B.A., Davis, K.B. and Liu, Q. (1995)
Production of gynogenetic and polyploid catfish by pressure-
induced chromosome set manipulation. Aquaculture 133,
185–198.
Graham, M.S., Fletcher, G.L. and Benfey, T.J. (1985) Effect of
triploidy on blood oxygen content of Atlantic salmon.
Aquaculture 50, 133–139.
Henken, A.M., Brunink, A.M. and Richter, C.J.J. (1987)
Differences in growth rate and feed utilization between
diploid and triploid African catfish, Clarias gariepinus
(Burchell 1822). Aquaculture 63, 233–242.
Hussain, M.G., Chatterji, A., McAndrew, B.J. and Johnstone,
R. (1991) Triploidy induction in Nile tilapia, Oreochromis
niloticus L. using pressure, heat and cold shocks. Theor. Appl.
Genet. 81, 6–12.
Hussain, M.G., Rao, G.P.S., Humayun, N.M., Randall, C.F.,
Penman, D.J., Kime, D., Bromage, N.R., Myers, J.M. and
McAndrew, B.J. (1995) Comparative performance of growth,
biochemical composition and endocrine profiles in diploid
and triploid tilapia, Oreochromis niloticus L. Aquaculture 138,
87–95.
Hyndman, C.A., Kieffer, J.D. and Benfey, T.J. (2003) The
physiological response of diploid and triploid brook trout to
exhaustive exercise. Comp. Biochem. Physiol. 134A, 167–179.
Inada, Y. and Taniguchi, N. (1992) Studies on jumping,
territorial and algae feeding behaviors of induced triploid
ayu Plecoglossus altivelis. Suisanzoshoku 40, 99–104.
Johnson, O.W., Dickhoff, W.W. and Utter, F.M. (1986)
Comparative growth and development of diploid and triploid
coho salmon, Oncorhynchus kisutch. Aquaculture 57, 329–336.
Johnstone, R., Knott, R.M., McDonald, A.G. and Walsingham,
M.V. (1989) Triploidy induction in recently fertilized Atlantic
salmon ova using anaesthetics. Aquaculture 78, 229–236.
Johnston, I.A., Strugnell, G., McCracken, M.L. and Johnstone,
R. (1999) Muscle growth and development in normal sex-
ratio and all-female diploid and triploid Atlantic salmon.
J. Exp. Biol. 202, 1991–2016.
Jungalwalla, P.J. (1991) Production of non-maturing Atlantic
salmon in Tasmania. Can. Tech. Rep. Fish. Aquat. Sci. 1789,
47–71.
Kavumpurath, S. and Pandian, T.J. (1992a). Effects of induced
triploidy on aggressive display in the fighting fish, Beta
splendens Regan. Aquacult. Fish. Mgmt. 23, 281–290.
Kavumpurath, S. and Pandian, T.J. (1992b) The development
of all-male sterile triploid fighting fish (Beta splendens Regan)
400
by integrating hormonal sex reversal of broodstock and
chromosome-set manipulation. Isr. J. Aquacult. 44, 111–119.
Kerby, J.H., Eversion, J.M., Harrell, R.M., Starling, C.C.,
Ravels, H. and Geiger, J.G. (1995) Growth and survival
comparisons between diploid and triploid sunshine bass.
Aquaculture 137, 355–358.
Kim, D.S., Jo, Y.Y. and Lee, T.Y. (1994) Induction of triploidy
in mud loach (Misgurnus mizolepis) and its effect on gonad
development and growth. Aquaculture 120, 263–270.
Kobayashi,T., Fushiki, S., Sakai, N., Hara, A., Amano, M.,
Aida, K., Nakamura, M. and Nagahama, Y. (1998) Oogen-
esis and changes in the levels of reproductive hormones in
triploid female rainbow trout. Fish. Sci. 64, 206–215.
Koteeswaran, R., Sheela, S.G. and Pandian, T.J. (1995)
Testosterone biosynthesis in triploid sterile male tilapia.
Curr. Sci. 69, 545–547.
Krasznai, Z. and Marian, T. (1986) Shock induced triploidy
and its effect growth and gonadal development of the
European catfish Silurus glanis L. J. Fish Biol. 29, 519–527.
Krieger, M.J. and Keller, L. (1998) Estimation of the propor-
tion of triploids in populations with diploid and triploid
individuals. J. Hered. 89, 275–279.
Lamatsch, D.K., Steinlein, C., Schmid, M. and Schartl, M.
(2000) Non-invasive determination of genome size and ploidy
level in fishes by flow cytometry detection of triploid Poecilia
formosa. Cytometry 39, 91–95.
Leary, R.F., Allendorf, F.W., Knudsen, K.L. and Thorgaard,
G.H. (1985) Heterozygosity and developmental stability in
gynogenetic diploid and triploid rainbow trout. Heredity 54,
219–225.
Lincoln, R.F. (1981a) The growth of female diploid and triploid
plaice (Pleuronectes platessa) · flounder (Platichthys flesus)
hybrids over one spawning season. Aquaculture 25, 259–268.
Lincoln, R.F. (1981b) Sexual maturation in female triploid
plaice, Pleuronectes platessa, and plaice · flounder, Platich-
thys flesus, hybrids. J. Fish Biol. 19, 499–507.
Lincoln, R.F. and Scott, A.P. (1983) Production of all-female
triploid rainbow trout. Aquaculture 30, 375–380.
Lincoln, R.F. and Scott, A.P. (1984) Sexual maturation in
triploid rainbow trout, Salmo gairdneri Richardson. J. Fish
Biol. 25, 385–392.
Linhart, O. and Flajshans, M. (1995) Triploidization of
European catfish, Silurus glanis L., by heat shock. Aquacult.
Res. 26, 367–370.
Linhart, O., Flajshans, M. and Kvasnicka, P. (1991). Induced
triploidy in the common carp (Cyprinus carpio L.): a
comparison of two methods. Aquat. Liv. Res. 4, 139–145.
Liu, S., Sezaki, D., Hashimoto, K., Kobayashi, H. and
Nakamura, M. (1978) Simplified techniques for determina-
tion of polyploidy in ginbuna, Carassius auratus langsdorfi.
Bull. Jpn. Soc. Sci. Fish. 44, 601–606.
Lou, Y.D. and Purdom, C.E. (1984) Polyploidy induced by
hydrostatic pressure in rainbow trout, Salmo gairdneri
Richardson. J. Fish Biol. 25, 345–351.
Malison, J.A., Procarione, L.S., Held, J.A., Kayes, T.B. and
Amudson, C.H. (1993) The influence of triploidy and heat
and hydrostatic shocks on the growth and reproductive
development of juvenile yellow perch (Perca flavescens).
Aquaculture 116, 121–133.
McCarthy, I.D., Carter, C.G., Houlihan, D.F., Johnstone, R.
and Mitchell, A.I. (1996) The performance of all female
diploid and triploid Atlantic salmon smolts on transfer
together to sea water. J. Fish Biol. 48, 545–548.
McGeachy, S.A., Benfey, T.J. and Friars, G.W. (1995) Fresh-
water performance of triploid Atlantic salmon (Salmo salar)
in New Brunswick aquaculture. Aquaculture 137, 333–341.
McGeachy, S.A., O’Flynn, F.M., Benfey, T.J. and Friars, G.W.
(1996) Saltwater performance of triploid Atlantic salmon in New
Brunswick aquaculture, Bull. Aquacult. Assoc. Can. 96, 24–28.
Nakamura, M., Nagahama, Y., Iwahashi, M. and Kojima, M.
(1987a). Ovarian structure and plasma steroid hormones in
triploid female rainbow trout. Nippon Suisan Gakkaishi 53,
1105.
Nakamura, M., Nagahama, Y., Iwahashi, M. and Kojima, M.
(1987b). Reproduction of triploid rainbow trout, Salmo
gairdneri. Oh Tsu Chi Marine Biological Laboratory Report
(University of Tokyo) 12, 167–171 (in Japanese).
Nakamura, M., Tsuchiya, F., Iwahashi, M. and Nagahama, Y.
(1993) Reproductive characteristics of a precociously mature
triploid male masu salmon (Oncorhynchus masou). Zool. Sci.
10, 117–125.
Okada, H. (1985) Studies on the artificial sex control in rainbow
trout, Salmo gairdneri. Sci. Rep. Hokkaido Fish Hatchery 40,
1–49.
Oliva-Teles, A. and Kaushik, S.J. (1987a) Nitrogen and energy
metabolism during the early ontogeny of diploid and triploid
rainbow trout (Salmo gairdneri R.). Comp. Biochem. Physiol.
87A, 157–160.
Oliva-Teles, A. and Kaushik, S.J. (1987b) Metabolic utilization
of diets by polyploid rainbow trout (Salmo gairdneri). Comp.
Biochem. Physiol. 88A, 45–47.
Oliva-Teles, A. and Kaushik, S.J. (1990a) Effect of temperature
on utilization of endogenous energy reserves during embry-
onic development of diploid and triploid rainbow trout
(Salmo gairdneri R.). Aquaculture 84, 373–382.
Oliva-Teles, A. and Kaushik, S.J. (1990b) Growth and nutrient
utilization by 0+ and 1+ triploid rainbow trout, Oncorhyn-
cus mykiss. J. Fish Biol. 37, 125–133.
Parsons, G.R. (1993) Comparisons of triploid and diploid white
crappies. Trans. Am. Fish. Soc. 122, 237–243.
Piferrer, F., Benfey, T.J. and Donaldson, E.M. (1994a)
Production of female triploid coho salmon (Oncorhynchus
kisutch) by pressure shock and direct estrogen treatment.
Aquat. Liv. Res. 7, 127–131.
Piferrer, F., Benfey, T.J. and Donaldson, E.M. (1994b)
Gonadal morphology of normal and sex reversed triploid
and gynogenetic diploid coho salmon (Oncorhynchus kisutch).
J. Fish Biol. 45, 541–553.
Purdom, C.E. (1972) Induced polyploidy in plaice (Pleuronectes
platessa) and its hybrid with the flounder (Platichthys flesus).
Heredity 29, 11–24.
Purdom, C.E. (1993) Genetics and Fish Breeding, Chapman &
Hall, London.
Refstie, T., Vaasvic, V. and Gjedrem, T. (1977) Induction of
polyploidy in salmonids by Cytochalasin B. Aquaculture 10,
65–74.
Richter, C.J.J., Eding, E.H., Verreth, J.A.J. and Fleuren,
W.L.G. (1994) African catfish. In: Bromage, N. and Roberts,
R.J. (eds.), Broodstock Management and Egg and Larval
Quality. Blackwell Science, Oxford, pp. 242–276.
Schafhauser-Smith, D. and Benfey, T.J. (2003a) The effects of
long term estradiol 17 beta treatment on the growth and

physiology of female triploid brook trout (Salvelinus fonti-
nalis). Gen. Comp. Endocrinol. 131, 9–20.
Schafhauser-Smith, D. and Benfey, T.J. (2003b) In vitro steroid
production by triploid ovarian follicles. Gen. Comp. Endo-
crinol. 133, 279–286.
Sezaki, K., Watabe, S. and Hashimoto, K. (1983) A compar-
ison of chemical composition between diploids and triploids
of ‘‘ginbuna’’ Carassius auratus langsdorfi. Bull. Jpn. Soc. Sci.
Fish. 49, 97–101.
Sezaki, K., Watabe, S. and Hashimoto, K. (1988) Haemato-
logical parameters and erythrocyte enzyme activities associ-
ated with increase in ploidy status of the spinous loach,
Cobitis biwae Jordon & Snyder. J. Fish Biol. 32, 149–150.
Sezaki, K., Watabe, S., Tsukamoto, K. and Hashimoto, K.
(1991) Effects of increase in ploidy status on respiratory
function of ginbuna, Carassius auratus langsdorfi (Cyprini-
dae). Comp. Biochem. Physiol. 99A, 123–127.
Small, S.A. and Benfey, T.J. (1987) Cell size in triploid salmon.
J. Exp. Zool. 241, 339–342.
Small, S.A. and Randall, D.J. (1989) Effects of triploidy on the
swimming performance of coho salmon (Oncorhynchus kis-
utch). Can. J. Fish. Aquat. Sci. 46, 243–245.
Smith, L.T. and Lemoine, H.L. (1979) Colchicine-induced
polyploidy in brook trout. Prog. Fish-Cult. 41, 86–88.
Solar, I.I. and Donaldson, E.M. (1985) Studies on genetic and
hormonalsex control in domesticated rainbow trout. I. The effect
of heat shock treatment for induction of triploidy in cultured
rainbow trout. Can. Tech. Rep. Fish. Aquat. Sci. 1379, 15pp.
Solar, I.I., Donaldson, E.M., and Hunter, G.A. (1984) Induction
of triploidy in rainbow trout (Salmo gairdneri Richardson) by
heat shock and investigations on early growth. Aquaculture
42, 57–67.
Stillwell, E.J. and Benfey, T.J. (1994) Haemoglobin level,
metabolic rate and swimming performance in triploid brook
trout (Salvelinus fontinalis). In: MacKinlay, D.D. (ed.) High
Performance Fish, Fish Physiology Association. Vancouver,
BC, July 16–21, 1994. pp. 288–293.
Stillwell, E.J. and Benfey, T.J. (1996) Haemoglobin level,
metabolic rate, opercular abduction rate and swimming
efficiency in female triploid brook (Salvelinus fontinalis). Fish
Physiol. Biochem. 15, 377–383.
Streisinger, G., Walker, C., Dower, N., Knauber, D. and
Singer, F. (1981) Production of clones of homozygous diploid
zebra fish (Brachydanio rerio). Nature 291, 293–296.
Sugama, K., Taniguchi, N., and Seki, S. (1992) Survival,
growth and gonad development of triploid red sea bream,
Pagrus major (Temmink et Schlegel): use of allozyme marker
for ploidy and family identification. Aquacult. Fish. Mgmt.
23, 149–159.
Sumpter, J.P., Scott, A.P., Baynes, S.M. and Witthame, P.R.
(1984) Early stages of the reproductive cycle in virgin female
rainbow trout (Salmo gairdneri Richardson). Aquaculture 43,
235–242.
Sutterlin, A.M. Holder, J. and Benfey, T.J. (1987) Early survival
rates andsubsequent morphologicalabnormalities in landlocked
anadromous and hybrid (landlocked · anadromous) diploid
and triploid Atlantic salmon. Aquaculture 64, 157–164.
Sutterlin, A.M. and Collier, C. (1991) Some observation on the
commercial use of triploid rainbow trout and Atlantic salmon
in Newfoundland, Canada. Can. Tech. Rep. Fish. Aquat. Sci.
1789, 89–96.
401
Suzuki, R., Nakanishi, T. and Oshiro, T. (1985) Survival,
growth and sterility of induced triploids in the cyprinid loach,
Misgurnus anguillicaudatus. Bull. Jap. Soc. Sci. Fish. 51, 889–894.
Swarup, H. (1959). Effect of triploidy on the body size, general
organization and cellular structure in Gasterosteus aculeatus
(L.). J. Genet. 56, 143–155.
Tave, D. (1993) Growth of triploid and diploid bighead carp,
Hypophthalmichthys nobilis. J. Appl. Aquacult. 2, 13–25.
Thititananukij, S., Vejaratpimol, R., Pewnim, T. and Fast,
A.W. (1996) Ethidium bromide nuclear staining and fluores-
cence microscopy: an alternative method for triploidy detec-
tion in fish. J. World Aquacult. Soc. 27, 213–217.
Thorgaard, G.H. (1983) Chromosome set manipulation and sex
control in fish. In: Hoar, W.S., Randall, D.J. and Donaldson,
E.M. (eds.), Fish Physiology, Vol. IX, Part B. Academic
Press, New York, pp. 405–434.
Thorgaard, G.H. and Gall, G.A.E. (1979) Adult triploids in a
rainbow trout family. Genetics 93, 961–973.
Thorgaard, G.H., Robinovitch, P.S., Shen, M.W., Gall,
G.A.E., Propp, J. and Utter, F.M. (1982) Triploid rainbow
trout identified by flow cytometry. Aquaculture 29,
305–309.
Tiwary, B.K. and Ray, A.K. (2004) Alterations in air sac and
skeleton of triploid Heteropneustes fossilis. J. Fish Biol. 64,
268–272.
Tiwary, B.K., Kirubagaran, R. and Ray, A.K. (1997) Induction
of triploidy by cold shock in catfish, Heteropneustes fossilis
(Bloch). Asian Fish. Sci. 10, 123–129.
Tiwary, B.K., Kirubagaran, R. and Ray, A.K. (1999) Altered
body shape as a morphometric indicator of triploidy in
Indian catfish, Heteropneustes fossilis (Bloch). Aquacult. Res.
30, 907–910.
Tiwary, B.K., Kirubagaran, R. and Ray, A.K. (2000) Gonadal
development in triploid Heteropneustes fossilis. J. Fish Biol.
57, 1343–1348.
Tiwary, B.K., Kirubagaran, R. and Ray, A.K. (2001) Plasma
levels of Gonadotropin-II and gonadal sex steroids in triploid
catfish, Heteropneustes fossilis (Bloch). Fish Physiol. Biochem.
24, 9–14.
Tiwary, B.K., Kirubagaran, R. and Ray, A.K. (2002) Gona-
dotropin releasing hormone (GnRH) neurones of triploid
catfish, Heteropneustes fossilis (Bloch): an immunocytochem-
ical study. Comp. Biochem. Physiol. 132A, 375–380.
Ueda, T., Sato, R. and Kobayashi, J. (1988) Triploid rainbow
trout induced by high pH multiplied by high calcium. Nippon
Suisan Gakkaishi 54, 2045.
Ueno, K. (1984) Induction of triploid carp and their haema-
tological characterstics. Jap. J. Genet. 59, 585–591.
Valenti, R.J. (1976) Induced polyploidy in Tilapia aurea
(Steindachner) by means of temperature shock treatments.
J. Fish Biol. 7, 519–528.
Varadaraj, K. and Pandian, T.J. (1988) Induction of triploids in
Oreochromis mossambicus by thermal, hydrostatic pressure
and chemical shocks. Proc. Aquacult. Inter. Congress Expo,
pp. 531–535.
Virtanen, E., Forsman, L. and Sundley, A. (1990) Triploidy
decreases the aerobic swimming capacity of rainbow trout
(Salmo gairdneri). Comp. Biochem. Physiol. 96A, 117–121.
Wattendorf, R.J. (1986) Rapid identification of triploid grass
carp with coulter counter and channelyzer. Prog. Fish-Cult.
48, 125–132.
 402
Wiley, M.J. and Wike, L.D. (1986) Energy balances of diploid,
triploid, and hybrid grass carp. Trans. Amer. Fish. Soc. 115,
853–863.
Withler, R.E., Beacham, T.D., Solar, I.I. and Donaldson, E.M.
(1995) Freshwater growth, smolting and marine survival and
growth of diploid and triploid coho salmon (Oncorhynchus
kisutch). Aquaculture 136, 91–107.
Wolters, W.R., Chrisman, C.L. and Libey, G.S. (1982a)
Erythrocyte nuclear measurements of diploid and triploid
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Channel catfish, Ictalurus punctatus. J. Fish Biol. 20,
253–258.
Wolters, W.R., Libey, G.S. and Chrisman, C.L. (1982b) Effects
of triploidy on growth and gonadal development of Channel
catfish. Tans. Am. Fish. Soc. 111, 102–105.
Yamamoto, A. and Iida, T. (1994) Haematological characters-
tics of triploid rainbow trout. Fish Pathol. 29, 239–243.
Yamazaki, F. (1983) Sex control and manipulation in fish.
Aquaculture 33, 329–354.