ELSEVIER Aquaculture 137 (1995) 313-322

Growth performance trials of genetically male

tilapia (GMT) derived from YY-males in
Oreochromis niloticus L.: On station comparisons
with mixed sex and sex reversed male populations
Graham C. Mair a,*, Jose S. Abucay b, John A. Beardmore , David
O.F. Skibinski a
~School ofBiological Sciences, Universiry College of Swansea, Swansea SA2 8PP, UK
Freshwater Aquaculture Center, Central Luzon State University, Nueva Ecija 3I20, Philippines

Abstract

Genetic manipulation of predominantly monofactorial sex determining mechanisms in the Nile tilapia
Oreochromis niloticus is proposedas a technology for the mass production of all-male progeny for
culture. Existing alternative methods for producing all-male tilapia, principally direct hormonal
masculinization, have produced inconsistent results and are largely the preserve of scientists and
developed country aquaculturists. Successful adoption of these techniques in developing countries is
seriously limited by the number and complexity of factors involved in their effectiveness.
Novel YY-male genotypes have been shown to be viable and to produce all- or nearly all-male
(XY) progeny in crosses with normal XX females. This paper presents the results from a series of
on-station growth performance trials of these genetically male tilapia (GMT) in three environments.
Two comprehensive and replicated trials in extensively managed earthen ponds, compared the per-
formance of GMT with sex reversed male (SRT) and normal mixed sex tilapia (MST). Three further,
smaller experiments, compared the growth of GMT with that of MST populations in rice-cum-fish
culture and under intensive culture in closed recirculating systems. These comparisons were made
within the Egypt-UCS strain of 0. niloticus.
Results indicate that culture of GMT populations can improve marketable yields under all environ-
ments. In the first experiment in ponds, GMT had significantly (P= 0.003) higher yields than both
MST ( + 58.8%) and SRT ( + 3 1.03%). The second experiment, following the same design, showed
a similar trend, although differences were smaller and not significant. GMT produced yields 11.9%
greater than MST in rice-cum-fish culture and 33.6-47.0% greater in recirculating systems. There
was no recruitment in GMT pond populations whereas recruits were present in MST and SRT
populations. Coefficient of variance for individual weight in ponds was significantly (P <0.05)

* Corresponding author

0044.8486/95/$09.50 0 1995 Elsevier Science B.V. All rights reserved

SSD10044-8486(95)01 I IO-2
314 G.C. Mair et al. /Aquaculture 137 (1995) 313-322

lower for GMT than for MST for both pond experiments. GMT produced from novel YY-males in
this strain has considerable potential for increasing yields from tilapia culture.

Ke.yword.s: Oreochromis niloticus; Growth; Sex control; YY-males

1. Introduction

Early sexual maturation resulting in unwanted reproduction and overcrowding has long
been accepted as a major limitation in the culture of most tilapia species, particularly the
commonly cultured Oreochromis niloticus and 0. mossambicus. This unwanted reproduc-
tion generally results in suppression of growth and reduction in yields in cultured popula-
tions. Males, and especially females, divert energy which could be utilised for somatic
growth, into gamete production and behavioural interactions. In addition, competition with
recruits in confined environments further suppresses the growth of stocked fish and can
result in 30-50% of harvested biomass consisting of largely unmarketable recruits.
Numerous solutions to this problem have been proposed including manual sexing and
separation of the sexes, culture in cages, controlled use of predator species, production of
monosex hybrids, and direct hormonal sex reversal (Mair and Little, 1991). The latter two
technologies of hybridisation and sex reversal have become the most widely used and are
successfully applied by many farmers. However, both technologies have significant disad-
vantages, particularly for application in developing countries. Hybridisation necessitates
loss of species purity and introgression of cultured strains which can combine less desirable
characteristics from one of the parent species. This method was used for some time in
commercial production of all- or nearly all-male tilapia in Israel but is now less commonly
used due to inconsistencies in the proportions of males in the hybrid progeny (Wohlfarth,
1994). Today, commercial production of all-male tilapia is more commonly achieved
through hormonal sex reversal and techniques have been developed and described for
conducting this on a commercial scale (Guerrero and Guerrero, 1988; Popma and Green,
1990). Mair and Little ( 1991) listed the numerous factors involved in consistently obtaining
successful androgen sex reversal. Failure to optimise the necessary parameters such as age
and size of fry and feeding frequency, can result in lower than expected rates of sex reversal,
as is commonly observed in commercial hatcheries in developing countries.
Comprehensive studies on the genetics of sex determination in 0. niloticus (as reviewed
by Mair et al., 199 1 and Trombka and Avtahon, 1993) led to the conclusion that this species
exhibits predominantly monofactorial sex determination with homogametic females (XX)
and heterogametic males (XY). Based on this hypothesis, a model was proposed for the
production of monosex male 0. niloticus through the production of the novel YY-male
genotypes (Scott et al., 1989). This technology was developed and tested in the Philippines
under a collaborative research project between the University of Wales, Swansea and the
Freshwater Aquaculture Center of Central Luzon State University (Mair et al., 1993).
Under this research project YY-males in an Egyptian strain of 0. niloticus were demon-
strated to produce 67-100% male populations (with a mean of 96.5% male) and could be
mass produced in YY X YY crosses, indicating the feasibility of application of this tech-
nology on a commercial scale.
 G. Mair et al. /Aquaculture 137 (1995) 313-322 315

Considering the large volume of literature on techniques of hybridisation and sex reversal
for the production of monosex male tilapia, there are relatively few published studies
comparing the growth performance and yields of these populations compared to normal
mixed sex tilapia, under culture. A few studies have demonstrated the enhanced yield of
monosex male populations created by hybridisation, sex reversal and manual sexing (Han-
son et al., 1983; Wohlfarth et al., 1983; McAndrew and Majumdar, 1989; Vera Cruz and
Mair, 1994) under experimental conditions. In an evaluation of growth of 0. niloticus raised
in earthen ponds (Macintosh et al., 1988)) mean harvest body weight and survival of sex-
reversed male populations (93.1% male) were respectively 22.7% and 2.9% greater than
for mixed sex controls (54.5% male), although neither of these values were significant.
There is as yet little published information on the yield characteristics and potential eco-
nomic benefit of these monosex populations under commercial culture conditions.
This study aimed to evaluate the culture performance (in terms of yield, recruitment,
survival, sex ratio, gonadosomatic indices and size variability) of genetically male tilapia
(GMT) derived from YY-males, compared with that of normal mixed sex tilapia (MST)
and hormonally sex reversed male tilapia ( SRT) . Evaluations were carried out in a range
of culture environments including those approximating to tilapia farming systems in the
Philippines. All comparisons were made within the same Egyptian strain, in which the YY-
male technology has been developed.

2. Materials and methods

Origin of stocks: The Egyptian strain, known in our strain nomenclature as Egypt-
Swansea, in which the YY-male technology has been developed, and from which all GMT
are derived, originates from L. Manzala, N. Egypt. The strain was collected and transferred
to the University of Stirling in 1979. Several introductions were made to create a laboratory
strain at the University of Wales, Swansea in the 1980s. Several adults and several hundred
fry were transferred to the Philippines in 1989. This strain has passed through a number of
genetic bottlenecks and is considered to be inbred. All control fish were derived from crosses
of normal broodstock in the same strain.

2.1. Growth performance trials

The comparison of the performance of mixed-sex and monosex populations necessitates

separate stocking, preferably in replicated culture units, due to interactions that would take
place under communal stocking. Where possible, trials were replicated in three culture units
for each treatment (GMT, MST and SRT) , however limitations of fish or of facilities limited
the number of replicates and/or the number of treatments that could be used in each of the
trials. These trials were undertaken between 1987 and 1994.
The emphasis of this paper is on the culture performance of the three treatments in
replicated, extensively managed ponds, located at Central Luzon State University in the
Philippines. Results from an experiment conducted in a rice-cum-fish culture system in the
Philippines and two experiments in a closed recirculating system at the University of Wales,
Swansea, are also presented.
316 G. Mair et al. /Aquaculture 137 (1995) 313-322

I
Table
Details
of set up of on station performance trials of GMT, SRT and MST

Trial Environment Control Unit Reps. per stocking Management Feeding CUltWe
number size treatment Density period
(m2) (fish m-2) (days)

FAC Ponds I SRTand 200 3 2 Extensive Pond 231

MST fertilization
FAC Ponds 2 SRT and 200 3 2 Extensive Pond 168
MST fertilization
Rice-Fish MST 100 3 1.7 Exten sive Pond 80
fertilization
Recirc. 1 MST 0.78 2b 51 Intensive Trout pellets 136
Recirc. 2 MST 0.78 1 63.6 Intensive Trout pellets 119

a In extensive systems no feeding was applied, relying on primary productivity based on pond fertilization with chicken manure
and inorganic fertilizer at 5 kg N ha- month-. Fish in recirculating systems were fed to satiation.b One replicate of MST was
lost during grow-out due to water management problems.

With the exception of the experiments conducted in recirculating systems, all GMT and
control fish were derived from pooled spawning of YY-male or normal male broodstock,
stocked in fine mesh cages, at a ratio of 1:3 with normal females from the same strain. Fry
or eggs were collected every 5 or 7 days. Incubation for eggs was completed in incubating
trays and all fry on-grown at equivalent densities. Fry for the SRT treatment groups were
sex reversed using minor modifications to the standard protocol for commercial sex reversal
as described by Popma and Green ( 1990) and Vera Cruz and Mair ( 1994). Following
completion of incubation, fry were transferred to circular concrete tanks at a density of 1000
m-*. Fry were treated for 25 days, by application of hormone treated commercial fry diet
containing 40 mg kg- of 17a-methyltestosterone. The maximum age difference for fry
compared in a single trial was 7 days and initial weights were not significantly different for
the treatments in all replicated trials. Fingerlings were stocked in ponds at a mean weight
of approximately 5 g. In the rice-fish experiment, fish were marked by clipping of left or
right pelvic fins, prior to stocking in the fields in order to detect immigration of unmarked
feral fish during the grow-out.
Fish in pond Trials 1 and 2 were not on-grown for a fixed period but to a size considered
as marketable in the Philippines ( 130-190 g wet weight). At the end of each trial all stocked
fish and recruits were harvested although recruits collected from the MST and SRT ponds
in Trial 1 were accidentally mixed before they could be weighed. All fish were counted,
sexed and individually weighed and measured. In Trial 1, 50 males from each treatment
were dissected and their gonads weighed from determination of male gonadosomatic index
(GSI) . All harvested fish were checked for their fin clip markings and recruit number and
biomass was recorded. Recruits represented juveniles from reproduction within the pond
population as unwanted immigrants were considered to be negligible.
Trial 3 was conducted in a rice-cum-fish field utilising the pond refuge system described
by Sevilleja et al. (1992). Fish were harvested from the refuge after rice-harvest with a
short grow-out period of only 80 days.
Two trials (4 and 5) were conducted under conditions of intensive culture in plastic bins
in a closed recirculating system at the Tilapia Genetics Laboratory of the University of
 G. Mair et al. /Aquaculture 137 (1995) 313-322 317

Wales, Swansea. In recirculating systems MST were half-sibs of the GMT, created by
fertilizing a divided batch of eggs from a single female with sperm from a normal and a
YY-male. Fry were stocked immediately after incubation in bins in the same recirculating
system.
Trials 1 and 2 compared the performance of GMT with MST and SRT whilst MST alone
was compared with GMT in Trials 3,4 and 5. Full details of these experiments are presented
in Table 1.

2.2. Data analysis

Data from all replicated trials were analyzed using a two-way ANOVA on treatment and
replicate means. Duncans multiple range test was used to determine significance of differ-
ence (P < 0.05) between treatment means. Percentage data for sex ratio and survival was
transformed using the arcsin square root transformation. Mean harvest weight and length
data from the experiments conducted in recirculating systems were compared using the
students t-test.

3. Results

Table 2 shows the results from the two pond growth trials ( 1 and 2). No data is available
on recruit biomass (and thus total biomass) for Trial 1 and for GSI (not measured) in Trial
2. ANOVA models for sex ratio, were significant in both Trials 1 (P= 0.007) and 2
(P = 0.000). In the first of these trials the sex reversal treatment was very effective, giving
almost identical sex ratios to those observed in the GMT (98.4-98.5% male). In the second
trial sex reversal was less effective, producing only 7 1.6% male progeny, significantly lower
than the GMT (99.6% male). In both experiments, sex ratio of SRT and GMT was signif-
icantly greater than in MST, although the latter was biased to male. The ANOVA model
for survival was significant (P=O.O15) in Trial 1, with GMT having significantly higher
survival than MST. There were no significant differences in survival in Trial 2.
Similar trends were seen for mean harvest weight in the two trials, with significant
differences in both trials when both sexes were considered together (P = 0.003 and P = 0.037
for Trials 1 and 2, respectively). The mean weight of fish in the GMT treatment was
significantly greater than MST in both experiments. In Trial 1, the mean weight of GMT
was also significantly greater than that of SRT. There were significant differences
(P = 0.0 16) for mean harvest weight of males in Trial 1, with GMTmales being significantly
larger than MST and SRT males. There were no significant differences in male weight in
Trial 1, and in female weight in both trials. Trends in length differences were similar to
those for weight.
There were significant differences for GSI (P = 0.044) between the males in the three
treatments, being significantly lower in MST than in SRT. There were no significant dif-
ferences in coefficients of variance (CV) for harvest weight or standard length in Trial 1.
These differences were significant for Trial 2 (P=O.O02), with CV being significantly
higher in MST than in SRT or GMT.
Table 2
c:
Data (means of three replicates) from harvest of repeated on-station performance trials of GMT. SRT and MST in ponds.

e
Trial Treat. Sex ratio Survival (7%)Mean harvest Wt. (g) Male GSI CV of Mean CV of SL Yield (kg) Recruit Total
:
(% male) weight harvest SL (%) Biomass biomass
?
(S) (cm) (kg) (kg)
6&P c?d 99 z
9
6
136.6a 153.7a 80.2a 0.79a 26.24a 15.30 a 9.84a 28.50a a
I MST 75.33a 83.46a N\A N\ A 9
SRT 98.36b 90.27ab 152.5a 152.9a 88.4a 1.58a 16.10b 15.81ab 6.26a 34.38b N\A N\A r;:
GMT 98.46b 96.40b 187.0b 188.7b 54.la 1.19a 17.97ab 16.86b 6.74a 45.05c N\A N\A
34.17a 15.27a 5
2 MST 59.13a 95.20a 138.8a 176.9a 93.6a N\A I 1.23a 33.04a 7.55 40.59
=
SRT 71.61b 93.47a 1576ab 180.5a 100.3a N\A 30.83a 16.03b 10.37a 36.74ab 4.5P 41.24
%
GMT 99.58~ 95.07a 175.7b 1759a 117Sa N\ A 206Ob 16.20b 6.73b 41.7Ob OS@ 41.7P z!

$
Representing marketable yield being that of marked fish that were initially stocked.Within trial values superscripted with different letters are signiticantly different (P < 0.05); &
Analysis not applied to non-replicated treatments.N\A, not applicable or not available. tXl
 G. Mair et al. /Aquaculture 137 (1995) 313-322 319

ElMSI- WSRT n GMT

FAC Ponds 1 FAC Ponds 2 Recirc. 1 Recirc. 2 Rice-fish

Note: Yields from recirculating systems x 10m2

Fig. 1. Yield of marketable fish in performance trials of MST, SRT and GMT within the Egypt-UCS strain, in
earthen ponds, rice-cum-fish and recirculating systems ( X lo-*) culture environments.

Marketable yields were significantly different in both trials (P = 0.000 and P = 0.018 for
Trials 1 and 2 respectively) as illustrated in Fig. 1. Yield of GMT was significantly greater
than both MST and SRT in Trial 1 but GMT and SRT were not significantly different in
Trial 2. There was no recruitment in the GMT stocked ponds in either of the trials despite
the presence of small numbers of females, whereas recruits were recovered from MST and
SRT stocked ponds in both trials. It was interesting to note that females recovered from
GMT populations had failed to attain sexual maturation whereas this had occurred in females
from MST and SRT populations. In Trial 2, where recruits were weighed, there were no
significant differences in recruit biomass between the three treatments. When recruit biomass
was added to the biomass of stocked fish, there were no significant differences in total
biomass between the three treatments in Trial 2.
Table 3 summarises the results from somewhat less comprehensive trials in rice-cum-
fish culture and under intensive culture in recirculating systems. Recovery\survival rates
were low in the rice-cum-fish culture (53.2-55.4%) and there were no significant differ-
ences between GMT and MST for any of the measured parameters. The comparative growth
and survival trends for GMT and MST in this trial were consistent with the pond trials as
were those from the experiments conducted in recirculating systems. In both recirculating
system experiments, the mean harvest weight of GMT was significantly greater (P < 0.001)
than for MST, resulting in 33.6A7.0% increases in yield.
Fig. 1 illustrates the differences in yield (converted to kg ha year-) for the five trials,
demonstrating that culture of GMT in the same strain, consistently enhances production in
all of the culture systems tested, with yields being on average 23% and 34% greater than
SRT and MST respectively.
320 G. Mair et al. /Aquaculture 137 (1995) 313-322

Table 3
Data from harvest of on-station performance trials of GMT and MST in rice-cum-fish and recirculating culture
systems

Environment Treatment Sex Survival Mean cv of Mean CV of Yield Total

(trial number) ratio (% (%) harvest weight harvest SL (kg) biomass
male) weight (%) SL(cm) (S) (kg)
(8)

Rice-Fish MST N/A 53.25 14.8 25.3 9 1.26 9.39 1.34= N/A
(3) GMT N/A 55.44= 15.98 24.82 8.08 8.37 1.50 N/A
Recirc. 1 MST 47.22 90.00 5 1.39a 26.22 N/A N/A 1.85 1.85
(4) GMT lOQ.00 91.25 74.63 22.46 N/A N/A 2.72 2.72
Recirc. 2 MST 53.19 94.00 50.02 20.75 N/A N/A 2.35 2.35
(5) GMT 100.00 92.0 0 68.41b 26.36 N/A N/A 3.14 3.14

4. Discussion

It is apparent from the results of these growth comparisons, especially from the pond
trials which approximate to culture environments commonly used in developing countries,
that the genetically male tilapia derived from YY-male broodstock in 0. niloticus have
attributes potentially well suited to aquaculture. It is generally perceived that all-male
populations will produce higher yields of marketable fish than normal mixed sex populations
and this study confirms this to be the case. The harvested yields observed in these trials
demonstrate that culture of genetically male tilapia can increase production of marketable
fish, compared with normal mixed sex tilapia, by between 12% and 58%, with a mean of
34%. Perhaps more surprising is the increase in yield of GMT populations when compared
to androgen sex reversed fish in the two comprehensive pond trials, these differences being
significant in the first of these trials. The main reason for this may be the lower sex ratios
in SRT although, in the case of the first pond trial (FAC Ponds Trial 1), sex ratios of GMT
and SRT were the same. For this reason this trial represents the best comparison of GMT
and SRT, and demonstrated significant improvements in mean harvest weight and yield of
the former. This can be considered a somewhat surprising result as the YY-male technology
has originally been proposed merely as an alternative method to produce all- or nearly all-
male populations and not as a means to produce better performing all-male tilapia. These
differences may be due to a number of factors, perhaps the most likely being the effect of
competition from recruits in SRT populations. In this trial a small percentage of females
( 1.5 - 1.7%) were present in both GMT and SRT populations. However, females in GMT
populations did not attain sexual maturity during the grow-out period whereas non-reversed
females in the SRT populations did mature and recruits were generated. This apparent
phenomenon late maturation of females in GMT populations is presently undergoing further
investigation. Additional factors may include differential growth of female genotypes in
phenotypically male SRT. Also, it is possible that the initial hormone treatment to induce
sex reversal may have a long term deleterious effect on the growth of treated fish as has
been suggested in carp (Komen et al., 1993), although this treatment with methyltestoster-
one is normally considered as a growth promoter as described in 0. mossambicus (Kuwaye
et al., 1993).
 G. Mair et al. /Aquaculture 137 (1995) 313-322 321

The absence of significant differences in weight and yield between GMT and SRT in the
repeated trial (FAC Ponds 2) may be due to a greater variation in performance between
replicates. Failure to induce sufficiently high sex ratios, whilst emphasizing the problems
of obtaining consistent results using this technique, limited the value of this trial as a direct
comparison of GMT and SRT.
As might be expected the presence of significant proportions of females (which were
smaller than males) increased the coefficient of variation for both weight and length com-
pared to the all-male populations. The lower size variation and particularly the absence of
recruitment in all ponds containing GMT are potentially significant benefits in tilapia culture
as these minimize or eliminate competitive effects on growth and facilitate pond manage-
ment, harvesting and marketing.
The results from trials conducted in a rice-cum-fish environment were inconclusive due
to high variation in growth and survival between replicates. However, the trend of improved
yield performance of GMT over MST was evident in this trial. Growth performance was
very poor in this environment and it cannot be considered as a typical culture system for
tilapia. Improved growth rates and yields of GMT were evident in recirculating systems
although survival and size variation did not appear to differ. Whilst these recirculating
systems clearly represented small experimental units, these results might suggest that GMT
could also have significant benefits for culture in intensive farming systems.
An unexpected finding was the low level of recruitment in GMT populations. It had
previously been suggested for cultured tilapia, that proportions of females in excess of 5%
would lead to very high level of recruitment and significant growth depression of stocked
fish under culture in ponds (Mair and van Dam, 1995). The data from culture of SRT in
these trials indicates that recruitment can be considerable in these populations but that any
increase in the proportions of males, over that of the control, will lead to increases in yield
of stocked fish at harvest.
Total biomass (stocked + recruit) was similar between all treatments in most trials
suggesting that this may be limited by the carrying capacity of the culture unit, under given
levels of input. Under such conditions, GMT, and probably well sex reversed SRT, would
have the economic advantage that the carrying capacity of the pond would be made up
almost entirely of marketable fish.

5. Conclusions

YY-males were successfully used to produce genetically male tilapia for a series of within
strain growth trials under a range of culture systems. Fish in GMT populations grew to a
larger size, had marginally greater survival and produced highly significant increases in
yield, when compared to mixed-sex populations. In one experiment in which sex ratios of
GMT and SRT were comparable, yields of the former, were significantly greater than the
latter, suggesting that the YY-male technology may have additional benefits for tilapia
culture other than simply as an alternative method for production of males. In addition,
there was no recruitment in GMT populations and any females in GMT populations (mean
sex ratio of 99.5% male across all trials) did not become sexually mature during the period
of grow-out. The culture of GMT may be more promising for enhancing yields in tilapia
322 G. Mair et al. /Aquaculture 137 (1995) 313-322

culture, than the use of hormonally sex reversed tilapia (SRT) . Further on-station trials of
GMT are underway to compare their performance in culture systems closely resembling
those commonly applied in developing countries such as the Philippines. Growth charac-
teristics are being compared with those of MST and SRT of locally culture Philippines
strains in order to determine the likely benefits to the Philippine fish farmer. On-farm trials
are also planned as are trials of YY-male broodstock in commercial hatcheries.
It can be concluded that the YY-male technology appears to be a highly promising genetic
technology for application in tilapia culture across a range of environments and could lead
to substantial increases in yield. It is important that the performance of GMT be further
evaluated in on-farm environments to determine if the same kind of increases in yield can
be achieved under less controlled conditions.

Acknowledgements

This paper is a contribution of the FAC\CLSU-UCS Genetic Manipulations for Improved

Tilapia (GMIT) Project supported by a grant (R.4803) from the British Overseas Devel-
opment Administrations Fish Genetics Programme to G.C. Mair and D.O.F. Skibinski. The
authors thank Mary Ann Abiado, Lilia Dahilig, and the involved GMIT project staff, for
their invaluable assistance in conducting these growth trials and for collation and analysis
of the data.

References

Guerrero III, R.D. and Guerrero, L.A., 1988. Feasibility of commercial productions of sex-reversed Nile tile
fingerlings in the Philippines, In: R.S.V. Pullin, T. Bhukaswan, K. Tonguthai and J.L. Maclean (Editors),
Proceedings of the Second International Symposium on Tilapia in Aquaculture, Bangkok, Thailand. ICLARM
Conf. Proc. 15, pp. 183-186.
Komen, J., Eding, E.H., Bongers, A.B.J. and Richter, C.J.J., 1993. Gynogenesis in common carp (Cyprinus
carpio) IV. Growth, phenotypic variation and gonad differentiation in normal and methyltestosterone-treated
homozygous clones and F, hybrids. Aquaculture, 111: 271-280.
Kuwaye, T.T., Okimoto, D.K., Shimoda, S.K., Howerton, R.D., Hoa-Ren, L., Pang, P.K.T. and Grau, E.G., 1993.
Effect of 17-methyltestosterone on the growth of the euryhaline tilapia, Oreochromis mossambicus, in fresh
water and in sea water. Aquaculture, 113: 137-152.
Hanson, T.R., Smitherman, R.O., Shelton, W.L. and Dunham, R.A., 1983. Growthcomparisons of monosex tilapia
produced by separation of sexes, hybridization, and sex reversal. In: L. Fishelson and Z. Yaron (Editors),
Proceedings of the International Symposium on Tilapia in Aquaculture, Nazareth, Israel Tel Aviv University,
Tel Aviv, Israel, pp. 570-579.
Macintosh, D.J., Singh, T.B., Little, D.C. and Edwards, P., 1988. Growth and sexual development of ll-methyl-
testosterone- and progesterone-treated Nile tilapia (Oreochromis niloricus) red in earthen ponds. In: R.S.V.
Pullin, T. Bhukaswan, K. Tonguthai and J.L. Maclean (Editors), The Second International Symposium on
Tilapia in Aquaculture. ICLARM Conf. Proc. 15, pp. 457-463.
Mair, G.C. and Little, DC., 1991. Population control in farmed tilapias. NAGA, 14:8-13.
Ma?, G.C., Capili, J.B., Beardmore, J.A. and Skibinski, D.O.F., 1993. The YY male technology for production
of monosex male tilapia, Oreochromis niloticus L. In: D.J. Penman, N. Roongratri and B.J. McAndrew,
(Editors), Proceedings of the International Workshop on Genetics in Aquaculture and Fisheries Management,
University of Stirling, 3 1 August4 September 1992 ASEAN-EEC Aquaculture Development And Coordi-
nation Programme ( AADCP) , Bangkok, Thailand, pp. 93-95.
 G. Mair et al. /Aquaculture I37 (1995) 313-322 323

Mair, G.C. and van Dam, A.A., 1995. The effect of sex ratio at stocking on recruitment in Nile tilapia (Oreochromis
niloticus L.) ponds. Proc. Third Int. Symp. Tilapia in Aquaculture, Abidjan, CBte dIvoire, 11-16 November
1991.
Mair, G.C., Scott, A., Penman, D.J. Beardmore, J.A. and Skibinski, D.O.F., 1991. Sex determination in the genus
Oreochromis 1: Sex reversal, gynogenesis, and triploidy in 0. niloticus L. Theoret. Appl. Genet., 82: 144
152.
McAndrew, B.J. and Majumdar, KC., 1989. Growth studies on juvenile tilapia using pure species, hormone-
treated and nine interspecific hybrids. Aquacult. Fish. Manage., 20:3547
Popma, T.J. and Green, B.W., 1990. Research and Development Series, 35: Sex reversal of tilapia in earthen
ponds, International Center for Aquaculture, Alabama Agricultural Experiment Station, Auburn University,
Auburn, AL, 15 pp.
Scott, A.G., Penman, D.J., Beardmore, J.A. and Skibinski, D.O.F., 1989. The YY supermale in Oreochromis
niloticus (L.) and its potential in aquaculture. Aquaculture, 78: 237-25 1.
Sevilleja, R.C., Cagauan, A.G., Lopez, E.A., dela Cruz, C.R. and van Dam, A.A., 1992. Component technology
research in rice-fish systems in the Philippines. In: C.R. dela Cruz, C. Lightfoot, B.A. Costa-Pierce, V.R.
Carangal and M.P. Bimbao (Editors), Rice-fish Research and Development in Asia. ICLARM Conf. Proc.
24, pp. 373-384.
Trombka, D. and Avtalion, R., 1993. Sex determination in tilapia - a review. The Israeli J. Aquacult. Bamidgeh,
45: 2637.
Vera Cruz, E.M. and Mair, G.C., 1994. Conditions for optimum androgen sex reversal in Oreochromis niloticus
(L.). Aquaculture, 122: 237-248.
Wohlfarth, G.W., 1994. The unexploited potential of tilapia hybrids in aquaculture. Aquacult. Fish. Manage., 25:
781-788.
Wohlfarth, G., Hulata, G., Rothbard, S., Itzkovich, J. and Halevy, A., 1983. Comparisons between interspecific
tilapia hybrids for some production traits. In: L. Fishelson and Z. Yaron (Editors), Proceedings of the
International Symposium on Tilapia in Aquaculture, Nazareth, Israel, Tel Aviv University, Tel Aviv, Israel,
pp. 559-569.