Journal of Thermal Biology 27 (2002) 433–437
Effect of temperature on spawning of Penaeus merguiensis
Tung Hoanga,*, S.Y. Leea, Clive P. Keenanb, Gay E. Marsdenc
a School of Environmental & Applied Sciences, Griffith University, Qld 9726, Australiab Bribie Island Aquaculture Research Centre, P.O. Box 2066, 144 North Street, Bribie Island, Qld 4507, Australia
c NFIEC, TAFE, Locked Bag 5, Grafton, NSW 2460, Australia
Received 22 September 2001; accepted 1 February 2002
Abstract
The maturation rate and spawning performance of ablated Penaeus merguiensis were significantly (Po0:05) higher at
271C than at 231C. Increasing the temperature from 231C to 271C further improved spawning performance (Po0:05),
whereas reducing the temperature from 271C to 231C impaired spawning. The effect of temperature during the inducing
period was more critical than during the holding period. Prawn growth (% of weight gain), however, was not different
between 231C and 271C over the 60-day experimental period (P > 0:05). r 2002 Elsevier Science Ltd. All rights
reserved.
Keywords: Temperature; Ovarian maturation; Spawning; Penaeus merguiensis; Growth; Prawn
1. Introduction
The banana prawn, Penaeus merguiensis, is an
important species for commercial fisheries in the Indo-
Pacific region (Grey et al., 1983). This species, however,
has been ignored by modern shrimp farmers because it
does not grow as fast as P. monodon, the major culture
species. Thus, information about reproductive biology
of captive P. merguiensis is limited compared with other
culture species. However, the recent shortage of quality
P. monodon post-larvae for grow-out in southeast Asia
and Australia has prompted more farmers to culture P.
merguiensis; and the change has been good (Hoang,
2001). Therefore, further research on its reproductive
biology is needed to aid hatchery production.
Water temperature is one of the environmental factors
that strongly influence prawn reproduction (Muncy,
1984; Primavera, 1985; Staples, 1991; Bray and Lawr-
ence, 1992). In natural context, an increase of tempera-
ture from late winter to spring is believed as one of the
cues that trigger spawning of prawns (Dall et al., 1990;
Yano, 1993). Simultaneously increasing temperature
and day length has been reported to stimulate ovarian
maturation and spawning of P. japonicus (Laubier-
Bonichon and Laubier, 1979, cited by Primavera, 1985),
P. duorarum (Cripe, 1994) and P. stylirostris (Chamber-
lain and Gervais, 1984). Rothlisberg and Jackson (1985),
based on field sampling of larval abundance, suggested
that 291C is optimal for reproduction of P. merguiensis
in the tropical northern Australia. However, the effect of
temperature on reproduction of subtropical P. mer-
guiensis has not been reported. Dredge (1985) reported
the absence of ripe P. merguiensis females during winter
in the central Queensland coast, Australia. This in-
dicates that a low temperature is unsuitable for ovarian
maturation and spawning of this species. Recent study
on the cold tolerance of P. merguiensis suggests that the
species can cope with low temperatures, enabling two
culture crops per year in the subtropical southeast
Queensland, Australia (Hoang et al., 2002). Breeding P.
merguiensis to produce post-larvae for stocking, there-
fore, may require temperature manipulation as this
environmental factor varies seasonally.
In this study, the effect of low and high temperatures
(in the holding and inducing periods) on ovarian
maturation and spawning of P. merguiensis was
*Corresponding author. Present address: Bribie Island
Aquaculture Research Centre, P.O. Box 2066, 144 North
Street, Bribie Island, QLD 4507, Australia. Tel.: +61-7-3400-
2020; fax: +61-7-3408-3535.
E-mail address: [email protected] (T. Hoang).
0306-4565/02/$ - see front matter r 2002 Elsevier Science Ltd. All rights reserved.
PII: S 0 3 0 6 - 4 5 6 5 ( 0 2 ) 0 0 0 1 3 - X
investigated. The effect of short-term raising and
reducing temperature was also examined. Other envir-
onmental parameters such as photoperiod, light
intensity, salinity and pH were maintained at optimal
levels in order to demonstrate the profound effect of
temperature.
2. Materials and methods
Female prawns were randomly collected from the
grow-out tanks (60 m3 in volume) at Bribie Island
Aquaculture Research Centre, Queensland, Australia
in early spring 2000. Average carapace length (CL) and
weight (W ) at collection time was 22.971.2 mm (SD)
and 7.671.2 g (SD), respectively. Prawns were then
stocked in twelve 1-m3 tanks (fiber glass, painted black,
40 cm water depth) at a density of 5 individuals tank�1.
The experiment used a flow-through system; water used
in the experimental period was continuously exchanged
at a rate of 300% volume day�1. During the experi-
mental period, photoperiod, light intensity, salinity and
pH were maintained at 14L:10D, 2–3 lux, 34.5–35.1 ppt
and 8.0–8.1, respectively. Prawns were fed with 10% of
biomass day�1 by the BIARC maturation diet (a
mixture of fresh frozen green mussel, squid, calf liver
and other essential nutrients; 54.6% crude protein;
Marsden et al., 1997) and 2% of biomass day�1 by
EBI STAR pellets (50% crude protein, Higashimaru,
Japan) at 09:30 and 17:00. The experimental tanks were
cleaned twice a day prior to feeding.
The experiment had two consecutive periods: holding
(39 days) and inducing (21 days). Two levels of water
temperature (high: 271C and low: 231C) were tested in
each period, establishing four combinations of treat-
ments namely: high–high (HH), high–low (HL), low–
low (LL) and low–high (LH). Each treatment had three
replicate tanks. In the HH and LL treatments, water
temperature was maintained at 271C and 231C, respec-
tively, throughout the experimental period. In the HL
treatment, water temperature was 271C during the first
28 days, then reduced to 231C (within 24 h) from day 29
and maintained at this level until the end of the
experiment. In the LH treatment, water temperature
was 231C during the first 28 days, then increased to 271C
(within 24 h) on day 29 and was maintained at this level
until the end of the experiment. To induce spawning, all
the prawns were unilaterally ablated on day 39 (10 days
after the water temperature was increased in the LH
treatment and reduced in the HL treatment).
Ovarian maturation was checked daily at 17:30,
following the descriptions provided by Tuma (1967)
and Primavera (1985). Females with stage-IV ovaries
were then transferred to 150-l spawning tanks and
returned to the corresponding experimental tank early
next morning. Water temperature in the spawning tank
was similar to that in the experimental tank where the
spawner was taken from. Sampling for fecundity was
conducted by vigorously agitating the tank water to
ensure a homogenous suspension of eggs, then quickly
taking 100-ml aliquot samples and counting the number
of eggs in each sample under a stereo microscope.
Data were analyzed with Genstat 4.1 for Windows
(Payne et al., 1993). Analysis of variance (ANOVA) and
least significant difference (LSD) multiple-range proce-
dure were used to detect the statistical difference among
the treatments. The percentage of spawners and the
percentage of prawns reaching stage IV were analyzed
by a generalized linear model (modeling of binomial
proportions by logits) (McCullagh and Nelder, 1989).
Due to the limited number of observations, no statistical
test was conducted for latency period and fecundity.
Means were calculated and presented for only prawns
that spawned. The latency period is defined as the
number of days from eyestalk ablation to the first
spawning of a spawner. The number of spawns
produced by prawn individuals (i.e. number of spawns
female�1) was adjusted by the number of survival days,
presumably capable of spawning.
3. Results
Prawns started to spawn 6 days after eyestalk
ablation. More prawns reached full ovarian maturation
(stage IV) and were spawned at 271C, while increasing
the temperature from 231C to 271C further stimulated
ovarian maturation and spawning (Table 1). The
percentage of spawners was significantly (Po0:05)
highest in the LH treatment (73.3%), followed by the
HH (40.0%) and HL (26.7%) treatments, and lowest in
the LL treatment (6.7%) (Table 1). No significant
difference (P > 0:05) in the percentage of spawners was
found between the HH and HL treatments (treatments
that had a high temperature during the holding period),
and between the HL and LL treatments (treatments that
had a low temperature during the inducing period). The
numbers of spawns female�1 in the LH and HH
treatments was significantly (Po0.05) higher than the
HL and LL treatments, averaging at 0.061 and 0.048
spawns day�1, respectively (Table 1). Prawns in the HL
and LL treatments produced only 0.017 and 0.003
spawns day�1 on average, respectively. The total
number of eggs produced female�1 averaged at 63,137
in the LH and 42,870 in the HH treatments; significantly
(P > 0:05) higher than the HL and LL treatments.
Overall, the LH and HH treatments each produced
almost a million eggs during 21 days, while the HL and
LL treatments produced only 164,450 and 18,200 eggs,
respectively.
For prawns that spawned, the latency period was
similar among the HH, HL and LH treatments with
T. Hoang et al. / Journal of Thermal Biology 27 (2002) 433–437434
mean values of 12.0–13.8 days. Only one prawn spawned
in the LL treatment 17 days after eyestalk ablation
(Table 1). This prawn produced 18,200 eggs. Mean
fecundity of spawners was higher in other treatments,
ranging from 32,175 to 42,945 eggs spawn�1.
The interval between two successive spawns was usually
3 days within an intermoult period; and was extended to
5–8 days if moulting occurred between two successive
spawns.
Growth of prawns (% of weight gain) was not
significantly different (P > 0:05) among the treatments
(Table 1). However, there was a trend of reducing
growth in treatments with the lower temperature (231C).
Growth of prawns in the LH treatment was likely to be
affected by high spawning rate. Survival was high in all
the treatments: 86.7%, 93.3%, 93.3% and 100% in the
LL, LH, HH and HL treatments, respectively.
4. Discussion
From the results of this study, it is clear that water
temperature has a strong effect on ovarian maturation
and spawning of P. merguiensis. The percentage of
spawners, number of spawns female�1 and total eggs
produced female�1 were all significantly higher
(Po0:05) in treatments with the higher water tempera-
ture (271C) during the period that prawns were induced
to spawn, i.e. during ovarian maturation. The formation
of oocytes during ovarian maturation requires a
substantial mobilization of nutrients from other tissues,
e.g. hepatopancreas and food intake for the synthesis of
new materials, e.g. vitellogenin (Harrison, 1990; Wou-
ters et al., 2001), which is regulated by hormones (Yano,
1998; Huberman, 2000). The lower temperature (231C)
in this study may have retarded these processes, thus
compromising the completion of ovarian maturation or
reducing fecundity. Although photoperiod, light inten-
sity, salinity and pH were generally optimal for penaeid
reproduction (Primavera, 1985; Bray and Lawrence,
1992), spawning performance of prawns was poor in
treatments that had lower water temperature (231C)
during the inducing period. This may indicate that the
effect of temperature can override those aforementioned
environmental factors. Alternatively, ovarian maturation
can be induced by eyestalk ablation, but a suitable
temperature is still a requisite for the success of the
process and spawning. This finding is in agreement with
previous studies on penaeid species. Rothlisberg et al.
(1987) suggested that temperature might be the factor that
controls spawning pattern of penaeid prawns in natural
conditions. During winter ripe P. merguiensis females were
absent in the central Queensland coast (Dredge, 1985) and
were significantly reduced in number in the Gulf of
Carpentaria, northern Australia (Crocos and Kerr, 1983).
Similarly, Penn (1980) reported neither maturation nor
spawning of P. latisulcatus in Cockburn Sound when
water temperatures were o171C. In laboratory condi-
tions, high maturation and spawning rate of P. esculentus
were achieved at 261C, while only a few spawnings were
achieved at 201C (Crocos and Kerr, 1986).
Furthermore, the water temperatures in which the
prawns had been held previously (before spawning was
induced) and its patterns (increasing or reducing) also
influenced the spawning performance of prawns. The
HL treatment had a favorable result compared to the
LL treatment, suggesting that the higher temperature
during the first 28 days (holding period) in this treatment
was beneficial to ovarian maturation and spawning
performance of the prawns later. However, when
compared with the HH treatment, it is clear that
reducing the temperature from 271C to 231C severely
affected spawning performance of prawns. In contrast,
despite the fact that prawns were held in 231C during the
first 28 days, increasing the temperature from 231C to
271C in the LH treatment significantly improved the
Table 1
Maturation and spawning performance of prawns
Variables Treatment
HH HL LL LH
Percentage of prawns reaching stage IV 6079.1 ab 33.378.7 bc 13.376.3 c 8077.4 a
Percentage of spawners 4078.6 b 26.777.7 bc 6.774.4 c 73.377.7 a
Latency period (days)a 12.070.9 13.870.4 17.0 12.871.6
Number of spawns female�1 day�1 0.04870.005 a 0.01770.003 b 0.003 b 0.06170.006 a
Fecunditya 40,17173962 32,17575461 18,200 42,94577750
Total eggs produced female�1 42,87078726 a 10,96373337 b 1213 b 63,137712,186 a
Total eggs produced treatment�1 899,050 164,450 18,200 947,050
Weight gain (%) 129730.8 127724.4 119711.7 105719.8
Data are means7SE; within rows; means with different alphabets are significantly different (Po0:05).a No statistical test was conducted due to the limited number of observations. Means were calculated only for prawns that spawned.
Only one spawn was recorded in the LL treatment.
T. Hoang et al. / Journal of Thermal Biology 27 (2002) 433–437 435
percentage of spawners and spawning performance of
prawns; it even appeared better than the HH treatment.
This somehow suggests that the varying pattern of
temperature may be as important as, if not more
important than, its static levels (high or low) on
spawning of P. merguiensis. Furthermore, the results
of this study suggest that the effect of temperature
during the inducing period is more important than the
holding period.
Similar results have been reported for other penaeid
species. Simultaneously increasing temperature and
day length resulted in ovarian maturation of P. japonicus
(from 151C to 281C; Laubier–Bonichon and Laubier,
1979, cited by Primavera, 1985), P. stylirostris
(from 181C to 281C; Chamberlain and Gervais, 1984)
and P. duorarum (from 211C to 281C; Cripe, 1994).
Increasing the temperature from 201C to 281C alone
stimulated mating in P. stylirostris (Robertson et al.,
1991).
Therefore, the induction of spawning in P. merguien-
sis should involve raising water temperature to assist
eyestalk ablation; and must be maintained at a favorable
level for high ovarian maturation and spawning rate.
Field survey in the tropical northern Australia suggests
the optimal temperature for spawning and larval
development of P. merguiensis is 291C (Rothlisberg
and Jackson, 1985). Success in breeding P. merguiensis
in tropical areas was also reported at 25–311C (Beard
et al., 1977); 26–301C (Nurjana and Yang, 1976). In
subtropical areas natural water temperature is rarely
over 271C. Our observations show that ablated P.
merguiensis had a higher mortality when maintained at
29–301C. Therefore, 271C is more suitable for breeding
this species in subtropical areas.
Within the range of temperature from 231C to 271C in
this study, prawn growth was not significantly (P > 0:05)
different among the treatments although there was a
trend of reducing growth in treatments with the lower
temperature (231C). It is probable that, for P. merguien-
sis in subtropical areas, 231C might not adversely affect
prawn growth. Staples and Heales (1991) also reported
that a combination of 201C and 20 ppt resulted in
highest survival and greatest biomass increases of P.
merguiensis juveniles. In this present study, low growth
rate of prawns in the LH treatment appeared to be a
combined result of a lower temperature during the first
28 days and their higher spawning rate later on. With an
interval between two successive spawns as short as 3
days, prawn spawners are believed to mobilize their own
energy more than from daily intake for reproduction
(Harrison, 1990). Higher spawning rate, therefore, is
likely to result in depressed growth. Since prawn growth
was not affected at 231C, this may facilitate holding
prawn broodstock at this level before raising tempera-
ture up to 271C to stimulate spawning of ablated P.
merguiensis.
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