carp, Megalobrama amblycephala (Yih), (Pisces,

Aquaculture Research, 1998, 29, 643–648
Preferred temperature of grass carp,
Ctenopharyngodon idella (Valenciennes), and brema
carp, Megalobrama amblycephala (Yih), (Pisces,
Cyprinidae) in horizontal and vertical gradients
F Dı´az1, S Espina2, C Rodrı´guez2 & F Soto3
1Departmento
de Acuicultura, Centro de Investigacio´n Cientı´fica y de Educacio´n Superior de Ensenada (CICESE),
Kilo´metro 107, Carretera Tijuana Ensenada, Ensenada, Baja California, Me´xico, 2Laboratorio de Ecofisiologı´a,
Departmento de Biologı´a, Facultad de Ciencias, Universidad Nacional Auto´noma de Me´xico, Me´xico D.F 04510, and
3Instituto
Nacional de la Pesca, Chilpancingo #70, Me´xico D.F 06100
Correspondence: Dr Fernando Dı´az Herrera, Department of Aquaculture (CICESE), P.O. Box 434844, San Diego, CA 92143–4844,
USA
Abstract
The preferred temperatures of grass carp,
Ctenopharyngodon idella (Valenciennes), and brema
carp, Megalobrama amblycephala (Yih), were
determined individually and together in both
horizontal and vertical gradients. No significant
difference was found by two methods between the
preferred temperatures in either species of carp
(28.0–28.5 °C) when the fish were placed
individually. However, when the two species of
carp were placed together, C. idella was displaced
thermally, preferring a temperature of 23.5 °C,
which was significantly lower than that obtained
when it was held separately. This suggests that, if
both species of carp are placed in ponds forming
part of a Chinese polyculture system, C. idella will
seek out a lower temperature as a mechanism to
avoid competition, and therefore, the growth of this
species will be reduced.
Introduction
Organisms inhabiting an aquatic environment that
is thermally heterogeneous in space and time possess
physiological and behavioural mechanisms which
permit them to live in changing environments.
Thermoregulatory behaviour is a coordinated
activity that results in a preference for a temperature
© 1998 Blackwell Science Ltd.
where the physiological functions operate with a
maximum efficiency, and also in which the animals
are exposed to minimal thermal stress. These are
the changing environments in which organisms
grow successfully (Hutchison & Maness 1979;
Prosser & Nelson 1981; Nichelmann 1983).
The grass carp, Ctenopharyngodon idella
(Valenciennes), and the brema carp, Megalobrama
amblycephala (Yih), were selected as test animals
because these fish are potentially important in
aquaculture since they form part of the Chinese
polyculture system. Because of their nutritional
habits, herbivorous carp are also used to biologically
control aquatic weeds (Cui, Liu & Chen 1992). In
Me´xico, carp are well accepted as a source of protein
by the communities which inhabit the central
plateau (Arredondo & Jua´rez 1986).
The devices and methods which determine
thermoregulatory behaviour in aquatic organisms
have been developed extensively, but because of
the fundamental differences in geometry between
horizontal and vertical gradients, the available
information on comparisons of the temperature
preferred by fish in both types of gradient is scarce
(McCauley 1977).
The aim of the present study was to determine the
preferred temperature of C. idella and M. amblycephala
when held in single-species and mixed populations
in both horizontal and vertical gradients.
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Preferred temperature of carp F Dı´az et al.
Aquaculture Research, 1998, 29, 643–648
Figure 1 Thermoregulatory
behaviour of Ctenopharyngodon idella
in (a) horizontal and (b) vertical
gradients. The shaded zone
represents the 95% confidence
interval of the median. The clear
bars show the 50% limits of
distribution.
Materials and methods
Juveniles of C. idella and M. amblycephala were
collected in the polyculture integral farm at
Tezontepec de Aldama, Hidalgo, Me´xico (20° 03’ N,
99° 17’ W). The carp were maintained for 15 days
in 60-L aquaria at a density of 15 organisms per
aquarium and at the temperature registered at
collection of 25 6 1 °C. The wet weight intervals of
C. idella and M. amblycephala were 2.3–14.1 and
3.6–14.4 g, respectively. The fish were fed daily with
a 1:1 mixture of chopped alfalfa and a balanced
meal, and the percentage of the diet varied from
10% to 14%, depending on the wet weight of the
animals. Food was provided for an hour and any
remaining food was then withdrawn by means of a
siphon. Two-thirds of the water volume was changed
644
daily after feeding. The photoperiod was maintained
at 12 light h:12 h dark.
The horizontal gradient consisted of an asbestos
tube (3.1 m long and 0.3 m in diameter) with a
channel in the upper part coated with liquid plastic
on its interior. The tube was divided into 16 sections
and a thermometer was placed in each one. There
were two separate compartments fitted at each end:
one was a cold chamber which was connected to a
thermoregulator; and the other contained a 500-W
heater. The gradient had a temperature range of
13–38 °C. Airstones were placed at the bottom of
the tube in order to avoid stratification of the
water column and maintain dissolved oxygen at
concentrations above 5 ppm.
The vertical gradient was a rectangle of galvanized
sheets (2.4 m high and 0.4 m long) with five 15-
© 1998 Blackwell Science Ltd, Aquaculture Research, 29, 643–648
Preferred temperature of carp F Dı´az et al.
Aquaculture Research, 1998, 29, 643–648
Figure 2 Thermoregulatory
behaviour of Megalobrama
amblycephala in (a) horizontal and
(b) vertical gradients. The shaded
zone represents the 95% confidence
interval of the median. The clear
bars show the 50% limits of
distribution.
cm thick plexiglass windows which were coated
internally with liquid plastic. The trough was divided
into 15 sections and thermometers were placed in
each one. The lower part of the apparatus was the
cold chamber and the upper one was the hot
chamber. The gradient had an temperature range
of 5–40 °C. An aeration system was installed in the
lower part of the device in order to avoid water
stratification and to maintain oxygen concentrations
of 4–5.2 p.p.m in the water column.
For each carp species, 10 individuals in the same
weight range were introduced into the horizontal
and vertical gradients via the section with the same
temperature as that of the acclimation conditions.
The position of the animals and the temperature
were recorded every 2 h over a 24-h period. Three
replicates were carried out for each experimental
condition.
Three C. idella and three M. amblycephala in same
weight range were introduced into both gradients
to discover if the preferred temperature was modified
© 1998 Blackwell Science Ltd, Aquaculture Research, 29, 643–648
by the interaction of the two species of carp. The
position of carp and the temperature in each of the
gradients sections was recorded every 2 h over a
24 h cycle. Three replicates were carried out for
each experimental condition.
The temperature preference data were processed
using the exploratory data analysis of Tukey (1977),
which was represented in a parallel box plot. The
data on temperature selection when C. idella and M.
amblycephala were placed individually and together
in both gradients were compared using the Mann–
Whitney rank sum test.
Results
Neither C. idella nor M. amblycephala exhibited a diel
cycle of preferred temperature when placed in the
horizontal and vertical gradients. Grass carp had a
preferred temperature of 25.8–30.2 °C (median 5
28 °C) in a horizontal gradient. In the vertical
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Preferred temperature of carp F Dı´az et al.
Aquaculture Research, 1998, 29, 643–648
Figure 3 Thermoregulatory
behaviour of (a) Ctenopharyngodon
idella and (b) Megalobrama
amblycephala in a horizontal
gradient. The shaded zone
represents the 95% confidence
interval of the median. The clear
bars show the 50% limits of
distribution.
trough, the temperature selected by grass carp was in
the 26.3–29.6 °C range (median 5 28 °C) (Fig. 1).
There was no difference when the temperatures
selected by grass carp in both types of gradients
were compared (P 5 0.7605). The preferred
temperature of brema carp placed in both troughs
was in the 27.4–30.6 °C range (median 5 28.5 °C)
(Fig. 2). No differences were observed when the
preferred temperature of brema carp in the
horizontal and vertical gradients were compared
(P 5 0.4872).
When both species of carp were placed together
in the horizontal and vertical gradients, the preferred
temperature of C. idella was 20.4–25.6 °C (median 5
23.5 °C) (Figs 3 & 4). When this result was compared
with that obtained when grass carp were placed
individually in both gradients, it was found to be
significantly lower (P 5 0.0001) by a mean value
646
of 4.5 °C. Megalobrama amblycephala had a selected
temperature range of 25.7–30.4 °C (median 5
27.5 °C) (Figs 3 & 4) when placed together with
grass carp. This result was not significantly different
(P 5 0.6229) from that obtained when brema carp
were placed individually in both gradients.
Discussion
Predictive information about the temperature
responses of fish that inhabit heterothermal
environments can be obtained by studying
thermoregulatory behaviour since devices and
methods for analysing this data have been
extensively developed (McCauley 1977). No
differences in preferred temperature (28–28.5 °C)
were observed in grass and brema carp when both
© 1998 Blackwell Science Ltd, Aquaculture Research, 29, 643–648
Aquaculture Research, 1998, 29, 643–648
Preferred temperature of carp F Dı´az et al.
Figure 4 Thermoregulatory
behaviour of (a) Ctenopharyngodon
idella and (b) Megalobrama
amblycephala in a vertical gradient.
The shaded zone represents the 95%
confidence interval of the median.
The clear bars show the 50% limits
of distribution.
species were placed individually in horizontal and
vertical gradients. McCauley & Pond (1971) found
that the preferred temperature (18–19 °C) of
fingerling rainbow trout, Oncorhynchus mykiss
(Walbaum), was similar in both horizontal and
vertical troughs. This suggests that the nature and
geometry of the gradients did not have any effect
on the preferred temperature values which were
determined in these fish.
The temperature range selected by the two species
of carp in both types of gradient (28–29 °C) is close
to that reported by Pitt, Garside & Hepburn (1956),
who used the acute method to obtain a final preferred
temperature of 32 °C for Cyprinus carpio (L.).
Reynolds & Casterlin (1977) reported a preferred
temperature of 29 °C for the same species in a
shuttlebox. In grass carp, Alcaraz, Rosas & Espina
(1993) used the acute method and obtained a
© 1998 Blackwell Science Ltd, Aquaculture Research, 29, 643–648
temperature preference of 29 °C. McCauley &
Casselman (1981) and Kellog & Gift (1983)
demonstrated that the final preferred temperature
values of 16 species of freshwater fish were close to
or coincided with the optimum temperature for
growth. Therefore, fish-culturists should use the
preferred temperature obtained in the present study
as a guide for adjusting rearing temperatures to
favour maximum growth and optimize the culture
of C. idella and M. amblycephala in Mexico. This is
in agreement with Magnuson, Crowder & Medvik
(1979), who examined temperature preference in
relation to the thermal niche and body growth in
the three species of fish, and can be viewed as
an ecological resource which can be successfully
exploited by fish.
Giattina & Garton (1982) suggested four
hypotheses to unify concepts in the study of the
647
Preferred temperature of carp F Dı´az et al.
thermoregulatory behaviour of fish: (1) final
preferred temperature is a specific response; (2) the
partition of habitat by thermoregulatory behaviour
is both an intra- and an inter-specific response in
segregation; (3) the preferred temperature may
reflect thermal optima for certain physiological
processes; and (4) fish generally avoid thermal
extremes before these become lethal. When two
species of carp were placed together, it was observed
that grass carp were equally displaced to 23.5 °C in
both the horizontal and vertical gradients. This
finding is in agreement with the second hypothesis.
Therefore, C. idella use thermoregulatory behaviour
as an environmental segregation mechanism to
avoid competition for space and food with M.
amblycephala.
Martinez & Abrego (1984) mention that C. idella
occupies the middle layer of the water column in
Chinese polyculture systems. The nutritional habits
of grass carp are omnivorous. Megalobrama
amblycephala also occupies the middle layer of the
water column and feeds on macrophytes. It was
found that carp do not fulfil the assumption of the
Chinese polyculture system, which states that a
complementary relationship exists between these
species, when there is no competition for food or
space. When the thermoregulatory behaviour of
both fish together was studied in horizontal and
vertical gradients, C. idella were found to prefer a
lower temperature, which significantly reduces their
growth in culture conditions. Therefore, it is
recommended that only one of these species should
be included in a Chinese polyculture system if
optimum utilization of the resource is to be achieved.
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© 1998 Blackwell Science Ltd, Aquaculture Research, 29, 643–648

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