Feeding behaviour of fast- and slow-growing strains

of rainbow trout, Oncorhynchus mykiss (Walbaum),

during rst feeding
L M P Valente
1
, P Saglio
2
, L M Cunha
3
& B Fauconneau
4
1
Departamento de Engenharia Biologica e Ambiental, Universidade de Tras-os-Montes e Alto Douro, Apartado 202,
5001 Vila Real codex, Portugal
2
INRA, Ecologie des Poissons, 65 rue de St Brieuc, 35042 Rennes Cedex, France
3
Departamento de Ciencia e Tecnologia, Universidade Fernando Pessoa, Praca 9 de Abril 349, 4249004 Porto,
Portugal
4
INRA, Laboratoire de Physiologie des Poissons, Campus de Beaulieu, 35042 Rennes Cedex, France
Correspondence: Lu sa Valente, DEBA UTAD, Apartado 202, 5001 Vila Real codex, Portugal. E-mail: lvalente@utad.pt
Abstract
The development of feeding behaviour in two
rainbow trout strains and their hybrid was com-
pared at the transition from endogenous nutrition
to exogenous feeding. After 60 min of acclimatiza-
tion, the swimming pattern, spatial distribution,
agonistic behaviour and snapping of sh were
quantied by making 5-min observations on groups
of ve sh. Behavioural patterns were recorded in
four replicate groups per treatment, before and after
the introduction of dry trout feed or Artemia nauplii
(actometric tests) or extracts of these (olfactometric
tests). Fish of the slow growing strain were less
active than those of the other two strains. Greater
swimming activity observed in the sh of the fast
growing strain was associated with more rapid
feeding and a higher number of snapping responses.
Such behavioural precocity is of interest because it
could provide the basis for the growth differences
between the fast and slow growing strains.
Keywords: feeding behaviour, aggression,
swimming activity, growth, rainbow trout
Introduction
The transition from endogenous nutrition to exo-
genous feeding is important for the survival of the
post-embryonic stages of sh and some of the high
mortality which occurs at this stage can be
attributed to starvation (Mearns 1986). Although
salmonids are believed to be predominantly visual
feeders (Mearns 1986), several studies (McBride,
Idler, Jonas & Tomlinson 1962; Adron & Mackie
1978; Atema 1980; Shparkovskiy, Pavlov &
Chinarina 1983; Mearns 1985, 1986) indicate
that olfactory and gustatory senses probably play
an important role in the detection and recognition
of potential food (Saglio & Blanc 1983; Mearns
1985). Fish may perceive food at a distance either
visually or chemically but the sense of taste is used
in making the decision whether to swallow or reject
the potential food material (Adron & Mackie 1978;
Atema 1980).
The start of exogenous feeding in rainbow trout,
Oncorhynchus mykiss (Walbaum), occurs at the time
of absorption of the yolk sac (usually 2029 days
from hatch) and follows the disappearance of the
oesophageal tissue plug and maturation of the taste
buds (Twongo & MacCrimmon 1976, 1977). On
emergence from the substrate, the young sh
exhibit a shift from a strong to a weak photo-
negative response (MacCrimmon & Twongo 1980)
and there is a synchrony in the anatomical,
physiological and behavioural development
involved in the transition from endogenous to
exogenous nutrition after emergence. A failure to
2001 Blackwell Science Ltd 471
Aquaculture Research, 2001, 32, 471480
initiate feeding or the inability to feed adequately for
survival seem to have a behavioural basis
(MacCrimmon & Twongo 1980).
The aim of the present work was to examine the
development of feeding behaviour during the early
life-history stages of two rainbow trout strains and
their hybrid. Responses to olfactory and visual
stimuli (live and formulated feed) were examined
in an attempt to assess whether differences in
feeding behaviour could provide a basis for observed
growth differences.
Materials and methods
Experimental animals
Fish from two strains of rainbow trout and a hybrid
were used in this study. The fast-growing strain,
strain C, originated from a French farm (Cornec 29)
and the slow growing strain, strain M (Mirwart),
originated from a Belgium farm. Full-sib matings
were obtained from the spawning of 30 females and
30 males. The hybrid, strain H, resulted from a cross
of strain M male and strain C female.
Eggs were fertilised on the same date, incubated
at 1112 C at INRA in Rennes, and hatched in
early January 1997. Behavioural observations were
carried out during January and February 1997, at
the end of yolk-sac reabsorption (T1) and then one
(T2) and 2 weeks later (T3). The young sh used in
the two rst series of tests had not been fed but those
used in the third test had been fed a commercial
trout feed (Aquastart, Biomar).
Treatments
Fish of each age were submitted to actometric or
olfactometric tests and two feed treatments were
applied in each type of test: live brine shrimp nauplii
(Artemia salina, 400500 m) or dry trout pellets
(Aquastart Biomar: 0.5 mm diameter). The food
particles for use in the actometric test were diluted
in 60 mL of freshwater (18 nauplii mL
1
of Artemia
or 227 particles mL
1
of pellets) immediately before
use. The extracts used for the olfactometric tests
were obtained by placing the same number of
particles as in the previous test in 60 mL freshwater
for 30 min. The mixtures were then ltered to
remove particulate matter. The ltrates were pre-
pared immediately before use. The 60 mL solutions
of food particles (actometric test) or the respective
ltrates (olfactometric test) were then injected in the
water inlet of the tank (Fig. 1) at a controlled rate
(ve injections of 12 mL at 1 min intervals). Five
injections of water were used as controls.
Procedure
The sh were observed in glass test tanks
(60 3 10 3 10 cm) with a shelter (screen) cover-
ing the third of the tank furthest from the water
inlet (Fig. 1). A group of ve naive sh was used in
each trial and there were four trials per treatment
and strain for each of T1, T2 and T3. Each group of
sh was left in the tank for 60 min before
observation. The behavioural patterns of the sh
were recorded prior to the introduction of the test
stimulus. Fish in each tank were then exposed to
one of the two types of food (actometric test) or their
respective ltrates (olfactometric test), and further
recordings of behavioural were undertaken. Given
groups of sh were tested only once and tanks were
cleaned thoroughly between tests.
Recorded response
The reactions of the ve sh in each tank were
observed directly, from behind a black cloth blind,
over 10 periods of 30 s each (5 min). The following
behavioural patterns were recorded at the begin-
ning of each 30-s interval, before and after
introducing the stimulus:
Immobility (I): number of sh resting motionless on
the bottom of the tank;
Active swimming (AS): number of sh exhibiting
body movements to propel themselves forwards,
backwards or upwards;
Figure 1 Illustration of a test tank. Test solutions (dry
pellets, Artemia or respective ltrates) were injected into
the water inlet. Test tanks were isolated with silent-
blocks and black plastic sheeting. Behavioural patterns
were observed from the side through a small (3-cm
diameter) opening in the sheeting. 1, 2, 3 are zones of
the aquarium; a shelter (shadow screen) was placed
above zone 3.
472 2001 Blackwell Science Ltd, Aquaculture Research, 32, 471480
Feeding behaviour in strains of trout L M P Valente et al. Aquaculture Research, 2001, 32, 471480
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2001 Blackwell Science Ltd, Aquaculture Research, 32, 471480 473
Aquaculture Research, 2001, 32, 471480 Feeding behaviour in strains of trout L M P Valente et al.
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474 2001 Blackwell Science Ltd, Aquaculture Research, 32, 471480
Feeding behaviour in strains of trout L M P Valente et al. Aquaculture Research, 2001, 32, 471480
Station holding (SH): number of sh exhibiting body
movements but without any change in position;
Shelter: number of sh in the downstream zone of
the tank, under the shelter.
Other behavioural activities were recorded during
the 30-s intervals:
Movement (M): number of times that transient jerks
or twitches of the body were observed;
Dart (D): number of times that short, sudden bursts
of rapid swimming were observed;
Agonistic behaviour (AG): number of sh exhibiting
aggressive acts, which include nipping and chasing
other sh;
Snap: number of times that a sh quickly opens and
closes the mouth, presumably directed at a particle
in the water.
Each set of data consisted of the total number of
observations over the 10 periods.
Data analysis
At each time of testing (T1, T2 and T3), the data
recorded for the different behavioural patterns in
the two types of test (actometric and olfacto-
metric) were subjected to a three-factor analysis
Figure 2 Mean number of observations of immobile sh
at the end of yolk-sac absorption (T1), after the
introduction of the stimulus. *P < 0.05.
Table 3 Mean number of observations (mean 6 SD), during 5-min periods of the different behavioural patterns in two
rainbow trout strains (C and M) and their hybrid (H), before and after the introduction of food items (Actometric test) or
their ltrates (Olfactometric test) 1 week after the end of yolk sac absorption, T2. N = 4
Growing
phase
Type
of test
Feeding
time Strain
Type
of food I AS SH Shelter M + D AG Snap
T2 Actometric Before C
M
H
Artemia
Pellets
Artemia
Pellets
Artemia
Pellets
0 (0)
0 (0)
0 (0)
0 (0)
0 (0)
0 (0)
34.5 (9.3)
28.5 (9.8)
13.2 (6.6)
23.8 (2.2)
29.8 (15.7)
28.3 (12.3)
15.5 (9.3)
21.5 (9.8)
36.8 (6.6)
26.2 (2.2)
20.2 (15.7)
21.7 (12.3)
6.8 (4.6)
8.5 (4.9)
4.0 (4.6)
3.0 (2.4)
2.5 (1.0)
4.6 (5.5)
0 (0)
0.8 (1.0)
0.2 (0.5)
0.8 (1.5)
0 (0)
0.3 (0.6)
0.5 (0.6)
0 (0)
0.8 (1.5)
0 (0)
1 (0.8)
0.3 (0.6)
11.0 (4.2)
14.2 (11.7)
8.5 (5.5)
12.5 (8.8)
13.5 (7.0)
12.6 (11.0)
After C
M
H
Artemia
Pellets
Artemia
Pellets
Artemia
Pellets
0 (0)
0 (0)
0.2 (0.5)
0 (0)
0 (0)
0 (0)
33.2 (4.4)
16.0 (12.0)
24.0 (9.3)
18.5 (7.8)
30.5 (4.4)
24.3 (5.0)
16.8 (4.4)
34.0 (12.0)
26.0 (9.3)
31.5 (7.8)
19.5 (4.4)
25.7 (5.0)
4.2 (4.6)
5.5 (5.3)
3.0 (4.7)
0.8 (0.5)
2.0 (4.0)
1.7 (2.9)
0 (0)
0.2 (0.5)
0 (0)
0.2 (0.5)
0 (0)
0 (0)
1.5 (1.0)
0.8 (1.0)
1.0 (0.8)
0.5 (1.0)
0 (0)
0.7 (0.6)
138 (21)
107 (25)
94 (17)
81 (17)
128 (23)
91 (1)
Olfactometric Before C
M
H
Artemia
Pellets
Artemia
Pellets
Artemia
Pellets
0 (0)
0 (0)
0 (0)
0 (0)
0 (0)
0 (0)
31.8 (7.5)
33.0 (11.0)
29.2 (7.8)
26.0 (7.3)
33.2 (3.2)
12.5 (9.8)
8.2 (7.5)
17.0 (11.0)
20.8 (7.8)
24.0 (7.3)
16.8 (3.2)
37.5 (9.8)
4.5 (4.8)
8.5 (7.1)
6.2 (4.4)
6.8 (8.5)
9.0 (6.0)
5.5 (5.3)
0.2 (0.5)
0.2 (0.5)
0.8 (1.0)
1.2 (0.5)
0.8 (1.0)
1.2 (1.3)
0 (0)
0 (0)
0.8 (1.0)
0.8 (1.0)
0 (0)
0.8 (1.5)
7.2 (5.5)
13.5 (6.9)
7.0 (2.9)
11.5 (7.8)
8.2 (3.8)
13.5 (9.9)
After C
M
H
Artemia
Pellets
Artemia
Pellets
Artemia
Pellets
0 (0)
0 (0)
0 (0)
0 (0)
0 (0)
0 (0)
24.0 (10.7)
24.7 (11.4)
25.5 (12.8)
18.5 (5.7)
32.0 (7.9)
11.8 (2.5)
26.0 (10.7)
25.2 (11.4)
24.5 (12.8)
31.5 (5.7)
18.0 (7.9)
38.2 (2.5)
2.8 (2.2)
5.8 (5.7)
6.2 (6.1)
3.8 (6.2)
2.0 (0.8)
7.0 (8.0)
1.2 (1.9)
5.5 (3.1)
2.2 (2.2)
3.2 (2.6)
2.2 (2.2)
5.0 (4.4)
0.5 (0.6)
0.5 (0.6)
0.5 (0.6)
1.2 (1.0)
0.5 (1.0)
0.7 (1.0)
8.5 (3.1)
14.0 (6.2)
11.5 (4.2)
9.2 (6.1)
13.2 (7.5)
13.0 (5.0)
2001 Blackwell Science Ltd, Aquaculture Research, 32, 471480 475
Aquaculture Research, 2001, 32, 471480 Feeding behaviour in strains of trout L M P Valente et al.
of variance (ANOVA) with blocks. Fish strain (C, M
and H) and food type (Artemia and pellets) were
the main factors and observation period (before
and after introduction of stimulus) the block.
Statistical analysis followed methods outlined by
Zar (1996) and data were analysed with the
SPSS 6.0 program. When F-values indicated
signicance, individual means were compared
using Tukey's Honest Signicant Difference
(HSD) test. Signicant differences were considered
when P < 0.05.
Overall evaluation of the different behavioural
patterns was undertaken using multivariate analy-
sis of variance (MANOVA) in which I, AS, SH, M + D,
shelter, AG and snap were taken as dependent
variables on a model similar to the one applied to
the univariate analysis. Wilks' Lambda was used as
the test value. Signicant differences were consid-
ered when P < 0.05.
Results
Behavioural patterns
In the `blank' tests, run as control, the injection of
water did not result in any change in behavioural
Table 4 Mean number of observations (mean 6 SD) during 5-min periods of the different behavioural patterns in two
rainbow trout strains (C and M) and their hybrid (H), before and after the introduction of food items (Actometric test) or
their ltrates (Olfactometric test), 2 weeks after the end of yolk sac absorption, T3. N =4
Growing
phase
Type
of test
Feeding
time Strain
Type
of food I AS SH Shelter M + D AG Snap
T3 Actometric Before C
M
H
Artemia
Pellets
Artemia
Pellets
Artemia
Pellets
0 (0)
1.8 (3.5)
0 (0)
0 (0)
1.5 (3.0)
0 (0)
14.2 (7.9)
17.0 (11.2)
13.5 (11.3)
12.0 (10.0)
14.5 (9.4)
21.8 (12.8)
35.8 (7.9)
31.2 (10.3)
36.5 (11.3)
38.0 (10.0)
34.0 (8.0)
28.2 (12.8)
9.0 (5.0)
3.8 (3.5)
12.8 (6.3)
22.5 (10.4)
10.0 (7.2)
4.2 (3.1)
1.8 (1.7)
1.0 (1.4)
0.8 (1.0)
0.5 (1.0)
1.2 (1.2)
1.2 (0.5)
2.6(3.8)
4.2 (2.6)
4.5 (3.9)
2.5 (3.0)
3.8 (7.5)
1.2 (1.5)
5.0 (5.7)
12.5 (8.0)
13.0 (7.4)
9.5 (5.5)
11.2 (12.0)
16.0 (6.0)
After C
M
H
Artemia
Pellets
Artemia
Pellets
Artemia
Pellets
0 (0)
2.0 (4.0)
0 (0)
0 (0)
0.2 (0.5)
0 (0)
30.2 (7.0)
29.2 (18.5)
37.8 (6.3)
24.0 (15.7)
31.0 (6.7)
32.0 (18.0)
19.8 (7.0)
18.8 (17.6)
12.2 (6.3)
26.0 (15.7)
18.7 (6.5)
18.0 (18.0)
9.8 (7.4)
1.0 (0.0)
9.8 (4.9)
13.0 (8.5)
5.2 (4.6)
0.8 (0.5)
0 (0)
0 (0)
0 (0)
0.2 (0.5)
0 (0)
0.2 (0.5)
5.0 (4.4)
2.8 (1.3)
2.0 (2.4)
2.5 (1.7)
0.8 (1.0)
1.0 (0.8)
171 (26)
152 (76)
162 (25)
115 (63)
174 (49)
161 (72)
Olfactometric Before C
M
H
Artemia
Pellets
Artemia
Pellets
Artemia
Pellets
0 (0)
0 (0)
0 (0)
0 (0)
1.0 (2.0)
0 (0)
16.3 (5.1)
17.8 (11.5)
15.9 (13.1)
11.5 (10.4)
20.8 (17.4)
26.8 (18.3)
33.8 (5.1)
32.2 (11.5)
35.0 (13.1)
38.5 (10.4)
28.2 (19.0)
23.2 (18.3)
6.5 (7.0)
10.8 (6.0)
17.0 (15.7)
8.5 (6.6)
10.2 (9.1)
8.5 (3.7)
1.0 (1.4)
1.5 (1.3)
1.2 (1.5)
1.2 (1.3)
2.2 (3.9)
1.8 (2.2
2.8 (4.3)
2.2 (2.2)
4.2 (4.0)
3.2 (2.4)
1.0 (1.2)
1.8 (2.4)
8.2 (7.3)
10.8 (6.5)
8.5 (3.7)
8.8 (5.2)
11.0 (6.3)
12.8 (7.8)
After C
M
H
Artemia
Pellets
Artemia
Pellets
Artemia
Pellets
0 (0)
0 (0)
0 (0)
0 (0)
0 (0)
0 (0)
19.5 (10.6)
20.0 (10.4)
13.5 (15.6)
22.0 (13.0)
12.7 (7.5)
27.5 (13.1)
30.5 (10.6)
30.0 (10.4)
36.5 (15.6)
28.0 (13.0)
37.2 (7.5)
22.5 (13.1)
5.8 (4.3)
6.7 (4.6)
18.5 (18.6)
9.5 (7.1)
5.5 (5.3)
8.0 (3.6)
3.8 (3.8)
2.0 (1.4)
2.8 (2.2)
1.8 (1.0)
2.0 (1.1)
1.0 (1.4)
5.8 (6.6)
5.8 (4.6)
4.5 (3.1)
8.0 (5.1)
1.0 (0.8)
5.8 (1.5)
18.0 (8.3)
23.5 (17.2)
9.0 (4.1)
15.8 (6.8)
18.0 (2.8)
19.5 (5.7)
Figure 3 Mean number of observations of sh under the
shelter, 2 weeks after the yolk-sac absorption (T3), before
and after the introduction of food items (actometric test).
*P < 0.05; **P < 0.01.
476 2001 Blackwell Science Ltd, Aquaculture Research, 32, 471480
Feeding behaviour in strains of trout L M P Valente et al. Aquaculture Research, 2001, 32, 471480
responses compared to the pre-injection condition.
This indicates that injections of water, per se, did
not inuence the behaviour of the sh. However,
rainbow trout responded to both visual and
chemical stimuli at the start of exogenous feeding
(Table 1).
Immobility
Exposure to feeding stimuli did not affect the
immobility of the sh (Tables 24). The mean
number of immobile sh differed among strains
when tested at the rst ontogenetic stage. This was
the case both before and after the introduction of
food items (actometric test) or of the respective
ltrates (olfactometric test). The slow growing strain
(M) had the highest number of immobile sh
(Fig. 2). Furthermore, the sh were less immobile
when Artemia was used than when dry pellets were
used.
Active swimming and station holding
Exposure to food items or their respective ltrates
affected the swimming activity of the sh in different
ways. During the olfactometric tests, 1 week after
yolk-sac absorption (T2), the number of sh
swimming actively decreased, whereas the number
of sh holding station increased. The reverse was
observed in the last period (T3), when food items
were introduced into the tank. Fish were then
mainly swimming actively.
During the two rst ontogenetic stages the
swimming activity of the sh was signicantly
affected by both the type of food and the genetic
origin of the sh The number of sh exhibiting
active swimming was signicantly higher when
the food was Artemia than when dry pellets were
used. On the other hand, pellets induced a
signicant increase in the number of sh holding
station. At the rst ontogenetic stage (T1), strain
C showed signicantly higher swimming activity
than the other two strains. At the second stage
(T2), more strain M sh held station but fewer
swam actively during the actometric test. No
strain differences in activity were detected in the
tests carried out at stage 3 (T3).
Shelter
The presence of stimuli only affected the number of
sh under the shelter at the last stage (T3)
(Table 1). The presence of food items induced sh
to leave this zone. The number of sh under the
shelter differed among strains when actometric tests
were performed. At the rst ontogenetic stage (T1),
strain C had the least number of sh under the
shelter but the highest number 1 week later (T2).
During T3, there were a signicantly higher
(P < 0.001) number of sh of the slow growing
strain (M) under the shelter than the other two
strains. In addition, these differences were observed
both before and after the introduction of food items
(Fig. 3).
Movement and dart
Data were pooled before analysis because of the low
number of sh exhibiting this behaviour. During the
olfactometric tests, at the two rst ontogenetic
stages (T1 and T2), the number of sh performing
movement or darting increased signicantly after
the introduction of the stimuli (Table 1). The strain
of the sh did not inuence movement and dart but
the number of responses was higher when pellets
were used.
Agonistic behaviour
The agonistic behaviour of the sh was not strongly
affected by either strains or food type (Table 1).
However, the number of aggressive interactions
increased signicantly during the olfactometric test
carried out at stage 3 (T3)
Figure 4 Mean number of snaps, 1 week after the yolk-
sac absorption (T2), before and after the introduction of
food items (actometric test). **P < 0.01.
2001 Blackwell Science Ltd, Aquaculture Research, 32, 471480 477
Aquaculture Research, 2001, 32, 471480 Feeding behaviour in strains of trout L M P Valente et al.
Snap
Snapping increased signicantly (P < 0.001) after
the introduction of food items (Table 1). When sh
were tested 1 week after yolk-sac absorption (T2),
snapping was found to be markedly inuenced
(P < 0.05) by both the type of food and the genetic
origin of the sh. Fish of the slow growing strain (M)
exhibited a signicantly lower response than those
of the other two strains (Fig. 4). The sh displayed
signicantly higher snapping responses towards
Artemia than towards pellets.
Overall response
The results of the MANOVA (Table 1) indicated that
feeding behaviour was inuenced by sh strain,
particularly during the rst two stages (T1 and T2)
of ontogenetic development and the effects were
more pronounced after the introduction of food
items (actometric test) than in olfactometric tests.
Discussion
There were differences in activity between the
strains, with sh of the slow growing strain (strain
M) being consistently less active than those of the
other two strains (strain C and H). The greater
swimming activity observed in the fast-growing
strain may be related to a higher number of bres in
the red muscle (Valente, Rocha, Gomes, Silva,
Oliveira, Moteiro & Fauconneau 1999). Such bres
are involved in sustained swimming activity (Nag &
Nursall 1972) and are recruited for low speed,
steady, undulatory swimming (Johnston, Davison &
Goldspink 1977; Bone 1978).
The position of the sh in the tank at the time of
introduction of the food items (actometric test)
differed among strains when sh were tested
2 weeks after yolk-sac absorption (T3). More of
the sh of the slow growing strain (M) tended to be
in zone 3 (shelter) and therefore did not have the
most rewarding location with respect to prey
availability. In contrast, sh of the fast growing
strain were more often in the upstream part of the
tank, where food items arrived. Fausch (1984)
observed that juvenile salmonids which held opti-
mal positions in the water current, in terms of
maximizing feeding in relation to energy expendi-
ture, also had the highest growth rate. The
establishment of territories often involves agonistic
behaviour, leads to the monopolization of food by
dominant individuals and results in an increase in
size variation with time (Jobling & Baardvik 1994).
However, in our study, there were few strain-related
differences in agonistic behaviour, irrespective of the
stage at which tests were carried out.
Our observations indicate that unfed rainbow
trout show innate snapping responses to the
presence of food at the start of exogenous
feeding. Similar results have been reported pre-
viously for brown trout Salmo trutta L. and
Atlantic salmon Salmo salar L. fry (Mearns
1986). Exposure to amino acids 5 days prior to
the rst exogenous feeding has been reported to
release biting and snapping behaviour in salmo-
nid alevins (Valentincic, Lamb & Caprio 1999).
In addition, Mearns (1985; 1989) observed both
increased snapping responses and a general
augmentation of movement, induced by amino
acids and shrimp extracts in unfed salmonids.
However, during the olfactometric tests, the sh
used in our study did not initiate clear feeding
responses to any of the food extracts tested,
unless they had previously been fed (T3).
Nevertheless, sh at the end of yolk sac absorp-
tion (T1) responded to the introduction of
chemical stimuli by increasing the number of
movements and darting.
The signicantly greater number of snapping
reactions observed in the sh of the fast growing
strain, 1 week after yolk-sac absorption (T2), could
provide the basis for the growth differences among
strains. Studies carried out on Japanese crucian
carp, Carassius auratus langsdori, Temminck &
Shlegel, demonstrated that size differences formed
during the rst feeding stage were maintained
(Umino, Sakae, Morishima, Arai & Nakagawa
1996). This is also consistent with observations
that interstrain differences in the capacity to
synthesize proteins were established soon after rst
feeding and were maintained throughout develop-
ment (Valente, Gomes & Fauconneau 1998). The
snapping reaction is an important indicator of
ingestion, but very little is known about its genetic
variation. Differences in growth between juveniles
of fast and slow growing rainbow trout strains have
been attributed to the capacity to ingest food
(Valente, Fauconneau & Gomes 1998) and feed
intake seems to have a heritable basis in channel
catsh, Ictalurus punctatus, Ranesque (Silverstein,
Bosworth, Waldbieser & Wolters, 2000).
When Artemia was the food, the sh displayed
signicantly more snapping responses than when
478 2001 Blackwell Science Ltd, Aquaculture Research, 32, 471480
Feeding behaviour in strains of trout L M P Valente et al. Aquaculture Research, 2001, 32, 471480
pellets were used. It is of little value to attempt direct
comparisons of frequencies of snap for the tested
foods because of appreciable differences in size,
texture, composition and number of particles
(MacCrimmon & Twongo 1980). However, the
present results are in accordance with the conten-
tion of Rimmer & Power (1978), who implied that
prey motility was a factor of major importance in
releasing feeding behaviour in fry. Furthermore,
Stradmeyer & Thorpe (1987) showed that naive
hatchery-reared Atlantic salmon readily consumed
wild prey and quickly developed a preference for it
over pelleted food. The same trends were observed in
rainbow trout (MacCrimmon & Twongo 1980) and
coho salmon Oncorhynchus kisutch Walbaum (Kim,
Massee & Hardy 1996). Moreover, Holm & Mller
(1984) suggested that sh preference could be
explained by different capture probabilities and
Ware (1972) observed that feeding rate could be
stimulated by increasing either prey density or prey
size.
In conclusion, the results of the present study
indicate that the swimming activity and the
feeding behaviour of rainbow trout are under
some genetic inuence and, during the rst
ontogenetic stage, the sh of the slow growing
strain were consistently less active than sh of
the other two strains. Furthermore, among fast
growing sh, the higher swimming activity and
occupation of a more rewarding position in the
tank, where food items arrived, seemed to be
associated with more snapping responses. Such
behavioural precocity is of interest because it
could contribute to growth differences between
fast and slow growing strains.
Acknowledgments
This research programme was supported by a grant
provided by the Cooperation JNICT/French
Embassy. The authors wish to express their
appreciation to Malcolm Jobling for his comments
on the manuscript.
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