Aquaculture 195 2001.

121131
www.elsevier.nlrlocateraqua-online

Feed intake and growth of fast and slow growing

strains of rainbow trout Oncorhynchus mykiss / fed
by automatic feeders or by self-feeders
L.M.P. Valente a,) , B. Fauconneau b, E.F.S. Gomes c , T. Boujard d
a

Departamento de Engenharia Biologica
e Ambiental, Uniersidade de Tras-os-Montes e Alto Douro,
Apartado 202 5001, Vila Real, codex, Portugal
b
INRA, Laboratoire de Physiologie des poissons, Campus de Beaulieu, 35042 Rennes Cedex, France
c

Instituto Ciencias
Biomedicas Abel Salazar, Uniersidade do Porto, 4000-Porto, Portugal
d
INRA, Unite mixte INRA-IFREMER de Nutrition des Poissons, B.P. 3, 64310 Saint-Pee-sur-Nielle,
France
Received 17 July 2000; received in revised form 5 September 2000; accepted 5 September 2000

Abstract

Rainbow trout 8.59.5 g. of two strains C and M. differing in growth potential were
compared with respect to feeding motivation and feeding rhythms, over a 65-day experimental
period, employing self-feeding or automatic feeding. Growth rate, feed gain ratio, feed intake and
pattern of feeding activity of fish fed with self-feeders, were recorded, as was body composition of
both strains. The final weight of fish of the fast-growing strain, strain C, fed using self-feeders,
was significantly higher 82.6 g. than that observed for fish of the slow-growing strain, strain M
69.3 g.. When the automatic feeders were used, no significant differences were found between
the strains in terms of body weight gain 65 g.. Results observed for feed gain ratio were also
similar between the two strains. Although the voluntary feed intake VFI. did not vary signifi-
cantly with the genetic origin of the fish, strain C displayed a consistently higher VFI compared to
strain M. The retention efficiency of nutrients and energy were similar between strains but
significantly different between feeding systems. With regard to body composition, when fish were
fed by means of self-feeders, no significant differences were found between the two rainbow trout
strains. However, when fed automatically, dry matter and lipid content were highest in the strain
M fish. The energy and protein content of the whole fish were not significantly influenced either
by the feeding system or by the strain of the fish used. q 2001 Elsevier Science B.V. All rights
reserved.

Keywords: Growth; Self-feeding; Feed intake; Rainbow trout

)
Corresponding author. Tel.: q351-59-350000; fax: q351-259-350480.
E-mail address: lvalente@utad.pt L.M.P. Valente..

0044-8486r01r$ - see front matter q 2001 Elsevier Science B.V. All rights reserved.
PII: S 0 0 4 4 - 8 4 8 6 0 0 . 0 0 5 3 6 - 6
122 L.M.P. Valente et al.r Aquaculture 195 (2001) 121131

1. Introduction

When an animal starts to eat it is said to do so because it is in a state of hunger; when

it stops eating it does so because it is satiated Forbes, 1995.. The satiation time and
subsequent return of appetite limits the number of meals a fish will take each day
Grove et al., 1978.. The expression of feeding behaviour and the satiation time are both
of importance to fish farmers because of the need to ensure that feeding regimes
feeding frequencies, ration sizes and time over which they are dispensed. are adjusted
to maximize consumption, growth and conversion efficiencies. Furthermore, wastage
must be minimized because of high feed costs and the potentially deleterious effects of
waste feed on water quality Hyatt, 1979; Alanara, 1996.. The initiation of feeding
response and the modulation of food intake in fish depends on factors whose relative
importance differ with the species of fish concerned Boujard and Leatherland, 1992a..
Mechanisms that control satiety and food intake are complex, multifactorial and are not
yet clearly defined Forbes, 1995; Le Bail and Boeuf, 1997.. Nevertheless, environmen-
tal and physiological factors are considered important in the control of feeding be-
haviour Fletcher, 1984; Kaushik, 1996..
Rainbow trout display an endogenous rhythm of feeding behaviour synchronized by

the lightrdark alternation Boujard and Leatherland, 1992a; Sanchez-Vazquez and
Tabata, 1998.. These rhythms, which influence daily food requirements, are difficult to
predict. The ideal feeding method may therefore be a demand-triggered feeding system
that allows the fish themselves to control the rate of food supply Alanara, 1992a;
Boujard et al., 1992.. There are several factors that may influence the demand feeding
activity of a fish and thereby the ability to optimise growth and feed conversion rates:
the ability of the fish to regulate such a feeding system Adron et al., 1973; Landless,
1976; Boujard and Leatherland, 1992b; Alanara, 1994., temperature Alanara, 1992a,b,
1994., photoperiod Boujard and Leatherland, 1992b; Boujard and Luquet, 1996.,
reward level Landless, 1976; Alanara, 1994; Brannas
and Alanara, 1994., dietary
energy content Grove et al., 1978; Boujard and Medale, 1994 . , stocking density

Alanara and
Brannas, 1996 . and social dominance Adron et al., 1973; Landless, 1976;

Brannas and
Alanara, 1994;
Alanara and
Brannas, 1996;
Gelineau et al., 1998..
Examples of genetic effects on the feeding behaviour are not well documented, although
under certain circumstances differences in growth between triploid and diploid Atlantic
salmon parr were attributed to feeding behaviour Carter et al., 1994..
Differences in growth performance between rainbow trout strains have partly been
attributed to feed intake Valente et al., 1998; Valente, 1999.. While the distinct feeding
behaviour displayed by those strains could be only based on a distinct capacity to ingest
food, differences in appetite or feeding rhythms could also be involved. An appraisal of
these factors depends on the development of model systems by which the fish response
to the food can be assessed in quantitative terms Adron et al., 1973.. Self-feeders have
been shown to be useful for this kind of study as it is assumed that the fish themselves
accurately control the feeding level by activating a trigger in the water Alanara, 1996..
Furthermore, each feed can be recorded as a discrete event, providing information about
the feeding rhythms of fish that is also of practical interest, as feeding schedules out of
 L.M.P. Valente et al.r Aquaculture 195 (2001) 121131 123

phase with natural rhythm can affect growth Boujard et al., 1995; Gelineau et al.,
1996..
The aim of this work was to evaluate whether differences in growth performance due
to the genetic origin of the fish could be attributed to differences in appetite or feeding
rhythms.

2. Material and methods

Fish from two strains of rainbow trout Oncorhynchus mykiss . were used in this
study. The fast-growing strain, strain C, was originally obtained from a French farm
Cornec 29. and had been raised by INRA for six generations, whereas the slow-grow-
ing strain, strain M Mirwart., obtained from a private farm in Belgium by the
University of Louvain, had been raised by INRA for three generations. The genetic
distance between the strains, determined by analysis of five highly polymorphic
microsatellite, was 0.388 Nei, 1972. and 0.054 Cavalli-Sforza and Edwards, 1967..
Although differences were not too big, the two strains were always found in two distinct
clusters.
Eight groups of 100 fish weight range of 8.59.6 g. were randomly distributed
among eight square fibreglass tanks 180 l., in an open system. Fish were fed a
commercial pelleted feed Aqualife 17, Biomar: 48.5% crude protein, 19.5% crude fat;
23.5 kJrg DM gross energy; 90.1% dry matter.. Water flow and quality characteristics
were as follows: flow 0.6 lrmin; temperature 12138C; O 2 7 mgrl, NOy 3 7 mgrl,
PO43y 0.10 mgrl. Tanks were under a photoperiod regime of 12L:12D dawn at 0600 h
and dusk at 1800 h..

2.1. Feed and feeding

Four groups of fish two for each strain. were fed by automatic feeders programmed
to continuously deliver feed, for 12 h during the light phase, each day. The amount of
feed distributed was 2.5% of the fish biomass per tank per day. The other four groups
were fed by means of electronic self-feeders which delivered approximately 0.06 g of
feed each time a fish activated the trigger located 1 cm above the water surface. This
feeding system consisted of three parts: the detector, the feed hopper, and the interface
between the detector and the feed hopper Boujard et al., 1992.. The self-feeders were
connected to a computer recording time, the day and the tank from which each feed
demand originated. Demand for food was satisfied on a 24-h basis. The amount of feed
distributed was assessed by regular weighing of the feed remaining in the feed hoppers.
Feed waste was recovered, dried and weighed, both in tanks with self-feeders and
automatic feeders.
Data on distributed feed and weight gain of fish were collected every 2 weeks during
the 65-day experiment. All fish from each tank were weighed individually, at each time,
to the nearest 0.01 g. The diel profile of feeding activity of the fish fed by self-feeders
124 L.M.P. Valente et al.r Aquaculture 195 (2001) 121131

was recorded during a period of 20 days, 3 weeks after starting the experiment. A
pooled sample of 10 fish of each strain at the beginning of the experiment and five fish
per tank at the end were taken for whole-body proximate composition analyses.

2.2. Analytical methods

Chemical analyses of feed and whole fish were carried out on deep-frozen ho-
mogenates according to standard methods: dry matter 1048C for 24 h., crude protein
Kjeldahl, total nitrogen= 6.25. after acid digestion, fat content by petroleum ether
extraction at 40608C Soxhlet. and energy using an adiabatic bomb calorimeter Parr..

2.3. Calculation

The growth performance, feed intake and feed utilisation were described using the
following parameters:
DGC y Daily growth coefficients 100 final weight 1r3 y initial weight 1r3 . rdays;
Weight gain g . s final weight g . y initial weight g . ;
FGR y Feed gain ratio s feed intake g DM . rweight gain g . ;
VFI y Voluntary feed intake g DMrmean BWrd.
s feed intake g DM . rmean body weight BWi q BWf . r2 . rdays;
Nutrient or energy. intake grmean BWrd.
s VFI = Nutrient or energy. content of the diet g . ;
Nutrient retention efficiency % .
s 100 final weight= % nutrient of final whole body .
y initial weight= % nutrient of initial whole body .
rtotal nutrient intake.

2.4. Statistical analysis

Statistical analyses followed methods outlined by Zar 1996.. Data were analysed by
ANOVA with the Statistics 5.0 for Windows package. Arcsine transformations of
percentage data were performed to achieve homogeneity of variance. When F values
showed significance, individual means were compared using Tukeys honest significant
difference HSD. test. All differences were considered significant at P - 0.05.

3. Results

The performance characteristics of the fish are presented in Table 1. When fed by
means of self-feeders, fish of strain C exhibited significantly higher growth than those
 L.M.P. Valente et al.r Aquaculture 195 (2001) 121131 125

Table 1
Growth and feed intake of fast and slow-growing strains of rainbow trout C and M. fed by means of
self-feeders or automatic feeders. Values are the mean of duplicate groups"SD
Self-feeders Automatic feeders
C M C M
Growth performance
Initial BW g. 8.57"0.11 9.66"0.03 8.42"0.01 9.69"0.01
Final BW g. 82.60"6.22 a 69.30"1.98 b 64.95"2.19 b 64.75"0.21b
Weight gain g. 74.04"6.12 a 59.64"2.01b 56.54"2.18 b 55.06"0.23 b
DGC 3.55"0.16 a 3.05"0.07 b 3.04"0.06 b 2.90"0.01b
FGR 0.62"0.03 a 0.61"0.04 a 0.78"0.04 b 0.80"0.02 b

Intake (g or kJr kg per day)

Dry matter 15.3"0.85a,b 13.9"0.99 b 18.05"0.78 a 17.70"0.42 a
Energy 336.6"18.7 a,b 305.8"21.8 b 397.1"17.1a 389.4"9.3 a
Protein 7.41"0.41a,b 6.73"0.48 b 8.75"0.38 a 8.58"0.21a
Lipid 2.97"0.16 a,b 2.70"0.18 b 3.51"0.15a 3.44"0.08 a

Figures with different superscripts in the same line are significantly different from each other ANOVA,
P - 0.05..

fed using automatic feeders, but growth of strain M fish was not affected by the feeding
system. Significant differences in growth between trout of the two strains were observed
when fish were fed using self-feeders. Fish fed by automatic feeders were significantly
less efficient at converting feed to growth than were fish fed by self-feeders. No
differences were observed between the strains efficiency, regardless of the feeding
system used.
The VFI of the fish was influenced by the feeding system, with fish fed using
automatic feeders having the highest VFI. Although the VFI did not vary significantly
with the genetic origin of the fish, when trout were fed by self-feeders, strain C
displayed a higher VFI compared to strain M. However, taking into account that there is
a general decrease of VFI with body weight, strain C always ingested more feed than
strain M. In fact, fish from strain C ingested 46 g of feed weight gain = feed gain ratio.,
whereas slow-growing fish only took up 36 g. When automatic feeders were used, 44 g
of feed were distributed to both strains. Moreover, feed waste % feed intake. for fish
fed either by means of self-feeders or automatically, was 2.68% or 0.70%, for strain C,
and 8.36% or 3.54% for strain M, respectively, during the overall period.
The diel changes in the pattern of feeding demands are presented in Fig. 1. Almost all
the feeding activity occurred during the photophase in both strains. The pattern of
feeding demand in fish of strain C showed a clear major peak of food demand at dawn
P - 0.001. and a smaller peak at dusk P - 0.05.. In fish of strain M, the increase of
activity at dawn was gradual, the frequency of demands being lower than that observed
for strain C. At dusk, however, they also exhibited a small peak of activity P - 0.05..
Furthermore, the percentage of feed refusals % feed demand. during that particular
period was 2.42 for strain C and 22.75 for strain M.
 126
L.M.P. Valente et al.r Aquaculture 195 (2001) 121131
Fig. 1. Diel profile of feeding activity of the two strains of rainbow trout C and M., fed by means of self-feeders, under a photoperiod regime of 12L:12D dawn at
0600 h and dusk at 1800 h.. Values are the mean of duplicate groups"SD.
 L.M.P. Valente et al.r Aquaculture 195 (2001) 121131 127

Table 2
Whole body composition % body weight or kJrg body weight. and retention efficiency %. of fast and
slow-growing strains of rainbow trout C and M., fed by means of either self-feeders or automatic feeders, at
the beginning and end of the experiment. Values are the mean"SD of duplicate groups of five pooled fish
Self-feeders Automatic feeders
C M C M
Whole body composition (% BW or kJr g BW)
Dry matter 30.81"0.30 a 31.42"0.64 a 29.63"0.08 b 30.92"0.30 a
Energy 8.58"0.02 8.84"0.36 8.05"0.13 8.67"0.03
Protein 15.08"0.11 15.13"0.17 14.72"0.21 15.25"0.24
Lipid 10.75"0.67 a,b 12.12"0.39 b 9.95"0.15a 11.72"0.31b

Retention efficiency (%)

Energy 61.79"2.70 a 64.70"2.16 a 46.84"3.14 b 49.60"1.12 b
Protein 51.29"1.86 a 51.90"3.35a 40.26"2.68 b 40.86"0.35 b
Lipid 95.20"1.79 a 109.57"4.85a 71.75"4.63 b 82.81"4.57 a,b

The initial whole-body composition % wet weight. of a group of 10 fish was for strain C and M, respectively:
dry matter 22.79 and 24.62; energy 5.79 and 6.28; protein 13.68 and 14.16; lipid 5.36 and 6.91. Figures with
different superscripts in the same line are significantly different from each other ANOVA, P - 0.05..

3.1. Body composition

The body compositions of the fish at the start and end of the experiment are presented
in Table 2. The percent dry matter and lipid varied with the origin of the fish and the
feeding system. When fish were fed by self-feeders, no significant differences were
found between the two rainbow trout strains. However, when they were fed automati-
cally, dry matter and lipid were highest in fish of strain M. The percent energy and
protein of the whole fish were not significantly influenced by the feeding system or
strain of the fish.

3.2. Retention efficiency

The retention efficiency of nutrients and energy Table 2. were similar between
strains but significantly different between feeding systems. Fish fed by means of
self-feeders retained significantly more energy, protein and lipid than fish fed automati-
cally.

4. Discussion

Trout of different strains, fed by means of automatic feeders did not display growth
differences. Valente et al. 1998. have previously observed that fast and slow-growing
strains of trout grow at similar rates when a restricted level is manually distributed,
leading us to conclude that, in the present study, when the automatic feeders were used,
fish of strain C did not ingest the quantity of feed that was necessary to display maximal
growth rate. This resulted in increased feed waste and poorer feed gain ratios. When fed
128 L.M.P. Valente et al.r Aquaculture 195 (2001) 121131

by means of a self-feeder, strain C exhibited a significantly better growth performance

than strain M, but similar FGR, suggesting it has a greater motivation for food. The
motivation to eat, which is dependent on appetite and endogenous activity rhythms,
determines the strength of the feeding response Groot, 1996.. Growth differences
between the two rainbow trout strains have already been attributed to a higher intake
capacity of strain C, when fed manually to satiation Valente et al., 1998.. The VFI of
the fish when the self-feeder was used, although not significantly different between the
strains, was higher in trout of strain C. Taking into account the general decrease of VFI
with increasing body weight Kaushik and Luquet, 1984., and considering the signifi-
cantly higher weight and the faster growth of strain C, this means that fast growing trout
have consumed more feed than slow growing ones of equal size. In fact, the motivation
to feed was highest in fish from strain C, since these fish gained 19% more weight and
consumed 21% more feed weight gain = feed gain ratio. than fish from strain M. We
can conclude that the increased growth rate of strain C was not only due to a higher
capacity of the fish to ingest feed Valente et al., 1998., but also to a bigger appetite as
expressed by the activation of the trigger in the self-feeding system. In contrast to strain
C, the feeding method did not influence the growth performance of strain M. This
suggests that these fish ingested as much feed as they could under both systems,
displaying maximal growth rates.
With self-feeding it was possible to obtain good growth simultaneously with low feed
gain ratio. Good feed conversion FGR 0.70.8. was also obtained by Alanara 1994.
when rainbow trout were fed high energy diets using demand feeders. This was in
agreement with earlier observations on rainbow trout reared in tanks and fed using other
techniques Storebakken et al., 1991.. When the automatic-feeding method was used, a
poorer feed gain ratio was observed in both strains. Similarly, Paspatis et al. 1999.
found the highest feed gain ratio values in automatically fed sea bass compared to
self-fed or hand-fed fish.
In this report it was observed that rainbow trout were able to adjust their self-feeding
activity to fit food demand and thereby maintain a high growth rate, confirming previous
data in salmonids Boujard and Leatherland, 1992b; Paspatis and Boujard, 1996..

However, Gelineau et al. 1998. reported both lower VFI and specific growth rate in
groups of rainbow trout self-fed than in groups fed to satiation by hand. Furthermore,
Paspatis and Boujard 1996. found that Atlantic salmon fed high energy diets by means
of automatic feeders exhibited a lower growth rate than those fed by self-feeders.
Similarly, in sea bass it was observed that fish grew faster when they had access to a
self-feeder than when they were fed an identical amount of feed automatically Azzaydi
et al., 1998; Paspatis et al., 1999, 2000.. Thus, feeding strategy itself may affect feed
intake and growth.
When food is available at all times, rainbow trout synchronize the feeding rhythm
according to the cycles of light and dark Boujard and Leatherland, 1992a.. Concerning
the diel pattern of feed demand, fish from strain C showed a higher activity at dawn
0600 h. and again at dusk 1800 h.. Strain M exhibited a more equilibrated pattern
throughout the day, with no clear peaks. Boujard and Leatherland 1992b. have already
reported that food demand in rainbow trout occurred mainly during the photophase with
a major peak of feeding activity shortly after dawn and a secondary one associated with
 L.M.P. Valente et al.r Aquaculture 195 (2001) 121131 129

dusk. However, a high level of nocturnal feeding activity may be observed under natural
lighting conditions, as long as low background light intensity allows the fish to locate
and activate a trigger Landless, 1976..
Within each feeding method, nutrient and energy retention did not differ between the
two strains, confirming previous data Valente et al., 1998.. Medale 1993. has also
observed similar protein and energy retention efficiencies in rainbow trout strains with
different growth rates. The results on proximate body composition did not differ
between strains fed by means of self-feeders, although strain C showed significantly less
dry matter and lipid when fed automatically. This tends to support earlier findings
Valente et al., 1998. that reported similar body compositions but different body weights
between the two strains when fed to satiation, by hand; however, among fish of equal
weight there was a general tendency strain M to be fatter than strain C, if a dietary
restriction were to be imposed. Some studies have, however, shown a correlation
between lipid content of carcass and growth rate Ayles et al., 1979; Kinghorn, 1983;
Gjerde and Schaeffer, 1989., whereas other authors found higher body fat and lower
protein content in slow-growing rainbow trout strains than in fast-growing fish Smith et

al., 1988; Medale, 1993.. The significant differences in the retention efficiency between
feeding strategies seem to be a reflection of the poorer feed gain ratio in the automatic-
feeding method. The body composition of the fish fed automatically was similar to that
observed in self-fed fish. This is not in accordance with the results reported by Paspatis
and Boujard 1996., who found the lowest growth rates and the highest final adiposity in
Atlantic salmon fed with high energy diets by means of automatic feeders. In contrast,
however, Azzaydi et al. 1998. observed that in European sea bass the feeding system
had no effect on body composition.
From the experiments described here it can be concluded that the increased growth
performance of the fast-growing strain was due to the expression of a bigger appetite
that gave rise to a higher capacity of the fish to ingest food. The feed gain ratio and
retention efficiency of nutrients and energy were similar between strains.

Acknowledgements

The authors wish to thank Yannick Labreuche and Laurence Labbe for their technical
assistance during the trial and Paulette Peyrotte for performing the body composition
analysis. We would also like to thank Jorge Dias for his critical reading of the
manuscript. This research program was supported by a grant provided by the COST
Action 827 for short-term scientific missions.

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