Available online at www.sciencedirect.

com

Aquaculture 275 (2008) 163 – 168

www.elsevier.com/locate/aqua-online

Growth, production and fin damage in cage-held 0+ Atlantic salmon pre-smolts

(Salmo salar L.) fed either a) on-demand, or b) to a fixed
satiation–restriction regime: Data from a commercial farm
Chris Noble a,b,⁎, Sunil Kadri a,c , David F. Mitchell a,d , Felicity A. Huntingford a
a
Fish Biology Group, Division of Environmental and Evolutionary Biology, Institute of Biomedical and Life Sciences,
University of Glasgow, Glasgow, Scotland, G12 8QQ, UK
b
Norwegian Institute of Fisheries and Aquaculture Research, Muninbakken 9-13, Postboks 6122, N-9291 Tromsø, Norway
c
Aquaculture Innovation, 34 Lawrence Street 1/L, Glasgow, Scotland, G11 5HD, UK
d
Huon Aquaculture Company Pty Ltd, Hideaway Bay, Dover, Tasmania, Australia 7117

Received 25 April 2007; received in revised form 17 December 2007; accepted 26 December 2007

Abstract

This experiment investigated the impact of feeding regime upon the growth performance, production efficiency and fin damage of cage-held 0+
Atlantic salmon pre-smolts. Six groups of pre-smolts (n = 61847 ± 2620 fish group− 1) were held in six 12 × 12 × 4 m production cages for 64 days
(23rd August until 26th October) at initial and final densities of 4.6 ± 0.2 kg m− 3 and 9.5 ± 0.5 kg m− 3, respectively (mean ± SEM). Fish were
subjected to ambient photoperiod (11.67–16.05 h min–max) until 16th October, before being held under 24 h light. Fish were fed throughout the
light phase and comparisons were made between three groups fed to an imposed regime (scheduled fixed ration feeding every 10 min) and three
groups fed on-demand using commercial interactive feedback systems. The study was divided into primary and secondary phases: during the
primary phase (part I) each regime was fed to satiation, whereas during the secondary phase (part II) fish under the imposed regime received a
restricted ration that was ca. 20% less than those fed on-demand. During part I there were no differences in ration (g fish− 1), fish condition and
Specific Growth Rate (Gw) between regimes (Gw: 1.45 ± 0.06 vs. 1.44 ± 0.04 for imposed vs. on-demand fed fish, respectively). During part II
underfeeding in the imposed regime had a significant and deleterious impact upon fish condition, FCR and growth (Gw: 0.45 ± 0.04 vs. 0.84 ± 0.09
for imposed vs. on-demand fed fish, respectively). Fish under the imposed regime also had significantly more dorsal fin splitting than those fed on-
demand for the duration of the study, and significantly more dorsal fin erosion at the end of Parts I and II. In addition, the smallest fish within each
cage suffered the greatest amount of fin splitting and erosion at the end of Parts I and II, irrespective of feeding regime. Daily ration amongst the
on-demand cages decreased as winter approached; stepwise multiple regression analyses demonstrated that this was significantly related to a
seasonal decrease in temperature.
© 2008 Elsevier B.V. All rights reserved.

Keywords: Atlantic salmon; FCR; Feeding regime; Fin damage; Growth; Welfare

1. Introduction
Feed is a substantial cost driver in aquaculture and accounts for
50–60% of the total operational costs of a seawater salmon farm
⁎ Corresponding author. Current address: Norwegian Institute of Fisheries and
Aquaculture Research, Muninbakken 9-13, Postboks 6122, N-9291 Tromsø,
Norway. Tel.: +47 77 62 90 11; fax: +47 77 62 91 00.
E-mail address: chris.noble@fiskeriforskning.no (C. Noble).
0044-8486/$ - see front matter © 2008 Elsevier B.V. All rights reserved.
doi:10.1016/j.aquaculture.2007.12.028
(Sveier and Lied, 1998). Feed management strategies must match
feed supply (feed delivery) to demand (appetite and the nutritional
requirements of fish) incorporating such factors as feeding rate,
frequency, duration and the choice of feeding regime (see Talbot
et al., 1999). For a number of years, salmon growers have fed fish
according to feed tables, which feed a fixed daily ration in response
to changes in fish size, photoperiod and water temperature.
Although based upon comprehensive research into fish nutrition,
feed tables assume that food is consumed whenever it is offered
164 C. Noble et al. / Aquaculture 275 (2008) 163–168
irrespective of time of day or feeding regime and this can lead to
over- or under-feeding, as the appetite of Atlantic salmon (Salmo
salar L.) varies both within (Ang and Petrell, 1997) and between
days (Blyth et al., 1993).
Overfeeding can lead to food wastage and can have a sub-
stantial impact on farm profits (Thorpe and Cho, 1995), can cause
environmental degradation (Cho and Bureau, 1998) and can also
increase FCR (Talbot et al., 1999). Alternatively, underfeeding
can reduce growth and increase FCR (Cho, 1992) equating to
lower yields and lost profits. It can also increase competition
(Davis and Olla, 1987), size heterogeneity (Johansen and Jobling,
1998) and fin damage (Damsgård et al., 1997). Dorsal fin damage
may be caused by aggression in salmonids (Turnbull et al., 1996)
and can be used as an indirect index of aggression and com-
petition (Moutou et al., 1998). Fin damage may also affect
perceived product quality (Winfree et al., 1998 and references
therein) and is a potential operational welfare indicator.
On-demand feeding technology, where fish regulate the
timing, frequency and size of feed deliveries, offers the
opportunity to evaluate existing feed management strategies and
potentially improve feeding methods. The primary objective of
this commercial scale experiment was to compare the production
performance and fin damage of cage-held 0+ pre-smolts fed to a)
satiation using on-demand feeders, or b) fed a fixed satiation–
restriction regime using automatic feeding systems. Additional
information was provided on inter-day variability in daily ration
for cages under the on-demand feeding regime, in relation to
changes in environmental variables, including a short 10 day
period where the 0+ pre-smolts were held under 24 h light.
2. Materials and methods
2.1. Fish husbandry and experimental set-up
The experiment was carried out at the Marine Harvest Scotland Glenfinnan
smolt production site (56.9°N 4.7°W). Six groups of Atlantic salmon pre-smolts,
S. salar L. were held in six 12× 12× 4 m (576 m3) cages (n = 61847 ± 2620 fish
group− 1, mean ± SEM). On the 13th August fish were given a 10 day acclimation
period to become conditioned to each feeding regime. The experiment began on the
23rd August using fish weighing 43.0 ± 0.4 g and held at an initial density of 4.6 ±
0.2 kg m− 3 (mean ± SEM). The study was split into two parts: Part I (23rd August–
27th September) and Part II (28th September–26th October) and lasted 64 days.
Final density was 9.5± 0.5 kg m− 3 (mean ± SEM). All cages were subjected to an
ambient photoperiod (11.67–16.05 h: min–max that incorporated civil twilight)
until the 16th October. From 16th October until seawater transfer during the week
beginning 15th November, fish were exposed to additional night-time illumination
between 1630 h and 0800 h using twin 400 W metal halide lights (Pisces 6;
Aquabeam Ltd., Grantham, UK) submerged at 2 m. Fish were subjected to ambient
temperature (10.25–15.25 °C: min–max) for the duration of the study.
Due to the high numbers of fish used in the experiment, it was not possible to
select all fish from an identical strain. Two cages of fish were from a low
maturing strain of fish and four groups were from a medium maturing strain.
Each regime had one cage from the low maturing strain and two cages from the
high maturing strain. The experiment was terminated on the 26th October in
response to a farm management protocol requiring minimal handling of the fish
during the two weeks prior to seawater transfer.
2.2. Feeding regimes and feed management strategies
Each cage was fitted with a 50 kg centralised feed hopper and a vibrator
dispenser attached to a circular spreader (Sterner Aquatech UK Ltd., Inverness,
UK) which spread feed pellets in a 10 m diameter circle. Fish were fed
commercial diets in relation to fish size: a 2 mm extruded pellet (Ecostart,
Biomar Ltd., Grangemouth, UK) until 30th September and from the 1st October
a 2.3 mm extruded pellet (Royale, Skretting, Northwich, UK).
Three cages (hereafter termed the imposed regime) were fed a fixed ration
every 10 min using Aquatess KC20 Automatic Feed Controllers (Aquatess Ltd.,
Dingwall, UK). Food was dispensed for 15 s at each feed delivery. Fish under
the imposed regime were subject to a satiation–restriction regime. During the
primary phase of the experiment (Part I) fish under the imposed regime were fed
to perceived daily satiation using a combination of in-house feed tables
supplemented by daily hand feeding at 08.00 h and 15.00 h. During hand
feeding a surface feeding response was used to determine satiation. During the
secondary phase of the experiment (Part II), fish under the imposed regime were
fed using only the automatic feed controllers and received a restrictive ration in
response to a production directive from farm management (this ration was
retrospectively ascertained to be ca. 80% of the total ration delivered to the on-
demand treatment).
A further three cages (hereafter termed the on-demand regime) were fed to
daily satiation during Parts I and II using AQ1 on-demand feeding systems
(AQ1 Systems Pty Ltd., Hobart, Australia). Each AQ1 system consisted of a
submerged infra-red sensor (located at the base of a 0.5 m diameter conical pellet
trap and suspended at 3 m) which regulated the amount of food delivered in
relation to pellet wastage (as described by Blyth et al., 1993). Ration size per
feeder spin was synchronised between cages, and feed delivery per spin
increased or decreased in response to pellet wastage (30–230 g spin− 1).
From 23rd August until the 16th October, fish under each regime were fed
from dawn until dusk (beginning and end of twilight). After this point (when all
cages were subject to 24 h light) fish under the on-demand feeding regime were
given the opportunity to feed for 24 h, whilst control fish under the imposed
regime had their fixed daily ration extended throughout the 24 h period. In
addition, the percentage of feed delivered by on-demand feeders during the
ambient day and illuminated night (when fish were held under 24 h light) was
also recorded.
2.3. Sampling methods
Fish were sampled on the following dates: 23rd–26th August (n = 300 fish
sample− 1), 28th–30th September (n = 400 fish sample− 1) and 25th–26th
October (n = 500 fish sample− 1). During sampling, all fish were crowded into
one quarter of the cage for b3 min and fish were netted out at random. Fish were
anaesthetised using a stock solution of benzocaine (b50 mg L− 1) and body
weight (wet weight ± 0.5 g), fork length (±0.5 cm), the number of fish with
dorsal fin splitting (presence/absence of splitting) and dorsal fin erosion (N50%
tissue loss) was recorded.
2.4. Environmental variables
Daylength (h) was measured using sunrise and sunset data for Fort William
(56.8° N 5.1° W) obtained from the Royal Observatory (Edinburgh, UK).
Changes in daylength from the previous day (h) were also calculated (after
Smith et al., 1993). Water temperature (±0.1 °C, at 2 m) and clarity (measured
using a Secchi disc) were taken at ca. 0800 h each morning. Average daily wind
speed (Knot, recorded at the Aonach Mor weather station 56.8° N 5.0° W) was
kindly provided by The British Atmospheric Data Centre (Chilton, UK), and
used as a proxy for the strength of wind-driven currents and an index of water
conditions at the sampling site (George, 1981).
2.5. Data manipulation and statistical analysis
Statistics were performed using Minitab (v. 13-0. Minitab Inc., Philadelphia,
USA) or SPSS (v. 13-0. SPSS Inc., Chicago, USA). Normality was tested for
using an Anderson–Darling test. If requirements for normality were not satisfied,
data were either subjected to natural log transformation or to non-parametric
statistics. All production performance and fin damage data satisfied homogeneity
of variance requirements, irrespective of cage or fish strain (Levene's test: all
analyses P N 0.05) allowing the pooling of data from different strains. Any
percentage data were transformed prior to analysis using the arcsine square root
transformation. Weight gain (% of initial weight) was used to express growth.
Size heterogeneity (weight) within each cage was calculated using the
 C. Noble et al. / Aquaculture 275 (2008) 163–168 165

Coefficient of Variation (CV) and differences between regimes were tested for
using a Mann–Whitney test. Potential differences in condition factor (K) between
regimes were tested for using a Kruskal Wallis test. Specific Growth Rate (Gw)
was calculated using the formula: Gw = 100 (ln W2 − ln W1) t− 1, where W1 and
W2 are the initial and final body weights (g) and t is the number of days between
sampling points. Feed Conversion Ratio (FCR) was calculated using the amount
of food delivered divided by wet weight gain (kg). Ration size (g fish− 1) for each
experimental period was also calculated. Any potential differences in mortality,
Gw, FCR and ration size between regimes were tested for using one-way
ANOVA's. Differences in the amount of a) fin splitting and b) fin erosion
between regimes were tested for using 2 × 2 Chi2 tests. The weight of fish with fin
splitting and erosion was compared within five percentiles (1–20 containing the
smallest fish and 81–100 the largest) for each sampling event and feeding regime
and 2 × 5 Chi2 tests were used to test for any differences within each regime.
Potential effects of environmental variables on the daily ration of cages fed on-
demand were tested for using stepwise multiple regression analyses. Independent
variables were: daylength (h), change in daylength (h), water temperature (°C),
water clarity (m), and average daily wind speed (Knots). The dependent variable
was daily ration, expressed as percentage body weight day− 1. Residuals of
significant variables and daily ration were then used to create a ‘seasonally
corrected daily ration’ index, and post-hoc regression analyses of non-significant
predictors were used to investigate whether these factors had any additional effect
upon daily ration. A significance level of P b 0.05 was used for all statistical tests.

3. Results

For a summary of production variables during Parts I and II see Table 1.

Mortality was low (b 0.5% for both treatments) and unrelated to feeding
regime (Part I: F1, 4 = 3.52, P = 0.134; Part II: F1, 4 = 0.46, P = 0.534).

3.1. Production variables Part I Fig. 1. The effect of feeding regime on the incidence of dorsal fin damage. Filled
symbols represent the imposed regime (mean + SEM) and open symbols
During Part I, feeding regime had no significant effect on ration represent the on-demand regime (mean − SEM). Graphs a) and b) represent data
(F1, 4 = 1.82, P = 0.249, Table 1), growth rate (F1, 4 = 0.01, P = 0.932) or on the incidence of fin splitting and erosion, respectively. ⁎, ⁎⁎, ⁎⁎⁎ denote
significant differences at P b 0.05, b0.01 and b0.001, respectively.
fish condition at both the start (Hadj = 2.21, P = 0.137) and end
(Hadj = 0.00, P = 0.972) of the period. In addition, there were no significant
differences in size heterogeneity between regimes at either the start 3.2. Production variables Part II
(W = 10.0, P = 1.000) or end (W = 13.0, P = 0.383) of Part I. Further, each
regime achieved consistently low FCR's (b1.0) and there were no During Part II, when fish under the imposed regime were subject to
significant differences between treatments (F1, 4 = 3.33, P = 0.142). feed restriction, the reduced ration had an adverse effect on a number of
production variables. Fish under the imposed regime received
Table 1 significantly less feed than those fed on-demand (79.2 ± 3.3% of the
Mortality (%), ration size (g fish− 1), growth rate (Gw), weight gain (%), body total on-demand ration, mean ± SD, F1, 4 = 20.13, P = 0.011, Table 1).
weight (BW initial and final), condition factor (K), size heterogeneity (CV Growth rate was also significantly lower in fish under the imposed
weight) and the FCR of fish fed to an imposed or on-demand regime during parts regime (F1, 4 = 16.59, P = 0.015) and these fish were also in significantly
I and II, where fish under the imposed regime were fed to satiation or restriction, poorer condition at the end of Part II (Hadj = 40.70, P b 0.001). Feeding
respectively regime had no effect on growth heterogeneity (W = 11.0, P = 1.000),
Part I Part II although there was a weak significant trend for higher FCR in fish fed to
the imposed regime (F1, 4 = 7.84, P = 0.049).
Imposed On-demand Imposed On-demand
Mortality (%) 0.24 ± 0.07 0.11 ± 0.01 0.06 ± 0.02 0.04 ± 0.01 3.3. Fin damage
Ration (gram fish− 1) 24.58 ± 1.28 22.45 ± 0.93 13.78 ± 0.70 17.38 ± 0.39⁎
Gw 1.45 ± 0.06 1.44 ± 0.04 0.45 ± 0.04 0.84 ± 0.09⁎ The total number of fish affected by dorsal fin splitting remained
Weight gain (%) 66.1 ± 3.2 67.4 ± 4.0 12.9 ± 0.9 25.0 ± 3.1
relatively constant throughout the experiment, although the incidence of
BWi (g) 42.4 ± 0.5 43.7 ± 0.1 – –
BWf (g) 70.4 ± 0.8 73.1 ± 1.6 79.4 ± 0.5 91.4 ± 2.4
dorsal fin erosion increased with time, irrespective of regime (Fig. 1).
CVi 22.5 22.8 – – Although fish under the on-demand regime had significantly less fin
CVf 43.3 37.1 50.4 47.2 splitting at the start of the experiment, this trend became highly sig-
Ki 1.24 1.24 – – nificant at the end of Part I and Part II (for the significance of Chi2
Kf 1.40 1.40 1.33 1.38⁎⁎⁎ analyses see Fig. 1). Significantly more fish exhibited fin erosion under
FCR 0.89 ± 0.06 0.77 ± 0.04 1.55 ± 0.18 0.98 ± 0.11⁎ the imposed feeding regime at the end of Parts I and II, in comparison to
All fish were also subject to 24 h during the last 10 days of Part II. All data are those fish fed on-demand.
presented as mean ± SEM with the exception of condition factor and growth There was a relationship between fish size and the incidence of
heterogeneity data (median values). ⁎ and ⁎⁎⁎ denote inter-regime differences at dorsal fin splitting and erosion (Fig. 2). At the end of Parts I and II, fin
P b 0.05 and P b 0.001, respectively. splitting was most prevalent amongst the smallest fish within each cage
166 C. Noble et al. / Aquaculture 275 (2008) 163–168

Fig. 2. The incidence of dorsal fin damage within each regime in relation to fish size. Values represent the mean ± SEM. Filled bars represent the imposed regime and
open bars represent the on-demand regime. Numeric titles 1), 2) and 3) represent the sampling periods 23rd–26th August, 28th–30th September and 25th–26th
October. a) and b) represent fin splitting or erosion, respectively. ⁎⁎⁎ denotes significant differences at P b 0.001 for each feeding regime.

(for the significance of Chi2 analyses see Fig. 2), irrespective of feeding
regime. There was a similar trend for fin erosion, although differences
were only significant at the end of part II.
3.4. Variability in daily on-demand feed delivery
Daily ration in all on-demand cages decreased as winter approached
(Fig. 3). Forward and backward stepwise regressions (as two or more
independent variables were correlated, Table 2) showed that directional
changes in temperature were the only significant predictor of daily
ration for all cages fed on-demand (cage 1: P = 0.001, R2adj = 35.8;
cage 2: P = 0.013, R2adj = 15.0; cage 3: P b 0.001, R2adj = 49.0).
Residuals of temperature and daily ration were then used to create a
‘seasonally corrected daily ration’ index. Regression analyses of this
dataset (independent variables: water clarity, daylength, change in
daylength and wind speed; dependent variable: residuals of daily ration
and temperature) show that average daily wind speed was the only
significant predictor of seasonally corrected daily ration for all cages
(cage 1: P = 0.043, R2adj = 12.0; cage 2: P = 0.008, R2adj = 19.8; cage 3:
P = 0.027, R2adj = 9.5). On-demand daily ration was therefore correlated
with changes in temperature and additional variation around this trend
was significantly correlated with daily wind speed.

Table 2
The Spearman's rank-order correlation coefficients showing the relationships
between daylength (h), change in daylength (h), water temperature (°C), water
clarity (m) and average daily wind speed (Knot) of factors included the
regression analysis of on-demand daily ration data
Daylength Change in Water Water
daylength temperature clarity
Change in − 0.906 ⁎⁎⁎
daylength
Water − 0.578 ⁎⁎⁎ −0.756 ⁎⁎⁎
temperature
Water clarity 0.273 0.375 ⁎ − 0.565 ⁎⁎⁎
Fig. 3. Variation in daily ration (% body weight day − 1) in cage-held 0+ parr fed
Wind speed − 0.128 −0.206 0.391 ⁎⁎ −0.419 ⁎⁎
on-demand, in addition to water temperature (°C). The arrow indicates the onset
of 24 h artificial light. ⁎P b 0.05, ⁎⁎P b 0.01, ⁎⁎⁎P b 0.001.
 C. Noble et al. / Aquaculture 275 (2008) 163–168
When held under 24 h light, on-demand replicates 1, 2 and 3 con-
sumed 71.04 ± 11.29, 58.80 ± 3.06 and 69.90 ± 30.06% of their ration
during the ambient day, respectively (mean ± SD). The onset of 24 h
light had neither a positive nor negative effect on daily ration size (see
Fig. 3) and mean water temperature and ambient daylength during this
period was 11.30 ± 0.43 °C and 11.29 ± 0.24 h (mean ± SD).
4. Discussion
The primary objective of this experiment was to compare the
production performance of commercial scale groups of fish fed
on-demand in comparison with fish fed according to existing farm
practices. Part II also investigated the effect of feeding regime and
moderate feed restriction on production performance.
The results of Part I suggested that the imposed feeding
regime, based upon a combination of in-house feed tables and
hand feeding, accurately matched the daily ration requirements
of Atlantic salmon pre-smolts during autumn. A previous study
on cage-held Atlantic salmon parr (Noble et al., 2007a) was used
to revise the farming company's feed tables during the one year
inter-study period, and data from the current study show this
revision was successful. During Part II, fish under the imposed
regime were intentionally and significantly underfed, although
feed restriction during this period was relatively moderate (ca.
80% of fish fed on-demand) and short-lived (b 1 month).
During Part I, there were no significant differences in growth
rate (GW) or condition factor between feeding regimes. When fish
under the imposed regime were fed a restrictive ration, growth rate
and condition were significantly worse in comparison to those
fed to satiation under the on-demand regime. Other studies have
also reported the detrimental effects of underfeeding upon growth
(Quinton and Blake, 1990; Gaylord et al., 2001) and fish
condition (Gregory and Wood, 1999; Pierce et al., 2001).
Aquaculturists have a vested interest in reducing size hetero-
geneity, as market forces demand a high quality product of uniform
size. In the current study, size heterogeneity was not affected by
feeding regime or feed restriction, a finding that is supported by a
comparable study on cage-held parr (Noble et al., 2007a). This
suggests that group size (Li and Brocksen, 1977) or high intra-
specific variation in metabolic rate (Cutts et al., 1998) plays a
greater role in regulating growth heterogeneity than behavioural
interactions and competition under commercial conditions.
During Part I, FCR's were low (b 1.0, see also Noble et al.,
2007a) and did not differ between regimes. Low FCR equates to
efficient farm practice, which is especially important in a com-
petitive and somewhat volatile Atlantic salmon aquaculture sector,
where market prices can change on a weekly basis (Forsberg and
Guttormsen, 2006). During Part II, moderate feed restriction
(ca. 80% of satiation) had a detrimental effect upon FCR. This is
supported by some studies (Cho, 1992; Bureau et al., 2006), but
not others (Storebakken and Austreng, 1987; Rasmussen et al.,
2000) that suggest feed restriction (N 50% of satiation) has no
impact upon FCR.
With consumer and retailer awareness of farm animal welfare
now expanding to fish, an increasing number of producers are
incorporating welfare considerations and stewardship into their
management practices. Fin damage is a welfare problem and can
167
be acerbated by a number of on-farm risk factors such as high
stocking density (North et al., 2006) or handling (Bosakowski
and Wagner, 1994). This experiment suggests that a farmer can
reduce the incidence of dorsal fin erosion by feeding fish on-
demand. Dorsal fin splitting was also less prevalent in fish fed
on-demand. However, this difference was also discernible at the
start of the study and may have been related to fish having the
opportunity to feed on-demand during the pre-study condition-
ing period. Other comparable studies on cage-held Atlantic
salmon parr (Noble et al., 2007a) and post-smolts (Noble et al.,
2007b) also report reduced fin damage in fish fed on-demand.
At the end of Parts I and II, the smallest fish within each cage had
significantly more fin splitting than their conspecifics, irrespective
of feeding regime. A similar trend was also observed with respect to
fin erosion, although it did not become significant until the end of
Part II. This suggests that the smallest fish within each group are the
greatest recipients of aggression, a finding supported by other
studies (Damsgård et al., 1997; Moutou et al., 1998). In a
comparable study using 1+ parr (Noble et al., 2007a), there was no
relationship between fish size and fin damage at the same time of
the year. However, when comparing the timing of smolt transfer
(UK 0+ and 1+ pre-smolts are transferred to sea during November
and April, respectively) the smallest fish exhibited significantly
more fin damage during the month prior to seawater transfer.
Daily ration in fish under the on-demand regime varied between
days (see also Blyth et al., 1993) and decreased as winter
approached. This was significantly related to change in tempera-
ture in all cages, although there were strong cross-correlations
between temperature and both daylength and change in daylength.
Additional variation was positively correlated with wind speed
(used as an index for wind-driven water conditions at the cage site)
for all cages (George, 1981). Other studies have reported that
temperature regulates feed intake in Atlantic salmon (Brett, 1979;
Noble et al., 2007c) and other species (Azzaydi et al., 1998; Bégout
Anras, 1995), although the latter three studies reported that
daylength had an additional effect. In the current study, increased
food delivery on windy days may be a response to increased feed
consumption at higher water velocities (Jørgensen and Jobling,
1993). Alternatively, increased feed delivery may be related to
both the nature of cage culture and the feeding system, which relies
upon waste pellet detection to terminate feed delivery. During
periods of high water velocity, pellets can be carried away from
both the cage and the pellet sensor (pers. obs.) driving the system to
step up feed delivery (under the assumption that fish are hungry)
when fish may actually be satiated.
Under 24 h light, fish consumed only 30–40% of their daily
ration during the illuminated nocturnal period. Water tempera-
ture during this period was around 11 °C and near, but not
below, the temperature threshold when parr switch from being
diurnal to nocturnal (Fraser et al., 1993). This suggests some
fish may be taking the opportunity to feed during the
illuminated nocturnal period around this time. Further, fish
fed on-demand do not appear to consume more food during the
extended photoperiod and only extend the time required to
consume their daily ration. However, the 10 day monitoring
period may have been too short to detect a change in feed
demand.
168 C. Noble et al. / Aquaculture 275 (2008) 163–168
In summary, when fish were fed to satiation, feeding regime
had no effect on growth rate, FCR or a number of other production
variables and ration was closely matched between regimes.
However, when cage-held pre-smolts were subjected to a
moderate period of feed restriction (ca. 80% of satiation for
b 1 month) it had a significant and detrimental effect upon fish
condition, growth and FCR. Aquaculturists should be wary of this
when implementing feed management strategies that incorporate
periods of feed restriction. Feeding fish on-demand also reduced
the incidence of dorsal fin erosion, which may improve the
marketability and consumer perception of farmed fish. Variability
in the daily ration of fish fed on-demand was significantly related
to water temperature and wind speed, and fish fed on-demand
consumed only 30–40% of their total daily ration during the
illuminated nocturnal period when held under 24 h light.
Acknowledgments
This study was funded by The Biotechnology and Biological
Sciences Research Council (BBSRC) and Marine Harvest
Scotland. The authors would like to thank David Gray, Mark
Fowler and all the staff at Glenfinnan for excellent fish
husbandry, and everybody who helped with measuring the fish.
References
Ang, K.P., Petrell, R.J., 1997. Control of feed dispensation in seacages using
underwater video monitoring: effects on growth and food conversion.
Aquac. Eng. 16, 45–62.
Azzaydi, M., Madrid, J.A., Sánchez-Vázquez, F.J., Martínez, F.J., 1998. Effect
of feeding strategies (automatic, ad libitum demand-feeding and time-
restricted demand-feeding) on feeding rhythms and growth in European sea
bass (Dicentrarchus labrax L.). Aquaculture 163, 285–296.
Bégout Anras, M.L., 1995. Demand feeding behaviour of sea bass kept in
ponds: diel and seasonal patterns, and influences of environmental factors.
Aquac. Int. 3, 186–195.
Blyth, P.J., Purser, G.J., Russell, J.F., 1993. Detection of feeding rhythms in
seacaged Atlantic salmon using new feeder technology. In: Reinertsen, H.,
Dahle, L.A., Jørgensen, L., Tvinnereim, K. (Eds.), Fish Farming
Technology. Balkema, Rotterdam, pp. 209–216.
Bosakowski, T., Wagner, E.J., 1994. A survey of trout fin erosion, water quality,
and rearing conditions at state fish hatcheries in Utah. J. World Aquac. Soc.
25, 308–316.
Brett, J.R., 1979. Environmental factors and growth. In: Hoar, W.S., Randall, D.J., Brett,
J.R. (Eds.), Fish Physiology, vol. VIII. Academic Press, New York, pp. 599–675.
Bureau, D.P., Hua, K., Cho, C.Y., 2006. Effect of feeding level on growth and
nutrient deposition in rainbow trout (Oncorhynchus mykiss Walbaum)
growing from 150 to 600 g. Aquac. Res. 37, 1090–1098.
Cho, C.Y., 1992. Feeding systems for rainbow trout and other salmonids with
reference to current estimates of energy and protein requirements. Aquaculture
100, 107–123.
Cho, C.Y., Bureau, D.P., 1998. Development of bioenergetic models and the
Fish-PrFEQ software to estimate production, feeding ration and waste
output in aquaculture. Aquat. Living Resour. 11, 199–210.
Cutts, C., Metcalfe, N.B., Taylor, A., 1998. Aggression and growth depression
in juvenile Atlantic salmon: the consequences of individual variation in
standard metabolic rate. J. Fish Biol. 52, 1026–1039.
Damsgård, B., Arnesen, A.M., Baardvik, B.M., Jobling, M., 1997. State
dependent feed acquisition among two strains of hatchery-reared Arctic
charr. J. Fish Biol. 50, 859–869.
Davis, M.W., Olla, B.L., 1987. Aggression and variation in growth of chum
salmon juveniles in seawater: effect of limited rations. Can. J. Fish. Aquat.
Sci. 44, 192–197.
Forsberg, O.I., Guttormsen, A.G., 2006. The value of information in salmon farming.
Harvesting the right fish at the right time. Aquac. Econ. Manag. 10, 183–200.
Fraser, N.H.C., Metcalfe, N.B., Thorpe, J.E., 1993. Temperature-dependent
switch between diurnal and nocturnal foraging in salmon. Proc. R. Soc.
Lond. B 252, 135–139.
Gaylord, T.G., MacKenzie, D.S., Gatlin, D.M., 2001. Growth performance, body
composition and plasma thyroid hormone status of channel catfish (Ictalurus
punctatus) in response to short-term feed deprivation and refeeding. Fish
Physiol. Biochem. 24, 73–79.
George, D.G., 1981. Wind-induced water movements in the South Basin of
Windermere. Freshw. Biol. 11, 37–60.
Gregory, T.R., Wood, C.M., 1999. Interactions between individual feeding behaviour,
growth, and swimming performance in juvenile rainbow trout (Oncorhynchus
mykiss) fed different rations. Can. J. Fish. Aquat. Sci. 56, 479–486.
Johansen, S.J.S., Jobling, M., 1998. The influence of feeding regime on growth
and slaughter traits of cage-reared Atlantic salmon. Aquac. Int. 6, 1–17.
Jørgensen, E.H., Jobling, M., 1993. The effect of exercise on growth, food
utilisation and osmoregulatory capacity of juvenile Atlantic salmon, Salmo
salar. Aquaculture 116, 233–246.
Li, H.W., Brocksen, R.W., 1977. Approaches to the analysis of energetic costs of
intraspecific competition for space by rainbow trout (Salmo gairdneri).
J. Fish Biol. 11, 329–341.
Moutou, K.A., McCarthy, I.D., Houlihan, D.F., 1998. The effect of ration level
and social rank on the development of fin damage in juvenile rainbow trout.
J. Fish Biol. 52, 756–770.
Noble, C., Kadri, S., Mitchell, D.F., Huntingford, F.A., 2007a. The influence of
feeding regime on intraspecific competition, fin damage and growth in 1+
Atlantic salmon parr (Salmo salar L.) held in freshwater production cages.
Aquac. Res. 38, 1137–1143.
Noble, C., Kadri, S., Mitchell, D.F., Huntingford, F.A., 2007b. The effect of feed
regime on the growth and behaviour of 1+ Atlantic salmon post-smolts (Salmo
salar L.) in semi-commercial sea cages. Aquac. Res. 38, 1686–1691.
Noble, C., Kadri, S., Mitchell, D.F., Huntingford, F.A., 2007c. The impact of
environmental variables on the feeding rhythms and daily feed intake of cage-
held 1+ Atlantic salmon parr (Salmo salar L.). Aquaculture 269, 290–298.
North, B.P., Turnbull, J.F., Ellis, T., Porter, M.J., Migaud, H., Bron, J., Bromage,
N.R., 2006. The impact of stocking density on the welfare of rainbow trout
(Oncorhynchus mykiss). Aquaculture 255, 466–479.
Pierce, A.L., Beckman, B.R., Shearer, K.D., Larsen, D.A., Dickhoff, W.W.,
2001. Effects of ration on somatotropic hormones and growth in coho
salmon. Comp. Biochem. Phys. B 128, 255–264.
Quinton, J.C., Blake, R.W., 1990. The effect of feed cycling and ration level on
the compensatory growth response in rainbow trout, Oncorhynchus mykiss.
J. Fish Biol. 37, 33–41.
Rasmussen, R.S., Ostenfeld, T., McLean, E., 2000. Effect of growth rate on quality
traits and feed utilisation of rainbow trout and brook trout. Aquaculture 184,
327–337.
Smith, I.P., Metcalfe, N.B., Huntingford, F.A., Kadri, S., 1993. Daily and
seasonal patterns in the feeding behaviour of Atlantic salmon (Salmo salar)
in a sea cage. Aquaculture 117, 165–178.
Storebakken, T., Austreng, E., 1987. Ration level for salmonids, I. Growth,
survival, body composition, and feed conversion in Atlantic salmon fry and
fingerlings. Aquaculture 60, 189–206.
Sveier, H., Lied, E., 1998. The effect of feeding regime on growth, feed
utilisation and weight dispersion in large Atlantic salmon (Salmo salar)
reared in seawater. Aquaculture 165, 333–345.
Talbot, C., Corneillie, S., Korsøen, O., 1999. Pattern of feed intake in four
species of fish under commercial farming conditions: implications for
feeding management. Aquac. Res. 30, 509–518.
Thorpe, J.E., Cho, C.Y., 1995. Minimising waste through bioenergetically and
behaviourally based feeding strategies. Water Sci. Technol. 31, 29–40.
Turnbull, J.F., Richards, R.H., Robertson, D.A., 1996. Gross, histological and
scanning electron-microscopic appearance of dorsal fin rot in farmed
Atlantic salmon, Salmo salar L., parr. J. Fish Dis. 19, 415–427.
Winfree, R.A., Kindschi, G.A., Shaw, H.T., 1998. Elevated water temperature,
crowding, and food deprivation accelerate fin erosion in juvenile steelhead
trout. Prog. Fish Cult. 60, 192–199.