Energy Requirements and Feeding

Behaviour of Salmonids in Culture

Jason Bailey
Department of Aquaculture
Umeå, Sweden

Doctoral Thesis
Swedish University of Agricultural
Sciences
Umeå 2003
Abstract
Bailey, J. 2003. Energy requirements and feeding behaviour of salmonids in culture.
ISSN, 1401-6230, ISBN 91-576-6348-3

This thesis deals with the feeding management of salmonids, from the calculation of the
daily feed requirement to the presentation and delivery of that ration taking into
consideration the feeding behaviour. The digestible energy need (DEN) to grow 1 kg of wet
body weight was calculated for different farmed fish species. In general, the DEN increased
with increasing body weight. Using this information, a feed ration model was created and
tested using rainbow trout (Oncorhynchus mykiss). Fish fed using this model showed a high
rate of growth and a good feed conversion ratio (FCR). The model accurately predicted
growth in trout and can be easily applied as a method of calculating the daily ration. For
comparison, 3 groups were fed using self-feeders. These groups showed a pattern of
fluctuating trigger-biting activity across days and this type of pattern was further studied
using time series analysis for groups of rainbow trout. Peaks in activity occurred in the
morning and evening at lights on and lights off. Across days, peaks in activity are
significantly higher every second day. This pattern is probably a result of an “hourglass”
mechanism such as the time required for gastric evacuation and the return of appetite.
Knowing how much feed to give each day, one must then present it in an optimal way.
Rainbow trout were fed the daily ration using various portion sizes and at different
temperatures. At 5 ºC, growth and FCR were best when small portion sizes were used. At
higher temperatures, good growth rates and FCR were observed over a broader range,
indicating a high level of plasticity in the trout’s ability to adjust to different portion sizes at
higher temperatures. To further evaluate feed presentation, Atlantic salmon (Salmo salar
L.) were raised in sea cages, and fed using different combinations of pellet sizes and
delivery rates. Treatment groups were compared using growth rate. No significant linear
effect on growth due to pellet size or delivery rate could be detected. For pellet size,
however, indications point to a non-linear relationship with growth rate. Salmon grew
equally well irrespective of treatment further indicating a high level of plasticity in the
salmon’s ability to adjust to changes in feed presentation.

Keywords: feed management, energy requirements, feed budget, self-feeders, feeding

behaviour, feeding intensity, portion size, pellet size

Author’s address: Jason Bailey, Department of Aquaculture, Swedish University of

Agricultural Sciences, S-901 83 Umeå, Sweden.
To Albert & Arthur
‘Tis in me blood’
Contents
Introduction, 7
The Daily Ration, 7
Variation in Feed Intake, 10
Feed Presentation, 11
Delivery Rate, 11
Portion Size, 12
Pellet Size, 12
Objectives and Aims, 13

Materials and Methods, 14

Fish and Rearing, 14
DEN Calculation, 14
Feed Budget Model, 15
Time Series, 15
Feed Presentation, 15
Feed Portion Size, 15
Feed Delivery Rate and Pellet Size, 15

Results – Summary of Papers, 16

Discussion, 17

Conclusions, 23

References, 24

Acknowledgements, 28
Appendix

Papers I-V
The present thesis is based on the following papers, which will be referred to by
their Roman numerals:

I. Bailey, J. & Alanärä, A. Digestible energy need (DEN) of selected farmed fish
species. Manuscript.

II. Bailey, J. & Alanärä, A. 2001. A test of a feed budget model in rainbow trout,
Oncorhynchus mykiss (Walbaum). Aquaculture Research 32: 465-469.

III. Bailey, J. & Alanärä, A. Mapping the demand-feeding pattern of hatchery-

reared rainbow trout, Oncorhynchus mykiss (Walbaum). Manuscript.

IV. Bailey, J. & Alanärä, A. Effect of feed portion size on growth of rainbow
trout, Oncorhynchus mykiss (Walbaum), reared at different temperatures.
Submitted.

V. Bailey, J., Alanärä, A. & Crampton, V. Does delivery rate and pellet sizes
affect growth rate in Atlantic salmon (Salmo salar L.) raised under semi-
commercial farming conditions? Accepted in Aquaculture.

Papers II and V are reproduced with permission of the journal concerned.

Introduction
The food and agriculture organization (FAO) of the United Nations estimated
aquaculture production in 1999 to be 32.9 million metric tonnes (mt) with the
farming of salmonids making up approximately 900,000 mt (or 2.7 %) (FAO,
2000). Norway and Chile lead the way in production of mainly Atlantic salmon
(Salmo salar L.) with 462,000 mt and 222,900 mt respectively. Norway’s salmon
exports of 8.3 million kilograms to the United States alone in 1999 held a value of
51 million US$ (FAO, 2000). Slight changes in production efficiency of an
industry of this magnitude will mean profits of millions of dollars. Feed costs
account for about 50% of the total yearly production costs on large commercial
farms. More efficient production will translate into more biomass for a smaller
amount of feed. This will lead to further reductions in environmental impacts and,
therefore, increased economic gains. The aquaculture industry is one of the few
where direct economic gains can be realised through a decrease in pollution.
A fish farmer’s primary objective is to produce high quality, disease-free fish in
as short a time as possible. The three principal factors that govern this objective
are the physical environment of the fish, fish physiology, and behaviour.
Throughout development, at least one of these factors, and most often an
interaction of all three, is affecting fish growth and ultimately the cost of
production of the final product.
As stated, feed costs are by far the principal expense a fish farmer has to deal
with. Thus, it can be contended that the act of feeding is the single most important
undertaking for the economy of the farming industry. The size of the daily ration,
the nutritional and physical make-up of the feed, and the method by which the
feed is delivered can have significant effects on farm economy. This thesis deals
with some of these parameters.

The Daily Ration

If the act of feeding is the most important step for the fish farmer, then the part of
that step that includes the calculation of the daily ration is perhaps the most
difficult. An optimal ration will give maximal growth while maintaining a good
feed conversion ratio (FCR), the ratio of feed ingested to body weight gained. One
may assume that once the daily ration is known, that merely weighing up the daily
amount as a proportion of the biomass of the fish to be fed, will be a simple
procedure. However, fish species, individual fish size, water temperature, time of
year, and an array of behavioural, physiological, and environmental variables can
affect the daily and seasonal requirements of the fish (Farbridge & Leatherland,
1987; Langhans & Scharrer, 1992; Heggenes et al., 1993; Jobling, 1994; De Silva
& Anderson, 1995; Saether, Johnsen & Jobling, 1996; Tveiten, Johnsen &
Jobling, 1996; Thetmeyer et al., 1999).
Feeding fish on demand, either through fish feeding themselves (self-feeders) or
by automatic cessation or reduction of feed delivery when feeding activity
declines (interactive feedback system), has received much attention within the

7
aquaculture industry (Alanärä, Kadri, & Paspatis, 2001). Demand feeding systems
operate on the premise that the fish themselves “decide” when and how much they
will eat by providing direct and constant feedback to a computer-controlled
device. Since they incorporate feedback from the fish, they are thought to take
variations in appetite into account. These systems can, however, become
expensive and complex and are not without their own problems. It has even been
suggested that these systems may induce variation in daily appetite level (Paper
III).
An effective means of calculating the daily ration should be based on the daily
energetic requirements of the fish. Even if demand feeding systems are in place,
theoretical calculations of the daily ration should be conducted on a regular basis
as a control of the amount of feed delivered by these systems. Moreover, if feeding
models are based on accurate calculations of growth, they can also be used in the
calculation of detailed production plans, aiding farm management and planning.
Fish farmers who use automatic, timer-controlled feeders often rely on feeding
charts or growth models for calculation of the daily feed requirements of the fish (e.g.
Iwama & Tautz, 1981; Jobling, 1983; From & Rasmussen, 1984; Austreng,
Storebakken & Åsgård, 1987; Cho & Bureau, 1998). These are frequently based on
temperature and fish size and are often in very generalised form with little room
for adjustments based on local conditions and the strain of fish being farmed.
Using growth data readily available from previous production cycles, the farm
manager can easily calculate his/her own feed budget model. Since this model will
be based on local conditions, a very accurate value can be calculated for the
theoretical daily requirements of the fish for one unit of biomass gain (g or kg).
Alanärä, Kadri, & Paspatis (2001) described a feed budget model that utilises
the digestible energy need (DEN) for a 1 kg gain in wet body weight of the fish
(MJ DE·kg-1). This can be written mathematically as:

DEN =
(FI ⋅ DE ) (Equation 1)
(Wi )
where FI is the feed intake (kg), Wi is the increment of weight gain (kg), and DE
is the digestible energy content of the feed (MJ·kg-1).
The energy budget of an organism can be described as:
C=F+U+R+P (Equation 2)
where C is the energy of the feed consumed, F is the energy lost in the faeces, U is
the energy lost to excretion (urinary, gill, body surface energy loss), R is the
energy used for metabolism, and P is the remaining energy deposited as growth
(De Silva & Anderson, 1995). The energy budget can be described figuratively as
shown in Fig. 1 and the relationship between this energy budget and Equation 1
can be described as follows:

DEN =
(C − F ) (Equation 3)
(U + R + P )

8
 From a fish farmer’s point of view, the details concerning where energetic
expenditures occur are less important (Fig. 1). What is important is obtaining the
maximum return (gain in biomass) per unit input (feed intake). The advantage
with this view of the energy budget is that, since the values for F, U, and R need
not be quantified, the fish can be raised under normal culture conditions when the
estimation of DEN is performed. By doing so, energetic costs due to the ingestion
and digestion of feed, and to swimming activity are more similar to those used in
culture than to those used when a complete energy budget model, as that described
in Equation 2, is used (Alanärä, Kadri, & Paspatis, 2001).

Feed Intake, C

FI*DE Faeces, F

Digestible Energ y (DE)

Urinary Loss
Gill Excretion U

Metabolisable Energy (ME)

Heat Energy
•Digestion & Absorption
•Product Formation
Wi •Waste Formation & Excretion

R
Net Energy (NE)

Net Energy for Maintenance

•Bas al Metabolism
•Voluntary Activity
•Thermal Regulation

Net Ene rgy for Production (NEP) , P

Fig. 1. Energy flow in fish. Arrows indicate the direction of energetic transfer. Large
brackets to the left show the parts of the energy budget encompassed by the equation
DEN=(FI*DE)/Wi (see text). C, F, U, R and P make up the energy balance indicated by the
equation: C=F+U+R+P, where C is the energy of the feed consumed, F is the energy lost
in the faeces, U is the energy lost to excretion (urinary, gill, body surface energy loss), R is
the energy used for metabolism, and P is the remaining energy deposited as growth (De
Silva & Anderson, 1995).
Due to its ease and relatively low cost of calculation, the DEN can be estimated
for a particular strain of fish and under local farm conditions, providing the farmer
with a very specific method of evaluating the energetic requirements of the fish.

9
Variation in Feed Intake
Demand feeders have been used extensively in behavioural studies on salmonids
(Landless, 1976; Boujard & Leatherland, 1992; Brännäs & Alanärä, 1993; Alanärä
& Brännäs, 1993; Juell, Furevik, & Bjordal, 1993; Alanärä, 1994; Alanärä &
Brännäs, 1996; Sanchez-Vazquez & Tabata, 1998). Time series plots of the
feeding activity data obtained from these kinds of studies often display a variable
pattern of peaks and troughs across days (Boujard & Leatherland, 1992; Brännäs
& Alanärä, 1993; Alanärä & Brännäs, 1993; Alanärä, 1994; Alanärä & Kiessling,
1996). In general, peaks observed in the series on one day, are followed by
troughs on the following day. It appears that these peaks in activity occur every
second day (Paspatis & Boujard, 1996; Alanärä, Kadri, & Paspatis, 2001) and that
perhaps the demand feeding system itself may be causing this pattern through
changes in stomach fullness and the time required for emptying (gastric
evacuation time) (Grove, Loizides & Nott, 1978; Bromley, 1994).
Trout fed using demand feeders may also be consuming more than they need to
cover their daily metabolic requirements (Bres, 1986). Appetite is controlled partly
by stretch receptors in the gut and partly by the level of various metabolites in the
blood system (Grove, Loizides & Nott, 1978; Dill, 1983; Juell et al., 1994; De
Silva & Anderson 1995). It has been suggested that if fish eat until the gut is full
of pelleted feed, much of the energy might be unused and excreted undigested or
stored as excess visceral and muscular lipid (Jobling, 1986; Storebakken et al.,
1991) although no studies have been conducted to fully support this suggestion.
Bres (1986) and Alanärä (1994) showed that rainbow trout were unable to
distinguish between pellets of different energetic contents and were therefore
unable to, or do not need to regulate their daily energetic intake in the range
studied.
The high-energy pelleted feed used in salmonid aquaculture has a very low
volume to energy ratio. Pellets used in today’s aquaculture industry have as high
as 4 or 5 times the energy value for an equivalent volume of wild food (Ruohonen
& Grove, 1996) due, in part, to the low water content of pelleted feed. It follows
logically that fish fed high energy diets require less feed in terms of volume, than
do those fed low energy diets. However, fish fed using demand feeders often eat to
satiation and the time required to digest this feed might be very long. Ruohonen,
Grove, & McIlroy (1997) suggested that the longer time required for the gastric
evacuation of pelleted feed is due to the time needed to moisten the feed in the
stomach, creating a high lag time before digestion can continue. The variation in
feed intake seen in rainbow trout fed using demand feeders, then, could be due to
the time required for gastric evacuation and the return of appetite. If the time
required for gastric evacuation is more than 24 hours as suggested by Ruohonen,
Grove, & McIlroy (1997) and Grove, Loizides & Nott (1978), then the pattern of
daily activity should reveal a period between peaks of 2 days. This may have
implications on how the feed should be delivered using demand feeders and
further strengthens the necessity of using a model to calculate energy need, even if
demand feeders are in use. Perhaps restrictions on the amount available per day
should be applied using the feed budget model as a guide so as to prevent
excessive wastage.

10
 In addition to the pattern in daily activity levels, recurring activity patterns have
been shown to occur within days (Cuenca & de la Higuera, 1994; Alanärä &
Brännäs, 1997; Sanchez-Vazquez & Tabata, 1998; Bolliet, Aranda & Boujard,
2001). These studies show that peaks in activity usually occur under conditions of
changing light levels, at dawn and at dusk, probably as a result of evolutionary
selection for activity when there is increased prey abundance and decreased
predation risk (Eriksson & Alanärä, 1992). Some fish farmers have decided to
adapt their feeding regimes around these patterns in activity, feeding the daily
ration in the morning and evening, while others feed constantly throughout the
day.

Feed Presentation
Alanärä, Kadri & Paspatis (2001) described a meal as the amount of food
consumed in a single feeding bout, usually with a duration of between 30 min and
2 h. A meal can be made up of one to many “feed portions”, consisting of short,
periodic feed delivery periods followed by a short pause. The size, rate of delivery
and temporal spacing of these portions will affect the meal length and/or the
feeding intensity of the daily ration. As these definitions are interrelated, a change
in one parameter in the schedule will affect each of the other parameters.

Delivery Rate
Using a model to calculate the daily feed ration one must still face the problem of
how to deliver the feed so that the fish can utilise it in an optimal way. Feeding
schedules may vary in number of meals per day, number of portions in each meal,
the time between portions and meals, the size of each portion, and the speed of
delivery of each portion.
The delivery rate can be described in two ways: (1) the number and temporal
spacing of portions given per day, often referred to as the feeding intensity, or (2)
the delivery rate of the feeder itself, measured in number of pellets per unit time.
Both approaches will likely affect the ability of the fish in catching pellets and
consequently, growth, and so both should be considered when presenting feed to
salmonids.
The number of feed items an individual fish can catch is related to its swimming
activity and the temperature of the water (Brett, 1969; Alanärä, 1994; Jobling,
1994; Tang & Boisclair, 1995). In salmonids, activity level increases and the
proportion of time spent hiding during daylight hours decreases as temperatures
rise between ca. 4-8 ºC (Cunjak, 1988; Elliott, 1975; Fraser, Metcalfe & Thorpe,
1993; Fraser et al., 1995; Graham, Thorpe & Metcalfe, 1996; Rimmer, Saunders
& Paim., 1985) dependent, of course, upon the acclimation history of the fish and
the strain or species being considered. In aquaculture, decreased swimming
activity may affect the rate of feed capture and feed delivery rate, therefore, should
perhaps reflect this change in activity level at different temperatures.

11
 Portion Size

With respect to portion size, fish are poikilothermic and at low temperatures,
when fish are less active, many small feed portions will likely allow the fish to
catch a larger proportion of the feed than if large portion sizes are used. At higher
temperatures, the fish are more active and will likely be able to catch all pellets in
the feed portions up to a point where the portion size becomes too large and waste
occurs. It is suspected that the relationship between feed portion size and growth
rate follows a quadratic function as shown in Fig. 2. Assuming that the ability to
catch pellets is limited by the activity level of the fish, at different temperatures the
optimal portion size should shift along the x-axis (Fig. 2). In addition, because
swimming activity increases, the plateau of the quadratic function flattens as a
broader range of portion sizes (i.e. larger and larger portion sizes) give equivalent
growth rates as small portion sizes, assuming fish receive the same amounts of
feed each day and energetic expenditures associated with feeding under longer
periods are relatively insignificant.

Increased Temperature
Low Temperature
Growth Rate (TGC)

Portion Size
Fig. 2. Schematic relationship showing the effect of changes in temperature on the range of
portion sizes giving good growth rates. At the low end of the normal range of growing
temperatures, small portion sizes give highest biomass increases (solid curve). As
temperatures increase, the range of portion sizes giving good growth increases as fish
activity increases and more and more pellets in the larger portion sizes can be caught
(dashed line).

Pellet Size
The size of the feed pellets and the rate at which they are delivered should also
affect the amount of feed an individual fish can ingest over a period of time. The
effects of pellet size on growth or feeding behaviour has been examined in

12
juvenile to adult (over 3 kg) Atlantic salmon (Stradmeyer, Metcalfe & Thorpe,
1988; Smith, Metcalfe & Huntingford, 1995) and Arctic charr (fork length 20-24
cm) (Linnér & Brännäs, 1994). Atlantic salmon appear to be rather plastic in their
behavioural responses to different pellet sizes. In salmon a more immediate
response to larger pellets was observed when compared to smaller pellets, but the
former were more likely to be rejected than the latter. Smith, Metcalfe &
Huntingford (1995) also found that pellets slightly smaller than the “normal”
commercial size were eaten at the fastest rate, thereby indicating that the salmon
are perhaps adjusting their feeding behaviour to compensate for a smaller feed
pellet size. For Arctic charr reared under hatchery conditions, the optimal pellet
size has been suggested to be around 2% of the fish’s length (Linnér & Brännäs,
1994). Others have suggested that for a range of fish species, the optimal feed size
appears to be 25-50% of the mouth width (Wankowski, 1979; Tabachek, 1988).
Pellets of sub-optimal size or pellets that are delivered at a high rate may cause
wastage, as fish may be unable to catch large numbers of pellets before they sink
through the net pen.
As a result of suggestions for an “optimal” pellet size based on fish length,
salmonid farmers use many different sizes of feed pellets during the growout
period. Each time the size is changed new calculations for the optimal number of
pellets per fish and delivery must be done. It would therefore save both time and
money if fewer sizes of pelleted feed can be used for larger parts of the growout
period. Fewer sizes of pellets will also mean lower warehouse costs and more
storage space for the farm operators as well as a reduced working capital for the
salmon farming business.

Objectives and Aims

The objective of this thesis was to develop and test practical methodology in the
various stages of feeding management for salmonid culture, specifically the
calculation and presentation of the daily feed ration. In papers I-V, I have
attempted to:
1) develop and test a reliable and practical method of calculating the daily
ration of farmed fish using rainbow trout, Oncorhynchus mykiss
(Walbaum), as a test species.
2) map the feeding activity patterns of rainbow trout fed using self-feeders
and held under hatchery conditions.
3) examine the effects of altering methods of delivery of the daily ration under
hatchery and commercial conditions using variation in pellet size, feed
portion size, and delivery rate.

13
Materials & Methods

Fish and Rearing

Studies II and IV were conducted at UMF (Umeå Marine Research Centre;
63°35'N, 19°50'E) using rainbow trout under hatchery conditions in small tanks.
Study V was carried out using Atlantic salmon in Norway and under semi-
commercial farming conditions in sea cages. It was conducted under 2 trials and
over a period of 2 years with salmon ranging in size between 1.25 and 5.0 kg.
Rainbow trout used in the experiments (Papers II, III, and IV) weighed between
30-500g.

DEN Calculation
The digestible energy need (DEN) was required as part of the feed budget model
tested in Paper II. For paper II, 12 trout were held individually in 170 l aquaria.
These were individually hand-fed twice per day to satiation for three weeks at
10°C and feed intake (FI) was registered. The increment of weight gain (Wi) for
these fish was measured. The digestible energy (DE) content of the feed (MJ·kg-1)
was estimated from its nutrient composition, supplied by the manufacturer, and
apparent nutrient digestibility coefficients (ADC). The following values, from
digestibility studies with Atlantic salmon, were used for calculation: protein, 20.9
kJ·g-1; fat, 35.1 kJ·g-1; and carbohydrate, 11.0 kJ·g-1 (Hillestad, Åsgård & Berge,
1999). Based on Equation 1, the DEN was calculated to be 12.4 MJ DE·kg-1 for
the strain of rainbow trout used. In comparison, the calculated DEN for a rainbow
trout strain of similar size in Canada was found to be approximately 15 MJ DE ·
kg-1 (Cho & Bureau, 1998).
Due to ontogenetic changes in metabolism and body composition (Jobling,
1994), the DEN changes as fish grow. A juvenile fish has the possibility to grow
faster and to use less energy to achieve this growth than an adult fish. In order to
fully evaluate the relationship between body size and DEN for different fish
species, a literature search was performed (Paper I). FI, average weight (We), DE,
ADC, temperature, and species data were collected. DEN was calculated using
equation 1 above, for 26 different species important for aquaculture.
DE content was chosen as the energy unit as it is a more reliable measurement to
use in nutrition and production studies than gross energy or metabolisable energy
(Jobling, 1983b). Apparent digestibility coefficients were collected for a large
number of species, but the values varied markedly both within and between
species (see appendix II) depending on experimental conditions and the content of
the feed. Therefore, for estimating the digestible energy in the feed, the ADC
values from Hillestad et al. (1999) were also used here. These values lie within the
range of ADCs found in the literature for carnivorous fish and have been generally
accepted by the Norwegian salmon farming industry (Hillestad et al., 1999).
However, from a practical point of view, and in lieu of the fact that variation in the
ADCs and calculation of DE may be difficult, the GE might also be a suitable
alternative for fish farmers.

14
Feed Budget Model
A feed budget model, which enables farmers to adapt a general model to local
conditions, was tested on rainbow trout (Paper II). The model uses the theoretical
daily energetic requirements of the fish and an estimation of growth to calculate
the daily ration required. The advantage is that this model can be adjusted in
relation to prevailing environmental conditions and the specific species or strain
being farmed. Fish fed using self-feeders were used as a comparison to the results
obtained using the model.

Time Series
Rainbow trout were organised into 10 groups fed using self-feeders (Paper III).
The feeders incorporated the use of a trigger suspended 2 cm below the water
surface. Biting the trigger caused a small feed portion to be delivered to the fish.
Fish were allowed to feed unrestricted for about 60 days. Using the information
collected in this way, within- and between-day trigger-biting datasets were
prepared. The Fast Fourier Transform (FFT) (Cooley & Tukey, 1965) time-series
analysis was used to analyse the trigger-biting activity of groups of fish on a daily
and hourly scale (Paper III).

Feed Presentation
Groups of rainbow trout (Paper IV) and Atlantic salmon (Paper V) were subjected
to variations of the method of presentation of the daily feed ration.

Feed Portion Size

In Paper IV, rainbow trout were fed using different portion sizes, measured in
number of pellets per fish in each portion over a 25-day period and growth rate,
FCR, and coefficient of variation of growth rate, were compared between groups.
This was repeated for temperatures of 5, 10, and 15 ºC and any changes in
performance with temperature were analysed.

Feed Delivery Rate and Pellet Size

Atlantic salmon reared on a commercial farm site in Norway were arranged into
14 groups with various combinations of pellet sizes (0.03 - 2.68g) and delivery
rates (82.5 - 1320 g·min-1) (Paper V). The growth rate and FCR were used to
compare differences between treatments.

15
Results – Summary of Papers
Paper I makes use of data from studies on various fish species common to
aquaculture. The FI, We, DE, ADC, temperature, and species were collected from
scientific journal articles and used to examine the relationship between DEN, and
body size and temperature for different species. There seems to be a positive
relationship between DEN and log body weight for salmonids and pleuronectids
while no such relationship was found for percid fishes. In addition, no effect of
temperature was found for either of the salmonid, pleuronectid, or percid Orders.
When examined by species, a significantly positive, linear relationship could be
found between the body weight and the DEN for rainbow trout, Atlantic salmon,
Atlantic halibut (Hippoglussus hippoglossus), and Arctic charr. No linear models
were found to relate body weight, temperature, or a combination of both body
weight and temperature to the DEN for sea bass (Dicentrarchus labrax L.), sea
bream (Sparus aurata L.), turbot (Scophthalmus maximus L.) or tilapia
(Oreochromis spp.).
In order to explore the effect of strain, a separate regression relating body weight
to DEN was calculated using “Scandinavian” rainbow trout. Different strains of
rainbow trout appear to have different abilities in converting the energy content of
the feed into body mass gain, with data from Scandinavia being superior over data
from North America. In addition, data taken from Larsson (2002) and Thodessen
et al. (1999) and used to calculate the DEN, suggest that there may be an effect of
selection on the utilisation of energy in feed. Larsson (2002) showed 2 strains of
Arctic charr having different mean DEN and Thodessen et al. (1999) indicate that
wild Atlantic salmon have a higher DEN than cultured strains.
Paper II was designed to test a feed budget model, which incorporated the idea
of DEN as described above, and which farmers can use and adapt to local
conditions. Rainbow trout fed using the model as a guide had a high rate of growth
and good feed conversion. After 49 days the model accurately predicted growth of
300g rainbow trout to within 5g of the measured value. Growth of fish fed using
self-feeders is presented for comparison with the results obtained using the model.
During both growth periods, the fish fed by automatic feeders according to the
model had higher rates of growth than those fed using self-feeders. The daily
ration for self-fed groups in the second part of the experiment fluctuated around
the ration level predicted by the model with what looked like an apparent
frequency of about 2 days between peaks in activity.
In Paper II, a subjective analysis was made suggesting that the daily trigger-
biting activity for self-fed groups fluctuated around the ration level predicted by
the feeding model at a frequency of about 2 days. Paper III was designed to test
this observation quantitatively using time series analysis. The self-feeding activity
for groups of rainbow trout showed a repeating pattern recurring every second
day. FFT analyses revealed peaks in activity for free-running rainbow at
frequencies between 0.312 and 0.469. The period, P, is defined as the length of
time required to complete one cycle and is the reciprocal of the frequency, υ. This
corresponds to periods between peaks in activity in a range of 2.1 and 3.2 days

16
(mean 2.4). Period values between 2.1 and 2.5 days are likely the result of leakage
and should, therefore be considered as a period of 2. Although padding the series
to 64 will help reduce the effects of leakage, a loss of power from frequency bands
to adjacent spectral lines due to the limited number of frequencies over which the
periodogram is estimated is inevitable.
When examined hourly, peaks in activity occurred at a period of 24 hours
corresponding to the two crepuscular peaks in activity recurring at a period of 24
hours each. Two daily peaks in trigger activation occur at about lights on (dawn)
and lights off (dusk).
In Paper IV, groups of 125 rainbow trout were fed using various portion sizes
(0.23 – 4.29 pellets·fish-1·portion-1) at rearing temperatures of 5, 10, and 15 ºC. It
should be pointed out, however, that because this study did not use replicate
groups, the results should be taken as preliminary indications in this question. The
study does give an indication that portion size influences growth and variation in
growth between individuals as temperatures change. At 5 ºC, good growth and
FCR were observed using a low portion size. However, the range of portion sizes
where good growth was observed expanded with temperatures of 10 ºC and 15 ºC
(0.52 - 3.11 pellets·fish-1), possibly suggesting a high level of plasticity in the
trout’s ability to adjust to different portion sizes at higher temperatures.
In Paper V salmon were raised in commercial cages and fed using 14 different
combinations of pellet sizes and delivery rates. No consistent pattern with respect
to TGC and pellet size or delivery rate could be found using multiple regression
analysis. Salmon performed equally well irrespective of treatment, having similar
final sizes at the end of the experiment. Atlantic salmon showed a high level of
behavioural plasticity in their ability to adjust to changes in pellet size and
delivery rates without experiencing negative affects on growth or survival. That
said, indications of a small effect of pellet size on the mean TGC for both trials
were seen when a second-order plot was drawn of this data.

Discussion
The digestible energy need (DEN) changed with changes in body size for
salmonid and pleuronectid fishes (Paper I) whereas no such relationship was
found for percid fishes. The reason for this is unknown and an attempt to draw any
broadly encompassing conclusions based on the limited information available will
not be attempted here. A wide variety of uncontrollable variables as well as the
relatively low number of samples likely contribute to the lack of relationship. In
fact, it is suspected that such a relationship does exist for all fishes since, as stated,
metabolic rates follow such an ontogenetic pattern (Jobling, 1994). Although
metabolic processes are not directly measured for the calculation of DEN, it is
understood that these metabolic processes are included in the final value of the
DEN. Therefore, since metabolism has been shown to change exponentially with
temperature (Brett, 1964; Brett & Groves, 1979) and allometrically with body
weight (Jobling, 1994) for fish, DEN should also change similarly.

17
 For all species where a model could be fit, temperature had an insignificant
effect on DEN. This result can be explained by looking at the source of the growth
and feed intake information. Most of the studies examined come from aquaculture-
related studies where good growing conditions are simulated. Seldom are fish
subjected to temperature extremes unless this is the purpose of the study. The
effect of temperature around the optimal growing temperature of the fish is both
linear and very small when compared to the effect of body size (Paper I). In
addition, much more information is available in the literature for fish of different
body size than for temperatures over a wide growing range, making an effect of
body size easier to detect than an effect of temperature.
It was stated in the introduction that fish farmers could use the DEN to calculate
their own feed budget models based on previous growth records. To illustrate how
the energy requirements and daily feed allowance are calculated an example will
be given using Atlantic salmon.
Suppose the group of salmon is made up of 50 individuals weighing
approximately 200g each. The temperature of the water is 10ºC. From growth
information from previous growout years (or alternately through a preliminary
growth study), the growth rate (expressed here using the TGC) is 2.50. Using this
information the farm manager can calculate the weight of the fish after 1 day by
using the formula for growth rate (in this case the TGC):
TGC = [(BWf1/3 –BWi1/3)/ (T x D)] x 1000 (Equation 4)
and rearranging to give:
BWf = [BWi1/3 + (TGC/1000 x T x D)]3 (Equation 5)
where BWf is the final body weight, BWi is the initial body weight, T is the
temperature, and D is the number of days (in this case 1). Filling in the available
information:
BWf = [(200g)1/3 + (2.50 /1000 x 10 x 1)]3 (Equation 6)
= 202.6g
The increment of weight gain (Wi) in kg after one day then is:
Wi = BWf - BWi (Equation 7)
= 0.2026 – 0.2000 = 0.0026 kg · day-1
The farm manager can calculate the DEN for the particular strain or stock of fish
being farmed, but for simplicity the regression line for Atlantic salmon will be
used to estimate the DEN for a 200g fish (Paper I).
DEN = 10.77 + 1.05 x LN(BW) (Equation 8)
= 10.77 + 1. 05 x LN(200g) = 16.3 MJ·kg-1.
Since the DEN and the daily weight increase are known, the theoretical
energetic requirement (TER) to achieve this growth can be estimated using:
TER = 16.3 MJ · kg-1 x 0.0026 kg · day-1 (Equation 9)
= 0.042 MJ · day-1

18
 The digestible energy content of the feed was estimated as 19.0 MJ·kg-1 and so
finally, the daily feed allowance can be calculated using:
FA = (n x TER)/DE (Equation 10)
where FA is the daily feed allowance and n is the number of fish, giving:
FA = (50 x 0.042)/19.0 (Equation 11)
= 0.111 kg = 111g day-1
Table 1 shows an example of a feeding chart for Atlantic salmon. Feeding charts
such as this one can be easily created for the particular species and stock of fish
being farmed, and under the local farm conditions.
Table 1. An example of a feeding chart for Atlantic salmon (Salmo salar L.) constructed
using fish sizes of 50 and 200 g, a TGC of 0.75 during autumn and 2.5 during spring, and
DE of 19 MJ · kg-1. Amounts given are in g·fish-1·day-1

Spring Autumn
Temperature 50g 200g 50g 200g

4 0.28 0.88 0.08 0.26

8 0.57 1.77 0.17 0.53

10 0.71 2.21 0.21 0.66

12 0.85 2.65 0.26 0.79

The feed ration model tested in Paper II and described above predicted the
growth of rainbow trout extremely well, and good feed conversion indicated that
there was little feed waste. These results can likely be attributed to the flexibility
of the model. Both components of the model: (1) the calculated energy needed to
grow one kg (DEN) and (2) the growth data, come from calculations using the
stock of fish used to test the model, and are based on local conditions. This is the
biggest advantage of the present model when compared to other, similar models.
Most feed budget models are based on either growth rate (e.g. Austreng,
Storebakken & Åsgård, 1987) or energy requirement (e.g. Cho, 1992) estimates,
but generally lack the flexibility to allow adjustments to be made in relation to
local conditions. In practice, fish-farmers make somewhat arbitrary adjustments,
without knowing how factors like temperature, fish growth, body composition,
season, etc. influence the feed budget during each growth period. Using a model
such as the one tested in Paper II, however, the farmer can adjust the daily ration
more often and with more certainty.
The day-to-day variation in feed demand observed in the self-fed groups in
Paper II is consistent with earlier observations on salmonids (Grove, Loizides &
Nott, 1978; Juell, Furevik & Bjordal, 1993; Juell et al., 1994). A plot of the daily
self-fed activity with the amount predicted by the model (Paper II) shows variation
of the self-fed amount around the predicted value. The reason for this variation is
unclear, but it may be related to changes in stomach fullness and the time required
for emptying (gastric evacuation time) (Grove, Loizides & Nott, 1978; Bromley,

19
1994). Using self-feeders with unlimited access to feed then, may cause fish to eat
more than is energetically required for maximal growth every second day.
Variation in feeding for groups of fish fed using automatic feeders at rates
predicted by the model were not registered, but any such variations were probably
minor since major variation would have resulted in feed waste and high FCR
values.
In Paper III the daily variation in self-feeding activity for groups of rainbow
trout was mapped using time-series analysis. It was concluded that a 2-day pattern
between peaks in daily trigger-biting activity exists in rainbow trout weighing
between 30 and 300g and held in 1m3 freshwater tanks. The implication of the 2-
day period is not that trout fast every second day. On the contrary, trout activate
the trigger every day, but the results suggest that the total biting activity is greater
every other day. Whether the rhythmic pattern seen is controlled by some kind of
internal clock or whether it is simply an artefact of the feeding system used is
unknown. Whatever the cause of the pattern, fish farmers should perhaps adjust
feed delivery to match feeding activity and thereby reduce overfeeding.
When the trigger-biting activity was examined within days, an interesting
observation was made. As expected, the morning peak in activity always appears
as trout probably initially associate the conditioned stimulus of lights on with the
availability of food. As the study progressed under consistent photoperiod
conditions, trout learned when “dawn” occurred and anticipated this by biting
before the lights came on. A second and larger peak appeared in the evening,
contrary to other studies showing the major peak occurring during the dawn in
Atlantic salmon (Kadri et al., 1991; Paspatis & Boujard, 1996). Eriksson &
Alanärä (1992) suggested that the crepuscular activity (at dawn and dusk) of
stream-living salmonids is due to a trade-off between food availability and
predation risk. They explained that locomotory activity patterns of Atlantic salmon
corresponded to organic drift of mayfly larvae. At the same time, lowered light
levels impair the ability of visual predators on salmonids. However, since salmon
are also visual predators, a trade-off must exist between predation risk and the
efficiency of catching abundant prey; that time being at dusk and dawn. Although
not the same species, the behavioural patterns observed in rainbow trout might
also share these explanations.
From a fish farming point of view, it is obvious that the best feeding times
should be at dawn and dusk, when fish-feeding activity is highest. However, fish-
feeding schedules are often built around human convenience and the fish are
therefore fed at various times of the day. It may be difficult, for example, to get to
a remote farm-site at dawn and dusk every day to feed fish. In this capacity, timer-
controlled automatic or demand-feeding systems might be used to feed at optimal
times during the day.
Paper IV indicates that portion size may affect the growth rate and food
conversion efficiency of rainbow trout held in small tanks. As temperatures
increase from 5 to 15 ºC, trout may be able to feed effectively over a broader
range of portion sizes. The group fed using 0.52 pellets·fish-1·portion-1 grew at a
high rate while maintaining good FCR throughout the study, however, due to the
confounding effect of group it is difficult to determine if this effect is significant.

20
Groups fed using between 1.89 and 3.11 pellets·fish-1·portion-1 did not grow well
and had a poor FCR at 5 ºC. When temperatures were increased fish probably
became more active and caught a larger proportion of the pellets in larger portions
as reflected by improvements in growth rate and FCR. The positive result in terms
of growth performance with increasing portion size at 10 ºC and 15 ºC would be
expected, since, like other ectothermic animals, fish become sluggish at low
temperatures. This is in agreement with studies on feeding and changes in
swimming activity at different temperatures (Elliott, 1975; Rimmer, Saunders &
Paim., 1985; Cunjak, 1988; Fraser, Metcalfe & Thorpe, 1993; Fraser et al., 1995;
Graham, Thorpe & Metcalfe, 1996).
While these results are interesting, the authors suggest some caution be taken in
interpreting them due to the confounding effect of group in the experimental
design. The results do give an indication of an effect of portion size on the growth
and FCR of small rainbow trout and so are presented here in that context.
However, further work will be required before suggestions are made as to exact
portion sizes at different temperatures.
In this study, group 2, fed using a small feed portion size, gave good growth and
FCR throughout the study. However, increasing temperatures increase the energy
and feed demand of fish, and without a corresponding change in feed portion size,
the fish-farmer is forced to feed the fish essentially throughout the entire day. A
reduced period of feeding, however, would be beneficial from a practical
viewpoint. Commercial farms using a feed barge, for example, would benefit from
reduced feeding time at every cage. Thus, a better strategy would be to feed fish
using feed portions that change in size with changes in temperature, large at high
temperatures and small at low temperatures, thereby keeping meal length
relatively constant over temperatures. In this way, the daily feed budget can be
delivered in a much shorter time without decreasing performance. The increase in
growth performance with larger portion sizes at higher temperatures as has been
found in Paper IV supports this theory.
In Paper V the effects of delivery rate and pellet size were examined on a semi-
commercial salmon farm. Fish reared in commercial aquaculture operations are
sometimes fed at a relatively low intensity over prolonged periods (Alanärä,
Kadri, & Paspatis, 2001). This ‘little-and-often’ strategy has partly been the result
of technical limitations in some of the feeding systems and partly because many
farmers think that intense feeding will result in feed waste. In Paper V, fish were
fed using one intense meal per day of approximately two hours duration. Other
studies have shown that feeding fish using 1-4 meals per day under high intensity
have resulted in good growth rates in salmonids (Elliot, 1975; Grayton &
Beamish, 1977; Jobling, 1983; Juell et al., 1994), and Cho (1990) suggested that
1-2 meals per day is sufficient for optimal growth in rainbow trout weighing over
200g. The ‘little-and-often’ feeding regime is probably associated with increased
swimming activity and energy expenditures (Alanärä, 1992; Johansen & Jobling,
1998), whereas the main positive effect of a ‘large-and-seldom’ regime is the
inclusion of longer non-feeding periods during the rest of the day, as well as the
aforementioned, practical benefits of reduced feeding time.

21
 Assuming, according to the above, that the number or length of meals decrease,
the rate at which feed is supplied must increase accordingly. Since salmon have
been shown to be extremely flexible in adopting their feeding rate to the rate of
delivery (Paper V), the fastest delivery rate that still produces little waste should
be chosen in order to fulfil this requirement. In addition, the faster the supply rate
the better in avoiding large variation in growth rates, because slow rates may allow
feed monopolisation by more competitive individuals (Grant, 1993). As shown in
Paper III, it appears that the optimal times for feeding in rainbow trout are for
short periods in the morning and evening. A fish farmer should take advantage of
this intense feeding activity period and feed at as high a rate as is possible without
causing wastage.
Growth Rate (TGC)

Pellet Size
Fig. 3. Schematic illustration of plasticity in salmon feeding behaviour. The solid line
shows the “preferred” pellet size for salmon of a given body size. The dashed line indicates
the ability of the salmon to make adjustments in their behaviour and accept pellets of an
inoptimal size without compromising growth.
As stated in the introduction, the relationship between portion size and growth
rate should follow a quadratic function. Similarly, it was hypothesised that pellet
size and growth rate should follow the same type of function. This type of plot
was examined and indications of a small effect of pellet size on the mean TGC for
both trials were seen. It is probable that this relationship exists but that the pellet
sizes tested here were not large or small enough to fully challenge the salmon’s
ability to adjust feeding activity. A “preferred” pellet size might exist but, under
conditions where this size is unavailable, salmon display their remarkable ability
to adjust, and show no decreases in growth rates (Fig. 3). The full effect of pellet
size on the growth rate may have been masked by the long growout periods in the
study and led to results seen as contradictory to previous work with Atlantic
salmon (Stradmeyer, Metcalfe & Thorpe, 1988; Smith, Metcalfe & Huntingford,
1995). Pellet sizes that were very large for a particular fish size at the beginning of
a ‘Period’ may have become only slightly oversized by the end of that same period
due to fish growth. The results from Paper V, however, are seen as a testament to

22
the extraordinary ability of Atlantic salmon to adjust to changes in feed
presentation. At some extreme, pellet size, for example, will obviously have an
affect on fish performance. A pellet that is larger than the gape width of the fish or
is so large that handling time becomes a limiting factor in the fish’s ability to
ingest enough pellets to maintain good growth will clearly have adverse effects.
Within a reasonable range of pellet sizes and delivery rates, however, large
Atlantic salmon raised in sea cages appear to adjust and grow relatively well
regardless if the size/rate is ¼ to 4 times that which is recommended by the
industry.

Conclusions
It has been emphasised throughout this thesis that the cost of feed is the highest
returning expense in salmonid aquaculture. Therefore, in order for the business to
be successful, it is paramount (1) to have access to an accurate model for
calculation of daily feed requirements, including an understanding of how these
requirements may change over time, and (2) that the feed is presented in a manner
that is acceptable to the target species in order to ensure utilisation. Uningested or
undigested feed translates directly into a reduction in the farm’s economic viability
and an increase in environmental loading.
The model used for the calculation of the daily feeding ration in Paper II was
based on the amount of energy required by that strain of fish, reared under those
particular conditions, and at that particular time of the year. The advantage with
this model is that although it is based on energetics, direct calculation of a
complete energetic budget is unnecessary. Farmers can use source information
such as that presented in Paper I to estimate the digestible energy needed for a 1
kg gain in biomass, or may calculate this value specifically for the fish species and
strain being farmed (Paper II). Information on annual growth cycles is often
available from previous production years and so the farmer can adjust accordingly
for changes in appetite throughout the year. Even if a demand feeding system is in
place, the utilisation of a model to calculate the daily ration is useful to aid in
feeding management.
As demonstrated in Paper III, fish appetite varies both within and across days. It
has also been shown that this variation occurs on a seasonal basis (Rowe &
Thorpe, 1990; Jobling & Baardvik, 1991; Eriksson & Alanärä, 1992; Thorpe,
1994; Sæther et al., 1996). Within-day variation can be easily adjusted for, simply
by feeding during times of the day when fish are most willing to accept feed. For
rainbow trout, this appears to be during short periods in the morning and evening,
under conditions of changing light (dawn and dusk). The feed ration model
presented in Paper II does not regulate daily feed intake to account for daily
variations in appetite. However, as shown in Paper II, it does calculate an amount
of feed at the approximate midpoint between the peaks and troughs of the amount
fed using demand feeders. Therefore, over a production period, the amount fed
using demand feeders and that fed using the model should be approximately equal.

23
 Papers IV and V demonstrate, probably more than anything, plasticity in
salmonids. Although the feed delivery rates and pellet sizes were varied between
¼x and 4x that considered ‘normal’, no significant effect on growth rate was
detected (Paper V). In Paper IV, a significant effect of portion size was found at 5
ºC. At low temperatures, physiology and swimming activity is reduced and so fish
have difficulty in catching all pellets in a large portion before they sink out of
reach. At higher temperatures, fish could be fed, with good production results,
using larger and larger portion sizes. It was suggested, therefore, that, if possible,
to save time and reduce the length of the stressful feeding periods, that the largest
possible feed portion be used at different temperatures.
The thesis began with some production figures presented by the FAO. It was
stated that small advances in production efficiency for such a large industry would
give large economic returns. Consider this: A farm having 12 cages each stocked
with 20,000 Atlantic salmon weighing 500g each, giving a total of 120 mt.
Production for the previous year gave an FCR of 1.0. Fish final size was 4kg
(biomass 960 mt) and total feed used throughout the year was therefore 840 mt.
Improvements in feeding management have lowered the FCR to 0.90 during the
current production year. Assuming no mortality and the same final slaughter size,
the farmer would have used 756 mt of feed; a reduction of almost 100 mt!
Considerable economic returns for small changes in feed management practices
and yet, further benefits from time saved through, for example, increased feed
delivery rates or portion sizes are not included in this simple illustration.

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Acknowledgements
Papers II and IV were funded by contributions from the European Community
Structural Funds: Financial Instrument of Fisheries Guidance (FIFG). All other
funding provided by the Oscar & Lili Lamms Minne scholarship.
Given the only space where I am risk-free in expressing my own feelings, I
choose to use the words of others. In this way unnamed names are never forgotten
but remain unnamed with the unforgotten.
“In order to make an apple pie from scratch, you must first create the universe”
(Sagan, Cosmos). Having seen that the work was already ‘published’, I proceeded
to work on my own ‘pie’. Thank-you to my friends and co-workers at the Dept. of
Aquaculture. You’ve made a foreign cook feel at home among the Swedish chefs
(and even made me learn that crazy language “U putt de morn-y mort in de fishy
dishy!”). Special thanks to the ‘maître d'Vattenbruk.’ Utan henne skulle vi inte
klara oss! “Discovery consists of seeing what everybody has seen and thinking
what nobody has thought” (von Szent-Gyorgyi). Thank you to my supervisor ‘Chef
(Ankan) Alanärä’ for sharing your ability to perform this task with me. Thanks
also to Ewos and ‘Chef (Laksen) Crampton’ for the “salmon projects”.
Not everything can be explained by science. It’s been said that, “Only love can
be divided endlessly and still not diminish” (Lindbergh). Varför kan jag “inte ens
stå när du inte ser på?” (Berg). Tack för att du ser på. And, in stereo, “Yours are
the sweetest eyes, I’ve ever seen” (Taupin)… Often “The best effect of fine
persons is felt after we have left their presence” (Emerson) and meeting is a
feeling like no other. “Nothing makes the earth seem so spacious as to have
friends at a distance” (Thoreau). Sweden, Newfoundland, & Canada… the earth
will always be huge to me. Thanks to all my friends!
Now, “Sooner or later we all quote our mothers” (Williams)… and fathers for
that matter. I remember, “When I was 14 I thought my father was the dumbest
human being alive. By the time I reached 24 I couldn't believe how smart he had
become in 10 years” (Twain). Sometimes we take our family for granted but I am
forever indebted to my family for standing behind me through everything. What
can I say?.. Other than, “Thanks, you’re the best!” I’ve been in school for some 25
years (it’s true!) but no amount of reading will ever make me as smart as my
mother’s mother, who left school in Grade 8 to take care of her family. If there’s
one thing I’ve learned from her it’s that “We do not stop playing because we are
old, we grow old because we stop playing” (Hayes). She’s the youngest 91-year
old in the world. Ginger tea, raison buns, and late night yarns at the kitchen table,
where the beach still sings, the tides still change and the gulls still rise in the
landwash. Those memories have both hurt me and helped me uncountable times
during my studies here in Sweden.
Back to the Cosmos! “Ah, but a man's reach should exceed his grasp - or what's
a heaven for?” (Browning). I have a tendency to fly away, hands clasping for
everything in sight. Wanting to try everything at once. Thanks to everyone for
keeping me grounded (or trying to?). Lastly, from one of the best: “Imagination is
more important than knowledge” (Einstein). Words to live by.

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