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Aquaculture Research, 1997, 28, 43–49
Effects of decreases in photoperiod on growth and
bimodality in Atlantic salmon Salmo salar L.
O T Skilbrei
Department of Aquaculture, Institute of Marine Research, Bergen-Nordnes, Norway
T Hansen
Matre Aquaculture Research Station, Institute of Marine Research, Matredal, Norway
S O Stefansson
Department of Fisheries and Marine Biology, University of Bergen, High Technology Centre, Bergen, Norway
Correspondence: O T Skilbrei, Department of Aquaculture, Institute of Marine Research, POB 1870, N-5024 Bergen-Nordnes, Norway
Abstract
Two experiments were performed to study the
relative significance of the absolute daylength and
the change in photoperiod on the growth and
development of bimodality in juvenile Atlantic
salmon Salmo salar L. In Experiment A juveniles
were reared on 24 h daily light until they were
seven months old (65–82 mm in length after size
grading). They were then divided into six groups
and subjected to six photoperiods (6, 9, 12, 15, 18
and 24 h of light). In Experiment B the decrease in
photoperiod was made in two steps. First, the day
length was reduced to 18 and 21 hours, three
months after first feeding when the weight of the
juveniles averaged 2.5 g and one group was kept
under 24 hour daily light. Two months later, each
of these treatment groups was subdivided to produce
new groups of juveniles (65–82 mm in length)
under 6, 9 and 12 h of daily light. Irrespective of
whether the photoperiods were reduced in one or
two steps, groups held under short-day photoperiods,
6–12 h, grew significantly slower (Exp. A) and
showed higher proportions of lower modal group
fish (Exp. B) than groups treated with long-day
photoperiods, greater than 12 h. There were low
proportions of lower modal group fish among
juveniles larger than 75 mm at the dates of decreases
in daylength irrespective of photoperiod (Exp. B, 0–
16%), and high or variable proportions among fish
smaller than 75 mm, depending on photoperiod
(Exp B, 32–71%). It is concluded that the growth
© 1997 Blackwell Science Ltd.
response of juvenile Atlantic salmon changes in the
range of 12–15 hours of daily light. This mechanism
is probably linked to the size of the parr and is one
important reason for the development of bimodal
length-frequency distributions.
Introduction
In the culture of Atlantic salmon Salmo salar L.,
growth and developmental rates until smolting are
influenced by photoperiod (Folmar & Dickhoff 1980;
McCormick, Saunders, Henderson & Harmon 1987;
Hoar 1988; Komourdjian, Fenwick & Saunders
1989). Endogenous rhythms regulate growth rate
and smolting in the absence of natural photoperiod
changes, which act as synchronizers of these
processes (Eriksson & Lundqvist 1982). For the fish
farming industry, variation in percentages of one-
year smolts (Knutsson & Grav 1976; Skilbrei 1991)
and in the timing of smolting (Saunders & Henderson
1970; Duston & Saunders 1990; Stefansson,
Björnsson, Hansen, Haux, Taranger & Saunders
1991) are of practical interest. Although growth
rates are improved by rearing the fish under
artificially extended daylengths (Saunders,
Henderson & Harmon 1987; Stefansson, Nævdal &
Hansen 1989), a decrease in photoperiod is
necessary to initiate the smolting process which is
completed under increasing photoperiod (Björnsson,
Thorarensen, Hirano, Ogasawara & Kristinsson
1989).
43
Effects of decreases in photoperiod O T Skilbrei et al.
The reduction in photoperiod may also induce
growth changes which are undesirable in a culture
situation. There is evidence that the development of
bimodal length-frequency distributions, the result
of processes which separate smolting from non-
smolting fish in the wild (Nicieza, Braña & Toledo
1991) and in culture (Simpson & Thorpe 1976;
Thorpe 1977; Thorpe, Morgan, Ottaway & Miles
1980), is influenced by photoperiod (Villarreal,
Thorpe & Miles 1988; Stefansson, Nævdal & Hansen
1990). Elson (1957) proposed that juveniles had to
reach a critical threshold length during the autumn
to smolt the following spring. There are observations
that this mechanism may operate through a
relationship between threshold length and decrease
in photoperiod (Thorpe et al. 1980; Skilbrei 1991;
Saunders, Duston & Benfey, 1994).
Groups of Norwegian Atlantic salmon parr of
70–80 mm separated into lower and upper mode
fish after being transferred from continuous to
natural light in December and January (Skilbrei
1991). At that time the natural daylength was
approximately 7 h (latitude 60°159) and
photoperiods other than continuous and natural
light were not examined. Even though there may
be a relationship between a critical length of about
75 mm, photoperiod and bimodality the relationship
between the magnitude of decrease in photoperiod
and the absolute daylength after decrease is not
known. This study reports the findings from two
experiments which were conducted to investigate
the effects of reducing the photoperiod during
autumn on the growth rates and length-frequency
distributions. The aim was to examine the effects of
specific photoperiods and also the importance of the
size of the reduction in photoperiod. To faciliate
comparison between these two aspects and with the
results from the referred study (Skilbrei 1991), the
experimental designs were standardized so as to
include juveniles of a narrow length range (65–82
mm) at the time of exposure to reduced photoperiods.
Materials and methods
Experiment A
Juvenile Atlantic salmon of mixed genetic origin
were obtained from the commercial hatchery Salar
A/S, Ålvik, western Norway. The fish were first
fed in June 1990 and then reared under elevated
temperatures (. 10°C) and continuous light in 3 m
diameter fibreglass tanks. The fish were size graded
44
Aquaculture Research, 1997, 28, 43–49
regularly. Light intensities were 120 lux immediately
below the surface and 70 lux at the bottom. In early
February 1991, fish of 65–82 mm (fork length)
from one tank were transferred to the Institute of
Marine Research and divided into six groups. Each
group contained 110 juveniles individually marked
by a combination of adipose fin clips and blue spots
(Alcian blue) in one or two of nine ventral positions
relative to the fins. From February 18 the six groups
were reared separately in 200 l covered rectangular
fibreglass tanks and exposed to 6, 9, 12, 15, 18 or
24 h of daily light (P6, P9, P12, P15, P18 and
P24). Light was provided by 15 W fluorescent tubes
giving 2.1–3.4 W m–2 10 cm below the water
surface and 1.5–2.2 W m–2 at the bottom (LI-COR
LI1000 Underwater Sensor). The quality of the
freshwater, treated with NaOH to increase pH to
about 6.5 before it was supplied to the laboratories,
was further improved by adding seawater to increase
salinity to approximately 0.5‰. Mean temperatures
were 6.1, 6.5 and 7.2°C during the first (4 April),
second (11 May) and third period (14 June) of
measurement, respectively, and between 9 and 13°C
for the rest of the experiment.
From the start of the experiment in early February,
all fish were measured (fork length to the nearest
mm) and weighed (total weight to nearest 0.1 gram)
five times until early September. On September 1–2
problems with the NaOH-treatment equipment at
the Institute resulted in the death of 35% of the fish
and the experiment was terminated.
Experiment B
The experiment was performed at Matre
Aquaculture Research Station, Matredal, Western
Norway. Approximately 3000 fish of one family
group were reared together under continuous light
from first feeding in mid-April. On July 2 1990,
when the fish had a mean weight of 2.5 g, they
were separated randomly into three 1 m2 fibreglass
tanks and reared under 24, 21 or 18 h of daily light
(P24, P21 and P18) at 260–300 lux light intensities.
Temperature was maintained at about 12°C from
first feeding through to late June. The maximum
rearing temperature was about 15°C in early August
1990 and decreased gradually to a minimum of 6°C
in winter. On September 12 1990, fish smaller than
65 and larger than 82 mm were removed from the
tanks. The remaining fish (n 5 1060) were fin-
clipped according to photoperiod and fork length
© 1997 Blackwell Science Ltd, Aquaculture Research, 28, 43–49
Aquaculture Research, 1997, 28, 43–49
(below and above 75 mm) by using combinations
of clipping of adipose and pelvic fins (6 possible
alternatives). A quarter of the fish were maintained
under photoperiods of 24, 21 or 18 h daily light
while the rest were mixed in three additional 1 m2
fibreglass tanks and exposed to 12, 9 or 6 h of light
daily (P24–12, P24–9, P24–6, P21–12, P21–9,
P21–6, P18–12, P18–9 and P18–6). Fork lengths
were measured on January 9 1991. The level of
mortality during the period of the experiment was
1.9% (20 fish).
In both experiments a commercial dry diet
(Skretting) was dispensed according to temperature
and fish size (Austreng, Storebakken & Åsgård 1987)
in both experiments using automatic feeders which
operated during the last 6 h of each experimental
photoperiod at 7 min intervals. During
measurements the fish were anaesthetized using
benzocaine.
Results
Experiment A
Photoperiod affected the mean growth rate of the
groups differently, resulting in two clusters of growth
curves depending on whether the groups had been
exposed to ‘short’; (P6–12) or ‘long’ (P15–24)
photoperiods (Fig. 1). The mean lengths of the three
groups within each of these two categories in early
September were not statistically different (t-tests;
P . 0.05, Fig. 1), but the mean lengths between
the two categories were significantly different (P ,
0.001). There was no clear bimodality in P15–24,
but the length-frequencies were skewed to the left,
partly because of a small number of fish that were
less than 120 mm in length (Fig. 2).
The individual lengths in September were
positively related to initial lengths in February,
especially for the three groups held under short-day
photoperiod. Additionally, the difference in length
increment between P6–12 and P15–24 decreased
with increased initial length (Fig. 3).
Experiment B
In January 1991, bimodal length-frequency groups
had been established (Fig. 4). The proportion of
lower modal group fish (, 100 mm) was higher
under 6–12 h d–1 light compared with 18–24 h
d–1, especially for fish initially smaller than 75 mm
© 1997 Blackwell Science Ltd, Aquaculture Research, 28, 43–49
Effects of decreases in photoperiod O T Skilbrei et al.
(Table 1). When the photoperiod was reduced by 6
or more hours, i.e. to 12 h daily light or lower,
there was no correlation between the reduction in
photoperiod and the proportions of lower modal
group fish among the fin-clipped small (linear
regression on arcsin transformed data of Table 1
where photoperiod reduction is 6 h or more, slope
P 5 0.52; intercept P 5 0.93) or large fish (slope
P 5 1.0; intercept P 5 0.06).
Discussion
The present results show that a decrease in
photoperiod between two constant levels strongly
influences the subsequent growth and development
of five to seven month old juvenile Atlantic salmon.
Further, the effect of the decrease is dependent on
the size of the juveniles at the date of decrease and
on the magnitude of the reduction in photoperiod.
The separation into slowly and rapidly growing
parr during autumn is usually observed through
the establishment of bimodal length-frequency
distributions. The lack of bimodality in Experiment
A may have several explanations; the experiment
may have been terminated too early or the
bimodality of the mixed genetic material may have
been hidden because of overlap of modal groups of
different mean sizes as both proportions and size of
the modal groups are partly under genetic control
(Bailey, Saunders & Buzeta 1980). The feeding
regime (6 h per day) used in this study may also
have influenced the growth performance of the
treatment groups, but the clear differences between
the short- and the long-day photoperiods and the
similarities between groups held under 6 to 12 h of
daily light in both experiments do not indicate that
feeding regime was a major factor which could
explain differences in growth.
From the growth differences in Experiment A (P6–
12 vs P15–24) it was not possible to conclude
whether the change (ù 12 h reduction) or the
magnitude of the new photoperiod (ø 12 h
daylength) was the most important growth
regulating factor. The corresponding results between
the 6–12 h d–1 treatments in Experiment B,
irrespective of the previous photoperiod, strongly
indicate that daylengths of magnitude 12 h or less
have a pronounced effect on growth and the
development of bimodality. The present results,
which suggest that the new absolute daylength is
more important as a winter signal in juvenile
45
Effects of decreases in photoperiod O T Skilbrei et al.
Atlantic salmon than the difference between the
two photoperiods, differ from the results of Bromage,
Randall, Davies, Thrush, Duston, Carillo & Zanuy
(1991) on maturation and timing of spawning of
rainbow trout and smoltification. They concluded
that these developmental rates are not directly
controlled by photoperiod, but rather by changing
daylength. Differences in photoperiodic regulation
46
Aquaculture Research, 1997, 28, 43–49
Figure 1 Growth curves for each
treatment group in Experiment A.
95 % confidence intervals of mean
length are included from June.
Figure 2 Length-frequency
distributions (n) in Experiment A on
September 1–5.
of developmental rates may possibly vary among
salmonid species.
The response to photoperiod also seems to be
related to the size of the parr. The increase in length
increment with initial length in P6–12 in Experiment
A and the clearly higher proportions of upper mode
fish among the initially largest fin-clipped group
(75–80 mm) in Experiment B agree well with the
© 1997 Blackwell Science Ltd, Aquaculture Research, 28, 43–49
Aquaculture Research, 1997, 28, 43–49 Effects of decreases in photoperiod O T Skilbrei et al.

Table 1 Percentages of lower modal group fish (LM) among

initially small (65–74 mm) and large fin-clipped fish
(75–82 mm) in Experiment B. The reduction in the number
of hours of light for each photoperiod in September is
also given

Treatment Photoperiod LM (%) LM (%)

group reduction Small Large

P24–24 0 37.5 0.0

P21–21 0 39.5 2.4
P18–18 0 31.6 2.9
P18–12 6 53.9 4.1
P21–12 9 47.6 4.4
Figure 3 Individual length on February 12–17 versus P18–9 9 68.5 15.7
length increment until September 1–5 in Experiment A. P24–12 12 54.8 11.4
Initial lengths of fish in each treatment group (See Fig. 1 for P21–9 12 56.8 13.3
symbols) are grouped in 4 mm length intervals. Common P18–6 12 58.9 8.9
regression lines are calculated for P6–12 (R2 5 0.77, P24–9 15 71.0 4.1
P , 0.001) and P15–24 (R2 5 0.60, P , 0.01). P21–6 15 60.5 8.7
P24–6 18 55.2 6.7

Figure 4 Length-frequency distri-

butions (%) in Experiment B on
January 9 under the 6 final
photoperiods ranging from 6 to 24 h
of daily light. The bars for each of
the 6 groups summarize to 100%,
and they are shown stacked. See text
for further explanation of the
photoperiods of subgroups during the
first part of Experiment B.

proposal of Skilbrei (1988) and the results of Skilbrei
(1991) that in parr of cultured stocks of Norwegian
salmon the probability of becoming upper mode fish
is increased if a threshold length of approximately
75 mm is attained before photoperiod is reduced
during the autumn. Further support for this
argument is that the dates for the reductions in
photoperiod were arbitrarily chosen and there was
no reason to believe that the length range of
65–82 mm should contain high proportions of both
potential lower and upper mode fish at those specific
dates. Daylengths of less than 12 h do not appear
to cause a further depression in growth.
The establishment of lower modes under long
photoperiods (18–24 h in Experiment B) show that
there are also other causes of bimodality beside
the interactions between length and photoperiod.
Metcalfe (1994) has shown that differences in social
status between individuals develop before autumn
and that this behaviour evidently influences whether
an individual enters the upper mode or not. Such
relationships, in addition to genetic differences, may
have contributed to the differences in proportions of
modal groups under long photoperiods between
Experiments A and B.
Based on the two clusters of growth curves in

© 1997 Blackwell Science Ltd, Aquaculture Research, 28, 43–49 47

Effects of decreases in photoperiod O T Skilbrei et al.
Experiment A, there seems to be a transition zone
or threshold level that causes the juveniles to
distinguish between ‘summer’ and ‘winter’ days,
or between long and short days, in the range
between 12 and 15 h of daily light. To be able
to study the growth response of specific size groups
the reductions in photoperiod were given as single
or double cues. This experimental set-up deviates
from the gradually declining natural photoperiod.
However, the results may still be representative of
the basal growth dynamics of Atlantic salmon.
Bimodal size distributions have been observed in
wild populations (Nicieza et al. 1991) and in
groups planted as fry in a Norwegian stream
(Heggenes & Metcalfe 1991). The relationships
between parr size, photoperiod and growth are
probably ecological mechanisms, preparing the
parr for smolting and ensuring that the individual
has a high probability of reaching a minimum
smolt size the next spring. Accordingly, smolts are
of relatively homogeneous size when leaving the
stream in schools (Solomon 1982; Hansen &
Jonsson 1985). In this way, the opportunity for
smolting the next spring may partly be determined
by the size of the parr in mid-autumn when the
natural photoperiod approaches 12 h of daily light.
The establishment of a relationship between
parr size, photoperiod and smolt age has several
implications for the aquaculture industry. If it is
a goal to reduce the mean smolt age, by maximizing
the proportion of the upper modal group, then
the parr should be kept under long photoperiod
until they grow larger than the threshold length
(approximately 75 mm for Norwegian salmon) to
maintain sustainable growth rates during autumn
and winter. The present findings may also be
applied in out-of-season smolt production. Recent
research (including the present results) have led
to production schemes for zero-age smolts, and
many Norwegian hatcheries now produce zero-
age smolts if they can meet the temperature
requirements. By combining artificial long, short
and then increasing (or long) photoperiod (Berge,
Berg, Fyhn, Barnung, Hansen & Stefansson 1995)
the fish reach smolt stage during their first
autumn, and they are transferred to sea-cages at
least a half year earlier than one-year-smolts
produced under natural light. For practical
purposes it is usually recommended that the parr
reach 10 cm before the photoperiod is reduced to
increase the smolt size.
48
Aquaculture Research, 1997, 28, 43–49
Acknowledgment
The authors thank Svein Aleksandersen, Rolf Sundt
and the staff at Matre Aquaculture Research Station
for practical assistance and care of the fish and John
Thorpe who gave valuable comments about the
manuscript. The authors are also grateful to BP
Salar Ålvik A/S for providing some of the fish needed
for the experiments. The study was supported by
the Department of Aquaculture, Institute of Marine
Research (IMR), Matre Aquaculture Research
Station (IMR) and the Norwegian Fisheries Research
Council (NFFR).
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