Effects of photoperiod and light intensity on

growth and activity of juvenile haddock

(Melanogrammus aeglefinus)
Edward A. Trippel
*
, Steven R.E. Neil
Fisheries and Oceans Canada, St. Andrews Biological Station, 531 Brandy Cove Road, St. Andrews,
New Brunswick, Canada E5B 2L9
Received 11 May 2001; received in revised form 28 March 2002; accepted 14 April 2002
Abstract
Enhancement of growth of juvenile haddock (Melanogrammus aeglefinus) was achieved
through photomanipulation. After 24 weeks (August January), hatchery-reared haddock under 24 h
light were 5360% heavier than those under natural photoperiod. In a second 24-week experiment,
haddock were grown under five photoperiod regimes (natural photoperiod, 12, 16, 20 and 24 h
light) with two light intensities (30 and 100 lx) at 24 h. Continuous light (and 20 h light) resulted in
the greatest growth response, though other seasonally unchanging photoperiods (12 and 16 h light)
also resulted in faster growth than natural photoperiod. Reduced light intensity, from 100 to 30 lx,
at 24 h, led to a further 11% improvement in body mass. The effects of photomanipulation declined
as temperatures decreased in late autumn and winter. Locomotor activity was the greatest under
natural photoperiod (100 lx), less at 24 h (100 lx) and lowest at 24 h (30 lx). Lower swimming
activity under continuous dim light may translate into metabolic savings and increased body mass.
Integrating these findings with research on larval haddock suggests a period in the ontogeny exists
during which bright light should be dimmed to maximize growth and this perhaps coincides with
changes in body morphology and behaviour associated with benthic foraging.
D 2003 Elsevier Science B.V. All rights reserved.
Keywords: Photoperiod; Light intensity; Growth; Activity; Temperature; Juvenile; Haddock
0044-8486/03/$ - see front matter D 2003 Elsevier Science B.V. All rights reserved.
PII: S0044- 8486( 02) 00198- 9
*
Corresponding author. Tel.: +1-506-529-8854; fax: +1-506-529-5862.
E-mail address: trippele@mar.dfo-mpo.gc.ca (E.A. Trippel).
www.elsevier.com/locate/aqua-online
Aquaculture 217 (2003) 633645
1. Introduction
Photoperiod and light intensity manipulation have been successfully used to improve
growth of larval and juvenile stages of a number of fish species. Continuous illumination
has enhanced juvenile growth of Atlantic salmon (Salmo salar) (Saunders et al., 1985);
Atlantic cod (Gadus morhua) (Folkvord and Ottera, 1993), turbot (Scophthalamus
maximus) (Imsland et al., 1995, 1997), barramundi (Lates calcarifer) (Barlow et al.,
1995), and halibut (Hippoglossus hippoglossus) (Simensen et al., 2000), but for several
other species juvenile growth was not influenced by extended daylength (sole (Solea
solea), Fuchs, 1978; black porgy (Mylio macrocephalus), Kiyono and Hirano, 1981;
yellowtail flounder (Pleuronectes ferrugineus), Purchase et al., 2000). Larval haddock
(Melanogrammus aeglefinus) grew faster when reared under continuous bright compared
to dim light (Downing and Litvak, 1999a), though wide variability in feeding success and
growth in relation to light intensity have been reported in larval fishes (Blaxter, 1986;
Huse, 1994) and these variations are believed to be a function of species-specific natural
light niches and stock origin (Job and Bellwood, 2000; Puvanendran and Brown, 2000).
In the marine environment, as haddock change from larval to juvenile phases, they shift
from the pelagic to demersal zone. This occurs at a body length of
f
23 cm when
juveniles are characterized by a subterminal mouth and a natural diet consisting primarily
of benthic invertebrates (Scott and Scott, 1988). The requirement of bright visual feeding
conditions associated with planktivory presumably diminishes with this ontogenetic shift.
Thus, the effect of light intensity on growth of haddock and other demersal marine fishes
may be important and different cultivation strategies may be required for different life
history phases. Improved fish growth in relation to photo regime has been attributed to a
number of factors including higher food conversion efficiency, lower activity and lower
oxygen consumption (Imsland et al., 1995; Appelbaum and Kamler, 2000).
The objectives of the present study were to examine the effects of photoperiod and light
intensity on growth and locomotor activity of juvenile haddock. In the first experiment, we
examined the effects of natural photoperiod and continuous light on juvenile growth under
two feeding regimes. In the second experiment, we broadened our study and examined
juvenile growth in relation to natural photoperiod, 12, 16, 20, and 24 h illumination (in
addition two light intensities were examined at 24 h light). Locomotor activity was
evaluated under natural photoperiod and continuous light.
2. Method
2.1. Experiment 1
Haddock used in the first experiment were fish of mixed parental background raised
from eggs produced by broodstock maintained at the St. Andrews Biological Station, New
Brunswick, Canada. Prior to experimentation, larvae were maintained from May to July,
1998 in 7 m
3
conical tanks containing filtered seawater (salinity
f
30 ppt; 710 jC), fed
wild zooplankton ad libitum, and weaned to a formulated dry food diet (Moore-Clark,
Perla Marine, 400700 Am) and maintained under continuous light (100 lx (
f
1.72 AE/s/
E.A. Trippel, S.R.E. Neil / Aquaculture 217 (2003) 633645 634
m
2
) at waters surface). On July 13, at a weight of 12 g, fish were transferred to 450-l
tanks kept under continuous light (100 lx). On August 12, 360 juveniles of similar size
were anaesthetized in MS-222 (60 mg/l) and wet weight (F0.1 g) and fork length (F0.1
cm) recorded (initial mean weight (SD) 4.38 g (F0.05 g)). Sixty fishes were randomly
distributed into each of six square 450-l (110.45 m deep (water depth 0.30 m)) grey
tanks with two replicates for each of three treatments (i) simulated natural photoperiod
(NP), dry food delivered continuously during daylight phase using automatic belt feeder;
(ii) 24 h light:0 h dark (LD 24:0), food delivery same as (i); and (iii) 24 h light:0 h dark
(LD 24:0), food delivered continuously over 24 h. Fish were fed to satiation at
f
3% body
weight per day (Corey Feed, Haddock Grower, pellet size 1 mm, 3 mm). In all treatments,
fish foraged on pellets as they descended to the tank bottom as well as off the bottom. Fork
length and body weight were recorded once every 4 weeks until the end of the 24-week
experiment on January 27, 1999. Fultons condition factor (body weight/fork length
3
) x
100 was calculated once every 4 weeks. Specific growth rate (SGR) over 4-week periods
was calculated based on means of treatment weights as:
SGR 100 lnW
2
lnW
1
=t
2
t
1

where W
2
and W
1
are body weights (g) at days t
2
and t
1
, respectively.
Lights used were two fluorescent 60 cm tubes (Sylvania white 20 W bulbs F 20T12)
per tank placed 1 m above the waters surface. Simulated natural photoperiod was adjusted
weekly using timers and changed from 14 h light:10 h dark at the beginning of the
experiment to 10 h light:14 h dark at its cessation with the shortest daylength in December
of
f
8 h 15 min (Fig. 1). Tanks were checked daily for mortalities and any dead fish
removed. Water temperature was recorded throughout the experiment and changed
seasonally (Fig. 1).
Fig. 1. Seasonal changes in water temperature and natural photoperiod at the St. Andrews Biological Station
during the experimental periods of 1998 and 1999.
E.A. Trippel, S.R.E. Neil / Aquaculture 217 (2003) 633645 635
2.2. Experiment 2
Haddock used in the second experiment were fish of mixed parental background raised
from eggs produced by broodstock maintained at the Aquarium and Marine Center,
Shippagan, New Brunswick. Prior to experimentation, larvae were maintained on a diet of
n3 HUFA enriched rotifers and Artemia and weaned to a dry food diet (Biokiowa, 400
700 Am). Fish were maintained from May to July, 1999 in 1.2 m
3
circular tanks containing
filtered seawater (salinity 30 ppt, 1012 jC) under continuous light (100 lx). On July 8,
they were transferred by truck (6 h trip) in insulated tanks containing oxygenated water
(
f
100% saturation) to the St. Andrews Biological Station where they were placed in a 5-
m
3
tank under continuous light (100 lx) and maintained on a formulated dry food diet. On
August 4, 600 juveniles of similar size were anaesthetized in MS-222 (60 mg/l) and wet
weight (F0.1 g) and fork length (F0.1 cm) recorded (initial mean weight (SD) 5.38 g
(F0.06 g)). Sixty fish were randomly distributed in each of 12 square 450-l tanks with two
replicates for each of six treatments (i) simulated natural photoperiod (NP), (ii) 12 h
light:12 h dark (LD 12:12), (iii) 16 h light:8 h dark (LD 16:8), (iv) 20 h light:4 h dark (LD
20:4), (v) 24 h light:0 h dark (LD 24:0), (i v at 100 lx,
f
1.72 AE/s/m
2
), and (vi) 24 h
light:0 dark (LD 24:0) at 30 lx,
f
0.20 AE/s/m
2
. Lower light intensity was achieved by
covering the two fluorescent 60-cm tubes with shade cloth. Fish were fed to satiation at
f
3% body weight per day (Corey Feed, Haddock Grower, pellet size 1 mm, 3 mm)
partitioned over three meals with food administered using automatic belt feeders (0800
0900 h 45% of ration, 12001300 h 20%, and 16001700 h 35%). Initial fish size (46 g)
corresponded with the size at which juvenile haddock are customarily transferred from
hatcheries to marine cage sites. Fork length and body weight were recorded once every 4
weeks until the end of the 24-week experiment on January 19, 2000.
Locomotor activity was estimated
f
1 week after final body measurements and were
recorded by counting the number of fish that passed through a fixed station in each of six
tanks representing three treatments. Fixed stations were erected by vertically fixing two
rods 30 cm apart to the bottom of a tank such that the space between them was centrally
located in the tank. The rods were placed into tanks 3 days before counts were conducted.
The number of fish that passed through the 3030 cm space between the rods (water
depth in tanks was 30 cm) was recorded over a 1-min period. From January 2428, 1 min
activity measurements were recorded five times per day: 0830, 0930, 1030, 1530, and
2000 h for three treatments; natural photoperiod (100 lx), 24 h (100 lx), and 24 h (30 lx). A
red light source was used to conduct night (2000 h) observations under natural photo-
period (40 W incandescent bulb CSA LR94262 located 1 m above waters surface, light
intensity 15 lx (
f
0.13 AE/s/m) at waters surface). Fish showed no signs of being startled
when the red light was turned on
f
1 h before movements were recorded as the vision of
marine teleosts occurring in blue-green coastal waters is commonly not red-sensitive
(Nicol, 1989).
Data were tested for normality (KolmogorovSmirnov test) and homogeneity of
variance (Levene statistic) (SPSS version 10.0). Single classification analysis of variance
was used to determine if significant differences ( P<0.05) in fork length, body weight,
condition factor and locomotor activity occurred between treatments. For attributes where
significant differences occurred, the Duncans multiple range test (a=0.05) was used to
E.A. Trippel, S.R.E. Neil / Aquaculture 217 (2003) 633645 636
determine whether treatments differed from one another. Tank effects were examined for
each variable on each sampling date using a series of Bonferroni corrected t-tests (Trippel
and Hubert, 1990). The tanks effects were minor (in 1998, 95% of the replicate tanks
showed no significant difference ( P<0.05), and in 1999, 88% showed no difference).
When differences did occur, they were small (
f
10%). For this reason, we pooled data of
replicate tanks to conduct subsequent statistical analyses (e.g., ANOVA).
3. Results
3.1. Experiment 1
Haddock grew faster under continuous light than under simulated natural photoperiod.
After 4 weeks, significantly greater body weights occurred under the two 24-h treatments
compared to natural photoperiod (significant differences in fork length and condition
Table 1
Fork length, body weight, and condition factor of haddock maintained for 24 weeks at three light/feeding regimes
(NP=natural photoperiod)
Photoperiod
Week Date NP (100 lx) NP Feed 24 h (100 lx) NP Feed 24 h (100 lx) 24 h Feed
Fork length (cm)
0 12/08/98 7.25
a
F0.09 7.29
a
F0.09 7.31
a
F0.08
4 09/09/98 10.52
b
F0.12 10.96
a
F0.13 10.98
a
F0.12
8 07/10/98 12.78
b
F0.16 13.89
a
F0.14 13.89
a
F0.13
12 05/11/98 14.40
b
F0.18 16.10
a
F0.14 16.09
a
F0.13
16 02/12/98 15.56
b
F0.20 17.75
a
F0.16 17.66
a
F0.14
20 30/12/98 16.34
b
F0.21 18.82
a
F0.16 18.63
a
F0.14
24 27/01/99 17.01
b
F0.21 19.33
a
F0.17 19.19
a
F0.14
Body weight (g)
0 12/08/98 4.32
a
F0.17 4.38
a
F0.16 4.43
a
F0.14
4 09/09/98 13.71
b
F0.47 15.78
a
F0.52 15.79
a
F0.49
8 07/10/98 25.13
b
F0.95 33.50
a
F0.99 33.48
a
F0.91
12 05/11/98 35.62
b
F1.43 52.42
a
F1.43 51.76
a
F1.32
16 02/12/98 45.77
b
F1.89 73.02
a
F1.94 71.05
a
F1.73
20 30/12/98 53.79
b
F2.19 88.63
a
F2.30 86.06
a
F2.09
24 27/01/99 57.34
b
F2.26 91.47
a
F2.38 87.72
a
F2.19
Condition factor
0 12/08/98 1.06
a
F0.01 1.06
a
F0.01 1.09
a
F0.01
4 09/09/98 1.12
a
F0.01 1.14
a
F0.01 1.14
a
F0.01
8 07/10/98 1.14
b
F0.01 1.20
a
F0.01 1.21
a
F0.01
12 05/11/98 1.12
b
F0.01 1.22
a
F0.01 1.21
a
F0.01
16 02/12/98 1.14
b
F0.01 1.27
a
F0.01 1.26
a
F0.01
20 30/12/98 1.16
b
F0.01 1.29
a
F0.01 1.30
a
F0.01
24 27/01/99 1.10
b
F0.01 1.23
a
F0.01 1.21
a
F0.01
MeanFstandard error values are shown. Means sharing the same letter superscript on given dates are not
significantly different (Duncans multiple range test, a=0.05).
E.A. Trippel, S.R.E. Neil / Aquaculture 217 (2003) 633645 637
factor commenced at 8 weeks) (Table 1). After 24 weeks, haddock under 24 h light
(natural photoperiod (NP) food delivery) and 24 h light (24 h food delivery) were
significantly heavier (60% and 53%, respectively) than those under NP (Table 1). Timing
Fig. 2. Changes in daily specific growth rate (SGR) of haddock exposed to different photo regimes in 1998 and
1999. Unless indicated otherwise, all treatments were conducted at a light intensity of 100 lx. In 1998, two
feeding regimes were used at 24 h, one in which food was administered over 24 h and the other during the
daylight phase of natural photoperiod (NP). In 1999, a single feeding regime comprised of three meals per day
was used for the six treatments.
E.A. Trippel, S.R.E. Neil / Aquaculture 217 (2003) 633645 638
of food delivery did not significantly influence body growth at 24 h light (Table 1).
Specific growth rate (weights) calculated over 28-day periods declined seasonally as fish
became larger and water temperature cooled (Figs. 1 and 2). Weight gain was <7% during
the final 4 weeks of the experiment at
f
2.5 jC. Percent survival over 24 weeks ranged
from 82% to 92% among the three treatments.
3.2. Experiment 2
After 16 weeks, significantly greater body weights occurred at 20 and 24 h light than at
12 and 16 h light and natural photoperiod (Table 2). Body weight was significantly lower
under natural photoperiod than at other light regimes and this difference was noted early (4
weeks) and maintained through the experiment (24 weeks). Low light intensity resulted in
comparatively faster growth. After 24 weeks, significantly greater body weights (11.1%)
occurred at 30 lx (24 h) than at 100 lx (24 h). Specific growth rates reflected the trend of
faster growth in dim light (Fig. 2). The initial higher specific growth rates in 1998
Table 2
Fork length, body weight, and condition factor of haddock maintained for 24 weeks at six light regimes
(NP=natural photoperiod)
Photoperiod
Week Date NP (100 lx) 12 h (100 lx) 16 h (100 lx) 20 h (100 lx) 24 h (100 lx) 24 h (30 lx)
Fork length (cm)
0 04/08/99 7.61
a
F0.07 7.64
a
F0.07 7.67
a
F0.07 7.67
a
F0.07 7.69
a
F0.07 7.54
a
F0.08
4 01/09/99 10.38
a
F0.09 10.37
a
F0.09 10.40
a
F0.09 10.38
a
F0.09 10.40
a
F0.10 10.23
a
F0.11
8 29/09/99 12.90
a
F0.08 13.03
a
F0.08 13.00
a
F0.08 13.09
a
F0.09 13.04
a
F0.09 13.08
a
F0.10
12 27/10/99 14.76
c
F0.09 15.23
ab
F0.09 15.07
b
F0.08 15.24
ab
F0.09 15.04
b
F0.10 15.39
a
F0.11
16 24/11/99 16.19
d
F0.10 16.78
bc
F0.10 16.62
c
F0.09 16.93
b
F0.11 16.76
bc
F0.10 17.26
a
F0.11
20 22/12/99 17.52
d
F0.11 18.13
bc
F0.11 17.95
c
F0.10 18.36
b
F0.12 18.26
bc
F0.11 18.50
a
F0.11
24 19/01/00 18.20
d
F0.13 18.86
c
F0.12 18.76
c
F0.11 19.26
b
F0.12 19.20
b
F0.12 19.79
a
F0.12
Body weight (g)
0 04/08/99 5.54
a
F0.13 5.63
a
F0.14 5.56
a
F0.13 5.60
a
F0.14 5.84
a
F0.15 5.46
a
F0.16
4 01/09/99 13.20
a
F0.33 13.25
a
F0.34 13.43
a
F0.33 12.80
a
F0.31 13.61
a
F0.38 13.02
a
F0.42
8 29/09/99 26.94
b
F0.55 28.37
ab
F0.59 28.16
ab
F0.57 28.12
ab
F0.54 28.62
ab
F0.63 29.33
a
F0.67
12 27/10/99 40.78
c
F0.86 45.70
b
F0.96 44.78
b
F0.94 46.94
ab
F0.88 44.70
b
F0.96 49.46
a
F1.05
16 24/11/99 54.99
d
F1.17 62.99
bc
F1.24 59.81
c
F1.27 65.16
b
F1.19 64.50
b
F1.25 69.56
a
F1.39
20 22/12/99 64.77
d
F1.46 72.26
c
F1.54 71.23
c
F1.54 78.02
b
F1.51 77.29
b
F1.57 85.05
a
F1.52
24 19/01/00 74.56
d
F1.71 82.61
c
F1.86 82.07
c
F1.77 91.11
b
F1.78 92.05
b
F1.81 102.26
a
F2.00
Condition factor
0 04/08/99 1.24
ab
F0.01 1.24
ab
F0.01 1.21
a
F0.01 1.22
a
F0.01 1.26
b
F0.01 1.23
ab
F0.01
4 01/09/99 1.15
b
F0.01 1.16
b
F0.01 1.16
b
F0.01 1.12
a
F0.01 1.17
b
F0.01 1.16
b
F0.01
8 29/09/99 1.23
c
F0.01 1.26
ab
F0.01 1.26
ab
F0.01 1.24
bc
F0.01 1.27
a
F0.01 1.28
a
F0.01
12 27/10/99 1.25
d
F0.01 1.27
cd
F0.01 1.29
bc
F0.01 1.31
ab
F0.01 1.29
bc
F0.01 1.33
a
F0.01
16 24/11/99 1.27
c
F0.01 1.31
b
F0.01 1.28
c
F0.01 1.33
ab
F0.01 1.35
a
F0.01 1.34
ab
F0.01
20 22/12/99 1.18
d
F0.01 1.19
cd
F0.01 1.21
c
F0.01 1.24
b
F0.01 1.25
b
F0.01 1.34
a
F0.01
24 19/01/00 1.21
c
F0.01 1.21
c
F0.01 1.22
c
F0.01 1.26
b
F0.01 1.28
ab
F0.01 1.30
a
F0.01
MeanFstandard error values are shown. Means sharing the same letter superscript on given dates are not
significantly different (Duncans multiple range test, a=0.05).
E.A. Trippel, S.R.E. Neil / Aquaculture 217 (2003) 633645 639
compared to 1999 occurred under warmer water temperatures, i.e., natural photoperiod
SGRs were 4.2% and 3.2%/day at 14.1 and 12.4 jC, respectively (Figs. 1 and 2). Percent
survival over 24 weeks ranged from 94% to 96% among the six treatments.
Haddock locomotor activity was the greatest under natural photoperiod (100 lx), less at
24 h (100 lx), and lowest at 24 h (30 lx) (Fig. 3). Fish under natural photoperiod (100 lx)
passed through the fixed station approximately twice as often as those at 24 h (30 lx).
Under natural photoperiod, activity rates varied through the day with activity being lowest
in the mid-afternoon and evening and highest during the loading of belt feeders in the
morning. Night activity levels under natural photoperiod remained higher than those at 24
h light.
4. Discussion
Extended daylength significantly improved growth of juvenile haddock. Improved
appetite, greater ration and higher food conversion efficiency are factors commonly
reported to be responsible for faster teleost growth under continuous light (Saunders
and Henderson, 1970; Folkvord and Ottera, 1993). Changes in endocrine functioning as a
Fig. 3. Number of haddock (meanFstandard error) that moved through a fixed station under three different photo
regimes during the period January 2428, 2000. Counts were conducted during five periods: Before Feeding: just
prior to loading of belt feeders, Feed Preparation: immediately after belt feeders have been loaded, First Feeding:
coincides with first meal, After Feeding: between second and third meals, and Evening: natural photoperiod (NP)
represents darkness (red light used to conduct counts). Statistically significant differences in activity occurred
between each of the three photo regimes at each time of day evaluated (Duncans multiple range test, a=0.05).
E.A. Trippel, S.R.E. Neil / Aquaculture 217 (2003) 633645 640
result of photo regime likely play a key role in this response (Porter et al., 2001). Whether
extended daylength continues to have a beneficial effect on haddock growth after the first
summer and autumn is uncertain and requires further investigation. Given the cold winter
temperatures in the Bay of Fundy (
f
23 jC) and associated slow growth, any extended
gains in growth would presumably occur during the subsequent spring to autumn period.
Although the application of continuous light (and 20 h light) resulted in the greatest
growth response, it is worth noting that the other seasonally unchanging photoperiods (12
and 16 h light) resulted in faster growth than natural photoperiod; a result similar to that
observed in Atlantic salmon (Saunders et al., 1985). Moreover, it is of interest that two
discrete growth responses were observed (12 and 16 h compared to 20 and 24 h light)
(Table 2). It is unclear why an additional 4 h of light from 16 to 20 h improved growth, but
an extra 4 h had no effect between other treatments (i.e., from 12 to 16 h and 20 to 24 h).
In juvenile turbot (maintained at natural photoperiod, 16 h light:8 h dark, and 24 h
light:0 h dark), continuous light slightly enhanced the growth rate above that of other
regimes, after 3 months of exposure at 10 and 16 jC, but not throughout the 6 month
experiment (Imsland et al., 1995). However, in a recent paper (Imsland et al., 1997), better
long-term growth (18 months) of turbot occurred when exposed to extended daylength
during the first winter. In Sebastes diploproa, the positive effects of constant illumination
on growth were related to a greater scope for growth and due to their lower standard
metabolic rate (Boehlert, 1981).
It is often not possible to determine if the light effect on growth depends on food
consumption or better food utilization. Similar growth of haddock between the two feeding
regimes in Experiment 1 suggests that an extended feeding period does not enhance
growth under equivalent daily rations. Thus, haddock may be capable of reaching satiation
during a feeding period equivalent to natural daylength, as observed for Atlantic salmon in
sea cages (Krakenes et al., 1991). In green sunfish (Lepomis cyanellus) maintained for 6
weeks at four photoperiods (constant 8 h light:16 h dark, 16 h light:8 h dark, increasing 8
16 h light and decreasing 168 h light), Gross et al. (1965) demonstrated greater food
intake occurred under longer daylength. Fish growth and food conversion efficiency also
were closely correlated and were generally highest in the increasing photoperiod. As noted
in Boeuf and LeBails (1999) review of the subject, Gross et al. (1965) were the first to
specify that growth might be influenced by light through a better food conversion
efficiency and not just stimulated food intake.
Reduced light intensity, from 100 to 30 lx, at 24 h light, led to improved juvenile
growth of haddock. Consequently, dim continuous light appears to be the best strategy of
those investigated. This low light level is more characteristic of their natural habitat at
60120 m on offshore banks in the Northwest Atlantic (Beamish, 1966; Jerlov, 1968;
Scott and Scott, 1988; Nicol, 1989; Puvanendran and Brown, 2000). Locomotor activity
indicated that juvenile haddock swim less at low light and this may translate into metabolic
savings and increased body mass. Swimming activity is a bioenergetic component not
frequently related to improved growth efficiency and larger body mass of cultivated fish
(Jorgensen and Jobling, 1991). African catfish (Clarias gariepinus) under reduced light,
swam less and used less oxygen with more energy channeled into growth (Appelbaum and
Kamler, 2000). In another demersal species, Atlantic halibut, Hole and Pittman (1995)
observed the best growth at 1 to 10 lx, compared to 500 lx (12 h light at 11 and 14 jC).
E.A. Trippel, S.R.E. Neil / Aquaculture 217 (2003) 633645 641
Effects of light intensity on Atlantic salmon postsmolt growth in net pens have shown
mixed results, with enhanced summer and autumn growth occurring when light-reducing
black polyethylene netting covers were deployed for 1 year (Huse et al., 1990). In another
experiment, Atlantic salmon postsmolts exposed to high light intensity in sea pens showed
significantly better growth (Oppedal et al., 1997).
Swimming activity among individual haddock was less variable (low standard errors) at
24 h light (30 lx) relative to 24 h light (100 lx) and natural photoperiod (100 lx) (Fig. 3).
Further observations of haddock following Experiment 2 in FebruaryMay, 2000 revealed
that by shifting photoperiod and light intensity, their activity patterns could be altered (E.A.
Trippel, unpublished data). When measurements of activity were made 3 days after shade
cloth was removed from the 24 h light (30 lx) treatment (raising it to 100 lx), the activity rate
had increased and was equivalent to the 24 h light (100 lx) treatment (the reverse was also
achieved). Thus, the behavioural responses to light appear to be quite flexible.
Interestingly, haddock activity was greater during darkness than in light under, either of
the two 24 h light regimes (i.e., during the 2000 h observation period, which was initiated
f
4 h after darkness in natural photoperiod) (Fig. 3). Thus, the dark phase of natural
photoperiod did not have a great impact in reducing swimming activity. The greater
movement during light and dark phases under natural photoperiod compared to 24 h light
presumably resulted in additional oxygen consumption and energy expended, which could
have been diverted to growth. Although not specifically measured, haddock at 24 h light
(30 lx) relative to other treatments appeared to require lower rations (via visual
observations of remaining uneaten food) and this facet of their diet requires further
examination. If true, then food conversion efficiency is better under dim constant
illumination and would help to reduce food costs for commercial culture.
Consequently, haddock photo regime (periodicity and intensity) could have a signifi-
cant effect on the culture of haddock during the 3100 g phase when seasonal water
temperatures support good growth. The seasonal duration that photo regime enhances
growth could be further extended in a land-based facility in which water temperature can
be controlled. Differences in specific growth rates between 1998 and 1999 may in part be
related to differences in seasonal water temperatures between the two years (Figs. 1 and 2).
In Experiment 1, the summer temperature was higher and winter temperature lower than in
Experiment 2. The growth differences between natural photoperiod and 24 h light were
greater in Experiment 1, perhaps initiated by the higher early period temperature. At the
completion of Experiment 2, there was much less of a growth difference between natural
photoperiod and 24 h light (22% compared to 5360% in Experiment 1), but the higher
temperature by
f
2 jC during winter was associated with faster December January
growth compared to Experiment 1. Fish culturists need to recognize that the scope for
improving growth through photo manipulation is apparently associated with ambient water
temperature. Inspection of the specific growth rates in Fig. 2 suggests that the effect of
photoperiod declined as temperatures decreased in late autumn and winter. It would be of
interest to further explore the interaction between photoperiod and temperature under
controlled temperature conditions. Other contributing factors include diet composition and
parental origin. As a preliminary analysis, Iwama and Tautzs (1981) growth coefficient
( G
c
) was calculated for each treatment from each experimental year. This measure is
useful in that it allows the comparison of the growth of fish at different sizes, reared at
E.A. Trippel, S.R.E. Neil / Aquaculture 217 (2003) 633645 642
different temperatures. Growth coefficients for 1998 (all at 100 lx) were: 36.4 (NP, NP
feed), 62.4 (24 h light, NP feed), and 60.0 (24 h light, 24 h feed). Growth coefficients for
1999 (all at 100 lx unless indicated) were: 42.4 (NP), 48.1 (12 h light), 47.2 (16 h light),
53.3 (20 h light), 52.8 (24 h light), and 59.6 (24 h light, 30 lx). These trends reflect the
results presented. Note, however, at natural photoperiod, G
c
was substantially lower in
1998 (36.4) than in 1999 (42.4) suggesting that other factors in addition to temperature and
fish size could have contributed to the growth differences in 1998 and 1999.
To maximize production, the present study on juveniles and the results of Downing and
Litvak (1999a,b) on larval haddock indicate there is a period in the ontogeny during which
bright light which increases forage success of highly visual feeders should be dimmed and
this perhaps coincides with changes in body morphology and behaviour adapted for
benthic feeding. Continuous light also has been reported to reduce the incidence of sexual
maturity in some teleosts (Hansen et al., 1992; Oppedal et al., 1997; Jourdan et al., 2000),
and supports the widespread applicability of photoperiod manipulation to finfish aqua-
culture (Boeuf and LeBail, 1999). Recently, many Atlantic salmon farmers in Norway and
Scotland began using continuous lighting during the autumn or winter (October April) to
improve growth, as growth of fish subjected to natural daylight is lowered during the
autumn and winter, while, conversely, no such growth depression is observed during
winter under a continuous light regime (Forsberg, 1995). In a recent study (Oppedal et al.,
1997), when light intensity was sufficient, abrupt changes in exposure of Atlantic salmon
from natural short daylength to continuous additional illumination (JanuaryJune)
promoted growth without initiating maturation.
Besides application to aquaculture, the findings on light intensity are also relevant to our
understanding of fish behaviour in the natural environment. Aquatic ecosystems, both
freshwater and marine, often undergo changes in clarity which affect light transmission.
Thus, fish in part may alter their depth distribution in relation to preferred light intensities
(Job and Bellwood, 2000). The degree that light intensity and photoperiod affect locomotion
and growth is worthy of study for other fishes from both aquaculture and environmental
perspectives.
Acknowledgements
We thank J. Bowers, P. Harmon, K. Madsen, T. Shepherd, and T. Taylor for laboratory
assistance and appropriate care of fish. C. Lanteigne of the Aquarium and Marine Center
located in Shippagan, NB, kindly provided haddock for experimental use. J. Castell and P.
Harmon provided helpful comments on an earlier draft. Two anonymous reviewers also
assisted in improving the manuscript for which the authors are grateful. Financial support
for the project was provided in part by the Department of Fisheries and Oceans Science
Youth Internship Program and Heritage Aquaculture, Blacks Harbour, NB.
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