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Aquacultural Engineering 38 (2008) 3642

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Effects of light spectrum on growth and stress response

of rainbow trout Oncorhynchus mykiss reared under
recirculating system conditions
Nafsika Karakatsouli a, Sofronios E. Papoutsoglou a,*,
Georgios Panopoulos a, Eustratios S. Papoutsoglou a,
Stella Chadio b, Dimitris Kalogiannis b
a

Department of Applied Hydrobiology, Faculty of Animal Science and Aquaculture,

Agricultural University of Athens, Iera Odos 75, 118 55 Athens, Greece
b
Department of Anatomy and Physiology of Farm Animals, Faculty of Animal Science and Aquaculture,
Agricultural University of Athens, Iera Odos 75, 118 55 Athens, Greece
Received 4 April 2007; accepted 24 October 2007

Abstract
Fish growth and physiology may be affected by light spectrum, which can be easily manipulated in indoor aquaculture facilities,
and especially recirculating water systems, with little cost. Since data related to light spectrum and widely reared fish are still few,
the present study aimed to investigate the effects of coloured light on growth performance and stress response to confinement of
rainbow trout Oncorhynchus mykiss. Fish (145.3  1.5 g) were reared under white (full spectrum, fluorescent lamps), red (605 nm)
and blue (480 nm) light (lamps covered with appropriate filters) for 111 days under recirculating water systems (150 lx, 12L12D).
At the end of the experimental period and for each light treatment, fish were either subjected for 1 h to confinement stress or
remained undisturbed (control groups). Total length of the fish reared under red light was greater than that of the other regimes,
whereas other growth parameters showed a similar trend but were not significantly different from one another. Carcass proximate
composition was not affected by light spectrum. Fish exposed to confinement showed typical primary (high cortisol) and secondary
(high glucose and haematocrit, liver lipid mobilization, osmotic and ionic disturbances, blood acidosis, etc.) stress responses.
Nevertheless, in fish reared under blue light, stress-induced cortisol increase was lower and liver lipids mobilization was absent
compared with white light (significant interaction). Present results indicate that if stressors are kept to a minimum then red light
could be suggested for the intensive rearing of rainbow trout.
# 2007 Elsevier B.V. All rights reserved.
Keywords: Light spectrum; Oncorhynchus mykiss; Growth; Confinement; Stress

1. Introduction
Intensively reared fish may often be subjected to
several chronic and acute stressors of endogenous (e.g.

* Corresponding author. Tel.: +30 2105294401;

fax: +30 2105294401.
E-mail address: sof@aua.gr (S.E. Papoutsoglou).
0144-8609/$ see front matter # 2007 Elsevier B.V. All rights reserved.
doi:10.1016/j.aquaeng.2007.10.006

social interactions) or exogenous (e.g. unfavourable

water quality, high densities, grading, handling) origin
that may have detrimental effects on fish welfare and
growth especially when imposed together (Huntingford
et al., 2006). Among other advantages, the use of
recirculating water systems in fish farming, through a
major control of fish as well as their environment, can
help to minimize the occurrence of stressful conditions
and is strongly encouraged (Papoutsoglou, 1993, 1996;

N. Karakatsouli et al. / Aquacultural Engineering 38 (2008) 3642

Blancheton, 2000; Funge-Smith and Phillips, 2001).

Nevertheless, since an artificial environment has to be
involved, the knowledge of applying appropriate
rearing conditions for each species is necessary for
the most effective and productive operation of these
systems.
Lighting is one of the rearing conditions that can be
easily manipulated in recirculating water systems and
most of the related research papers are focused on
photoperiod and intensity (Boeuf and Le Bail, 1999).
Despite the fact that fish visual system has been
adequately described for several species and proven to
be sufficiently equipped to respond to different
wavelengths (Neumeyer, 1992; Flamarique and Hawryshyn, 1996; Cheng and Flamarique, 2004), light
spectrum effects on farmed fish have only recently
been investigated. Depending on fish species natural
habitat characteristics and their specific visual abilities,
light spectrum has been reported to affect multiple
physiological aspects such as growth, neuro-hormonal
system, reproduction, behaviour, etc. (e.g. Downing,
2002; Bayarri et al., 2002; Naor et al., 2003; Ruchin,
2004; Karakatsouli et al., 2007).
To our knowledge only a limited number of studies
are related to coloured light effects on fish stress
response either to chronic stressors such as daily
disturbance (Head and Malison, 2000) and high density
(Van der Salm et al., 2004) or to acute stressors such as
confinement (Volpato and Barreto, 2001) and chasing
(Barcellos et al., 2006). Although mechanisms involved
are not yet elucidated, the above-mentioned studies
indicate that light spectrum can differentiate fish stress
response (evaluated by plasma cortisol levels) and may
be a useful means to alleviate adverse effects of stress.
It has been previously reported that rearing of
rainbow trout Oncorhynchus mykiss for 11 weeks under
blue light (480 nm) had negatively affected growth and
led to increased brain neurotransmitters indicating that
blue light was perceived as stressful (Karakatsouli et al.,
2007). In that experiment light intensity was 300 lx and
although red light seemed to improve fish performance
it was not possible to distinguish significant differences
compared to white light. In another experiment, the
same authors demonstrated that O. mykiss reared under
red light of 150 lx grew better than under red light of
300 or 600 lx (unpublished results). Thus, it seems that
the effect of light spectrum could be modified under
specific rearing conditions.
The present study aimed to investigate whether light
of specific spectral composition could favour rainbow
trout growth and alleviate stress response, so that an
appropriate light colour for intensively reared trout

37

under recirculating water system could be definitely

suggested. For this reason, fish were reared under white
(full spectrum), red and blue light at light intensity of
150 lx and then an acute stressor (confinement) was
imposed to evaluate fish stress response.
2. Materials and methods
Rainbow trout O. mykiss purchased from a commercial Greek hatchery and raised under laboratory
conditions for at least 12 months were used. One
hundred and twenty fish of mean initial body weight
(S.E.M.) 145.3  1.5 g and total length 23.6 
0.09 cm were randomly distributed (10 fish per group)
in 12 tanks (glass, length  height  width: 83 cm 
42 cm  49 cm, volume capacity 171 l, rearing density
59 fish m 3). Experimental tanks were part of an indoor
recirculating freshwater system provided with mechanical and biological filters, UV-sterilization, compressed
air supply and cooling water apparatus. Water flow
rate was 1.8 l min 1 and all tanks were thoroughly cleaned once a week. Water physiochemical
properties were monitored daily and water quality
was maintained as follows: temperature, 16.6 
0.03 8C; dissolved oxygen, 9.1  0.01 mg l 1 (93.4 
0.08% saturation); pH, 7.18  0.005; total ammonia
nitrogen, 0.503  0.0139 mg l 1; unionized ammonia
nitrogen, 0.0024  0.00008 mg l 1; nitrite nitrogen,
0.187  0.0084 mg l 1; chloride, 37.3  0.88 mg l 1;
total hardness, 216.8  2.44 mg l 1 CaCO3.
Fish were acclimated to experimental tanks for 5
days under room ambient lighting. After the acclimation period white, red and blue light colour was applied
(four tanks per treatment) and fish remained in these
conditions for 111 days. Light colour was achieved by
covering light source (Cool White fluorescence lamps,
OSRAM DULUX D/E 26W/840 G24Q-3) with
coloured filters (red #024, blue #165; LEE Filters,
Andover, Hampshire, England, UK) while no filter was
used for white light colour (full spectrum). Light
spectrum was specified using a constant slit Kruss
spectroscope equipped with a graduated scale that was
wavelength calibrated. Red filter had peak transmission
at approximately 605 (90% relative transmission) and
blue filter at 480 nm (83% relative transmission). In
order to avoid complications due to room lighting, all
tank sides were covered with opaque covers and light
source was placed, through appropriate opening in top
cover, above each tank, at approximately 10 cm from
water surface. All experimental populations were
subjected to photoperiod 12L12D (with half hour
dawn and dusk simulation) and light intensity, in all

38

N. Karakatsouli et al. / Aquacultural Engineering 38 (2008) 3642

treatments, was adjusted to 150 lx. Light manipulation

was controlled with winDim 4.0e PC software and light
intensity measured by means of digital light meter (RS
180-7133, RS Components Ltd., Corby, Northants,
UK).
Fish were fed by hand a commercial pelleted diet
(moisture, 7.9%; crude protein, 44.0%; crude lipid,
11.0%; ash, 9.5%; nitrogen-free extract, 27.6%) twice
daily from Monday to Saturday, while no food was
given on Sunday. All fish populations were fed at
feeding level 1.5% b.w.t. throughout the experiment and
were individually weighed every 2 weeks.
At the end of the experimental period and following
48 h fasting, half fish populations (two tanks from each
light colour) were subjected for 1 h to confinement
stress by lowering water level at 10 cm height (rearing
density 246 fish m 3). The other half fish populations
remained undisturbed and used as controls. All fish
were anaesthetized, in their tanks, using 2-phenoxyethanol at a dose of 0.018 ml g 1 l 1. After complete
anaesthetization (within 1 min) blood sampling took
place from the ventral aorta. About 0.3 ml of each blood
sample was immediately used for measuring haematocrit (12,000  g for 10 min) and blood electrolytes
(Na+, K+ and Ca2+), pH and pCO2 taking into
consideration fish temperature (Rapidpoint 400 Blood
Gas Analyzer, Bayer, Tarrytown, NY, USA). The
remaining blood was centrifuged (12,000  g for
10 min) for plasma separation. Plasma was used for
the determination of glucose, triacylglycerides, cholesterol (enzymatic colorimetric methods, Elitech diagnostics, Sees, France), osmolality (cryoscopic
osmometer, Gonotec Osmomat 010) and cortisol, which
was measured by radioimmunoassay, using a commercially available kit (Coat-A-Count Cortisol, DPC, Los
Angeles, CA, USA) that has been previously validated
for fish (Ainsworth et al., 1985). In the present study, the
sensitivity of the assay was 0.2 mg dl 1 and intra- and
inter-assay coefficient of variation was 3.2 and 6.5%,
respectively.
All fish were individually weighed and total length
was measured to calculate condition factor
[K = 100  (body weight, g)  (total length, cm) 3].
Liver was removed from each fish and used for the
determination of moisture, total lipids (Folch et al.,
1957), as well as for the calculation of hepatosomatic
index. Spleen was also isolated, weighed and expressed
as percent body weight. Sampled fish of each population
were minced (without viscera) in two groups of five
specimens each, and lyophilized for carcass proximate
composition determination according to Kjeldahl and
Soxhlet methods (AOAC, 1984). Specific growth rate

[SGR = (ln Wfn ln Win)  100  t 1, Wfn mean final

body weight (g), Win mean initial body weight (g), t is
the days of rearing], weight gain (% initial body weight)
and food conversion ratio [FCR = (food consumed,
g)  (weight gain, g) 1] were calculated for the whole
fish population in each tank.
Data concerning carcass and liver composition,
organosomatic indices and bloodplasma parameters
were analyzed by two-way analysis of variance
(ANOVA) with light colour and stress as factors. For
growth related data (final weight, total length, condition
factor, SGR, weight gain, FCR), no significant effect of
stress or significant interaction were detected and
therefore these data were analyzed by one-way analysis
of variance with light colour as factor. There was no
significant difference (P > 0.05) between duplicated
tanks, so data for each species concerning replicate
treatments were pooled. Where P values were
significant (P < 0.05) multiple comparisons were
carried out using the Duncan test. Wherever necessary,
data were transformed (logarithm or square root) in
order to obtain normal distribution and/or homogeneity
of variance. All values presented in the text and tables
are untransformed means  S.E.M. (Sokal and Rohlf,
1995).
3. Results
Light colour did not have a significant effect on fish
weight, SGR, weight gain or food utilization, although
these parameters presented their best values for fish
under red light (Fig. 1; Table 1). On the other hand, red
light led to significantly higher total length compared
with blue or white light, while condition factor was not

Fig. 1. Mean weight (S.E.M.) during rainbow trout (Oncorhynchus

mykiss) rearing (tank-acclimation period included) under white, red
and blue light.

N. Karakatsouli et al. / Aquacultural Engineering 38 (2008) 3642

39

Table 1
Growth performance of rainbow trout Oncorhynchus mykiss reared under white, red and blue light for 111 days
Light colour

Final weight (g)

Total length (cm)a
Condition factora
SGR (% day 1)b
Weight gain (%)b
FCRb

White

Red

Blue

370.9  9.2
29.9  0.2 a
1.37  0.01
0.82  0.02
155.14  4.58
1.69  0.05

396.2  9.7
30.7  0.2 b
1.36  0.01
0.87  0.01
172.33  1.42
1.57  0.02

378.9  9.0
29.8  0.2 a
1.42  0.03
0.83  0.03
160.47  9.44
1.66  0.07

NS
*
NS
NS
NS
NS

All values are means (S.E.M.); P: significance level; NS: non significant; *P < 0.05. Means with the same superscripts are not significantly
different.
a
n = 40 fish.
b
n = 4 tanks per colour.

differentiated among experimental light treatments

(Table 1). No differences were observed for carcass
proximate composition apart from a significant interaction for ash content (Table 2). However, differences
were small and probably not biologically important.
Liver water content was increased in confined fish
(Table 2). Also, fish under white light had significantly
higher liver water content than fish under red light,
while fish under blue light presented the lowest levels. A
significant interaction was detected for liver total lipids
(Table 2). For fish under white or red light liver total
lipids were reduced when fish were subjected to
confinement, response that was not observed when fish
were reared under blue light (Table 2). Confinement
also resulted in increased hepatosomatic and decreased
spleenosomatic indices irrespective of light colour
(Table 2).
A significant interaction was also observed for
plasma cortisol levels. Cortisol was increased when fish
were confined but this increase was less in fish under
blue light (Table 2). Confinement resulted in elevated
plasma glucose, osmolality, blood haematocrit, reduced
blood pH and increased blood pCO2, sodium and
potassium irrespective of light colour (Table 2). On the
other hand, white light resulted in lower levels of
plasma glucose than red or blue light (Table 2). Plasma
osmolality levels were higher in fish under red light
followed by those under blue and white light (Table 2).
Also, plasma triacylglycerides and cholesterol levels
were highest in fish under blue light compared with
white light, while fish under red light presented
intermediate values (Table 2). No significant effects
of light colour were detected for the other blood and
plasma parameters, apart from blood calcium levels
(interaction term P < 0.05, Table 2). Unconfined
(control) fish under red light had significantly lower
calcium levels compared with white or blue light. When

fish were subjected to confinement, calcium levels were

increased only in fish under red light (Table 2).
4. Discussion
Present results showed that growth of rainbow trout
was favoured when reared under red light, although
significant differences were detected only for fish total
body length. Similarly, Head and Malison (2000)
reported that yellow perch Perca fluviatilis under red
light grew more in body length than fish under blue or
white light. Also, larval rearing of Wallago attu under
red light resulted in increased survival and total biomass
compared with white light (Giri et al., 2002). Present
growth performance of rainbow trout under red light
confirms what has been previously reported as a trend
(Karakatsouli et al., 2007). In the latter study, a negative
effect of blue light was detected which was not observed
in the present study and this may be related to
differences in experimental design and mostly to the
combination of fish initial size and light intensity.
Karakatsouli et al. (2007) used a light intensity of
300 lx, while in another experiment of the same authors
studying red light effects of 150, 300 and 600 lx on
rainbow trout growth, better growth was observed at
150 lx (unpublished results). Thus, the possibility that
the effect of light wavelength itself may be differentiated under different light intensities cannot be
excluded. It should also be emphasized that these two
light characteristics combined with fish biological stage
have been indicated to interact for larval phototactic
behaviour (Gehrke, 1994), first feeding success (Downing and Litvak, 2001), survival (Downing, 2002) and
reaction distance to prey (Utne-Palm and Bowmaker,
2006).
In the present study, fish subjected to confinement
showed typical primary (high cortisol) and secondary

40
Table 2
Carcass proximate and liver composition, organosomatic indices, blood and plasma parameters of rainbow trout reared under white (W), red (R) and blue (B) light for 111 days and subjected to
confinement
Unconfined fish (U)
White

Two-way ANOVAa

Confined fish (C)

Red

Blue

White

Red

Blue

Stress (S) Light colour (LC)

S  LC

NS
NS
NS
NS

NS
NS
NS
**

72.2  0.2
19.2  0.1
6.3  0.2
2.3  0.1 a

71.5  0.1
19.6  0.3
6.5  0.1
2.6  0.1 b

72.3  0.2
19.3  0.2
6.0  0.2
2.3  0.1 ab

71.9  0.3
19.1  0.1
6.5  0.2
2.2  0.1 a

72.0  0.2
19.3  0.1
6.3  0.3
2.2  0.0 a

Liver parametersc
Water (%)
Total lipids (% wet weight)

79.0  0.2
3.10  0.05 bc

78.0  0.2
3.16  0.05 c

79.8  0.2
2.91  0.05 a

79.1  0.2
2.99  0.05 ab

78.7  0.2
U < C** B < R < W***
3.21  0.05 c U > C*** W = R < B*

0.954  0.031
0.155  0.009

1.146  0.037
0.111  0.006

79.2  0.2
3.09  0.04 bc

Organosomatic indices (% body weight)c

Liver
0.982  0.030
Spleen
0.174  0.015

1.004  0.028
0.148  0.008

Blood and plasma parameters c

Cortisol (ng ml 1)
2.5  0.4 a
Glucose (mg 100 ml 1)
73.12  1.14
Osmolality (Osmol kg 1)
0.313  0.001
Triacylglycerides (mg 100 ml 1) 202.11  8.07
Cholesterol (mg 100 ml 1)
234.09  7.58
Haematocrit (%)
31.1  0.5
pH
7.576  0.019
pCO2 (mmHg)
7.1  0.3
Sodium (mmol l 1)
143.7  1.6
Potassium (mmol l 1)
3.88  0.16
Calcium (mmol l 1)
0.82  0.06 b

2.1  0.3 a
80.13  2.72
0.324  0.001
241.34  14.85
238.63  7.88
30.9  0.7
7.531  0.085
8.2  0.3
139.4  1.8
4.08  0.20
0.60  0.04 a

0.999  0.031
0.100  0.008

3.2  0.3 a
36.8  4.8 c
35.2  4.5 bc
82.30  2.22
92.22  3.47
102.95  3.83
0.318  0.001
0.326  0.002
0.337  0.002
258.13  11.90 239.52  14.92 250.59  13.31
261.60  6.10
241.79  7.24
252.73  7.51
30.7  0.6
37.0  0.8
35.7  0.8
7.592  0.082
7.304  0.145
7.332  0.139
7.7  0.2
12.3  0.6
12.2  0.5
140.5  1.3
145.9  0.9
146.6  2.0
3.75  0.15
4.70  0.34
5.09  0.25
0.83  0.05 b
0.95  0.05 b
0.92  0.05 b

1.049  0.051
0.111  0.008

26.6  3.4 b
100.06  4.32
0.333  0.002
272.09  23.63
255.89  6.33
35.9  0.6
7.326  0.112
12.8  0.4
146.2  1.6
4.96  0.20
0.87  0.04 b

NS
NS
NS
NS

NS
*

U < C** NS
U > C*** NS

NS
NS

U < C***
U < C**
U < C***
NS
NS
U < C***
U > C***
U < C***
U < C***
U < C*
U < C*

*
NS
NS
NS
NS
NS
NS
NS
NS
NS
*

NS
W < R = B***
W < B < R***
W  R  B; W < B**
W  R  B; W < B*
NS
NS
NS
NS
NS
R  B  W; R < W*

a
Significant treatment effects are shown as symbols; < or >, means are significantly lower or higher;  or =, means are not significantly different; significant interaction is shown as letters;
*P < 0.05; **P < 0.01; ***P < 0.001; NS, non-significant.
b
Results represent means  S.E.M.; n = 4 groups of 5 fish each.
c
n = 20 fish.

N. Karakatsouli et al. / Aquacultural Engineering 38 (2008) 3642

Carcass composition (% wet weight)

Water
72.1  0.2
Protein
19.3  0.1
Lipid
6.3  0.2
Ash
2.1  0.1 a

N. Karakatsouli et al. / Aquacultural Engineering 38 (2008) 3642

(high glucose and haematocrit, liver lipid mobilization,

osmotic and ionic disturbances, blood acidosis) stress
responses (Mommsen et al., 1999; Barton, 2002), which
are in accordance to what has been previously reported
for rainbow trout under confinement or other acute
stressors (Pottinger et al., 1992; Kakizawa et al., 1995;
Davidson et al., 2000; Kulczykowska, 2001). Focusing
on our objective, confined fish under blue light showed a
better stress response as it is indicated by the lower
increase in plasma cortisol which may also explain the
absence of liver lipid utilization. Volpato and Barreto
(2001) subjected Nile tilapia Oreochromis niloticus for
48 h to confinement under blue, green or white light and
reported lower stress-induced cortisol increase in fish
under blue light. Nevertheless, their results are
considered with certain reservations since they used
different light intensity for white and coloured lights.
Contrary to present results, jundia Rhambia quelen
juveniles subjected to chasing for 60 s after rearing for
10 days under 480 nm (which corresponds to our blue
filter) presented the highest cortisol increase compared
with light of 580 and 436 nm (Barcellos et al., 2006). On
the other hand, no effect of lighting spectrum on plasma
cortisol was detected for yellow perch P. fluviatilis
subjected to daily routine disturbances for 87 days
(Head and Malison, 2000) and for red porgy Pagrus
pagrus subjected to crowding for 30 days (Van der Salm
et al., 2004).
Although involved mechanisms remain still to be
elucidated, light spectrum effects have been generally
related to fish natural habitat and specific visual
abilities. Clean fresh water is a medium that permits
the penetration of longer wavelengths and visual
pigments of rainbow trout are sensitive to long
wavelength (Hawryshyn and Harosi, 1994). Lighting
conditions under red light may be perceived by rainbow
trout as more natural and thus favouring growth. On the
other hand present light spectrum effects on growth as
well as on stress response are probably related to neurohormonal mechanisms and the interaction among
several hormones affected by both acute stress and
lighting conditions cannot be excluded. For example,
plasma or pineal melatonin has been reported to be
influenced by light spectrum (Bayarri et al., 2002; Naor
et al., 2003; Ziv et al., 2007) but also by acute stress in
rainbow trout (Kulczykowska, 2001). Furthermore,
thyroid hormones, somatolactin, growth hormone and
a-melanocyte-stimulating hormone have been suggested to be affected by lighting conditions (Boeuf
and Le Bail, 1999; Szisch et al., 2002; Van der Salm
et al., 2004) and by acute stressors (Kakizawa et al.,
1995; Reddy et al., 1995; Ruane et al., 1999).

41

Present light spectrum treatment had also some other

physiological effects irrespective of confinement.
Higher liver water content observed in fish under white
light is in contrast to previously reported results for
rainbow trout (Karakatsouli et al., 2007). Differences in
experimental design may explain this discrepancy.
Nevertheless, since osmolality was lower in fish under
white light while no differences were detected for
plasma electrolytes, a probable effect of light spectrum
on osmotic regulation is indicated. Plasma glucose
results are also in contrast with previously reported for
rainbow trout (Karakatsouli et al., 2007) or red porgy P.
pagrus (Van der Salm et al., 2004). However, higher
glucose levels in fish under blue or red light seem not to
be indicative of a stress effect of light spectrum, since
cortisol levels in unconfined fish were similar and low
among light colours. On the other hand, glucose levels
especially in fish under blue light, along with the
observed higher levels of plasma cholesterol and
triacylglycerides could reflect a better nutritional status
(Congleton and Wagner, 2006) and may be related to the
better stress response these fish exhibited when
confined.
In conclusion, present results indicated that red light
favoured growth and blue light favoured fish acute
stress response. Providing that stressors are kept to a
minimum then red light (605 nm, 150 lx, 12L12D)
could be suggested for the intensive rearing of rainbow
trout. Light spectrum effects on fish physiological
condition require further research especially in the field
of neuro-hormonal mechanisms involved. However, it is
clear that light spectrum should not be neglected when
artificial light in indoor fish farming facilities and
recirculating water systems is concerned.
Acknowledgments
We are most grateful to G. Konstantinou, X. Vrettos,
P. Louizos, H. Louizos, J. Garofalakis for their
laboratory and technical assistance and to Mr. P. Dimou
for providing the fish.
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