Effects of different light sources and illumination methods on

growth and body color of shrimp Litopenaeus vannamei

Kui You
a,b,c
, Hongsheng Yang
a,
*
, Ying Liu
a
, Shilin Liu
a
,
Yi Zhou
a
, Tao Zhang
a
a
Institute of Oceanology, Chinese Academy of Sciences, Qingdao 266071, China
b
Graduate School, Chinese Academy of Sciences, Beijing 100039, China
c
Ocean University of China, Qingdao 266003, China
Received 17 May 2004; received in revised form 10 June 2005; accepted 29 June 2005
Abstract
Shrimps Litopenaeus vannamei with initial body weight of 2.108F0.036 g were sampled for specific growth rates (SGR)
and body color measurements for 50 days under different light sources (incandescent lamp, IL; cool-white fluorescent lamp, FL;
metal halide lamp, MHL; and control without lamp) and different illumination methods (illumination only in day, IOD, and
illumination day and night, IDN). Body color of L. vannamei was measured according to the free astaxanthin concentration
(FAC) of shrimp. The SGR, food intake (FI), feed conversion efficiency (FCE) and FAC of shrimps showed significant
differences among the experimental treatment groups ( Pb0.05). Maximum and minimum SGR occurred under IOD by MHL
and IDN by FL, respectively (difference 56.34%). The FI of shrimp for the control group did not rank lowest among treatments,
confirming that shrimp primarily use scent, not vision, to search for food. FI and FCE of shrimps were both the lowest among
treatment groups under IDN by FL and growth was slow, thus FL is not a preferred light source for shrimp culture. Under IOD
by MHL, shrimps had the highest FCE and the third highest FI among treatment groups ensuring rapid growth. FAC of shrimp
were about 3.31F0.20 mg/kg. When under IOD by MHL and IDN by FL, FAC was significantly higher than the other
treatments ( Pb0.05). To summarize, when illuminated by MHL, L. vannamei had not only vivid body color due to high
astaxanthin concentration but also rapid growth. Therefore, MHL is an appropriate indoor light source for shrimp super-
intensive culture. SGR of shrimp was in significantly negative correlation to FAC of shrimp ( Pb0.05). Thus, when FAC
increased, SGR did not always follow, suggesting that the purpose of astaxanthin accumulation was not for growth promotion
but for protection against intense light.
D 2005 Elsevier B.V. All rights reserved.
Keywords: Litopenaeus vannamei; Light source; Illumination method; Growth; Body color
0044-8486/$ - see front matter D 2005 Elsevier B.V. All rights reserved.
doi:10.1016/j.aquaculture.2005.06.041
* Corresponding author. Tel./fax: +86 532 82898582.
E-mail address: hshyang@ms.qdio.ac.cn (H. Yang).
Aquaculture 252 (2006) 557565
www.elsevier.com/locate/aqua-online
1. Introduction
As light is an important environmental factor for
animals living in water, many studies have been
undertaken on its effect. Significant differences in
behavior, food intake and growth of aquatic animals
appeared under different light conditions (Blaxter,
1968; Gehrke, 1994; Giri et al., 2002). Light also
impacted on ovarian maturation, reproduction and
juvenile growth of some aquatic animals (Kelemec
and Simth, 1980; Wallace, 1988; Hillier, 1984;
Fanjul-Moles and Fuentes-Pardo, 1988; Fanjul-
Moles et al., 1992; Primavera and Caballero,
1992; Wang et al., 2003a,b). Fluorescent lamp
(FL) has been used as the main light source for
trials involving shrimps (Wang et al., 2003a,b) and
there are few studies on the effect of different light
sources on shrimp.
Body color is one of the major factors deter-
mining quality and price of shrimp (Chien and
Jeng, 1992; Boonyaratpalin et al., 2001). Concen-
tration of astaxanthin is the main factor controlling
shrimp body color (Crozier, 1967; Howell and
Matthews, 1991; Menasveta et al., 1993; Pangan-
tihon-Ku hlmann et al., 1998; Ne`gre-Sadargues
et al., 2000; Stepnowski et al., 2004a). In this
study, free astaxanthin concentration (FAC) in
shrimp was chosen as the indicator of shrimp
body color. Astaxanthin is also a powerful antiox-
idant; indeed its antioxidant ability is much stron-
ger than that of h-carotene, vitamin E or C.
Animals are unable to synthesize these pigments,
therefore they must accumulate these pigments
through the diet (Shahidi et al., 1994; Ne`gre-
Sadargues et al., 2000; Velu et al., 2003). Astax-
anthin can also act to enhance the immune system
of aquatic animals, prevent disease, increase survi-
val rate and improve growth (Chien and Jeng,
1992; Coral-Hinostroza and Bjerkeng, 2002; Step-
nowski et al., 2004b; Velu et al., 2003). However,
the mechanism of astaxanthin accumulation is less
studied. It has been observed that the body color
of Penaeus monodon becomes faint when cultured
indoors (Tseng et al., 1998). Therefore there may
be some relationship between astaxanthin accumu-
lation of shrimp and light condition.
In this paper, different types of lighting (incan-
descent lamp, IL; fluorescent lamp, FL; metal
halide lamp, MHL; and control without lamp)
were chosen to investigate the effect of light
conditions on growth and body color of Litope-
naeus vannamei and the influence of light on
astaxanthin accumulation was also investigated.
2. Material and methods
2.1. Source and acclimation of shrimp
The experiment was conducted from July 7 to
August 25, 2003 at the Laboratory of Marine Ecol-
ogy and Environmental Sciences, Institute of Ocea-
nology, Chinese Academy of Sciences, Qingdao,
China. Shrimp L. vannamei used in the experiment
was collected from a coastal shrimp farm in Qing-
dao, Shandong Province, China. Before starting the
experiment, all shrimp selected were healthy and
had been acclimated for 15 days in a 10 m
3
con-
crete tank with specific experimental sea water (see
in the next part) under natural light. During both the
acclimation and experimental period shrimps were
fed with a formulated diet (44.39F0.27% crude
protein, 8.74F0.32% fat, 10.91F0.06% ash, and
9.41F0.07% moisture, bought from Haiyue Feed
Stuffs Company, Qingdao) at satiation level twice
daily (at about 6:00 and 18:00 hours).
2.2. Rearing condition
The shrimp were maintained in glass aquaria
(453525 cm, water volume of 35 l). Each
rearing unit contained six shrimps. Different light-
ing treatments (shaded from each other) were con-
ducted in well-ventilated separate wooden boxes
(15013550 cm). Water exchange was con-
ducted in all treatments simultaneously from an
identical water source filtered by a common water
filter tower filled with fine-grained sand. Aeration
was provided continuously and 90% of water
volume in the aquaria was replaced every other
day in order to maintain water quality. Dissolved
oxygen was maintained above 6.0 mg/l, pH was
approximately 8.0; concentration of ammonia was
under 0.24 mg/l, salinity was between 28 to 32,
and water temperature varied from 23~30 8C
according to air temperature.
K. You et al. / Aquaculture 252 (2006) 557565 558
2.3. Experiment setup
Three types of lamps (incandescent lamp, IL; cool-
white fluorescent lamp, FL; and metal halide lamp,
MHL) were chosen as light sources and the control
groups were placed in darkness. Detailed information
regarding the experiment is shown in Table 1. Lumi-
nance on the bottom of the aquaria was measured by an
underwater illumination photometer (ZDS-10, Shang-
hai Xuelian Instruments). Three replicates were set up
for each treatment. Normally, the lamps were hung 60
80 cmabove the aquaria. The illumination intensity was
controlled by adjusting the number and position of the
lamps in order to maintain as equal luminance as pos-
sible at the bottom of each aquaria.
2.4. Experimental procedure and samples collection
After 12-h food deprivation, L. vannamei from the
acclimating tank were randomly selected, weighed and
put into 24 aquaria, each containing 6 individuals.
Individual shrimp weight varied from 1.909 to 2.250
g (2.108F0.036 g). The shrimps were fed twice daily
(at 6:00 and 18:00 hours). Excess food and feces were
collected separately by siphoning within 3.5 h after
feeding. The collected excess food and feces settled,
the water above was drained carefully and replaced with
clean water. This procedure was repeated 4 times to
remove the salts. The molted shells were also collected.
The collected uneaten food, feces and shells were then
dried at 65 8C, separated, and kept in a desiccator for
weighing. At the end of the 50-day experimental period,
all test shrimps were weighed individually and con-
centration of free astaxanthin was determined.
2.5. Calculation of data and FAC determination
Specific growth rates (SGRw), feed intake
(FIw) and food conversion efficiency (FCEw) in
terms of wet weight of shrimp were calculated as
follows:
SGRw %day
1

100 lnW
2
lnW
1
=T
FIdw %BW day
1

100 C= T W
2
W
1
=2 f g
FCEw % 100 W
2
W
1
=C:
Here, W
2
and W
1
are the final and initial wet
body weight of the shrimp (g), respectively; T, the
time span of the experiment in days (d); and C,
total food consumed by shrimp (g). Sample weight
was determined by electronic balance (JA2003N,
Shanghai).
The method for determining astaxanthin concentra-
tion is based on previously published research (Torris-
sen, 1986; Christiansen et al., 1994, 1995; Negro and
Garrido-Fernandez, 2000; Boonyaratpalin et al., 2001).
Samples for astaxanthin determination were freeze-
dried and crushed. Then about 1.5 g of shrimp powder
was homogenized by acetone. The mixed solution was
settled for 1 h and the supernatant liquid was trans-
ferred to a volumetric flask and refilled with acetone.
This procedure was repeated at least nine times, until
the mixed solution became colorless. The clear super-
natant collected in the flask was made up to a known
volume with acetone. Extraction was then centrifuged
at 3500 rpm for 8 min. Finally, FAC were measured
by HPLC (Type: 2010, Waters) with the external
standards (A9335, 98%, Sigma, Germany).
2.6. Statistical analysis
Data on SGR, FI and FCE were converted by
arcsine transformation. The homogeneity of variances
among data groups was tested and then the possible
differences of data groups were analyzed by one-way
ANOVA. Statistics were done with SPSS 10.0 statis-
tical software. Duncans multiple range tests were
used to test differences among the treatment groups.
Table 1
Experimental setup
Treatments 1 2 3 4 5 6 7 8
Lighting type LPIL LPIL MHL Control HPIL HPIL FL FL
Illumination mode IOD IDN IOD / IOD IDN IOD IDN
Number of lamps 2 2 1 2 2 1 1
Power (W) 15 15 200 200 200 40 40
Intensity (lx) 18 18 2500 450 450 210 210
K. You et al. / Aquaculture 252 (2006) 557565 559
Differences are considered significant at a probability
level of 0.05.
3. Results
3.1. Growth
No significant differences in mean individual
shrimp weight was found between the groups
( PN0.05) from experimental onset to conclusion
(Table 2). However, SGR showed significant differ-
ences between the groups ( Pb0.05).
The order of SGR based on wet shrimp weight
from highest to lowest is as follows: #3N#6N#5N
#7N#4N#2N#1N#8 (Fig. 1). SGR varied from 1.42%
to 2.22%. The SGR of treatment under FL with IDN
was the lowest while the SGR of treatment under
MHL with IOD was the highest ( Pb0.05), the latter
being about 1.5 times that of the former. There was no
significant difference between the other treatment
groups and the control group ( PN0.05).
3.2. Food intake (FI)
FI, based on wet shrimp weight in descending
order, is as follows: #6N#5N#3N#4N#1N#7N#2N#8
(Fig. 2). FI varied from2.76%to 3.72%. When shrimps
were illuminated IOD by FL or LPIL or illuminated
IDN by LPIL, FI of L. vannamei showed no obvious
difference from that of the control ( PN0.05, Fig. 2).
Even when shrimp were illuminated IDN by FL, FI of
L. vannamei was significantly lower than that of the
control groups ( Pb0.05). Therefore, light is not neces-
sary for shrimps to locate food; indeed, shrimps use
odors as the main method of food retrieval. When
shrimps were illuminated by HPIL or MHL, FI was
significantly higher than that of the other groups
( Pb0.05).
3.3. Food conversion efficiency (FCE)
FCE based on wet shrimp weight in descending
order is as follows: #3N#7N#4N#2N#5N#6N#1N#8
(Fig. 3). FCE varied from 11.57% to 14.13%. When
shrimp were illuminated IOD by LPIL and IDN by
HPIL or FL, FCE of L. vannamei was significantly
lower than FCE of other groups ( Pb0.05, Fig. 3).
There were no obvious differences between the other
treatment groups and the control regarding FCE
Table 2
Body wet weight of shrimp and survival rates (means FS.E.)
Treatment
group
Body wet weight (g) Survival rate
(%)
Initial Final
1 2.133F0.109 4.717F0.397 93.33F4.08
2 2.185F0.114 5.485F0.333 90.00F6.67
3 2.008F0.075 6.171F0.231 90.00F6.67
4 2.053F0.098 5.453F0.249 90.00F6.67
5 2.233F0.112 5.897F0.218 93.33F4.08
6 2.088F0.085 5.472F0.483 90.00F6.67
7 1.909F0.098 4.949F0.343 100.00F0.00
8 2.250F0.115 4.522F0.270 93.33F4.08
a
cd
bde
ab
bc
e
bcd
cde
0.00
0.50
1.00
1.50
2.00
2.50
1 5 7 8
Treatment groups
6 4 3 2
S
G
R

(
%

d
a
y
-
1
)
Fig. 1. SGR of L. vannamei in experiment. Means with different superscript letters are significantly different ( Pb0.05) and bars indicate
standard errors of the means.
K. You et al. / Aquaculture 252 (2006) 557565 560
( PN0.05), but FCE of #3 group with MHL was the
highest among the remaining groups.
3.4. Free astaxanthin concentration (FAC)
FAC in the experimental shrimp varied from 2.06
to 5.52 mg/kg (3.31F0.20 mg/kg, meanFS.E.)
(Fig. 4). FAC in wild shrimp was higher than that
of the experimental shrimp. Shrimps illuminated IOD
by MHL or FL and IDN by FL or HPIL had FAC
obviously higher than that of the other treatments
( Pb0.05). There were no significant differences
among the remaining treatments ( PN0.05). When
this FAC was compared with those for salmon
and shrimp recorded in other literature (Torrissen,
1986; Christiansen et al., 1994, 1995; Pangantihon-
a
b
d
cd
b
c
ab
b
0.00
0.50
1.00
1.50
2.00
2.50
3.00
3.50
4.00
1 2 3
F
I

(
%
B

d
a
y
-
1
)
Treatment groups
8 7 6 5 4
Fig. 2. FI of L. vannamei during experimental period. Means with different superscript letters are significantly different ( Pb0.05) and bars
indicate standard errors of the means.
a
bc
a ab
bc
a
ab
c
0
2
4
6
8
10
12
14
16
1 2 3 4 7
F
C
E

(
%
)
8 6 5
Treatment groups
Fig. 3. FCE of L. vannamei during experimental period. Means with different superscript letters are significantly different ( P b0.05) and bars
indicate standard errors of the means.
K. You et al. / Aquaculture 252 (2006) 557565 561
Ku hlmann et al., 1998), it was lower when compared
with total carotenoid concentrations (including free
monoester, diester astaxanthin, h-carotene and other
carotenoids) measured by spectrophotometer and as
published elsewhere (Boonyaratpalin et al., 2001).
3.5. Relationship between SGR and FAC
Relationship between SGR and FAC is shown in
Fig. 5. SGR is in significant negative correlation with
FAC for shrimps in this experiment ( Pb0.05).
4. Discussion
4.1. Illumination by MHL improving shrimp growth
Previous research has been reported on relation-
ships between fish behavior and light (Blaxter,
1968; Gehrke, 1994) and the effect of light on
growth and reproduction of crustaceans and their
vision system has also been reported (Kelemec
and Simth, 1980; Hillier, 1984; Wurts and Stickney,
1984; Fanjul-Moles and Fuentes-Pardo, 1988; Fan-
bc
bc
ab
ab
a
a
b
a a
0.00
1.00
2.00
3.00
4.00
5.00
6.00
7.00
1 2 3 4 7 9
C
o
n
c
e
n
t
r
a
t
i
o
n

o
f

f
r
e
e

a
s
t
a
x
a
n
t
h
i
n

(
m
g
/
k
g
)
Treatment groups
8 6 5
Fig. 4. Concentration of free astaxanthin in L. vannamei under different treatments. Treat group 9 is wild shrimp. Means with different
superscript letters are significantly different ( Pb0.05) and bars indicate standard errors of means.
y = -0. 1183x + 2. 1989
r = 0. 477, F=6.497,P=0.018
(P=95%,n=23, r=0.396)
1.00
1.50
2.00
2.50
1 4 6
Concentration of free astaxanthin (mg/kg)
5 3 2
S
G
R

(
%

d
a
y
-
1
)
Fig. 5. Relationship between SGR and concentration of free astaxanthin.
K. You et al. / Aquaculture 252 (2006) 557565 562
jul-Moles et al., 1992; Primavera and Caballero,
1992). It is also reported that light color and inten-
sity have a significant effect on the growth of
Fenneropenaeus chinensis (Wang et al., 2003a,b).
In this study, the growth promotional ability of
MHL over FL on L. vannamei is related to the
particular characteristics of the lights. IL has a
wide spectrum but IL has low energy transformation
efficiency (An and Kim, 2000) and is rarely used
due to the high energy cost. The spectrum of MHL
has a larger infrared composition helpful to shrimp
growth. Some evidence shows MHL to be helpful to
plant growth. MHL is most efficient at promoting plant
growth among the three lamps (Warrington and Mitch-
ell, 1975). Although there is no evidence showing that
MHL is helpful for animal growth, the results of this
experiment showed MHL was beneficial to shrimp
growth. It is reported that the spectrum of FL contains
ultraviolet (Gro f et al., 2002), which may have a
negative effect on shrimp growth. F. chinensis grew
slowly when illuminated by intense FL light (Wang
et al., 2003a,b). When MHL was the chosen light
source, shrimp had high FI level and FCE thus these
shrimp grew faster than the other groups (Figs. 2 and
3). When FL was used the opposite occurred suggest-
ing that MHL is a better light source than FL for
shrimp culture.
4.2. Intense light illumination increasing body color
of shrimp
Body color of shrimp varies according to photo-
period (Lakshmi et al., 1976). Body color of P.
monodon would become faint when cultured indoor
under low light intensity less than 1000 lx (Tseng
et al., 1998). Astaxanthin concentration is an appro-
priate indicator of body color of shrimp (Crozier,
1967; Menasveta et al., 1993; Ne`gre-Sadargues
et al., 2000; Stepnowski et al., 2004a). Firstly,
FAC in wild shrimp was higher than in lab-cultured
shrimps as natural sunlight intensity (no less than
10,000 lx) was much higher than light intensity in
the labs (max 2500 lx) (Table 1). Secondly, when
shrimp were illuminated IOD by MHL or IDN by
FL, FAC of shrimp was higher than that of other
lighting conditions and the light intensity in these
two treatments was higher than that of other treat-
ments. Therefore, shrimp would have vivid body
color when illuminated by high-intensity light. More-
over, when illuminated by MHL, shrimp had not
only vivid body color but also faster growth rates
showing that MHL is a good light source for shrimp
culture.
4.3. Astaxanthin accumulation as a means of avoiding
damage caused by excess light
As an ideal antioxidant, astaxanthin has a strong
capability to eliminate free radicals and singlet oxy-
gen and is used widely in aquaculture to increase
survival rate, enhance the immune system, improve
growth and prevent diseases (Coral-Hinostroza and
Bjerkeng, 2002; Chien et al., 2003; Pan et al., 2003;
Stepnowski et al., 2004b; Velu et al., 2003). Con-
siderable research has shown that fish or shrimps
grew faster when astaxanthin was added to their diet
compared to control groups where no astaxanthin
was added (Christiansen et al., 1995; Boonyaratpalin
et al., 2001). The results of the present paper, how-
ever, appear to contradict these published results.
The diets used in this experiment contained no
deliberate addition of astaxanthin. Therefore, under
natural conditions shrimp would not always grow
better when FAC levels were high. In other words,
FAC level of shrimp is related to not only astax-
anthin concentration in the diet but also light con-
dition. What then is the purpose of FAC
accumulation? One explanation is that some types
of algae accumulate astaxanthin in their body in
order to avoid damage caused by intense light,
especially ultraviolet light. Strong light intensity or
ultraviolet light increases astaxanthin content in
those algae (Gerber et al., 1994; Barbosa et al.,
1999). Results of this study, where SGR had a
strong negative correlation to FAC, suggest that
this phenomenon may also be applicable to shrimp.
It is widely understood that shrimp are easily
harmed by excess light, especially ultraviolet light,
due to the transparency of their bodies. Therefore,
the principal purpose of astaxanthin accumulation
may be to avoid damage caused by excess light.
High FAC in shrimp may reject the intense light of
the shrimps natural habitat. This result suggests that
light condition should be taken into account when
astaxanthin is added in the diet to improve growth
and body color of shrimp.
K. You et al. / Aquaculture 252 (2006) 557565 563
Acknowledgements
This work was supported financially by CAS
Project (ZKCX2-211) and a project of the Institute
of Oceanology, Chinese Academy of Sciences
(L400223108). We also acknowledge Mr. Zhuang
Baibing and Mr. Zhang Jingyou of the Dalian Fish-
eries University for their help with the experiment
and Dr. Roger Z. Yu and Mrs. Kursty A. Martrnson
for their professional revision of the manuscript.
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