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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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