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Department of Food Science, National Taiwan Ocean University, Keelung, Taiwan 202, Republic of China
2To whom correspondence and reprint request should be addressed.
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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30,000 RE vitamin A/kg. Survival rate was higher
in shrimp fed diets supplemented with 1,500–30,000 RE vitamin A/kg
than shrimp fed the control diet. Highest blood triglyceride
concentration and body lipid concentration were in shrimp fed diets
supplemented with 45,000 and 60,000 RE vitamin A/kg, respectively. Eye
vitamin A concentration and hepatopancreatic total lipid concentration
in shrimp generally increased as dietary vitamin A supplementation
increased. In expt. 2, feed efficiency was highest in shrimp fed diets
supplemented with 2,400, 3,000, 3,600 and 4,500 RE vitamin A/kg,
followed by shrimp fed diets with 600 and 1,200 RE vitamin A/kg and
finally the unsupplemented control group. Shrimp fed diets supplemented
with vitamin A had significantly higher survival percentages than those
fed the unsupplemented control diet. Weight gain percentage of the
shrimp analyzed by broken-line regression indicated that the
minimal dietary vitamin A concentration in growing P.
monodon is 2,511 RE/kg (~8,400 IU/kg).
KEY WORDS: • vitamin A • shrimp • Penaeus monodon
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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),
guppy (Shim and Tan 1990
) and common carp (Aoe et al. 1968
, Suhenda and Djajadiredja 1985
),
respectively. These values for teleost species are somewhat different
from those established for homeotherms such as chickens and pigs in
which a requirement of 1,500 and 2,000 IU/kg, respectively,
were reported (NRC 1984
and 1988
).
For crustaceans, only limited qualitative data concerning vitamin
requirements are available. He et al. (1992)
and Chen and Li (1994)
reported that penaeid shrimp, Penaeus vannamei and P.
chinensis, required dietary vitamin A for optimal growth. However,
quantitative information on the requirements of vitamin A for penaeid
shrimp is lacking. The purpose of this study was to define the minimal
dietary vitamin A requirement of juvenile grass shrimp P.
monodon, the most widely cultured shrimp worldwide.
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MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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Experimental diet formulation is given in Table 1
. The formulation is similar to that of Shiau and Hwang (1994)
, which
has been shown to be adequate for P. monodon.
Vitamin-free casein (Sigma Chemical, St. Louis, MO), cod liver oil
(Scott and Bowne, London, United Kingdom) and corn starch (Sigma
Chemical) were used as dietary protein, lipid and carbohydrate sources,
respectively. A mixture of amino acids [including glycine,
L-alanine, L-glutamate and betaine (Sigma
Chemical)] was included in the diets as an attractant (Shiau and Chin 1998
). Sodium alginate (Hayashi Pure Chemical, Osaka,
Japan) was used as a binder. This constituent is not digested by the
shrimp, and it is essential for shrimp diet as it holds the ingredients
in a stable pellet form in water. The vitamin mixture was similar to
that used by Shiau and Hsu (1999)
, except that it did not contain
vitamin A. In expt. 1, vitamin A (retinyl acetate) (Sigma Chemical) was
added to the test diets at the expense of small amounts of cellulose to
provide concentrations of 0, 1,500, 3,000, 15,000, 30,000, 45,000 and
60,000 retinol equivalent (RE)/kg diet (0, 5,000, 10,000, 50,000,
100,000, 150,000, 200,000 IU/kg). In expt. 2, retinyl
acetate was added at 0, 600, 1,200, 1,800, 2,400, 3,000, 3,600 and
4,500 RE/kg diet (0, 2,000, 4,000, 6,000, 8,000, 10,000, 12,000, 15,000
IU/kg). The vitamin A concentrations of the experimental
diets were determined by HPLC (Larry et al. 1991
) to be
54 (unsupplemented control), 1,380 (1,500 RE/kg), 2,856 (3,000 RE/kg),
13,466 (15,000 RE/kg), 28,246 (30,000 RE/kg), 39,653 (45,000 RE/kg) and
55,978 (60,000 RE/kg) in expt. 1 and 54 (unsupplemented), 572 (600
RE/kg), 1,166 (1,200 RE/kg), 1,747 (1,800 RE/kg), 2,338 (2,400 RE/kg),
2,909 (3,000 RE/kg), 3,521 (3,600 RE/kg) and 4,346 (4,500 RE/kg) in
expt. 2. The diets were prepared and stored as previously described
(Shiau and Yu 1998
).
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Juvenile P. monodon were supplied by the Tungkang Marine Laboratory (Tungkang, Pingtung, Taiwan). Upon arrival, they were acclimated to laboratory conditions for 2 wk in a plastic tank [74 cm (w) x 95 cm (l) x 45 cm (h), 290 L] and fed a commercial diet (Yung-Hsien, Taipei, Taiwan). The laboratory conditions during the acclimation period were similar to those at the initiation of the experiment. The proximate composition (g/100 g) of the commercial diet was as follows: moisture, 10.74; crude protein (N x 6.25), 40.90; lipid, 7.19; and ash, 12.46. Shrimp were then fed the basal diet without vitamin A supplementation for 7 d prior to the start of the growth trial. At the beginning of the experiment, 16 shrimp [mean weight: 0.97 ± 0.01 g (SD)] in expt. 1 and 20 shrimp [mean weight: 0.68 ± 0.01 g (SD)] in expt. 2 were stocked in each experimental aquarium (60 x 60 x 45 cm3, 125 L). Each experimental diet was fed to three groups of shrimp. Each aquarium received continuous aeration. In each aquarium, impurities from uneaten feed and fecal pellets in the water were removed by siphoning every day, and 75% of the water was exchanged at 2, 4 and 6 wk in expt. 1 and 2, and 4 wk in expt. 2 to maintain water quality. Dissolved oxygen concentration was monitored weekly and maintained at 7.5 mg of O2/L throughout the experimental period. Water temperature ranged from 26 to 28°C, pH of 6.6–6.8, and salinity of 19–21 g/kg. The ammonia level was about 1.5 mg/kg. The water quality variables were recorded weekly. A photoperiod of 12 h light, 12 h dark (0800–2000 h) was used. Shrimp were fed their respective diets at a rate of 8 g/100 g body weight. This daily ration was subdivided into two equal feedings given at 0830 and 1730 h. Shrimp were weighed biweekly, and the daily ration was adjusted accordingly. Any dead shrimp were not replaced during the experiment. The shrimp were fed the test diets for 8 wk in expt. 1, and 6 wk in expt. 2. Throughout the experiment, the shrimp were maintained and used under humane conditions.
At the end of the feeding trial, the shrimp were weighed. Growth was
measured by the percentage of body weight gain [100(final body
wt-initial body wt)/initial body wt], feed efficiency (FE) and
protein efficiency ratio (PER) were calculated as described previously
(Shiau and Chou 1991
, Shiau and Liu 1994
). After the final weighing, three shrimp were randomly
removed from each aquarium, blood samples were collected from the
ostium of each shrimp and pooled for blood triglyceride and cholesterol
concentrations estimation (Carson and Goldfard 1979
).
Hepatopancreata and eyes were removed and pooled for total lipid
determination (Folch et al. 1957
) and vitamin A
(retinol) determination (Alava et al. 1993
),
respectively. Three other shrimp were then taken randomly from each
aquarium and pooled for body composition analysis (AOAC 1995
).
Statistical analysis.
Each experimental diet was fed to three groups of shrimp. Growth data
were means of three groups of shrimp, with 16 shrimp per group
(n = 3) in expt. 1 and 20 shrimp per group
(n = 3) in expt. 2. Blood triglyceride
concentration, hepatopancreatic total lipid concentration and eye
vitamin A concentration were means of the three groups of shrimp, with
three shrimp randomly selected from each group and pooled
(n = 3). Results were analyzed by one-way
ANOVA. When the ANOVA identified differences among groups, multiple
comparisons among means were made with Duncan’s new multiple range
test. Statistical significance of difference was determined by setting
the aggregate type I error at 5% (P < 0.05) for
each set of comparisons. Dietary vitamin A requirements for juvenile
P. monodon were estimated by the broken-line method
(Robbins 1986
).
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RESULTS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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The weight gains were significantly higher (P < 0.05)
in shrimp fed the diet supplemented with 3,000 RE vitamin A/kg than
shrimp fed the control group and shrimp fed diets supplemented with
30,000 RE vitamin A/kg (Table 2
). Patterns of difference in FE and PER were similar to those of the
weight gain. Survival of shrimp fed the control diet was significantly
lower (P < 0.05) than shrimp fed diets supplemented
with 1,500–30,000 RE vitamin A/kg.
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15,000 RE vitamin A/kg, followed by shrimp fed diet with 30,000 RE
vitamin A/kg and lowest in shrimp fed diets with 45,000 RE vitamin
A/kg. The differences between the highest groups and the lowest groups
were significant (Table 3
). Hepatopancreatic total lipid concentration was highest for shrimp fed
diets supplemented with 45,000 RE vitamin A/kg, intermediate for
shrimp fed the diet supplemented with 15,000 and 30,000 RE vitamin
A/kg, and lowest for shrimp fed the diet supplemented with 3,000 RE
vitamin A/kg. The differences between the three groups were all
significantly different from one another. Eye vitamin A concentration
was highest in shrimp fed the diet supplemented with 60,000 RE vitamin
A/kg, followed by shrimp fed the diets supplemented with 15,000, 30,000
and 45,000 RE vitamin A/kg and lowest in shrimp fed the diets
supplemented with 3,000 RE vitamin A/kg. The differences among the
three groups were significant.
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15,000 RE vitamin A/kg.
The differences between the highest groups and the lowest groups were
significant (Table 4
). The moisture, crude protein and ash concentrations of shrimp were not
affected by the dietary treatments.
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FE was highest in shrimp fed the diets supplemented with 2,400, 3,000,
3,600 and 4,500 RE vitamin A/kg diet, followed by the groups fed 600
and 1,200 RE vitamin A/kg, and finally the unsupplemented control group
(Table 5
,P < 0.05). Patterns of difference in PER were similar
to those of the FE ratio. The survival percentage was significantly
higher in shrimp fed diets supplemented with vitamin A than in shrimp
fed the unsupplemented control diet.
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). Based on the weight gain percentage, the regression equations were
Y = 0.055X + 112.64 (r = 0.98) and Y = -0.0069X + 268.06
(r = 0.73). Because the point of intersection between
the two lines is 2,511, the adequate amount of dietary vitamin A for
juvenile P. monodon was estimated to be 2,511 RE/kg (about
8,400 IU/kg).
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DISCUSSION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONREFERENCES |
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).
Retinoids, or their precursors, are essential in the diet of
vertebrates (Blomhoff et al. 1991
, Wolf 1984
). Their function as a component of retinal pigment in the
eye has long been known. Recent research showed however, that they have
a more fundamental function in gene transcription and thus are
essential in cell division and differentiation (Blomhoff et al. 1991
, Ross 1993
,Wolf 1984
).
Hence, retinoids are of particular importance in young, rapidly growing
vertebrates. Since invertebrates cannot synthesize carotenoids, and
thus cannot synthesize retinoids, it would be expected that retinoids
are essential to their metabolism, but in crustaceans the situation is
unclear. Fisher et al. (1952)
found very large quantities of retinoids
in the eyes of euphausiid species but only small amounts in the rest of
the body. In various other crustacean species, retinoids were either
absent or constituted <1µg/g wet mass. In eight species of
Penaeidea, some pelagic or deep-sea species had no retinoids in the
eyes and only trace amounts or none in the body. Shallow-water
species such as Penaeus aztecus had none in the body, but 4
µg/g in the eyes (Fisher et al. 1957
). A similar
situation was found in the freshwater crayfish Orconectes
rusticus (Wolfe and Cornwell 1965
). Thus, retinoids
if present at all in crustaceans, are concentrated in the eyes
suggesting that their principal function is in vision. It was also
suggested that vitamin A supplementation is unnecessary when fish oils
and carotenoids are added to diets for nonfeeding stage of P.
semisulcatus (Dall 1995
). In the present study 5%
fish oil was used in the basal diet to meet the requirement of the
shrimp (Sheen et al. 1994
). Nevertheless, vitamin A
supplementation is needed. He et al. (1992)
used 4,800 IU
of vitamin A/kg in their investigation with P. vannamei and
qualitatively evaluated the dietary essentiality of fat-soluble
vitamins, A, D, E and K. There was only one dietary vitamin A inclusion
level in their study (i.e., 4,800 IU/kg), and they did not
provide formal estimates of dietary requirement.
Shrimp fed the vitamin A-unsupplemented diet showed signs of
deficiency including light coloration beginning at wk 4 and increasing
mortality at wk 6 of the experiment. We cannot speculate about possible
reasons for the deficiency signs in the shrimp because no histological
examinations on the shrimp were conducted. Information on vitamin
deficiencies in shrimp is scarce. The only well-documented vitamin
deficiency in a shrimp species is the Black Death syndrome related to
vitamin C deficiency in penaeid shrimp (Lightner et al. 1977
).
The most marked effect of retinoid feeding seen in animal studies is
that of hypertriglyceridemia. In rats, oral
all-trans-retinoic acid, 13-cis-retinoic
acid and natural vitamin A resulted in hypertriglyceridemia
(Gerber and Erdman 1982
). Singh et al. (1969)
found elevated plasma-free fatty acids and increased liver lipids
in vitamin A-fed rats and postulated that these changes were caused
by a mobilization of fatty acids from adipose tissue. Ramachandran et al. (1986)
indicated that fatty acid homeostasis in rats can be greatly
influenced by their vitamin A status in which hypervitaminosis A caused
a decrease in the fatty acid release from adipose. On the other hand,
adipose tissue from vitamin A-deficient animals showed an increased
lipolytic rate as compared to that of the controls. A similar
phenomenon was also reported in fish. The reduced percentage of body
fat in the trout fed excess vitamin A may have resulted from a similar
mobilization of fatty acids in body fat (Poston 1970
). High body content of crude fat was also found in
vitamin A-deficient guppies (Shim and Tan 1990
).
However, in the present study, decreased blood triglyceride
concentrations in shrimp were associated with higher vitamin A in their
diets (Table 3)
and high body fat content was found in shrimp fed high
vitamin A diets (Table 4)
. Generally speaking, body fat content in
crustaceans is in the range of 1–2% which is much lower than other
animals. The extent to which the low body lipid affects fatty acid
mobilization is not known. The mechanism of dietary vitamin A
concentration on lipid nutrition in crustaceans requires further
investigation.
We could not measure the vitamin A concentration in the hepatopancreas
of shrimp. Vitamin A concentration in the eyes of shrimp fed vitamin A
supplemented diets generally increased as the vitamin A supplementation
level increased (Table 3)
, indicating that the tissue vitamin A
concentration of shrimp did not plateau. It was reported that higher
levels of nutrients are needed to maximize tissue concentration. For
example, studies of the riboflavin requirement of rainbow trout
indicated a growth requirement (dietary level needed to achieve maximal
growth) of 3.6 mg/kg, a requirement of 4.6 mg/kg for liver flavin
saturation and a requirement of 6.6 mg/kg for saturation of spleen and
head kidney with flavin compounds (Woodward 1985
).
However, it is probably undesirable to consider tissue vitamin A
saturation level as an indication of the requirement because excess
tissue vitamin A results in toxicity.
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FOOTNOTES |
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3 Abbreviations used: FE, feed efficiency; IU,
international unit; PER, protein efficiency ratio; RE, retinol equivalent.
Manuscript received August 30, 1999. Initial review completed September 10, 1999. Revision accepted September 16, 1999.
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