Effects of feeding level on the growth, energy budget and body biochemical composition of Chinese shrimp Fenneropenaeus chinesis.

ABSTRACT Chinese shrimp Fenneropenaeus chinensis with wet weight of 3.0 g, 8.0 g, 14.2 g, and 19.0 g were fed at 5 feeding levels of 0, 0.5%, 1.0%, and 1.5% of the body weight, and ad libitum to investigate the growth and food conversion efficiency under different feeding levels. The relationship among daily growth coefficient (DGC) (in terms of wet weight, dry weight, protein, and energy), initial body weight (IBW) and feeding level (RL) was well described with the dualistic linear model DGC = B0 + B1 x I[S.sub.W] + B2 x FL. The relationship between food efficiency (FE) in terms of dry weight, protein, and energy of shrimp with different body weight and feeding level was described with the model FC = B0 + B1 x lnFL. DGC and IBW were negatively correlated, regardless of term of wet weight, dry weight, protein, or energy of the shrimp body weight. The maintenance feeding level (MFL) of Chinese shrimp was positively linear correlated to IBW and the maintainable energy intakes (MEI, J/g BW/d) for shrimp with different IBW were: 3.0 g, 218.3/g BW/d; 8.0 g, 213.5 J/g BW/d; 14.2 g, 207.5 J/g BW/d; 19.0 g, 202.9 J/g BW/d.

KEY WORDS: feeding level, daily growth coefficient, energy budget, body biochemical composition, Chinese shrimp

INTRODUCTION

Either in natural environment or culture ponds, the crustacean experience starvation and under-nutrition occasionally. Studies have found that starvation caused low metabolic rate (Wallace 1973, Regnault 1981, Whyte et al. 1986, Chapelle et al. 1994) and high body moisture content (Dall 1974, Cuzon et al. 1980, Barclay et a1.1983, Sasaki et al. 1986, Stuck et al. 1996a, Wu & Dong 2000, Wu & Dong 2002a, Wu & Dong 2002b, Lin et al. 2004) in crustacean. Other items of shrimp body composition such as protein, lipid, and energy content decrease gradually through the fasting period (Barclay et a1.1983, Stuck et al. 1996a, Wu et al. 2000, Wu & Dong 2002a, Wu & Dong 2002b, Lin et al. 2004). Positive correlation existed between the growth and feeding level in shrimp and crabs (Bartley et al. 1980, Miao & Tu 1993, Gu et al. 1996). Besides, the feeding level affected body composition, such as moisture, protein, and lipid content of American lobster Homarus, americanus (Bartley et al. 1980) and redclaw crayfish, Cherax, quadricarinatus (Gu et al. 1996). To explore the physio-ecologic characters of shrimp in different nutritional conditions, the responses of growth, FE, energy budget, body biochemical composition, and body gross energy content to feeding levels of Chinese shrimp, Fenneropenaeus chinensis with different body weight, were studied in this experiment.

MATERIALS AND METHODS

Diet Used in This Experiment

The diet used in this study was a commercial formulated shrimp pellet (Sea-Horse Brand; Producer: Fujian Mawei Unite Feed Ltd. Co., China). The proximal composition of the diet was: moisture, 7.70%; crude protein, 42.57%; crude lipid, 9.93%; ash, 10.75%; and gross energy 19.23 KJ.[g.sup.-1].

Rearing Conditions

Chinese shrimp were kept in glass aquaria (45 x 30 x 30 [cm.sup.3], water volume of 35 [dm.sup.3]), and each rearing unit was stocked with 1 shrimp. The room temperature was thermostatically controlled, and water temperature was 25[degrees] [+ or -] 05[degrees]. Aeration was provided continuously and 1/2 to 2/3 of water was exchanged every other day. Seawater used in the experiment was filtered by composite sand filter. During the experiment, dissolved oxygen water was maintained above 5.5 mg/L, pH was about 8.0, the water salinity was between 30[per thousand] to 33[per thousand], and the photoperiod of 14 hours of light: 10 hours of darkness was used.

Source and Acclimation of Shrimp

The experiment was carried out at the Mariculture Research Laboratory, Ocean University of China, People's Republic of China. The shrimp used in the experiment were collected from the Tianheng Shrimp Farm, Qingdao. Prior to the experiment, the shrimp were transferred into aquaria and underwent a 7-day acclimatization period during which the shrimp were fed with the formulated diet (FD) at satiation level twice daily (at about 6:00AM and 6:00PM).

Experiment Design

After 24 hours starvation, shrimp with wet weight of 3.0 [+ or -] 0.2 g, 8.0 [+ or -] 0.5 g, 14.2 [+ or -] 0.9 g, and 19.0 [+ or -] 1.4 g (Mean [+ or -] SD) were selected from acclimated shrimp. For every body weight size, shrimp were fed at 5 feeding levels (0, 0.5%, 1.0%, 1.5% wet body weight, and satiation level) for 28 days. Six groups for every treatment were designed and a complete randomized block design was used to arrange aquaria.

Samples Collection and Analysis

Three groups (8 shrimp each) were sampled from the acclimated shrimp to determine the initial body composition of experimental shrimp. After 28 days all groups of shrimp were starved for 24 hours, and then sampled. The treated shrimp were pooled as three samples randomly (2 shrimp for a sample to get enough material for analyzing) and there were 60 samples (3 per treatment) of final shrimp.

During the course of the experiment the daily food supplied to shrimp was recorded and uneaten food and feces were collected within 3 hours after feeding. Shrimp and food were weighted using an electronic scale after carefully blotted with paper towel to remove excess moisture.

After weighing all samples of shrimp, feces, and food were dried in an oven at 70[degrees] to constant weight, homogenized with a glass mortar, and stored at -20[degrees]. Before chemical compositions were analyzed, the samples were re-dried at 70[degrees] to constant weight.

Nitrogen content was measured using the MicroKjeldahl methods and the crude protein content was calculated (N x 6.25) (AOAC, 1984). Crude lipid was determined by the Soxthlet method (AOAC, 1984), ash was determined by combusting dry samples in a muffle furnace at 550[degrees] for 12 h (AOAC, 1984), and the gross energy content of dry samples was determined with PARRI281 bomb calorimeter (PARR Instrument Company, USA). Analyses of each sample were conducted in triplicate (3 subsamples for each sample).

Calculation of Data

The FE, % was calculated as follow:

FE = 100 x (FBW - IBW) / I

Where the shrimp body weight was dry weight (DW, g), the diet was expressed in terms of dry weight (DW, g), FBW was the final shrimp weight, IBW was the initial shrimp weight, I was the quantity of ingested diet during the experiment period.

All samples in terms of dry weight (DW, g), protein (P, g), and energy (E, KJ) were used to calculate the FE (F[E.sub.d], F[E.sub.p], F[E.sub.e]) respectively.

The DG[C.sub.W] was calculated as follow (Cowey 1992):

DGC = (FB[W.sup.1/3] - IB[W.sup.1/3]) / D

Where D was the days of experiment period. All samples in terms of dry weight (DW, g), protein (P, g), and energy (E, K J) were used to calculate the DGC (DG[C.sub.d], DG[C.sub.p], DG[C.sub.e]) respectively.

The feeding level (mg/g BW/d) was calculated as follow:

FL = 2 x 1000 x I/(FBW + IBW)/D

Items in the energy budget equation ([C.sub.e] = [G.sub.e] + [F.sub.e] + [E.sub.e] + [U.sub.e] + [R.sub.e]) (Klein 1975) of shrimp were calculated as follow:

Energy intake ([I.sub.e]) = [I.sub.d] x G[E.sub.d]

Growth energy ([G.sub.e]) = F[S.sub.e] - I[S.sub.e]

Energy of feces ([F.sub.e]) = [F.sub.d] x G[E.sub.f]

Energy of exuviae ([E.sub.e]) = [E.sub.d] x G[E.sub.e]

Energy of excretion ([U.sub.e]) = [U.sub.N] x 24.83

Energy of metabolism ([R.sub.e]) = [I.sub.e] - [G.sub.e] - [F.sub.e] - [E.sub.e] - [U.sub.e]

Where G[E.sub.d], G[E.sub.f], and G[E.sub.e] were gross energy of diet, feces, and exuviations respectively, [F.sub.d], [E.sub.d], and [U.sub.N] were dry weight of diet, dry weight of feces and nitrogen of excretion respectively. F[S.sub.e] and I[S.sub.e] were energy content of final shrimp and initial shrimp. 24.83 means the energy cost of 1g nitrogen excreting (in form of NH3) is 24.83 KJ. The daily unit body weight energy partitions were calculated by dividing 28 x (FBW + IBW)/2.

The nitrogen of excretion ([U.sub.N]) was calculated as the follow formula (Levine & Sulkin 1979, Lemos and Phan 2001):

[U.sub.N] = [I.sub.N] - [G.sub.N] - [F.sub.N] - [E.sub.N]

Where [I.sub.N], [G.sub.N], [F.sub.N], and [E.sub.N] were nitrogen content in ingested diets, body growth, feces and exuviations.

The percentage of growth energy ([G.sub.e]) in energy consumption ([C.sub.e]) was calculated as 100 x [G.sub.e]/[I.sub.e], and the percentages of [F.sub.e], [E.sub.e], [U.sub.e], and [R.sub.e] in [I.sub.e] were calculated similarly.

Statistical Analysis

Statistics were performed using SPSS10.0 statistical software with possible differences among diet treatment being tested by l-way ANOVA. Tukey's b-multiple range tests were used to test differences between treatment groups of the same size. The square-root transformation of the sine-arc before analyzing the values given in percentages was used. Differences were considered significant at a probability level of 0.05. The curve estimation was used in regression analysis.

RESULTS

Relationship Between Daily Growth Coefficient and Feeding Level

The dualistic linear model (DGC = [B.sub.0] + [B.sub.1] x I[S.sub.W] + [B.sub.2] x FL) could describe the relationship among the DGC of Chinese shrimp, its initial body weight (IBW) and feeding levels (FL) (Table 1). Whatever the shrimp body weight and feeding levels were expressed by wet weight, dry weight, protein or energy, the values of parameter [B.sub.1] and [B.sub.2] were positive, which means the DGC of shrimp were positively linear correlative to FL and IBW.

The Relationship Between the Food Efficiency and Feeding Level

The FE of Chinese shrimp was significantly correlative to FL, and the linear model FE = [B.sub.0] + [B.sub.1] x lnFL could describe the relationships between FE and lnFL of the shrimp with four body weight sizes (Table 2). However, with the decreasing of the initial body weight the correlativity (see [R.sup.2] in Table 2) was improved.

The Energy Allocation of Chinese Shrimp Fed at Different Feeding

Table 3 denoted the models of energy allocation of Chinese shrimp with different body weight at different feeding levels. The maximal of feeding levels was different in shrimp with different body weight, and it declined when the body weight rose (Table 3). When the shrimp were fed at the satiation level, the average gross energy intake of single shrimp over the experimental period was 1.74 KJ, 3.37 KJ, 3.89 KJ, and 5.43 KJ respectively for shrimp with different IBW. The fasting energy loses during the experimental period varied from 8.6 KJ to 43.2 KJ according to different sizes (Table 4). The energy loss through respiration (%IE) was greater in small shrimp than that in large shrimp, and energy loss of exuviations (%IE) was greater in large shrimp than that in small shrimp (Table 4). For daily unit body weight energy allocation (J/g BW/d), [I.sub.e], [G.sub.e], [R.sub.e], [F.sub.e], and [U.sub.e] increased as FL increased, regardless of IBW. It seemed that FL did not significantly affected [E.sub.e] in all IBW.

The Biochemical Composition and Gross Energy Content of Final Shrimp Fed at Different Feeding Levels

The differences of biochemical composition of four size shrimp fed with different feeding levels were listed in Table 5. In the same size, the body moisture contents of the shrimp fed low feeding level were higher than that of initial shrimp, and it decreased with the increasing of feeding level. The range of the body moisture content of all treatments was from 75.1% to 87.9%. In low FL treatments, the protein content, lipid content and gross energy of shrimp body were significantly lower than those of the initial shrimp, and they increased along with the increasing of feeding level. The body protein content, lipid content and gross energy of four sizes shrimp fed at satiation level (from small to large) were listed below: protein, 15.2%; 19.3%; 17.9%; and 16.4%; lipid, 2.1%; 2.1%; 1.8%; and 2.8%; gross energy, 4.5KJ/g; 4.7 KJ/g; 4.6 KJ/g; and 5.1 KJ/g. It was found that no significant difference existed among different body weight sizes in the protein content, lipid content and gross energy when the shrimp were fed at satiation level.

It was found in the starved and low feeding level treatment that the body moisture content of final shrimp was higher than that of initial shrimp and high feeding level treatment. But the body protein content, lipid content and gross energy of starved and low feeding level treatment were lower than those of initial shrimp and high feeding level treatment. These indicated that the ingested nutrients and energy could not meet the requirement of minimal metabolism in low feeding level treatment, which resulted in the decreasing of body nutrient and energy deposits.

Energy Loss of Fasting Shrimp and Maintainable Feeding Level

It was found that the total energy loss for individual shrimp during the experimental period was directly related to IBW. For unit body weight energy loss per day, the small shrimp (3.0g and 8.0g) lose more energy than large shrimp (14.2 g and 19.0 g). [R.sub.e] accounted for most of the energy loss regardless of IBW. The relationship between MFL and IBW could be achieved from the relationship between DGC, FL, and IBW (supposed DGC = 0). A linear model was achieved (Table 6) and it inferred that MFL was inversely correlated to IBW. The MIE, (J/g BW/d) for shrimp with different IBW is listed in Table 7. The linear model indicated MIE was also inversely correlated to IBW.

DISCUSSION

Growth Model of Chinese Shrimp

In this study, it was found that the compound relationship among the special growth rate DGC of Chinese shrimp, feeding level (FL) and initial body weight (IBW) were duality linear. It is to say that in every body weight size the DGC and FL were significantly linearly correlated. Similar positive correlations between growth and ration size have been found in studies on shrimp and crabs (Lee 1971, Venkataramiah et al. 1975, Caillouet et al. 1976, Sedgwick 1979, Bartley et al. 1980, Viayaraghavan et al. 1982, Maguire & Leedow 1983, Gu et al. 1996). In this model, when DGC was calculated in wet weight, [B.sub.1] was less than one third and [B.sub.1] was less than one twentieth of those when DGC calculated in dry weight, protein, or energy (Table 1). This particularity resulted from variability of body moisture content in different conditions. The increase of body moisture content in under-nutritional condition and the reduction of body moisture content in nutrition satiation condition would conceal the alteration of wet body weight.

Effect of Feeding Level on Energy Allocation

In this study, dally unit body weight energy allocation (J/g BW/d) of [R.sub.e] and [U.sub.e] were positively correlated to FL regardless of IBW. This result was in agreement with results found in studies on the relationship between metabolic rate or excrete rate and feeding level or ration size in other crustacean (Hewitt & Iving 1990, Du Preez et al. 1992, Burggren et al. 1993, Mcgaw & Reiber 2000, Peck & Veal 2001). Directly positive relationship existed between [G.sub.e] and FL. This relationship supported the relationship between DGC and FL. It indicated that the more shrimp ate, the faster the shrimp grew. This was also similar to other studies (Lee 1971, Venkataramiah et al. 1975, Calllouet et al. 1976, Sedgwick 1979, Bartley et al. 1980, Viayaraghavan et al. 1982, Maguire & Leedow 1983, Gu et al. 1996).

Effect of Feeding Level on Body Composition

In this study, starvation and low feeding levels caused increase in body moisture content and reduction in protein, lipid, and energy content of Chinese shrimp, which is in agreement with previous studies on Chinese shrimp (Wu & Dong 2002a, Wu & Dong 2002b) and other species of crustacean (Dall 1974, Cuzon et al. 1980, Barclay 1983, Sasaki et al. 1986, Stuck et al. 1996a, Lin et al. 2004). It could be concluded that starvation and limited food supply would bring similar effects on body composition caused by other environmental stress such as pollution (Wang & Stickle 1988), infection (Stuck et al. 1996b), and nutrition stress (Lim 1997, Stuck et al. 1996a). There was an inverse relationship between body moisture content and growth, whereas the protein, lipid, and energy content were directly related to growth. Similar results have been found in other studies (Stuck et al. 1996a, Lim 1997). Besides, the protein, lipid, and energy content were negatively correlated to moisture content. Weatherley (1981) proposed to form a significant relationship between moisture and other composition such as protein, lipid, and energy in fish. It was easy to determine the moisture content. Then other composition would be easily calculated. But this method is unsuitable for shrimp. Except for nutritional conditions, the smolting cycle also affected the body composition of shrimp (Read & Caulton 1980).

Maintenance Feeding Level of Chinese Shrimp

The metabolic level of unit body weight of animals decline along with the rising of body weight, therefore the MFL of their unit body weight is negatively correlated to their actual body weight. It could be concluded from the values of [A.sub.1] that MFL is positively linear correlated to IBW, regardless of terms described in the data. The MEI (J/g BW/d) is inversely related to IBW. It is also a sign of reduction in metabolic level as BW increased. This fact correlates with results of other studies (Nelson et al. 1977, Emmerson 1985, Dong et al. 1994, Zhang et al. 1998). It was found that maintenance energy intakes (MIE) are higher than unit body weight energy loss of fasting shrimp. For large shrimp, the difference was more obvious. This could attribute to the increase of metabolic rate after ingestion of food and the decrease of unit body weight energy loss.

TABLE 1.

The relationship among the daily growth coefficient (DGC), feeding
levels (mg/g shrimp/day), and initial shrimp weight (g).

DGC = [B.sub.0] + [B.sub.1] x FL + [B.sub.2] x IBW (n = 120)

Model          [B.sub.0] (Mean [+ or -]    [B.sub.1] (Mean [+ or -]
                         SE)                         SE)

DG[C.sub.w]     -0.236 [+ or -] 0.028        0.021 [+ or -] 0.002
DG[C.sub.d]     -1.227 [+ or -] 0.059        0.067 [+ or -] 0.003
DG[C.sub.p]     -1.487 [+ or -] 0.073        0.078 [+ or -] 0.004
DG[C.sub.e]     -1.611 [+ or -] 0.072        0.083 [+ or -] 0.004

Model          [B.sub.2] (Mean [+ or -]
                         SE)                 R        F         P

DG[C.sub.w]      0.001 [+ or -] 0.001      0.780     90.63    <0.01
DG[C.sub.d]      0.024 [+ or -] 0.004      0.888    218.94    <0.01
DG[C.sub.p]      0.033 [+ or -] 0.005      0.879    198.33    <0.01
DG[C.sub.e]      0.037 [+ or -] 0.004      0.893    231.53    <0.01

TABLE 2.

The relationship between the food conversion and feeding level
(mg/g shrimp/day).

FC = [B.sub.0] + [B.sub.1] x InFL (n = 30)

      Model                      [B.sub.   [B.sub.   [R.sup.
     IBW (g)            FC         0]        1]        2]        P

3.0 [+ or -] 0.2    F[C.sub.d]   -174.2      60.6     0.90     <0.01
                    F[C.sub.p]   -317.3     110.6     0.90     <0.01
                    F[C.sub.e]   -203.5      70.0     0.91     <0.01
8.0 [+ or -] 0.5    F[C.sub.d]    -76.5      29.2     0.65     <0.01
                    F[C.sub.p]   -155.2      57.1     0.72     <0.01
                    F[C.sub.e]    -87.1      32.7     0.66     <0.01
14.2 [+ or -] 0.9   F[C.sub.d]   -146.8      58.1     0.87     <0.01
                    F[C.sub.p]   -217.0      86.1     0.87     <0.01
                    F[C.sub.e]   -155.5      61.1     0.88     <0.01
19.0 [+ or -] 1.4   F[C.sub.d]   -102.6      39.8     0.84     <0.01
                    F[C.sub.p]   -150.0      57.7     0.81     <0.01
                    F[C.sub.e]   -111.6      43.9     0.86     <0.01

TABLE 3.

The energy intake (IE, KJ) and allocation of Chinese shrimp at
different feeding levels (mg/g shrimp/day).

                                  Energy ingested and allocation

                                               IE

       IBW           FL        KJ/shrimp                J/gBW/d

 3.0 [+ or -] 0.2    4.4   0.24 [+ or -] 0.01    84.7 [+ or -] 2.8 (a)
                     7.9   0.44 [+ or -] 0.04   152.1 [+ or -] 14.5 (b)
                    17.7   1.13 [+ or -] 0.06   344.8 [+ or -] 23.4 (c)
                    25.4   1.74 [+ or -] 0.15   488.9 [+ or -] 36.2 (d)
 8.0 [+ or -] 0.5    8.7   1.31 [+ or -] 0.03   167.4 [+ or -] 3.4 (a)
                    13.1   2.05 [+ or -] 0.11   251.5 [+ or -] 11.3 (b)
                    17.2   2.82 [+ or -] 0.05   331.8 [+ or -] 9.5 (c)
                    21.2   3.37 [+ or -] 0.07   407.5 [+ or -] 11.8 (d)
14.2 [+ or -] 0.9    4.7   1.22 [+ or -] 0.04    90.3 [+ or -] 1.7 (a)
                     9.1   2.45 [+ or -] 0.19   175.6 [+ or -] 11.9 (b)
                    12.1   3.34 [+ or -] 0.13   233.2 [+ or -] 7.4 (c)
                    14.2   3.89 [+ or -] 0.16   272.8 [+ or -] 9.1 (d)
19.0 [+ or -] 1.4    4.4   1.61 [+ or -] 0.04   84.69 [+ or -] 1.2 (a)
                     9.0   3.23 [+ or -] 0.56   173.8 [+ or -] 25.9 (b)
                    12.0   4.44 [+ or -] 0.10   231.7 [+ or -] 4.4 (c)
                    14.3   5.43 [+ or -] 0.26   279.5 [+ or -] 8.2 (d)

                              Energy ingested and allocation

                                            GE

       IBW                    %IE                      J/gBW/d

 3.0 [+ or -] 0.2   -110.3 [+ or -] 6.2 (a)    -92.5 [+ or -] 4.7 (a)
                     -47.3 [+ or -] 8.8 (b)    -67.1 [+ or -] 6.6 (b)
                       1.2 [+ or -] 0.2 (c)     15.8 [+ or -] 5.6 (c)
                      14.0 [+ or -] 2.6 (d)     65.0 [+ or -] 11.6 (d)
 8.0 [+ or -] 0.5    -19.5 [+ or -] 2.6 (a)    -32.2 [+ or -] 4.1 (a)
                      -0.2 [+ or -] 0.1 (b)      8.0 [+ or -] 0.6 (b)
                      10.7 [+ or -] 0.8 (c)     33.7 [+ or -] 1.9 (c)
                      12.1 [+ or -] 4.3 (c)     34.8 [+ or -] 3.6 (c)
14.2 [+ or -] 0.9    -64.1 [+ or -] 4.l (a)    -57.8 [+ or -] 3.9 (a)
                     -16.3 [+ or -] 19.1 (b)   -23.7 [+ or -] 7.8 (b)
                      -4.5 [+ or -] 0.9 (bc)   -10.9 [+ or -] 6.2 (c)
                       2.5 [+ or -] 0.8 (c)      6.6 [+ or -] 0.9 (d)
19.0 [+ or -] 1.4    -44.3 [+ or -] 4.2 (a)    -37.4 [+ or -] 3.4 (a)
                     -22.9 [+ or -] 2.0 (b)    -30.8 [+ or -] 2.9 (a)
                       1.5 [+ or -] 0.6 (c)      3.7 [+ or -] 0.8 (b)
                       2.4 [+ or -] 0.6 (c)      5.8 [+ or -] 0.9 (b)

                              Energy ingested and allocation

                                            FE

       IBW                    %IE                      J/gBW/d

 3.0 [+ or -] 0.2    13.4 [+ or -] 1.6 (b)      11.3 [+ or -] 1.3 (a)
                      9.0 [+ or -] 1.0 (a)      13.1 [+ or -] 0.4 (a)
                     13.8 [+ or -] 1.2 (b)      48.1 [+ or -] 6.0 (b)
                     17.5 [+ or -] 1.7 (b)      77.2 [+ or -] 6.0 (c)
 8.0 [+ or -] 0.5    13.1 [+ or -] 0.4 (a)      21.8 [+ or -] 0.5 (a)
                     13.2 [+ or -] 3.8 (a)      39.8 [+ or -] 2.1 (b)
                     14.0 [+ or -] 0.5 (a)      45.2 [+ or -] 1.4 (c)
                     12.8 [+ or -] 0.5 (a)      47.7 [+ or -] 2.0 (c)
14.2 [+ or -] 0.9    14.0 [+ or -] 0.3 (a)      12.7 [+ or -] 0.4 (a)
                     15.8 [+ or -] 0.4 (a)      27.5 [+ or -] 1.6 (b)
                     18.0 [+ or -] 1.O (b)      42.2 [+ or -] 3.4 (c)
                     23.3 [+ or -] 0.8 (c)      63.6 [+ or -] 3.5 (d)
19.0 [+ or -] 1.4     9.4 [+ or -] 0.5 (a)       8.0 [+ or -] 0.3 (a)
                     14.1 [+ or -] 0.9 (bc)     24.2 [+ or -] 3.3 (b)
                     12.6 [+ or -] 0.5 (b)      29.3 [+ or -] 1.2 (b)
                     14.4 [+ or -] 0.5 (c)      40.0 [+ or -] 1.6 (c)

                              Energy ingested and allocation

                                            EE

       IBW                    %IE                      J/gBW/d

 3.0 [+ or -] 0.2    12.2 [+ or -] 1.4 (b)     10.3 [+ or -] 1.2 (a)
                     10.6 [+ or -] 1.4 (b)     15.5 [+ or -] 0.9 (b)
                      4.7 [+ or -] 0.3 (a)     16.7 [+ or -] 1.2 (b)
                      4.2 [+ or -] 0.5 (a)     17.2 [+ or -] l.O (b)
 8.0 [+ or -] 0.5     8.6 [+ or -] 0.8 (b)     14.3 [+ or -] 1.3 (a)
                      4.5 [+ or -] 0.7 (a)     13.3 [+ or -] 0.5 (a)
                      3.7 [+ or -] 0.2 (a)     11.7 [+ or -] 0.9 (a)
                      3.5 [+ or -] 0.1 (a)     13.4 [+ or -] 0.4 (a)
14.2 [+ or -] 0.9    15.1 [+ or -] 0.7 (c)     13.6 [+ or -] 0.6 (a)
                      9.1 [+ or -] 1.2 (b)     15.0 [+ or -] 0.5 (a)
                      5.9 [+ or -] 0.6 (ab)    13.6 [+ or -] 1.2 (a)
                      5.0 [+ or -] 0.8 (a)     13.5 [+ or -] 2.1 (a)
19.0 [+ or -] 1.4    11.3 [+ or -] 1.O (b)      9.5 [+ or -] 0.8 (a)
                     10.8 [+ or -] 2.3 (b)     17.1 [+ or -] 2.3 (b)
                      6.2 [+ or -] 0.4 (a)     14.3 [+ or -] 0.9 (b)
                      5.7 [+ or -] 0.6 (a)     15.6 [+ or -] 1.4 (ab)

                              Energy ingested and allocation

                                            RE

       IBW                    %IE                      J/gBW/d

 3.0 [+ or -] 0.2    166.0 [+ or -] 5.1 (d)    140.0 [+ or -] 2.6 (a)
                     117.5 [+ or -] 5.8 (c)    175.5 [+ or -] 9.0 (ab)
                      74.4 [+ or -] 1.9 (b)    252.8 [+ or -] 20.4 (b)
                      59.9 [+ or -] 3.2 (a)    306.3 [+ or -] 30.5 (c)
 8.0 [+ or -] 0.5     88.2 [+ or -] 2.4 (c)    147.5 [+ or -] 3.9 (a)
                      75.2 [+ or -] 3.8 (b)    173.6 [+ or -] 11.9 (a)
                      65.7 [+ or -] 1.2 (a)    221.2 [+ or -] 7.6 (b)
                      65.6 [+ or -] 3.9 (a)    285.2 [+ or -] 2.0 (c)
14.2 [+ or -] 0.9    121.7 [+ or -] 3.3 (c)    109.9 [+ or -] 3.7 (a)
                      82.8 [+ or -] 4.5 (b)    142.1 [+ or -] 6.3 (b)
                      73.8 [+ or -] 1.5 (b)    172.3 [+ or -] 7.9 (c)
                      63.7 [+ or -] 1.4 (a)    173.9 [+ or -] 8.3 (c)
19.0 [+ or -] 1.4    111.5 [+ or -] 3.4 (c)     94.4 [+ or -] 2.8 (a)
                      88.6 [+ or -] 7.8 (b)    147.9 [+ or -] 13.4 (b)
                      72.3 [+ or -] 1.5 (a)    167.2 [+ or -] 3.4 (bc)
                      70.4 [+ or -] 3.2 (a)    197.7 [+ or -] 13.7 (c)

                              Energy ingested and allocation

                                            UE

       IBW                    %IE                      J/gBW/d

 3.0 [+ or -] 0.2    18.7 [+ or -] 0.9 (c)     15.7 [+ or -] 0.4 (a)
                     10.2 [+ or -] 0.7 (b)     15.1 [+ or -] 0.5 (a)
                      5.9 [+ or -] 0.3 (a)     19.6 [+ or -] 2.0 (ab)
                      4.4 [+ or -] 0.3 (a)     23.1 [+ or -] 2.9 (b)
 8.0 [+ or -] 0.5     9.6 [+ or -] 0.4 (c)     16.0 [+ or -] 0.7 (a)
                      7.3 [+ or -] 0.3 (b)     16.6 [+ or -] 1.4 (a)
                      5.9 [+ or -] 0.2 (a)     20.0 [+ or -] 0.8 (a)
                      6.0 [+ or -] 0.4 (a)     26.3 [+ or -] 2.3 (b)
14.2 [+ or -] 0.9    13.3 [+ or -] 0.6 (c)     12.0 [+ or -] 0.6 (a)
                      8.7 [+ or -] 0.8 (b)     14.7 [+ or -] 0.7 (ab)
                      6.8 [+ or -] 0.3 (a)     15.9 [+ or -] 1.1 (b)
                      5.6 [+ or -] 0.2 (a)     15.2 [+ or -] 1.0 (b)
19.0 [+ or -] 1.4    12.0 [+ or -] 0.6 (a)     10.2 [+ or -] 0.5 (a)
                      9.5 [+ or -] 1.3 (a)     15.4 [+ or -] 0.1 (b)
                      7.4 [+ or -] 0.3 (b)     17.2 [+ or -] 0.5 (bc)
                      7.2 [+ or -] 0.5 (c)     20.3 [+ or -] 1.8 (c)

Values without same letters in the same parameter of the same body wet
weight were significantly different from each other.

The superscript letters a through d in this table were markers of
differences of values.

TABLE 4.

Energy loss during experimental period ([e.sub.1], KJ/shrimp)
and allocation of starved shrimp (Mean [+ or -] SE).

                          Energy loss

 IBW          KJ/shrimp                  J/g BW/d

 3.0     8.6 [+ or -] 04 (a)      173.7 [+ or -] 8.6 (b)
 7.9    12.7 [+ or -] 0.5 (a)     171.3 [+ or -] 24.8 (b)
14.5    32.6 [+ or -] 2.0 (b)      99.6 [+ or -] 5.3 (a)
19.1    43.2 [+ or -] 2.8 (c)     101.3 [+ or -] 6.9 (a)

                           [R.sub.e]

 IBW          %[E.sub.1]                 J/g BW/d

 3.0    84.0 [+ or -] 1.1 (b)     145.7 [+ or -] 6.7 (b)
 7.9    77.1 [+ or -] 2.0 (a)     132.2 [+ or -] 19.7 (b)
14.5    75.6 [+ or -] 1.7 (a)      75.4 [+ or -] 4.7 (a)
19.1    76.7 [+ or -] 1.2 (a)      77.7 [+ or -] 5.6 (a)

                           [U.sub.e]

 IBW          %[E.sub.1]                 J/g BW/d

 3.0    12.7 [+ or -] 0.2 (bc)    22.0 [+ or -] 1.0 (b)
 7.9    13.4 [+ or -] 0.4 (c)     23.0 [+ or -] 3.4 (b)
14.5    10.5 [+ or -] 0.4 (a)     10.5 [+ or -] 0.7 (a)
19.1    12.1 [+ or -] 0.2 (b)     12.3 [+ or -] 0.8 (a)

                           [E.sub.e]

 IBW          %[E.sub.1]                 J/g BW/d

 3.0     3.4 [+ or -] 1.3 (a)      6.0 [+ or -] 2.3 (a)
 7.9     9.5 [+ or -] 2.3 (ab)    16.1 [+ or -] 5.1 (a)
14.5    13.9 [+ or -] 2.0 (b)     13.8 [+ or -] 2.0 (a)
19.1    11.2 [+ or -] 1.3 (b)     11.3 [+ or -] 1.6 (a)

TABLE: 5.

The body biochemical composition and gross energy of shrimp fed at
different feeding levels (mg/g shrimp/day).

        IBW             FL           Moisture

 3.0  [+ or -] 0.2      IS    76.9 [+ or -] 0.5 (a)
                        0     87.9 [+ or -] 0.9 (c)
                        4.4   86.8 [+ or -] 0.5 (bc)
                        7.9   84.7 [+ or -] 0.8 (b)
                       17.7   78.5 [+ or -] 0.7 (a)
                       25.4   76.1 [+ or -] 0.8 (a)
  8.0 [+ or -] 0.5      IS    78.2 [+ or -] 0.6 (b)
                        0     84.0 [+ or -] 0.6 (d)
                        8.7   80.8 [+ or -] 0.4 (c)
                       13.1   75.9 [+ or -] 0.2 (ab)
                       17.2   78.2 [+ or -] 0.6 (b)
                       11.2   75.1 [+ or -] 1.6 (a)
14.2  [+ or -]  0.09    IS    75.9 [+ or -] 0.6 (a)
                        0     83.7 [+ or -] 1.0 (b)
                        4.7   82.4 [+ or -] 0.6 (b)
                        9.1   77.1 [+ or -] 0.8 (a)
                       12.1   76.8 [+ or -] 0.6 (a)
                       14.2   77.5 [+ or -] l.0 (a)
 19.0 [+ or -] 1.4      IS    75.9 [+ or -] 1.2 (a)
                        0     85.3 [+ or -] 0.3 (c)
                        4.4   80.5 [+ or -] 0.6 (b)
                        9.0   76.7 [+ or -] 0.9 (a)
                       12.0   78.8 [+ or -] 0.6 (ab)
                       14.3   76.8 [+ or -] 1.1 (a)

        IBW                    Protein                   Lipid

 3.0  [+ or -] 0.2     15.9 [+ or -] 0.4 (c)      1.8 [+ or -] 0.1 (c)
                        6.8 [+ or -] 0.1 (a)      1.0 [+ or -] 0.1 (a)
                        7.2 [+ or -] 0.1 (a)      1.0 [+ or -] 0.1 (a)
                       11.1 [+ or -] 0.4 (b)     11.1 [+ or -] 0.1 (a)
                       14.6 [+ or -] 1.3 (c)      1.5 [+ or -] 0.1 (b)
                       15.2 [+ or -] 0.6 (c)      2.1 [+ or -] 0.1 (c)
  8.0 [+ or -] 0.5     15.0 [+ or -] 0.2 (cd)     1.2 [+ or -] 0.1 (a)
                       10.0 [+ or -] 0.1 (a)      1.0 [+ or -] 0.1 (a)
                       11.8 [+ or -] 0.2 (b)      1.2 [+ or -] 0.1 (a)
                       14.5 [+ or -] 0.1 (c)      1.4 [+ or -] 0.1 (b)
                       16.2 [+ or -] 0.1 (d)      1.5 [+ or -] 0.1 (b)
                       16.3 [+ or -] 0.1 (e)      2.1 [+ or -] 0.1 (c)
14.2  [+ or -]  0.09   15.8 [+ or -] 0.2 (d)      1.8 [+ or -] 0.1 (e)
                       10.8 [+ or -] 0.1 (a)      0.8 [+ or -] 0.1 (a)
                       11.7 [+ or -] 0.3 (b)      1.0 [+ or -] 0.1 (b)
                       14.9 [+ or -] 0.1 (e)      1.3 [+ or -] 0.1 (c)
                       16.3 [+ or -] 0.1 (d)      1.6 [+ or -] 0.1 (d)
                       17.9 [+ or -] 0.2 (e)      1.8 [+ or -] 0.1 (e)
 19.0 [+ or -] 1.4     16.2 [+ or -] 0.2 (d)      1.7 [+ or -] 0.1 (bc)
                        8.8 [+ or -] 0.1 (a)      1.0 [+ or -] 0.1 (a)
                       13.0 [+ or -] 0.3 (b)      1.0 [+ or -] 0.1 (a)
                       13.2 [+ or -] 0.1 (b)      1.4 [+ or -] 0.1 (b)
                       15.7 [+ or -] 0.1 (c)      2.3 [+ or -] 0.1 (c)
                       16.4 [+ or -] 0.1 (d)      2.8 [+ or -] 0.2 (d)

        IBW                      Ash                  Gross energy

 3.0  [+ or -] 0.2      3.2 [+ or -] 0.1 (a)     4.4 [+ or -] 0.1 (c)
                        3.1 [+ or -] 0.1 (a)     1.9 [+ or -] 0.1 (a)
                        3.4 [+ or -] 0.1 (a)     1.9 [+ or -] 0.1 (a)
                        3.1 [+ or -] 0.1 (a)     2.9 [+ or -] 0.1 (b)
                        3.1 [+ or -] 0.3 (a)     4.0 [+ or -] 0.4 (c)
                        3.0 [+ or -] 0.1 (a)     4.5 [+ or -] 0.2 (c)
  8.0 [+ or -] 0.5      3.3 [+ or -] 0.1 (a)     4.0 [+ or -] 0.1 (cd)
                        2.9 [+ or -] 0.1 (b)     2.7 [+ or -] 0.1 (a)
                        3.3 [+ or -] 0.1 (a)     3.3 [+ or -] 0.1 (b)
                        2.5 [+ or -] 0.1 (d)     3.8 [+ or -] 0.1 (c)
                        3.6 [+ or -] 0.1 (e)     4.4 [+ or -] 0.1 (d)
                        2.9 [+ or -] 0.1 (e)     4.7 [+ or -] 0.1 (e)
14.2  [+ or -]  0.09    3.1 [+ or -] 0.1 (ab)    4.5 [+ or -] 0.1 (e)
                        3.2 [+ or -] 0.1 (b)     2.7 [+ or -] 0.1 (a)
                        2.6 [+ or -] 0.1 (a)     3.2 [+ or -] 0.1 (b)
                        3.0 [+ or -] 0.1 (ab)    4.0 [+ or -] 0.1 (c)
                        3.2 [+ or -] 0.4 (b)     4.3 [+ or -] 0.1 (d)
                        3.4 [+ or -] 0.2 (b)     4.6 [+ or -] 0.1 (e)
 19.0 [+ or -] 1.4      3.0 [+ or -] 0.1 (c)     4.6 [+ or -] 0.1 (d)
                        3.1 [+ or -] 0.1 (c)     2.4 [+ or -] 0.1 (a)
                        3.2 [+ or -] 0.1 (d)     3.5 [+ or -] 0.1 (b)
                        2.7 [+ or -] 0.1 (a)     3.8 [+ or -] 0.1 (e)
                        2.9 [+ or -] 0.1 (b)     4.5 [+ or -] 0.1 (d)
                        3.0 [+ or -] 0.1 (c)     5.1 [+ or -] 0.1 (e)

IBW. Initial body weight (g): FL. Feeding level (mg/g shrimp/day).

Values without same letters in the same parameter of the same Body wet
weight were significantly different from each other.

The superscript letters (a) through (e) in this table were markers of
differences of values.

TABLE 6.

The relationship between maintainable feeding level (MFL, mg/g
shrimp/day) and initial body weight (IBW).

Model                                          Terms of IBW

                                     Wet      Dry
MFL = [A.sub.0] + [A.sub.1] x IBW   weight   weight   Protein   Energy

Parameters              [A.sub.0]    11.5     18.3     19.0      19.5
                        [A.sub.1]    -0.05    -0.36    -0.42     -0.45

TABLE 7.

Maintainable energy intake (MIE, J/g BW/d) for shrimp.

                                      MIE

       IBW           Wet weight    Dry weight    Protein    Energy

 3.0 [+ or -] 0.2       218.3         331.1       341.1      349.0
 8.0 [+ or -] 0.5       213.5         296.5       300.8      305.8
14.2 [+ or -] 0.9       207.5         253.6       250.7      252.1
19.0 [+ or -] 1.4       202.9         220.4       220.4      210.6

Wet weight, dry weight, protein, and energy means maintain zero wet
weight, dry weight, protein, and energy growth or loss.

ACKNOWLEDGMENT 

This work was supported by funds from the Project under the Major State Basic Research of China (Grant no. G1999012011) and the State Agriculture Program (Grant no. K2002-16).

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GUOQIANG HUANG, SHUANGLIN DONG * AND FANG WANG

* Corresponding author. E-mail: [e-mail]

Mariculture Research laboratory, Fisheries College, Ocean University of China, Qingdao 266003, People's Republic of China