JOURNAL OF THE WORLD AQUACULTURE SOCIETY

Vol. 27, No. 1 March, 1996

Effect of Dietary Protein Level on Apparent Heat Increment and Post-Prandial Nitrogen Excretion of Penueus setiferus, P. schmitti,

P. duorarum, and P. notialis Postlarvae

CARLOS ROSAS' AND ADOLFO SANCHEZ Laboratorio de Ecofriolgia. Departmento de Biologia. Facultad de Ciencias,

Universidad Nacional de Autonoma de Mexico (UNAM). Mexico 0451 0 D.F., Mexico

EUGENIO DIAZ Centro de Investigaciones Marinas, Universidad de la Habana, Habana. Cuba

LUIS A. SOTO Laboratorio de Ecologia del Bentos. Instituto de Ciencias del Mar y Limnologia,

UNAM, Mexico 04510 D.F., Mexico

GABRIELA GAXIOLA Laboratorio de Ecofriologia, UNAM, Mexico 0451 0 D.F.. Mexico

ROBERTO BRITO Centro de Investigaciones Marinas. Universidad de la Habana. Habana, Cuba

Abstract The calorigenic effect of feeding (apparent heat increment, AHI) and post-prandial nitrogen

excretion (PPNE) were measured in postlarval (PL 25-30) Penaeus setiferus, P. schmitti, P. duor- arum and P. notialis fed a fixed ration of 3 mg/animal using purified diets with 40, 50,60, or 65% protein. Both AH1 and PPNE increased with increasing dietary protein. The contribution of PPNE to AH1 varied from 6.1 to 94%, with lesser values for P. setiferus and greater ones for P. duorarum. Also, the AH1 coefficient (percentage of ingested energy) increased with increasing dietary protein. The AH1 and PPNE coefficients for the four shrimp species ranged from 0.3 to 6.5% and 0.02 to 5.04% of ingested energy, respectively. These results suggest close relationships among protein requirements, the capacity to use dietary proteins as a source of energy, and adaptation by different species to different types of food. The amount of energy used for production of ammonia is proposed as an adequate measure of the part played by dietary proteins in food cost.

Apparent heat increment (AHI), also et al. 1992). Beamish and Trippel (1990) showed that AH1 reflects the metabolism of proteins, carbohydrates, and lipids; and that the deamination and synthesis of proteins are the greatest contributing factors to AHI.

In marine crustaceans, ammonia-nitro- gen constitutes between 60 and 100°/o of the final products of metabolism of proteins (Regnault 1979, 1986). Amino acids in ex- cess of those needed for growth and main- tenance, as well as those derived from nu- tritionally incomplete diets, are degraded to ammonia (Campbell 199 1). Due to the high energy cost of metabolizing amino acids (Hochachka 199 l), protein metabolism and

I Corresponding author. the consequent production of ammonia

known as the specific dynamic action, has been associated with the calorigenic effect of food. It is a measure of the metabolic work of the post-absorptive processes that follow the ingestion of food (Beamish and McMahon 1988). The AH1 in crustaceans and fish depends on the quantity, quality, and balance of the energy components of the diet (Hamada and Ida 1973; Beamish 1974; Hamada and Maeda 1983; Hewitt and Irving 1990; Du-Preez et al. 1992; Koshio

Q Copyright by the World Aquaculture Society 1996

92

NUTRITIONAL PHYSIOLOGY OF PENAE US POSTLARVAE 93

TABLE I . Composition of test diets.

Crude protein (96 dry weight)

40 50 60 65

Ingredients (Yo dry weight) Casein L-Arginine HCI Dextrin Cod liver oil Sunflower seed oil Soybean lecithin Cholesterol Vitamin and mineral mixa Carboxymethyl cellulose

Digestible energy (DE: kcal/g diet)b Protein/DE (g protein/kcal) Lipid (Yo dry weight) g Carbohydratelg lipid

43.3 1.1

37.9 2.3 2.3 1 .o 0.5 2.5 5.0

3.83 10.40 6.5 5.83

54.1 1.4

26.8 2.3 2.3 1 .o 0.5 2.5 5.0

3.83 13.05 6.5 4.12

64.9 1.7

15.7 2.3 2.3 1 .o 0.5 2.5 5.0

3.83 15.70 6.5 2.41

70.4 1.8

10.1 2.3 2.3 1 .o 0.5 2.5 5.0

3.83 16.97 6.5 1.55

~~ ~

a Purina de Mexico, Mexico City, Mexico. Calculated from digestible energy values provided by Nose (1979).

should make considerable contributions to the AH1 of a particular diet. Reduced post- prandial nitrogen excretion (PPNE) asso- ciated with a diet that ensures maximum growth would be beneficial to shrimp cul- ture.

Penaeus setiferus, P. schmitti, P. duora- rum and P. notialis are of interest for mari- culture (Hopkins et al. 1993; Sandifer et al. 1993). They inhabit the coasts of the Gulf of Mexico and the Caribbean Sea (Williams 1984) and two species, P. setiferus and P. schmitti, are widely cultured in those regions (Weidner and Wells 1992; Weidner et al. 1992). This study evaluates the AH1 and PPNE of the postlarvae of these four species of penaeids to define the metabolic costs related to the processing of purified diets with different levels of protein and to es- tablish the role of nitrogen metabolism in the AHI.

Materials and Methods Randomly selected postlarvae of P. seti-

ferus, P. schmitti, P. notialis, and P. duor- arum were reared under controlled labora- tory conditions to PL15 and then trans- ferred to plastic tanks for a 1 O-d acclimation period. Penaeus setiferus and P. duorarum

were transferred to 30-L plastic tanks and held at densities of 1.6 shrimp/L. P. schmitti and P. notialis were moved to 50-L tanks and held at densities of 5 shrimp/L. During acclimation, water temperature was 28 k 2 C, salinity was 36 f 2 ppt, the pH was 8.5 f 0.1, and dissolved oxygen was main- tained above 5 mg/L by aeration. Shrimp were given free access to feed (55% protein) twice a day. To maintain water quality, 50% of the water in the tanks was replaced twice a day during the 10-d acclimation period.

Once the 1 O-d period of acclimation was over, 200 PL25 of P. setiferus (mean live weight f SD = 23.4 k 2.3 mdanimal), P. schmitti (28.2 f 1.2 mglanimal), P. duor- arum (31.3 f 2.3 mdanimal), and P. no- tialis (27.3 f 3.3 mglanimal) were separat- ed into four groups, each of which was as- signed a purified diet with a different level of protein (40, 50, 60, and 65% protein; Table 1). Casein, with 90% protein, was used as the main source of protein in the purified diets. L-Arginine HC1, was added to the ca- sein to improve the amino acid balance and to approximate the balance to the amino acid composition of penaeid abdominal muscle (Teshima et al. 1986). Diets were prepared by thoroughly mixing the dry in-

94 ROSAS ET AL.

gredients with oil and then adding water until a stiff dough was formed. The dough was then passed through a meat grinder to form 5-mm diameter pellets. The pellets were then dried at 60 C in an electric oven. After drying, the feeds were broken up, sieved to obtain a convenient pellet size, and stored at -4 C. Groups of shrimp were given free access to the experimental diets twice a day for a conditioning period of not less than 5 d. Water quality during the con- ditioning period was the same as during ac- climation.

Oxygen Consumption Oxygen consumption of individual

shrimp in the different groups was used to evaluate the AH1 associated with each diet (Medland and Beamish 1985). Closed res- piration chambers of 50 mL were used to hold individual shrimp. The chambers were placed in a heat-regulated bath and kept at 28 k 0.2 C. Postlarvae unfed for 24 h were acclimated to the chambers for 60 min to reduce the effects of handling. Three oxygen consumption measurements made with 10 shrimp each demonstrated that this period of food deprivation and acclimation was suf- ficient to obtain a low and uniform metab- olism.

The water (36 ppt salinity) was carefully changed after acclimation, a first sample of water in the chambers was taken, and the chambers were then closed. This was ac- complished by installing an input and out- put siphon (2 mm diameter) in each cham- ber in such a way as to ensure that the vol- ume of the chamber remained constant. Ox- ygen consumption of the unfed shrimp was calculated from the difference between the dissolved oxygen concentration before clo- sure of the chambers and the dissolved ox- ygen concentration after 2.5 h. Dissolved oxygen concentrations were measured to within 0.01 mg/L with a digital meter (YSI 54 ARC, Yellow Springs Instruments, Yel- low Springs, Ohio, USA) with a polaro- graphic sensor that was calibrated with air- saturated sea water. The results were cor-

rected with data obtained from two control chambers containing no shrimp.

Shrimp in the chambers were fed the test diets at 3 mg/animal (a mean of 10% of body weight) after metabolism of unfed animals was determined. The same amount of food was placed in two control chambers to es- timate the oxygen loss due to food decom- position. In most cases, the dissolved oxy- gen consumption in the control chambers was negligible. The process described above to determine metabolism of unfed animals was used to measure the oxygen consump- tion in fed animals. Preliminary experi- ments demonstrated that highest oxygen consumption occurred during the first 3 h after feeding. In most cases, the shrimp in- gested all the presented food within 30 min. Data obtained from animals that did not ingest all the food offered were not used. The respirometric chambers were covered to eliminate possible interference by activ- ity in the laboratory. Disturbances were limited to those necessary to observe feed- ing activity. All measurements were con- ducted during daylight between 1000 and 1600 h.

The AH1 was calculated from the differ- ence between the oxygen consumption of the unfed and fed shrimp. Following the trials, the shrimp were killed, dried at 60 C, and weighed. The conversion factor of 3.53 cal/mg O2 consumed was used to convert the data to cal/g dry weight per h (Brody 1945).

Nitrogen Excretion Nitrogen excretion of unfed and fed an-

imals was measured to assess PPNE. Water samples were taken from the chambers be- fore and after closing, and the concentration of total ammonia-nitrogen (TAN) was mea- sured. Ammonia was measured with a gas- sensing ammonia electrode connected to an Orion Model 720 ion multianalyzer (Orion Research, Cambridge, Massachusetts, USA). The conversion factor of 5.94 cal/mg NH3-N excreted was used to express the results as

NUTRITIONAL PHYSIOLOGY OF PENAEUS POSTLARVAE 95

cal/g dry weight per h (Brafield and Solo- mon 1972).

To determine the contribution of nitro- gen metabolism to AHI, the PPNE/AHI ra- tio was calculated. This ratio was expressed as a percentage and was obtained for each individual trial. The AH1 and PPNE coef- ficients were calculated as percentage of in- gested energy (Chakraborty et al. 1992; Ross et al. 1992):

AH1 and PPNE (To) = AH1 or PPNE/energy content of

food ingested,

where AHI, PPNE, and energy of food in- gested were expressed in cal/g dry weight.

The Kruskal-Wallis non-parametric test was used to detect significant differences be- tween each experimental group for oxygen consumption and nitrogen excretion. Val- ues for PPNE/AHI, O/o AHI, and O/o PPNE were arcsin-square root transformed prior to analysis (Zar 1974).

Results Oxygen consumption by postlarvae of the

four shrimp species was significantly affect- ed by diet. In general terms, a similar result was observed for nitrogen excretion. For all four species, unfed animals showed a low and uniform metabolism. Once food was provided, the animals displayed intensive muscular activity (pleopod motion) that contrasted with the tranquil behaviour ob- served for shrimp in the respirometric chambers during the 60-min acclimatiza- tion period. As the first three pairs of per- iopods secured the food materials, food was fragmented and passed into the mouth parts for ingestion. At that time, movement of the pleopods stopped. Shrimp in respiro- metric chambers ingested the food that was offered within 30 min. Once food was in- gested, shrimp were relatively inactive for the rest of the 2-h test period.

The metabolic rate of unfed P. setiferus was between 2 1.5 and 83.9 cal/g dry weight per h and varied with protein level in the conditioning diet (Table 2; P < 0.05). The

metabolic rate of fed animals increased by 1.4 to 2.5 times that of the unfed animals. The AH1 was clearly related to protein level (Table 2). Nitrogen metabolism among unfed and fed P. setiferus was similar to respiratory metabolism, with the lowest lev- els obtained for shrimp fed diets with 40 and 50% protein and the highest levels at 60 and 65% protein (Table 2). Food inges- tion resulted in an increase in nitrogen ex- cretion rates between 1.5 and 2.5 times that of the unfed animals. The PPNE increased from 0.6 to 18.9 cal/g dry weight per h, corresponding to increased protein in the diet (Table 2; P < 0.05).

In P. schmitti, the oxygen consumption rate of unfed animals was between 19.1 and 34.2 cal/g dry weight per h and differed with protein level in the conditioning diet (Table 2; P < 0.05). The maximum metabolic rate of fed animals was 2.6 to 3.6 times that of the unfed animals. The AH1 was related to protein level, with values of 29.8 cal/g dry weight per h for animals fed with 40% pro- tein to 68.2 cal/g dry weight per h in shrimp fed with 65% protein (Table 2). Nitrogen metabolism in unfed P. schmitti was lowest for shrimp fed conditioning diets with 40 and 60% protein. Feeding resulted in an in- crease in nitrogen excretion of between 1.7 and 4.2 times that of the unfed animals. The PPNE increased from 6.8 to 32.9 cal/g dry weight per h, corresponding to increasing protein in the diet between 40 to 60% pro- tein. The PPNE in shrimps fed with 65% protein was 12.2 caVg dry weight per h (Ta- ble 2).

The metabolic rate of unfed P. duorarum was highest for shrimp fed conditioning di- ets with 40,50 and 65% protein (an average of 39.9 cal/g dry weight per h; Table 2). Shrimp fed the conditioning diet with 60% protein diet had a lower value of 19 cal/g dry weight per h. Feeding produced an in- crease in metabolic rate between 2.4 and 9.6 times that for unfed animals in this spe- cies, with the lowest values obtained for an- imals fed 40% protein (88.9 cal/g dry weight per h) and the highest for animals fed 65%

96 ROSAS ET AL

TABLE 2. Mean (f SEM) oxygen consumption. apparent heat increment (AHI). nitrogen excretion, and post- prandial nitrogen excretion of postlarvae of four species of shrimp. AN values are expressed as callg dry weight per h. Significant diferences existed among protein levels for all variables measured for all species (Kmkal- Wallis Test; P < 0.05).

Rate Pro- tein

Variable (96) N Unfed Fed AH1 PPNE

Penaeus setifem Oxygen consumption

Nitrogen excretion

Penaeus schmitti Oxygen consumption

Nitrogen excretion

Penaeus duorarum Oxygen consumption

Nitrogen excretion

Penaeus notiatis Oxygen consumption

Nitrogen excretion

40 50 60 65 40 50 60 65

40 50 60 65 40 50 60 65

40 50 60 65 40 50 60 65

40 50 60 65 40 50 60 65

8 8 8 7 8 8 8 7

20 18 19 19 20 18 19 18

10 10 10 6

10 10 9 6

20 19 19 20 20 19 18 19

23.2 f 1.4 21.5 f 2.8 83.9 f 7.9 61.2 f 6.4

1.2 * 0.1 1.3 f 0.1

10.7 f 1.0 15.1 f 2.2

19.1 f 2.8 23.7 f 2.2 24.1 f 1.9 34.2 f 6.6

8.6 f 0.7 14.0 f 3.6 10.2 f 1.3 16.4 f 3.3

36.5 f 3.9 42.8 f 7.6 19.0 f 2.9 31.5 f 5.5 19.9 f 1.6 23.4 f 2.6 15.3 f 1.2 15.2 f 1.3

28.1 f 1.7 52.7 f 5.5 58.0 f 2.6 66.6 f 4.9 18.2 * 4.5 48.7 f 3.8 25.1 f 2.4 51.6 f 4.6

33.1 f 2.9 45.3 f 4.6

181.5 f 17.6 153.8 f 22.8

1.8 f 0.1 2.4 f 0.1

20.4 f 0.6 34.0 f 1.2

48.9 f 3.4 67.3 f 7.9 85.5 f 4.5

102.4 f 2.8 15.4 f 1.0 24.2 f 1.7 43.1 f 3.6 28.6 f 2.9

88.9 f 6.9 123.9 f 4.4 182.7 f 3.1 202.3 f 7.7

53.4 f 5.2 90.6 f 11.2

158.7 f 12.4 175.8 f 5.6

101.4 f 3.2 164.6 f 7.9 251.3 t 7.9 312.4 f 8.6 64.0 _+ 5.6

120.2 f 6.7 162.5 f 9.6 244.3 f 6.3

9.9 f 1.4 23.8 k 5.7 97.6 f 97.6 92.6 f 10.2 -

- - -

29.8 f 1.9 43.7 f 1.9 61.4 f 4.5 68.2 f 2.8

- - - -

52.4 f 4.9 81.1 f 7.7

163.7 ? 3.9 170.8 f 7.7

- - - -

73.3 f 3.5 119.9 f 5.9 193.3 ? 8.3 246.8 f 6.3

- - - -

- - - -

0.6 f 0.1 1.1 f 0.1 9.7 f 1.5

18.9 f 3.5

- - - -

6.8 f 0.7 10.2 f 2.0 32.9 5 3.1 12.2 f 2.7

- - - -

33.5 * 5.2 67.2 f 7.6

143.4 f 14.2 160.7 f 16.0

- - - -

45.8 2 3.4 71.5 f 5.3

137.4 f 10.4 193.1 f 9.9

protein (202.3 cal/g dry weight per h; Table 2). The AH1 increased with increasing di- etary protein, from 52.4 to 170.8 cal/g dry weight per h for 40 and 65% protein, re- spectively (P < 0.05). Nitrogen metabolism of unfed P. duorarurn was lowest for shrimp fed conditioning diets of 40, 60 and 65% protein (an average value of 16.8 cal/g dry

weight per h) but was higher (23.4 cal/g dry weight per h) for shrimp fed 50% protein. Feeding caused an increase in nitrogen ex- cretion rate related to the increase in the protein of the diet (Table 2). Maximum ni- trogen excretion rates for fed animals were 2.7 to 11.5 times greater than those ob- served among unfed animals. Similarly, the

NUTRITIONAL PHYSIOLOGY OF PENAEUS POSTLARVAE 97

TABLE 3. Efects of dietary protein level on apparent heat increment coeflcient (AHIlenergy content of.food ingested in cal per g dry weight) and post-prandial nitrogen excretion coemient (PPNElenergy content of food ingested in callg dry weight) for four species of shrimp. All values are mean f SEM. Significant diferences existed among protein levels for both coeflcients for all four species.

Dietary protein (Yo) Species 40 50 60 65

AH1 Coefficient Penaeus setiferus 0.3 f 0.02 0.6 f 0.07 2.5 f 0.31 2.4 f 0.37 Penaeus schmitti 0.8 f 0.08 1.1 f 0.18 1.6 f 0.13 1.8 f 0.18 Penaeus dourarum 1.4 f 0.14 2.1 f 0.20 4.3 f 0.53 4.5 2 0.31 Penaeus notiafis 1.9 f 0.13 2.9 f 0.43 5 . 5 & 0.44 6.4 f 0.93

PPNE Coefficient Penaeus setiferus 0.02 * 0.002 0.03 f 0.005 0.03 & 0.001 0.49 f 0.030 Penaeus schmitti 0.17 f 0.007 0.27 f 0.017 0.86 f 0.009 0.32 f 0.008 Penaeus dourarum 0.87 f 0.004 1.75 f 0.020 3.74 f 0.042 4.25 f 0.026 Penaeus notialis 1.20 f 0.016 1.87 f 0.053 3.95 ? 0.054 5.04 f 0.190

PPNE was highest for animals fed diets with 60 and 65% protein compared to those ob- served at 40 and 50% protein (Table 2). The maximum average value obtained with 60 and 65% protein (152 cal/g dry weight per h) was 4.5 and 2.26 times greater than those obtained with 40 and 50% protein, respec- tively.

The metabolic rate of unfed P. notialis increased with protein in the diet, from 28.1 cal/g dry weight per h to 66.6 cal/g dry weight per h (Table 2; P < 0.05). The metabolic rate of the fed animals increased to between 3.1 and 4.7 times that of the unfed animals. The metabolic rate of shrimp fed 65% pro- tein was 2.37 times greater than that for shrimp fed 40% protein. The AH1 was clear- ly related to protein level with lowest values with shrimps fed with 40% protein and highest values with shrimps fed with 65% protein (P < 0.05). The pattern of nitrogen metabolism in unfed P. notialis was not consistent with protein level in the condi- tioning diet. High values were obtained with 50 and 65% protein and low ones with 40 and 60% protein (Table 2). In fed animals, nitrogen excretion increased 3.5, 2.5, 6.5, and 4.7 times over that for unfed animals for 40,50,60 and 65% protein, respectively. The PPNE increased significantly (P < 0.05) as dietary protein increased from 40% (45.8

cal/g dry weight per h) to 65% ( 1 93.14 cal/g dry weight per h).

The AH1 and PPNE coefficients of the four species were found to be a small pro- portion of the ingested energy (Table 3). Highest values of AH1 coefficients were for P. duorarum and P. notialis fed the diet with 65% protein (5 and 6.4%, respectively) and the lowest were for P. setiferus and P. schmitti fed diets with 40% protein (0.3 and O.6%, respectively). The highest values for PPNE coefficients were for P. duorarum and P. notialis postlarvae fed 65% protein diets (4.2 and 5.04%, respectively) and the lowest for P. setiferus and P. schmitti fed 40% protein diets (0.02 and 0.17%, respectively). For all four species, AH1 and PPNE coefficients varied with the protein of the diet, being lowest for animals fed with 40% protein di- ets and highest for animals fed the 65% pro- tein diet.

The AH1 and PPNE were higher for P. duorarum and P. notialis than for P. seti- ferus and P. schrnitti (Figs. 1,2). The PPNE/ AH1 ratio (Fig. 3) for P. setiferus was be- tween 6 to 9% among diets of 40 and 60% protein, with an increase at 65% protein. The PPNEIAHI ratios for P. schmitti were similar for animals fed 40, 50, or 65% pro- tein diets (mean = 23.l%), but was higher (mean = 53.6%) for animals fed the 60%

98 ROSAS ET AL.

SPECIES FIGURE 1. Apparent heat increment (AHI, caNg dry weight per h) for four species of penaeid shrimp fed diets

with four dietary protein levels.

protein diet. The highest PPNE/AHI ratios were observed in P. duorarum with values between 64% (40% protein) and 94% (65% protein). In P. notialis the PPNE/AHI ratio varied between 62 and 78%, and were in- termediate to values for P. setiferus, P. schmitti, and P. duorarum.

Discussion The four species of shrimp showed an

increase in AH1 related to the level of pro- tein in the diet. This pattern has also been reported for Macrobrachium rosenbergii (Clifford and Brick 1983) and tilapia and carp (Chakraborty et al. 1992; Ross et al. 1992). However, changes in the AH1 value relative to changes in dietary protein will be related to the adaptative capacity of each species to use protein as a metabolic sub- strate. Evaluations made with the oxygen: nitrogen (0:N) ratio of various species have

demonstrated striking variation in the ca- pability of crustaceans to catabolize dietary protein. Thus, it has been reported that the metabolism of carbohydrates and lipids dominates in the omnivorous crustaceans Macrobrachium rosenbergii and Palaemo- netesvarians(Snowand Williams 197 1; Clif- ford and Brick 1983), while the metabolism of protein is dominant in the carnivorous crustaceans Penaeus esculentus, Homarus americanus, and Crangon crangon (Capuz- zo and Lancaster 1979; Regnault 1979; Dall and Smith 1986; Hewitt and Irving 1990; Koshio et al. 1992). Recently, Rosas et al. (1995) have pointed out that there is an inverse relationship between the O:N ratio and the requirement of proteins of some Penaeus species. In that study the greater values of O:N were obtained in P. setiferus postlarvae (40% protein requirement) and lower values in P. schmitti postlarvae, a spe-

NUTRITIONAL PHYSIOLOGY OF PENAEUS POSTLARVAE 99

W-

a a z

SPECIES FIGURE 2. Post-prandial nitrogen excretion (PPNE, callg dry weight per h) for four species of penaeid shrimp

fed diets with four protein levels.

cies with a higher protein requirement of 60% (Gaxiola 1994). Although both P. se- tiferus and P. schmitti had similar values of AH1 and PPNE, the O:N results suggest that each species has a different strategy to use energy from food. The AHI, PPNE, and O:N data indicate that in P. setiferus, amino ac- ids are used more for growth than metabolic substrate, whereas in P. schmitti the amino acids are used both for growth and meta- bolic substrate.

With regard to protein requirement of the four shrimp species studied in the present study, Garcia and Galindo (1 990), Gaxiola et al. (in press), and Gaxiola (1994) have reported a protein requirement for growth of 40, 50 and 60% for P, setiferus, P. duor- arum, and P. schmitti postlarvae, respec- tively. Peneaus setiyerus appears to be better adapted to process diets with a low protein

120/ loot

60 w z n

40

20

0

I? duorarum T

: & I? notialis

-

P. setiferw

1 I

40 45 50 55 60 65

% DIETARY PROTEIN

FIGURE 3. Contribution of post-prandial nitrogen ex- cretion (PPNE) to apparent heat increment (AHI) for four species of penaeid shrimp fed diets with four protein levels.

100 ROSAS ET AL.

level than P. duorarum or P, schmitti which have a greater protein requirement.

According to Chakraborty et al. (1992) and Ross et al. ( 1 992), the AH1 coefficient may be used as an indicator of efficiency of transformation of digestible energy. In the present study the PPNE coefficient was also used. The AH1 coefficients ranged from 0.3 to 6.4% and the PPNE coefficients from 0.02 to 5.04%, and increased with the protein content of the diet in all four species (Table 3). In fish, AH1 coefficients change in rela- tion to protein levels in the diet (Beamish and Trippel 1990) confirming that the effi- ciency of transformation of digestible en- ergy to growth is reduced when protein ca- tabolism increases. The balance between energy cost and protein requirement for growth is thus the key to obtaining an effi- cient diet.

The role of nitrogen metabolism in the overall metabolism of shrimps and fish is a key factor in the AHI. According to Beam- ish and Trippel ( 1 990), even when the me- tabolism of protein, lipids, and carbohy- drates together contributes to the AHI, de- amination and synthesis of protein are probably the greatest contributors. Produc- tion of ammonia results mainly from the catabolism of amino acids of both alimen- tary and metabolic origin. Ammonia ac- counts for 60 to 100% of the products of nitrogen metabolism and its formation may result directly from deamination or trans- amination. The formation of glucose through aminoacidic gluconeogenesis is the most common way of forming ammonia (Campbell 199 1).

In this study, calculated values for PPNE showed distinct differences between the four species of shrimp. In view of the levels of PPNE shown in Fig. 2 and Table 3, two groups of species can be formed: one with P. setiferus and P. schmitti (low ammonia production and low PPNE coefficients) and another with P. duorarum and P. notialis (high ammonia production and high PPNE coefficients). It can be seen in Fig. 3 that the role played by PPNE in AH1 is also greater

in P. duorarum and P. notialis than in P. setiferus and P. schmitti, which suggests the possibility of a metabolism differently adapted to the use of sources of energy of proteic origin. Thus, the highest percentages could signify that P. duorarum and P. no- tialis make greater use of protein to obtain metabolic energy as opposed to P. setijerus and P. schmitti. The tendency towards more carnivorous habits in P. duorarum than in P. setiferus (Guitart and Hondares 1980) or the preference for an animal protein:plant protein ratio of 38:62 in P. schmitti(Gaxio1a et al., in press) might be indirect evidence to support such a hypothesis.

Acknowledgments The organisms used in this study were

provided partly by the Ministerio de la In- dustria Pesquera de Cuba (P. schmitti and P. notialis) and by the Centro Regional de Investigaciones Pesqueras de Campeche (P. duorarum). We are grateful for the partici- pation of the personnel of the Centro de Investigaciones Marinas of the Universidad de la Habana. This project was developed with funds provided by the Direccion Gen- eral de Atencion a1 Personal Academic0 of the UNAM for the project IN-20 1292 given to Luis A. Soto and Carlos Rosas. Further- more, help was received from the Facultad de Biologia of the Universidad de la Ha- bana, from the Instituto Nacional de la Pes- ca through the collaboration agreement with the Facultad de Ciencias of the UNAM, from the Departamento de Nutricion of the Universidad Iberoamericana and from the Third World Academy of Science for the research visit of Dr. E. Diaz to the UNAM during 1992.

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NUTRITIONAL PHYSIOLOGY OF PENAEUS POSTLARVAE 101

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