Aust. J . Mar. Freshwater Res.

, 1981, 32, 245-51

Carbon and Nitrogen Content of Food
and the Assimilation Efficiencies of
Penaeid Prawns in the Gulf of Carpentaria
D. J . W, Moriarty and M. C. Barclay
Division of Fisheries and Oceanography, CSIRO, P.O. Box 120, Cleveland, Qld 4163.
Abstract
The food of seven species of penaeid prawns from the Gulf of Carpentaria consists predominantly of
Foraminifera, small molluscs, crustaceans and polychaetes. Measurements of organic and inorganic carbon,
organic nitrogen and bacterial biomass were made. Foregut contents of adult prawns contained between 72
and 223 mg organic carbonlg dry wt. Protein constituted between 43 and 64% of the organic matter.
Approximate assimilation efficiencies of food in prawns caught in the gulf, determined for four species,
varied from 48 to 77% of organic carbon and from 42 to 77% of organic nitrogen. The food of juvenile
Penaeus merguiensis was examined for two growing seasons. In the 1976-1977 season the foregut contents
contained a mean of 41 mg organic nitrogenig dry wt and 18 1 mg organic car bon~g dry wt. In the 1977-1978
season, significantly lower proportions of organic nitrogen and carbon were eaten, viz, 21 mg organic
nitrogenlg dry wt and 101 mg organic carbon/g dry wt. Improved assay procedures for muramic acid have
shown that bacteria are less important in the food of prawns than previously reported. Bacteria constituted
less than 2% of the organic matter in the adults of all species, but in many juvenile P, merguiensis bacteria
were more important, constituting up to 14% of organic matter.
Introduction
The Gulf of Carpentaria supports an important commercial fishery of penaeid
prawns. A study is being made of the biology and ecology of the major species,
particularly Penaeus merguiensis de Man, and this paper describes the food
components and assimilation efficiencies of these prawns.
Preliminary work indicated that penaeid prawns are omnivores feeding mainly on
live prey, but also eating some micro-organisms including bacteria (Moriarty 1977).
Bacteria were thought to be an important part of their diet, based on estimates of
biomass made by measurement of muramic acid. Recent improvements in the assay
technique for muramic acid (Moriarty 1980) have shown that many of the early results
overestimated bacterial biomass, particularly in samples containing CaCO,. Revised
estimates are published here. It is difficult to determine the food of prawns either
quantitatively or qualitatively because they break much of it up with their mandibles or
gastric mill. Other workers (e.g. Williams 1955; Hall 1962) noted that, in general,
prawns are omnivores and that much of the unidentifiable material in the gut was
probably animal remains. We have combined microscopic observations with a more
objective analysis of carbon and nitrogen in the gut contents of the following species:
P. merguiensis, P. esculentus Haswell, P. semisulcatus de Hann, P. latisulcatus
Kishinouye, P, monodon Fabricius, Metapenaeus endeavouri (Haswell), and M. ensis
(de Haan).
D. J. W. Moriarty and M. C. Barclay
Condrey et al. (1972) have reported assimilation efficiencies of 5547% of organic
matter for two species of Penaeus that were fed in the laboratory on diatoms, bacteria
and compounded diets. We have used the ratios of organic carbon and nitrogen to
inorganic matter in the foregut, midgut and hindgut to estimate assimilation
efficiencies of food of prawns caught in the field.
Materials and Methods
Adult prawns were collected by trawling in south-eastern, eastern and north-eastern areas of the Gulf of
Carpentaria. Juvenile P. merguiensis were caught at low tide with a small beam trawl in the Norman River at
Karumba, northern Queensland (cf. Staples and Vance 1979). All prawns were frozen within half an hour of
capture and brought to the laboratory on dry ice. Foregut contents were dissected out while frozen and were
then freeze-dried. Sample sizes ranged from 17 to 100 jubeniles and from 10 to 30 adult prawns at each
locality and time. They were pooled to provide sufficient material for analysis. To obtain midgut and hindgut
contents, prawn tails were partly thawed and the intestine was removed and immediately dissected. The
contents were frozen and freeze-dried.
A Perkin-Elmer CHN analyser was modified for measurement of organic carbon and nitrogen and
carbonate. The main furnace was operated at 550C and the high-heat zone was extended slightly and
packed with insulation to operate at 1000C. Samples were burnt for 20 min at 550C to give organic carbon
and nitrogen and then comhusted at 1000C to give carbonate (Moriarty and Barclay 1981). The uptake of
water by CaO and MgO after combustion made it difficult to determine ash weights by weighing. We found it
simpler to estimate ash weight by subtraction of the dry weight of organic matter from the original weight.
Organic matter was calculated from the organic carbon values, assuming that these were 45% of organic
matter. These calculations were checked by carefully weighing some ash samples; the results agreed with the
estimated ash content. Organic carbon and nitrogen values are expressed in terms of dry weight and ash
weight.
Assimilation efficiencies (U) in the digestive gland were calculated from the ratios of organic carbon and
nitrogen to ash weight as follows:
U = [(CJA,) -(C2/A2)]/(C,IA,) x 100,
where C,/A, is the ratio of organic carbon to ash in the foregut and C21A2 is the ratio of organic carbon to ash
in the midgut or hindgut. Assimilation efficiencies of organic nitrogen were calculated similarly. The results
that were obtained were the same as those obtained by the more complicated equation for assimilation of a
dietary component given by Condrey et al. (1972). As it is obviously impossible to obtain a representative
sample of the food of prawns in their natural habitat before they eat, analysis of foregut contents is the only
practicable method available. Animals selected were those that had full guts.
Chitin was estimated approximately by extracting samples with 1 M NaOH at 100'C for 24 h and
comparing organic nitrogen with untreated samples. Chitin obtained from B.D.H., Melbourne, Vic., was
treated similarly as a control. Muramic acid was estimated and bacteria examined microscopically as
described by Moriarty (1980). Foregut contents were examined under the microscope and the predominant
food items were noted.
Results
Food Analyses in Adult Prawns
Foraminifera tests were the most common items observed in the gut contents of
many adult prawns. Fragments of small crustaceans, molluscs and polychaetes were
often present. Much of the CaCO, in the gut contents was due to Foraminifera tests,
particularly in P. esculentus and P. semisulcatus which fed mainly on Foraminifera. In
a few cases, molluscan shells predominated. Organic carbon constituted generally
10-20% of the diet and organic nitrogen about 2 4 % (Table 1). The carbon to nitrogen
ratios ranged from 4. 4 to 6. 7, which indicates that between 43 and 64% of the organic
matter was protein (assuming that protein = N x 6.25). About 40% of the nitrogen
was extracted by NaOH from control samples of chitin, whereas 93-97% was extracted
by NaOH from prawn gut contents. The residual 3-7% of nitrogen in the gut contents,
Food Components and Assimilation Efficiencies of Penaeid Prawns
when multiplied by the factor of 40% for nitrogen extracted from chitin, gives a value
of about 5-10% for nitrogen present in chitin or other nitrogenous material insoluble in
NaOH. No seasonal or geographical trends were apparent in the nature of the food
within species. Under the microscope, more particles of sediment with attached
bacteria and less Foraminifera and molluscan fragments were observed in the food of
the Metapenaeus species than in the Penaeus species. Values for organic nitrogen and
carbon were higher and for CaCO, lower in animals that had eaten few Foraminifera
and molluscs.
Table 1. Analysis of food of adult penaeid prawns from the Gulf of Carpentaria
Each sample consisted of the pooled gut contents of 10-30 individuals. Food types are listed in order of
abundance. Concentrations are given as mgig dry wt
Species Date LocalityA FoodB Organic Organic CaCO,
nitrogen carbon concn
concn concn
P. rnerguiensis 20.x.77
10.xi.77
P. esculentus 20.x.77
20.xi.77
6.xi.77
4.xi.77
P. rnonodon 3.iv.78
29.iii.78
20.x.77
10.xi.77
P. latisulcatus 17.x.77
5.i.78
2.xi.77
P. sernisulcatus 6.xi.77
I1 . xi77
17.x.77
M. endeavouri 15.xi.77
6.xi.77
8.x.77
M. ensis 6.xi.77
12.x.77
l.xi.77
A S, south-eastern gulf; M, midway along east coast; N, north-eastern gulf.
F, Foraminifera; P, polychaetes; M, molluscs; C, crustaceans; S, sand: D, detritus.
Approximate assimilation efficiencies, calculated for four species, show that for
most animals over 50% of the organic matter was assimilated (Table 2). Most
assimilation occurred between the foregut and midgut. In the P. merguiensis sample
there was no further nitrogen assimilation, but in the other species a further 8-14% of
organic nitrogen and organic carbon was assimilated. The efficiency of assimilation in
P. esculentus was lower than 50%. These animals had fed mainly on Foraminifera,
many of which were not broken up. The other species had fed principally on
crustaceans or polychaetes and molluscs. There was little alteration in the carbon to
nitrogen ratio as food material moved through the gut, indicating that the efficiency of
digestion and assimilation of protein was similar to that of other digestible organic
material.
D. J. W. Moriarty and M. C. Barclay
Food Analysis of Juvenile Prawns
Particles of sediment with bacteria attached were commonly observed in the foregut
of juvenile P. merguiensis. Fragments of Foraminifera, crustaceans, molluscs and
polychaetes were also observed. The food ofjuveniles in the 1976-77 season had twice
Table 2. Minimum assimilation efficiencies of food in penaeid prawns from the Gulf of Carpentaria
Samples of gut contents from 14-19 individuals with full guts were pooled. Values are means for duplicate or
triplicate analyses. Values for assimilation efficiency between foregut and either midgut or hindgut are
shown. Unit of concentration is mglg dry wt
Species Gut Organic nitrogen Organic carbon Ratio of CaCO,
region Concn % assimi- Concn % assimi- carbon to concn
lation lation nitrogen
P. latisulcatus Fore 64 69 283 68 4. 4 139
Mid 20 77 89 77 4. 6 367
Hind 14 - 64 - 4. 4 331
P. merguiensis Fore 34 54 161 57 4. 7 353
Mid 16 54 69 52 4. 5 327
Hind 16 - 77 - 4. 9 393
P. esculentus Fore 22 34 114 44 5. 2 520
Mid 15 42 64 48 4 . 4 637
Hind 12 - 59 - 4. 8 629
M. endeavouri Fore 56 56 264 60 4 . 7 120
Mid 24 7 1 106 74 4. 4 139
Hind 16 - 68 - 4 . 2 210
the mean concentration of nitrogen as that of those sampled in the 1977-78 season
(Table 3). The concentration of organic carbon was similarly greater. Detritus and
Crustacea were predominant in their gut contents. Foraminifera were more prominent
in the gut contents of the animals caught in the 1977-78 season. The carbon to nitrogen
ratios indicate that protein constituted about 40-76% of the organic matter.
Table 3. Analysis of food of juvenile P. merguiensis
Samples of prawns were collected about once per month in 1976-77 and about once each fortnight
in the 1977-78 season. Carapace length varied from 5 to 20 mm. Each sample consisted of 17-100
prawns. Unit of concentration is mg,g dry wt
Season Organic Organic CaCO, Ratio of No. of
nitrogen carbon concn carbon to samples
concn concn nitrogen
Sept. 1976- Mean 41A 181A 250 4 . 4 13
March 1977 Range 24-62 105-270 80-650 3. 8-5. 0
Dec. 1977- Mean 21A 101 A 410 5. 0 28
March 1978 Range 12-39 58-270 80-750 3. 7-6. 8
A Significantly different P = 0,001, rank test (Sokal and Rohlf 1969).
Of the juveniles dissected, only about 50% had a foregut that was more than half
full. Many had empty guts, although they were collected at low tide which is probably
the time of greatest activity (Staples and Vance 1979). There was no noticeable
difference in the composition of the diet of juvenile prawns in the size range 5-20 mm
carapace length.
Food Components and Assimilation Efficiencies of Penaeid Prawns
Bacteria in the Food
Bacteria were observed microscopically in the foregut contents, usually in
association with particles of sediment. Samples which obviously contained large
amounts were analysed quantitatively and it was found that in juvenile prawns bacteria
constituted 2-14% of the organic matter (Table 4). Juvenile prawns with large amounts
of bacteria generally had lower values for organic carbon and nitrogen. In adult
Metapenaeus spp., 1-3% of organic matter was derived from bacteria. Other samples
of prawns contained less bacteria.
Table 4. Bacteria in the food of penaeid prawns
Mean values are given for duplicate analyses of foregut contents. P, merguiensis samples were juveniles from
the Norman River; the other species were adults from the Gulf of Carpentaria
Species Organic Organic Bacterial CaCO,
nitrogen carbon carbon dry wt)
( W g dry wt) (mg,/g dry wt) (% organic
carbon)
P. merguiensis 15
17
17
18
3 5
3 5
30
M, endeavouri 42
44
44
M. ensls 39
45
45
Discussion
Bacteria are less important in the food of most of the adult prawns than reported
previously (Moriarty 1977). Measurement of trace amounts of muramic acid in
sediment samples, particularly in those containing CaCO,, has proved difficult, but
these problems have been resolved and the method checked against another technique
(Moriarty 1980). The Metapenaeus species did appear to be selecting sediment particles
with bacteria, but the amounts involved (up to 3% of organic matter) are small. These
species were caught in shallow water closer to shore than most adult Penaeus spp., so it
is not clear whether the difference in diet between the two genera of prawns is due to
differences in feeding selectivity. The juvenile P, merguiensis do ingest bacteria,
although bacteria were not the predominant items in their diet.
The main components of the diet of all the prawns examined are meiofauna. As
concluded previously, the low bacterial density and high protein content of their food
indicates that most of these prawns are not feeding on detritus, but rather on living
animals (Moriarty 1977). Bell and Coull (1978) have shown that the shrimp
Palaemonetespugio is an important controller of meiofauna abundance in salt-marsh
environments. It is not unlikely that the penaeid prawns would have a similar effect on
meiofauna. Few trends were noted in the types of food organisms eaten by the various
species in different parts of the Gulf of Carpentaria, but P. esculentus consumed more
D. J. W. Moriarty and M. C. Barclay
Foraminifera than did the other species. In Moreton Bay, P. esculentus also ate mainly
Foraminifera (Moriarty 1977).
The content of protein and organic matter in the food varied over about a fivefold
range. Without information on feeding rates, it is not possible to say whether those
with small amounts were short of food. In the two seasons that were investigated,
juvenile P. merguiensis contained significantly less nitrogen in the 1977-78 growing
period. Staples (1980) has shown that the juveniles in the Norman River grow rapidly
from the postlarval stage to a carapace length of about 10-20 mm over the period
November-February. A point which could be investigated further, therefore, is
whether food might be one of the factors limiting the growth rate of the juveniles and
perhaps also the numbers entering the fishery. White (1978) has pointed out that the
numbers of many insect populations are regulated by the supply of nitrogen to the
juveniles, and as prawns select food with a high nitrogen content, a similar effect might
occur.
It is obviously impossible to accurately measure assimilation efficiencies of prawns
in their natural environment, but an approximation can be obtained from analyses of
foregut and hindgut contents. The values recorded here are minimum estimates. These
values were calculated on the assumption that mineral content remains constant. In
three of the four groups of prawns analysed, CaCO, content increased from foregut to
midgut, and in all groups it increased in the hindgut (Table 2). This suggests that little,
if any, CaCO, was absorbed. Forster and Gabbott (1971) have discussed this problem
in their study of assimilation in two species of carid prawns. They found that 32% of
inorganic salts were apparently assimilated, although they could not rule out
regurgitation as a possible explanation. In their experiments, however, inorganic
matter was only a small percentage of the total and thus a small change would
introduce a large error in calculations, whereas in this study, inorganic matter
constituted 60-88% of the prawns' diet and it is most unlikely that a large proportion
would be assimilated. Thus, errors due to ash assimilation would not be large.
Variation in composition of the diet is another source of error, and we have assumed
that the average composition oker the period of time that food took to pass from
foregut to hindgut was similar for a group of prawns caught in one area.
Another factor which would lead to underestimation of assimilation efficiency is the
digestive process in the foregut. Much of the digestion in decapods probably occurs in
the foregut. Digestive enzymes are secreted dorsally into the foregut and soluble
products are filtered into ventral tubes and then pass into the digestive gland (Powell
1974; Dall and Moriarty 1981). We have minimized this problem by selecting prawns
with full guts (i.e. those which had been feeding actively at the time of capture). Thus,
although digestion would have been proceeding, much of the products would have
remained in the frozen pellet of contents that was removed for analysis. The range of
values recorded here is only a little less than the range of 52-87%, measured by analysis
of food and faeces, for P. setiferus and P. aztecus that were fed on defined diets and an
algal mat (Condrey et al. 1972). We may conclude therefore that the prawns are
assimilating a high proportion of their diet in the field.
In their studies on the assimilation of various components of food by Astacus spp.,
Speck and Urich (1970) found that most assimilation occurred from the foregut, and
only a small amount (about 2 4 %) occurred from the midgut. The results that we have
obtained with penaeid prawns support their observations (Table 2). In three of the
samples, 10% ofthe assimilation occurred in the midgut, and in one (P. merguiensis) no
Food Components and Assimilation Efficiencies of Penaeid Prawns
further food was assimilated in the midgut. These results also agree with Powell's
(1974) model for digestive physiology in decapods, as discussed above.
Acknowledgments
We are grateful to J. A. Redfield, J. P. Salini and T. Wassenberg for supplying the
adult prawns, and to D. J. Staples, D. J. Vance and D. S. Heales for the juvenile
prawns. We thank Drs W. Dall and D. J. Staples for their helpful criticism of the
manuscript.
References
Bell, S. S., and Coull, B. C. (1978). Field evidence that shrimp predation regulates meiofauna. Oecologia
(Berlin) 35, 141-8.
Condrey, R. E., Gosselink, J. G., and Bennett, H. J. (1972). Comparison of the assimilation of different diets
by Penaeus setiferus and P. azrecus. U.S. Fish Wildl. Serv. Fish. Bull. No. 70. pp. 1281-92.
Dall, W., and Moriarty, D. J. W. (1981). Functional aspects of nutrition and digestion. In 'The Biology of
Crustacea'. Vol.,II. (Eds Linda H. Mantel and D. Bliss.) In press. (Academic Press: New York.)
Forster, J. R. M. , and Gabbott, P. A. (1971). The assimilation of nutrients from compounded diets by the
prawns Palaemon serratus and Pandalus platyceros. J. .&r. Biol. Assoc. U.K. 51, 943-61.
Hall, D. hr. F. (1962). Observations on the taxonomy and biology of some Indo-West Pacific Penaeidae
(Crustacea. Decapoda). Fish. Publ. Colonial Off. (London). No. 17. pp. 1-229.
Moriarty, D. J. W. (1977). Quantification of carbon, nitrogen and bacterial biomass in the food of some
penaeid prawns. Aust. J. ,Mar. Freshwater Res. 28, 11 3-1 8.
Moriarty, D. J. W. (1980). Measurement of bacterial biomass in sandy sediments. In 'Biogeochemistry of
Ancient and Modern Environments'. (Eds P. A. Trudinger, M. R. Walter and B. J. Ralph.) pp. 131-8.
(Australian Academy of Science: Canberra.)
Moriarty, D. J. W.. and Barclay, M. C. (1981). Determination of organic carbon and carbonate in the same
sample with an elemental analyser. Lab. Pract. (In press.)
Powell, R. R. (1974). The functional morphology of the foreguts of the thalassinid crustaceans, Callianassa
cal~forniensis and Upogebiapugettensis. Univ. Calif. Berkeley Publ. Zool. No. 102. pp. 1-41.
Staples, D. J., and Vance, D. J. (1979). Effects of changes in catchability on sampling of juvenile and
adolescent banana prawns. Penaeus merguiensis de Man. Aust. J. Mar. Fresh~vater Res. 30, 511-19.
Staples, D. J. (1980). Ecology of juvenile and adolescent banana prawns, Penaeus merguiensis de Man, in a
mangrove estuary and adjacent off-shore area of the Gulf of Carpentaria. 11. Emigration, population
structure and growth of juveniles. Aust. J. Mar. Freshwater Res. 31, 653-66.
Sokal, R. R., and Rohlf, F. J. (1969). 'Biometry.' (W. H. Freeman and Company: San Francisco.)
Speck, U. , and Urich, K. (1970). The metabolic fate of nutrients in the crayfish, Orconectes limosus. 11.
Absorption of U-14C-labelled nutrients and their distribution among the organs. 2. Vgl. Physiol. 68,
318-33.
White, T. C. R. (1978). The importance of a relative shortage of food in animal ecology. Oecologia (Berlin)
33, 71-86.
Williams, A. B. (1955). A contribution to the life histories of commercial shrimps (Penaeidae) in North
Carolina. Bull. Mar. Sci. Gulf' Caiibb. 5, 11646.
Manuscript received 18 August 1980, accepted 6 November 1980