,4quaculmral Engineering 5 ( 1986 ) 183-197

New Techniques for the Assessment and Optimal Management of Growth and Standing Crop Variation in the Cultured Freshwater Prawn, Macrobrachium

rosenbergii
Spencer R. Malecha
Aquaculture Research Program, Department of Animal Sciences. University of Hawaii. 1800 East-West Road, Honolulu, Hawaii 96822, USA

ABSTRACT The current 'traditional' prawn pond management +3"stem really does not 'manage', but rather accommodates prawn production characteristics attd was instituted with a minimum knowledge of prawn biology and husbandr3: A s sttch, the traditional system is not optimal because it does trot rely on a strong empirical knowledge base. Prawn culture has succeeded up until now because Macrobrachium rosenbergii is easily cult,tred attd will give 500-1000 kg ha -I year -t without much effort. This is su[ficient in many areas of the worm but trot in other+" where ecottomic conditions have break-even production requirements of nearly 1 tonne greater than this. The traditional system relying on selective harvesting with large seine nets is seriously inefficient which not only leads to lost reventte but ttnder-manages pond growth sittce unctdled large animals suppress the growth of unculled smaller ones. The degree to which this occttrs was trot k/town to the designers of the traditional system who had no way oJ"knowing (as we do now front our research results) the extremely large compensatory growth capability of small prawns in the absence of large ones. The traditional system also does not manage sexual dintorphic growth because no technolo~' exists which can be used to create monosex brood+" or manipulate the sex ratio in ponds. Accurate production models are not available because traditional ponds are rarely sampled attcb'or rarely drained. This, along with inefficient han,ests, results in a co-mingling of cohort stocking classes. It is impossible to get accurate survival attd growth dam because of this attd because it is so time con183 Aquactdtural Engineering 0 1 4 4 - 8 6 0 9 / 8 6 / S 0 3 . 5 0 - Elsevier Applied Science Publishers Ltd. England. 1986. Printed in Great Britain

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S. R. Malecha

surning to sample commercial ponds and process the dam on a regular basis. This paper describes four new engineering and 'bioengineering" techniques which are under development and can overcome all the drawbacks of the traditional system: (1) surgical sex reversal to create rnonosex broods; (2) genetic tagging of stocking and resident cohorts to assess survival and growth; (3) size grading and 100% ejficient harvesting using pond draining and a machine grader-hare'ester: and [4) a serni-automated computer assisted prawn sample data m a n a g e m e n t system which uses sonic digitization of prawn sample data.

INTRODUCTION The purposes of this paper are to point out the limitations of the current prawn pond culture system and to give a brief description of components of one system which is predicted upon a more efficient management of prawn growth variation and standing crop through efficient harvesting, computer assisted assessment of inventory, control of sex ratio, and more accurate computer based production models which use genetic tagging of prawn age and size classes and a semi-automated, computer based prawn sampling data management system. All the components of the new management system are by no means immediately at hand; however, we can expect future prawn pond production systems to be much different from what we have today.

THE 'TRADITIONAU PRAWN PRODUCTION SYSTEM The current prawn pond management system has been termed the "traditional' system (Malecha, 1983a) and involves stocking postlarval prawns once or twice a year into earthen ponds and then selectively harvesting market-sized animals using seine nets. Originally the traditional system was designed in Hawaii by Fujimura (1974) to take advantage of the fact that individual prawns in a pond population exhibit highly variable growth rates, so that by culling large animals from the population with nets, smaller animals could grow into the size classes 'vacated' by the larger animals and market-sized animals could be produced uniformly throughout the year. It had been assumed from pilot studies that periodic stocking and periodic seine-

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18 5

and-cull harvesting could sustain production throughout many years of stocking and ha~'esting. Unfortunately this does not appear to be true since stocking cohorts do not completely differentiate into clear model classes when harvesting is inefficient. The pilot studies on which the traditional system is based involved only two stockings, made roughly' one ,,ear apart, which distinguished themselves as the modes in a bimocl'al size-frequency distribution tFujimura, 1974). Apparently the periodic seine-and-cull harvest practiced in pilot studies gradually eliminated the first stocked cohort and then was applied to the second cohort as the latter grew to include harvestable animals. However, we believe that inter-size-class competition between stocking cohorts was minimized in the Fujimura pilot studies but is maximized in continuously stocked and harvested, but rarely drained, commercial prawn ponds. Indeed, net harvests capture 50% or less of the large harvest-eligible animals (Lam, 1984). The large animals remaining in the pond utilize resources and suppress the growth of smaller animals, delaying the latter group's entry into the harvest class. Over time a mixture of year classes comprise the pond population and newly stocked cohorts must compete with slowgrowing long-term residents and production is lowered significantly. In addition, the harvestable class may cause the entire population to exceed the critical standing crop in inefficiently harvested ponds so that competition is increased causing increased mortality'.

DEPENSATORY AND H E T E R O G E N E O U S INDIVIDUAL GROWTH (HIG) Inasmuch as the prawn's growth pattern constitutes one of its most visible and economically important biological characteristics we decided to study its genetic and environment determinants. Like most aquatic species the variance of the size distribution of Macrobrachium rosenbergii increases as its mean increases, i.e. it 'depensates" i Ricker, 1975), due to the fact that weight gain is a power function of initial weight. Most aquatic populations with a non-zero variance in their size distributions undergo depensation, but the ratio of the first and second moments usually remains relatively constant through time. Prawn and carp populations on the other hand display heterogeneous individual growth (HIG) characterized by an increased first to second moment

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S. R. Malecha

ratio and an elevated third moment, skewness, due to the fact that a spectrum of growth functions is represented in a growing M. rosenbergii population resulting in skewed (to the right) size distributions (Malecha et al., 1981b; Raanan, 1982). Since the prices paid for crustaceans are a function of their size, the prawn's growth pattern affects its product value to a degree not found in other aquatic animals and livestock. The prawns in the right-hand tail of the distribution are usually males and have been referred to as 'bulls' (Fujimura and Okamoto, 1972) and are analogous to "jumpers' in the common carp (Moav and Wohlfarth, 1974) and to "Tobi koi' in the ornamental carp (Wohlfarth, 1977). Individual prawns whose sizes lie around the population mode have been called 'runts' (Fujimura and Okamoto, 1972).

GENETIC AND ENVIRONMENTAL SOURCES OF HIG We have found that HIG is an equilibrium state of prawn populations and probably cannot be changed through domestication since it has high adaptive value. Males could vary genetically in their growth rates and consequently the stage in which they assume a certain role, but our research has shown that heterogeneous individual growth of males appears to be a non-genetic, socially dependent phenomenon (Malecha et al., 1984). In addition, we have conducted a series of water table experiments in the laboratory to study HIG (Malecha et al., 1981b). We found that HIG does not develop in populations of individually housed animals to the degree that it does in free-living populations and that populations of individually housed animals with individual water are not self-growth-inhibitory and depensate at the same rate as flow-through portion-segregated individuals sharing common water. We also found that tactile, visual and water contact across a barrier does not induce large animals to suppress the growth of smaller animals and that there were no significant differences between the HIG patterns in monosex populations and in populations of the same sex in bisexual populations. We concluded that the characteristic depensation patterns develop in males and females regardless of the presence of the opposite sex. In a recent experiment we tested whether water from a free living population of animals in rearing tanks could effect the HIG in a 'population' of individually housed animals when passed over them. HIG did develop in these control popula-

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18 7

tions, suggesting that a long-lived water borne factor could be involved with the animal-to-animal communication necessary for HIG to develop. Overall our results show that small prawns retain an inherent capacity, for compensatory, growth; consequently their sizes are largely ephemeral conditions brought on by non-genetic, extrapopulational behavioral factors. This means that HIG is best managed bv completely removing all the dominant males since their presence restricts the transformation of subdominant males into dominant males and small males into subdominant ones. Inefficient removal of large animals by inefficient harvesting which characterizes the traditional system diminishes this desired effect. Translated into practical terms, inefficient harvesting not only represents lost revenue by leaving animals 'behind' but also allows a growth suppression of the smaller animals by the larger ones. We now are convinced that the major limiting factor in prawn production is inefficient harvesting which under-manages the prawns' social structure. Unlike other agricultural species whose production limitation is usually a function of energy-based inputs (feeds, fertilizer) and animal biomass (density and size), a prawn population's biomass and yield is controlled and limited by its social structure.

OPTIMUM MANAGEMENT OF H I G - - S I Z E GRADED, STOCK DIVIDED AND EFFICIENTLY HARVESTED SYSTEMS On the basis of the results described above we have designed and tested in research ponds a new prawn stock management system theoretically described in Malecha et al. (1981a). Our results from over four years of field trials have shown that efficient harvesting has a dramatic effect on the compensatory growth capability of smaller prawns and that size-grading of prawns and stocking by size classes shortens the time to first harvest and the overall turnover time of the pond population. High production rates were achieved because the test populations were harvested with 100% efficiency using pond draindown and a machine grader (designed and fabricated by the University of Hawaii, Department of Agricultural Engineering). All animals were 'harvested out' of both experimental and control populations within 12 months. Production per unit time and area was greatest

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in the size-graded system due to the high turnover of the large sizegraded classes and the increased compensatory growth of the smaller animals. Also, time to first harvest was sooner in the size-graded system.

SEXUAL DIMORPHIC GROWTH AND SEX DETERMINATION In z~4acrobrachium rosenbergii female growth begins to sharply level off much earlier than male growth. Sexual dimorphic growth following the onset of sexual maturity, a common phenomenon, is usually managed directly in animal production. For example, in poultry sexual dimorphic growth is managed by the slaughtering of broiler hens earlier than cocks. Analogously, the opposite happens in M. rosenbergii. Males are harvested first and females are allowed to grow until they reach market weight at a size larger than their optimum. This represents the worst possible management of sexual dimorphic growth. Better management of sexual dimorphic growth is needed. This could be done if there were some means of generating all-male and/or all-female broods. In vertebrates, steroids produced in the gonads cause sex differentiation. Indeed, monosex broods are produced in fishes either through direct application of sex steroids to fry or from matings of normal parents and parents whose sex has been reversed by sex steroid application. Unfortunately this technology is not useful since readily available vertebrate sex steroids have limited effect in crustaceans. However, unlike fishes, in Malacostracan crustaceans the gametogenic and endocrine functions are separated respectively into distinct organs, the androgenic gland and gonad. The androgenic gland produces a hormone responsible for the differentiation of male secondary sexual characteristics and internal genital and gametogenic organs (Charniaux-Cotton et al., 1966; Thampy and John, 1973). The gland was first discovered by Charniaux-Cotton (1954)and shown by her (1959) and others (Katakura, 1959, 1960) to be capable of masculinizing a female when transplanted into the latter. Recently Nagamine et al. (1980a) have shown these effects in M. rosenbergii. Nagamine et al. (1980b) showed that andrectomized males develop into females with an ovatestes capable

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189

of oogenesis. Katakura (1961) produced viable all-female progeny from the mating of a masculinized and normal female isopod. It is possible that monosex, or altered sex ratio populations could be created by mating a phenotypically surgically sex reversed parent with a genetically but not phenotypically identical normal parent. Depending how sex control is inherited, there are many possible outcomes from such crosses. It is safe to assume that sex differentiation in prawns, as it is in other Decopod crustaceans, is under the control of a genic and/or chromosexual homo- and hetero-gametic system. We have been directing research toward rapidly performing the surNcal implantation of androgenic gland tissue into small sexually undifferentiated putative females. Our approach is to perform as many implants as possible and to progeny test the putative sex reversed parents. Table 1 shows putative genotypes of parents, their progeny, and progeny sex ratios in crosses between a normal parent and transplanted females reversed to neo-males, ablated males reversed to neofemales, or incorrectly sexed recipients or ablatees. Only chromosomal genotypes are shown. XX, XY, YY, XO are analogous to HH, Hh, hh, Ho genotypes where H and h are sex determining genes. Table 2 depicts a possible scenario for hypothetical commercial scale application of culturing monosex broods based on surgical sex reversal. 'Bottleneck level' (Column 1) signifies certain production or mortality assumptions given in the footnote. The scenario is based on a farm of 4500 ha (100 acres) with adult standing crop of 10 individuals m - " or 100000 h a - ~ = 4 - 4 x l 0 ~' farm -~ (1 individual ft -2 = 4 4 0 0 0 a c r e - l = 4 . 4 x 10 ~ farm-t).

M E T H O D S FOR D E V E L O P I N G M O R E ACCURATE PRODUCTION MODELS

Genetic tagging of stocking cohorts

There is a lack of accurate production models in prawn culture due to the fact that a strong empirical data base does not exist to support the assumptions of the models. The reason for this is twofold. First, prawn farmers do not frequently sample their ponds because of high labor

m.,.

TABLE 1 Possible O u t c o m e s of Sex-Reversal T r a n s p l a n t a t i o n s and Ablations

0

(a) Trtmsplanmtio, (sex reversal crosses)

N ( ' o illlll(' h

I"emah's of normal mule'X

Xy;, X O j, XX;' XY' XX' XY XO XX XX XY -" ~ ~ ~ ~ 1:2:t 1:2:1 1:0:0 I:t:0 I : 1:0

/1(;" itz (1)

Normal fi'mah,

I~,lative progeny ( )bserved l)henotypic progeny genoopic ratio sex ratio XA'.'XY(XO)." YY(O0) mules.'j'emales (comntenO
I : 3 ( Y Y lives), 1:2 (YY dies) 1:3 (OO lives), 1:2 ( O O dies) 0:1 (all females) 1:1 (normal ratio) 1:1 (normal ratio)

I Icterogamelic, XY Heterogametic, XO Homoganlctic, XX Homogametie, XX Heterogametie, XY

(b) Ablation (sex n, vel:sion crosse.s)

Males Nee female t' oJ'normal female' X

XY t' XO ~ XX I' XY' XX' XY XO XX XX XY ~ ~ ~ ~ ~

/1(;" ottt (1)

Normal male

l'utative plvgeny Observed l)henoO,l~ic progeny genotypic ratio sex ratio XX:XY(XO) : Y Y ( O 0 ) males.'J'emales (comment)
1:2:1 1:2:1 1:0:0 1 : 1:0 1:1:0 3:1 3:1 1:0 1:1 1: I (YY lives), 2:1 (YY dies) ( O O lives), 2:1 ( O O dies) (all nmles) (normal ratio) (normal ratio)

t'.,

Heterogametic, XY Heterogametic, XO Homogametic, XX Homogametic, XX Heterogametic, XY

~ ~ ~ ~ ~

" A n d r o g e n i c gland tissue. ;'Indicates surgically manipulated animals: n e e females = ablatces; nee males = transplant recipients. ' Indicates +mis-sexing' normal animals, i.e. putative n e e - m a l e s are really n o r m a l males and putative nee-females arc really normal females.

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191

TABLE 2
Bottleneck level ~ Estimated numbers Description

6 5 4 3

4400000 t 5 860 000 9 766600 t 10851 778 t "~17 271

542

Standing crop number per 100 acres (approx. 45 ha) 75% pond survival PLS stocked 60% hatchery survival Viable larvae hatched 90% hatch rate (egg to larvae) Viable fertilized eggs spawned 50 000 eggs per females Gravid females 80% mating success Reproductive competent parents "50% success rate" Surgically manipulated animals

"Assumptions and/or source for each bottleneck level: 6 - - Smith et al. ( 1981 ); 5 - estimate based on performances of three commercial and one government hatcheries in Hawaii (Malecha, 1983a); 4 - - assumption only - - no published data available on the relationship of number of fertilized spawned eggs and number of viable larvae hatched; 3 - - based on about 5 0 0 0 0 eggs per female. Malecha (1983a) showed that a laboratory-reared 45 g female can produce over 40 000 eggs per spawn, and fecundity, K is related to length. X, by the equation Y= 0-006 X ~ : ; 2 - - from Malecha et al. (1984); t - - estimated achievable project goal - - 50% of surgically manipulated animals survive to produce viable offspring.

costs and the inconvenience of obtaining, recording, reducing and analyzing measurement data. The cost of obtaining animals by seining is minor compared to data management. Second, almost all prawn farms use the traditional pond management system (Malecha, 1983b) whereby ponds are periodically stocked and harvested without complete pond draining and restocking. This results in an admixture of stocking to resident pond cohorts and it is impossible to assess the degree to which this is occurring and to determine growth rate, growth pattern, survival, and time of entry to harvest of these cohorts once multiple stocking has progressed. Because of this, commercial prawn farming is characterized by unquantified assumptions regarding the percent harvestable animals and the movements of the size distribution in resident and stock cohorts model classes. In effect, commercial

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.5. R. Malecha

prawn farmers have really no economically viable precise way of assessing their standing crop, predicting their harvests and developing accurate production models and cash flow projections based on inventory, assessment. All-in-all, prawn farming remains largely a 'hitor-miss" affair dependent upon "what comes out' at the harvest. An efficient way of tagging prawns according to their stocking cohort type is clearly needed. However, marking thousands of stocking juveniles needed for assessing growth, survival and harvesting patterns is impossible with current tagging technology. For this reason we have developed a genetic tag utilizing naturally occurring, electrophoretically detectable genetically controlled enzyme variants called 'allozymes'. First we have established broodstock carrying rare allozymes in a large electrophoretic survey of Hawaiian prawns. Controlled crosses among these broodstock are producing populations of genetically tagged stocking cohorts which can be easily distiguished from other stocking cohorts and resident pond cohorts. We have shown (Hedgecock et al., 1979; Stelmach, 1980) four allozyme systems to be variable in the Hawaiian cultivar of Macrobrachium rosenbergii called the "Anuenue strain' (Malecha, 1980). We have selected glutamate oxaloacetate transaminase (Got-l) and phosphoglucomutase (Pgm) for this study since they are the most polymorphic allozyme systems, they are easiest to assay, and thev are rare. The nomenclature for the two allozyme systems is shown in Table 3.

TABLE 3

Gore system

Alleles g'getws)

Got-1 ~m

103, 100, 94 100, 97

Each allelic gene, as a segment of the genetic material, controls the synthesis of one protein enzyme which is revealed as a zone or band by histochemical staining on a starch gel following electrophoresis of muscle homogenates extracted from walking legs of sampled and harvested prawns.

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193

TABLE 4 Observed Frequencies of Common and Rare Genotypes at the Got-1 and Pgm Loci Based on a Survey of 2000 Adult Prawns Collected from Commercial Ponds
Gene system Common geno~'pes Rare geno~. 'pes

Got-I

100/100

100/103
100/94 Frequency 96"74% 103/103 0.7% 94/94 1-07% 103/94 1-49%

Pgm
Frequency

lO0/100 lOO/97
99.0%

97,/97
1.0%

For Got-l, six diploid genotypes may be formed from the three alleles, while three different diploid genotypes are possible for the Pgm system. These are listed and grouped in Table 4 into the common and rare genotypes detected in a survey we conducted of 2000 adult prawns collected from commercial ponds. As can be seen in Table 4, over 96% of the commercial Hawaiian prawn pond populations have common Got-1 genotypes and 99% have common Pgm genotypes. Therefore, stocked cohorts which are homozygous or heterozygous for any of the rare genotypes can be readily distinguished from resident populations with an error proportional to the naturally occurring frequency of the rare genotypes involved. The genetic tagging of cohorts will allow a very accurate assessment of the degree to which admixture occurs within and between stocking and pond resident cohorts and it will then be possible to assign accurate resident times and probabilities of survival and growth among stocking cohorts and between size classes within a stocking cohort. This information is essential for developing production simulation models for a pond system where harvesting is not efficient, the ponds are not drained and the age structure of the resident is not known.

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S. R. Malecha

COMPUTER BASED SAMPLING DATA MANAGEMENT We have developed components of a computer based data acquisition and analysis system which could greatly reduce the cost of commercial prawn pond sampling data management. The main feature of the system is that prawn len~hs are digitized instantaneously and recorded, compiled and stored in a microcomputer. Labor is reduced by half and the materials handling (e.g. prawns, rulers, data sheets, hand calculators, nets, etc.)is reduced to a fraction of that required in the present system. Figure 1 shows the general organization of the system which we are developing. A slide image of prawns is obtained by photographing individuals scattered onto a fiat surface using a waterproof camera containing Polaroid 35 mm transparency (slide) film which can be developed on the spot using the Polaroid Autoprocessor. High contrast, high quality images are not required for the system to work; only the prawn's "orbit length' (distance between the eyestalk orbit posterior surface and the tip of the telson) has to be discernible on the projected image. In practice, even a cloudy or poor quality image can be measured at least as accurately as is presently done with a ruler or caliper. Slide images are projected (using a standard carousel projector) onto the back of a semi-opaque window mounted in the side of a projection cabinet. Two mirrors inside the cabinet carry the slide

I I I I sonic digitization I t I I I storage R232 analysis output~ input IBM PC L. I compilation f f I 1 _1 Fig.1. Flow diagram forsonicdigitization ofprawnlengths anddataanalysis.
@
sli i e

J1projected w image

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19 5

image from the slide projector located on the floor of the cabinet to the projection window. A Graf/Bar Sonic Digitizer (Model GP-7, Science Accessories Corporation, Westport, Conn, USA) is mounted on the projection cabinet out in front of and on top of the image projection window. A stylus, electronically connected to the Graf/Bar main chassis, is capable of emitting an electronic signal and a sonic sound when its tip is depressed by touching a point within the active area. The electronic signal is cabled back to the chassis. Microphones located on the chassis ends pick up the sonic signal from a stylus. The relative time the signal takes to reach each of the microphones is then recorded by the device and the Cartesian coordinates of the point relative to an oriNn in the reference corner of the active area is determined. These coordinates are then transmitted as ASCII data to the asynchronous communications port of a microcomputer by an RS232 connector. The program which we wrote then records these coordinates. All software for input management and analysis of the data is written in Advanced IBM PC BASIC. The software is user-friendly and menu driven. When two points (at either end of the prawn image) have been digitized the program calculates the distance between the points and converts this to an actual length by the scaling rule or a standard of known length, and then calculates the animal weight. Analyses programs which convert the lengths to weights generate weight frequency histograms and statistics, plot the mean weight of the pond population against time, and print out hardcopies of a juxtaposition of histograms and statistics in chronological order, pond name and start and end dates.

REFERENCES Charniaux-Cotton, H. (1954). Ddcouverte chez un Crustace Amphipode ( Orchestia gamarella) d'une glande endocrine responsable de la diffdrenciation des caract~res sexuels premieres et secondaires males. C. R. Hebd. SOances Acad Sci. S&. D, Sci. Nat., 239,780-2. Charniaux-Cotton, H. (1959). Masculinization des femelles de la Crevette/i hermaphrodisme prot&andrique Lysmata seticaudata, par greffe de D~capodes. Note prdliminaire. C. R. Hebd. S~ances Acad. Sci. SSr. D, Sci. Nat., 249, t580-2. Charniaux-Cotton, H., Zerbib, C. & Meusy, J. J. (1966). Monographie de la glande androg~ne des crustaces supdrieurs. Cmstaceana, 10, 113-36.

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Fujimura, T. (1974). Job completion r e p o r t - - D e v e l o p m e n t of a prawn industry in Hawaii, July 1, 1969 to June 30. 1972. US Dept. Commerce, N O A A NMFS, 21 pp. Fujimura, T. & Okamoto, H. (1972). Notes on progress made in developing a mass culturing technique for Macrobrachium rosenbe~ii in Hawaii. In: Coastal Aquactdture in the Indo-Pacific Region, ed. T. V. R. Pillar. Fishing News Books, Ltd, London. Hedgecock, D., Stelmach, D. J., Nelson, K., kindenfelser, M. E. & Malecha, S. R. (1979). Genetic divergence and biogeography of natural populations of Macrobrachiurn rosenbergii. Proc. World Maricul. Soc., 11,500-28. Katakura, Y. (1959). Masculinization through implanting testes into the female Arrnadillidium vtdgare, as isopod crustacean. Proc. Jpn. Acad., 35, 95-8. Katakura, Y. (1960). Transformation of ovary into testes following implantation of androgenous glands in Armadillidiurn vulgare, an isopod crustacean. Annot. Zool. Jpn., 33, 241-4. Katakura, Y. ( 1961). Progeny from the mating of the normal female and the masculinized female of Armadilliditml vulgare, an isopod crustacean. Annot. Zool. Jpn., 34 (4), 197-9. Lam, C. Y. (1984). Forecasting freshwater prawn harvest yields. MSc Thesis. Dept. Agri. Eng., University of Hawaii, Hawaii. 61 pp. Malecha, S. R. (1980). Development and general characterization of genetic stocks of Macrobrachium rosenbergii and their hybrids for domestication. Universi~' of Hawaii, Sea Grant Quarterl), 2 (4), 6 pp. Matecha, S. R. (1983a). Commercial seed production of the freshwater prawn Macrobrachium rosenbergii. In: CRC Handbook of Marine Science Vol. L Aquaculture of Crustaceans, ed. J. P. McVey, CRC Press, Boca Raton. Florida, pp. 205-30. Malecha, S. R. (1983b). Commercial pond production of the freshwater prawn Macrobrachium rosenbergii. In: CRC Handbook of Marine Science Vol. L Aquaculture of Crustaceans, ed. J. P. McVey, CRC Press, Boca Raton, Florida, pp. 231-59. Malecha, S. R., Polovina, J. & Moav, R. (198 l a). A multi-stage rotational stocking and harvesting system for year round culture of the freshwater prawn, Macrobrachitml rosenbergii. UNIHI-SEAGRANT-TR-81-01. University of Hawaii. Malecha, S. R., Bigger, D., Brand, T., Levitt, A., Masuno, S. & Weber, G. ( 1981 b). Genetic and environmental sources of growth pattern variation in the cultured freshwater prawn, Macrobrachium rosenbe~ii. Paper presented at World Conference on Aquaculture, September 21-25, 1981, Venice, Italy. Malecha, S. R., Masuno, S. & Onizuka, D. (1984). The feasibility of measuring the heritability of growth pattern variation in juvenile freshwater prawns, Macrobrachium rosenbergii (de Man). Aquaculture, 38,347-63. Moav, R. & Wohlfarth, G. (1974). Carp breeding in Israel. In: Aqllacultura/ Genetics, ed. R. Moav, John Wiley and Sons, New York, pp. 295-318.

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Nagamine, C., Knight. A. W., Maggenti. A. & Pa.xmen, G. ~1980ai. Masculinization of female Macrobrachium rosenbergii (de Man)(Decapoda: Palaemonidae) by androgenic gland implantation. Gen, Comp. Endocrin., 41,442-57. Nagamine, C.. Knight, A. W., Maggenti, A. & Paxmen, G. (1980b). Effects of androgenic gland ablation on male primary and secondary sexual characteristics in the Malaysian prawn, 3,lacrobrachium rosenbergii (de Man) (Decapoda: Palaemonidae), with first evidence of induced feminization in a non-hermaphroditic decapod. Gen. Comp. Endocrin., 41,423-41. Raanan, Z. (1982). The ontogeny of social structure in the freshwater prawn Macrobrachium rosenbergii !de Man). PhD Thesis, Hebrew University, Jerusalem, Israel, l 01 pp. Picker. W. E. (1975). Computation and interpretation of biological statistics for fish populations. Bull. Fish. Res. Bd. Can., 191, 38 l. Smith. T. I. J., Sandifer, P. A., Jenkins, W. E. & Stockes, A. D. ( 1981 ). Effect of population structure at stocking and density on production and economic potential of prawn (Macrobrachium rosenbergii) farming in temperate climates. Proc. World Maricul. Soc., 12,233-50. Stelmach, D. J. (1980). Electrophoretic variation in Macrobrachium rosenbergii: evidence for subspecific variation. MSc Thesis, Dept. of Genetics, University of Hawaii. Hawaii, 40 pp. Thampy, D. M. & John, P. A. (1973). Observations on variations in the male sex characters and their relation to the androgenic gland in the shrimp Macrobrachium idae(Heller). Acta Zool. (Stockh.), 54, 193-200. Wohlfarth, G. (1977). Shoot carp. Bamidgeh, 29, 35-9.