Penaeid Genetics and Biotechnology

Aquaculture 164 1998.
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Penaeid genetics and biotechnology

J.A.H. Benzie
) Australian Institute of Marine Science and Cooperatie Research Centre for Aquaculture, P.M.B. 3, Townsille, M.C. QLD 4810, Australia
Abstract The application of biotechnology to penaeid prawn aquaculture is increasing in importance. Early doubts that such approaches were unnecessary, given the crude state of development of the prawn aquaculture industry, have been replaced by the recognition that sophisticated molecular approaches will be required to tackle key problems of disease recognition and control, and to achieve higher production through the development of domesticated strains. Progress in penaeid genetic and biotechnological research has been slow because of a lack of knowledge on fundamental aspects of their biology. However, data is beginning to emerge from research projects started in the last decade, and is likely to increase rapidly in the next 10 years. Highly variable markers have been developed that allow the genetic variation in prawn stocks to be assessed, even in highly inbred cultured lines. The extent to which growth rate is under genetic control has now been assessed in some species using rigorous experimental designs to estimate heritabilities. Highly sensitive techniques that allow the isolation and characterisation of very small quantities of peptides are being used to investigate the endocrine control of reproduction, and work has begun on the isolation and characterisation of genes that play an important role in growth and reproduction. Greater attention is being paid to developing cell lines for prawns, and to the development of molecular probes for the identification of pathogens. These tools will provide a basis for a more detailed description of the penaeid prawn genome, for understanding disease resistance, and for developing effective control of reproduction and the development of disease free or disease resistant domesticated lines. q 1998 Elsevier Science B.V. All rights reserved.
Keywords: Genetics; Biotechnology; Maturation; Endocrinology; Prawns; Disease; Selective breeding; Penaeids; Aquaculture; Larval quality
Corresponding author. Tel.: q61-7-47534444; fax: q61-7-47725852.
0044-8486r98r$19.00 q 1998 Elsevier Science B.V. All rights reserved. PII S 0 0 4 4 - 8 4 8 6 9 8 . 0 0 1 7 5 - 6
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J.A.H. Benzier Aquaculture 164 (1998) 2347
1. Introduction The potential for the application of genetics and biotechnology to prawn aquaculture has been recognised for some time Primavera, 1985; Malecha and Hedgecock, 1989; Hedgecock and Malecha, 1991.. Areas in which biotechnologies were thought to be most likely to have a practical impact were in the control of reproduction, the development of domesticated strains and in the recognition and control of diseases. However, it was also thought a decade ago that many of the practical problems would be overcome by focusing on simpler approaches to husbandry and nutrition. The major commercial emphasis then was on the problems of pond production. The fact that a lack of information on basic aspects of prawn biology meant that biotechnological research needed to focus on developing the information and basic tools required for investigation of the genome and the physiology of penaeid prawns, also extended the time scale required for the practical application of biotechnology Hedgecock and Malecha, 1991.. The growing problem of diseases which have devastated, and continue to threaten, production of several species throughout the world, has emphasised the need to develop tools for the rapid recognition and control of pathogens. It has also been recognised that the only long-term solution to maintaining production in the face of endemic pathogens is the development of disease-free or disease-resistent domesticated stocks Wyban et al., 1993; Carr et al., 1996a,b.. This realisation has also emphasised the problems which remain with respect to the effective control of reproduction of cultured stocks. While a number of species have been spawned in culture, for some such as Penaeus monodon, pond-reared animals remain difficult to spawn effectively. Even in species which are considered easy to breed in culture, such as Penaeus japonicus and Penaeus annamei, there appear to be problems in the quality of larvae from domesticated stocks. It is telling that, even where it is considered easy to obtain pond-reared spawners, the prawn farmers prefer, or still rely upon, wild populations for their supply of spawners. The fact that these wild stocks are now threatened in many countries, or lost in some, has added urgency to the need to develop domesticated strains Liao, 1990; Primavera, 1993; Laubier and Laubier, 1993.. As yet, there are few large scale prawn breeding programs based on sound animal breeding practices which could provide a base for the development of basic information on the genome, and a base for the development of commercial breeds Wyban et al., 1993; Benzie, 1994; Bedier et al., 1996.. It is clear that the problems now confronting the prawn aquaculture industry will require the combined application of biotechnologies targeted at several areas, and in several species, in order to develop a sustainable prawn farming industry. The considerable advances made in recent years in the rapidity and ease of use of molecular tools will assist the development of penaeid biotechnologies. However, the notion that super-prawn biotechnologies will appear very quickly does not take adequate account of the primitive state of knowledge currently available for penaeids compared with that known for species established in agriculture. Nevertheless, work by several research groups over the last decade has started to provide the information required to begin this process, and is reviewed in the present paper.
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2. Genetic variation 2.1. Molecular ariation The importance of genetics for the continued productivity of cultured stocks of penaeid prawns was starkly revealed by the close association of a decline in spawner productivity with reducing genetic variability of allozyme variants in P. japonicus farmed in Italy Sbordoni et al., 1986, 1987.. It is remarkable that this relationship was observed using allozymes, as penaeid prawns usually show very little allozyme polymorphism Hedgecock et al., 1982; Koh et al., 1983; Tam and Chu, 1993.. Nevertheless, the finding focused attention on the possible deleterious consequences of uncontrolled inbreeding, and engendered attempts to maintain larger effective population sizes in breeding programs Malecha and Hedgecock, 1989.. Sunden and Davis 1991. subsequently reported reduced allele numbers, but no significant depression of heterozygosity, in cultured populations of P. annamei where management procedures kept a high effective population size. This finding suggested it is possible to maintain sufficient levels of genetic variation in cultured stocks over the short term by maximising the variation in the base populations, and in the longer term by including progeny from a large number of broodstock to prevent inbreeding. Surprisingly, Laubier et al. 1984. found that levels of genetic variation in fourth generation stocks of P. japonicus originally derived from crosses of two separate sets of progeny, each derived from an unknown number of females, were similar to that observed in natural populations, but the long term genetic status of this population is unknown. Early allozyme work emphasised the lack of significant geographic structure in wild populations of penaeids Lester, 1979; Mulley and Latter, 1980, 1981a,b; Villaescusa et al., 1984. although some variation was observed among populations of P. latisulcatus Richardson, 1982. and Metapenaeus bennettae Mulley and Latter, 1981a; Salini, 1987. in Australia, and P. kerathurus in the Mediterranean Mattoccia et al., 1987.. More recent allozyme surveys over large geographical scales subsequently showed highly significant variation among wild stocks of P. monodon in Australia Benzie et al., 1992. and Indonesia Sugama and Haryanti, 1997.. The populations sampled from the western-most site in each country were differentiated from populations to the east. Further work with more highly variable mtDNA markers is revealing greater structuring of P. monodon populations in Australia Benzie et al., 1993. and south-east Asia Sodsuk et al., 1992, 1996., demonstrating the diversity of wild genetic resources that the industry might access for breeding programs. It is likely that the transfer of large numbers of spawners from the Andaman Sea to farms in the Gulf of Thailand has resulted in a change in the genetic structure of the wild stocks of P. monodon in the Gulf of Thailand Sodsuk et al., 1992, 1996. emphasising the need for care in the distribution of cultured stocks that might endanger wild genetic resources Benzie, 1996.. High genetic diversity in the mtDNA of Penaeus notialis and P. schmitti populations has also been recorded Machado et al., 1993.. The desire to assess the genetic diversity within prawn stocks led to attempts to develop DNA-based markers that were likely to show higher levels of variation, and to have greater sensitivity in detecting genetic variability in inbred cultured stocks, than
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allozyme variants. Considerable variation in randomly amplified polymorphic DNA RAPD. was found in families of P. annamei Garcia et al., 1994. and P. monodon Garcia and Benzie, 1995., and their potential for use as family specific markers has been demonstrated in these studies. Mitochondrial DNA mtDNA. and short sequence repeat DNA SSR or microsatellite DNA. was also shown to vary considerably within and between families of P. annamei Garcia et al., 1996.. Variants at the COI mitochondrial gene in these P. annamei families were associated with individuals showing high growth, providing positive evidence that molecular variants could be used as markers for traits of commercial importance, an important goal for developing information on molecular variation in prawns. 2.2. Morphological ariation The development of sound data concerning the level of genetic variation for quantitative genetic characters in cultured prawn populations has been slowed by the fact that environmental influences on prawn growth are considerable. Early work concentrated on examining morphometric data to determine simple and reliable characters that could be used to assess broodstock quality mainly defined as tail size., to assist in the choice of individuals for inclusion in breeding programs Lester, 1983; Huang et al., 1990; Goswami et al., 1990.. The difficulty of producing large numbers of defined matings, and of providing adequately controlled rearing conditions has slowed efforts to estimate the heritability of quantitative traits of commercial importance. The fact that hatcheries using different genetic stock differ in their performance Chow and Sandifer, 1991., and that different families vary in growth performance Lester, 1988; Lester and Lawson, 1990., does not provide a sound means of estimating the extent to which growth is under genetic control because of the effects of environmental differences between hatcheries and because of large environmental variances between rearing tanks Benzie, 1997.. Estimates of the heritability of weight of 6- and 10-week-old P. monodon postlarvae have been made by Benzie et al. 1997. using a half sib design where each of 18 male P. monodon were mated with two females, and the heritability estimated from the male component of variance which is relatively free of environmental effects. This showed that about 10% of the difference in growth between progeny was a result of genetic factors, an estimate likely to be a minimum, since heritability estimates for size tend to be low in young animals, and to increase with age. Recent estimates obtained from 57 full-sib families, and therefore including some environmental effects, showed heritability of weight in a small pathogen-free population of P. annamei to be about 0.42 " 0.05 Carr et al., 1996a,b.. This was similar to values of full-sib estimates for P. monodon 0.56 " 0.03 at 6 weeks, 0.39 " 0.004 at 10 weeks., with the drop in heritability over time thought to be the result of reducing maternal effects Benzie et al., 1997.. The appearance of strong resistance to IHHNV in an inbred stock of P. stylirostris Bedier et al., 1996. and preliminary reports of considerable variation in survival among families challenged with taura virus in P. annamei Pruder et al., 1996. give good indications that disease resistance could be selected in penaeids. Success has now been achieved in rearing large numbers of families from defined matings for P. annamei Wyban et al., 1993. and P. monodon Benzie et al., 1997.,
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and work is in progress with P. stylirostris Bedier et al., 1996.. This technical capacity will now provide the means to assess responses to selection and to define breeding programs to achieve the general breeding goals for prawns outlined years ago Lester, 1983; Malecha and Hedgecock, 1989.. 3. Genome structure 3.1. Genome size and chromosome number Some additional information on the structure of the penaeid genome has appeared since the review of crustacean genetics by Hedgecock et al. 1982., but the data base is still limited and highly fragmented. The genome size of four penaeid species Penaeus aztecus, P. duorarum, P. setiferus and P. annamei . was estimated by flow cytometry to be approximately 70% that of the human genome Chow et al., 1990.. Information on the number of chromosomes present is available for several penaeid taxa with the diploid chromosome number ranging from 2 n s 6492 Hayashi and Fujiwara, 1988; Dai et al., 1989; Chow et al., 1990; Kong and Zhang, 1993; Xiang et al., 1991, 1993.. Sicyonia ingentis has 2 n s 64 Xiang et al., 1991., Metapenaeopsis barbata 2 n s 68 Hayashi 1988. referenced in Murofushi and Deguchi, 1990., and Xiphopenaeus kroyesi 2 n s 78 Xiang et al., 1993.. P. japonicus has most likely a 2 n s 86, obtained from many counts of air dried preparations from blastema cells Hayashi and Fujiwara, 1988. rather than 2 n s 92 obtained from squashes of other tissues with fewer dividing cells Niiyama, 1948.. P. semisulcatus has 2 n s 90 Xiang et al., 1993. as has P. setiferus Milligan, 1976; Chow et al., 1990., P. stylirostris, P. occidentalis and P. californiensis 2 n s 92 Mayorga, 1982.. All other species investigated had 2 n s 88: P. aztecus Milligan, 1976; Goswami, 1985; Chow et al., 1990., P. chinensis s P. orientalis . Xiang, 1988; Dai et al., 1989., P. duorarum Milligan, 1976; Chow et al., 1990., P. monodon Kong and Zhang, 1993; Xiang et al., 1993., P. esculentus, P. merguiensis, and P. penicillatus Xiang et al., 1993.. Differences in the chromosome numbers reported for some species w Trachypenaeus curirostris 2 n s 70 Xiang et al., 1993., 2 n s 80 Hayashi 1988. referenced in Murofushi and Deguchi, 1990.; P. setiferus 2 n s 90 Milligan, 1976; Chow et al., 1990., 2 n s 88 Xiang et al., 1993.; and P. annamei 2 n s 92 Mayorga, 1982., 2 n s 88 Chow et al., 1990.x, are likely to have resulted from the large number of chromosomes present and the small number of dividing cells available from which to obtain counts in squashes of most tissues Hayashi and Fujiwara, 1988. rather than geographic variation in chromosome numbers. The more reliable reports include tables of the frequency distribution of the range of counts obtained. The report of 2 n s 28 for P. duorarum in Xiang et al. 1993. was most probably a misprint. Chromosome spreads were most readily attained from testes but large numbers of dividing cells from which to obtain somatic metaphase spreads for determining chromosome morphology were found in regenerating blastemas Hayashi and Fujiwara, 1988.. Most of the chromosomes in all penaeids studied to date are metacentric or sub-metacentric in type with some sub-telocentric Dai et al., 1989.. Suggestions continue to be made that the changes in chromosome numbers observed in the Decapoda arise from polyploidisation events Murofushi and Deguchi, 1990.. This
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is despite the fact that electrophoretic surveys have shown no evidence of gene duplication and that the relative amount of non-satellite DNA declines or remains constant with phylogenetic position, suggesting polyploidisation is not the principal mechanism for increasing chromosome number in the crustacea Hedgecock et al., 1982.. The data bases are still relatively small, however, and generalisations concerning the evolution of penaeid chromosome structure must await the collection of further information. One point in common for the few species for which detailed information on the relative size and structure of penaeid chromosomes is available, is that no defined sex chromosome has been identified Hayashi and Fujiwara, 1988; Dai et al., 1989.. 3.2. Gene sequence data Comparison of mtDNA sequence data from the 12S ribosomal gene and cytochrome oxidase I subunit from P. annamei, P. stylirostris and P. esculentus and Metapenaeus endeaouri demonstrated a surprising degree of genetic divergence between these taxa Palumbi and Benzie, 1991.. Differences in transversions at silent sites between subgenera of shrimp 18%. were larger than that between families of mammals 10%., and intergeneric differences in shrimp 25%. were more divergent than between some orders of mammals 23%.. Sequence from a section of the 16S ribosomal gene in P. notialis and P. schmitti Machado et al., 1993., although less than that recorded by Palumbi and Benzie 1991., confirmed the relatively large genetic differences between these shrimp species 9.4% genome divergence between subgenera.. Crude restriction fragment length polymorphism RFLP. maps of the mitochondrial genome show marked differences between P. monodon and P. japonicus, and vary between populations from different parts of P. monodons range Bouchon et al., 1994.. The size of the mitochondrial genome falls in the range of 1517 kb in all species for which data exists Bouchon et al., 1994.. Apart from the mitochondrial gene segments sequenced by Palumbi and Benzie 1991. and Machado et al. 1993., sequence data exists only for the 18S ribosomal nuclear gene of P. aztecus Kim and Abele, 1990., a cDNA encoding a serine protease in P. annamei Sellos and van Wormhoudt, 1992., a cDNA encoding a moult-inhibiting-like peptide in P. annamei Sun, 1994. and for two 1.5 kb and 2.0 kb strands of unknown genes from P. annamei Garcia et al., 1996.. cDNA sequence of arginine kinase has been used to determine amino acid sequences for that enzyme in P. japonicus and established a 51% homology in amino acid sequence between P. japonicus and the abalone Nordotis madaka Furukohri et al., 1994; Suzuki and Furukohri, 1994.. There is no information concerning the chromosomal location of genes for any penaeid. 4. Genome manipulation 4.1. Interspecies hybridisation Natural service matings between species are often prevented by behavioural mechanisms, as illustrated by the failure to cross P. annamei and P. setiferus Misamore and
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Browdy, 1994., although in vitro studies suggested that there might also be post-mating isolation mechanisms acting to prevent interspecific crosses in that species pair. The development of techniques of artificial insemination have allowed a number of successful attempts at interspecies hybridisations in the penaeidae Benzie et al., 1995.. Listing the female first, the following species pairs have produced hybrid larvae: P. setiferus= P. stylirostris Lawrence et al., 1984., P. setiferus= P. schmitti Bray et al., 1990a,b., P. monodon= P. penicillatus and P. penicillatus= P. monodon Lin et al., 1988. and P. monodon= P. esculentus Benzie et al., 1995.. The proportion of matings which led to spawnings, the proportion of these which hatched, the proportion of eggs in a spawn which hatched, and the survival to the postlarval stage varied considerably between these crosses. However, the survival to postlarvae stage 20 was low - 1%. in all cases, with no correlation between the success of the hybridisation and the closeness of phylogenetic relationship of the taxa involved. Most of the hybrids were not reared for long enough to test their fecundity, but where they were, no maturation of females was observed and no matings of the hybrids was obtained Lin et al., 1988; Bray et al., 1990a,b.. 4.2. Ploidy manipulation and cryopreseration A detailed description of the timing of meiotic divisions after spawning, and of the timing of the fusion of the pronuclei and later mitotic divisions, has been published for S. ingentis by Clark and Griffin 1993. who have summarised the limited data available for other species P. japonicus and P. aztecus .. In S. ingentis, the first polar body is released 35 min after spawning and the second polar body is released 45 min after spawning Clark and Griffin, 1993.. The success of ploidy manipulations in other species also suggest that both polar bodies are extruded in the first hour after hatching in P. chinensis and Penaeus indicus Xiang et al., 1991, 1992; AQUACOP et al., 1993.. Polyploid penaeids have been successfully produced using chemical and temperature shock and the animals reared to early larval, and in one case, to late post larval stages Xiang et al., 1991, 1992; AQUACOP et al., 1993.. Tetraploid and triploid P. indicus were produced by heat shock of 17 min duration applied 546 min after the eggs were spawned AQUACOP et al., 1993.. At the respective optimal conditions, details of which were not reported, 47% triploid and 93% tetraploid nauplii were produced with 19% and 0.3% survival in these groups relative to diploid controls at the end of the naupliar phase. Haploid zygotes of S. ingentis were produced by fertilizing eggs in vitro with UV irradiated sperm, and both triploid and tetraploid zygotes produced by chemical and temperature shock of normally fertilized eggs Xiang et al., 1991.. The success of the operation was reported to vary considerably on the time between hatching and treatment, temperature and the duration of the treatment, but optimal conditions were not reported. The note that eggs were too fragile to survive early treatment well, and that the extrusion of the second polar body was easier to inhibit than that of the first, suggested later times of treatment were more effective. Heat shock and treatment with cytochalasin B produced 3360% and 2166% tetraploid P. chinensis respectively in a series of trials Xiang et al., 1992.. Heat and chemical treatments lasted 1520 min and, in the case of cytochalasin B was applied 5065 min after fertilization. The duration of cold
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shock, and time at which cold and heat shocks were administered, was not stated. Tetraploids reared to 6 months old demonstrated a growth rate 20% higher than the diploid controls, but all experimental animals died soon after. There was no report on the sexual development of these specimens and there are no confirmed reports of polyploid animals being reared to maturity. The formation of haploid zygotes of P. chinensis s P. orientalis. using sperm irradiated with 49 Co g Dai et al., 1993., and the successful production of 14.9% and 37.2% diploid gynogens by later inhibiting the extrusion of one polar body in P. orientalis Naner and Feng, 1993. has been reported. Work in this area is still in its infancy. Part of the difficulty of research in ploidy manipulation is the fact that the unfertilised gametes are generally difficult to obtain in vitro with the exception of S. ingentis, which has therefore been used to investigate penaeid gamete activation and early development Clark and Griffin, 1993.. A block to fertilisation resulting from the development of a hard egg membrane within 1215 min of contacting seawater, also places constraints on in vitro fertilisation experiments. Cryopreservation of sperm has been achieved Clark and Griffin, 1993. and further development of this technique will be of use in future breeding programs, but reports of the cryopreservation of eggs and early larval stages have not been substantiated. 4.3. Gene expression and transgenesis No detailed molecular information exists on gene expression in penaeids. Variation in allozyme patterns at different stages of larval development in P. aztecus, P. setiferus, P. stylirostris and P. annamei Lester and Cook, 1987. and P. monodon Tong et al., 1995. have been suggested to result from differential gene expression through development. Two dimensional electrophoresis has been used to identify different patterns of proteins in male and female P. annamei, and has revealed tissue specific patterns, thought to be the result of differential expression of protein coding genes Cariolou and Flytzanis, 1993.. Immunological techniques have identified several genetically distinct types of collagen in P. japonicus Mizuta et al., 1992., but the nature of the genetic control of the production of these has not been investigated. Variable staining intensity of two mRNA transcripts from unknown nuclear genes in P. annamei at different larval stages and between adults from different populations was thought to indicate differential gene expression with development which was also dependent upon genetic background Garcia et al., 1996.. Immunological detection of vitellogenin and vitellin Shafir et al., 1992a,b; Tom et al., 1992. and a cortical granule peptide Bradfield et al., 1989. has revealed variable levels of these proteins at different times in the reproductive cycle and differential expression of the genes encoding these proteins has been inferred. Changes in the levels of vertebrate-like growth factor peptides detected in P. annamei, P. stylirostris and P. indicus with developmental stage suggest differential expression of the genes coding for these during development Toullec et al., 1991.. Using northern blotting, Sun 1994. identified that RNA for an MIH-like peptide is located only in eyestalk extracts and not in muscle. Increased levels of mRNA transcripts coding for presumptive vitellin during ovarian synthesis of P. semisulcatus has provided more direct evidence of variation in gene transcription Khayat et al., 1994a.. The fact that
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this transcript was found in both hepatopancreas and ovary led to the suggestion that vitellogenin and vitellin were coded for by the same gene in P. semisulcatus. In addition to interest in the prawns themselves, there has been a little work on their potential for providing molecular tools of use in molecular biology. The fact that the optimum pH for physiological activity of P. monodon alkaline phosphatase is different to that in mammals has led to suggestions that P. monodon alkaline phosphatase may provide a useful reporter gene in mammalian systems Chuang and Yang, 1990.. No report of the successful development of transgenic prawns has been published, but DNA has successfully been introduced into prawn eggs by microinjection Cadoret et al., 1991; Gendreau et al., 1991; Preston and Atkinson, 1995., and b-glucuronidase GUS. reporter genes have been expressed in prawn eggs after injection of foreign DNA Rothlisberg, 1998..
5. Reproduction 5.1. Sex determination and sex reersal Sex determination in the crustacea has been reviewed by Legrand et al. 1987. who noted that the genetic basis for sex determination had been studied in only a few species, none of which were decapods. The strong influence of environmental factors including temperature, food supply and social environment. on sex determination in some crustacean groups has been reviewed by Korpelainen 1990.. No sex chromosome has been observed in penaeids, nor any environmental sex determination reported Malecha, 1983; Korpelainen, 1990.. Experimental sex reversal has been used to identify the genetic sex determination mechanism of a number of crustacean species Legrand et al., 1987.. Implantation of androgenic gland in sexually undifferentiated female Macrobrachium has been used to produce masculinised females neomales. that were reproductively competent Sagi and Cohen, 1990.. The presence of males in the progeny from matings of the neomales with normal females suggested the female sex could not be homogametic XX. but was heterogametic WZ.. However, this result cannot be extrapolated to all decapods as sex determination mechanisms can vary considerably within crustacean groups Legrand et al., 1987.. Indeed, Legrand et al. 1987. report results from crossing two subspecies of isopod, one of which had an XXrXY mechanism and the other WWrWZ sex determination mechanism. They demonstrated that major genes were associated with sex determination XrY,WrZ. but that these effects were modified by an undetermined number of minor genes on other chromosomes. Variable results from sex manipulation experiments in Macrobrachium rosenbergii provide conflicting evidence of control mechanisms 100% male progeny suggest ZZ males, and the presence of some females in other experiments suggest the control cannot be simply WZrZZ. and suggest sex control in this species may also result from the interaction of both major and minor genes Sagi and Cohen, 1990.. There are no published reports on experimental manipulations of this type for penaeids, and their sex determination mechanism remains unknown. Little data on the developmental processes
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of penaeids has been published, but the differentiation of the genital organs and the androgenic gland in P. japonicus has been described Nakamura et al., 1992.. 5.2. Control of reproduction The minimum requirements to bring animals into breeding condition have been established for a number of penaeid species and have been reviewed by Primavera 1985., Ogle 1992. and McVey 1993.. The role of nutrient metabolism in crustacean reproduction, and the role of genetics and environment in determining penaeid reproduction and larval quality, have also been reviewed recently by Harrison 1990. and Benzie 1997., respectively. Environmental factors such as temperature, daylength and light quantity, have an influence on maturation, spawning, egg and larval quality of penaeids, and the inclusion of fresh or fresh-frozen seafood in the maturation diets of penaeid shrimp found necessary for effective maturation Primavera, 1985; Harrison, 1990; McVey, 1993.. However, a principal requirement for the production of large numbers of penaeid larvae on a precise time schedule in the prawn farming industry is removal of the eyestalk eyestalk ablation.. This removes the source of a substance inhibiting egg production, providing evidence of the importance of hormonal control in shrimp reproduction. Eyestalk ablation also removes the controls on a large number of body functions, so that females so treated divert all their energies into ever more frequent bouts of egg production, leading to a loss in egg quality and eventual death. The development of a reversible technique that would allow the production of high quality eggs over time without these side effects would be of immense benefit to the penaeid prawn culture industry. 5.3. Endocrine control Results of research on the crustacean endocrine system over the last 20 years has been extensively reviewed Cooke and Sullivan, 1982; Quackenbush, 1986; Meusy and Payen, 1988; Beltz, 1988; Charniaux-Cotton and Payen, 1988; Van-Herp, 1988; VanHerp and Payen, 1991.. The sinus gland in the eyestalk has been determined to be the source of a number of peptides affecting a range of physiology including moulting, glucose metabolism, pigment dispersion, various aspects of carbohydrate and protein metabolism, ionic regulation, respiration and reproduction. A general model of endocrine control in crustacea has been proposed Van-Herp and Payen, 1991.. Although a useful starting point from which to infer events, this model is based on information from very different crustacean groups, and direct evidence from penaeids is limited. Several vertebrate steroid hormones have been detected in penaeids and levels of some, such as 17-b-estradiol and estrone, increase in the early stages of ovary growth in P. monodon Fairs et al., 1990; Young et al., 1992a,b, 1993; Quinitio et al., 1994.. Attempts to induce spawning in penaeids by the addition of crude extracts from the mandibular organ or thoracic ganglion of ovigerous lobster or prawn containing these, and other, active compounds have been reported to induce ovarian development or
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vitellogenesis in P. japonicus Yano and Chinzei, 1987; Yano et al., 1988; Yano, 1992., P. annamei Tsukimura and Kamemoto, 1991; Yano and Wyban, 1992. and P. monodon Chan, 1991.. These results have been used to suggest that prawn brain may release a gonad stimulating hormone, which is speculated to have an antagonistic control of reproduction in tandem with vitellogenin inhibiting hormone VIH.. Attempts to isolate and characterise the neuropeptides from the eyestalk are beginning to yield results with Yang et al. 1995. reporting the amino acid sequences for crustacean hyperglycaemic hormone CHH. in P. japonicus, and a peptide with moult inhibiting hormone MIH. activity having been described by Yang et al. 1996.. As yet, VIH has not been characterised in any penaeid but the considerable homology between VIH, CHH and MIH suggests the peptide may soon be identified. However, assessing the biological activity of these peptides and their mode of action and interaction may be difficult to disentangle. This stems from their similarity, lack of knowledge of normal physiological levels and sites of action, and the possibility of cross activity between CHH MIH and VIH at various concentrations. 5.4. Molecular characterisation and regulation of itellinr itellogenin There have been many attempts to isolate and characterise vitellin andror vitellogenin from penaeids Vazquez-Boucard et al., 1986; Tom et al., 1987; Quinitio et al., 1989, 1990; Tom et al., 1992, 1993; Mendoza et al., 1993; Chang et al., 1993a,b; Chen and Chen, 1993; Chang et al., 1994; Chen and Chen, 1994.. However, variable results for the number and size of subunits detected in some species casts some doubt as to the true identity and function of these isolates. The different immunogenic reactions between isolates from one species with those from other species indicates the speciesspecific roles particular molecules may play. Both findings also emphasise the complex structural changes that occur through vitellogenesis Tom et al., 1992; Laverdure et al., 1994.. However, the extent to which the results obtained to date reflect true species differences, fundamental differences in the protein fractions which have been isolated, or variation in experimental procedure is unknown. The physiological and biochemical processes underlying egg development are best understood in P. semisulcatus where vitellin and vitellogenin have both been isolated, characterised, and shown to be synthesised in the ovary and in the hepatopancreas, respectively Fainzilber et al., 1992; Khayat et al., 1994a.. The dynamics of vitellin and vitellogenin production have also been traced throughout ovarian development Browdy et al., 1990.. Transport of vitellogenin from the hepatopancreas to the ovary via the haemolymph has been demonstrated Shafir et al., 1992a,b. and the bulk of the synthesis of these proteins shown to occur in the ovary Fainzilber et al., 1992.. Despite relatively detailed molecular characterisation of these processes in P. semisulcatus the mechanism of regulation is still unknown Khayat et al., 1994b.. There are also conflicting results concerning the site of synthesis of vitellogenin in other species which has been demonstrated to occur in the ovary in P. japonicus Yano and Chinzei, 1987. and in the hepatopancreas of P. annamei and P. semisulcatus Quackenbush, 1989; Fainzilber et al., 1992..
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5.5. Molecular characterisation of neuropeptides The cDNA encoding the moult-inhibiting hormone isolated from P. annamei showed considerable homology with other known crustacean MIH moult inhibiting hormone. sequences Sun, 1994.. The known amino acid sequences for MIH, VIH vitellogenin inhibiting hormone. and CHH crustacean hypoglycaemic hormone. in crustaceans are similar, and each appears to consist of a peptide family Keller, 1992; Subramaniam and Keller, 1993; Yang et al., 1995, 1996.. Several peptides with CHH activity and close amino acid sequence homologies to each other and other crustacean CHHs 4050% homology., all of which demonstrated some hyperglycaemic activity, have been isolated and sequenced from P. japonicus Yang et al., 1995.. Subsequently, one of these peptides, Pej-SGP-IV, which had the weakest CHH activity, was demonstrated to have strong MIH activity, inhibiting ecdysteriod synthesis Yang et al., 1996.. The homology between MIH and VIH molecules was found to be greater than that between either of these and the CHH peptides. A neurohormonal influence by dopamine on CHH has been demonstrated in P. monodon Kuo et al., 1995.. 5.6. Enironmental influences A feature of pond-reared penaeids is low reproductive success, even with eyestalk ablation, although the extent of depression varies with species Primavera, 1985; Dong, 1990; Chen et al., 1991; Lotz and Ogle, 1994; Menasveta et al., 1993, 1994.. This has been thought to be the result of genetic effects such as inbreeding depression Sbordoni et al., 1987. but more often of environmental factors, such as poor water quality or, particularly, the lack of key nutritional elements Primavera, 1985.. Prawns cannot elongate linolenic acid to PUFA or synthesise cholesterol, both of which must therefore be supplied in the diet by means of fresh seafood. to support biomembrane and steroid hormone formation, and as yolk constituents Teshima et al., 1989, 1992; Cuzon et al., 1994; Cahu et al., 1994; Cruz-Suarez et al., 1992.. Most detailed information is from both P. japonicus, in which the radioactive uptake of cholesterol, a precursor of steroid hormones, was greatest in the ovary Kanazawa et al., 1988., and in P. semiculcatus where lipid synthesis has been described in the ovaries Shenker et al., 1993.. The principal focus, therefore, has been on the female nutrition, but poor male performance has also been identified in P. annamei, P. stylirostris, P. setiferus and P. monodon Beard and Wickens, 1980; AQUACOP, 1983; Gomes and Honculada-Primavera, 1993; Chamberlain and Gervais, 1984; Alfaro and Lozano, 1993; Alfaro et al., 1993; Makinouchi and Hirata, 1995.. The fact that specially formulated feeds improved sperm quality in P. annamei Heitzmann et al., 1993., and that Pratoomchat et al. 1993. found sperm quality of pond-reared P. monodon males was no different from wild males, suggests that husbandry methods can provide adequate conditions to achieve normal male fertility levels. However, complex interactions between gametes determining fertilisation success and egg quality. may occur as evidenced by lower egg production in matings between wild female and cultured male P. monodon than matings between cultured males and females Menasveta et al., 1993.. The importance of nutritional and hormonal balance during the juvenile phase, when reproductive organs
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are forming, in determining the reproductive capacity of both male and female prawns in adulthood Nakamura et al., 1992., and of having a mature hormonal balance for high sperm quality in P. stylirostris Alfaro, 1993. has also been identified but little studied. Stress may also reduce reproductive rate and gamete quality in male and female penaeid prawns but there is no firm data on its effects on titre levels of vitellogenin, or of the hormonal mechanisms regulating these Chen et al., 1991; McVey, 1993.. Stress might act through changing levels of key hormones regulating reproduction and development either directly, or indirectly through competition for precursor molecules. The close molecular relationship suggested by the limited data for hormones controlling moulting, reproduction, mineral metabolism and stress response in prawns raises questions about their inter-relationships and regulation at the gene and physiological levels Quinitio et al., 1993.. However, specific information on the role of particular hormones, and indeed the identity of these, in penaeids, is almost non-existent. There is little published on the detailed physiological effects of stress in penaeids although Bishop and Burton 1993. give some information on changes in amino acid synthesis in hyperosmotically stressed P. aztecus.
6. Growth While there have been innumerable studies measuring the growth rate of penaeids in production systems, the fundamental controls on growth are largely unstudied. Many peptides related to vertebrate growth factors have been isolated from crustaceans, but few have been observed from penaeids as yet Toullec et al., 1991.. Vertebrate-like growth factor peptides have been detected in P. annamei, P. stylirostris and P. indicus, and their levels shown to change with developmental stage Toullec et al., 1991.. Supplementation of the diet of P. annamei larvae with human growth hormone had a positive effect on growth of the larvae and on their resistance to salinity stress. The role of moult and juvenile hormones JH. and JH-like molecules such as methyl farneosate in growth, and their possible effects on reproduction, have yet to be elucidated Laufer and Sagi, 1991..
7. Tissue and cell culture Primary cell cultures derived from several tissues have been made but no immortal cell lines have yet been established, with the exception of prawn cells transformed with SV-40 Bols, 1991; Tapay et al., 1995.. Explant outgrowth from haematopoietic tissues, primary lymphoid cells, and ovaries have been used to obtain cell cultures maintained on Leibovitzs L-15 basal medium supplemented by fetal bovine serum and prawn haemolymph Chen et al., 1988, 1989; Rosenthal and Diamant, 1990; Lu et al., 1995. and lately, with some modifications, have been successfully applied to P. annamei and P. indicus Toullec et al., 1996.. A range of media has been tested for P. chinensis Tong and Miao, 1996.. These cell cultures, and whole tissue biopsies have been used to
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assay extract activities on the ovary, and to assess the effects of viruses on prawn cells e.g., Browdy et al., 1990; Shenker et al., 1993.. The primary cell cultures suffer from the difficulty that the longest they have been maintained is several months through 23 passages., but sometimes only for 3 weeks. This leads to considerable difficulties in obtaining consistent assays of viral effects, and markedly increases work loads Lu et al., 1995.. Toullec et al. 1996. review the most recent information on primary cell cultures, and Sano 1998. describes examples of tissue organisation in primary cell cultures.
8. Disease Prawns are affected by a number of disease organisms including viruses, bacteria, protozoans, microsporidians, gregarines and haplosporidians, nematodes and cestodes McVey, 1993; Lightner, 1993, 1998.. The diagnosis of most of these diseases still depends largely on anatomical and histological examination. The lack of molecular probes has considerably hampered the rapidity of diagnosis, and prevented the development of a deeper understanding of the epidemiology of bacterial and, particularly, viral diseases Lightner et al., 1983; Miahle et al., 1992.. The lack of prawn cell lines has also slowed the isolation and characterisation of prawn viruses Mari et al., 1993; Lu et al., 1995; Tapay et al., 1995.. Infectious hypodermal and hematopoietic necrosis virus IHHNV. was characterised molecularly several years after it was first described morphologically Bonami et al., 1990., and a probe developed only after ten years had elapsed Mari et al., 1993.. Molecular probes are commercially available for only two of the eight described prawn viruses, IHHNV and type-A baculovirus BP.. However, a number of research groups have completed the early stages of development of molecular probes for systemic ectodermal and mesodermal baculovirus white spot. SEMBV., monodon baculovirus MBV., taura virus TSV., hepatopancreatic parvovirus HPV. and yellowhead virus YHV. Chang et al., 1993a,b.. DNA-based, PCR and immunological probes have been developed for white spot disease Wongteerasupaya et al., 1996. and are being developed for other viruses and bacteria Vibrio spp... Gene sequence similarities and cross reaction of BP polyhedrin with Autographa californica nuclear polyhedrosis virus AcNPV. antisera suggest some relationship of the BP prawn virus with insect viruses Vickers et al., 1994.. However, the last decade has seen a large increase in the number of viruses isolated from crustaceans and penaeids in particular, and most of these are uncharacterised Lightner, 1998.. For example, a virus originally isolated from Australian P. monodon and thought originally to be IHHNV Owens et al., 1992. does not react with the IHHNV probe developed from american strains, and may be an entirely different virus. The increasing application of molecular techniques, and a concerted effort on the virology of penaeids will lead to more rapid diagnosis and a greater understanding of the epidemiology of these diseases and therefore to more rapid development of effective control strategies. The physiological response to diseases in shrimp are also relatively unknown. Although they demonstrate an enhanced response to diseases to which they have already been exposed, they do not have a vertebrate cell-mediated immunological response but a
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prophenoloxidase proPO. activating system for nonself recognition system has been described McVey, 1993; Sung et al., 1996.. It is this system that may be activated by the addition of immunostimulants. The enhancement of microbiocidal activity following the addition of immunostimulants has been described in P. monodon Sung et al., 1996.. Two lectins and one agglutinin have been characterised for Parapenaeus longirostris Fragkiadakis and Stratakis, 1995. and were speculated to contribute to defense against specific microorganisms.
9. Discussion The principal advantages arising from the application of genetics and biotechnology to aquaculture are usually identified as optimization of the production cycle through control of development and maturation, gamete production and storage, sex and ploidy manipulation gynogenesis, androgenesis, polyploidy and sex reversal., transgenesis, improved disease diagnosis, control of diseases through better vaccine production, selective breeding and marker assisted selection Maclean and Penman, 1990; Bols, 1991; Hedgecock and Malecha, 1991.. Key elements in achieving these goals are specific applications of techniques of gene manipulation, immunological and biochemical methods for the isolation, characterisation, detection and manipulation of molecules and their effective introduction into whole animals, and cell culture. The advantages of such approaches are clearly demonstrated in animal agriculture McConnell, 1986; Ward and Nancarrow, 1991; Muller and Brem, 1991.. The advances in mammalian and vertebrate systems has resulted from the large arsenal of molecular and biochemical tools, and the deep background knowledge of the genome, cell biology, development and physiology available for these groups McConnell, 1986; Bols, 1991.. The relatively rapid advances in fish genetics and biotechnology has stemmed from the ability to use a number of the specific tools developed for mammalian systems. These have been used either directly or, because of strong homologies in genetic and developmental systems between fish and vertebrates, for the rapid probing and development of specific fish products Houdebine and Chourrout, 1991; Du et al., 1992.. Considerable advances in basic technologies such as mass spectrometry, 2-D electrophoresis, gene cloning, and monoclonal antibody production make the development of new molecular tools easier than in the past Van-Herp and Payen, 1991, Van-Herp, 1993.. It will, however, take time to develop effective tools on penaeids Hedgecock and Malecha, 1991.. The lack of knowledge on the genetics, physiology, molecular and cellular biology of penaeids means work has had to begin almost from scratch. While the goals may be extremely applied, as in for example the development of hormonal controls on prawn reproduction, the capacity to use this powerful method to assist farmers will demand fundamental work on the penaeid endocrine system. An understanding of disease control mechanisms will demand a focus on the basic cell biology of penaeids. Sex manipulation is often discussed in terms of producing single sex populations to improve production by allowing culture of only the faster growing sex, or to increase economic returns through the more consistent size of the single-sex crop produced. However, sex manipulation might be far more important as an experimental
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tool in the development of other prawn biotechnologies Hedgecock and Malecha, 1991.. The determination of genes affecting key production characteristics, including disease resistance, would be greatly speeded by the ability to produce lines of genetically identical prawns through gynogenesis andror androgenesis, and associated ploidy manipulation to create haploid lines. The ability to reverse sex, a key part of producing genetically identical lines, will depend on the extent to which the factors controlling sex and reproduction are understood. Until such time as assays are developed for key hormones and their receptors, the role hormones play in reproduction and development, and the nature of their interaction with environmental variables and the genotype cannot be dissected. The interdependence of technical advances in any one area of penaeid biology is further illustrated by the fact that the effective maintenance of transgenic populations demands the ability to reproduce animals reared in culture, and to have a sound quantitative breeding program within which to place particular genes and assess their performance. The potential of linking interspecies hybridisation and ploidy manipulation to create animals extremely unlikely to be fertile because of their triploidy, and because of hybrid sterility. has been highlighted as a means of reducing the potential impact of finfish escapees Scheerer et al., 1987.. The ability of molecular technologies to track the provenance of animals through genetic markers, and the ability to develop means of making production stock sterile through ploidy manipulation, will become important in determining the impact of aquaculture operations on wild stocks which is of growing concern Skaala et al., 1990; Bentsen, 1991.. This review of the published literature has illustrated the fragmented and preliminary nature of the data available on the genetics of, and biotechnological applications for, penaeids. However, there is much work in progress. Research aimed at isolating and characterising penaeid hormones, vitellogenin and vitellin in order to elucidate their role in reproduction and lipid metabolism is taking the first steps towards characterising key active compounds. Immunological probes are already being used to identify the sites of vitellogenin synthesis and accumulation. The fact that stocks based on specific matings in which family pedigrees are traced are now available, provides a basis for mapping quantitative traits, and their association with molecular markers, something not possible in mass selection programs. Comparison with agricultural programs demonstrates how little has yet been done with shrimp, and the considerable amount of work that lies ahead to obtain the large number of markers required for effective attempts at marker assisted selection. Nevertheless, the finding that a mtDNA COI gene variant is associated with high growth, at least in the populations of P. annamei held by the US Consortium Program, is a promising start. Work on the isolation and characterisation of genes controlling growth and reproduction will first provide strong experimental tools to assist the identification of the sites of synthesis of key molecules, and in the longer term may provide technologies for control of reproduction and disease. The practical application of many of these approaches may seem far off to prawn farmers struggling with the day to day issues of production, where more immediate solutions may be sought in changing husbandry methods. However, ad hoc changes to husbandry systems are unlikely to alleviate the loss of wild supplies of spawners and the devastating effects of disease. These events have focused attention on the vulnerability
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of wild supplies, and the need to develop domesticated stock that are either disease free or disease resistant. This has focused attention on the necessity of research on fundamental aspects of the biology of penaeid prawns in order to provide practical solutions to these problems, as evidenced by the papers in this symposium. As a result it is likely that significant advances will be made in the next decade as molecular and biotechnological tools become available to dissect the genome and biology of penaeids. Acknowledgements I thank M. Hall, K. Wilson, M. Thurn and M. Kenway for comments on drafts of the manuscript. This is contribution no. 861 from the Australian Institute of Marine Science. References
Alfaro, J., 1993. Reproductive quality evaluation of male Penaeus stylirostris from a grow-out pond. J. World Aquacult. Soc. 24, 611. Alfaro, J., Lozano, X., 1993. Development and deterioration of spermatophores in pond-reared Penaeus annamei. J. World Aquacult. Soc. 24, 522529. Alfaro, J., Lawrence, A.L., Lewis, D., 1993. Interaction of bacteria and male reproductive system blackening disease of captive Penaeus setiferus. Aquaculture 117, 18. AQUACOP, 1983. Constitution of broodstock, maturation, spawning and hatching system for penaeid shrimps in the Centre Oceanologique du Pacifique. In: McVey, J.P. Ed.., CRC Handbook of Mariculture, Vol. 1: Crustacean Aquaculture. CRC Press, Miami, FL, USA, pp. 105121. AQUACOP, Ledu, C., Diter, A., 1993. Induction of polyploid nauplii in Penaeus indicus. Aquaculture, 111, 315 abstract.. Beard, T.W., Wickens, J.F., 1980. Breeding of Penaeus monodon Fabricius in laboratory recirculation systems. Aquaculture 20, 7989. Bedier, E., Ottogalli, L., Patrois, J., Weppe, M., AQUACOP, 1996. Development of the Penaeus stylirostris Aquacop SPR43 strain: in situ experiments and genetic enhancement. World Aquaculture 96. Book of Abstracts, World Aquaculture Society, pp. 3233 Beltz, B.S., 1988. Crustacean neurohormones. In: Laufer, H., Downer, R.G.H. Eds.., Endocrinology of Selected Invertebrate Types: Invertebrate Endocrinology, Vol. 2. Alan R. Liss, New York, USA, pp. 235258. Bentsen, H.B., 1991. Quantitative genetics and management of wild populations. Aquaculture 98, 263266. Benzie, J.A.H., 1994. Genetic and reproduction research on giant tiger prawns Penaeus monodon.: pond-reared spawners achieved in Australia. Australasian Biotech. 4, 222224. Benzie, J.A.H., 1996. The genetic structure of cultured marine species in the Indo-Pacific: implications for aquaculture and biodiversity. In: Penman, D.J., Pongthana, N., McAndrew, B., Bell, A. Eds.., Proceedings of the Second ASEAN-EEC Aquaculture Development and Coordination Program, AADCP, Phuket, Thailand. Benzie, J.A.H., 1997. A review of the effect of genetics and environment on the maturation and larval quality of the giant tiger prawn, Penaeus monodon. Aquaculture 155, 6985. Benzie, J.A.H., Frusher, S., Ballment, E., 1992. Geographical variation in allozyme frequencies of Penaeus monodon Crustacea: Decapoda. populations in Australia. Aust. J. Mar. Freshwat. Res. 43, 715725. Benzie, J.A.H., Ballment, E., Frusher, S., 1993. Genetic structure of Penaeus monodon in Australia: preliminary data from allozymes and mtDNA. Aquaculture 111, 8993. Benzie, J.A.H., Kenway, M., Ballment, E., Frusher, S., Trott, L., 1995. Interspecific hybridization of the tiger prawns Penaeus monodon and Penaeus esculentus. Aquaculture 133, 103111. Benzie, J.A.H., Kenway, M., Trott, L., 1997. Estimates for the heritability of size in juvenile P. monodon from half-sib matings. Aquaculture 152, 4953.
40
Bishop, J.S., Burton, R.S., 1993. Amino acid synthesis during hyperosmotic stress in Penaeus aztecus postlarvae. Comp. Biochem. Physiol. A 106, 4956. Bols, N.C., 1991. Biotechnology and aquaculture: the role of cell cultures. Biotechnol. Adv. 9, 3149. Bonami, J.R., Trumper, R., Mari, J., Brehelin, M., Lightner, D.V., 1990. Purification and characterization of the infectious hypodermal and haematopoietic necrosis virus of penaeid shrimps. J. Gen. Virol. 71, 26572664. Bouchon, D., Souty-Grosset, C., Raimond, R., 1994. Mitochondrial DNA variation and markers of species identity in two penaeid shrimp species: Penaeus monodon Fabricius and P. japonicus Bate. Aquaculture 127, 131144. Bradfield, J.Y., Berlin, R.L., Rankin, S., Keeley, L.L., 1989. Cloned cDNA and antibody for an ovarian cortical granule polypeptide of the shrimp Penaeus annamei. Biol. Bull. 177, 344349. Bray, W.A., Lawrence, A.L., Lester, L.J., 1990a. Reproduction of eyestalk-ablated Penaeus stylirostris fed various levels of total dietary lipid. J. World Aquacult. Soc. 21, 4152. Bray, W.A., Lawrence, A.L., Lester, L.J., Smith, L.L., 1990b. Hybridization of Penaeus setiferus and Penaeus schmitti. J. Crustacean Biol. 10, 278283. Browdy, C.L., Fainzilber, M., Loya, T.Y., Lubzens, E., 1990. Vitellin synthesis in relation to oogenesis. In vitro-incubated ovaries of Penaeus semisulcatus Crustacea Decapoda, Penaeidae.. J. Exp. Zool. 255, 205215. Cadoret, J.P., Mialhe, E., Gendreau, S., Ohresser, M., Le Deuff, R.M., 1991. Gene transfer: potential application to pathology of farmed marine invertebrates. International Council for the Exploration of the Sea, Mariculture Communication. ICES 1991 F:21. 5 pp. Cahu, C., Guillaume, J.C., Stephan, G., Chim, L., 1994. Influence of phospholipid and highly unsaturated fatty acids on spawning rate and egg and tissue composition in P. annamei fed semi-purified diets. Aquaculture 126, 159170. Cariolou, M.A., Flytzanis, C.N., 1993. Specific gene expression in distinct tissues of the shrimp Penaeus annamei. Comp. Biochem. Physiol. B 106, 705716. Carr, W.H., Fjalestad, K.T., Godin, D. Swingle, J., Sweeney, J.N., Gjedrem, T., 1996a. Genetic variation in weight and survival in a population of specific pathogen-free shrimp, Penaeus annamei. World Aquaculture 96, Book of Abstracts. World Aquaculture Society, p. 63. Carr, W.H., Sweeney, J.N., Nunan, L., Lightner, D.V., Hirsch, H., Reddington, J.J., 1996b. The use of an infectious hypodermal and hematopoietic necrosis virus gene probe serodiagnostic field kit for the screening of candidate specific pathogen-free Penaeus annamei broodstock. Aquaculture 147, 18. Chamberlain, G.W., Gervais, N.F., 1984. Comparison of unilateral eyestalk ablation with environmental control for ovarian maturation. J. World Maricult. Soc. 15, 2930. Chan, H.H., 1991. Effects of eyestalk ablation, thoracic ganglion extract and gonad extract from spent spawners on ovarian maturation in pond-reared shrimps Penaeus monodon Fabricius. Aquacult. Fish. Manage. 22, 463471. Chang, C.F., Lee, F.Y., Huang, Y.S., 1993a. Purification and characterization of vitellin from the mature ovaries of prawn Penaeus monodon. Comp. Biochem. Physiol. B 105, 409414. Chang, P.-S., Lo, C.-F., Kou, G.-H., Lu, C.-C., Chen, S.-N., 1993b. Purification and amplification of DNA from Penaeus monodon-type baculovirus. J. Invertebr. Pathol. 62, 116120. Chang, C.F., Lee, F.Y., Huang, Y.S., Hong, T.H., 1994. Purification and characterization of the female-specific protein vitellogenin. in mature female hemolymph of the prawn Penaeus monodon. Invertebr. Reprod. Dev. 25, 185192. Charniaux-Cotton, H., Payen, G.G., 1988. Crustacean reproduction. In: Laufer, H., Downer, R.G.H. Eds.., Endocrinology of Selected Invertebrate Types: Invertebrate Endocrinology, Vol. 2. Alan R. Liss, New York, USA, pp. 279303. Chen, C.C., Chen, S.N., 1993. Isolation and partial characterization of vitellin from the egg of giant tiger prawn Penaeus monodon. Comp. Biochem. Physiol. B 106, 141146. Chen, C.C., Chen, S.N., 1994. Vitellogenesis in the giant tiger prawn Penaeus monodon Fabricius, 1789. Comp. Biochem. Physiol. B 107, 453460. Chen, S.N., Jong, K.J., Kou, G.H., 1988. Cell cultures from hematopoietic tissue and ovary of penaeid shrimp, Penaeus monodon. In: Kuroda, J., Kurstak, E., Maramorosch, K., Eds.., Invertebrate and Fish Tissue Culture. Springer-Verlag, New York, pp. 195198.
41
Chen, S.N., Jong, K.J., Kou, G.H., 1989. Cell cultures derived from tissue of penaeid shrimp, Penaeus penicillatus and hard clam, Metrix lusoria. In: Mitsuhashi, J. Ed.., Invertebrate Cell System Applications, Vol. 2. CRC Press, Baton Rouge, USA, pp. 253262. Chen, F., Reid, B., Arnold, C.R., 1991. Maturing, spawning and egg collecting of the white shrimp Penaeus annamei Boone in a recirculating system. J. World Aquacult. Soc. 22, 167172. Chow, S., Sandifer, P.A., 1991. Differences in growth, morphometric traits and male sexual maturity among Pacific white shrimp, Penaeus annamei, from different commercial hatcheries. Aquaculture 92, 165178. Chow, S., Dougherty, W.J., Sandifer, P.A., 1990. Meiotic chromosome complements and nuclear DNA contents of four species of shrimps of the genus Penaeus. J. Crustacean Biol. 10, 2936. Chuang, N.-N., Yang, B.-C., 1990. A comparative study of alkaline phophatases among human placenta, bovine milk, hepatopancreases of shrimp Penaeus monodon Crustacea: Decapoda. and clam Meretrix lusoria Bivalvia: Veneidae.: to obtain an alkaline phosphatase with improved characteristics as a reporter. Comp. Biochem. Physiol. B 96, 787789. Clark, W.H., Griffin, F.J., 1993. Acquisition and manipulation of penaeoidean gametes. In: McVey, J.P. Ed.., CRC Handbook of Mariculture, Vol. I: Crustacean Aquaculture. CRC Press, Boca Raton, FL, USA, pp. 133151. Cooke, I.M., Sullivan, R.E., 1982. Hormones and neurosecretion. In: Atwood, H.L., Sandeman, D.C. Eds.., The Biology of Crustacea, Vol. 3. Academic Press, New York, USA, pp. 205289. Cruz-Suarez, L.E., Ricque, D., AQUACOP, 1992. Effect of squid meal on growth of Penaeus monodon juveniles reared in pond pens and tanks. Aquaculture, 106, 293299. Cuzon, G., Guillaume, J., Cahu, C., 1994. Composition, preparation and utilization of feeds for Crustacea. Aquaculture 124, 253267. Dai, J., Zhang, Q., Bao, Z., 1989. Karyotype studies on Penaeus orientalis. J. Ocean. Univ. Qingdao 19, 97103. Dai, J., Bao, Z., Zhang, Q., Liu, J., 1993. An observation of gynogenesis induced by 49 Co g rays in chinese prawn Penaeus chinensis. J. Ocean. Univ. Qingdao 23, 151155. Dong, Z., 1990. Overwintering and sexual maturation of Penaeus penicillatus Alcock in an outdoor earth pond. Aquaculture 86, 327331. Du, S.S., Gong, Z., Fletcher, G.L., Shears, M.A., King, M.J., Idler, D.R., Hew, C.L., 1992. Growth enhancement of transgenic Atlantic salmon by the use of an all Fish chimeric growth hormone gene construct. BiorTechnology 10, 176180. Fainzilber, M., Tom, M., Shafir, S., Applebaum, S.W., Lubzens, E., 1992. Is there extraovarian synthesis of vitellogenin in penaeid shrimp?. Biol. Bull. 183, 233241. Fairs, N.J., Quinlan, P.T., Goad, L.J., 1990. Changes in ovarian unconjugated and conjugated steroid titers during vitellogenesis in Penaeus monodon. Aquaculture 89, 8399. Fragkiadakis, G., Stratakis, E., 1995. Characterization of hemolymph lectins in the prawn Parapenaeus longirostris. J. Invertebr. Pathol. 65, 111117. Furukohri, T., Okamoto, S., Suzuki, T., 1994. Evolution of phosphagen kinase: III. Amino acid sequence of arginine kinase from the shrimp Penaeus japonicus. Zool. Sci. 11, 229234. Garcia, D.K., Benzie, J.A.H., 1995. RAPD markers of potential use in penaeid prawn Penaeus monodon. breeding programs. Aquaculture 130, 137144. Garcia, D.K., Faggart, M.A., Rhoades, L., Alcivar-Warren, A., Wyban, J., Carr, W.H., Sweeney, J.N., Ebert, K.M., 1994. Genetic diversity of cultured Penaeus annamei shrimp using three molecular genetic techniques. Mol. Mar. Biol. Biotechnol. 3, 270280. Garcia, D.K., Dhar, A.K., Alcivar-Warren, A., 1996. Molecular analysis of a RAPD marker B20. reveals two microsatellites and differential mRNA expression in Penaeus annamei. Mol. Mar. Biol. Biotechnol. 5, 7183. Gendreau, S., Delecheneau, J.M., Cadoret, J.P., Mialhe, E., 1991. Methodology for microinjection of embryos of the shrimp Penaeus indicus. European Aquaculture Society Special Publication No 14, pp. 115116 abstract.. Gomes, L.A.O., Honculada-Primavera, J., 1993. Reproductive quality of male Penaeus monodon. Aquaculture 112, 157164. Goswami, U., 1985. Chromosomal studies in Penaeus aztecus Ives prawn larvae. Mahasagar Bull. Natl. Inst. Oceanol. 18, 7577.
42
Goswami, U., Goswami, S.C., Dalal, S.G., 1990. Morphometric studies in Penaeus merguiensis from Ratnagiri waters for selection of the broodstock in genetic improvement programmes. J. Ind. Fish. Assoc. 20, 15. Harrison, K.E., 1990. The role of nutrition in maturation, reproduction and embryonic development of decapod crustaceans: a review. J. Shellfish Res. 9, 128. Hayashi, K., Fujiwara, Y., 1988. A new method for obtaining metaphase chromosomes from the regeneration blastema of Penaeus Marsupenaeus. japonicus. Nippon Suisan Gakkaishi 54, 15631565. Hedgecock, D., Malecha, S.R., 1991. Prospects for the application of biotechnology to the development and improvement of shrimp and prawns. In: Sandifer, P.A. Ed.., Shrimp culture in North America and the Caribbean. Advances in World Aquaculture, Vol. 4. World Aquaculture Society, Baton Rouge, LA, USA, pp. 161200. Hedgecock, D., Tracey, M.L., Nelson, K., 1982. Genetics. In: Abele, L.G. Ed.., The Biology of Crustacea. Academic Press, New York, USA, pp. 284403. Heitzmann, J.-C., Diter, A., AQUACOP, 1993. Spermatophore formation in the white shrimp, Penaeus annamei Boone 1931: dependence on intermoult cycle. Aquaculture, 116, 9198. Houdebine, L.M., Chourrout, D., 1991. Transgenesis in fish. Experientia 47, 891897. Huang, C.M., Tsai, M.Y., Liao, I.C., 1990. Morphometric studies on choice of broodstock for genetic improvement in Penaeus penicillatus. In: Hirano, R., Hanyu, I. Eds.., The Second Asian Fisheries Forum. Asian Fisheries Society, Manila, Philippines, pp. 523526. Kanazawa, A., Chim, L., Laubier, A., 1988. Tissue uptake of radioactive cholesterol in the prawn Penaeus japonicus Bate during induced ovarian maturation. Aquat. Living Resour. 1, 8591. Keller, R., 1992. Crustacean neuropeptides: structures functions and comparative aspects. Experientia 48, 439448. Khayat, M., Lubzens, E., Tietz, A., Funkstein, B., 1994a. Are vitellin and vitellogenin coded by one gene in the marine shrimp Penaeus semisulcatus?. J. Mol. Endocrinol. 12, 251254. Khayat, M., Lubzens, E., Tietz, A., Funkstein, B., 1994b. Cell-free synthesis of vitellin in the shrimp Penaeus semisulcatus de Haan.. Gen. Comp. Endocrinol. 93, 205213. Kim, W., Abele, L.G., 1990. Molecular phylogeny of selected decapod crustaceans based on 18S rRNA nucleotide sequences. J. Crustacean Biol. 10, 113. Koh, G.L., Pasteur, N., Bonhomme, F., AQUACOP, SEAFDEC, Liao, D., 1983. Variabilite genetique de quelques especes de crevettes peneides d interet aquicole. Bases biologiques de laquaculture, Montpellier, 1983. Actes de Colloques, Vol. 1, 175180. Kong, F., Zhang, D., 1993. The karyotype of the tiger shrimp Penaeus monodon. J. Fish. China 17, 8384. Korpelainen, H., 1990. Sex ratios and conditions required for environmental sex determination in animals. Biol. Rev. 65, 147184. Kuo, C-M., Hsu, C.-R., Lin, C.-Y., 1995. Hyperglycaemic effects of dopamine in tiger shrimp Penaeus mondon. Aquaculture 135, 161172. Laubier, A., Laubier, L., 1993. Marine crustacean farming: present status and perspectives. Aquat. Living Resour. 6, 319329. Laubier, A., Pasteur, N., Moriyasu, M., 1984. Estimation du polymorphisme enzimatique dune population de Penaeus japonicus maintenue en elevage depuis quatre generations. Oceanol. Acta 7, 451456. Laufer, H., Sagi, A., 1991. Juvenile hormone-like compounds and reproduction in male and female crustaceans: with implications for aquaculture. Bull. Inst. Zool. Academia Sinica, Monogr. 16, 541551. Laverdure, A-M., Carette-Desmoucelles, C., Breuzet, M., Descamps, M., 1994. Neuropeptides and related nucleic acid sequences detected in penaeid shrimps by immunohistochemistry and molecular hybridizations. Neuroscience 60, 569579. Lawrence, A.L., Bray, W.A., Wilkenfield, J.S., Lester, L.J., 1984. Successful interspecific crosses of two species of marine shrimp, Penaeus stylirostrus and Penaeus setiferus. World Maricult. Soc. 1984, 39, abstract. Legrand, J.J., Legrand-Hamelin, E., Juchault, P., 1987. Sex determination in crustacea. Biol. Rev. 62, 439470. Lester, L.J., 1979. Population genetics of penaeid shrimp from the Gulf of Mexico. J. Hered. 70, 175180. Lester, L.J., 1983. Developing a selective breeding program for penaeid shrimp mariculture. Aquaculture 33, 4150.
43
Lester, L.J., 1988. Difference in larval growth among families of Penaeus stylirostris Stimpson and Penaeus annamei Boone. Aquacult. Fish. Manage. 19, 243251. Lester, L.J., Cook, J.P., 1987. Ontogenic changes in isozyme patterns of Penaeus species. Comp. Biochem. Physiol. B 86, 253258. Lester, L.J., Lawson, K.S., 1990. Inheritance of size as estimated by principal component analysis at two temperatures in Penaeus annamei. Aquaculture 85, 323, abstract. Liao, I.C., 1990. The worlds marine prawn culture industries: today and tomorrow. In: Hirano, R., Hanyu, I. Eds.., The Second Asian Fisheries Forum. Asian Fisheries Society, Manila, Philippines, pp. 1127. Lightner, D.V., 1993. Diseases of cultured penaeid shrimp. In: McVey, J.P. Ed.., CRC Handbook of Mariculture, Vol. I: Crustacean Aquaculture. CRC Press, Boca Raton, FL, USA. pp. 393486. Lightner, D.V., 1998. Shrimp diseases and current diagnostic methods. Aquaculture 164, 201220. Lightner, D.V., Redman, R.M., Bell, T.A., 1983. Observations on the geographic distribution, pathogenesis and morphology of the baculovirus from Penaeus monodon Fabricius. Aquaculture 32, 209233. Lin, M-N., Ting, Y-Y., Hanyu, I., 1988. Hybridization of two close-thelycum penaeid species Penaeus monodon= P. penicillatus and P. penicillatus= P. monodon, by means of spermatophore transplantation. Bull. Taiwan Fish. Res. Inst. 45, 83101. Lotz, J.M., Ogle, J.T., 1994. Reproductive performance of the white-legged shrimp Penaeus annamei in recirculating systems. J. World Aquacult. Soc. 25, 477482. Lu, Y., Tapay, L.M., Loh, P.C., Brock, J.A., Gose, R., 1995. Development of a quantal assay in primary shrimp cell culture for yellowhead baculovirus YBV. of penaeid shrimp. J. Virol. Meth. 52, 231236. Machado, E.G., Dennebouy, N., Suarez, M.O., Mounolou, J.-C., Monnerot, M., 1993. Mitochondrial 16S-rRNA gene of two species of shrimps: sequence variability and secondary structure. Crustaceana 65, 279286. Maclean, N., Penman, D., 1990. The application of gene manipulation to aquaculture. Aquaculture 85, 120. Makinouchi, S., Hirata, H., 1995. Studies on maturation and reproduction of pond-reared Penaeus monodon for developing a closed-cycle culture system. Isr. J. Aquacult.-Bamidgeh 47, 6877. Malecha, S.R., 1983. Crustacean genetics and breeding: an overview. Aquaculture 33, 395413. Malecha, S.R., Hedgecock, D., 1989. Prospects for the domestication and breeding of marine shrimp. Sea Grant Technical Report UNIHI-SEAGRANT-TR-89-01. University of Hawaii Sea Grant College Program, Honolulu, USA, 40 pp. Mari, J., Bonami, J.-R., Lightner, D.V., 1993. Partial cloning of the genome of infectious hypodermal and haematopoietic necrosis virus, an unusual parvovirus pathogenic for penaeid shrimps; diagnosis of the disease using a specific probe. J. Gen. Virol. 74, 26372643. Mattoccia, M., La Rosa, G., De Matthaeis, E., Cobolli-Sbordoni, M., Sbordoni, V., 1987. Patterns of genetic variability in Mediterranean populations of Penaeus kerathurus Crustacea, Decapoda.. In: Tiews, K. Ed.., Selection, Hybridization and Genetic Engineering in Aquaculture. Heenemann Verlag, Berlin, pp. 131142. Mayorga, Z., 1982. Genetica de crustaceos. Documenta 10, 36. McConnell, D.J., 1986. Methods and achievements of genetic engineering: prospects in agriculture, a review. Livest. Prod. Sci. 15, 109131. McVey, J.P. Ed.., 1993. CRC Handbook of Mariculture, Vol. I: Crustacean Aquaculture. CRC Press, Boca Raton, FL, USA, 526 pp. Menasveta, P., Piyatiratitivorakul, S., Rungsupa, S., Moree, N., Fast, A.W., 1993. Gonadal maturation and reproductive performance of giant tiger prawn Penaeus monodon Fabricius. from the Andaman Sea and pond-reared sources in Thailand. Aquaculture 116, 191198. Menasveta, P., Sangpradub, S., Piyatiratitivorakul, S., Fast, A.W., 1994. Effect of broodstock size and source on ovarian maturation and spawning of Penaeus monodon Fabricius from the Gulf of Thailand. J. World Aquacult. Soc. 25, 4149. Mendoza, R., Guillaume, J.-C., Fauvel, C., 1993. Homologous ELISA procedure for determination of penaeid shrimp vitellogenin. Aquat. Living Resour. 6, 3948. Meusy, J-J., Payen, G.G., 1988. Female reproduction in malacostracan crustacea. Zool. Sci. 5, 217265. Miahle, E., Boulo, V., Bachere, E., Hervio, D., Cousin, K., Noel, D., Noel, T., Ohresser, M., le Deuff, R.M., Despres, B., Gendreau, S., 1992. Development of new methodologies for diagnosis of infectious diseases in mollusc and shrimp aquaculture. Aquaculture 107, 155164. Milligan, D.J., 1976. A method for obtaining metaphase chromosome spreads from marine shrimp with notes
44
on the karyotypes of Penaeus aztecus, P. setiferus and P. duorarum. Proc. World Maricult. Soc. 7, 327332. Misamore, M., Browdy, C.L., 1994. Evaluation of the potential for hybridization between two litopenaeid species, Penaeus annamei and Penaeus setiferus. World Aquacult. Soc 1994, 85, Abstract. Mizuta, S., Yoshinaka, R., Sato, M., Suzuki, T., Sakaguchi, M., 1992. Immunohistochemical localization of genetically distinct types of collagen in muscle of kuruma prawn Penaeus japonicus. Comp. Biochem. Physiol. B 103, 917922. Muller, M., Brem, G., 1991. Disease resistance in farm animals. Experientia 47, 923934. Mulley, J.C., Latter, B.D.H., 1980. Genetic variation and evolutionary relationships within a group of thirteen species of penaeid prawns. Evolution 34, 904916. Mulley, J.C., Latter, B.D.H., 1981a. Geographic differentiation of eastern Australian prawn populations. Aust. J. Mar. Freshwat. Res. 32, 889895. Mulley, J.C., Latter, B.D.H., 1981b. Geographic differentiation of tropical Australian prawn populations. Aust. J. Mar. Freshwat. Res. 32, 895906. Murofushi, M., Deguchi, Y., 1990. Karyotype evolution in Decapoda, Crustacea. In: Hirano, R., Hanyu, I. Eds.., The Second Asian Fisheries Forum. Asian Fisheries Society, Manila, Philippines, pp. 549553. Nakamura, K., Matsuzaki, N., Yonekura, K., 1992. Organogenesis of genital organs and androgenic gland in the kuruma prawn. Nippon Suisan Gakkaishi 58, 22612267. Naner, C., Feng, L., 1993. Artificial induction of gynogenesis in Chinese shrimp, Penaeus orientalis Kishinouye. Aquaculture 111, 314, abstract. Niiyama, H., 1948. Oguma Commemoration volume on Cytology and Genetics, Part I. Hoppo Syuppansha, Sapporo, pp. 1920. Ogle, J.T., 1992. A review of the current 1992. state of our knowledge concerning reproduction in open thelycum penaeid shrimp with emphasis on Penaeus annamei. Invertebr. Reprod. Dev. 22, 267274. Owens, L., Anderson, I.G., Kenway, M., Trott, L., Benzie, J.A.H., 1992. Infectious hypodermal and haematopoietic necrosis virus IHHNV. in a hybrid penaeid prawn from tropical Australia. Dis. Aquat. Org. 14, 219228. Palumbi, S.R., Benzie, J.A.H., 1991. Large mitochondrial DNA differences between morphologically similar penaeid shrimp. Mol. Mar. Biol. Biotechnol. 1, 2734. Pratoomchat, B., Piyatiratitivorakul, S., Menasveta, P., Fast, A.W., 1993. Sperm quality of pond-reared and wild-caught Penaeus monodon in Thailand. J. World Aquacult. Soc. 24, 530540. Preston, N., Atkinson, P., 1995. The use of green fluorescent protein to assess the success of DNA delivery into prawn embryos. First International Workshop on Transgenesis of Invertebrates. Montpellier, France, April 1995, p. 2, abstract.. Primavera, J.H., 1985. A review of maturation and reproduction in closed thelycum penaeids. In: Taki, Y., Primavera, J.H., Lobrera, J.A. Eds.., Proceedings of the First International Conference on the Culture of Penaeid PrawnsrShrimps, 47 December 1984, at Iloilo City, Philippines, pp. 4764. Primavera, J.H., 1993. A critical review of shrimp pond culture in the Philippines. Rev. Fish. Sci. 1, 151201. Pruder, G., Sweeney, J., Carr, W., Gjedrem, T., Wyban, J., 1996. High Health and Genetically Improved Penaeus annamei: Fecundity, Growth Rate and Disease Resistance. World Aquaculture 96, Book of Abstracts. World Aquaculture Society, p. 321. Quackenbush, L.S., 1986. Crustacean endocrinology, a review. Can. J. Fish. Aquat. Sci. 43, 22712282. Quackenbush, L.S., 1989. Vitellogenesis in the shrimp, Penaeus annamei: in vitro studies of the isolated hepatopancreas and ovary. Comp. Biochem. Physiol. B 94, 253261. Quinitio, E.T., Hara, A., Yamauchi, K., Mizushima, T., Fuji, A., 1989. Identification and characterization of vitellin from the ovary in a hermaphrodite shrimp Pandalus kessleri. Comp. Biochem. Physiol. B 94, 445451. Quinitio, E.T., Hara, A., Yamauchi, K., Fuji, A., 1990. Identification and characterization of vitellin from the ovary of Penaeus monodon. Invertebr. Reprod. Dev. 17, 221227. Quinitio, E.T., Caballero, R.M., Gustilo, L., 1993. Ovarian development in relation to changes in the external genitalia in captive Penaeus monodon. Aquaculture 114, 7181. Quinitio, E.T., Hara, A., Yamauchi, K., Nakao, S., 1994. Changes in the steroid hormone and vitellogenin levels during the gametogenic cycle of the giant tiger shrimp Penaeus monodon. Comp. Biochem. Physiol. C 109, 2126.
45
Richardson, B.J., 1982. Geographical distribution of electrophoretically detected protein variation in the Australian commercial fishes: III. Western king prawn, Penaeus latisulcatus Kishinouye. Aust. J. Mar. Freshwat. Res. 33, 933937. Rosenthal, J., Diamant, A., 1990. In vitro primary cell cultures from Penaeus semisulcatus. Pathology in Marine Science. Academic Press, New York, pp. 713. Rothlisberg, P., 1998. Penaeid biology and ecologya review. Aquaculture 164, 4965. Sagi, A., Cohen, D., 1990. Growth, maturation and progeny of sex-reversed Macrobrachium rosenbergii males. World Aquacult. 21, 8790. Salini, J.C., 1987. Genetic variation and population subdivision in the greentail prawn Metapenaeus bennettae. Aust. J. Mar. Freshwat. Res. 38, 339349. Sano, T., 1998. A novel tissue organised in the primary hemolymph culture of penaeid shrimp. Aquaculture 164, 289296. Sbordoni, V., De Matthaeis, E., Cobolli-Sbordoni, M., La Rosa, G., Mattoccia, M., 1986. Bottleneck effects and the depression of genetic variability in hatchery stocks of Penaeus japonicus Crustacea: Decapoda.. Aquaculture 57, 239251. Sbordoni, V., La Rosa, G., Mattoccia, M., Cobolli-Sbordoni, M., De Matthaeis, E., 1987. Genetic changes in seven generations of hatchery stocks of the Kuruma prawn, Penaeus japonicus Crustacea, Decapoda.. In: Tiews, K. Ed.., Selection, Hybridization and Genetic Engineering in Aquaculture. Heenemann Verlag, Berlin, pp. 143155. Scheerer, P.D., Thorgaard, G.H., Seeb, J.E., 1987. Performance and developmental stability of triploid tiger trout Salmo trutta= Salelinus fontinalis.. Trans. Am. Fish. Soc. 116, 9297. Sellos, D., van Wormhoudt, A., 1992. Molecular cloning of a cDNA that encodes a serine protease with chymotryptic and collagenolytic activities in the hepatopancreas of the shrimp Penaeus annamei Crustacea: Decapoda.. FEBS Lett. 309, 219224. Shafir, S., Ovadia, M., Tom, M., 1992a. In vivo incorporation of labeled methionine into proteins, vitellogenin, and vitellin in females of the penaeid shrimp Penaeus semisulcatus de Haan. Biol. Bull. 183, 242247. Shafir, S., Tom, M., Ovadia, M., Lubzens, E., 1992b. Protein, vitellogenin, and vitellin levels in the hemolymph and ovaries during ovarian development in Penaeus semisulcatus de Haan.. Biol. Bull. 183, 394400. Shenker, O., Tietz, A., Ovadia, M., Tom, M., 1993. Lipid synthesis from acetate by the in vitro incubated ovaries of the penaeid shrimp Penaeus semisulcatus. Mar. Biol. 117, 583589. Skaala, ., Dahle, G., Jrstad, K.E., Naevdal, G., 1990. Interactions between natural and farmed fish populations: information from genetic markers. J. Fish Biol. 36, 449460. Sodsuk, S., McAndrew, B.J., Penman, D.J., 1992. Genetic population structure of the giant tiger prawn Penaeus monodon Fabricius, 1798. in the gulf of Thailand and the Andaman Sea. In: Penman, D., Roogratri, N., McAndrew, B. Eds.., Proceedings of the First ASEAN-EEC Aquaculture Development and Coordination Program, AADCP, Stirling, Scotland, pp. 161165. Sodsuk, S., McAndrew, B.J., Penman, D.J., 1996. Population structure and fisheries management of tiger shrimp Penaeus monodon. in SE Asia. In: Penman, D.J., Pongthana, N., McAndrew, B., Bell, A. Eds.., Proceedings of the Second ASEAN-EEC Aquaculture Development and Coordination Program, AADCP, Phuket, Thailand. Subramaniam, T., Keller, R., 1993. A new look at the endocrine regulation of egg maturation in the decapod crustaceans. Curr. Sci. 65, 619623. Sugama, K., Haryanti, 1997. Genetic variation and population structure of tiger shrimp Penaeus monodon Fabricius in Indonesia. Aquaculture, this volume.. Sun, P.S., 1994. Molecular cloning and sequence analysis of a cDNA encoding a molt-inhibiting hormone-like neuropeptide from the white shrimp Penaeus annamei. Mol. Mar. Biol. Biotechnol. 3, 16. Sunden, S.L.F., Davis, S., 1991. Evaluation of genetic variation in a domestic population of Penaeus annamei Boone.: a comparison with three natural populations. Aquaculture 97, 131142. Sung, H.H., Yang, Y.L., Song, Y.L., 1996. Enhancement of microbiocidal activity in the tiger shrimp Penaeus monodon via immunostimulation. J. Crustacean Biol. 16, 278284. Suzuki, T., Furukohri, T., 1994. Evolution of phosphagen kinase. Primary structure of glycocyamine kinase and arginine kinase from invertebrates. J. Mol. Biol. 237, 353357.
46
Tam, Y.K., Chu, K.H., 1993. Electrophoretic study of the phylogenetic relationships of some species of Penaeus and Metapenaeus Decapoda: Penaeidae. from the South China Sea. J. Crustacean Biol. 13, 697705. Tapay, L.M., Lu, Y., Brock, J.A., Nadala, E.C.B., Loh, P., 1995. Transformation of primary cultures of shrimp Penaeus stylirostris. lymphoid Oka. organ with simian virus-40 T. antigen. P.S.E.B.M. 209, 7378. Teshima, S., Kanazawa, A., Koshio, S., Horinouchi, K., 1989. Lipid metabolism of the prawn Penaeus japonicus during maturation: variation in lipid profiles of the ovary and hepatopancreas. Comp. Biochem. Physiol. B 92, 4549. Teshima, S., Kanazawa, A., Hitotsumatsu, K., Kim, K., Oshida, K., Koshio, S., 1992. Tissue uptake and bioconversion of icosapentaenoic acid and phosphatidylcholine in prawns Penaeus and Macrobrachium. Comp. Biochem. Physiol. B 102, 885890. Tom, M., Goren, M., Ovadia, M., 1987. Purification and partial characterization of vitellin from the ovaries of Parapenaeus longirostris Crustacea: Decapoda, Penaeidae.. Comp. Biochem. Physiol. B 87, 1723. Tom, M., Fingerman, M., Hayes, T.K., Johnson, V., Kerner, B., Lubzens, E., 1992. A comparative study of the ovarian protein from two penaeid shrimps, Penaeus semisulcatus de Haan and Penaeus annamei Boone.. Comp. Biochem. Physiol. B 102, 483490. Tom, M., Shenker, O., Ovadia, M., 1993. Partial characterization of three hemolymph proteins of Penaeus semisulcatus de Haan Crustacea: Decapoda, Penaeidae.. Comp. Biochem. Physiol. B 104, 811816. Tong, S-l., Miao, H-Z., 1996. Attempts to initiate cell cultures from Penaeus chinensis tissues. Aquaculture 147, 151157. Tong, J., Ballment, E., Benzie, J.A.H., 1995. Ontogenetic studies of allozymes in the giant tiger prawn Penaeus monodon Crustacea: Decapoda., Aust. Inst. Mar. Sci. Tech. Rep., 21, pp. 116. Toullec, J.-V., Le Moullac, G., Cuzon, G., Van Wormhoudt, A., 1991. Immunoreactive human growth hormone like peptides in tropical penaeid and the effect of dietary HGH on Penaeus annamei larval development. Aquat. Living Resour. 4, 125132. Toullec, J.Y., Crozat, Y., Patrois, J., Porcheron, P., 1996. Development of primary cell cultures from the penaeid shrimps Penaeus annamei and P. indicus. J. Crustacean Biol. 16, 643649. Tsukimura, B., Kamemoto, F.I., 1991. In vitro stimulation of oocytes by presumptive mandibular organ secretions in the shrimp Penaeus annamei. Aquaculture 92, 5966. Van-Herp, F., 1988. La neuroendocrinologie des crustaces: evaluation des progres obtenus ces dernieres annees. In: Le Gal, Y., Van Wormhoudt, A. Eds.., Aspect Recents de la Biologie des Crustaces, Actes de Colloques, 8, pp. 85100. Van-Herp, F., 1993. Importance of reproductive biological knowledge for application of biotechnology in aquaculture: crustaceans as an example. J. Mar. Biotechnol. 1, 1720. Van-Herp, F., Payen, G.G., 1991. Crustacean neuroendocrinology: perspectives for the control of reproduction in aquacultural systems. Bull. Inst. Zool., Academia Sinica, Monogr. 16, 513539. Vazquez-Boucard, C., Ceccaldi, H.J., Benyamin, Y., Roustan, C., 1986. Identification, purification et partial characterization de la lipovitellin chez un crustace decapode natantia Penaeus japonicus Bate.. J. Exp. Mar. Biol. Ecol. 97, 3750. Vickers, J.E., Lester, R.J.G., Spradbrow, P.B., Pemberton, J.M., 1994. Evidence for homology between polyhedrin genes of baculoviruses of a prawn and an insect. J. Invertebr. Pathol. 63, 207208. Villaescusa, A., Camacho, A., Rivalta, V., 1984. Polimorfismo de la fosfoglucosa isomerasa y de la fosfoglucomutasa en el camaron rosado Penaeus notialis .. Cienc. Biol. 12, 2330. Ward, K.A., Nancarrow, C.D., 1991. The genetic engineering of production traits in domestic animals. Experientia 47, 913922. Wongteerasupaya, C., Wongwisansri, S., Boonsaeng, V., Panyim, S., Patanpipat, P., Nash, G., Withyachumnarnkul, B., Flegel, T., 1996. DNA fragment of Penaeus monodon baculovirus PmNOBII gives positive in situ hybridization with white spot viral infections in six penaeid shrimp species. Aquaculture 143, 2332. Wyban, J.A., Swingle, J.S., Sweeney, J.N., Pruder, G.D., 1993. Specific pathogen free Penaeus annamei. World Aquacult. 24, 3945. Xiang, J.-H., 1988. Chromosome studies on Penaeus orientalis. Oceanologia et Limnologia Sinica 19, 205209. Xiang, J.H., Clark, W.H., Griffin, F., Hertzler, P., 1991. A study on feasibility of chromosome set manipulations in the marine shrimp Sicyonia ingentis. Mem. Qld. Mus. 31, 287.
47
Xiang, J.H., Zhou, L.H., Liu, R.Y., Zhu, J.Z., Li, F.H., Li, X.H., Liu, X.D., 1992. Induction of the tetraploids of the chinese shrimp, Penaeus chinensis. Proceedings of the Asia Pacific Conference on Agricultural Biotechnology, Beijing, 2024 Aug. 1992. China Science and Technology Press, Beijing, pp. 841846. Xiang, J.-H., Liu, R.-Y., Zhou, L.-H., 1993. Chromosomes of marine shrimps with special reference to different techniques. Aquaculture 111, 321. Yang, W-J., Aida, K., Nagasawa, H., 1995. Amino acid sequences of a hyperglycaemic hormone and its related peptides from the Kuruma prawn Penaeus japonicus. Aquaculture 135, 205212. Yang, W-J., Aida, K., Terauchi, A., Sonobe, H., Nagasawa, H., 1996. Amino acid sequence of a peptide with moult-inhibiting activity from the Kuruma prawn Penaeus japonicus. Peptides 17, 197202. Yano, I., 1992. Effect of thoracic ganglion on vitellogenin secretion in kuruma prawn Penaeus japonicus. Bull. Natl. Res. Inst. Aquacult. 21, 914. Yano, I., Chinzei, Y., 1987. Ovary is the site of vitellogenin synthesis in kuruma prawn Penaeus japonicus. Comp. Biochem. Physiol. B 86, 213218. Yano, I., Wyban, J.A., 1992. Induced ovarian maturation of Penaeus annamei by injection of lobster brain extract. Bull. Natl. Res. Inst. Aquacult. 21, 17. Yano, I., Tsukimura, B., Sweeney, J.N., Wyban, J.A., 1988. Induced ovarian maturation of Penaeus annamei by implantation of lobster ganglion. J. World Aquacult. Soc. 19, 204209. Young, N.J., Quinlan, P.T., Goad, L.J., 1992a. Cholesteryl esters in the decapod crustacean Penaeus monodon. Comp. Biochem. Physiol. B 102, 761768. Young, N.J., Quinlan, P.T., Goad, L.J., 1992b. Progesterone metabolism in vitro in the decapod crustaceans Penaeus monodon. Gen. Comp. Endocrinol. 87, 300311. Young, N.J., Webster, S.G., Rees, H.H., 1993. Ecdysteroid profiles and vitellogenesis in Penaeus monodon Crustacea: Decapoda.. Invertebr. Reprod. Dev. 24, 107118.

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Aquaculture 164 1998.

Penaeid genetics and biotechnology

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0044-8486r98r$19.00 q 1998 Elsevier Science B.V. All rights reserved. PII S 0 0 4 4 - 8 4 8 6 9 8 . 0 0 1 7 5 - 6

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

J.A.H. Benzier Aquaculture 164 (1998) 2347

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