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Artigos • Bol. Inst. Oceanogr. 41 (1-2) • 1993 • https://doi.org/10.1590/S0373-55241993000100005   copiar

Estudo eletroforético sobre a estrutura populacional do krill, Euphausia superba Dana, na região da Passagem Drake-Estreito de Bransfield - oeste da Península Antártica

Electrophoretic study on population structure of krill, Euphausia superba Dana, in the region of Drake Passage, Bransfield Strait and west of the Anterctic Peninaula

Autoria SCIMAGO INSTITUTIONS RANKINGS

Resumos

Foram analisadas amostras de krill, Euphausia superba, coletadas em 18 estações na região da Passagem Drake-Estreito de Bransfield-oeste da Península Antártica para estudar as variações genéticas através de eletroforese de enzimas em gel de poliacrilamida. Dezenove loci enzimáticos foram analisados, dos quais quatorze (73,6%) apresentam variações alélicas. Cinco dos loci polimórficos apresentam alelos nulos. A heterozigosidade média varia de 0,129-0,157 dependendo da localidade. Não foram observadas, no entanto, diferenças relacionadas ao sexo e ao tamanho do animal A conformidade de distribuição de fenótipos em relação ao equilíbrio de Hardy-Weinberg foi encontrada em todos os loci em todas as estações, exceto Aph-1 das estações 8 e 18, Est-1 das estações 9 e 11 e Mdh-1 da estação 18. As análises de distribuição de alelos indicam a homogeneidade entre todas as amostras coletadas em localidades diferentes. A distância genética entre as amostras varia de 0,000 - 0,002. Os resultados sugerem que os indivíduos de todas as amostras coletadas em todas as estações, pertencem a uma única população com acasalamento ao acaso. A contribuição de fatores físicos e biológicos na homogeneidade do krill na região e a necessidade de um estudo mais detalhado, utilizando não somente a eletroforese mas também outros métodos, foram discutidas.

Estrutura da população; Eletroforese; Isoenzimas; Euphausia superba; Passagem de Drake; Estreito de Bransfield; Península Antártica; Antártica

Samples of Antarctic krill, Euphausia superba, collected from 18 locations in the Drake Passage, Bransfield Strait and on the west coast of the Antarctic Peninsula were analyzed for protein variation using enzyme electrophoresis. Of 19 enzyme loci studied, 14 (73,6%) exhibit allelic variations. Average heterozigosity for a station varies from 0,129 to 0,157. Five loci are with null alleles. Differences in allelic frequency related to sex and size of animal were not observed. No significant deviations from random mating expectation according to the Hardy-Weinberg principle were found in any locus at any station except Aph-1 at the stations 8 and 18; Est-1 at the station 9 and 11, and Mdh-1 at the station 18. These deviations are likely due to the difficulty in .analysing the enzyme pattern or to the small sample sizes. Analysis of allele distribution indicate that samples from all locations are homogeneous in all polymorphic loci. Genetic distances between samples vary from 0.000 to 0.002. These results suggest that specimens from all locations of the investigated region belong to a single genetically homogeneous population. The contribution of physical and biological factors to the homogeneity of krill in the region and the necessity of a detailed study utilizing not only electrophoresis but also another techniques were discussed.

Population structure; Electrophoresis; Isoenzymes; Euphausia superba; Drake Passage; Bransfield Strait; Antarctic Peninsula; Antarctica

ARTIGOS

Estudo eletroforético sobre a estrutura populacional do krill, Euphausia superba Dana, na região da Passagem Drake-Estreito de Bransfield - oeste da Península Antártica

Electrophoretic study on population structure of krill, Euphausia superba Dana, in the region of Drake Passage, Bransfield Strait and west of the Anterctic Peninaula

Phan Van NganI; Vicente GomesI; Maria José de Arruda Campos Rocha PassosI; Hana SuzukiII

IInstituto Oceanográfico da Universidade de São Paulo (Caixa Postal 9075, 01065-970, São Paulo, SP, Brasil)

IICentro de Estudos do Mar, Universidade Federal do Paraná (Caixa Postal 43, 83200-970, Paranaguá, PR, Brasil)

RESUMO

Foram analisadas amostras de krill, Euphausia superba, coletadas em 18 estações na região da Passagem Drake-Estreito de Bransfield-oeste da Península Antártica para estudar as variações genéticas através de eletroforese de enzimas em gel de poliacrilamida. Dezenove loci enzimáticos foram analisados, dos quais quatorze (73,6%) apresentam variações alélicas. Cinco dos loci polimórficos apresentam alelos nulos. A heterozigosidade média varia de 0,129-0,157 dependendo da localidade. Não foram observadas, no entanto, diferenças relacionadas ao sexo e ao tamanho do animal A conformidade de distribuição de fenótipos em relação ao equilíbrio de Hardy-Weinberg foi encontrada em todos os loci em todas as estações, exceto Aph-1 das estações 8 e 18, Est-1 das estações 9 e 11 e Mdh-1 da estação 18. As análises de distribuição de alelos indicam a homogeneidade entre todas as amostras coletadas em localidades diferentes. A distância genética entre as amostras varia de 0,000 - 0,002. Os resultados sugerem que os indivíduos de todas as amostras coletadas em todas as estações, pertencem a uma única população com acasalamento ao acaso. A contribuição de fatores físicos e biológicos na homogeneidade do krill na região e a necessidade de um estudo mais detalhado, utilizando não somente a eletroforese mas também outros métodos, foram discutidas.

Descritores: Estrutura da população, Eletroforese, Isoenzimas, Euphausia superba, Passagem de Drake, Estreito de Bransfield, Península Antártica, Antártica.

ABSTRACT

Samples of Antarctic krill, Euphausia superba, collected from 18 locations in the Drake Passage, Bransfield Strait and on the west coast of the Antarctic Peninsula were analyzed for protein variation using enzyme electrophoresis. Of 19 enzyme loci studied, 14 (73,6%) exhibit allelic variations. Average heterozigosity for a station varies from 0,129 to 0,157. Five loci are with null alleles. Differences in allelic frequency related to sex and size of animal were not observed. No significant deviations from random mating expectation according to the Hardy-Weinberg principle were found in any locus at any station except Aph-1 at the stations 8 and 18; Est-1 at the station 9 and 11, and Mdh-1 at the station 18. These deviations are likely due to the difficulty in .analysing the enzyme pattern or to the small sample sizes. Analysis of allele distribution indicate that samples from all locations are homogeneous in all polymorphic loci. Genetic distances between samples vary from 0.000 to 0.002. These results suggest that specimens from all locations of the investigated region belong to a single genetically homogeneous population. The contribution of physical and biological factors to the homogeneity of krill in the region and the necessity of a detailed study utilizing not only electrophoresis but also another techniques were discussed.

Descriptors: Population structure, Electrophoresis, Isoenzymes, Euphausia superba, Drake Passage, Bransfield Strait, Antarctic Peninsula, Antarctica.

Texto completo disponível apenas em PDF.

Full text available only in PDF format.

Agradecimentos

Agradecemos ao Programa Antártico Brasileiro, PROANTAR/CNPq, à Comissão Interministerial para os Recursos do Mar, CERM, e ao Instituto Oceanográfico da Universidade de São Paulo, pela oportunidade, apoio logístico e financeiro para realizar este trabalho. Ao Conselho Nacional de Desenvolvimento Científico e Tecnológico pela bolsa de pesquisa concedida (Proc. 300299/77-O). À tripulação e equipes científicas das Expedições Antárticas do N/Oc. Trof. W. Besnard" pelo material coletado. À Estação Antártica Brasileira "Comandante Ferraz" pela hospitalidade e apoio durante os trabalhos de campo. À Srta. Lucy Teixeira pela padronização das referências bibliográficas e à Marizilda Magro pelas ilustrações.

Referencias bibliográficas

AKROYD, P. 1967. Acrylamide gel slab electrophoresis in a simple glass cell for improved resolution and comparison of serum proteins. Analyt. Biochem., 19:399-410.

ANDERSON, R. C. 1982. Isozyme variation in euphausiids. PhD. Thesis. University of East Anglia, Fisheries Laboratory. Lowestoft. 184p.

AVISE, J. C. 1974. Systematic value of electrophoretic data. Syst. Zool., 23:465-481.

AYALA, F. J.; VALENTINE, J. W. & ZUMWALT, G. S. 1975. An electrophoretic study of the Antarctic zooplankter Euphausia superba. Limnol. Oceanogr., 20:635-640.

BIOMASS. 1982. 1st Post-FIBEX Hydrographie Data Interpretation Workshop, Hamburg FRG, 20-26 September 1982. BIOMASS Rep. Ser., 30. lip.

______ 1983. 2nd Post-FIBEX Hydrographie Data Interpretation Workshop, Hamburg FRG, 16-20 May 1983. BIOMASS Rep. Ser., 31.26p.

BROWN, A. H. D. 1970. The estimation of Wright's fixation index from genotypic frequencies. Genetica, 41:399-406.

BUCKLIN, A. & WIEBE, P. H. 1986. Genetic heterogeneity in euphausiid populations: Euphausia krohnii and Nematoscelis megalops in North Atlantic slop water. Limnol. Oceanogr., 31:1346-1352.

BURTON, R. S. 1983. Protein polymorphisms and genetic differentiation of marine invertebrate populations. Mar. Biol. Letts, 4:193-206.

FAO. 1985. FAO species identification sheets for fishery purposes. Southern Ocean. CCAMLR Convention area, fishing areas 48, 58 and 88. Commission for the Conservation of Antarctic Marine Living Resources. Rome, Food and Agriculture Organization of the United Nations. 232p.

FERGUSON, A. 1980. Biochemical systematic and evolution. Glasgow, Blackie. 194p.

FEVOLDEN, S. E. 1979. Investigations on krill (Euphausiacea) sampled during the Norwegian Antarctic Research Expedition, 1976 -1977. Sarsia, 64:189-198.

______ 1986. Genetic variation of Euphausia superba Dana in the Antarctic Peninsula waters. Sarsia, 71:169-175.

______ 1988. Biochemical genetics and population structure of Euphausia superba. Comp. Biochem. Physiol, 90B:507 - 513.

______ & AYALA, F. J. 1981. Enzyme polymorphism in Antarctic krill (Euphausiacea): genetic variation between populations and species. Sarsia, 66:167-181.

______ & GEORGE, R. Y. 1984. Size frequency pattern of Euphausia superba in the Antarctic Peninsula waters in the Austral Summer of 1983. J. crustacean Biol., 4 (Spec. No. 1):107-122.

______ & SCHNEPPENHEIM, R. 1989. Genetic homogeneity of krill (Euphausia superba Dana) in the Southern Ocean. Polar Biol., 9:533 -539.

FLOWERDEW, M. W. & CRISP, D. J. 1975. Esterase heterogeneity and an investigation into racial differences in the cirripede Balanus balanoides using acrylamide gel electrophoresis. Mar. Biol., 33:33-39.

FUJIO, Y.; YAMANAKA, R. & SMITH, J. P. 1983. Genetic variation in marine molluscs. Bull, japan. Soc scient. Fish., 49:1809-1817.

GODLEWSKA, M. & RAKUSA-SUSZCZEWSKI, S. 1988. Variability of krill, Euphausia superba, Dana 1852 (Crustacea, Euphausiacea), distribution and biomass in the Western Antarctica (Bransfield Strait, Drake Passage, Elephant Island) during 1976-1977. Investigación pesq., Barcelona, 52:575-586.

HUBBY, J. L. & LEWONTIN, R. C. 1966. A molecular approach to the study of genic heterozigosity in natural populations. I. The number of alleles at different loci in Drosophila pseudoobscura. Genetics, 54:577-594.

KANDA, K.; TAKAGI, K. & SEKI, Y. 1982. Movement of the larger swarms of Antarctic krill Euphausia superba population off Enderby Land during 1976 - 1977 season. J. Tokyo Univ. Fish., 68:25-42.

KALINOWSKI, J. & WTTEK, Z. 1985. Scheme for classifying aggregations of Antarctic krill. BIOMASS Handb., (27), 9p.

KATSURAGAWA, M.; NONATO, L. V.; MADUREIRA, L.; AMARAL, J. C. & FERREDLA, S. L. B. L. 1989. Analysis of krill distribution in the Bransfield Strait and Drake Passage by means of hydroacoustic survey. Pesq. antárt. brasil, 1:11-23.

KÜHL, S. & SCHNEPPENHEIM, R. 1986. Electrophoretic investigations of genetic variation in two krill species Euphausia superba and E. crystallorophias (Euphausiidae). Polar Biol., 6:17-23.

LI, C C 1955. Population genetics. Chicago, University of Chicago Press. 366p.

LUBIMOVA, T. G.; MAKAROV, R. R.; MASLENNIKOV, V. V.; SHEVTSOV, V.V. & SHUST, K. V. 1982. The ecological peculiarities, stocks and role of E. superba in the trophic structure of the Antarctic ecosystem. Selected Papers Presented to the Scientific Committee of CCAMLR, 1982-1984. Part II: p. 391-505. Hobart, Australia.

MacDONALD, C. M. & SCHNEPPENHEIM, R. 1983. Breeding structure and stock identity in the Antarctic krill Euphausia superba Dana. In: Schnack, S. B., ed. On the biology of krill Euphausia superba. Proceedings of the Seminar and Report of the Krill Ecology Group. Bremerhaven, 12-16 May 1983. Ber. Polarfors. Alfred-Wegener-Inst. Polarfors., Sonderheft, 4:240-245.

______; WILLIAMS, R. & ADAMS, M. 1986. Genetic variation and population structure of krill (Euphausia superba Dana) from the Prydz Bay region of Antarctic waters. Polar Biol., 6:233-236.

MacKINTOSH, N. A. 1972. life cycle of Antarctic krill in relation to ice and water conditions.'Discovery' Repts, 36:1-94.

______ 1973. Distribution of post-larval krill in the Antarctic. 'Discovery' Repts, 36:95-156.

MAKAROV, R. R. 1974. Dominance of larval forms in Euphausiid (Crustacea: Eucaridae) ontogenesis. Mar. Biol., 27:93-99.

______; MASLENNIKOV, V. V.; SOLYANKIN, E. V.; SPIRIDONOV, V. A. & YAKOVLEV, V.N. 1987. Variability in population density of Antarctic krill in the western Scotia Sea in relation to hydrological conditions. In: Sahrhage, D., ed. Antarctic Ocean and resources variability. Berlin, Springer-Verlag. p. 231-236.

MARR, J. W.S. 1962. The natural history and geography of the Antarctic krill (Euphausia superba Dana). 'Discovery' Repts, 32:33-464.

MASLENNIKOV, V. V. & SOLYANKIN, E. V. 1987. Patterns of fluctuations in the hydrological conditions of the Antarctic and their effect on the distribution of Antarctic krill. In: Sahrhage, D., ed. Antarctic Ocean and resources variability. Berlin, Springer-Verlag. p. 209-213.

MAUCHLINE, J. 1982. Key for the identification of Antarctic euphausiids. BIOMASS Handb., (5),4p.

NEI, M. 1972. Genetic distance between populations. Am. naturalist, 106:283-292.

______ 1977. F-statistics and analysis of gene diversity in subdivided populations. Ann. Human Genet., 41:225-233.

PHAN, V. N. 1980. Estudo bioquímico e fisiológico sobre os bagres marinhos do Brasil. I. Sobre padrão eletroforético do plasma em gel de poliacrilamida dos bagres da região estuarino lagunar de Cananéia. Bolm Inst, oceanogr., S Paulo, 29:301-303.

PHAN, V. N.; GOMES, V.; SUZUKI, H. & PASSOS, M. J. A. C. R. 1989. Preliminary study on chromosomes of Antarctic krill, Euphausia superba Dana. Polar Biol., 10:149-150.

PRIDDLE, J.; CROXALL, J. P.; EVERSON, I.; HEYWOOD, R. B.; MURPHY, E. J.; PRINCE, P. A. & SEAR, C. B. 1987. Large-scale fluctuations in distribution and abundance of krill. A discussion of possible causes. In: Sahrhage, D., ed. Antarctic Ocean and resources variability. Berlin, Springer-Verlag, p. 169-182.

RAKUSA-SUSZCZEWSKI, S. 1978. Environmental conditions within krill swarms. Pol. Arch. Hydrobiol., 25:585-587.

ROHLF, F. J. 1988. NTSYS-pc Numerical taxonomy and multivariate analysis system. New York. Exeter Publ 156p.

SCHELTEMA, R. S. 1971. Larval dispersal as a means of genetic exchange between geographically separated populations of shoalwater benthic marine gastropods. Biol. Bull mar. biol. Lab., Woods Hole, 140:284-322.

SCHNEPPENHEIM, R. & MacDONALD, C. M. 1984. Genetic variation and population structure of krill (Euphausia superba) in the Atlantic sector of Antarctic waters and off the Antarctic Peninsula. Polar Biol., 3:19-28.

SCOPES, R. K. 1968. Methods for starch gel electrophoresis of sarcoplasmic proteins. An investigation of the relative variety of species. Biochem. J., 707:139-150.

SIEGEL, V. 1983. Population structure of Euphausia superba in the eastern part of the Bransfield Strait. In: Schnack, S. B., ed. On the biology of krill Euphausia superba. Proceeding of the Seminar and Report of the Krill Ecology Group. Bremerhaven, 12-16. May 1983. Ber. Polarfors. Alfred-Wegener-Inst. Polarfors., Sonderheft, 4:227-238.

SMITH, P. J.; FRANCIS, R. LC.C. & JAMESON, A. 1981. An excess of homozygotes at a serum esterase locus in the Atlantic mackerel Scomber scombrus. Anim. Blood Groups and Biochemical Genetics, 12:171-180.

SNEATH, P. H. & SOKAL, R. R. 1973. Numerical taxonomy. San Francisco, W. H. Freeman. 573p.

SWOFFORD, D. L. & SELANDER, R. B. 1981. BIOSYS-1: a FORTRAN program for the comprehensive analysis of electrophoretic data in population genetics and systematics. J. Hered., 72:282-283.

UTTER, F.; AEBERSOLD, P. & WINAS, G. 1986. Interpreting genetic variation detected by electrophoresis. In: Ryman, N. & Utter, F., eds Population genetics & fisheries management. Washington Sea Grant Program. Seattle, University of Washington Press, p. 21-45.

WOLNOMIEJSKI, N.; CZYKIETA, H.; STEPNIK, R. & JACKOWSKA, H. 1982. Biological characteristics of Euphausia superba Dana in the southern Drake Passage and the Bransfield Strait in February-March, 1981 (BIOMASS-FTBEX). Pol. Polar Res., 3:259-271.

WORKMAN, P. L. & NISWANDER, J. D. 1970. Population studies on southwestern Indian tribes. II. Local genetic differentiation in the Papago. Am. J. Human Genet., 22:24-49.

WRIGHT, S. 1965. The interpretation of population structure by F-statistics with special regard to systems of mating. Evolution, 19:395-420.

______ 1978. Evolution and the genetics of populations. Variability within and among natural populations. Chicago, University of Chicago Press. v.4.

YABU, H. & KAWAMURA, A. 1984. Chromosomes of Euphausia superba Dana. Bull. Plankt. Soc. Japan, 31:61-63.

(Manuscrito recebido 25 agosto 1992; revisto 13 dezembro 1993; aceito 17 dezembro 1993)

  • AKROYD, P. 1967. Acrylamide gel slab electrophoresis in a simple glass cell for improved resolution and comparison of serum proteins. Analyt. Biochem., 19:399-410.
  • ANDERSON, R. C. 1982. Isozyme variation in euphausiids. PhD. Thesis. University of East Anglia, Fisheries Laboratory. Lowestoft. 184p.
  • AVISE, J. C. 1974. Systematic value of electrophoretic data. Syst. Zool., 23:465-481.
  • AYALA, F. J.; VALENTINE, J. W. & ZUMWALT, G. S. 1975. An electrophoretic study of the Antarctic zooplankter Euphausia superba. Limnol. Oceanogr., 20:635-640.
  • BIOMASS. 1982. 1st Post-FIBEX Hydrographie Data Interpretation Workshop, Hamburg FRG, 20-26 September 1982. BIOMASS Rep. Ser., 30. lip.
  • ______ 1983. 2nd Post-FIBEX Hydrographie Data Interpretation Workshop, Hamburg FRG, 16-20 May 1983. BIOMASS Rep. Ser., 31.26p.
  • BROWN, A. H. D. 1970. The estimation of Wright's fixation index from genotypic frequencies. Genetica, 41:399-406.
  • BUCKLIN, A. & WIEBE, P. H. 1986. Genetic heterogeneity in euphausiid populations: Euphausia krohnii and Nematoscelis megalops in North Atlantic slop water. Limnol. Oceanogr., 31:1346-1352.
  • BURTON, R. S. 1983. Protein polymorphisms and genetic differentiation of marine invertebrate populations. Mar. Biol. Letts, 4:193-206.
  • FAO. 1985. FAO species identification sheets for fishery purposes. Southern Ocean. CCAMLR Convention area, fishing areas 48, 58 and 88. Commission for the Conservation of Antarctic Marine Living Resources. Rome, Food and Agriculture Organization of the United Nations. 232p.
  • FERGUSON, A. 1980. Biochemical systematic and evolution. Glasgow, Blackie. 194p.
  • FEVOLDEN, S. E. 1979. Investigations on krill (Euphausiacea) sampled during the Norwegian Antarctic Research Expedition, 1976 -1977. Sarsia, 64:189-198.
  • ______ 1986. Genetic variation of Euphausia superba Dana in the Antarctic Peninsula waters. Sarsia, 71:169-175.
  • ______ 1988. Biochemical genetics and population structure of Euphausia superba. Comp. Biochem. Physiol, 90B:507 - 513.
  • ______ & AYALA, F. J. 1981. Enzyme polymorphism in Antarctic krill (Euphausiacea): genetic variation between populations and species. Sarsia, 66:167-181.
  • ______ & GEORGE, R. Y. 1984. Size frequency pattern of Euphausia superba in the Antarctic Peninsula waters in the Austral Summer of 1983. J. crustacean Biol., 4 (Spec. No. 1):107-122.
  • ______ & SCHNEPPENHEIM, R. 1989. Genetic homogeneity of krill (Euphausia superba Dana) in the Southern Ocean. Polar Biol., 9:533 -539.
  • FLOWERDEW, M. W. & CRISP, D. J. 1975. Esterase heterogeneity and an investigation into racial differences in the cirripede Balanus balanoides using acrylamide gel electrophoresis. Mar. Biol., 33:33-39.
  • FUJIO, Y.; YAMANAKA, R. & SMITH, J. P. 1983. Genetic variation in marine molluscs. Bull, japan. Soc scient. Fish., 49:1809-1817.
  • GODLEWSKA, M. & RAKUSA-SUSZCZEWSKI, S. 1988. Variability of krill, Euphausia superba, Dana 1852 (Crustacea, Euphausiacea), distribution and biomass in the Western Antarctica (Bransfield Strait, Drake Passage, Elephant Island) during 1976-1977. Investigación pesq., Barcelona, 52:575-586.
  • HUBBY, J. L. & LEWONTIN, R. C. 1966. A molecular approach to the study of genic heterozigosity in natural populations. I. The number of alleles at different loci in Drosophila pseudoobscura. Genetics, 54:577-594.
  • KANDA, K.; TAKAGI, K. & SEKI, Y. 1982. Movement of the larger swarms of Antarctic krill Euphausia superba population off Enderby Land during 1976 - 1977 season. J. Tokyo Univ. Fish., 68:25-42.
  • KALINOWSKI, J. & WTTEK, Z. 1985. Scheme for classifying aggregations of Antarctic krill. BIOMASS Handb., (27), 9p.
  • KATSURAGAWA, M.; NONATO, L. V.; MADUREIRA, L.; AMARAL, J. C. & FERREDLA, S. L. B. L. 1989. Analysis of krill distribution in the Bransfield Strait and Drake Passage by means of hydroacoustic survey. Pesq. antárt. brasil, 1:11-23.
  • KÜHL, S. & SCHNEPPENHEIM, R. 1986. Electrophoretic investigations of genetic variation in two krill species Euphausia superba and E. crystallorophias (Euphausiidae). Polar Biol., 6:17-23.
  • LI, C C 1955. Population genetics. Chicago, University of Chicago Press. 366p.
  • LUBIMOVA, T. G.; MAKAROV, R. R.; MASLENNIKOV, V. V.; SHEVTSOV, V.V. & SHUST, K. V. 1982. The ecological peculiarities, stocks and role of E. superba in the trophic structure of the Antarctic ecosystem. Selected Papers Presented to the Scientific Committee of CCAMLR, 1982-1984. Part II: p. 391-505. Hobart, Australia.
  • MacDONALD, C. M. & SCHNEPPENHEIM, R. 1983. Breeding structure and stock identity in the Antarctic krill Euphausia superba Dana. In: Schnack, S. B., ed. On the biology of krill Euphausia superba. Proceedings of the Seminar and Report of the Krill Ecology Group. Bremerhaven, 12-16 May 1983.
  • Ber. Polarfors. Alfred-Wegener-Inst. Polarfors., Sonderheft, 4:240-245.
  • ______; WILLIAMS, R. & ADAMS, M. 1986. Genetic variation and population structure of krill (Euphausia superba Dana) from the Prydz Bay region of Antarctic waters. Polar Biol., 6:233-236.
  • MacKINTOSH, N. A. 1972. life cycle of Antarctic krill in relation to ice and water conditions.'Discovery' Repts, 36:1-94.
  • ______ 1973. Distribution of post-larval krill in the Antarctic. 'Discovery' Repts, 36:95-156.
  • MAKAROV, R. R. 1974. Dominance of larval forms in Euphausiid (Crustacea: Eucaridae) ontogenesis. Mar. Biol., 27:93-99.
  • ______; MASLENNIKOV, V. V.; SOLYANKIN, E. V.; SPIRIDONOV, V. A. & YAKOVLEV, V.N. 1987. Variability in population density of Antarctic krill in the western Scotia Sea in relation to hydrological conditions. In: Sahrhage, D., ed. Antarctic Ocean and resources variability. Berlin, Springer-Verlag. p. 231-236.
  • MARR, J. W.S. 1962. The natural history and geography of the Antarctic krill (Euphausia superba Dana). 'Discovery' Repts, 32:33-464.
  • MASLENNIKOV, V. V. & SOLYANKIN, E. V. 1987. Patterns of fluctuations in the hydrological conditions of the Antarctic and their effect on the distribution of Antarctic krill. In: Sahrhage, D., ed. Antarctic Ocean and resources variability. Berlin, Springer-Verlag. p. 209-213.
  • MAUCHLINE, J. 1982. Key for the identification of Antarctic euphausiids. BIOMASS Handb., (5),4p.
  • NEI, M. 1972. Genetic distance between populations. Am. naturalist, 106:283-292.
  • ______ 1977. F-statistics and analysis of gene diversity in subdivided populations. Ann. Human Genet., 41:225-233.
  • PHAN, V. N. 1980. Estudo bioquímico e fisiológico sobre os bagres marinhos do Brasil. I. Sobre padrão eletroforético do plasma em gel de poliacrilamida dos bagres da região estuarino lagunar de Cananéia. Bolm Inst, oceanogr., S Paulo, 29:301-303.
  • PHAN, V. N.; GOMES, V.; SUZUKI, H. & PASSOS, M. J. A. C. R. 1989. Preliminary study on chromosomes of Antarctic krill, Euphausia superba Dana. Polar Biol., 10:149-150.
  • PRIDDLE, J.; CROXALL, J. P.; EVERSON, I.; HEYWOOD, R. B.; MURPHY, E. J.; PRINCE, P. A. & SEAR, C. B. 1987. Large-scale fluctuations in distribution and abundance of krill. A discussion of possible causes. In: Sahrhage, D., ed. Antarctic Ocean and resources variability. Berlin, Springer-Verlag, p. 169-182.
  • RAKUSA-SUSZCZEWSKI, S. 1978. Environmental conditions within krill swarms. Pol. Arch. Hydrobiol., 25:585-587.
  • ROHLF, F. J. 1988. NTSYS-pc Numerical taxonomy and multivariate analysis system. New York. Exeter Publ 156p.
  • SCHELTEMA, R. S. 1971. Larval dispersal as a means of genetic exchange between geographically separated populations of shoalwater benthic marine gastropods. Biol. Bull mar. biol. Lab., Woods Hole, 140:284-322.
  • SCHNEPPENHEIM, R. & MacDONALD, C. M. 1984. Genetic variation and population structure of krill (Euphausia superba) in the Atlantic sector of Antarctic waters and off the Antarctic Peninsula. Polar Biol., 3:19-28.
  • SCOPES, R. K. 1968. Methods for starch gel electrophoresis of sarcoplasmic proteins. An investigation of the relative variety of species. Biochem. J., 707:139-150.
  • SIEGEL, V. 1983. Population structure of Euphausia superba in the eastern part of the Bransfield Strait. In: Schnack, S. B., ed. On the biology of krill Euphausia superba. Proceeding of the Seminar and Report of the Krill Ecology Group. Bremerhaven, 12-16. May 1983.
  • Ber. Polarfors. Alfred-Wegener-Inst. Polarfors., Sonderheft, 4:227-238.
  • SMITH, P. J.; FRANCIS, R. LC.C. & JAMESON, A. 1981. An excess of homozygotes at a serum esterase locus in the Atlantic mackerel Scomber scombrus. Anim. Blood Groups and Biochemical Genetics, 12:171-180.
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Datas de Publicação

  • Publicação nesta coleção
    14 Maio 2012
  • Data do Fascículo
    1993

Histórico

  • Aceito
    17 Dez 1993
  • Revisado
    13 Dez 1993
  • Recebido
    25 Ago 1992
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