Reviews in Fish Biology and Fisheries

, Volume 11, Issue 4, pp 301–319 | Cite as

Salmonid inbreeding: a review



We review the published literature oninbreeding and its consequences in salmonidfishes. Inbreeding reduces genetic variationwithin populations by decreasingheterozygosity, either through an increasedchance of sharing parental genes or a loss ofalleles from random genetic drift. Increasedinbreeding is often associated with a reductionin mean phenotypic value of one or more traitswith respect to fitness (inbreedingdepression). We identify several sources ofinbreeding in salmonids. Although inbreedingoccurs naturally, much of the evidence forinbreeding stems from direct or indirectresults of human activity. The potentialconsequences of inbreeding highlight theimportance of maintaining genetic diversity insalmonid populations. Our weak understandingof genetic interactions between cultured andwild salmonids has allowed widespread practicesthat can reduce genetic variability in naturalpopulations. Although studies have detectedinbreeding depression in salmonids, its geneticbasis has rarely been addressed in wild,anadromous salmon. The genetic basis ofinbreeding depression is complex, andevaluating its effects over the entire lifecycle remains challenging. The experimentalevidence nevertheless reinforces the importanceof maintaining genetic variation withinpopulations as a primary goal of conservationand management.

inbreeding depression inbreeding genetic variation salmon culture salmonids 


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  1. Allendorf, F.W. and Leary, R.F. (1986) Heterozygosity and fitness in natural populations of animals. In: Soulé, M.E. (ed.), Conservation Biology: The Science of Scarcity and Diversity. Sunderland, Massachusetts, Sinauer Associates, Inc. Publishers, pp. 57-76.Google Scholar
  2. Allendorf, F.W. and Phelps, S.R. (1980) Loss of genetic variation in hatchery stock of cutthroat trout. Trans. Am. Fish. Soc. 109, 537-543.Google Scholar
  3. Allendorf, F.W. and Ryman, N. (1987) Genetic management of hatchery stocks. In: Ryman, N. and Utter, F. (eds.), Population Genetics and Fishery Management. Seattle, University of Washington Press, pp. 141-159.Google Scholar
  4. Allendorf, F.W. and Seeb, L.W. (2000) Concordance of genetic divergence among sockeye salmon populations at allozyme, nuclear DNA, and mitochondrial DNA markers. Evolution 54, 640-651.PubMedGoogle Scholar
  5. Allendorf, F.W. and Thorgaard, G.H. (1984) Tetraploidy and the evolution of salmonid fishes. In: Turner, B.J. (ed.), Evolutionary Genetics of Fishes. New York, Plenum, pp. 1-53.Google Scholar
  6. Allendorf, F.W and Utter, F.M. (1979) Population genetics. In: Hoar, W.S., Randall, D.J., and Brett, J.R. (eds.), Fish Physiology, Vol. VIII. New York, Academic Press, pp. 407-454.Google Scholar
  7. Allendorf, F.W. and Waples, R.S. (1996) Conservation and genetics of salmonid fishes. In: Avise, J.C. and Hamrick, J.L. (eds.), Conservation Genetics: Case Histories from Nature. New York, Chapman & Hall, pp. 238-501.Google Scholar
  8. Altukhov, Yu. P. (1995) Intraspecific genetic diversity: Monitoring and conservation. Genetika 31, 1333-1357.PubMedGoogle Scholar
  9. Amos, W. and Balmford, A. (2001) When does conservation genetics matter? Heredity 87, 257-265.PubMedGoogle Scholar
  10. Armbruster, P., Hutchinson, R.A. and Linvell, T. (2000) Equivalent inbreeding depression under laboratory and field conditions in a tree-hole-breeding mosquito. Proc. Roy. Soc. Lond. Biol. Sci., Ser. B 267, 1939-1945.Google Scholar
  11. Aulstad, D., Gjedrem, T. and Skjervold, H. (1972) Genetic and environmental sources of variation in length and weight of rainbow trout (Salmo gairdneri). J. Fish. Res. Board Can. 29, 237-241.Google Scholar
  12. Aulstad, D. and Kittlesen, A. (1971) Abnormal body curvatures of rainbow trout (Salmo gairdneri) inbred fry. J. Fish. Res. Board Can. 28, 1918-1920.Google Scholar
  13. Avise, J.C. (1994) Molecular Markers, Natural History and Evolution. New York, Chapman & Hall, 511 pp.Google Scholar
  14. Avise, J.C. and Hamrick, J.L. (eds.) (1996) Conservation and Genetics: Case Histories from Nature. New York, Chapman & Hall, 512 pp.Google Scholar
  15. Ballou, J.D. (1997) Ancestral inbreeding only minimally affects inbreeding depression in mammalian populations. J. Heredity 88, 169-178.Google Scholar
  16. Barrett, S.C.H. and Charlesworth, D. (1991) Effects of a change in the level of inbreeding on the genetic load. Nature (London) 352, 522-524.PubMedGoogle Scholar
  17. Bateson, P. (1983) Optimal outbreeding. In: Bateson, P. (ed.), Mate Choice, Cambridge, Cambridge University Press, pp. 257-277.Google Scholar
  18. Bierne, N., Tsitrone, A. and David, P. (2000) An inbreeding model of associative overdominance during a population bottleneck. Genetics 155, 1981-1990.PubMedCentralPubMedGoogle Scholar
  19. Bijlsma, R., Bundgaard, J. and Boerema, A.C. (2000) Does inbreeding affect the extinction risk of small populations?: predictions from Drosophila. J. Evol. Biol. 13, 502-514.Google Scholar
  20. Bijlsma, R., Bundgaard, J. and Van Putten, W.F. (1999) Environmental dependence of inbreeding depression and purging in Drosophila melanogaster. J. Evol. Biol. 12, 1125-1137.Google Scholar
  21. Blanchfield, P.J. and Ridgway, M.S. (1998) The cost of peripheral males in a brook trout mating system. Anim. Behav. 57, 537-544.Google Scholar
  22. Burger, C.V., Spearman, W.J. and Cronin, M.A. (1997) Genetic differentiation of sockeye salmon subpopulations from a geologically young Alaskan lake system. Trans. Am. Fish. Soc. 126, 926-938.Google Scholar
  23. Busack, C.A., Halliburton, R. and Gall, G.A.E. (1979) Electrophoretic variation and differentiation in four strains of domesticated rainbow trout (Salmo Gairdneri). Can. J. Genet. Cytol. 21, 81-94.PubMedGoogle Scholar
  24. Busack, C.A. and Currens, K.P. (1995) Genetic risks and hazards in hatchery operations: Fundamental concepts and issues. Am. Fish. Soc. Symp. 15, 71-80.Google Scholar
  25. Byers, D.L. and Waller, D.M. (1999) Do plant populations purge their genetic load? Effects of population size and mating history on inbreeding depression. Annu. Rev. Ecol. Syst. 30, 479-513.Google Scholar
  26. Caballero, A. (1994) Developments in the prediction of effective populations size. Heredity 73, 657-679.PubMedGoogle Scholar
  27. Campton, D.E. (1995) Genetic effects of hatchery fish on wild populations of Pacific salmon and steelhead: What do we really know? Am. Fish. Soc. Symp. 15, 337-353.Google Scholar
  28. Campton, D.E. and Utter, F.M. (1987) Genetic structure of anadromous cutthroat trout (Salmo clarki clarki) populations in the Puget Sound area: Evidence for restricted gene flow. Can. J. Fish. Aquat. Sci. 44, 573-582.Google Scholar
  29. Campbell, R.B. (1995) The effect of mating structure and progeny distribution on heterozygosity versus the number of alleles as measures of variation. J. Theor. Biol. 175, 503-509.PubMedGoogle Scholar
  30. Caughley, G. and Gunn, A. (1996) Conservation Biology in Theory and Practice. Cambridge, MA, Blackwell Science, 459 pp.Google Scholar
  31. Charlesworth, B. (1998) The effect of synergistic epistasis on the inbreeding load. Genet. Res. Camb. 71, 81-89.Google Scholar
  32. Charlesworth, D. and Charlesworth, B. (1987) Inbreeding depression and its evolutionary consequences. Annu. Rev. Ecol. Syst. 18, 237-268.Google Scholar
  33. Chebanov, N.A. (1979) Behavior and mating of chum salmon spawners of similar size with a significant abundance domination of males on spawning grounds. Ekologiya 2, 73-79.Google Scholar
  34. Chebanov, N.A. (1984) Assortative mating in pink salmon Oncorhynchus gorbuscha (Walbaum). Ekologiya 4, 70-76.Google Scholar
  35. Chebanov, N.A. (1990) Spawning behavior, assortative mating, and spawning success of coho salmon, Oncorhynchus kisutch, under natural and experimental conditions. J. Ichthyol. 30, 1-12.Google Scholar
  36. Chesser, R.K. and Ryman, N. (1986) Inbreeding as a strategy in subdivided populations. Evolution 40, 616-624.Google Scholar
  37. Cooney, R.T. and Brodeur, R.D. (1996) Carrying capacity and North Pacific salmon production. Rosen. Sch. Mar. Atmos. Sci. 62, 443-464.Google Scholar
  38. Cooper, E.L. (1961) Growth of wild and hatchery strains of brook trout. Trans. Am. Fish. Soc. 90, 424-438.Google Scholar
  39. Cross, T.F. and King, J. (1983) Genetic effects of hatchery rearing in Atlantic salmon. Aquaculture 33, 33-40.Google Scholar
  40. Courtenay, S.C., Quinn, T.P., Dupuis, M.C., Groot, C. and Lapkin, P.A. (2001) Discrimination of family-specific odours by juvenile coho salmon: roles of learning and odour concentration. J. Fish. Biol. 58, 107-125.Google Scholar
  41. Crnokrak, P. and Roff, D.A. (1999) Inbreeding depression in the wild. Heredity 83, 260-270.PubMedGoogle Scholar
  42. Crow, J.F. and Kimura, M. (1970) An Introduction to Population Genetics Theory. New York, Harper & Row, Publishers, 591 pp.Google Scholar
  43. Crozier, W.W. and Moffett, I.J.J. (1989) Amount and distribution of biochemical-genetic variation among wild populations and a hatchery stock of Atlantic salmon, Salmo salar L., from northeast Ireland. J. Fish. Biol. 35, 665-677.Google Scholar
  44. Dahlgaard, J. and Hoffmann, A.A. (2000) Stress resistance and environmental dependency of inbreeding depression in Drosophila melanogaster. Conserv. Biol. 14, 1187-1192.Google Scholar
  45. Danielsdottir, A.K., Marteinsdottir, G., Arnason, F. and Gudjonsson, S. (1997) Genetic structure of wild and reared Atlantic salmon (Salmo salar L.) populations in Iceland. ICES J. Mar. Sci. 54, 986-997.Google Scholar
  46. DeRose, M.A. and Roff, D. (1999) A comparison of inbreeding depression in life-history and morphological traits in animals. Evolution 53, 1288-1292.Google Scholar
  47. Ehiobu, N.G., Goddard, M.E. and Taylor, J.F. (1989) Effect of rate of inbreeding on inbreeding depression in Drosophila melanogaster. Theor. Appl. Genet. 77, 123-127.PubMedGoogle Scholar
  48. Falconer, D.S. and Mackay, T.F.C. (1996) Introduction to Quantitative Genetics, 4th ed. Harlow, UK, Longman, 480 pp.Google Scholar
  49. Ferguson, M.M., Ihssen, P.E. and Hynes, J.D. (1991) Are cultured stocks of brown trout (Salmo trutta) and rainbow trout (Oncorhynchus mykiss) genetically similar to their source populations? Can. J. Fish. Aquat. Sci. 48(Suppl. 1), 118-123.Google Scholar
  50. Flagg, T.A., Waknitz, F.W., Maynard, D.J., Milner, G.B. and Mahnken, C.V.W. (1995) The effect of hatcheries on native coho salmon populations in the lower Columbia River. Am. Fish. Soc. Symp. 15, 366-375.Google Scholar
  51. Foote, C.J. (1989) Female mate preference in Pacific salmon. Anim. Behav. 38, 721-723.Google Scholar
  52. Foote, C.J., Brown, G.S. and Wood, C.C. (1997) Spawning success of males using alternative mating tactics in sockeye salmon, Oncorhynchus nerka. Can. J. Fish. Aquat. Sci. 54, 1785-1795.Google Scholar
  53. Foote, C.J. and Larkin, P.A. (1988) The role of male choice in the assortative mating of anadromous and nonanadromous sockeye salmon. Behaviour 106, 43-62.Google Scholar
  54. Foote, C.J., Wood, C.C. and Withler, R.E. (1989) Biochemical genetic comparison of sockeye salmon and kokanee, the anadromous and nonanadromous forms of Oncorhynchus nerka. Can. J. Fish. Aquat. Sci. 46, 149-158.Google Scholar
  55. Ford, M.J. (2002) Selection in captivity during supportive breeding may reduce fitness in the wild. Conservation Biology. 16, 815-825.Google Scholar
  56. Frankham, R. (1995a) Conservation genetics. Annu. Rev. Genet. 29, 305-327.PubMedGoogle Scholar
  57. Frankham, R. (1995b) Effective population size/adult population size ratios in wildlife: a review. Genet. Res. Camb. 66, 95-106.Google Scholar
  58. Frankham, R., Ballou, J.D. and Briscoe, D.A. (eds.) (2002) Introduction to Conservation Genetics. Cambridge, UK, Cambridge University Press, 617 pp.Google Scholar
  59. Franklin, I.R. and Frankham, R. (1998) How large must populations be to retain evolutionary potential? Anim. Conserv. 1, 69-73.Google Scholar
  60. Gall, G.A.E. (1987) Inbreeding. In: Ryman, N. and Utter, F. (eds.), Population Genetics and Fisheries Management, Seattle, University of Washington Press, pp. 47-80.Google Scholar
  61. Geiger, H.A., Smoker, W.W., Zhivotovsky, L.A. and Gharrett, A.J. (1997) Variability of family size and marine survival in pink salmon (Oncorhynchus gorbuscha) has implications for conservation biology and human use. Can. J. Fish. Aquat. Sci. 54, 2684-2690.Google Scholar
  62. Gharrett, A.J. and Shirley, S.M. (1985) A genetic examination of spawning methodology in a salmon hatchery. Aquaculture 47, 245-256.Google Scholar
  63. Gile, S.R. and Ferguson, M.M. (1990) Crossing methodology and genotypic diversity in a hatchery strain of rainbow trout (Oncorhynchus mykiss). Can. J. Fish. Aquat. Sci. 47, 719-724.Google Scholar
  64. Gjerde, B., Gunnes, K. and Gjedrem, T. (1983) Effect of inbreeding on survival and growth in rainbow trout. Aquaculture 34, 327-332.Google Scholar
  65. Grant, W.S., García-Marín, J.L. and Utter, F.M. (1999) Defining population boundaries for fishery management. In: Mustafa, S. (ed.), Genetics in Sustainable Fisheries Management. Oxford, UK, Fishing News Books, Blackwell Science, pp. 27-72.Google Scholar
  66. Hard, J.J. (1995a) Genetic monitoring of life-history characters in salmon supplementation: problems and opportunities. Am. Fish. Soc. Symp. 15, 212-225.Google Scholar
  67. Hard, J.J. (1995b) A quantitative genetic perspective on the conservation of intraspecific diversity. Am. Fish. Soc. Symp. 17, 304-326.Google Scholar
  68. Hard, J.J., Connell, L., Hershberger, W.K. and Harrell, L.W. (2000) Genetic variation in mortality of chinook salmon (Oncorhynchus tshawytscha) during a bloom of the marine alga Heterosigma akashiwo. J. Fish Biol. 56, 1387-1397.Google Scholar
  69. Hard, J.J., Jones, R.P., Delarm, M.R. and Waples, R.S. (1992) Pacific salmon and artificial propagation under the Endangered Species Act. U.S. Dept. Commer., NOAA Tech. Memo. NMFS-NWFSC-2, 56 pp.Google Scholar
  70. Hard, J.J. and Hershberger, W.K. (1995) Quantitative genetic consequences of captive broodstock programs for anadromous Pacific salmon (Oncorhynchus spp.). In: Flagg, T.A. and Mahnken, C.V.W. (eds.), An Assessment of the Status of Captive Broodstock Technology for Pacific Salmon. Bonneville Power Administration (Project No. 93-56), P.O. Box 3621, Portland, OR 92708-3621, pp. 2-1-2-75.Google Scholar
  71. Hard, J.J. and Hershberger, W.K. (in press) Research on quantitative genetic consequences of captive broodstock programs for Pacific salmon populations. Annual Report to Bonneville Power Administration, P.O. Box 3621, Portland, OR 97208-3621.Google Scholar
  72. Hartl, D.L. and Clark, A.G. (1989) Principles of Population Genetics. Sunderland, MA, Sinauer Associates, Inc. Publishers, 682 pp.Google Scholar
  73. Heard, W.R. (1998) Do hatchery salmon affect the North Pacific Ocean ecosystem? NPAFC Bulletin: assessment and status of Pacific Rim salmonid stocks. No. 1, pp. 405-411.Google Scholar
  74. Hedrick, P.W. (1984) Is there an inbreeding optimum? Zoo Biol. 3, 167-169.Google Scholar
  75. Hedrick, P.W. (1994) Purging inbreeding depression and the probability of extinction: full-sib mating. Heredity 73, 363-372.PubMedGoogle Scholar
  76. Hedrick, P.W. and Hedgecock, D. (1994) Effective population size in winter-run chinook salmon. Conserv. Biol. 8, 890-892.Google Scholar
  77. Hedrick, P.W. and Miller, P.S. (1992) Conservation genetics: techniques and fundamentals. Ecol. Appl. 2, 30-46.Google Scholar
  78. Hedrick, P. and Miller, P. (eds.) (1994) Endangered Pacific salmonids. Conserv. Biol. 8, 863-894.Google Scholar
  79. Hedrick, P.W., Hedgecock, D., Hamelberg, S. and Croci, S.J. (2000a) The impact of supplementation in winter-run chinook salmon on effective population size. J. Hered. 91, 112-116.PubMedGoogle Scholar
  80. Hedrick, P.W., Rashbrook, V.K. and Hedgecock, D. (2000b) Effective population size of winter-run chinook salmon based on microsatellite analysis of returning spawners. Can. J. Fish. Aquat. Sci. 57, 2368-2373.Google Scholar
  81. Hendry A.P., Wenburg, J.K., Bentzen, P., Volk, E.C. and Quinn, T.P. (2000) Rapid evolution of reproductive isolation in the wild: evidence from introduced salmon. Science 290, 516-518.PubMedGoogle Scholar
  82. Henson, A.J. and Smith, H.D. (1967) Mate selection in a population of sockeye salmon (Oncorhynchus nerka) of mixed age groups. J. Fish. Res. Board Can. 24, 1955-1977.Google Scholar
  83. Hilborn, R. and Eggers, D. (2000) A review of the hatchery programs for pink salmon in Prince Williams Sound and Kodiak, Alaska. Trans. Am. Fish. Soc. 129, 333-350.Google Scholar
  84. Hilborn, R. and Eggers, D. (2001) Response to Wertheimer et al. Trans. Am. Fish. Soc. 130, 720-724.Google Scholar
  85. Hilborn, R. and Winton, J. (1993) Learning to enhance salmon production: lessons from the salmonid enhancement program. Can. J. Fish. Aquat. Sci. 50, 2043-2056.Google Scholar
  86. Hindar, K., Ryman, N. and Utter, F. (1991) Genetic effects of cultured fish on natural fish populations. Can. J. Fish. Aquat. Sci. 48, 945-957.Google Scholar
  87. Jimenez, J.A., Hughes, K.A., Alaks, G. Graham, L. and Lacy, R.C. (1994) An experimental study of inbreeding depression in a natural habitat. Science 266, 271-273.PubMedGoogle Scholar
  88. Jonsson, B. (1997) A review of ecological and behavioural interactions between cultured and wild Atlantic salmon. ICES J. Mar. Sci. 54, 1031-1039.Google Scholar
  89. Jorde, P.E. and Ryman, N. (1995) Temporal allele frequency change and estimation of effective size in populations with overlapping generations. Genetics 139, 1075-1090.Google Scholar
  90. Kalinowski, S.T. and Hedrick, P.W. (1998) An improved method for estimating inbreeding depression in pedigrees. Zoo Biol. 17, 481-497.Google Scholar
  91. Kapuscinski, A.R.D. and Lannan, J.E. (1984) Application of a conceptual genetic fitness model for managing Pacific salmon fisheries. Aquaculture 43, 135-146.Google Scholar
  92. Kapuscinski, A.R.D. and Lannan, J.E. (1986) A conceptual genetic fitness model for fisheries management. Can. J. Fish. Aquat. Sci. 43, 1606-1616.Google Scholar
  93. Karkkainen, K., Kuittinen, H., Van Treuren, R., Vogl, C., Oikarinen, S. and Savolainen, O. (1999) Genetic basis of inbreeding depression in Arabis petraea. Evolution 53, 1354-1365.Google Scholar
  94. Kincaid, H.L. (1976a) Effects of inbreeding on rainbow trout populations. Trans. Am. Fish. Soc. 105, 273-285.Google Scholar
  95. Kincaid, H.L. (1976b) Inbreeding in rainbow trout (Salmo gairdneri). J. Fish. Res. Board Can. 33, 2420-2426.Google Scholar
  96. Kincaid, H.L. (1983) Inbreeding in fish populations used for aquaculture. Aquaculture 33, 215-227.Google Scholar
  97. Kincaid, H.L. (1995) An evaluation of inbreeding and effective population size in salmonid broodstocks in federal and state hatcheries. Am. Fish. Soc. Symp. 15, 193-204.Google Scholar
  98. Koljonen, M. (1986) The enzyme gene variation of ten Finnish rainbow trout strains and the relation between growth rate and mean heterozygosity. Aquaculture 57, 253-260.Google Scholar
  99. Knudsen, E.E., MacDonald, D.D. and Steward, C.R. (2000) Setting the stage for a sustainable Pacific salmon fisheries strategy. In: Knudsen, E.E., Steward, C.R., MacDonald, D.D., Williams, J.E. and Reiser, D.W. (eds.), Sustainable Fisheries Management: Pacific Salmon. New York, Lewis Publishers, CRC Press LLC, pp. 3-11.Google Scholar
  100. Lacy, R.C., Alaks, G. and Walsh, A. (1996) Hierarchical analysis of inbreeding depression in Peromyscus polionotus. Evolution 50, 2187-2200.Google Scholar
  101. Lacy, R.C., Ballou, J.D., Princee, F, Starfield, A. and Thompson, E.A. (1995) Pedigree analysis for population management. In: Ballou, J.D., Gilpin, M. and Foose, T.J. (eds.), Population Management of Survival and Recovery: Analytical Methods and Strategies in Small Population Conservation. New York: Columbia University Press, pp. 57-75.Google Scholar
  102. Laikre, L. (ed.) (1999) Conservation genetic management of brown trout (Salmo trutta) in Europe. Report by the concerted action on identification, management and exploitation of genetic resources in the brown trout (Salmo trutta) (TROUTCONCERT; EU FAIR CT97-3882).Google Scholar
  103. Lande, R. and Barrowclough, G.F. (1987) Effective population size, genetic variation, and their use in population management. In: Soulé, M.E. (ed.), Viable Populations for Conservation. Cambridge, Cambridge University Press, pp. 87-123.Google Scholar
  104. Lande, R. and Schemske, D.W. (1985) The evolution of self-fertilization and inbreeding depression in plants. I. Genetic models. Evolution 39, 24-40.Google Scholar
  105. Landweber, L.F. and Dobson, A.P. (eds.) (1999) Genetics and the Extinction of Species: DNA and the Conservation of Biodiversity. New Jersey, Princeton University Presss, 189 pp.Google Scholar
  106. Latter, B.D.H., Mulley, J.C., Reid, D. and Pascoe, L. (1995) Reduced genetic load reveled by slow inbreeding in Drosophila melanogaster. Genetics 139, 287-297.PubMedCentralPubMedGoogle Scholar
  107. Latter, B.D.H. and Robertson, A. (1962) The effects of inbreeding and artificial selection on reproductive fitness. Genet. Res. 3, 110-138.Google Scholar
  108. Loeschcke, V., Tomiuk, J. and Jain, S.K. (1994) Introductory remarks: Genetics and conservation biology. In: Loeschcke, V., Tomiuk, J. and Jain, S.K. (eds.), Conservation Genetics. Basel/Switzerland: Birkhäuser Verlag, pp. 3-8.Google Scholar
  109. Lynch, M. (1988) Design and analysis of experiments on random drift and inbreeding depression. Genetics 120, 791-807.PubMedCentralPubMedGoogle Scholar
  110. Lynch, M. and Walsh, B. (1998) Genetics and Analysis of Quantitative Traits. Sunderland, MA, Sinauer Associates, Inc. Publishers, 980 pp.Google Scholar
  111. McElligott, E.A., Maguire, T.M.F. and Cross, T.F. (1987) The amount and nature of electrophoretically-detectable genetic polymorphism in hatchery reared Atlantic salmon (Salmo salar). Copenhagen, Denmark ICES, 10 pp.Google Scholar
  112. McElhany, P., Ruckelshaus, M.H., Ford, M.J., Wainwright, T.C. and Bjorkstedt, E.P. (2000) Viable salmonid populations and the recovery of evolutionarily significant units. U.S. Department of Commerce, NOAA Technical Memorandum NMFS-NWFSC-42, 1-156.Google Scholar
  113. Miller, P.S. (1994) Is inbreeding depression more severe in a stressful environment? Zoo Biol. 13, 195-208.Google Scholar
  114. Mitton, J.B. (1993a) Enzyme heterozygosity, metabolism, and developmental stability. Genetica 89, 47-65.Google Scholar
  115. Mitton, J.B. (1993b) Theory and data pertinent to the relationship between heterozygosity and fitness. In: Thornhill, N.W. (ed.), The Natural History of Inbreeding and Outbreeding: Theoretical and Empirical Perspectives. Chicago, The University of Chicago Press, pp. 17-41.Google Scholar
  116. Mjolnerod, I.B., Refseth, U.H., Karlsen, E., Balstad, T., Jakobsen, K.S. and Hindar, K. (1997) Genetic differences between two wild and one farmed population of Atlantic salmon (Salmo salar) revealed by three classes of genetic markers. Hereditas 127, 239-248.Google Scholar
  117. Morton, N.E., Crow, J.F. and Muller, H.J. (1956) An estimation of the mutational damage in man from data on consanguineous marriages. Proc. Natl. Acad. Sci. USA. 42, 855-863.PubMedCentralPubMedGoogle Scholar
  118. Myers, J.M., Heggelund, P.O., Hudson, G. and Iwamoto, R.N. (2001) Genetics and broodstock management of coho salmon. Aquaculture 197, 43-62.Google Scholar
  119. Myers, J.M., Kope, R.G., Bryant, G.J., Teel, D., Lierheimer, L.J., Wainwright, T.C., Grant, W.S., Waknitz, F.W., Neely, K., Lindley, S.T. and Waples, R.S. (1998) Status review of chinook salmon from Washington, Idaho, Oregon and California. U.S. Dept. Commer., NOAA Tech. Memo. NMFS-NWFSC-35, 443 pp.Google Scholar
  120. Nehlsen, W., Williams, J.E. and Lichatowich, J.A. (1991) Pacific salmon at the crossroads: stocks at risk from California, Oregon, Idaho, and Washington. Fisheries 16, 4-21.Google Scholar
  121. Neigel, J.E. (1996) Estimation of effective population size and migration parameters from genetic data. In: Smith, T.B. and Wayne, R.K. (eds.), Molecular Genetic Approaches in Conservation. Oxford, Oxford University Press, pp. 329-346.Google Scholar
  122. Norris, A.T., Bradley, D.G. and Cunningham, E.P. (1999) Microsatellite genetic variation between and within farmed and wild Atlantic salmon (Salmo salar). Aquaculture 180, 247-264.Google Scholar
  123. NRC — National Research Council (1996) Upstream: Salmon and Society in the Pacific Northwest. Washington, D.C., National Academic Press, 452 pp.Google Scholar
  124. NRC — National Research Council (2002) Genetic Status of Atlantic Salmon in Maine. Washington, D.C., National Acedemic Press, 62 pp.Google Scholar
  125. Oosterhout, C.V., Zijlstra, W., Van Heuven, M.K. and Brakefield, P.M. (2000) Inbreeding depression and genetic load in laboratory metapopulations of the butterfly Bicyclus anynana. Evolution 54, 218-225.PubMedGoogle Scholar
  126. Pante, M.J.R., Gjerde, B. and McMillan, I. (2001) Effect of inbreeding on body weight at harvest in rainbow trout, Oncorhynchus mykiss. Aquaculture 192, 201-211.Google Scholar
  127. Partridge, L. (1983) Non-random mating and offspring fitness. In: Bateson, P. (ed.), Mate Choice. Cambridge, Cambridge University Press, pp. 227-255.Google Scholar
  128. Phelps, S.R., LeClair, L.L., Young, S. and Blankenship, H.L. (1994) Genetic diversity patterns of chum salmon in the Pacific Northwest. Can. J. Fish. Aquat. Sci. 51(Suppl. 1), 65-83.Google Scholar
  129. Policansky, D. and Magnuson, J.J. (1998) Genetics, metapopulations, and ecosystem management of fisheries. Ecol. Appl. 8(Suppl. 1), S119-S123.Google Scholar
  130. Pray, L.A. and Goodnight, C.J. (1995) Genetic variation in inbreeding depression in the red flour beetle Tribolium castaneum. Evolution 49, 176-188.Google Scholar
  131. Pray, L.A., Schwartz, J.M., Goodnight, C.J. and Stevens, L. (1994) Environmental dependency of inbreeding depression: implications for conservation biology. Conserv. Biol. 8, 562-568.Google Scholar
  132. Prodoehl, P.A., Walker, A.F., Hynes, R., Taggart, J.B. and Ferguson, A. (1997) Genetically monomorphic brown trout (Salmo trutta L.) populations, as revealed by mitochondrial DNA, multilocus and single-locus minisatellite (VNTR) analyses. Heredity 79, 208-213.Google Scholar
  133. Quinn, T.P., Unwin, M.J. and Kinninson, M.T. (2000) Evolution of temporal isolation in the wild: genetic divergence in timing of migration and breeding by introduced chinook salmon populations. Evolution 54, 1372-1385.PubMedGoogle Scholar
  134. Ralls, K. and Ballou, J. (1983) Extinction: lessons from zoos. In: Schonewald-Cox, C.M., Chambers, S.M., MacBryde, B. and Thomas, L. (eds.), Genetics and Conservation. Menlo Park, CA, Benjamin/Cummings, pp. 164-184.Google Scholar
  135. Ralls, K., Ballou, J.D. and Templeton, A.R. (1988) Estimates of lethal equivalents and the cost of inbreeding in mammals. Conserv. Biol. 2, 185-193.Google Scholar
  136. Reilly, A., Elliott, N.G., Grewe, P.M., Clabby, C., Powell, R. and Ward, R.D. (1999) Genetic differentiation between Tasmanian cultured Atlantic salmon (Salmo salar L.) and their ancestral Canadian population: comparison of microsatellite DNA and allozyme and mitochondrial DNA variation. Aquaculture 173, 459-469.Google Scholar
  137. Rye, M. and Mao, I.L. (1998) Nonadditive genetic effects of inbreeding depression for body weight in Atlantic salmon (Salmo salar L.). Livest. Prod. Sci. 57, 15-22.Google Scholar
  138. Ryman, N. (1970) A genetic analysis of recapture frequencies of released young of salmon (Salmo salar L.) Hereditas 65, 159-160.Google Scholar
  139. Ryman, N. (1983) Patterns of distribution of biochemical genetic variation in salmonids: Differences between species. Aquaculture 33, 1-21.Google Scholar
  140. Ryman, N. (1991) Conservation genetics considerations in fishery management. J. Fish. Biol. 39(Suppl. A), 211-224.Google Scholar
  141. Ryman, N. (1994) Supportive breeding and effective population size: differences between inbreeding and variance effective numbers. Conserv. Biol. 8, 888-890.Google Scholar
  142. Ryman, N. and Laikre, L. (1991) Effects of supportive breeding on the genetically effective population size. Conserv. Biol. 5, 325-329.Google Scholar
  143. Ryman, N., Utter, F. and Laikre, L. (1995) Protection of intraspecific biodiversity of exploited fishes. Rev. Fish Biol. Fish. 4, 417-446.Google Scholar
  144. Saccheri, I., Kuussaari, M., Kankare, M., Vikman, P., Fortelius, W. and Hanski, I. (1998) Inbreeding and extinction in a butterfly metapopulation. Nature (London) 392, 491-494.Google Scholar
  145. Schwartz, M.K., Tallman, D.A. and Luikart, G. (1998) Review of DNA-based census and effective population size estimators. Anim. Conserv. 1, 293-299.Google Scholar
  146. Selander, R.K. (1983) Evolutionary consequences of inbreeding. In: Schonewald-Cox, C.M., Chambers, S.M., MacBryde, F. and Thomas, L. (eds.), Genetics and Conservation: A Reference for Managing Wild Animal and Plant Populations. Menlo Park, California, Benjamin/Cummings, pp. 201-215.Google Scholar
  147. Sheffer, R.J., Hedrick, P.W. and Velasco, A.L. (1999) Testing for inbreeding and outbreeding depression in the endangered Gila tompinnow. Anim. Conserv. 2, 121-129.Google Scholar
  148. Shields, W.M. (1982) Philopatry, Inbreeding, and the Evolution of Sex. Albany, NY, State University of New York Press, 245 pp.Google Scholar
  149. Shields, W.M. (1993) The natural and unnatural history of inbreeding and outbreeding. In: Thornhill, N.W. (ed.), The Natural History of Inbreeding and Outbreeding: Theoretical and Empirical Perspectives. Chicago, The University of Chicago Press, pp. 143-169.Google Scholar
  150. Shrimpton, J.M. and Randall, D.J. (1992) Smolting and survival in wild and hatchery coho salmon. World Aquacult. 23, 51-54.Google Scholar
  151. Simon, R.C. (1991) Management techniques to minimize the loss of genetic variability in hatchery fish populations. Am. Fish. Soc. Symp. 10, 487-494.Google Scholar
  152. Slate, J., Kruuk, L.E.B., Marshall, T.C., Pemberton, J.M. and Clutton-Brock, T.H. (2000) Inbreeding depression influences lifetime breeding success in a wild population of red deer (Cervus elaphus). Proc. Roy. Soc. Lond. Biol. Sci., Ser. B 267, 1657-1662.Google Scholar
  153. Ståhl, G. (1983) Differences in the amount and distribution of genetic variation between natural populations and hatchery stocks of Atlantic salmon. Aquaculture 33, 23-32.Google Scholar
  154. Ståhl, G. (1987) Genetic population structure of Atlantic salmon. In: Ryman, N. and Utter, F. (eds.), Population Genetics and Fisheries Management. Seattle, University of Washington Press, pp. 121-140.Google Scholar
  155. Su, G.S., Liljedahl, L.E. and Gall, G.A.E. (1996) Effects of inbreeding on growth and reproductive traits in rainbow trout (Oncorhynchus mykiss). Aquaculture 142, 139-148.Google Scholar
  156. Tallman, R.F. (1986) Genetic differentiation among seasonally distinct spawning populations of chum salmon, Oncorhynchus keta. Aquaculture 57, 211-217.Google Scholar
  157. Tave, D. (1986) Genetics for Fish Hatchery Managers. Westport, Connecticut, AVI Publishing Company, Inc., 299 pp.Google Scholar
  158. Tave, D. (1991) Inbreeding. Aquaculture Magazine 17, 65-67.Google Scholar
  159. Taylor, E.B. (1991) A review of local adaptation in Salmonidae, with particular reference to Pacific and Atlantic salmon. Aquaculture 89, 185-207.Google Scholar
  160. Teel, D.J., Milner, G.B., Winans, G.A. and W.S. Grant. (2000) Genetic population structure and origin of life history types in chinook salmon in British Columbia, Canada. Trans. Am. Fish. Soc. 129, 194-209.Google Scholar
  161. Templeton, A.R. and Read, B. (1983) The elimination of inbreeding depression a captive herd of Speke's gazelle. In: Schonewald-Cox, C.M., Chambers, S.M., MacBryde, B. and Thomas, L. (eds.), Genetics and Conservation. Menlo Park, CA, Benjamin/Cummings.Google Scholar
  162. Templeton, A.R. and Read, B. (1994) Inbreeding: One word, several meanings, much confusion. In: Loeschcke, V., Tomiuk, J. and Jain, S.K. (eds.), Conservation Genetics. Basel/Switzerland, Birkhäuser Verlag, pp. 91-105.Google Scholar
  163. Tessier, N., Bernatchez, L. and Wright, J.M. (1997) Population structure and impact of supportive breeding inferred from mitochondiral and microsatellite DNA analyses in land-locked Atlantic salmon Salmo salar L. Mol. Ecol. 6, 735-750.Google Scholar
  164. Thomaz, D., Beall, E. and Burke, T. (1997) Alternative reproductive tactics in Atlantic salmon: factors affecting mature parr success. Proc. Roy. Soc. Lond. Biol. Sci., Ser. B 264(1379), 219-226.Google Scholar
  165. Thornhill, N.W. (ed.) (1993) The Natural History of Inbreeding and Outbreeding. Chicago, University of Chicago Press, 575 pp.Google Scholar
  166. Unwin, M.J. (1997) Fry-to-adult survival of natural and hatchery-produced chinook salmon (Oncorhynchus tshawytscha) from a common origin. Can. J. Fish. Aquat. Sci. 54, 1246-1254.Google Scholar
  167. Utter, F. (1998) Genetic problems of hatchery-rear progeny released into the wild, and how to deal with them. Bull. Mar. Sci. 62, 623-640.Google Scholar
  168. Utter, F. (2001) Patterns of subspecific anthropogenic introgression in two salmonid genera. Rev. Fish Biol. Fish. 10, 435-451.Google Scholar
  169. Utter, F.M., Chapman, D.W. and Marshall, A.R. (1995) Genetic population structure and history of chinook salmon of the upper Columbia River. Am. Fish. Soc. Symp. 17, 149-165.Google Scholar
  170. Utter, F., Hindar, K. and Ryman, N. (1993) Genetic effects of aquaculture on natural salmonid populations. In: Heen, K., Monahan, R.L. and Utter, F. (eds.), Salmon Aquaculture. Oxford, Fishing News Books, pp. 144-165.Google Scholar
  171. Utter, F., Milner, G., Ståhl, G. and Teel, D. (1989) Genetic population structure of chinook salmon, Oncorhynchus tschawytscha, in the Pacific Northwest. Fish. Bull. 85, 13-23.Google Scholar
  172. Verspoor, E. (1988) Reduced genetic variability in first-generation hatchery populations of Atlantic Salmon (Salmo salar). Can. J. Fish. Aquat. Sci. 45, 1686-1690.Google Scholar
  173. Vuorinen, J. (1982) Little genetic variation in the Finnish Lake salmon, Salmo salar sebago (Girard). Hereditas 97, 189-192.Google Scholar
  174. Wang, J.L. (2000) Effects of population structures and selection strategies on the purging of inbreeding depression due to deleterious mutations. Genetical Research, Cambridge 76, 75-86.Google Scholar
  175. Wang, S., Hard, J.J. and Utter, F. 2002. Genetic variation and fitness in salmonids. Conservation Genetics (in press).Google Scholar
  176. Waples, R.S. (1990a) Conservation genetics of Pacific salmon. II. Effective population size and the rate of loss of genetic variability. J. Hered. 81, 267-276.Google Scholar
  177. Waples, R.S. (1990b) Conservation genetics of Pacific salmon. III. Estimating effective population size. J. Hered. 81, 277-289.Google Scholar
  178. Waples, R.S. (1991) Genetic interactions between hatchery and wild salmonids: lessons from the Pacific Northwest. Can. J. Fish. Aquat. Sci. 48(suppl. 1), 124-133.Google Scholar
  179. Waples, R.S. (1999) Dispelling some myths about hatcheries. Fisheries 24, 12-21.Google Scholar
  180. Waples, R.S. (2002) Definition and estimation of effective population size in the conservation of endangered species. In: Beissinger, S.R. and McCullough, D.R. (eds.), Population Viability Analysis. Chicago, IL, University of Chicago Press, pp. 147-168.Google Scholar
  181. Waples, R.S. and Do, C. (1994) Genetic risk associated with supplementation of Pacific salmonids: Captive broodstock programs. Can. J. Fish. Aquat. Sci. 51(suppl. 1), 310-329.Google Scholar
  182. Waser, N.M. (1993) Population structure, optimal outbreeding, and assortative mating in angiosperms. In: Thornhill, N.W. (ed.), The Natural History of Inbreeding and Outbreeding: Theoretical and Empirical Perspectives. Chicago, The University of Chicago Press, pp. 173-199.Google Scholar
  183. Wertheimer, A.C., Smoker, W.W., Joyce, T.L. and Heard, W.R. (2001) Hatchery pink salmon in Prince William Sound: enhancement or displacement. Trans. Am. Fish. Soc. 130, 712-719.Google Scholar
  184. Winkler, F.M., Bartley, D. and Diaz, N.F. (1999) Genetic differences among year classes in a hatchery population of a coho salmon (Oncorhynchus kisutch (Walbaum, 1792)) in Chile. Aquaculture 173, 423-431.Google Scholar
  185. Withler, R.E. (1988) Genetic consequences of fertilizing chinook salmon (Oncorhynchus tshawytscha) eggs with pooled milt. Aquaculture 68, 15-25.Google Scholar
  186. Withler, R.E. and Beacham, T.D. (1994) Genetic consequences of the simultaneous or sequential addition of semen from multiple males during hatchery spawning of chinook salmon (Oncorhynchus tshawytscha). Aquaculture 126, 11-23.Google Scholar
  187. Wood, C.C. (1995) Life history variation and population structure in sockeye salmon. Am. Fish. Soc. Symp. 17, 195-216.Google Scholar

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© Kluwer Academic Publishers 2001

Authors and Affiliations

  1. 1.School of Aquatic and Fishery SciencesUniversity of WashingtonSeattleUSA
  2. 2.Conservation Biology DivisionNorthwest Fisheries Science Center, National Marine Fisheries ServiceSeattleUSA

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