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Inbreeding – lessons from animal breeding, evolutionary biology and conservation genetics

Published online by Cambridge University Press:  09 March 2007

T. N. Kristensen  and
A. C. Sørensen
T. N. Kristensen
Affiliation:
Department of Genetics and Biotechnology, Danish Institute of Agricultural Sciences, PO Box 50, DK-8830 Tjele, Denmark Aarhus Centre for Environmental Stress Research (ACES), Department of Ecology and Genetics, University of Aarhus, Building 540, Ny Munkegade, DK-8000 Aarhus C, Denmark
A. C. Sørensen*
Affiliation:
Department of Genetics and Biotechnology, Danish Institute of Agricultural Sciences, PO Box 50, DK-8830 Tjele, Denmark Institute of Production Animals and Horses, The Royal Veterinary and Agricultural University, Ridebanevej 12, DK-1870 Frederiksberg C, Denmark
*
†Corresponding author. E-mail: [email protected]
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Abstract

Abstract Increased rates of inbreeding are one side effect of breeding programmes designed to give genetic progress for traits of economic importance in livestock. Inbreeding leads to inbreeding depression for traits showing dominance, and will ultimately lead to a decrease in genetic variance within populations. Here we review theoretical and experimental literature from animal breeding, evolutionary biology and conservation genetics on the consequences of inbreeding in terms of trait means and genetic and environmental variance components. The genetic background for these effects is presented and the experimental literature interpreted in relation to them. Furthermore, purging of deleterious alleles and the variable nature of effects of inbreeding on populations are discussed. Based on the literature, we conclude that inbreeding in animal breeding must be controlled very efficiently to maintain long-term sustainable livestock production in the future. The tools to do this efficiently exist, and much can be learnt on inbreeding from the literature in fields only distantly related to animal breeding.

Keywords

animal breedinggenetic varianceinbreedingsustainability
Type
Research Article
Information
Animal Science , Volume 80 , Issue 2 , April 2005 , pp. 121 - 133
Copyright
Copyright © British Society of Animal Science 2005

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References

Abplanalp, H. 1990. Inbreeding. In Poultry breeding and genetics (ed. Crawford, R.D.), pp. 955–984. Elsevier, Amsterdam.Google Scholar
Andersen, B.B., Jensen, B., Nielsen, A., Christensen, L. G. and Liboriussen, T. 2003. Rød Dansk Malkerace – avlsmæssigt og kulturhistorisk belyst. Report no. 50 in animal husbandry from the Danish Institute of Agricultural Sciences.Google Scholar
Anderson, V.L. and Kempthorne, O. 1954. A model for the study of quantitative inheritance. Genetics 9: 883–898.CrossRefGoogle Scholar
Ballou, J.D. 1997. Ancestral inbreeding only minimally affects inbreeding depression in mammalian populations. Journal of Heredity 8: 169–178.CrossRefGoogle Scholar
Belonsky, G.M. and Kennedy, B.W. 1988. Selection on individual phenotype and best linear unbiased predictor of breeding value in a closed swine herd. Journal of Animal Science 66: 1124–1131.CrossRefGoogle Scholar
Bijlsma, R., Bundgaard, J. and Boerema, A.C. 2000. Does inbreeding affect the extinction risk of small populations?: predictions from Drosophila. Journal of Evolutionary Biology 3: 502–514.CrossRefGoogle Scholar
Bijlsma, R., Bundgaard, J. and Van Putten, W.F. 1999. Environmental dependence of inbreeding depression and purging in Drosophila melanogaster. Journal of Evolutionary Biology 12: 1125–1137.CrossRefGoogle Scholar
Bohren, B.B., Hill, W.G. and Robertson, A. 1966. Some observations on asymmetrical correlated responses to selection. Genetical Research 7: 44–57.CrossRefGoogle ScholarPubMed
Briskie, J.V. and Mackintosh, M. 2004. Hatching failure increases with severity of population bottlenecks in birds. Proceedings of the National Academy of Sciences of the United States of America 101: 558–561.CrossRefGoogle ScholarPubMed
Bruce, A.B. 1910. The Mendelian theory of heredity and the augmentation of vigor. Science 32: 627–628.CrossRefGoogle ScholarPubMed
Bryant, E.H., McCommas, S.A. and Combs, L.M. 1986. The effect of an experimental bottleneck upon quantitative genetic variation in the housefly. Genetics 114: 1191–1211.CrossRefGoogle ScholarPubMed
Bryant, E.H. and Meffert, L. M. 1988. Effect of an experimental bottleneck on morphological integration in the housefly. Evolution 42: 698–707.CrossRefGoogle ScholarPubMed
Buri, P. 1956. Gene frequency in small populations of mutant Drosophila. Evolution 10: 367–402.CrossRefGoogle Scholar
Charlesworth, B. 1998. The effect of synergistic epistasis on the inbreeding load. Genetical Research 71: 85–89.CrossRefGoogle ScholarPubMed
Charlesworth, B. and Charlesworth, D. 1999. The genetic basis of inbreeding depression. Genetical Research 74: 329–340.CrossRefGoogle ScholarPubMed
Charlesworth, D. and Charlesworth, B. 1987. Inbreeding depression and its evolutionary consequences. Annual Review of Ecology and Systematics 18: 237–268.CrossRefGoogle Scholar
Christensen, L. G. and Pedersen, J. 1988. [Crossbreeding of dairy cattle.] Report no. 650 from the National Institute of Animal Science, Denmark, Copenhagen.Google Scholar
Clayton, G.A., Morris, J.A. and Robertson, A. 1957. An experimental check on quantitative genetical theory. I. Short-term responses to selection. Journal of Genetics 55: 131–151.CrossRefGoogle Scholar
Cockerham, C.C. 1984. Additive by additive variance with inbreeding and linkage. Genetics 108: 487–500.CrossRefGoogle ScholarPubMed
Cockerham, C.C. and Weir, B.S. 1973. Descent measures for two loci with some applications. Theoretical Population Biology 4: 300–330.CrossRefGoogle ScholarPubMed
Cockerham, C.C. and Weir, B.S. 1983. Variance of actual inbreeding. Theoretical Population Biology 23: 85–109.CrossRefGoogle ScholarPubMed
Cockerham, C.C. and Weir, B.S. 1984. Covariances of relatives stemming from a population undergoing mixed self and random mating. Biometrics 40: 157–164.CrossRefGoogle ScholarPubMed
Crnokrak, P. and Barrett, S.C.H. 2002. Purging the genetic load: a review of the experimental evidence. Evolution 56: 2347–2358.Google ScholarPubMed
Crnokrak, P. and Roff, D.A. 1999. Inbreeding depression in the wild. Heredity 83: 260–270.CrossRefGoogle ScholarPubMed
Crow, J.F. 1993. Mutation, mean fitness, and genetic load. In Oxford surveys in evolutionary biology (ed. Futuyma, D. and Antonovics, J.), pp. 3–42. Oxford University Press, NY.Google Scholar
Crow, J.F. and Kimura, M. 1970. An introduction to population genetics theory. Harper and Row, NY.Google Scholar
Darwin, C. 1868. The variation of animals and plants under domestication. John Murray, London.Google Scholar
Darwin, C. 1876. The effects of cross and self-fertilization in the vegetable kingdom. Appleton, NY.CrossRefGoogle Scholar
Davenport, C.B. 1908. Degeneration, albinism, and inbreeding. Science 28: 454–455.CrossRefGoogle ScholarPubMed
Day, S.B., Bryant, E.H. and Meffert, L.M. 2003. The infuence of variable rates of inbreeding on fitness, environmental responsiveness, and evolutionary potential. Evolution 57: 1314–1324.Google Scholar
DeRose, M.A. and Roff, D.A. 1999. A comparison of inbreeding depression in life-history and morphological traits in animals. Evolution 53: 1288–1292.CrossRefGoogle ScholarPubMed
Dudash, M.R. 1990. Relative fitness of selfed and outcrossed progeny in a self-compatible, protandrous species, Sabatia angularis L. (Gentianaceae): A comparison in three environments. Evolution 44: 1129–1139.CrossRefGoogle Scholar
Ehiobu, N.G., Goddard, M.E. and Taylor, J.F. 1989. Effect of rate of inbreeding on inbreeding depression in Drosophila melanogaster. Theoretical and Applied Genetics 77: 123–127.CrossRefGoogle ScholarPubMed
Ellegren, H., Hartman, G., Johansson, M. and Andersson, L. 1993. Major histocompatibility complex monomorphism and low levels of DNA fingerprinting variability in a reintroduced and rapidly expanding population of beavers. Proceedings of the National Academy of Sciences of the United States of America 90: 8150–8153.CrossRefGoogle Scholar
Falconer, D.S. and Mackay, T.F.C. 1996. Introduction to quantitative genetics. Longman, Harlow.Google Scholar
Fernández, A., Toro, M.A. and López-Fanjul, C. 1995. The effect of inbreeding on the redistribution of genetic variance of fecundity and viability in Tribolium castaneum. Heredity 75: 376–381.CrossRefGoogle Scholar
Fowler, K. and Whitlock, M.C. 1999a. The distribution of phenotypic variance with inbreeding. Evolution 53: 1143–1156.CrossRefGoogle ScholarPubMed
Fowler, K. and Whitlock, M.C. 1999b. The variance in inbreeding depression and the recovery of fitness in bottlenecked populations. Proceedings of the Royal Society of London, series B 266: 2061–2066.CrossRefGoogle ScholarPubMed
Frankel, O.H. and Soulé, M.E. 1981. Conservation and evolution. Cambridge University Press, Cambridge.Google Scholar
Frankham, R. 1999. Resolving conceptual issues in conservation genetics: the roles of laboratory species and meta-analysis. Hereditas 130: 195–201.CrossRefGoogle Scholar
Frankham, R., Ballou, J.D. and Briscoe, D.A. 2002. Introduction to conservation genetics. Cambridge University Press, Cambridge.CrossRefGoogle Scholar
Frankham, R., Gilligan, D.M., Morris, D. and Briscoe, D.A. 2001. Inbreeding and extinction: effects of purging. Conservation Genetics 2: 279–285.CrossRefGoogle Scholar
Franklin, I.R. 1980. Evolutionary change in small populations. In Conservation biology: an evolutionary-ecological perspective (ed. Soulé, M.E. and Wilcox, B.A.), pp. 135–150. Sinauer, Sunderland, MA.Google Scholar
Franklin, I.R. and Frankham, R. 1998. How large must populations be to retain evolutionary potential? Animal Conservation 1: 69–70.CrossRefGoogle Scholar
Fu, Y.-B., Namkoong, G. and Carlson, J.E. 1998. Comparison of breeding strategies for purging inbreeding depression via simulation. Conservation Biology 12: 856–864.CrossRefGoogle Scholar
García, N., López-Fanjul, C. and García-Dorado, A. 1994. The genetics of viability in Drosophila melanogaster: effects of inbreeding and artificial selection. Evolution 48: 1277–1285.Google ScholarPubMed
Gething, M.-J. and Sambrook, J. 1992. Protein folding in the cell. Nature 355: 33–45.CrossRefGoogle ScholarPubMed
Good, R.L. and Hallauer, A.R. 1977. Inbreeding depression in maize by selfing and self-sibbing. Crop Science 17: 935–940.CrossRefGoogle Scholar
Goodnight, C.J. 1987. On the effect of founder events on epistatic genetic variance. Evolution 41: 80–91.CrossRefGoogle ScholarPubMed
Goodnight, C.J. 1988. Epistasis and the effect of founder events on the additive genetic variance. Evolution 42: 441–454.CrossRefGoogle ScholarPubMed
Hallauer, A.R. and Sears, J.H. 1973. Changes in quantitative traits associated with inbreeding in a synthetic variety of maize. Crop Science 13: 327–330.CrossRefGoogle Scholar
Hedrick, P.W. 1994. Purging inbreeding depression and the probability of extinction: full-sib mating. Heredity 73: 363–372.CrossRefGoogle ScholarPubMed
Hedrick, P.W. and Kalinowski, S.T. 2000. Inbreeding depression in conservation biology. Annual Review of Ecology and Systematics 31: 139–162.CrossRefGoogle Scholar
Hill, W.G. and Robertson, A. 1966. The effects of linkage on limits of artificial selection. Genetical Research 8: 269–294.CrossRefGoogle ScholarPubMed
Hoelzel, A.R., Halley, J., O'Brien, S.J., Campagna, C., Arnbom, T., Le Boeuf, B., Ralls, K. and Dover, G.A. 1993. Elephant seal genetic variation and the use of simulation models to investigate historical population bottlenecks. Journal of Heredity 84: 443–449.CrossRefGoogle ScholarPubMed
Hoeschele, I. 1991. Additive and nonadditive genetic variance in female fertility of Holsteins. Journal of Dairy Science 74: 1743–1752.CrossRefGoogle ScholarPubMed
Jones, D.F. 1917. Dominance of linked factors as a means of accounting for heterosis. Genetics 2: 466–479.CrossRefGoogle ScholarPubMed
Joron, M. and Brakefield, P.M. 2003. Captivity masks inbreeding effects on male mating success in butterflies. Nature 424: 191–194.CrossRefGoogle ScholarPubMed
Karkkainen, K., Kuittinen, H., Treuren, R. van Vogl, C., Oikarinen, S. and Savolainen, O. 1999. Genetic basis of inbreeding depression in Arabis petraea. Evolution 53: 1354–1365.CrossRefGoogle ScholarPubMed
Keeble, F. and Pellew, C. 1910. The mode of inheritance of stature and of time of flowering in peas Pisum sativum. Journal of Genetics 1: 869–876.CrossRefGoogle Scholar
Keller, L.F. and Waller, D.M. 2002. Inbreeding effects in wild populations. Trends in Ecology and Evolution 17: 230–241.CrossRefGoogle Scholar
Kolmodin, R., Strandberg, E., Jorjani, H. and Danell, B. 2003. Selection in the presence of a genotype by environment interaction: response in environmental sensitivity. Animal Science 76: 375–385.CrossRefGoogle Scholar
Kristensen, T.N., Dahlgaard, J. and Loeschcke, V. 2002. Inbreeding affects Hsp 70 expression in two species of Drosophila even at benign temperatures. Evolutionary Ecology Research 4: 1209–1216.Google Scholar
Kristensen, T.N., Dahlgaard, J. and Loeschcke, V. 2003. Effects of inbreeding and environmental stress on fitness-using Drosophila buzzatii as a model organism. Conservation Genetics 4: 453–465.CrossRefGoogle Scholar
Kristensen, T.N., Sørensen, A.C., Sorensen, D., Pedersen, K.S., Sørensen, J.G. and Loeschcke, V. 2005. A test of quantitative genetic theory using Drosophila – effects of inbreeding and rate of inbreeding on heritabilities and variance components. Journal of Evolutionary Biology In press.CrossRefGoogle ScholarPubMed
Laikre, L., Ryman, N. and Thompson, E.A. 1993. Hereditary blindness in a captive wolf (Canis lupus) population. Conservation Biology 7: 592–601.CrossRefGoogle Scholar
Lande, R. 1979. Quantitative genetic analysis of multivariate evolution, applied to brain:body size allometry. Evolution 33: 402–416.Google ScholarPubMed
Lande, R. 1995. Mutation and conservation. Conservation Biology 9: 782–791.CrossRefGoogle Scholar
Latter, B.D.H., Mulley, J.C., Reid, D. and Pascoe, L. 1995. Reduced genetic load revealed by slow inbreeding in Drosophila melanogaster. Genetics 139: 287–297.CrossRefGoogle ScholarPubMed
Latter, R. and Robertson, A. 1962. The effects of inbreeding and artificial selection on reproductive fitness. Genetical Research 3: 110–138.CrossRefGoogle Scholar
Lerner, I.M. 1954. Genetic homeostasis. Oliver and Boyd, Edinburgh.Google Scholar
Lewontin, R.C. 1974. The analysis of variance and the analysis of causes. American Journal of Human Genetics 26: 400–411.Google ScholarPubMed
Li, Z.K., Luo, L.J., Mei, H.W., Wang, D.L., Shu, Q.Y., Tabien, R., Zhong, D.B., Ying, C.S., Stansel, J.W., Khush, G.S. and Paterson, A.H. 2001. Overdominant epistatic loci are the primary genetic basis of inbreeding depression and heterosis in rice.I. Biomass and grain yield. Genetics 158: 1737–1753.CrossRefGoogle ScholarPubMed
López-Fanjul, C., Fernández, A. and Toro, M.A. 1999. The role of epistasis in the increase in the additive genetic variance after population bottlenecks. Genetical Research 73: 45–59.CrossRefGoogle Scholar
López-Fanjul, C., Fernández, A. and Toro, M.A. 2000. Epistasis and the conversion of non-additive genetic variance at population bottlenecks. Theoretical Population Biology 58: 49–59.CrossRefGoogle ScholarPubMed
López-Fanjul, C., Fernández, A. and Toro, M.A. 2002. The effect of epistasis on the excess of the additive and nonadditive variances after population bottlenecks. Evolution 56: 865–876.Google ScholarPubMed
López-Fanjul, C., Guerra, J. and García, A. 1989. Changes in the distribution of the genetic variance of a quantitative trait in small populations of Drosophila melanogaster. Genetics, Selection, Evolution 21: 159–168.CrossRefGoogle Scholar
López-Fanjul, C. and Villaverde, A. 1989. Inbreeding increases genetic variance for viability in Drosophila melanogaster. Evolution 43: 1800–1804.CrossRefGoogle ScholarPubMed
Lynch, M. 1988. Design and analysis of experiments on random drift and inbreeding depression. Genetics 120: 791–807.CrossRefGoogle ScholarPubMed
Lynch, M. and Walsh, B. 1998. Genetics and analysis of quantitative traits. Sinauer Associates, Sunderland, MA.Google Scholar
Madsen, T., Shine, R., Olsson, M. and Wittzell, H. 1999. Restoration of an inbred adder population. Nature 402: 34–35.CrossRefGoogle Scholar
Malécot, P. 1948. Les mathématiques de l'hérédité. Masson, Paris.Google Scholar
Man, W.Y.N., James, J.W. and Nicholas, F.W. 2002. Effect of inbreeding contribution from particular ancestors: a preliminary analysis of first lactation milk yields from Holstein Friesians in Australia. Proceedings of the seventh world congress on genetics applied to livestock production,Montpellier, CD-ROM communication[23–01].Google Scholar
Meagher, S., Penn, D.J. and Potts, W.K. 2000. Male-male competition magnifies inbreeding depression in wild house mice. Proceedings of the National Academy of Sciences of the United States of America 97: 3324–3329.CrossRefGoogle ScholarPubMed
Meuwissen, T.H.E. 1997. Maximizing the response of selection with a predefined rate of inbreeding. Journal of Animal Science 75: 934–940.CrossRefGoogle ScholarPubMed
Meuwissen, T.H.E. and Woolliams, J.A. 1994. Effective sizes of livestock populations to prevent a decline in fitness. Theoretical and Applied Genetics 89: 1019–1026.CrossRefGoogle ScholarPubMed
Miglior, F., Burnside, E.B. and Hohenboken, W.D. 1994. Heterogeneity among families of Holstein cattle in inbreeding depression for production traits. Proceedings of the fifth world congress on genetics applied to livestock production, Guelph, vol. 18, pp. 479–482.Google Scholar
Miglior, F., Burnside, E.B. and Kennedy, B.W. 1995. Production traits of Holstein cattle: estimation of nonadditive genetic variance components and inbreeding depression. Journal of Dairy Science 78: 1174–1180.CrossRefGoogle ScholarPubMed
Misztal, I., Lawlor, T.J. and Gengler, N. 1997. Relationships among estimates of inbreeding depression, dominance and additive variance for linear traits in Holsteins. Genetics, Selection, Evolution 29: 319–326.CrossRefGoogle Scholar
Mitton, J.B. 1993. Enzyme heterozygosity, metabolism, and developmental stability. Genetica 89: 47–65.CrossRefGoogle Scholar
Nicholas, F.W. and Smith, C. 1983. Increased rates of genetic change in dairy cattle by embryo transfer and splitting. Animal Production 36: 341–353.Google Scholar
Pedersen, K.S., Kristensen, T.N. and Loeschcke, V. 2005. Effects of inbreeding and rate of inbreeding in Drosophila melanogaster – Hsp70 expression and fitness. Journal of Evolutionary Biology In press.CrossRefGoogle ScholarPubMed
Phillips, P.C., Whitlock, M.C. and Fowler, K. 2001. Inbreeding changes the shape of the genetic covariance matrix in Drosophila melanogaster. Genetics 158: 1137–1145.CrossRefGoogle ScholarPubMed
Pirchner, F. 1985. Genetic structure of populations.I. Closed populations or matings among related individuals. In General and quantitative genetics (ed. Chapman, A.B.), pp. 227–250. Elsevier, Amsterdam.Google Scholar
Pusey, A. and Wolf, M. 1996. Inbreeding avoidance in animals. Trends in Ecology and Evolution 11: 201–206.CrossRefGoogle ScholarPubMed
Queitsch, C., Sangster, T.A. and Lindquist, S. 2002. Hsp90 as a capacitor of phenotypic variation. Nature 417: 618–624.CrossRefGoogle ScholarPubMed
Ralls, K. and Ballou, J. 1983. Extinction: lessons from zoos. In Genetics and conservation: a reference for managing wild animal and plant populations (ed. Schonewald-Cox, S.M. Chambers, B. MacBryde, B. and Thomas, L.), pp. 164–184. Benjamin/Cummings, Menlo Park, CA.Google Scholar
Ralls, K., Ballou, J.D. and Templeton, A. 1988. Estimates of lethal equivalents and the cost of inbreeding in mammals. Conservation Biology 2: 185–193.CrossRefGoogle Scholar
Reed, D.H., Briscoe, D.A. and Frankham, R. 2002. Inbreeding and extinction: the effect of environmental stress and lineage. Conservation Genetics 3: 301–307.CrossRefGoogle Scholar
Reed, D.H. and Bryant, E.H. 2000. Experimental tests of minimum viable population size. Animal Conservation 3: 7–14.CrossRefGoogle Scholar
Reed, D.H., Lowe, E.H., Briscoe, D.A. and Frankham, R. 2003. Inbreeding and extinction: effects of rate of inbreeding. Conservation Genetics 4: 405–410.CrossRefGoogle Scholar
Robertson, A. 1952. The effect of inbreeding on the variation due to recessive genes. Genetics 37: 189–207.CrossRefGoogle ScholarPubMed
Rutherford, S.L. and Lindquist, S. 1998. Hsp90 as a capacitor for morphological evolution. Nature 396: 336–342.CrossRefGoogle ScholarPubMed
Rye, M. and Mao, I.L. 1998. Nonadditive genetic effects and inbreeding depression for body weight in Atlantic salmon (Salmo salar L.). Livestock Production Science 57: 15–22.CrossRefGoogle Scholar
Sánchez, L., Bijma, P. and Woolliams, J.A. 2003. Minimizing inbreeding by managing genetic contributions across generations. Genetics 164: 1589–1595.CrossRefGoogle Scholar
Smith, L.A., Cassell, B.G. and Pearson, R.E. 1998. The effects of inbreeding on the lifetime performance of dairy cattle. Journal of Dairy Science 81: 2729–2737.CrossRefGoogle ScholarPubMed
Sonesson, A.K., Grundy, B., Woolliams, J.A. and Meuwissen, T.H.E. 2000. Selection with control of inbreeding in populations with overlapping generations: a comparison of methods. Animal Science 70: 1–8.CrossRefGoogle Scholar
Stearns, S.C., Kaiser, M. and Kawecki, T.J. 1995. The differential genetic and environmental canalization of fitness components in Drosophila melanogaster. Journal of Evolutionary Biology 8: 539–557.CrossRefGoogle Scholar
Sørensen, J.G., Kristensen, T.N. and Loeschcke, V. 2003. The evolutionary and ecological role of heat shock proteins. Ecology Letters 6: 1025–1037.CrossRefGoogle Scholar
Templeton, A.R. and Read, B. 1984. Factors eliminating inbreeding depression in a captive herd of Speke's gazelle (Gazella spekei). spekei Zoo Biology 3: 177–199.CrossRefGoogle Scholar
Templeton, A.R. and Read, B. 1994. Inbreeding: one word, several meanings, much confusion. In Conservation genetics (ed. Loeschcke, V., Tomiuk, J. and Jain, S.K.), pp. 91–105. Birkhäuser, Basel.CrossRefGoogle Scholar
Vermeulen, C.J. and Bijlsma, R. 2004. Characterization of conditionally expressed mutants affecting age-specific survival in inbred lines of Drosophila melanogaster: Lethal conditions and temperature sensitive periods. Genetics 167: 1241–1248.CrossRefGoogle ScholarPubMed
Verrier, É., Colleau, J.J. and Foulley, J.-L. 1993. Long-term effects of selection based on the animal model BLUP in a finite population. Theoretical and Applied Genetics 87: 446–454.CrossRefGoogle Scholar
Visscher, P.M., Smith, D., Hall, S.J.G. and Williams, J.L. 2001. A viable herd of genetically uniform cattle. Nature 409: 303.CrossRefGoogle ScholarPubMed
Waddington, C.H. 1942. Canalization of development and the inheritance of acquired characters. Nature 150: 563–565.CrossRefGoogle Scholar
Waller, D.M. 1993. The statics and dynamics of mating systems. In The natural history of inbreeding and outbreeding: theoretical and emperical perspectives (ed. Thornhill, N.W.), pp.97–117. University of Chicago Press, Chicago.Google Scholar
Walsh, B. and Lynch, M. 1998. Evolution and selection of quantitative traits. http://nitro.biosci.arizona.edu/zbook/volume_2/chapters/vol2_03.htmlGoogle Scholar
Wang, J., Caballero, A., Keightley, P.D. and Hill, W.G. 1998. Bottleneck effect on genetic variance: a theoretical investigation of the role of dominance. Genetics 150: 435–447.CrossRefGoogle ScholarPubMed
Wang, S., Hard, J.J. and Utter, F. 2002. Salmonid inbreeding: a review. Reviews in Fish Biology and Fisheries 11: 301–319.CrossRefGoogle Scholar
Wang, J., Hill, W.G., Charlesworth, D. and Charlesworth, B. 1999. Dynamics of inbreeding depression due to deleterious mutations in small populations: mutation parameters and inbreeding rate. Genetical Research 74: 165–178.CrossRefGoogle ScholarPubMed
Weir, B.S., Avery, P.J. and Hill, W.G. 1980. Effect of mating structure on variation in inbreeding. Theoretical Population Biology 18: 396–429.CrossRefGoogle Scholar
Weir, B.S. and Cockerham, C.C. 1969. Group inbreeding with two linked loci. Genetics 63: 711–742.CrossRefGoogle ScholarPubMed
White, J.M. 1972. Inbreeding effects upon growth and maternal ability in laboratory mice. Genetics 70: 307–317.CrossRefGoogle ScholarPubMed
Whitlock, M.C. and Fowler, K. 1996. The distribution among populations in phenotypic variance with inbreeding. Evolution 50: 1919–1926.CrossRefGoogle ScholarPubMed
Whitlock, M.C. and Fowler, K. 1999. The changes in genetic and environmental variance with inbreeding in Drosophila melanogaster. Genetics 152: 345–353.CrossRefGoogle ScholarPubMed
Whitlock, M.C., Phillips, P.C. and Fowler, K. 2002. Persistence of changes in the genetic covariance matrix after a bottleneck. Evolution 56: 1968–1975.Google ScholarPubMed
Whitlock, M.C., Phillips, P.C. and Wade, M.J. 1993. Gene interaction affects the additive genetic variance in subdivided populations with migration and extinction. Evolution 47: 1758–1769.CrossRefGoogle ScholarPubMed
Wiener, G., Lee, G.J. and Woolliams, J.A. 1992a. Effects of rapid inbreeding and of crossing of inbred lines on the body weight growth of sheep. Animal Production 55: 89–99.Google Scholar
Wiener, G., Lee, G.J. and Woolliams, J.A. 1992b. Effects of rapid inbreeding and of crossing of inbred lines on the growth of linear body dimensions of sheep. Animal Production 55: 101–114.Google Scholar
Wiener, G., Lee, G.J. and Woolliams, J.A. 1992c. Effects of rapid inbreeding and of crossing of inbred lines on conception rate, prolificacy and ewe survival in sheep. Animal Production 55: 115–121.Google Scholar
Wiener, G., Lee, G.J. and Woolliams, J.A. 1994. Consequences of inbreeding for financial returns from sheep. Animal Production 59: 245–249.Google Scholar
Willis, J.H. and Orr, H.A. 1993. Increased heritable variation following population bottlenecks: the role of dominance. Evolution 47: 949–957.CrossRefGoogle ScholarPubMed
Woolliams, J.A. and Bijma, P. 2000. Predicting rates of inbreeding in populations undergoing selection. Genetics 154: 1851–1864.CrossRefGoogle ScholarPubMed
Wray, N.R. and Goddard, M.E. 1994. Increasing long-term response to selection. Genetics, Selection, Evolution 26: 431–451.CrossRefGoogle Scholar
Wright, S. 1951. The genetical structure of populations. Annals of Eugenics 15: 323–354.CrossRefGoogle ScholarPubMed