Estimating genetic parameters in natural populations using the 'animal model'

Philosophical Transactions of the Royal Society B: Biological Sciences

Volumn 359, Issue 1446, 2004, Pages 873-890

  Kruuk, Loeske E B ^a  

^a UNIVERSITY OF EDINBURGH   (United Kingdom)

Author keywords

Animal model; BLUP breeding values; Heritability; Microevolution; Restricted maximum likelihood

Indexed keywords

ENVIRONMENTAL EFFECT; HERITABILITY; MAXIMUM LIKELIHOOD ANALYSIS; POPULATION GENETICS; POPULATION MODELING;

ANIMALIA;

EID: 2942724484     PISSN: 09628436     EISSN: None     Source Type: Journal    
DOI: 10.1098/rstb.2003.1437     Document Type: Review

References (160)

1. Arnold, S. J. & Wade, M. J. 1984 On the measurement of natural and sexual selection: theory. Evolution 38, 709-719.
2. Badyaev, A. V., Beck, M. L., Hill, G. E. & Whittingham, L. A. 2003 The evolution of sexual size dimorphism in the house finch. V. Maternal effects. Evolution 51, 384-396.
3. Barton, N. H. & Keightley, P. D. 2002 Understanding quantitative genetic variation. Nature Rev. Genet. 3, 11-21.
4. Barton, N. H. & Turelli, M. 1989 Evolutionary quantiatative genetics: how little do we know? A. Rev. Genet. 23, 337-370.
5. Blasco, A. 2001 The Bayesian controversy in animal breeding. J. Anim. Sci. 79, 2023-2046.
6. Breslow, N. E. & Clayton, D. G. 1993 Approximate inference in generalized linear mixed models. J. Am. Statist. Assoc. 88, 9-25.
7. Brommer, J. E., Merilä, J. & Kokko, H. 2002 Reproductive timing and individual fitness. Ecol. Lett. 5, 802-810.
8. Brommer, J. E., Pietiainen, H. & Kolunen, H. 2003 Natural selection on individual clutch size-laying date trends in the Ural owl. Evol. Ecol. Res. 5, 229-237.
9. Charlesworth, B. 1987 The heritability of fitness. In Sexual selection: testing the alternatives (ed. J. W. Bradbury & M. B. Andersson), pp. 21-40. New York: Wiley.
10. Charlesworth, B. 1994 Evolution in age-structured populations, 2nd edn. Cambridge University Press.
11. Charmantier, A. & Blondel, J. 2003 A contrast in extra-pair paternity levels on mainland and island populations of Mediterranean blue tits. Ethology 109, 351-363.
12. Charmantier, A., Kruuk, L. E. B., Blondel, J. & Lambrechts, M. M. 2004a Testing for microevolution in body size in three blue tit populations. J. Evol. Biol. 17. (In the press.)
13. Charmantier, A., Kruuk, L. E. B. & Lambrechts, M. M. 2004 Parasitism reduces the potential for evolution in a wild bird population. Evolution 58, 203-206.
14. Cheverud, J. M. & Dittus, W. P. J. 1992 Primate population studies at Polonnaruwa. 2. Heritability of body measurements in a natural population of Toque macaques (Macaca sinica). Am. J. Primatol. 27, 145-156.
15. Clement, V., Bibe, B., Verrier, E., Elsen, J. M., Manfredi, E., Bouix, J. & Hanocq, E. 2001 Simulation analysis to test the influence of model adequacy and data structure on the estimation of genetic parameters for traits with direct and maternal effects. Genet. Selection Evol. 33, 369-395.
16. Clutton-Brock, T. H. & Pemberton, J. M. (eds) 2003 Soay sheep: population dynamics and selection on St Kilda. Cambridge University Press.
17. Clutton-Brock, T. H., Guinness, F. E. & Albon, S. D. 1982 Red deer - behaviour and ecology of two sexes. University of Chicago Press.
18. Coltman, D. W., Bancroft, D. R., Robertson, A., Smith, A., Clutton-Brock, T. H. & Pemberton, J. M. 1999a Male reproductive success in a promiscuous mammal: behavioural estimates compared with a genetic paternity. Mol. Ecol. 8, 1199-1209.
19. Coltman, D. W., Pilkington, J. G., Smith, J. A. & Pemberton, J. M. 1999b Parasite-mediated selection against inbred Soay sheep in a free-living, island population. Evolution 53, 1259-1267.
20. Coltman, D. W., Pilkington, J., Kruuk, L. E. B., Wilson, K. & Pemberton, J. M. 2001 Positive genetic correlation between parasite resistance and body size in a free-living ungulate population. Evolution 55, 2116-2125.
21. Coltman, D. W., O'Donoghue, P., Jorgenson, J. T., Hogg, J. T., Strobeck, C. & Festa-Bianchet, M. 2003 Undesirable evolutionary consequences of trophy hunting. Nature 426, 655-658.
22. Conover, D. O. & Schultz, E. T. 1995 Phenotypic similarity and the evolutionary significance of countergradient variation. Trends Ecol. Evol. 10, 248-252.
23. Cooke, F., Taylor, P. D., Frances, C. M. & Rockwell, R. F. 1990 Directional selection and clutch size in birds. Am. Nat. 136, 261-267.
24. Coulson, T., Kruuk, L. E. B., Tavecchia, G., Pemberton, J. M. & Clutton-Brock, T. H. 2003 Estimating selection on neonatal traits in red deer using elasticity path analysis. Evolution 57, 2879-2892.
25. Coulson, T., Festa-Bianchet, M., Benton, T. G., Kruuk, L. E. B. & Coltman, D. W. 2004 Linking individuals, lineages and population growth to measure fitness and selection in the wild. (Submitted.)
26. de Jong, G. & Bijma, P. 2002 Selection and phenotypic plasticity in evolutionary biology and animal breeding. Livestock Prod. Sci. 78, 195-214.
27. Dohm, M. R. 2002 Repeatability estimates do not always set an upper limit to heritability. Funct. Ecol. 16, 273-280.
28. Duchesne, P., Godbout, M.-H. & Bernatchez, L. 2002 PAPA (Package for the Analysis of Parental Allocation): a computer program for simulated and real parental allocation. Mol. Ecol. Notes 2, 191-193.
29. Ducrocq, V. & Casella, G. 1996 A Bayesian analysis of mixed survival models. Genet. Selection Evol. 28, 505-529.
30. Ducrocq, V. & Solkner, J. 1998 'The survival kit - V3.0'. A package for large analyses of survival data. In 6th World Congr. Genet. Appl. Livest. Prod., vol. 27, pp. 447-448. University of New England, Armidale, Australia.
31. Edwards, A. W. F. 1972 Likelihood. Cambridge University Press.
32. Falconer, D. S. & Mackay, T. F. C. 1996 Introduction to quantitative genetics. Harlow: Longman.
33. Festa-Bianchet, M., Jorgenson, J. T. & Réale, D. 2000 Early development, adult mass, and reproductive success in bighorn sheep. Behav. Ecol. 11, 633-639.
34. Fikse, W. F., Rekaya, R. & Weigel, K. A. 2003 Assessment of environmental descriptors for studying genotype by environment interaction. Livestock Prod. Sci. 82, 223-231.
35. Fisher, R. A. 1921 On the mathematical foundations of theoretical statistics. Phil. Trans. R. Soc. Lond. A 222, 309-368.
36. Fisher, R. A. 1958 The genetical theory of natural selection, 2nd edn. New York: Dover.
37. Garant, D., Dodson, J. J. & Bernatchez, L. 2003 Differential reproductive success and heritability of alternative reproductive tactics in wild Atlantic salmon (Salmo salar L.). Evolution 57, 1133-1141.
38. Garant, D., Sheldon, B. C. & Gustafsson, L. 2004 Climatic and temporal effects on the expression of secondary sexual characters: genetic and environmental components. Evolution 58. (In the press.)
39. George, A. W., Visscher, P. M. & Haley, C. S. 2000 Mapping quantitative trait loci in complex pedigrees: a two-step variance component approach. Genetics 156, 2081-2092.
40. Gianola, D. & Fernando, R. L. 1986 Bayesian methods in animal breeding theory. J. Anim. Sci. 63, 217-244.
41. Gilmour, A. R., Gogel, B. J., Cullis, B. R., Welham, S. J. & Thompson, R. 2002 ASREML user guide release 1.0. Hemel Hempstead, UK: VSN International Ltd.
42. Gosler, A. G. & Harper, D. 2000 Assessing the heritability of body condition in birds: a challenge exemplified by the great tit Parus major L. (Aves). Biol. J. Linn. Soc. 71, 103-117.
43. Grafen, A. 1988 On the uses of data on lifetime reproductive success. In Reproductive success (ed. T. H. Clutton-Brock), pp. 454-471. University of Chicago Press.
44. Groeneveld, E., Kovac, M. & Wang, T. 1990 PEST, a general purpose BLUP package for multivariate prediction and estimation. Proc. 4th World Congr. Genet. Appl. Livestock 13, 488-491.
45. Groeneveld, E., Kovac, M., Wang, T. L. & Fernando, R. L. 1992 Computing algorithms in a general purpose BLUP package for multivariate prediction and estimation. Arch. Anim. Breed. 35, 399-412.
46. Gustafsson, L. 1986 Lifetime reproductive success and heritability: empirical support for Fisher's fundamental theorem. Am. Nat. 128, 761-764.
47. Henderson, C. R. 1950 Estimation of genetic parameters. Ann. Math. Stat. 21, 309-310.
48. Henderson, C. R. 1975 Best linear unbiased estimation and prediction under a selection model. Biometrics 31, 423-447.
49. Henderson, C. R. 1984 Applications of linear models in animal breeding. Guelph, Canada: University of Guelph Press.
50. Hendry, A. P. & Kinnison, M. T. 1999 Perspective: the pace of modern life: measuring rates of contemporary microevolution. Evolution 53, 1637-1653.
51. Hendry, A. P. & Kinnison, M. T. 2001 An introduction to microevolution: rate, pattern, process. Genetica 112, 1-8.
52. Hill, W. G. & Caballero, A. 1992 Artificial selection experiments. A. Rev. Ecol. Syst. 23, 287-310.
53. Hoffmann, A. A. & Merilä, J. 1999 Heritable variation and evolution under favourable and unfavourable conditions. Trends Ecol. Evol. 14, 96-101.
54. Hoffmann, A. A. 2000 Laboratory and field heritabilities: some lessons from Drosophila. In Adaptive genetic variation in the wild (ed. T. A. Mousseau, B. Sinervo & J. A. Endler), pp. 200-218. New York: Oxford University Press.
55. Höschele, I., Gianola, D. & Foulley, J. L. 1987 Estimation of variance components with quasi-continuous data using Bayesian methods. J. Anim. Breed. Genet. 104, 334-349.
56. Jensen, H. 2002 Causes and consequences of individual variation in fitness-related traits in house sparrows. PhD thesis, Norwegian University of Science and Technology, Trondheim, Norway.
57. Jensen, H., Sæther, B.-E., Ringsby, T. H., Tufto, J., Griffith, S. C. & Ellegren, H. 2003 Sexual variation in heritability and genetic correlations of morphological traits in house sparrow (Passer domesticus). J. Evol. Biol. 16, 1296-1307.
58. Jones, A. G. & Ardren, W. R. 2003 Methods of parentage analysis in natural populations. Mol. Ecol. 12, 2511-2523.
59. Kempenaers, B., Pinxten, R. & Eens, M. 1995 Intraspecific brood parasitism in 2 tit Parus species - occurrence and responses to experimental parasitism. J. Avian Biol. 26, 114-120.
60. Kennedy, B. W. 1989 Animal-model BLUP. In Erasmus intensive graduate course 20-26 August 1989. Trinity College, Dublin.
61. Kimura, M. 1958 On the change of population fitness by selection. Heredity 12, 145-167.
62. Kingsolver, J. G., Hoekstra, H. E., Hoekstra, J. M., Berrigan, D., Vignieri, S. N., Hill, C. E., Hoang, A., Gibert, P. & Beerli, P. 2001 The strength of phenotypic selection in natural populations. Am. Nat. 157, 245-261.
63. Kirkpatrick, M. & Lande, R. 1989 The evolution of maternal characters. Evolution 43, 485-503.
64. Knott, S. A., Sibly, R. M., Smith, R. H. & Møller, H. 1995 Maximum likelihood estimation of genetic parameters in life-history studies using the 'animal model'. Funct. Ecol. 9, 122-126.
65. Kolmodin, R., Strandberg, E., Madsen, P., Jensen, J. & Jorjani, H. 2002 Genotype by environment interaction in Nordic dairy cattle studied using reaction norms. Acta Agric. Scand. A 52, 11-24.
66. Konigsberg, L. W. & Cheverud, J. M. 1992 Uncertain paternity in primate quantitative genetic studies. Am. J. Primatol 27, 133-143.
67. Kruuk, L. E. B., Clutton-Brock, T. H., Rose, K. E. & Guinness, F. E. 1999 Early determinants of lifetime reproductive success differ between the sexes in red deer. Proc. R. Soc. Lond. B 260, 1655-1661. (DOI 10.1098/rspb.1999.0828.)
68. Kruuk, L. E. B., Clutton-Brock, T. H., Slate, J., Pemberton, J. M., Brotherstone, S. & Guinness, F. E. 2000 Heritability of fitness in a wild mammal population. Proc. Natl Acad. Sci. USA 97, 698-703.
69. Kruuk, L. E. B., Merilä, J. & Sheldon, B. C. 2001 Phenotypic selection on heritable size traits revisited. Am. Nat. 158, 557-571.
70. Kruuk, L. E. B., Sheldon, B. C. & Merilä, J. 2002a Severe inbreeding depression in collared flycatchers (Ficedula albicollis). Proc. R. Soc. Lond. B 269, 1581-1589. (DOI 10.1098/rspb.2002.2049.)
71. Kruuk, L. E. B., Slate, J., Pemberton, J. M., Brotherstone, S., Guinness, F. E. & Clutton-Brock, T. H. 2002b Antler size in red deer: heritability and selection but no evolution. Evolution 56, 1683-1695.
72. Kruuk, L. E. B., Slate, J., Pemberton, J. M. & Clutton-Brock, T. H. 2003 Fluctuating asymmetry in a secondary sexual trait: no associations with individual fitness, environmental stress or inbreeding, and no heritability. J. Evol. Biol. 16, 101-113.
73. Kunz, C. & Ekman, J. 2000 Genetic and environmental components of growth in nestling blue tits (Parus caeruleus). J. Evol. Biol. 13, 199-212.
74. Lande, R. & Arnold, S. J. 1983 The measurement of selection on correlated characters. Evolution 37, 1210-1226.
75. Larsson, K., van der Jeugd, H. P., van der Veen, I. T. & Forslund, P. 1998 Body size declines despite positive directional selection on heritable size traits in a barnacle goose population. Evolution 52, 1169-1184.
76. Lessells, C. M. 2002 Parentally biased favouritism: why should parents specialize in caring for different offspring? Phil. Trans. R. Soc. Lond. B 357, 381-403. (DOI 10.1098/ rstb.2001.0928.)
77. Lindström, J. 1999 Early development and fitness in birds and mammals. Trends Ecol. Evol. 14, 343-347.
78. Lummaa, V. & Clutton-Brock, T. 2002 Early development, survival and reproduction in humans. Trends Ecol. Evol. 17, 141-147.
79. Lynch, M. & Walsh, B. 1998 Genetics and analysis of quantitative traits. Sunderland, MA: Sinauer.
80. McAdam, A. G. & Boutin, S. 2003a Effects of food abundance on genetic and maternal variation in the growth rate of juvenile red squirrels. J. Evol. Biol. 16, 1249-1256.
81. McAdam, A. G. & Boutin, S. 2003b Variation in viability selection among cohorts of juvenile red squirrels (Tamiasciurus hudsonicus). Evolution 57, 1689-1697.
82. McAdam, A. G., Boutin, S., Réale, D. & Berteaux, D. 2002 Maternal effects and the potential for evolution in a natural population of animals. Evolution 56, 846-851.
83. McCleery, R. H., Pettifor, R. A., Armbruster, P., Meyer, K., Sheldon, B. & Perrins, C. M. 2004 Components of variance underlying fitness in a natural population of the great tit, Parus major. Am. Nat. 163. (In the press.)
84. MacColl, A. D. C. & Hatchwell, B. J. 2003 Heritability of parental investment in a passerine bird. Evolution 57, 2191-2195.
85. McCulloch, C. E. & Searle, S. R. 2001 Generalized, linear, and mixed models. New York: Wiley.
86. Marshall, T. C., Slate, J., Kruuk, L. E. B. & Pemberton, J. M. 1998 Statistical confidence for likelihood-based paternity inference in natural populations. Mol. Ecol. 7, 639-655.
87. Merilä, J. 1996 Genetic variation in offspring condition: an experiment. Funct. Ecol. 10, 465-474.
88. Merilä, J. 1997 Expression of genetic variation in body size of the collared flycatcher under different environmental conditions. Evolution 51, 526-536.
89. Merilä, J. & Sheldon, B. C. 1999 Genetic architecture of fitness and non-fitness traits: empirical patterns and development of ideas. Heredity 83, 103-109.
90. Merilä, J. & Sheldon, B. C. 2000 Lifetime reproductive success and heritability in nature. Am. Nat. 155, 301-310.
91. Merilä, J. & Sheldon, B. C. 2001 Avian quantitative genetics. Curr. Ornithol. 16, 179-255.
92. Merilä, J., Sheldon, B. C. & Ellegren, H. 1998 Quantitative genetics of sexual size dimorphism in the collared flycatcher, Ficedula albicollis. Evolution 52, 870-876.
93. Merilä, J., Kruuk, L. E. B. & Sheldon, B. C. 2001a Cryptic evolution in a wild bird population. Nature 412, 76-79.
94. Merilä, J., Kruuk, L. E. B. & Sheldon, B. C. 2001b Natural selection on the genetical component of variance in body condition in a wild bird population. J. Evol. Biol. 14, 918-929.
95. Merilä, J., Sheldon, B. C. & Kruuk, L. E. B. 2001c Explaining stasis: microevolutionary studies of natural populations. Genetica 112, 119-222.
96. Merilä, J., Kruuk, L. E. B. & Sheldon, B. C. 2004 Evolution and expression of genetic variation in nestling traits. In 23rd Int. Ornithol. Conf., vol., China. (In the press.)
97. Meyer, K. 1989a Estimation of genetic parameters. In Evolution and animal breeding (ed. W. G. Hill & T. F. C. Mackay), pp. 161-167. Oxford University Press.
98. Meyer, K. 1989b Restricted maximum-likelihood to estimate variance components for animal models with several random effects using a derivative-free algorithm. Genet. Selection Evol. 21, 317-340.
99. Meyer, K. 2000 Random regressions to model phenotypic variation in monthly weights of Australian beef cows. Livestock Prod. Sci. 65, 19-38.
100. Meyer, K. & Hill, W. G. 1997 Estimation of genetic and phenotypic covariance functions for longitudinal or 'repeated' records by restricted maximum likelihood. Livestock Prod. Sci. 47, 185-200.
101. Milner, J. M., Albon, S. D., Illius, A. W., Pemberton, J. M. & Clutton-Brock, T. H. 1999 Repeated selection of morphometric traits in the Soay sheep on St Kilda. J. Anim. Ecol. 68, 472-488.
102. Milner, J. M., Brotherstone, S., Pemberton, J. M. & Albon, S. D. 2000 Variance components and heritabilities of morphometric traits in a wild ungulate population. J. Evol. Biol. 13, 804-813.
103. Mousseau, T. A. & Fox, C. W. (eds) 1998 Maternal effects as adaptations. Oxford University Press.
104. Mousseau, T. A. & Roff, D. A. 1987 Natural selection and the heritability of fitness components. Heredity 59, 181-197.
105. Mousseau, T. A., Ritland, K. & Heath, D. D. 1998 A novel method for estimating heritability using molecular markers. Heredity 80, 218-224.
106. Mrode, R. A. 1996 Linear models for the prediction of animal breeding values. Wallingford, UK: Cabi Publishing.
107. Neumaier, A. & Groeneveld, E. 1998 Restricted maximum likelihood estimation of covariances in sparse linear models. Genet. Selection Evol. 30, 3-26.
108. Patterson, H. D. & Thompson, R. 1971 Recovery of interblock information when block sizes are unequal. Biometrika 58, 545-554.
109. Pemberton, J. M., Albon, S. D., Guinness, F. E., Clutton-Brock, T. H. & Dover, G. A. 1992 Behavioral estimates of male mating success tested by DNA fingerprinting in a polygynous mammal. Behav. Ecol. 3, 66-75.
110. Pigliucci, M. & Schlichting, C. D. 1997 On the limits of quantitative genetics for the study of phenotypic evolution. Acta Biotheor. 45, 143-160.
111. Pinheiro, J. C. & Bates, D. M. 2000 Mixed effects models in S and S-Plus. New York: Springer.
112. Pomiankowski, A. & Møller, A. P. 1995 A resolution of the lek paradox. Proc. R. Soc. Lond. B 260, 21-29.
113. Potti, J. 1999 Maternal effects and the pervasive impact of nestling history on egg size in a passerine bird. Evolution 53, 279-285.
114. Price, T. & Langen, T. 1992 Evolution of correlated characters. Trends Ecol. Evol. 7, 307-310.
115. Price, T., Kirkpatrick, M. & Arnold, S. J. 1988 Directional selection and the evolution of breeding date in birds. Science 240, 798-799.
116. Przybylo, R., Sheldon, B. C. & Merilä, J. 2000 Climatic effects on breeding and morphology: evidence for phenotypic plasticity. J. Anim. Ecol. 69, 395-403.
117. Qvarnström, A. 1999 Genotype-by-environment interactions in the determination of the size of a secondary sexual character in the collared flycatcher (Ficedula albicollis). Evolution 53, 1564-1572.
118. Qvarnström, A. & Price, T. D. 2001 Maternal effects, paternal effects and sexual selection. Trends Ecol. Evol. 16, 95-100.
119. Rausher, M. D. 1992 The measurement of selection on quantitative traits: biases due to environmental covariances between traits and fitness. Evolution 46, 616-626.
120. Réale, D. & Festa-Bianchet, M. 2000 Quantitative genetics of life-history traits in a long-lived wild mammal. Heredity 85, 593-603.
121. Réale, D., Festa-Bianchet, M. & Jorgenson, J. T. 1999 Heritability of body mass varies with age and season in wild bighorn sheep. Heredity 83, 526-532.
122. Réale, D., McAdam, A. G., Berteaux, D. & Boutin, S. 2003a Genetic and plastic responses of a northern mammal to climate change. Proc. R. Soc. Lond. B 270, 591-596. (DOI 10.1098/rspb.2002.2224.)
123. Réale, D., Berteaux, D., McAdam, A. G. & Boutin, S. 2003b Lifetime selection on heritable life history traits in a natural population of red squirrels. Evolution 57, 2416-2423.
124. Rieseberg, L. H., Widmer, A., Arntz, A. M. & Burke, J. M. 2002 Directional selection is the primary cause of phenotypic diversification. Proc. Natl Acad. Sci. USA 99, 12242-12245.
125. Ritland, K. 1996 A marker-based method for inferences about quantitative inheritance in natural populations. Evolution 50, 1062-1073.
126. Ritland, K. & Ritland, C. 1996 Inferences about quantitative inheritance based on natural population structure in the yellow monkey flower Mimulus guttatus. Evolution 50, 1074-1082.
127. Robertson, A. 1955 Selection in animals: synthesis. Cold Spring Harbor Symp. Quant. Biol. 20, 294-299.
128. Robertson, A. 1966 A mathematical model of the culling process in dairy cattle. Anim. Prod. 8, 95-108.
129. Roff, D. 2003 Evolutionary quantitative genetics: are we in danger of throwing out the baby with the bathwater? Annls Zool. Fenn. 40, 315-320.
130. Roff, D. A. 1996 The evolution of genetic correlations: an analysis of patterns. Evolution 50, 1392-1403.
131. Roff, D. A. 1997 Evolutionary quantitative genetics. New York: Chapman & Hall.
132. Roff, D. A. & Mousseau, T. A. 1987 Quantitative genetics and fitness: lessons from Drosophila. Heredity 58, 103-118.
133. Schaeffer, L. R. & Song, H. 1978 Selection bias and REML variance-covariance component estimation. J. Dairy Sci. 61, 91-92.
134. Schluter, D. 2000 The ecology of adaptive radiation. Oxford University Press.
135. Shaw, R. G. 1987 Maximum-likelihood approaches applied to quantitative genetics of natural populations. Evolution 41, 812-826.
136. Sheldon, B. C. & Ellegren, H. 1999 Sexual selection resulting from extrapair paternity in collared flycatchers. Anim. Behav. 57, 285-298.
137. Sheldon, B. C., Kruuk, L. E. B. & Merilä, J. 2003 Natural selection and inheritance of breeding time and clutch size in the collared flycatcher. Evolution 57, 406-420.
138. Simm, G. 1998 Genetic improvement of cattle and sheep. Ipswich: Farming Press.
139. Slate, J., Pemberton, J. M. & Visscher, P. M. 1999 Power to detect QTL in a free-living polygynous population. Heredity 83, 327-336.
140. Slate, J., Visscher, P. M., MacGregor, S., Stevens, D., Tate, M. L. & Pemberton, J. M. 2002 A genome scan for quantitative trait loci in a wild population of red deer (Cervus elaphus). Genetics 162, 1863-1873.
141. Smith, J. A., Wilson, K., Pilkington, J. G. & Pemberton, J. M. 1999 Heritable variation in resistance to gastro-intestinal nematodes in an unmanaged mammal population. Proc. R. Soc. Lond. B 266, 1283-1290. (DOI 10.1098/rspb.1999. 0776.)
142. Sorenson, D. A. & Kennedy, B. W. 1984 Estimation of genetic variances from unselected and selected populations. J. Anim. Sci. 58, 1097-1106.
143. Sorensen, D. A., Wang, C. S., Jensen, J. & Gianola, D. 1994 Bayesian analysis of genetic change due to selection using Gibbs sampling. Genet. Selection Evol. 26, 339-360.
144. Stinchcombe, J. R., Rutter, M. T., Burdick, D. S., Tiffin, P., Rausher, M. D. & Mauricio, R. 2002 Testing for environmentally induced bias in phenotypic estimates of natural selection: theory and practice. Am. Nat. 160, 511-523.
145. Stirling, D. G., Réale, D. & Roff, D. A. 2002 Selection, structure and the heritability of behaviour. J. Evol. Biol. 15, 277-289.
146. Stockwell, C. A., Hendry, A. P. & Kinnison, M. T. 2003 Contemporary evolution meets conservation biology. Trends Ecol. Evol. 18, 94-101.
147. Thomas, S. C. & Hill, W. G. 2000 Estimating quantitative genetic parameters using sibships reconstructed from marker data. Genetics 155, 1961-1972.
148. Thomas, S. C. & Hill, W. G. 2002 Sibship reconstruction in hierarchical population structures using Markov chain Monte Carlo techniques. Genet. Res. 79, 227-234.
149. Thomas, S. C., Coltman, D. W. & Pemberton, J. M. 2001 The use of marker-based relationship information to estimate the heritability of bodyweight in a natural population: a cautionary tale. J. Evol. Biol. 15, 92-99.
150. Thompson, R. 1973 The estimation of variance and covariance components with an application when records are subject to culling. Biometrics 29, 527-550.
151. van der Werf, J. H. J. & de Boer, I. J. M. 1990 Estimation of additive genetic variance when base populations are selected. J. Anim. Sci. 68, 3124-3132.
152. van Tienderen, P. H. 2000 Elasticities and the link between demographic and evolutionary dynamics. Evolution 81, 666-679.
153. van Tienderen, P. H. & de Jong, G. 1994 A general model of the relation between phenotypic selection and genetic response. J. Evol. Biol. 1, 1-12.
154. Visscher, P. M., Woolliams, J. A., Smith, D. & Williams, J. L. 2002 Estimation of pedigree errors in the UK dairy population using microsatellite markers and the impact on selection. J. Dairy Sci. 85, 2368-2375.
155. Wilham, R. L. 1963 The covariance between relatives for characters composed of components contributed by related individuals. Biometrics 19, 18-27.
156. Wilham, R. L. 1972 The role of maternal effects in animal breeding. J. Anim. Sci. 28, 74-81.
157. Wilson, A. J., Hutchings, J. A. & Ferguson, M. M. 2003a Selective and genetic constraints on the evolution of body size in a stream-dwelling salmonid fish. J. Evol. Biol. 16, 584-594.
158. Wilson, A. J., McDonald, G., Moghadam, H. K., Herbinger, C. M. & Ferguson, M. M. 2003b Marker-assisted estimation of quantitative genetic parameters in rainbow trout, Oncorhynchus mykiss. Genet. Res. 81, 145-156.
159. Wolf, J. B. & Wade, M. J. 2001 On the assignment of fitness to parents and offspring: whose fitness is it and when does it matter? J. Evol. Biol. 14, 347-356.
160. Wolf, J. B., Brodie, E. D., Cheverud, J. M., Moore, A. J. & Wade, M. J. 1998 Evolutionary consequences of indirect genetic effects. Trends Ecol. Evol. 13, 64-69.