Abstract.-The severity of inbreeding depression appears to vary among taxa, but few ecological or other patterns have been identified that predict accurately which taxa are most sensitive to inbreeding. To examine the causes of heterogeneity in inbreeding depression, the effects of inbreeding on reproduction, survival, and growth were measured in three replicate experimental stocks for each of three subspecies of Peromyscus polionotus mice. Inbreeding of the dam reduced the probability of breeding, the probability of producing a second litter, and litter size. Inbreeding of the litter caused depression of litter size, juvenile viability, and mass at weaning, and caused an increase in the withinlitter variance in mass. In spite of differences between the subspecies in natural population sizes, genetic variation, and mean rates of reproduction and survival, all variation observed between experimental populations in their responses to inbreeding could be attributed to random founder effects. The genetic load of deleterious alleles in each replicate was unequally partitioned among its founder pairs, and different founders contributed to the load affecting different fitness components. Thus, inbreeding depression for anyone fitness component, in our experimental environment, must be due to relatively few deleterious alleles with major effects. Genetic loads so comprised would be expected to diverge among natural populations due to both random drift and selective removal of recessive deleterious alleles during population bottlenecks. The near universality of inbreeding depression would be maintained, however, if different alleles contribute to inbreeding depression of different fitness components and in different environments.

Many wildlife species are propagated in captivity as models for behavioral, physiological, and genetic research or to provide assurance populations to protect threatened species. However, very little is known about how animals evolve in the novel environment of captivity. The histories of most laboratory strains are poorly documented, and protected populations of wildlife species are usually too small and too short-term to allow robust statistical analysis. To document the evolutionary change in captive breeding programs, we monitored reproduction and behavior across 18 generations in six experimental populations of Peromyscus leucopus mice started from a common set of 20 wild-caught founders. The mice were propagated under three breeding protocols: a strategy to retain maximal genetic diversity, artificial selection against stereotypic behaviors that were hypothesized to reflect poor adaptation to captivity, and random bred controls. Two replicates were maintained with each protocol, and inter-replicate crosses at generations 19 and 20 were used to reverse accumulated inbreeding. We found that one of the stereotypic behaviors (repetitive flipping) was positively associated with reproductive fitness, while the other (gnawing) was relatively invariant. Selection to reduce these stereotypic behaviors caused marked reduction in reproduction, and populations not under artificial selection to reduce these behaviors responded with large increases in flipping. In non-selected populations, there was rapid evolution toward much higher proportion of pairs breeding and more rapid conception. Litter size, pup survival, and weaning mass all declined slowly, to the extent that would be predicted based on inbreeding depression. Inter-crossing between replicate populations reversed these declines in fitness components but did not reverse the changes in behavior or the accelerated breeding. These findings indicate that adaptation to captivity can be rapid, affecting reproductive patterns and behaviors, even under breeding protocols designed to minimize the rate of genetic change due to random drift and inadvertent selection.

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The severity of inbreeding depression appears to vary among taxa, but few ecological or other patterns have been identified that predict accurately which taxa are most sensitive to inbreeding. To examine the causes of heterogeneity in inbreeding depression, the effects of inbreeding on reproduction, survival, and growth were measured in three replicate experimental stocks for each of three subspecies of Peromyscus polionotus mice. Inbreeding of the dam reduced the probability of breeding, the probability of producing a second litter, and litter size. Inbreeding of the litter caused depression of litter size, juvenile viability, and mass at weaning, and caused an increase in the within-litter variance in mass. In spite of differences between the subspecies in natural population sizes, genetic variation, and mean rates of reproduction and survival, all variation observed between experimental populations in their responses to inbreeding could be attributed to random founder effects. The genetic load of deleterious alleles in each replicate was unequally partitioned among its founder pairs, and different founders contributed to the load affecting different fitness components. Thus, inbreeding depression for any one fitness component, in our experimental environment, must be due to relatively few deleterious alleles with major effects. Genetic loads so comprised would be expected to diverge among natural populations due to both random drift and selective removal of recessive deleterious alleles during population bottlenecks. The near universality of inbreeding depression would be maintained, however, if different alleles contribute to inbreeding depression of different fitness components and in different environments.

Inbreeding effects on fitness have most often been quantified via juvenile mortality. However, inbred adults may suffer from inbreeding depression if their fertility or fecundity is compromised as a consequence of inbreeding. Here, the effects of inbreeding on male fertility in oldfield mice, Peromyscus polionotus, were examined. Testicular sperm concentration was assessed in 93 males, 68 of which were paired for breeding. Forty of the 68 paired males failed to produce offspring. Total testicular sperm count, sperm count (g testis)(-1), and testis mass all declined significantly with increasing inbreeding coefficient. Sperm concentration did not significantly impact reproductive success. Although sperm concentration in males of most species can decline to low levels before reproductive impairment is detectable, the declines in testicular sperm concentration found here suggest that inbreeding can affect fertility in adult males. Furthermore, monitoring testicular sperm concentration could provide a mechanism to monitor potential declines in reproductive performance before population-level reproductive success is irreparably impaired. The implications for the management of small, captive and wild populations may be substantial.