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Artificial selection affects phenotypes differently by natural selection. Domestic traits, which pass into the wild, are usually negatively selected. Yet, exceptionally, this axiom may fail to apply if genes, from the domestic animals, increase fertility in the wild. We studied a rare case of a wild boar population under the framework of Wright's interdemic selection model, which could explain gene flow between wild boar and pig, both considered as demes. We analysed the MC1R gene and microsatellite neutral loci in 62 pregnant wild boars as markers of hybridization, and we correlated nucleotide mutations on MC1R (which are common in domestic breeds) to litter size, as an evaluation of fitness in wild sow. Regardless of body size and phyletic effects, wild boar sows bearing nonsynonymous MC1R mutations produced larger litters. This directly suggests that artificially selected traits reaching wild populations, through interdemic gene flow, could bypass natural selection if and only if they increase the fitness in the wild.

Interdemic selection by the differential migration of individuals out from demes of high fitness and into demes of low fitness (Phase III) is one of the most controversial aspects of Wright's Shifting Balance Theory. I derive a relationship between Phase III migration and the interdemic selection differential, S, and show its potential effect on FST. The relationship reveals a diversifying effect of interdemic selection by Phase III migration on the genetic structure of a metapopulation. Using experimental metapopulations, I explored the effect of Phase III migration on F ST by comparing the genetic variance among demes for two different patterns of migration: (1) island model migration and (2) Wright's Phase III migration. Although mean migration rates were the same, I found that the variance among demes in migration rate was significantly higher with Phase III than with island model migration. As a result, F ST for the frequency of a neutral marker locus was higher with Phase III than it was with island model migration. By increasing F ST , Phase III enhanced the genetic differentiation among demes for traits not subject to interdemic selection. This feature makes Wright's process different from individual selection which, by reducing effective population size, decreases the genetic variance within demes for all other traits. I discussed this finding in relation to the efficacy of Phase III and random migration for effecting peak shifts, and the contribution of genes with indirect effects to among-deme variation.

We critically review the two major theories of adaptive evolution developed early in this century, Wright's shifting balance theory and Fisher's large population size theory, in light of novel findings from field observations, laboratory experiments, and theoretical research conducted over the past 15 years. Ecological studies of metapopulations have established that the processes of local extinction and colonization of demes are relatively common in natural populations of many species and theoretical population genetic models have shown that these ecological processes have genetic consequences within and among local demes. Within demes, random genetic drift converts nonadditive genetic variance into additive genetic variance, increasing, rather than limiting, the potential for adaptation to local environments. For this reason, the genetic differences that arise by drift among demes, can be augmented by local selection. The resulting adaptive differences in gene combinations potentially contribute to the genetic origin of new species. These and other recent findings were not discussed by either Wright or Fisher. For example, although Wright emphasized epistatic genetic variance, he did not discuss the conversion process. Similarly, Fisher did not discuss how the average additive effect of a gene varies among demes across a metapopulation whenever there is epistasis. We discuss the implications of such recent findings for the Wright-Fisher controversy and identify some critical open questions that require additional empirical and theoretical study.

Using demes from experimental metapopulations of the flour beetle,Tribolium castaneum, we investigated phase 3 of Wright's shifting balance process. Using parent demes of high, intermediate, and low mean fitness, we experimentally modeled migration of varying amounts from demes of high mean fitness into demes of lower mean fitness (like phase 3) as well as the reciprocal (the opposite of phase 3). In natural populations, some migration among demes occurs independently of deme fitness. In this case, demes of high mean fitness are likely to receive migrants from demes of lower mean fitness; these effects might limit the effectiveness of phase 3 but have not been studied experimentally. We estimated the populational heritability of mean fitness by the regression of offspring deme means on the weighted parental means and found moderate levels of demic heritability one (0.641‐0.690) and two (0.518‐0.552) generations after migration. We discuss our findings in relation to the role of interdemic migration in “adaptive peak shifts” in metapopulations and the controversies over group selection and the units of inheritance.

The relationship between social structure and partitioning of genetic variance was examined in two red howler monkey populations (W and G) in Venezuela, one of which (G) was undergoing rapid growth through colonization by new troops. Rates and patterns of gene flow had been determined through radiotelemetry and direct observation data on solitary migrants, and 10 years of troop censusing. Standard electrophoresis techniques were used to examine 29 loci in blood samples taken from 137 of the study animals. Analysis of genetic variance demonstrated: (1) a significantly high level of genetic variation among troops within populations (FST= 0.225 for W and 0.142 for G), and (2) a significant excess of heterozygosity within troops relative to expected (FIS = -0.136 for W and -0.064 for G), despite relatively high levels of observed and inferred inbreeding in W. Differences between the populations in FSTvalues conformed to those predicted based on differences in colonization rate. Comparison of partitioning of genetic variance among different genealogical subsets of troops demonstrated that the pattern of genetic differentiation observed among troops within populations was promoted by an essentially single-male harem breeding structure, a very low rate of random exchange of breeding males among troops, and a high degree of relatedness among troop females. Between-troop genetic differentiation (FST) was thereby increased relative to that expected from other types of socxal organization, while the correlation between uniting gametes within troops (Fis) was decreased. Genetic differentiation between populations (2%) corresponded to that predicted from migration rates. Such a mosaic of genetic variation, combined with differences in reproductive success observed among troops and a high troop failure rate, create conditions m which interdemic selection could result in more rapid spread of advantageous gene combinations than would be expected in a panmictic population, particularly in a colonizing situation m which the founder population is small.

.— Using data from three years (1994–1996), I tested whether differential migration occurs from demes of high mean fitness in the shining fungus beetle, Phalacrus substriatus. The results show evidence for differential migration, thus providing evidence from a natural population for a critical demographic assumption of many interdemic selection models. To predict the evolutionary response to interdemic selection through differential migration, the genetic basis of the variation among demes in mean fitness must be known because the observed patterns could also be explained by some demes having an intrinsically favorable habitat. Thus, the importance of differential migration through interdemic selection in natural populations cannot be unequivocally answered without experiments specifically addressing the question of what causes differences in mean fitness among demes.

In this article, we reply to Getty's (1999, in this issue) critique of laboratory studies of interdemic selection, particularly the study of Wade and Goodnight (1991). Getty (1999) raises several issues regarding the interpretation of experimental studies of group selection (reviewed in Goodnight and Stevens 1997). The first issue concerns offspring production, or demic productivity, the group-level trait used in several experimental studies of flour beetle metapopulations. This trait, unlike other traits, has the property that random sampling of offspring from within a deme is individual selection. The second issue is the different interpretations of hard and soft selection vis-a-vis group selection in Wade (1985b) and Goodnight et al. (1992). The last is the relationship between hard and soft selection and the appropriate experimental control for artificial interdemic selection on the trait, demic productivity. Getty (1999) argues that there was no group selection and only individual selection in the experimental treatments of Wade and Goodnight (1991) and that there was group selection in the controls. We address each of these issues below.

Animals might behave in ways that are detrimental to themselves if their behavior enhances the survival of the group in which they live. This model does not require that the animals in the group be close relatives. Group selection is an alternative to kin selection in explaining the evolution of altruism.

Using data from three years (1994–1996), I tested whether differential migration occurs from demes of high mean fitness in the shining fungus beetle, Phalacrus substriatus. The results show evidence for differential migration, thus providing evidence from a natural population for a critical demographic assumption of many interdemic selection models. To predict the evolutionary response to interdemic selection through differential migration, the genetic basis of the variation among demes in mean fitness must be known because the observed patterns could also be explained by some demes having an intrinsically favorable habitat. Thus, the importance of differential migration through interdemic selection in natural populations cannot be unequivocally answered without experiments specifically addressing the question of what causes differences in mean fitness among demes. Corresponding Editor: C. Boggs

Wright partitioned the shifting-balance process into three phases. Phase one is the shift of a deme within a population to the domain of a higher adaptive peak from that of the historical peak. Phase two is mass selection within a deme towards that higher peak. Phase three is the conversion of additional demes to the higher peak. The migration rate between demes is critical for the existence of phases one and three. Phase one requires small effective population sizes, hence low migration rates. Phase three is optimal under high migration rates that spread the most-fit genotype from deme to deme. Thus, a population-wide peak shift requires intermediate levels of migration. By altering the rates of phases one and three, migration affects the predominant direction of mass selection within a population. This study examines the degree to which migration, through its effects on phases one and three, determines the probability of a simulated population arriving at its genotypic optimum after 12,000 generations. These simulations reveal that there is a range of migration rates for which an entire population might be expected to shift to a higher peak. Below m = 0.001 peak shifts occur frequently (phases I and II) but are not successfully exported out of subpopulations (phase III), and above 0.01 peak shifts within demes (phase I and II), required to initiate phase III, become increasingly uncommon. Because it is unlikely that real populations will have uniform migration rates from generation to generation, the probable effects of varying migration rates on broadening the range of conditions producing peak shifts are discussed.