Class II DRB polymorphism and sequence diversity in two vesper bats in the genus Myotis.

Almost no studies have been done with respect to major histocompatibility complex (MHC) polymorphism and sequence diversity in bats, although they account for one in five living mammalian species. We analysed MHC Class II DRB polymorphism and sequence diversity in two Mexican verpertilionid bat species, the widespread continental species Myotis velifer and the narrowly distributed (and endangered) island endemic Myotis vivesi. We find extensive DRB polymorphism in the widespread M. velifer, similar to that commonly reported in other mammals. The geographically restricted M. vivesi by contrast shows only very limited polymorphism. We conclude that M. vivesi has undergone a dramatic loss of MHC polymorphism. The significance of this inference in light of other information on population structure and genetic diversity in this species is discussed.

Evidence for balancing selection at the DAB locus in the axolotl, Ambystoma mexicanum.

The axolotl (Ambystoma mexicanum) has been characterized as immunodeficient, and the absence of major histocompatibility complex (MHC) class II polymorphism has been cited as a possible explanation. Here we present evidence for considerable allelic polymorphism at the MHC class II DAB locus for a sample of wild-caught axolotls. Evidence that these sequences are the product of balancing selection for disease resistance is discussed.

Characterization of Peromyscus MHC class II beta sequences by ligation-anchored RT-PCR and denaturing gradient gel electrophoresis.

We characterized MHC class II beta sequences in deermice (genus Peromyscus), which are amongst the most common mammals in North America. We find evidence for two different families of class II beta loci, most closely related to either the A-type or E-type loci in Mus. Population studies suggest the presence of more than one A-type locus but only one E-type locus for the species investigated.

MHC class II beta sequence diversity in the deer mouse (Peromyscus maniculatus): implications for models of balancing selection.

We studied population polymorphism at a major histocompatibility complex (MHC) class II beta gene in the deer mouse (Peromyscus maniculatus). We found that: (i) a single population of P. maniculatus has significantly higher levels of DNA and protein sequence diversity than worldwide samples from homologous genes in other taxa, including humans and mice; and (ii) the genealogy of allelic sequences in P. maniculatus deviates significantly from theoretical expectation under a model of symmetric balancing selection, in that alleles are relatively more divergent than expected. We suggest that the observation of high levels of pairwise allelic sequence divergence and deviation of the genealogy from theoretical expectation in P. maniculatus together provide support for a divergent allele advantage model for the maintenance of MHC polymorphism.

Evolutionary genetics of self-incompatibility in the Solanaceae.

The self-incompatibility (S) gene in flowering plants has long been appreciated as an example of extreme allelic polymorphism maintained by frequency-dependent selection. Recent studies of population samples of S-allele sequences obtained by RT-PCR from five species of Solanaceae now reveal a picture of conspicuous inter-specific variation in both S-allele number and age. Explanations for this variation are examined with reference to current theory. We propose that changes in species' effective population size, particularly those associated with the evolution of different life histories, best account for interspecific differences in both the number and average age of S alleles.

Self-incompatibility alleles from Physalis: implications for historical inference from balanced genetic polymorphisms.

Balanced genetic polymorphism has been proposed as a source from which to infer population history complementary to that of neutral genetic polymorphism, because genetic polymorphism maintained by balancing selection permits inferences about population size over much longer spans of time. However, empirical data for both S genes and major histocompatibility complex genes do not fit expectations of coalescent theory. Species-specific gene genealogies have longer terminal branches than expected, indicating an apparent slowdown in the origination of new alleles. Here, we present evidence that divergent S alleles were selectively maintained in Physalis cinerascens during a reduction in population size, generating longer terminal branches in the S gene genealogy relative to the congener Physalis crassifolia. Retention of divergent alleles during reduction in the number of alleles violates assumptions of the coalescent model used to estimate effective population size. Recent theoretical and empirical results are consistent with the proposition that nonrandom sorting is a general property of balanced genetic polymorphisms, suggesting that studies of balanced polymorphism that infer the absence of population bottlenecks may overestimate effective population size.

Allelic diversity and gene genealogy at the self-incompatibility locus in the Solanaceae.

The self-incompatibility (S) locus of flowering plants offers an example of extreme polymorphism maintained by balancing selection. Estimates of recent and long-term effective population size (Ne) were determined for two solanaceous species by examination of S-allele diversity. Estimates of recent Ne in two solanaceous species differed by an order of magnitude, consistent with differences in the species' ecology. In one species, the evidence was consistent with historical population restriction despite a large recent Ne. In the other, no severe bottleneck was indicated over millions of years. Bottlenecks are integral to founder-event speciation, and loci that are subject to balancing selection can be used to evaluate the frequency of this mode of speciation.

S-allele diversity in a natural population of Physalis crassifolia (Solanaceae) (ground cherry) assessed by RT-PCR.

Allelic diversity at the self-incompatibility (S-) locus in the ground cherry, Physalis crassifolia (Solanaceae), was surveyed in a natural population occurring in Deep Canyon, CA, using a molecular assay to determine the genotype of individual plants. A total of 28 different S-alleles were identified and sequenced from a sample of 22 plants. All plants examined were heterozygous, as expected under gametophytic self-incompatibility (GSI). The estimated number of alleles in this population is 43-44, comparable to allelic diversity reported for other species, as determined by the standard diallel crossing method. Allele frequencies in the sample deviated from the expectation of equal frequency under GSI; it is suggested that this deviation may result from sampling of related individuals. Molecular analysis of genotypes within single pollen donor families indicates that, for all alleles examined, segregation is consistent with predictions for single-locus GSI. The implications of a reliable and efficient molecular assay for determining the S-genotype of plants are discussed.

Learning from rejection: the evolutionary biology of single-locus incompatibility.

The self-incompatibility (S-) locus of flowering plants is among the most polymorphic known. PCR methods can now be used to estimate both the number of alleles in natural populations and their sequence diversity. The number of alleles provides an estimate of recent effective population size, thus the S-locus provides a tool for examining how species characteristics affect population size. Sequence relationships among alleles provide another estimate of population size extending millions of years into the past. Relationships between S-alleles and related genes provide a means of dating the age of origin of incompatibility systems and determining which, if any, angiosperm families share incompatibility by homology.

S-allele sequence diversity in natural populations of Solanum carolinense (Horsenettle).

S-allele diversity in Solanum carolinense was surveyed in two natural populations, located in Tennessee and North Carolina, with a molecular assay to determine the genotype of individual plants. A total of 13 different S-alleles were identified and sequenced. There is high overlap between the two populations sampled, with 10 alleles shared in common, one allele found only in Tennessee, and two found only in North Carolina. The number of alleles in this species appears to be extremely low compared with other species with gametophytic self-incompatibility. Sequence comparisons show that most alleles are extremely different one from another in their primary sequence and a phylogenetic analysis indicates extensive trans-specific evolution of S-lineages. In addition, some alleles appear to be derived much more recently. The implications of these observations are discussed in the light of recent theoretical results on S-allele population diversity and persistence.

Evolution of ecological differences in the Old World leaf warblers.

Sympatric species that belong to the same ecological guild usually differ in their behaviour and morphology, and these differences are often interpreted as adaptations to having to make use of different resources. Evidence supporting this interpretation comes from association between ecology and morphology among species, in which an a priori functional relationship is reasonable. But one problem with such comparisons is that members of a guild may be closely related, so the more closely related species can share a greater similarity in their morphology and ecology simply as a result of the lingering legacy of a common ancestor. In principle, the importance of historical legacy can be evaluated from phylogenetic relationships and times since divergence for all species, but this is rarely possible because these data are not available. Here we use a phylogeny for eight sympatric species of warbler in the genus Phylloscopus, based on their mitochondrial DNA sequences, to remove the effects of historical legacy. Without these effects, we find strong support for adaptive interpretations of among-species variation in habitat selection, prey-size choice and feeding method. Ecological variation along any of these three niche axes is associated with predictable morphological variation. We also find evidence for historical legacy in that more closely related species are often more similar behaviourally and morphologically. This paradoxical result can be reconciled because the most closely related species tend to differ along only one niche axis, habitat choice. In contrast, the evolution of prey-size choice and feeding method occurred rapidly and early in the diversification of this group. Once a new ecological zone was occupied, subsequent morphological change along these niche axes was limited, accounting for the similarity of closely related species.