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Do quantitative trait loci (QTL) for a courtship song difference between Drosophila simulans and D. sechellia coincide with candidate genes and intraspecific QTL?

The genetic architecture of traits influencing sexual isolation can give insight into the evolution of reproductive isolation and hence speciation. Here we report a quantitative trait loci (QTL) analysis of the difference in mean interpulse interval (IPI), an important component of the male courtship song, between Drosophila simulans and D. sechellia. Using a backcross analysis, we find six QTL that explain a total of 40.7% of the phenotypic variance. Three candidate genes are located in the intervals bounded by two of the QTL and there are no significant QTL on the X chromosome. The values of mean IPI for hybrid individuals imply the presence of dominant alleles or epistasis. Because unisexual hybrid sterility prevents an F(2) analysis, we cannot distinguish dominant from additive genetic effects at the scale of QTL. A comparison with a study of QTL for intraspecific variation in D. melanogaster shows that, for these strains, the QTL we have identified for interspecific variation cannot be those that contribute to intraspecific variation. We find that the QTL have bidirectional effects, which indicates that the genetic architecture is compatible with divergence due to genetic drift, although other possibilities are discussed.

Alleles↗

Genic similarity of American and European species of the lobster Homarus.

European lobsters (Homarus gammarus) from the Norway coast and from the Irish Sea are examined for electrophoretically detectable genetic variation in seventeen functionally different proteins. Forty-one loci encoding these proteins are homologous with loci studied in a previous survey of eight populations of H. americanus. Progeny hatched from ovigerous Norway females show variation in three enzymes, but Mendelian inheritance is confirmed only for triosephosphate isomerase and for one of the phosphoglucose isomerases. Complex PGI phenotypes are described. The average amounts of genetic variability in European and American lobster populations appear to be equivalent. More than one allele is detected at 20% of the loci, the average number of alleles detected per locus is 1.2 and the average proportion of loci heterozygous per individual is 4.0%. While much less genically variable than other invertebrates, Homarus is not atypical when compared with eleven decapod species that average 5.8% heterozygosity. This is consistent with hypotheses relating genetic variability to adaptive strategy. At thirty loci H. gammarus is monomorphic for the common H. americanus allele. Two acid phosphatase systems are fixed or nearly fixed for alternative alleles in the two species while the remaining polymorphic loci show various degrees of interspecific divergence. Unique H. gammarus alleles are detected at five loci but only contribute significantly to species differences at the Acph-5, Me, and Pgi-4 loci. Acph-1, Est-2, Pgi-3, and Pgm-1 are polymorphic for the same alleles in both species, but again, with various differences in allelic frequencies. In sum, average genetic identity and average genetic distance are: I = 0.896 +/- 0.007 and D = 0.110 +/- 0.007, respectively. Compared to the values for conspecific population comparisons, I = 0.994 +/- 0.001 and D - 0.006 +/- 0.001, it is clear that a small but significant amount of genetic divergence separates the European and American lobster. Based on the premise that protein differences between existing species reflect the amount of time since they shared a common ancestor, it can be speculated that the European and American lobsters were isolated during the Pleistocene. The apparent weakness of reproductive isolating barriers suggests that these populations have evolved allopatrically. Finally, quantification of species' genetic differences, together with recent successes in interspecific laboratory matings, implicates species hybridization as a potentially important breeding practice in lobster aquaculture.

Animals↗

Hybridization and postzygotic isolation patterns in pigeons and doves.

The study of the patterns of reproductive isolation in relation to species divergence is critical for the understanding of the process of speciation. Comparative analyses of this kind were previously conducted in Drosophila, Lepidoptera, frogs, ducks, and birds in general. In the present study, we used information from the literature to analyze hybrid inviability in relation to species divergence in pigeons and doves. Four main patterns arose from this analysis: (1) as in the other groups studied, F1 hybrid inviability gradually increases as species diverge, the time needed to reach total inviability being higher in birds than in the other groups; (2) as expected, the presence of geographic overlap does not influence the evolution of postzygotic isolation; (3) the percentage of unhatched eggs does not differ between hybrids of the first generation and the backcrosses, but it increases in the second hybrid generation; and (4) pigeons and doves follow Haldane's rule, as found in the other groups studied so far. The similarity between the results of this and previous studies contributes to the growing evidence suggesting that the patterns of the evolution of postzygotic isolation, and the process of speciation in general, are shared among animal groups.

Animals↗

Correspondence between sexual isolation and allozyme differentiation: a test in the salamander Desmognathus ochrophaeus.

Ethological reproductive isolation and genetic divergence across 26 protein loci were measured among populations of the salamander Desmognathus ochrophaeus in the southern Appalachian Mountains. Levels of ethological isolation varied from none to complete and were statistically significant for all but two pairings between populations inhabiting different mountain ranges. When geographic and genetic distances were treated as independent variables in multiple correlation analyses, they accounted for about half the variance in levels of ethological isolation. When genetic distance is held constant, the remaining relationship between ethological isolation and geographic distance is still statistically significant. When geographic distance is held constant, the remaining relationship between genetic distance and levels of ethological isolation is nonsignificant, as is the relationship between geographic distance and genetic distance when ethological isolation is held constant. Ethological isolation and genetic divergence evidently both reflect the gradual divergence of allopatric populations, but genetic distance is a poor predictor of ethological isolation in these salamanders.

Animals↗

Genetics of sexual isolation in females of the Drosophila simulans species complex.

Genetic analysis of hybrids between Drosophila simulans and D. sechellia shows that sexual isolation in females is caused by at least two genes, one on each major autosome, while the X chromosome has no effect. These results are similar to those of a previous study of hybrids between D. simulans and another sibling species, D. mauritiana. In this latter hybridization, each arm of the second chromosome carries genes causing sexual isolation in females, implying a total divergence of at least three loci. The genetic similarity between the D. simulans/D. mauritiana and D. simulans/D. sechellia hybridizations probably results from independent evolution and not phylogenetic artifacts, because the dominance relationships and behavioural interactions differ between the two hybridizations. The lack of an X-chromosome effect on sexual isolation contrasts with genetic studies of post-zygotic reproductive isolation, which invariably show strong effects of this chromosome.

Animals↗

Reinforcement drives rapid allopatric speciation.

Allopatric speciation results from geographic isolation between populations. In the absence of gene flow, reproductive isolation arises gradually and incidentally as a result of mutation, genetic drift and the indirect effects of natural selection driving local adaptation. In contrast, speciation by reinforcement is driven directly by natural selection against maladaptive hybridization. This gives individuals that choose the traits of their own lineage greater fitness, potentially leading to rapid speciation between the lineages. Reinforcing natural selection on a population of one of the lineages in a mosaic contact zone could also result in divergence of the population from the allopatric range of its own lineage outside the zone. Here we test this with molecular data, experimental crosses, field measurements and mate choice experiments in a mosaic contact zone between two lineages of a rainforest frog. We show that reinforcing natural selection has resulted in significant premating isolation of a population in the contact zone not only from the other lineage but also, incidentally, from the closely related main range of its own lineage. Thus we show the potential for reinforcement to drive rapid allopatric speciation.

Animals↗

Further studies on gene polymorphism in the mainbody and geographically isolated populations of Drosophila pseudoobscura.

We have examined polymorphism at 22 additional loci in the populations from the mainbody of Drosophila pseudoobscura and an isolated population from Bogotá, Columbia, which also shows partial reproductive isolation from mainbody populations. These studies extend our previous observations of reduced gene polymorphism and apparent lack of unique allele in the Bogotá population.

Alleles↗

Host plant influences on sex pheromone behavior of phytophagous insects.

The sexual behavior of phytophagous insects is often integrated in a variety of ways with their host plants. This integration may be manifested as effects or influences of host plants on insect physiology and behavior, including sex pheromone communication, that reflect strategies by insects to optimize mating and reproduction. Certain insects sequester or otherwise acquire host plant compounds and use them as sex pheromones or sex pheromone precursors. Other insects produce or release sex pheromones in response to particular host plant cues. Chemicals from host plants often synergize or otherwise enhance insect responses to sex pheromones. By these means, host plants may be used by insects to regulate or mediate sexual communication. For many species of insects, host plant influences on insect sex pheromone communication may be important aspects of the formation of feeding and mating aggregations, of insect strategies to locate both hosts and mates, of behavioral reproductive isolation among sibling species, and of the regulation of reproduction to coincide with the availability of food and oviposition sites. Knowledge of these relationships is critical to understanding many different areas of the behavioral ecology of plant-feeding insects.

Journal Article↗

[Genetic divergence of chars of the genus Salvelinus from Kronotsky Lake (Kamchatka Peninsula)].

Chars from the genus Salvelinus, inhabiting lakes and lake-river systems, belong to morphologically and ecologically different forms whose taxonomic status is under dispute. In the present work, we have examined genetic variation and divergence in various chars from the Kronotsky lake basin: the lacustrine chars (white, nose, and long-head) and Dolly Varden char Salvelinus malma. The study was conducted using analysis of allozyme and microsatellite loci, myogens, RAPD, and restriction analysis of two mtDNA segments. The estimates of heterozygoisty at allozyme and microsatellite loci were similar to the corresponding parameters in populations of northern Dolly Varden and Arctic char. Heterozygote deficit was recorded in both samples of individual forms, and in the combined sample of all chars from Kronotsky Lake. For both markers, appreciable genetic differentiation among the samples of different char forms was found, which was comparable to that among the spatially isolated populations of northern Dolly Varden. This result indicates reproductive isolation among the char forms examined. However, this isolation is not complete, because no fixed differences between the forms by any of the genetic systems analyzed was found. The genetic differentiation among different forms of lacustrine chars, which corresponds to the interpopulation rather than interspecies level, is thought to be explained by their comparatively recent divergence.

Animals↗

Postzygotic isolating factor in sympatric speciation in Rhagoletis flies: reduced response of hybrids to parental host-fruit odors.

Rhagoletis pomonella is a model for sympatric speciation (divergence without geographic isolation) by means of host-plant shifts. Many Rhagoletis species are known to use fruit odor as a key olfactory cue to distinguish among their respective host plants. Because Rhagoletis rendezvous on or near the unabscised fruit of their hosts to mate, behavioral preferences for fruit odor translate directly into premating reproductive isolation among flies. Here, we report that reciprocal F(1) hybrids between the apple and hawthorn host races of R. pomonella, as well as between the host races and an undescribed sibling species infesting Cornus florida (flowering dogwood) do not respond to host fruit volatiles in wind-tunnel assays at doses that elicit maximal directed flight in parental flies. The reduced ability of hybrids to orient to fruit volatiles could result from a conflict between neural pathways for preference and avoidance behaviors, and it suggests that hybrids might suffer a fitness disadvantage for finding fruit in nature. Therefore, host-specific mating may play a dual role as an important postzygotic as well as a premating reproductive barrier to isolate sympatric Rhagoletis flies.

Animals↗

Mode of mating preference and the direction of evolution in Drosophila.

Sexual (or ethological) isolation constitutes the most important class among different means of reproductive isolation in animal species. The phenomenon of sexual isolation has been extensively studied in the genus Drosophila. The degree of sexual isolation and the mode of mating preference are often used to indicate the phylogenetic relationship between the species and also their evolutionary sequence. Asymmetrical or one-sided mating preference observed in a number of Drosophila species groups has been used to elucidate the direction of evolution. Based on the mode of mating preference, different models which are even opposite to each other have been proposed by different investigators to predict the direction of evolution among closely related species. One-sided mating preference observed among the geographical strains of the same species has also been used to predict the direction of evolution. This review documents several cases of asymmetrical mating preference (both inter- and intraspecific) including the interesting cases reported by Indian workers and in the light of these observations of asymmetry in mating preference, the general applicability of different models is discussed.

Animals↗

Divergence in replicated phylogenies: the evolution of partial post-mating prezygotic isolation in bean weevils.

By tradition, speciation research has been focused on processes leading to either premating or post-zygotic reproductive isolation. The processes which generate isolation after mating but before zygote formation are less well understood. Here, we study divergence in characters which contribute to post-mating prezygotic isolation, such as egg production and remating rate. We propose that 'replicated' laboratory phylogenies with known histories can be used to yield insights into the processes of divergence. We performed a series of cross-matings between populations within two strains of the bean weevil Callosobruchus maculatus. Each strain has a unique and independent origin and both have been kept in the same set of laboratories during the last few decades. Our results show that divergence has occurred between laboratory populations within strains with regards to the effects that mating has on female reproductive behaviour, showing that the evolution of partial post-mating prezygotic isolation can be rapid. More importantly, the pattern of divergence across populations was distinct in the two strains, suggesting that coevolutionary trajectories are not determined by environmental factors but are to some extent arbitrary. We discuss the limitations of the novel empirical strategy employed here, and conclude that our results lend support to the hypothesis that post-mating sexual selection is capable of rapidly generating post-mating prezygotic isolation.

Animals↗

Social isolation partially restores reproduction of male staggerer mutant mice.

Social isolation is generally known to promote sexual performances in male mice. We employ here other indices to investigate the positive effects of social isolation on sexual behavior. Mutant staggerer male mice generally do not mate. However some of their reproductive traits can be restored by environmental modifications. We demonstrate that a postpubertal period of social isolation can significantly enhance mutant male sexual behavior. After a period of partial social isolation of one or 2 mo a significantly larger number of mutant males were observed to attempt mounting receptive non mutant C57BL/6 females during a 30 min test encounter. When reproductive success is considered, a larger number of males who were isolated for a period of 2 mo and then allowed to remain with non mutant females were found to sire pups than control (non-isolated) males. The results show that staggerer males isolated for 2 mo can partially overcome the reproductive deficit usually associated with the mutation.

Animals↗

Quantitative trait loci for sexual isolation between Drosophila simulans and D. mauritiana.

Sexual isolating mechanisms that act before fertilization are often considered the most important genetic barriers leading to speciation in animals. While recent progress has been made toward understanding the genetic basis of the postzygotic isolating mechanisms of hybrid sterility and inviability, little is known about the genetic basis of prezygotic sexual isolation. Here, we map quantitative trait loci (QTL) contributing to prezygotic reproductive isolation between the sibling species Drosophila simulans and D. mauritiana. We mapped at least seven QTL affecting discrimination of D. mauritiana females against D. simulans males, three QTL affecting D. simulans male traits against which D. mauritiana females discriminate, and six QTL affecting D. mauritiana male traits against which D. simulans females discriminate. QTL affecting sexual isolation act additively, are largely different in males and females, and are not disproportionately concentrated on the X chromosome: The QTL of greatest effect are located on chromosome 3. Unlike the genetic components of postzygotic isolation, the loci for prezygotic isolation do not interact epistatically. The observation of a few QTL with moderate to large effects will facilitate positional cloning of genes underlying sexual isolation.

Animals↗

Males, parthenogenesis, and the maintenance of anisogamous sex.

The problem of the maintenance of anisogamous sex is addressed by considering the effect of fertilization on the fitness of parthenogenetic females when such fertilization yields inviable triploid progeny. We consider four types of parthenogenesis: (i) apomixis, (ii) homogametic amphimixis, (iii) heterogametic amphimixis, and (iv) homogametic automixis. Homozygous sexual populations are genetically stable if males or selection eliminate the excess females produced by heterozygous parthenogenetic genotypes. Homozygous parthenogenetic populations are stable if the parthenogenetic output of homozygotes exceeds that of heterozygotes. In turn, sex can only invade heterozygous parthenogenetic populations when sexual output of parthenogens is larger than their parthenogenetic output. The existence of interior stable equilibria generally requires the instability of at least one boundary and some degree of heterosis. In a two-locus model, we study the evolution of mechanisms protecting either sex or parthenogenesis in reproductively polymorphic populations. We find that males do not respond to the presence of parthenogenesis in such a way as to eliminate it, but parthenogenesis is subject to selective pressures increasing reproductive isolation, and thus the success of parthenogenesis. The results suggest that reproductively polymorphic populations are ephemeral.

Animals↗

Eukaryotic microbes, species recognition and the geographic limits of species: examples from the kingdom Fungi.

The claim that eukaryotic micro-organisms have global geographic ranges, constituting a significant departure from the situation with macro-organisms, has been supported by studies of morphological species from protistan kingdoms. Here, we examine this claim by reviewing examples from another kingdom of eukaryotic microbes, the Fungi. We show that inferred geographic range of a fungal species depends upon the method of species recognition. While some fungal species defined by morphology show global geographic ranges, when fungal species are defined by phylogenetic species recognition they are typically shown to harbour several to many endemic species. We advance two non-exclusive reasons to explain the perceived difference between the size of geographic ranges of microscopic and macroscopic eukaryotic species when morphological methods of species recognition are used. These reasons are that microbial organisms generally have fewer morphological characters, and that the rate of morphological change will be slower for organisms with less elaborate development and fewer cells. Both of these reasons result in fewer discriminatory morphological differences between recently diverged lineages. The rate of genetic change, moreover, is similar for both large and small organisms, which helps to explain why phylogenetic species of large and small organisms show a more similar distribution of geographic ranges. As a consequence of the different rates in fungi of genetic and morphological changes, genetic isolation precedes a recognizable morphological change. The final step in speciation, reproductive isolation, also follows genetic isolation and may precede morphological change.

Classification↗

Speciation driven by natural selection in Drosophila.

Reinforcement is the process by which natural selection strengthens sexual isolation between incipient species, reducing the frequency of maladaptive hybridization and hence completing reproductive isolation. Although this model of speciation was once widely accepted, its plausibility and experimental support have been recently attacked. Here we provide an example of speciation by reinforcement, in the North American fruitfly Drosophila pseudoobscura. The results suggest that females of D. pseudoobscura evolved increased sexual isolation from their sibling species, D. persimilis, by natural selection against maladaptive hybridization.

Animals↗

Differential selection to avoid hybridization in two toad species.

The fitness consequences of hybridization critically affect the speciation process. When hybridization is costly, selection favors the evolution of prezygotic isolating mechanisms (e.g., mating behaviors) that reduce heterospecific matings and, consequently, enhance reproductive isolation between species (a process termed reinforcement). If, however, selection to avoid hybridization differs between species, reinforcement may be impeded. Here, we examined both the frequency and fitness effects of hybridization between plains spadefoot toads (Spea bombifrons) and New Mexico spadefoot toads (S. multiplicata). Hybridization was most frequent in smaller breeding ponds that tend to be ephemeral, and heterospecific pairs consisted almost entirely of S. bombifrons females and S. multiplicata males. Moreover, in controlled experimental crosses, hybrid offspring from crosses in which S. multiplicata was maternal had significantly lower survival and longer development time than pure S. multiplicata offspring. By contrast, hybrid offspring from crosses in which S. bombifrons was maternal outperformed pure S. bombifrons offspring by reaching metamorphosis faster. These data suggest that, although S. multiplicata females are under selection to avoid hybridization, selection might favor those S. bombifrons females that hybridize with S. multiplicata if their breeding pond is highly ephemeral. Generally, the strength of selection to avoid hybridization may differ for hybridizing species, possibly impeding reinforcement.

Animals↗