Rapid divergence of microsatellite abundance among species of Drosophila

Among major taxonomic groups, microsatellites exhibit considerable variation in composition and allele length, but they also show considerable conservation within many major groups. This variation may be explained by slow microsatellite evolution so that all species within a group have similar patterns of variation, or by taxon-specific mutational or selective constraints. Unfortunately, comparing microsatellites across species and studies can be problematic because of biases that may exist among different isolation and analysis protocols. We present microsatellite data from five Drosophila species in the Drosophila subgenus: D. arizonae, D. mojavensis, and D. pachea (three cactophilic species), and D. neotestacea and D. recens (two mycophagous species), all isolated at the same time using identical protocols. For each species, we compared the relative abundance of motifs, the distribution of repeat size, and the average number of repeats. Dimers were the most abundant microsatellites for each species. However, we found considerable variation in the relative abundance of motif size classes among species, even between sister taxa. Frequency differences among motifs within size classes for the three cactophilic species, but not the two mycophagous species, are consistent with other studied Drosophila. Frequency distributions of repeat number, as well as mean size, show significant differences among motif size classes but not across species. Sizes of microsatellites in these five species are consistent with D. virilis, another species in the subgenus Drosophila, but they have consistently higher means than in D. melanogaster, in the subgenus Sophophora. These results confirm that many aspects of microsatellite variation evolve quickly but also are subject to taxon-specific constraints. In addition, the nature of microsatellite evolution is dependent on temporal and taxonomic scales, and some variation is conserved across broad taxonomic levels despite relatively high rates of mutation for these loci.     

Perspective: female remating,operational sex ratio,and the arena of sexual selection in Drosophila species

Abstract.— As commonly observed among closely related species within a variety of taxa, Drosophila species differ considerably in whether they exhibit sexual dimorphism in coloration or morphology. Those Drosophila species in which male external sexual characters are minimal or absent tend, instead, to have exaggerated ejaculate traits such as sperm gigantism or seminal nutrient donations. Underlying explanations for the interspecific differences in the presence of external morphological sexual dimorphism versus exaggerated ejaculate traits are addressed here by examining the opportunity for sexual selection on males to occur before versus after mating in 21 species of Drosophila . Female remating frequency, an important component of the operational sex ratio, differs widely among Drosophila species and appears to dictate whether the arena of sexual selection is prior to, as opposed to after, copulation. Infrequent female mating results in fewer mating opportunities for males and thus stronger competition for receptive females that favors the evolution of male characters that maximize mating success. On the other hand, rapid female remating results in overlapping ejaculates in the female reproductive tract, such that ejaculate traits which enhance fertilization success are favored. The strong association between female remating frequency in a given species and the presence of sexually selected external versus internal male characters indicates that the relationship be examined in other taxa as well.     

Developmental stability in hybrids between the sibling species pair,Drosophila melanogaster and Drosophila simulans

Drosophila melanogaster and its sibling species D. simulans were hybridized in the laboratory to test the hypothesis that developmental homeostasis in hybrids between two species having no prior gene flow would be significantly reduced. Developmental stability was assessed by measuring fluctuating asymmetry for three bilateral traits: sternopleural chaetae, wing length, and fronto-orbital plus frontal chaetae. Male F₁ hybrids showed no decrease in developmental stability compared to males of parental species. Female hybrids showed significant fluctuating asymmetry compared to other flies. The results are discussed with respect to ideas about coadaptation and gene flow based upon previous studies of hybrid developmental stability.     

Analysis of Drosophila species genome size and satellite DNA content reveals significant differences among strains as well as between species

The size of eukaryotic genomes can vary by several orders of magnitude, yet genome size does not correlate with the number of genes nor with the size or complexity of the organism. Although "whole"-genome sequences, such as those now available for 12 Drosophila species, provide information about euchromatic DNA content, they cannot give an accurate estimate of genome sizes that include heterochromatin or repetitive DNA content. Moreover, genome sequences typically represent only one strain or isolate of a single species that does not reflect intraspecies variation. To more accurately estimate whole-genome DNA content and compare these estimates to newly assembled genomes, we used flow cytometry to measure the 2C genome values, relative to Drosophila melanogaster. We estimated genome sizes for the 12 sequenced Drosophila species as well as 91 different strains of 38 species of Drosophilidae. Significant differences in intra- and interspecific 2C genome values exist within the Drosophilidae. Furthermore, by measuring polyploid 16C ovarian follicle cell underreplication we estimated the amount of satellite DNA in each of these species. We found a strong correlation between genome size and amount of satellite underreplication. Addition and loss of heterochromatin satellite repeat elements appear to have made major contributions to the large differences in genome size observed in the Drosophilidae.     

Multilocus nuclear sequences reveal intra- and interspecific relationships among chromosomally polymorphic species of cactophilic Drosophila

Drosophila mojavensis and Drosophila arizonae, a pair of sibling species endemic to North America, constitute an important model system to study ecological genetics and the evolution of reproductive isolation. This species pair can produce fertile hybrids in some crosses and are sympatric in a large part of their ranges. Despite the potential for hybridization in nature, however, evidence of introgression has not been rigorously sought. Further, the evolutionary relationships within and among the geographically distant populations of the two species have not been characterized in detail using high-resolution molecular studies. Both species have six chromosomes: five large acrocentrics and one 'dot' chromosome. Fixed inversion differences between the species exist in three chromosomes (X, 2 and 3) while three are colinear (4, 5 and 6), suggesting that were introgression to occur, it would be most likely in the colinear chromosomes. We utilized nucleotide sequence variation at multiple loci on five chromosomes to test for evidence of introgression, and to test various scenarios for the evolutionary relationships of these two species and their populations. While we do not find evidence of recent introgression, loci in the colinear chromosomes appear to have participated in exchange in the past. We also found considerable population structure within both species. The level of differentiation discovered among D. arizonae populations was unexpectedly high and suggests that its populations, as well as those of D. mojavensis, may be themselves undergoing incipient speciation and merit further attention.     

Elemental stoichiometry of Drosophila and their hosts

1. Nitrogen (N) and phosphorus (P) availabilities are important ecological determinants of resource use in nature. Despite the wide range of hosts used by species of the genus Drosophila , elemental composition of natural resources of these flies has never been investigated.
2. Total body N and P contents were determined in seven species of wild-caught Drosophila , their natural hosts, and artificial diets routinely used to rear these flies in the laboratory. The flies tested included D. hydei, D. arizonae, D. simulans and D. pseudoobscura collected from rotting fruit (melons), and the cactophilic D. nigrospiracula, D. mojavensis and D. pachea collected from their specific host plants, Saguaro, Organpipe and Senita cactus, respectively.
3. Natural hosts varied in elemental composition, with fruit showing higher N (2·8–4·3% dry mass) and P (0·50–0·67%) levels compared with cacti (0·5–1·6% N; 0·01–0·29% P). No consistent differences in N and P levels were found between healthy and necrotic cactus tissue.
4. Total body N and P also varied among Drosophila species. This variation mirrored the levels of N and P found in the respective hosts and laboratory diets. N:P ratios were consistently lower in female flies compared with conspecific males suggesting phosphorus demands during oogenesis are high.
5. Potential mechanisms by which Drosophila deal with N or P limitation in nature are discussed.     

Reproductive Isolation Among <Emphasis Type="Italic">Drosophila arizonae</Emphasis> from Geographically Isolated Regions of North America

A long-standing goal of speciation research is to describe how reproductive isolating barriers develop, when they arise along the ‘speciation continuum’, and to measure the strength with which they restrict gene flow. Drosophila arizonae and D. mojavensis are a recently diverged sister species pair distributed from the southwestern United States through southern Mexico. While incipient speciation in D. mojavensis has been studied for decades, relatively little attention has been directed toward D. arizonae, despite the fact that previous studies have revealed evidence for significant genetic differentiation among populations separated by geographic barriers. Here, we examine the potential for both pre- and post-mating reproductive isolation in D. arizonae from geographically isolated parts of North America. We find evidence for strong premating isolation between flies from northern mainland Mexico and southern mainland Mexico, but no evidence for postmating isolation in any cross. This study highlights the utility of the D. arizonae system for further investigation into the early evolution of premating isolation, and reinforces the potential of the D. arizonae/D. mojavensis system as a whole for studying the evolution of reproductive isolation at a range of divergence times.     

Human handedness and the concept of developmental stability

A model is proposed to explain the etiology of pathological handedness. Developmental instability, caused by elevated genotypic homozygosity, environmental disturbances, or their interaction, overrides programmed laterality and handedness in the same way that it perturbs the bilaterally symmetrical expression of morphological and metric traits. The model predicts that pathological handedness should be elevated among individuals with higher than average homozygosity and individuals who have developed under unfavorable uterine environments. Suggestions are offered for specific populations in which the predictions may be tested.