RESUMEN
Traits involved in post-copulatory interactions between the sexes may evolve rapidly as a result of sexual selection and/or sexual conflict, leading to post-mating-prezygotic (PMPZ) reproductive isolating barriers between diverging lineages. Although the importance of PMPZ isolation is recognized, the molecular basis of such incompatibilities is not well understood. Here, we investigate molecular evolution of a subset of Drosophila mojavensis and Drosophila arizonae reproductive tract genes. These include genes that are transcriptionally regulated by conspecific mating in females, many of which are misregulated in heterospecific crosses, and a set of male genes whose transcripts are transferred to females during mating. As a group, misregulated female genes are not more divergent and do not appear to evolve under different selection pressures than other female reproductive genes. Male transferred genes evolve at a higher rate than testis-expressed genes, and at a similar rate compared to accessory gland protein genes, which are known to evolve rapidly. Four of the individual male transferred genes show patterns of divergent positive selection between D. mojavensis and D. arizonae. Three of the four genes belong to the sperm-coating protein-like family, including an ortholog of antares, which influences female fertility and receptivity in Drosophila melanogaster. Synthesis of these molecular evolutionary analyses with transcriptomics and predicted functional information makes these genes candidates for involvement in PMPZ reproductive incompatibilities between D. mojavensis and D. arizonae.
Asunto(s)
Drosophila/genética , Evolución Molecular , Animales , Drosophila/fisiología , Proteínas de Drosophila/genética , Proteínas de Drosophila/metabolismo , Femenino , Regulación de la Expresión Génica/fisiología , Masculino , Filogenia , Reproducción/fisiología , Selección GenéticaRESUMEN
Lengths, widths and volumes of eggs from 11 species of Drosophila whose genomes have been fully sequenced exhibit significant variation that is not explained by their phylogenetic relationships. Furthermore, egg size differences are unrelated to embryonic development time in these species. In addition, two of the species, Drosophila sechellia and, to a lesser degree, D. yakuba, both ecological specialists, exhibit ovoviviparity, suggesting that female control over oviposition in these species differs from what is observed in D. melanogaster. The interspecific differences in these reproductive characters, coupled with the availability of whole genome sequences for each, provide an unprecedented opportunity to examine their evolution.
Asunto(s)
Drosophila/fisiología , Ovoviviparidad/fisiología , Cigoto/fisiología , Animales , Drosophila/embriología , Femenino , Masculino , Especificidad de la Especie , Factores de TiempoRESUMEN
The adaptive significance of enzyme variation has been of central interest in population genetics. Yet, how natural selection operates on enzymes in the larger context of biochemical pathways has not been broadly explored. A basic expectation is that natural selection on metabolic phenotypes will target enzymes that control metabolic flux, but how adaptive variation is distributed among enzymes in metabolic networks is poorly understood. Here, we use population genetic methods to identify enzymes responding to adaptive selection in the pathways of central metabolism in Drosophila melanogaster and Drosophila simulans. We report polymorphism and divergence data for 17 genes that encode enzymes of 5 metabolic pathways that converge at glucose-6-phosphate (G6P). Deviations from neutral expectations were observed at five loci. Of the 10 genes that encode the enzymes of glycolysis, only aldolase (Ald) deviated from neutrality. The other 4 genes that were inconsistent with neutral evolution (glucose-6-phosphate dehydrogenase [G6pd]), phosphoglucomutase [Pgm], trehalose-6-phosphate synthetase [Tps1], and glucose-6phosphatase [G6pase] encode G6P branch point enzymes that catalyze reactions at the entry point to the pentose-phosphate, glycogenic, trehalose synthesis, and gluconeogenic pathways. We reconcile these results with population genetics theory and existing arguments on metabolic regulation and propose that the incidence of adaptive selection in this system is related to the distribution of flux control. The data suggest that adaptive evolution of G6P branch point enzymes may have special significance in metabolic adaptation.
Asunto(s)
Adaptación Fisiológica/genética , Drosophila melanogaster/genética , Drosophila melanogaster/metabolismo , Evolución Molecular , Redes y Vías Metabólicas/genética , Animales , Datos de Secuencia MolecularRESUMEN
Fruit flies of the genus Drosophila have independently invaded deserts around the world on numerous occasions. To understand the physiological mechanisms allowing these small organisms to survive and thrive in arid environments, we performed a phylogenetic analysis of water balance in Drosophila species from different habitats. Desert (cactophilic) species were more resistant to desiccation than mesic ones. This resistance could be accomplished in three ways: by increasing the amount of water in the body, by reducing rates of water loss or by tolerating the loss of a greater percentage of body water (dehydration tolerance). Cactophilic Drosophila lost water less rapidly and appeared to be more tolerant of low water content, although males actually contained less water than their mesic congeners. However, when the phylogenetic relationships between the species were taken into account, greater dehydration tolerance was not correlated with increased desiccation resistance. Therefore, only one of the three expected adaptive mechanisms, lower rates of water loss, has actually evolved in desert Drosophila, and the other apparently adaptive difference between arid and mesic species (increased dehydration tolerance) instead reflects phylogenetic history.