Germline-dependent gene expression in distant non-gonadal somatic tissues of Drosophila.

BACKGROUND: Drosophila females commit tremendous resources to egg production and males produce some of the longest sperm in the animal kingdom. We know little about the coordinated regulation of gene expression patterns in distant somatic tissues that support the developmental cost of gamete production. RESULTS: We determined the non-gonadal gene expression patterns of Drosophila females and males with or without a germline. Our results show that germline-dependent expression in the non-gonadal soma is extensive. Interestingly, gene expression patterns and hormone titers are consistent with a hormone axis between the gonads and non-gonadal soma. CONCLUSIONS: The germline has a long-range influence on gene expression in the Drosophila sexes. We suggest that this is the result of a germline/soma hormonal axis.

Evolution of cuticular hydrocarbons of Hawaiian Drosophilidae.

Hawaiian Drosophila offer an excellent model for adaptive evolution. More than 500 species are reported in Hawaiian islands, and there is considerable diversity in behavior and morphology. Such diversity is mainly driven by sexual selection. In this study qualitative and quantitative chemical compositions of cuticular hydrocarbons (CHCs) in 138 flies belonging to 27 Hawaiian Drosophila species, picture-winged and non picture-winged, were analyzed regarding sexual dimorphism, differences in saturation, branching position, and lengths of CHCs. We found significant variation in the CHC patterns. In several subgroups, new species show decreases in unsaturated hydrocarbons, and gradual increases in branched compounds, monomethylalkanes and dimethylalkanes, not commonly found in Drosophila. Moreover, branching positions gradually shifted towards internal carbons, and chain lengths increased in the new species. The long-term evolution of CHCs in the light of the recent evolutionary migration and adaptation history of Hawaiian Drosophila species along the developing archipelago was discussed.

Ontogeny of Drosophila melanogaster female sex-appeal and cuticular hydrocarbons.

The chemical communication system in Drosophila melanogaster Meigen, 1830 plays a major role in courtship and consists of the male-specific cis-Vaccenyl acetate and sex-specific contact pheromones, cuticular hydrocarbons (CHC), which build up during ontogeny (first 4 days). They replace longer CHCs, common to both sexes and present only after the imaginal eclosion. A detailed quantitative description of the evolution of cuticular unsaturated hydrocarbons with age is presented here for males and females of different D. melanogaster strains, which have been bred in well controlled environments. Monoenes appear in both sexes at around 12 h, before female dienes. The present paper argues that this is likely linked to the switching on of a new set of genes. Ecdysone, which is more abundant in females than in males during this critical period, might control this switch. Parallel behavioral studies show that whereas female of all ages trigger early mature male courtship steps like wing vibration, only females older than 1 day trigger late courtship steps like attempted copulation. This supports the hypothesis that late male courtship steps might be triggered by the CHCs, which build up after this age, especially female-specific (Z,Z)-7,11 - and (Z,Z)-5,9-dienes.

Role of the ejaculatory bulb in biosynthesis of the male pheromone cis-vaccenyl acetate in Drosophila melanogaster.

In Drosophila melanogaster, the male ejaculatory bulb is the site of synthesis of a male-specific pheromone, cis-vaccenyl acetate, which functions as both an attractant and an anti-aphrodisiac. This long monounsaturated acetate is structurally similar to a number of shorter gland-synthesized moth pheromones. The cell monolayer that forms the Drosophila male ejaculatory bulb wall is responsible for the production and secretion of cis-vaccenyl acetate into the seminal fluid. When dissected bulbs were incubated with sodium [14-C]-acetate (or deuterated acetate), a labeled acetate ester was synthesized. The labeled acetate ester co-migrated with cis-vaccenyl acetate in thin layer chromatography. Incubation of the abdomens of males from which the ejaculatory bulbs had been removed, or the abdomens of females, with radiolabeled acetate did not yield any acetate ester, but did yield other lipid products, including hydrocarbons. When the isolated labeled acetate ester was hydrolyzed, no radioactive vaccenol was formed. This strongly suggests that the acetyl group is incorporated via a transacetylation reaction, but that the vaccenyl moiety is not synthetized in the blub. The transacetylation enzyme activity was localized in the microsomal subfraction of the bulb homogenate, and its affinity for vaccenol was not very different from that reported for monounsaturated alcohol substrates in moths.

Sexual isolation and cuticular hydrocarbon differences between Drosophila santomea and Drosophila yakuba.

Drosophila santomea and Drosophila yakuba are two sister species inhabiting Saõ Tomé island. Previous studies showed that both species display strong reproductive isolation, although they can produce a few viable hybrids. Our study tried to understand the mechanism of this ethological isolation between two allopatric strains. A strong sexual isolation was confirmed, with a marked asymmetry. Comparisons of latency times to either courtship or copulation suggest that males do not discriminate females, whereas D. yakuba females, but not D. santomea females, accept their homospecifics more quickly. Cuticular hydrocarbon compositions of both species and sexes were also established with gas chromatography (GC) and GC/mass spectrometry analysis. All have (Z)-7-tricosene as their major compound. There are several quantitative differences between species for few minor compounds. The largest difference concerns n-heneicosane, which is more abundant in D. santomea than in D. yakuba flies (up to seven times more between males). A similar quantitative difference was also found in a pair of sympatric strains. Furthermore, D. yakuba males artificially perfumed with n-heneicosane were discriminated negatively by D. yakuba females, suggesting a role for this compound in the sexual isolation between these two species.

Quantitative trait loci for cuticular hydrocarbons associated with sexual isolation between Drosophila simulans and D. sechellia.

The identification of genes with large effects on sexual isolation and speciation is an important link between classic evolutionary genetics and molecular biology. Few genes that affect sexual isolation and speciation have been identified, perhaps because many traits influencing sexual isolation are complex behaviors. Cuticular hydrocarbons (CHs) of species of the Drosophila melanogaster group play a large role in sexual isolation by functioning as contact pheromones influencing mate recognition. Some of the genes that play key roles in determining species-specific CHs have been identified. We have performed separate quantitative trait locus (QTL) analyses of 7-tricosene (7-T) and 7,11-heptacosadiene (7,11-HD), the two major female CHs differing between D. simulans and D. sechellia. We find that approximately 40% of the phenotypic variance in each CH is associated with two to four chromosomal regions. A region on the right arm of chromosome 3 contains QTL that affect both traits, but other QTL are in distinct chromosomal regions. Epistatic interactions were detected between two pairs of QTL for 7,11-HD such that if either were homozygous for the D. simulans allele, the fly was similar to D. simulans in phenotype, with a low level of 7,11-HD. We discuss the location of these regions with regard to candidate genes for CH production, including those for desaturases.

Female-specific regulation of cuticular hydrocarbon biosynthesis by dopamine in Drosophila melanogaster.

The role of dopamine (DA) is investigated in cuticular hydrocarbon biosynthesis in Drosophila melanogaster with three different approaches: use of DA-deficient mutants (dopa decarboxylase temperature sensitive mutants reared at restrictive temperature, and rescued by dopamine ingestion or by pale mutants partially rescued by a tyrosine hydroxylase construction), pharmacological treatments (tyrosine hydroxylase inhibitors) and topical application on decapitated flies. We report that DA specifically regulates diene hydrocarbon biosynthesis, which is female specific. Our results suggest that DA acts in adult flies within the first hours of imaginal life and that DA production from the brain is crucial for this process. Thus, DA contributes to reproduction in D. melanogaster by acting during a critical period during development of young adults.

Relations between cuticular hydrocarbon (HC) polymorphism, resistance against desiccation and breeding temperature; a model for HC evolution in D. melanogaster and D. simulans.

D. simulans and D. melanogaster present two types of polymorphism in their cuticular hydrocarbon (HC) composition. Especially both sexes of D. simulans, and D. melanogaster males display 7-tricosene (7T) as the major compound type [7T]s and [7T]m, or 7-pentacosene (7P) [7P]s and [7P]m. D. melanogaster females display 7,11-heptacosadiene (7,11HD) as the major compound: [7,11HD]m, or 5,9-heptacosadiene (5,9HD): [5,9HD]m. The [7P]s, [7P]m and [5,9HD]m are mainly present in central Africa. A significant correlation was found between latitude and the proportion of compounds with 23 and 25 carbon atoms, especially 7T and 7P in both sexes of D. melanogaster. [7P]m type of D. melanogaster, characterized with an excess of C25 compounds, presents a higher resistance against desiccation than [7T]m type, where C23 compounds are more abundant. These differences can be correlated with calculated HC fusion temperatures. Moreover, increasing the breeding temperature from 18 to 29 degrees C induces in D. melanogaster males an increase in 25C compounds and a decrease in 23C compounds, but the opposite effect in D. simulans. A mathematical model of biosynthesis, based on kinetics of elongation and decarboxylation enzymes, suggests that a simple variation of the efficiency of an elongation enzyme may account for the differences observed between the [7T]m and [7P]m types of D. melanogaster and [7T]s and [7P]s types D. simulans. Finally on the basis of the geographical distribution of the HC types of both Drosophila species, an evolutionary dispersal pathway is proposed and discussed in relation to the environment and reproductive behavior.

Why there is a one-way crossability between D. melanogaster and D. simulans? An ontogenic explanation.

In the Drosophila melanogaster complex, females D. melanogaster mate relatively easily with males Drosophila simulans but the reciprocal cross is rare. The species sexual isolation is mainly based on chemical and acoustic signal exchanges between partners. The male side of this communication is investigated in this paper in order to understand the asymmetry. In D. melanogaster the acoustic signature is highly significant, and is synergistically reinforced by the chemical signal. In D. simulans the importance of the two signaling channels seems to be reversed. This could explain why D. simulans males produce less precise interpulse interval (IPI) mean value in the courtship song, which can readily overlap those of D. melanogaster. As the males of the two species use the same chemical key, D. simulans males could be recognized by D. melanogaster females as a conspecific.

ALTERED MATING BEHAVIOR IN A CARSONIAN POPULATION OF DROSOPHILA SECHELLIA.

Mating behavior was studied in two laboratory populations of Drosophila sechellia and their reciprocal hybrids. The ancestral population was reared on a special medium, optimal for this species, while the derived population was reared on a standard Drosophila food, and underwent a bottleneck while adapting to this new medium, in a manner similar to the "founder-flush" process of Carson (1971). A significant tendency towards mating asymmetry was found, with ancestral females mating significantly less frequently with derived males than derived females with ancestral males. Analysis of hybrids suggested an important role for the male's X chromosome or for a maternal effect. No significant differences were found among parental types for their main female cuticular hydrocarbons, the proportion of courtship spent in various behavioral elements, body weight, or wing length. Significant differences were found in the structure of courtship, male locomotor activity, male cuticular hydrocarbon levels, and male courtship song inter-pulse interval (i.p.i.). None of these differences showed an X-linked effect in the reciprocal hybrids. Hypotheses put forward to explain interspecific mating asymmetries are discussed in the light of these results.

VARIATIONS IN CUTICULAR HYDROCARBONS AMONG THE EIGHT SPECIES OF THE DROSOPHILA MELANOGASTER SUBGROUP.

In addition to protecting against desiccation, some of the hydrocarbons of the waxy cuticle have previously been shown to be mating pheromones in Drosophila melanogaster and D. simulans. Therefore, cuticular hydrocarbons were compared among the eight species in the D. melanogaster subgroup. For the two cosmopolitan species and several geographic strains that were studied, all males are quite similar with very abundant monoenes. The major compound in most cases is 7-tricosene. Only three exceptions were found: D. sechellia, and the Afrotropical strains of D. melanogaster and D. simulans. A significant sexual dimorphism exists in three species: D. melanogaster, D. erecta, and D. sechellia. Greater variation was observed in females than in males. D. erecta is singular in the production of long-chain molecules (31-33 carbons). Only three species (D. melanogaster, D. erecta, and D. sechellia) produce diene in significant amounts. Such products, especially 7,11-heptacosadiene, are known to act as aphrodisiacs for D. melanogaster males. In the five other species, females show only quantitative differences from males, generally with 7-tricosene as the most abundant compound. This compound is an aphrodisiac for D. simulans males. Some species such as D. yakuba, D. teissieri, D. orena, D. mauritiana, and the Seychelles strain of D. simulans are almost identical in the chemical composition of cuticular hydrocarbons. In contrast, important variations are observed between geographic populations of D. melanogaster and D. simulans.