Yuzuru Oguma

A locus for female discrimination behavior causing sexual isolation in Drosophila

The genetic basis of sexual isolation that contributes to speciation is one of the unsolved questions in evolutionary biology. Drosophila ananassae and Drosophila pallidosa are closely related, and postmating isolation has not developed between them. However, females of both species discriminate their mating partners, and this discrimination contributes to strong sexual isolation between them. By using surgical treatments, we demonstrate that male courtship songs play a dominant role in female mate discrimination. The absence of the song of D. pallidosa dramatically increased interspecies mating with D. ananassae females but reduced intraspecies mating with D. pallidosa females. Furthermore, genetic analysis and chromosomal introgression by repeated backcrosses to D. pallidosa males identified possible loci that control female discrimination in each species. These loci were mapped on distinct positions near the Delta locus on the middle of the left arm of the second chromosome. Because the mate discrimination we studied is well developed and is the only known mechanism that prevents gene flow between them, these loci may have played crucial roles in the evolution of reproductive isolation, and therefore, in the speciation process between these two species.

Heterospecific inter-pulse intervals of courtship song elicit female rejection in Drosophila biauraria

Abstract The present study investigated the significance of song in sexual isolation in Drosophila biauraria . Interspecific copulation was not observed with a heterospecific D. triauraria male that had intact wings, although it was often observed when the males wings were removed, indicating that heterospecific courtship song elicits female rejection, and prevents copulation. Artificially synthesized songs were played to a female with a wingless, antennaless male. The conspecific inter-pulse interval of artificial song increased mating frequency, whereas the heterospecific, longer or shorter, inter-pulse interval significantly decreased the frequency. The mating frequency with random noise and silence showed intermediate values. Female rejection behaviour was greater with heterospecific than with conspecific inter-pulse intervals of artificial song; with random song and silence female responses were intermediate. This indicates that the inter-pulse interval is used as a species discriminator in D. biauraria females. It is suggested that the females ability to discriminate inter-pulse intervals and her responses, rejection and acceptance, have evolved under the influence of reinforcement, reproductive character displacement, or both, which triggered sexual selection.

(Z,Z)-5,27-Tritriacontadiene: Major sex pheromone ofDrosophila pallidosa (Diptera; Drosophilidae)

A crude cuticular extract from both sexes of 3660 fruit flies (Drosophila pallidosa) was subjected to SiO2 and AgNO3/SiO2 column chromatography, accompanied by bioassay for the sex pheromone activity. After three chromatographic steps, the active fraction was obtained. The main component of the active fraction was determined to be (Z,Z)-5,27-tritriacontadiene [(Z,Z)-5,27-C33:2, on the basis of gas-liquid chromatographic analysis, chemical derivatization and gas chromatography-mass spectrometry. Synthetic (Z,Z)-5,27-C33:2 at 5 female equivalents (FE) elicited a clear courtship response with a high courtship index amongD. pallidosa males. Therefore it was concluded that (Z,Z)-5,27-C33:2 was a major sex pheromone component in this species.

COURTSHIP SONG RECOGNITION IN THE DROSOPHILA MELANOGASTER COMPLEX: HETEROSPECIFIC SONGS MAKE FEMALES RECEPTIVE IN D. MELANOGASTER, BUT NOT IN D. SECHELLIA

Abstract. The courtship song emitted by male wing vibration has been regarded as one of the most important signals in sexual isolation in the species of the Drosophila melanogaster complex. Inter‐ and intraspecific crosses were observed using males whose wings were removed (mute) or females whose aristae were removed (deaf). Females of D. melanogaster, D. simulans, and D. mauritiana mated with heterospecific males in the song‐present condition (cross between normal females and winged males) more often than in the no‐song condition (cross between normal females and wingless males or between aristaless females and winged males) or they showed no preference between the two conditions. It is possible that in these females heterospecific courtship songs play a role as if they were conspecific. In contrast, the females of D. sechellia mated with D. melanogaster or D. simulans males in the no‐song condition more often than in the song‐present condition, suggesting that they reject males with heterospecific song. Female mate recognition depending on the courtship song in D. melanogaster, D. simulans, and D. mauritiana is considered to be relatively broader and that in D. sechellia narrower.

(Z)-11-Pentacosene is the major sex pheromone component inDrosophila virilis (Diptera)

SummaryA crude cuticular extract from 3450 virgin 9–13 day old female fruit flies(Drosophila virilis), was subjected to chromatography accompanied by bioassay for sex pheromone activity. After three chromatographic steps, fractions containing active monoenes and dienes were obtained. Chemical analysis by infrared absorption, gas liquid chromatography and gas chromatography/mass spectrometry of the active fraction indicated that active monoenes were comprised chiefly of (Z)-11-pentacosene (abbreviated (Z)-11-C25:1), (Z)-13-C27:1’, (Z)-13- and (Z)-14-C29:1. Synthetic monoenes were made, and only (Z)-11-C25:1 elicited good courtship behaviour in maleD. virilis. Therefore it was concluded that (Z)-11-C25:1 was a major sex pheromone. A total of 16.2±1.32 µg of cuticular hydrocarbons was isolated from 10 day old females, including 5.9±0.56 µg of (Z)-11-C25:1. An additive effect was suggested from the higher observed courtship response when using a mixture of active dienes with the active monoene.

Genetic basis and evolution of species-specific courtship song in the Drosophila auraria complex.

The interpulse interval (IPI) of courtship song in the Drosophila auraria complex is the only parameter that is consistently species-specific among the several courtship elements examined within the complex. The genetic basis of the species-specific courtship song was examined by analysing the song of interspecific hybrids and of backcross progeny. IPI of all interspecific hybrids except two showed intermediate values, suggesting autosomal control of species-specific IPI. However, significant deviation for shorter IPI from midparent was found in thirteen out of 20 crosses. The chromosomal analysis between D. auraria and D. biauraria revealed that the two major autosomes had significantly large effects on IPI, but the sex chromosome and cytoplasm had no effect. Since no interaction was detected, it is concluded that each autosome acts additively in the determination of species-specific IPI. The common ancestors of the D. auraria complex may also have had autosomal control of IPI, which has been conserved during speciation in the complex.

Behavioral Response of Males to Major Sex Pheromone Component, (Z,Z)-5,25-Hentriacontadiene, of Drosophila ananassae Females

To identify sex pheromone components in Drosophila ananassae, the cuticular hydrocarbons were analyzed and bioassayed. After silicic acid and two silver nitrate–silicic acid chromatographic steps, active fractions containing (Z,Z)-5,25- and (Z,Z)-4,26-hentriacontadiene as the most abundant components were obtained. Both hydrocarbons could elicit male courtship behavior. The former was the more predominant compound and elicited a higher courtship index value than the latter. Cuticular hydrocarbon differences between D. ananassae and its close relative, D. pallidosa, and their roles in species recognition are discussed.

Courtship behavior and sexual isolation between Drosophila auraria and D. triauraria in darkness and light

Sexual isolation between two species of the Drosophila auraria complex, D. auraria and D. triauraria is different in darkness and light. In darkness there is complete isolation while in light it is only partial. The sensory bases of these differences were investigated by behavioral studies in darkness and light. In darkness there was no normal courtship sequence but males of both species displayed attempted copulation to homospecific and heterospecific females, as well as homospecific males. After a few hours together, homospecific copulations were observed after females displayed a characteristic posture: spreading the wings. Such an acceptance posture, also observed in other Drosophila species, was probably released by the wing vibration of a homospecific male. In light, visual signals alone are able to elicit males orientation and following, whether flies are housed together or physically separated. Intense homosexual courtships were observed. All these data suggest that males have a low discrimination ability and females play the main role in sexual isolation.

Genetic analysis of Drosophila virilis sex pheromone: genetic mapping of the locus producing Z-(11)-pentacosene.

Z-(11)-pentacosene, Drosophila virilis sex pheromone, is predominant among the female cuticular hydrocarbons and can elicit male courtship behaviours. To evaluate the genetic basis of its production, interspecific crosses between D. novamexicana and genetically marked D. virilis were made and hydrocarbon profiles of their backcross progeny were analysed. The production of Z-(11)-pentacosene was autosomally controlled and was recessive. Of the six D. virilis chromosomes only the second and the third chromosomes showed significant contributions to sex pheromone production, and acted additively. Analysis of recombinant females indicated that the locus on the second chromosome mapped to the proximity of position 2-218.

Negative Correlations Between Resistance to Three Organophosphate Insecticides and Productivity Within a Natural Population of Drosophila melanogaster (Diptera: Drosophilidae)

Abstract To investigate the relationship between resistance to organophosphate insecticides and fitness components, we first measured resistance to three organophosphates, malathion, prothiophos, and fenitrothion, and productivity, a measure of fitness components, for each of the isofemale lines from the same natural population of Drosophila melanogaster (Meigen). Pearson correlation coefficients indicated that positive correlations among resistance to the organophosphates and negative correlations between resistance to each of the organophosphates and the productivity existed within the natural population. We further investigated the genetic basis of the correlations among resistance to the organophosphates and the productivity, by using chromosome-substituted lines between a resistant and a susceptible inbred line established from the same natural population. Chromosomal analyses indicated that the third chromosome from the resistant line exhibited not only significant, positive effects on resistance to all of the organophosphates tested but also a significant negative effect on the productivity, suggesting positive genetic correlations between resistance to each organophosphate and negative genetic correlations between resistance to each organophosphate and the productivity. In addition, a significant negative effect on the productivity was also detected from the second chromosome, which did not exhibit significant major effects on resistance to the organophosphates. This suggests that fitness components of resistant lines could be also affected by factors independent of insecticide resistance. The dynamics of genetic variation in resistance to the organophosphates within the natural population of D. melanogaster are discussed from the standpoint of negative genetic correlations between resistance to the organophosphates and the productivity.