果实蝇求偶行为与求偶歌

果实蝇Bactrocera是危害经济作物的一类重要农业害虫,研发有效的果实蝇防控技术是现代农业产业发展的迫切需求。雄性不育技术是防控果实蝇高效绿色的防控方法。深入研究果实蝇求偶行为是雄性不育技术顺利开展的前提,也能为开发新防控技术提供有效的生物学靶标。本文将以果蝇研究作为对照,对果实蝇求偶行为与求偶歌的研究相关进展进行总结。果实蝇求偶行为基本属于预交配求偶行为,但其求偶歌中是否具有物种特异性的特征参数还有待验证。这为下一步开展果实蝇求偶行为与求偶歌的研究工作提供了基本依据与方向性引导。     

Rapid evolution of courtship song pattern in Drosophila willistoni sibling species

Courtship behaviours may provide a more reliable means of identifying reproductively isolated taxa than traits such as morphology or many genetic markers. Here we describe the courtship songs of the Drosophila willistoni sibling species group, which consists of several species and subspecies. We find that song pattern is species-specific, despite significant differences among strains within species. D. paulistorum has the most variable song pattern, which reflects this species' traditional subdivision into semispecies. All the other species could be unambiguously identified by song. The major differences among these species was in the interpulse interval, as has been found in other studies of fly song. However, the interpulse intervals of the species studied here were often multimodal. This was partly due to the presence of multiple song types within the courtship repertoire, but it also reflected changes in interpulse interval within a song type by some males. Unusually, some species had distinctively patterned variation in interpulse interval. Song must have evolved rapidly within the species complex, probably due to sexual selection.     

The courtship song of African Drosophila melanogaster

For many years it was thought that Drosophila melanogaster was relatively panmictic, without differentiation in the Mate Recognition System. Recent studies have demonstrated that flies from Africa vary in pheromones and assortative mating. Strains from Zimbabwe show strong sexual isolation from others. We show that the interpulse interval (IPI) of courtship song, an important mating signal, is unusually short among African flies. Zimbabwean flies have the shortest IPI, but there is no correlation with assortative mating, suggesting little direct role in sexual isolation. Chromosome replacements show that the IPI difference is largely due to genes on chromosome III, with significant interactions involving other chromosomes. Several traits potentially influencing sexual isolation among the melanogaster group of Drosophila seem to be localized to this chromosome. A concentration of important genetic differences might mean that the interaction effects reflect secondary coadaptation of the genetic background to changes associated with chromosome III.     

To females of a noctuid moth,male courtship songs are nothing more than bat echolocation calls

It has been proposed that intraspecific ultrasonic communication observed in some moths evolved, through sexual selection, subsequent to the development of ears sensitive to echolocation calls of insectivorous bats. Given this scenario, the receiver bias model of signal evolution argues that acoustic communication in moths should have evolved through the exploitation of receivers'' sensory bias towards bat ultrasound. We tested this model using a noctuid moth Spodoptera litura, males of which were recently found to produce courtship ultrasound. We first investigated the mechanism of sound production in the male moth, and subsequently the role of the sound with reference to the female''s ability to discriminate male courtship songs from bat calls. We found that males have sex-specific tymbals for ultrasound emission, and that the broadcast of either male songs or simulated bat calls equally increased the acceptance of muted males by the female. It was concluded that females of this moth do not distinguish between male songs and bat calls, supporting the idea that acoustic communication in this moth evolved through a sensory exploitation process.     

Differences in male courtship song among the species of thequinaria group ofDrosophila

Courtship song is known to vary among several groups of closely related species ofDrosophila. The present study investigates differences in song among four members of thequinaria group of fungal-breedingDrosophila which are known to coexist in Britain and the near-continent. Results show that the species can be distinguished by several song parameters including interpulse interval.D. phalerata is very different from the other three species, while there is considerable overlap betweenD. kuntzei andD. limbata in several characters.D. transversa appears to be more similar to the latter species than it is toD. phalerata. The question of mate choice and the potential for reproductive character displacement are discussed.     

Condition-dependent traits and the capture of genetic variance in male advertisement song

The occurrence of additive genetic variance (VA) for male sexual traits remains a major problem in evolutionary biology. Directional selection normally imposed by female choice is expected to reduce VA greatly, yet recent surveys indicate that a substantial amount remains in many species. We addressed this problem, also known as the 'lek paradox', in Achroia grisella (Lepidoptera: Pyralidae), an acoustic moth in which males advertise to females with a pulsed ultrasonic song. Using a standard half-sib/full-sib breeding design, we generated F1 progeny from whom we determined VA and genetic covariance (COVA) among seven traits: three song characters, an overall index of song attractiveness, nightly singing period, adult lifespan, and body mass at adult eclosion. Because A. grisella neither feed nor drink as adults, the last trait, eclosion body mass, is considered a measure of 'condition'. We found significant levels of VA and narrow-sense heritabilities (h2) for all seven traits and significant genetic correlations (= COVAi,j / radical (VA i x VA j)) between most pairs of traits (i, j). Male attractiveness was positively correlated with body mass (condition), adult lifespan, and nightly singing period, which we interpret as an energy constraint preventing males in poor condition from singing attractively, from singing many hours per night, and from surviving an extended lifespan. The positive genetic correlation (r = 0.79) between condition and attractiveness, combined with significant levels of VA for both traits, indicates that much of the variation in male song can be explained by VA for condition. Finally, we discuss the morphological and physiological links between condition and song attractiveness, and the ultimate factors that may maintain VA for condition.     

Modification of the courtship song by visual stimuli in the grasshopper Gomphocerus rufus (Acrididae)

ABSTRACT. Males of Gomphocerus rufus L. perform a courtship song consisting of repetitive units, each of which is composed of three subunits (S1, S2, S3). S1 is characterized mainly by slow and fast head rolling; S2 and S3 are distinguished by different types of leg-stridulation. These movements and the associated sounds were recorded during presentation of visual stimuli, either linear displacement of a living female or optomotor stimuli generated by a striped drum. Females moved artificially through the binocular visual field of a courting male with a velocity of 1 cm/s or more are mounted by the male from any subunit S1, S2 or S3, although under natural conditions mounting occurs only from S2. Thus above a critical velocity the courtship programme can be modified. Rotation of a striped drum about the yaw axis of the male during the slow S1 induces asymmetrical leg position, following movements of the head, and prolongation of S1. During S2 the male is especially sensitive to optomotor stimuli and responds with marked changes in body position. In S3 the intensity of the song is reduced, and its duration shortened. Fast drum movements interrupt the courtship programme. Rotation of the drum about the roll axis elicits optomotor head turning that interferes with the head rolling of S1. The fast phase of S1 and the frequency of head-rolling during S1 cannot be modified by optomotor stimulation. The results can be interpreted by assuming certain interactions between three central nervous elements: a calling-song generator, a head-rolling generator, and an optomotor centre.     

Acoustic response to a pheromonal cue in the arctiid moth Pyrrharctia Isabella

ABSTRACT. The acoustic properties of the clicks emitted in response to male courtship pheromone by female Pyrrharctia Isabella (J. E. Smith) (Lepidoptera: Arctiidae) have been investigated. Extracts of male scent organs or synthetic male pheromones can be applied to a glass rod and used to stimulate females to produce these sounds. Power spectra, sound pressure readings, and oscillographic analyses show that the acoustic signals elicited by male extracts or synthetic male pheromones are not distinguishable from those produced in response to disturbance (handling). This is the first reported example of a sound produced by a female moth in a sexual context, and is the first reported example in moths of an acoustic response to a pheromonal stimulus.     

Male courtship song and female preference variation between phylogeographically distinct populations of Drosophila montana

Understanding the variation within and between populations in important male mating traits and female preferences is crucial to theories concerning the origin of sexual isolation by coevolution or other processes. There have been surprisingly few studies on the extent of variation and covariation within and between populations, especially where the evolutionary relationships between populations are understood. Here we examine variation in female preferences and a sexually selected male song trait, the carrier frequency of the song, within and between populations from different phylogeographic clusters of Drosophila montana. Song is obligatory for successful mating in this species, and both playback and field studies implicate song carrier frequency as the most important parameter in male song. Carrier frequency varied among three recently collected populations from Oulanka (Finland), Vancouver (Canada), and Colorado (central United States), which represent the main phylogeographic groups in D. montana. Males from Colorado had the most distinct song frequency, which did not follow patterns of genetic differentiation. There was considerable variation in preference functions within, and some variation between, populations. Surprisingly, females from three lines from Colorado seem to have preferences disfavoring the extreme male trait found in this population. We discuss sources of selection on male song and female preference.     

The best of both worlds: cicada males change costly signals to achieve mates while females choose a mate based on both calling and courtship songs

Cicadas usually sing and mate in the higher parts of trees. Studies addressing the effects of different acoustic signals on mate choice in Cicadidae are very limited. We investigated the effects of both acoustical features and morphological traits on mate choice in an East Asian cicada Platypleura kaempferi. Males produce high-rate calling songs that attract females, then produce low-rate courtship songs to secure mating when a female is attracted. Higher calling song rate (CR), shorter single-pulse duration, and shorter pulse period of the calling song, together with lower courtship song rate and longer echeme period of the courtship song, are the most desirable traits used by females to choose a mate. These traits indicate that the more a male can raise the rate of song production, the higher the probability he is sexually selected by the female. No correlation was found between morphological traits and mating success. After mating, a minority of males started emitting calling songs again, but the CR was significantly lower than before mating and none of them attracted a new mate later. This promotes females mating with unmated males. We hypothesize that P. kaempferi may have the best of both worlds due to the unique song modulation and the mechanism of female mate choice: males change energetically, costly acoustic signals to achieve mates, while females choose a mate based on males’ acoustic properties. Our results contribute to better understanding the diversity of mating preference and enrich the mechanism of mate choice in acoustic insects.     

Acoustic courtship communication inSyntomeida epilais Wlk. (Lepidoptera: Arctiidae,Ctenuchinae)

The polka-dot wasp moth,Syntomedia epilais Wlk. (Lepidoptera, Arctiidae, Ctenuchinae), engages in extensive acoustic signaling during courtship. The signals, which are produced by both sexes, consist of sexually dimorphic trains of ultrasonic clicks. Field and laboratory behavioral experiments demonstrated that (1) sound production from both the male and the female is required for mating success, (2) femaleS. epilais show a higher level of acoustic responsiveness to male acoustic signals than to female signals, and (3) males use the female acoustic signals to locate their potential mates. It is suggested that the existence of the system is made possible by a reduction in bat predation pressure, perhaps the result of extreme distastefulness combined with the distinctive acoustic image that flying ctenuchines present to foraging bats.     

Ultrasonic signals in the defense and courtship ofEuchaetes egle Drury andE. bolteri Stretch (Lepidoptera: Arctiidae)

Euchaetes egle Drury andE. bolteri Stretch produce ultrasound using paired thoracic tymbal organs in both defensive and sexual contexts. The defensive ultrasound produced in response to tactile stimulation is fully characterized. The sounds are sexually monomorphic and species specific in the number of sound pulses produced during each flexion and relaxation of the tymbal, peak frequency, peak intensity, and duration of the interval between flexion and relaxation. Ultrasonic signals play a role in the courtship of both species. Males produce ultrasound just prior to contact with females, and it is shown to be important to courtship success inE. egle. Ultrasonic courtship communication is mapped on a recently proposed cladogram for the family Arctiidae. The use of ultrasound in courtship has evolved on at least three occasions within the family.     

Two distinct genomic regions, harbouring the period and fruitless genes, affect male courtship song in Drosophila montana

Acoustic signals often have a significant role in pair formation and in species recognition. Determining the genetic basis of signal divergence will help to understand signal evolution by sexual selection and its role in the speciation process. An earlier study investigated quantitative trait locus for male courtship song carrier frequency (FRE) in Drosophila montana using microsatellite markers. We refined this study by adding to the linkage map markers for 10 candidate genes known to affect song production in Drosophila melanogaster. We also extended the analyses to additional song characters (pulse train length (PTL), pulse number (PN), interpulse interval, pulse length (PL) and cycle number (CN)). Our results indicate that loci in two different regions of the genome control distinct features of the courtship song. Pulse train traits (PTL and PN) mapped to the X chromosome, showing significant linkage with the period gene. In contrast, characters related to song pulse properties (PL, CN and carrier FRE) mapped to the region of chromosome 2 near the candidate gene fruitless, identifying these genes as suitable loci for further investigations. In previous studies, the pulse train traits have been found to vary substantially between Drosophila species, and so are potential species recognition signals, while the pulse traits may be more important in intra-specific mate choice.     

Context-sensitive dance–vocal displays affect song patterns and partner responses in a socially monogamous songbird

Multimodal signaling contributes to efficient communication by improving signal efficacy and increasing signal information. Songbirds often combine dance displays with songs according to the socio-sexual context; therefore, the song is assumed to function differently depending on dance displays. In this study, we tested how dance displays affect song patterns and the responses of paired partners using male and female blue-capped cordon-bleus (Uraeginthus cyanocephalus). Blue-capped cordon-bleus are a socially monogamous estrildid finch, and both males and females perform songs and distinct “tap dance”-like displays. Songs with dance displays were longer and more stereotyped than songs without dance displays in both males and females. Furthermore, both male and female paired partners showed more gestural responses to songs with dance displays than those without dance displays. Songs without dance displays were performed under both isolated and paired conditions, whereas songs with dance displays were only performed when the focal bird was housed with a paired partner. These results suggest that songs had different functions depending on dance displays and social contexts. The multimodal display of blue-capped cordon-bleus seems to draw the attention of paired partners to the physical abilities of the performer.     

Genetics of incipient speciation in Drosophila mojavensis. I. Male courtship song, mating success, and genotype x environment interactions

Few studies have examined genotype by environment (GxE) effects on premating reproductive isolation and associated behaviors, even though such effects may be common when speciation is driven by adaptation to different environments. In this study, mating success and courtship song differences among diverging populations of Drosophila mojavensis were investigated in a two-environment quantitative trait locus (QTL) analysis. Baja California and mainland Mexico populations of D. mojavensis feed and breed on different host cacti, so these host plants were used to culture F2 males to examine host-specific QTL effects and GxE interactions influencing mating success and courtship songs. Linear selection gradient analysis showed that mainland females mated with males that produced songs with significantly shorter L(long)-IPIs, burst durations, and interburst intervals. Twenty-one microsatellite loci distributed across all five major chromosomes were used to localize effects of mating success, time to copulation, and courtship song components. Male courtship success was influenced by a single detected QTL, the main effect of cactus, and four GxE interactions, whereas time to copulation was influenced by three different QTLs on the fourth chromosome. Multiple-locus restricted maximum likelihood (REML) analysis of courtship song revealed consistent effects linked with the same fourth chromosome markers that influenced time to copulation, a number of GxE interactions, and few possible cases of epistasis. GxE interactions for mate choice and song can maintain genetic variation in populations, but alter outcomes of sexual selection and isolation, so signal evolution and reproductive isolation may be slowed in diverging populations. Understanding the genetics of incipient speciation in D. mojavensis clearly depends on cactus-specific expression of traits associated with courtship behavior and sexual isolation.