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Search for species-specific mating signal in courtship songs of sympatric sibling species, Drosophila ananassae and D. pallidosa.

Sexual isolation is one of the most important mechanisms that may lead to speciation. Drosophila ananassae and D. pallidosa are useful for the study of sexual isolation, because of their sympatric distribution and no postmating isolation between them. Courtship songs are considered to play a crucial role in sexual isolation between D. ananassae and D. pallidosa. We recorded and analyzed male courtship songs of D. ananassae and D. pallidosa for eight and four geographical strains, respectively. Courtship behaviors of the two species were consistent with those previously described, however, male's middle leg shaking, which had not before been described, was observed in both species. Males sing by wing vibration only during courtship. Their song oscillograms were distinct between species, but those of conspecific strains were very similar, in spite of their different geographical derivation. We found species-specificity in burst length, pulse length, cycle number in a pulse, and frequency spectra of bursts; these results suggest that these song parameters may play a role in mate recognition that enforces their sexual isolation. The specific values of interpulse interval, cycle number in a pulse and intrapulse frequency were involved with the determination of specificity in frequency spectra of bursts. We discussed the possibility that the specific frequency spectra of bursts are recognized by females as the species-specific signal rather than each parameter individually.

Animals↗

Chemically reinforced conditioned courtship in Drosophila: responses of wild-type and the dunce, amnesiac and don giovanni mutants.

To test the hypothesis that associative learning is the basis for conditioned courtship, male Drosophila melanogaster were paired with virgin females in the presence of quinine, known to be a negative reinforcer in a learning paradigm independent of courtship. Such "experienced" wild-type males failed to court virgin females and remained refractory to them for 1-2 h. But experienced males from the learning-defective strain, dunce, continued to court females at high levels; and experienced males from the retention-defective strain, amnesiac, failed to demonstrate the wild-type refractory period. Finally, males from the don giovanni strain, defective with respect to fertilized-female conditioning, were conditioned by presentation of virgin females and quinine. Courtship-depressing effects of cis-vaccenyl alcohol, which can be recovered from fertilized females, were confirmed, but no evidence for its role as a negative reinforcer of male courtship was obtained.

Animals↗

Effects of the apterous4 mutation on Drosophila melanogaster males' courtship.

The apterous4 (ap4) mutation has pleiotropic effects on the morphology and lifespan of D. melanogaster adults. The mutation also affects one aspect of male sexual behaviour. Unlike normal, sexually mature males, three day old ap4/ap4 males synthesize courtship-stimulating hydrocarbons that immature males make; as a result they elicit vigorous courtship (Jallon and Hotta, 1979; Jallon et al., 1986). Data presented in this report indicate that expression of the apterous4 mutation does not affect all aspects of sexual maturation in males. Specifically, 24-30 hour old ap4/ap4 males are capable of performing all of the courtship behaviors, including abdominal vibration, a newly described response of mutant and wild-type males to virgin females. However, the mutant males spend less time courting and are less likely to perform some of the courtship behaviours than age-matched controls. These abnormalities are probably indirect effects of the apterous4 allele's previously unreported effects on the flies' motor functions, which include partial paralysis, lack of coordination, and sluggishness.

Animals↗

CaM kinase II and visual input modulate memory formation in the neuronal circuit controlling courtship conditioning.

In Drosophila, calcium/calmodulin-dependent protein kinase II (CaM kinase) has been shown to be important in the expression of both learning and memory for the associative behavior courtship conditioning. In this study we examine the role of visual input in producing this behavior and the effects of modifying visual input on CaM kinase-dependent memory formation. Inhibition of CaM kinase blocked apparent learning regardless of visual input. Visual input selectively affected the memory phase of courtship conditioning: normal visual input masked the memory effects of inhibition of CaM kinase resulting in generation of memory without apparent learning, whereas disruption of visual input revealed the CaM kinase-dependence of memory. Visual input was found to be important only during the training period, which implies that vision and CaM kinase are interacting in the formation rather than the retrieval of memory. Our results suggest a model for courtship conditioning in which multiple sensory inputs are integrated at a CaM-kinase-dependent neuronal switch to modulate courtship behavior.

Animals↗

Pinealectomy, melatonin, and courtship behavior in male red-sided garter snakes (Thamnophis sirtalis parietalis).

Activation of courtship behavior in male red-sided garter snakes is independent of androgens. Only exposure to extended periods of low temperature with subsequent warming stimulates courtship in males. The pineal gland is thought to transduce temperature as well as photoperiodic information in reptiles. Therefore, we explored the relationship of the pineal and melatonin to sexual behavior in this species. Pinealectomy of male garter snakes disrupted sexual behavior upon emergence from a 17-week period of low temperature in approximately 60% of treated individuals in each of the 3 years of study. However, 40% of the males were unaffected by the pinealectomy, engaging in vigorous courtship. Administration of exogenous, chronic melatonin did not significantly modulate the effect of pinealectomy. Upon pinealectomy in the autumn (before hibernation), plasma levels of melatonin fell. However, upon emergence from hibernation, melatonin levels in pinealectomized (PINX) and sham-treated (SHAM) animals were equivalent, indicating extrapineal source(s) of melatonin. However, PINX males did not exhibit a diel cycle in melatonin levels upon emergence. Instead, melatonin remained elevated through the subsequent 24-hr period. SHAMs did exhibit a diel cycle. Ten days after emergence, PINX animals either had a disrupted/abnormal melatonin cycle and were non-courters or had a cycle similar to SHAM males and courted. Therefore, a normal diel cycle of melatonin appeared necessary for the proper expression of courtship behavior. These results suggest that the pineal in snakes 1) is part of a complex, multi-oscillator system as it is in birds and lizards and 2) may play a role in maintaining polymorphism in timing of reproductive behavior.

Acclimatization↗

Variance in male mating success and female choice for persuasive courtship displays.

An important function of the courtship displays of male animals is to stimulate, or 'persuade', females to mate. While many studies of courtship success have shown that females choose stimulating males as partners, fewer have demonstrated that the most persuasive males in a population also enjoy the highest mating success (i.e. mate with the greatest numbers of partners). Such a relationship provides a stronger test of the hypothesis that male displays evolve by sexual selection via female choice. We tested this hypothesis in the laboratory using the North American plethodontid salamander, Desmognathus ocoee, in which females only elicit spermatophore deposition if rendered sexually responsive by male courtship. First, we determined variance among individual males in the numbers of females with which they mated across 35 encounters, in the absence of differences in partner encounter rates and direct intermale interactions. Second, we demonstrated that males with high 'historical' scores of mating success performed significantly higher frequencies of persuasive courtship displays, which provide tactile, chemical and visual stimulation to females. We conclude that sexual selection via female choice favours persuasive displays because they confer high mating success on the males that perform them at the highest frequencies. Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

Courtship songs of Drosophila pseudoobscura and D. persimilis: analysis of variation.

Differences in Drosophila courtship song elements are thought to confer species sexual isolation because of the low levels of variation within species, the large differences often seen between closely related species, and the results of experiments using synthetic songs in two species groups. In this study, we reanalyse the courtship song elements of D. pseudoobscura and D. persimilis. We confirmed the differences between these species in high-rate repetition song interpulse interval (IPI) and intrapulse frequency and in the amount of low-rate repetition song produced. We found that both X-chromosome and autosomal loci contribute to the differences between D. pseudoobscura and D. persimilis in their IPI and intrapulse frequency. No association was detected between IPI and hybrid male mating success to females of either species, although frequency was associated with mating success to D. persimilis females. Additionally, we found evidence of IPI periodicity in the song of D. persimilis, demonstrated differences in mean IPI between D. pseudoobscura populations, and showed a significant influence of temperature on IPI and intrapulse frequency. Such environmental effects may have obscured tests for population differentiation in courtship song parameters in other species, and we suggest the possibility that differences between Drosophila species in mean courtship song IPI might only rarely affect their sexual isolation. Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

Habituation and development of response specificity to a sign stimulus: male preference for female courtship posture in stickleback.

I examined biased patterns of habituation that occur when reproductive male three-spined stickleback, Gasterosteus aculeatus, respond to a choice of two dummy females, one displaying the head-up posture of a sexually receptive female and an otherwise identical dummy lacking this sign stimulus (i.e. a dummy in a 'neutral' horizontal posture). Males initially courted both dummies about equally, but within 5 min, their courtship to the horizontal dummy began to habituate and was eventually replaced by attack behaviour, which increased about four-fold during the 1-h trial. Courtship to the head-up dummy continued unabated throughout the trial and by 5 min exceeded courtship to the horizontal dummy. Attack to the head-up dummy increased slightly in the first half of the trial then leveled off to about one-third that directed to the horizontal dummy. Both dummies were stationary and unreactive so could not directly provide differential feedback to the males. It is speculated that the self-arousal males obtain when responding to head-up posture reinforces their response to that cue. The salience of this sign stimulus may develop through an interactive process wherein the male's predisposition to recognize and court the head-up posture and the sexual self-arousal he obtains by doing so counter habituation and help maintain his sexual response to that cue. The stimulus-response specificity of male courtship is assumed to have an innate basis but requires experience to become fully differentiated. The results also illustrate that rate or extent of habituation to different stimuli, even on the same dimension, can differ within the same individual and in an adaptive manner. Copyright 2000 The Association for the Study of Animal Behaviour.

Journal Article↗

Vasotocin stimulates appetitive responses to the visual and pheromonal stimuli used by male roughskin newts during courtship.

It is now well established that vasotocin (AVT) and its mammalian homologue vasopressin influence various social behaviors in vertebrates, but less is known about the mechanisms through which these peptides modulate behavior. In male roughskin newts, Taricha granulosa, AVT stimulates a courtship behavior, amplectic clasping. Three general explanations for how AVT affects male courtship behavior have been considered: by enhancing a central state of sexual motivation, by affecting sensorimotor integration mechanisms in individual sensory modalities, or by influencing a nonspecific state of attention, arousal, or anxiety. AVT administration enhanced appetitive responses to visual and olfactory sexual stimuli, as would be expected if AVT affects a state of sexual motivation that affects behavioral responses to sexual stimuli regardless of the sensory modality in which they are processed. However, AVT selectively enhanced responses to female olfactory stimuli (sex pheromones), but similarly enhanced responses to female and food-related visual stimuli (worms), thus questioning the utility of such a motivational mechanism, as responses to female stimuli were not selectively enhanced in all sensory modalities. We therefore propose that exogenous AVT independently influences olfactory processes associated with orientation/attraction toward a female sex pheromone and visual processes associated with orientation/attraction toward a visual feature common to females and worms. In further experiments AVT administration failed to stimulate feeding behavior but did decrease locomotor activity. Thus, AVT does not stimulate courtship behavior in this species by enhancing the animals' general state of attention or by decreasing general anxiety, as responses to nonsexual, attractive stimuli were not uniformly enhanced, nor by stimulating general arousal, as activity levels did not increase. Rather, the data support the conclusion that AVT affects courtship by influencing specific sensorimotor processes associated with behavioral responses to individual releasing stimuli, which suggests a mechanistic framework for understanding socially motivated behavior is this species.

Animals↗

Manipulations of the AVT system shift social status and related courtship and aggressive behavior in the bluehead wrasse.

Arginine vasotocin (AVT) and its mammalian homologoue arginine vasopressin (AVP) influence male sexual and aggressive behaviors in many species. We tested the effects of AVT and an AVP-V(1a) receptor antagonist on the display of alternative male tactics in a tropical coral reef fish, the bluehead wrasse Thalassoma bifasciatum. We gave AVT injections to territorial and nonterritorial males of the large and colorful phenotype (terminal phase) and an AVP-V(1a) receptor antagonist, Manning compound, to territorial males in the field. AVT increased courtship independent of status, while its effects on territoriality and aggression were dependent upon male status. In territorial males, AVT increased courtship and tended to decrease the number of chases toward initial phase individuals. In nonterritorial males, AVT increased courtship, chases toward initial phase individuals, and territorial behavior while decreasing feeding. These are all behaviors rarely seen in nonterritorial males, so AVT made these males act like territorial TP males. The AVP-V(1a) receptor antagonist had opposite effects. It decreased courtship and territorial defense, making these males act more like nonterritorial males. Manipulations of the AVT system shifted males within a single phenotype from the nonterritorial social status to the territorial social status and vice versa. Since the entire suite of behaviors related to territoriality was affected by AVT system manipulations, our results suggest that the AVT system may play a key role in motivation of behaviors related to mating.

Aggression↗

Courtship behavior of Drosophila genetically or surgically deprived of basiconic sensilla.

The lack of basiconic antennal sensilla in the mutant lozenge was used to assess the role of these olfactory receptors in the courtship behavior of Drosophila melanogaster. Under normal light conditions, lozenge males courted virgin females much less than wild-type males did. However, when visual courtship stimuli were eliminated by studying behavior under dim red light, the two kinds of males courted individual wild-type virgin females with the same intensity, and the latency to copulation was similar. Also, no difference in courtship vigor was observed if the two kinds of males were paired in red light with a mated female. These data suggest that antennal basiconic sensilla are important for neither the perception of the attraction pheromone(s) of virgin females nor the inhibitory pheromone(s) of mated females. Similar assays with males deprived of maxillary palps make it unlikely that the basiconic-like sensilla on these appendages are needed to perceive the attraction pheromones. However, the unexpectedly high courtship activity of palp-deprived males toward mated females suggests that basiconic-like maxillary sensilla may be receptors of inhibitory female compounds.

Animals↗

Identification of stimuli that mediate experience-dependent modification of homosexual courtship in Drosophila melanogaster.

A naive, sexually mature D. melanogaster male tested with a young, sexually immature male will perform vigorous courtship, but the mature male will perform much less courtship if he is subsequently tested with a second young male. This phenomenon is called experience-dependent courtship modification (EDCM). We have shown that exposure to either or both of the two courtship-stimulating pheromones that immature males synthesize is sufficient to induce EDCM.

Animals↗

The role of male courtship song in species recognition in Drosophila montana.

In many Drosophila species the male courtship song is an essential part of a successful courtship, playing a role in species-recognition and/or in sexual selection exercised by the females on conspecific males. In Drosophila montana, specific traits of the courtship song (the length and the carrier frequency of sound pulses) have earlier been shown to play an important role in intraspecific mate choice. Here we show that the male song (especially the interpulse interval) also plays a role in species-recognition, maintaining sexual isolation between sympatric species. We succeeded in breaking down sexual isolation between D. montana females and D. lummei males by playing the females simulated courtship song with a species-specific interpulse interval while they were courted by a mute (wingless) male. Involvement of different song traits in intra- and interspecific mate choice suggests that the song may be affected by both directional and stabilizing selection.

Animals↗

Characterization of a female-produced courtship pheromone in the parasitoid Nasonia vitripennis.

Males of the parasitoid Nasonia vitripennis (Walker) (Hymenoptera: Pteromalidae) show a characteristic courtship behavior. We demonstrate that male arrestment and key behavioral elements of the courtship sequence are mediated by a female-derived contact sex pheromone. Males were arrested on paper disks treated with female extracts but not on those treated with male extracts. Male responsiveness was influenced by the surface to which female extracts were applied. Extracts applied to an extracted beetle elytron arrested males more strongly than those applied to filter paper of comparable size. However, more complex behavioral elements, such as head nodding and copulation attempts, were shown only when extracts were applied to extracted male cadavers, suggesting that tactile or visual cues synergize the male response. The chemicals involved are stable, of low volatility, and nonpolar. Dead females arrested males and elicited courtship behavior for at least 8 d. Males showed no sign of attraction to live females at a distance of 3 cm in an olfactometer. Fractionation of female extracts demonstrated that the activity was exclusively located in the nonpolar fraction. Analysis of the active fraction by gas chromatography-mass spectrometry revealed that cuticular hydrocarbons with chain lengths between 25 and 37 carbon units were present. Comparison of hydrocarbon profiles from males and females showed qualitative and quantitative differences. These results suggest that sex-specific cuticular hydrocarbons are the key signals mediating the male courtship behavior in N. vitripennis.

Animal Communication↗

The role of the midbrain in courtship behavior of the female ring dove (Streptopelia risoria): evidence from radiofrequency lesion and hormone implant studies.

The involvement of the nucleus intercollicularis (ICo) region of the midbrain, an estrogen-sensitive area, in the expression of estrogen-dependent female courtship behavior was examined with radiofrequency lesion and intracranial hormone implant techniques. Bilateral lesions in the ICo region caused a reduction in the female's nest coos, and no reduction in other behaviors in response to male courtship. In addition, the follicles of the ICo lesioned females failed to grow in response to male courtship. A second experiment showed that the difference in follicular size between ICo-lesioned and sham-ICo-lesioned females was not observed if females were visually isolated from males, suggesting that tonic ovarian activity was not affected by ICo lesions. In the third experiment, unilateral 30-gauge implant containing estrogen (E) or estradiol benzoate (EB) in the ICo region of bilaterally ovariectomized females restored the nest-coo, though not to the level obtained with systemic EB injections. Other behaviors were not selectively affected by E or EB implants in the ICo region. Diffusion from the implant was probably not responsible for the elicitation of the nest-coo, since the oviduct weights of females implanted with E or EB and cholesterol did not differ. These data are consistent with the hypothesis that the ICo region is involved in the expression of vocal (cooing) courtship behavior in the female ring dove.

Animals↗

Stress-induced inhibition of sexual behavior: corticosterone inhibits courtship behaviors of a male amphibian (Taricha granulosa).

When male rough-skinned newts (Taricha granulosa) are exposed to presumptive stressors, the incidence of courtship decreases and plasma corticosterone concentration increases. When sexually active males are injected intraperitoneally with corticosterone (1, 5, 10, 15, 20, or 25 micrograms), the incidence of courtship decreases rapidly and in proportion to the dose of corticosterone. Intracerebroventricular infusion of synthetic corticotropin-releasing factor (CRF) elevates plasma corticosterone levels and suppresses courtship. When male newts receive an injection of metyrapone, a drug that interferes with corticosterone synthesis, the inhibitory effects of stress or CRF infusion on courtship are reduced. These results support the hypothesis that, in this amphibian, elevated levels of corticosterone associated with exposure to stressful stimuli inhibit sexual behaviors.

Adrenocorticotropic Hormone↗

Plasma testosterone and 17 beta-estradiol concentrations, and aromatase activity, during courtship in male Triturus carnifex.

Plasma testosterone and 17 beta-estradiol were monitored during the main phases of male Triturus carnifex courtship. "Inactive" males showed higher levels of testosterone with respect to those found during the various courtship phases. Estradiol was low in "inactive" males, and it reached the highest values at the beginning of the courtship. In addition aromatase activity was higher in the brain of the newts at the beginning of the courtship. These findings seem to support the "aromatization hypothesis" in Triturus carnifex.

Animals↗

fruitless splicing specifies male courtship behavior in Drosophila.

All animals exhibit innate behaviors that are specified during their development. Drosophila melanogaster males (but not females) perform an elaborate and innate courtship ritual directed toward females (but not males). Male courtship requires products of the fruitless (fru) gene, which is spliced differently in males and females. We have generated alleles of fru that are constitutively spliced in either the male or the female mode. We show that male splicing is essential for male courtship behavior and sexual orientation. More importantly, male splicing is also sufficient to generate male behavior in otherwise normal females. These females direct their courtship toward other females (or males engineered to produce female pheromones). The splicing of a single neuronal gene thus specifies essentially all aspects of a complex innate behavior.

Alleles↗