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Courtship behavior of brain mosaics in Drosophila.

Sites in the brain that show functional, sexual dimorphism in courtship behavior have been mapped at high resolution in male/female mosaics of Drosophila melanogaster. The sex mosaics were produced by enhancer-trap expression of GAL4 driving the female-spliced form of the transformer gene (tra), revealing sites in the dorsal brain, lateral protocerebrum, suboesophageal, thoracic and abdominal ganglia, and suggesting the importance of cross-talk between these regions in the implementation of the courtship sequence.

Animals↗

Abnormal courtship conditioning in males mutant for the RI regulatory subunit of Drosophila protein kinase A.

The previously described site-selected P-element mutagenesis of a Drosophila gene encoding the regulatory subunit of cAMP-dependent protein kinase generates mutants that have defective behavior in the olfactory learning test. Here we describe the effect of the same mutations in a courtship conditioning assay. Wild-type males can distinguish between virgin females (which they court vigorously), and fertilized females (which they court less vigorously). After exposure to fertilized females, wild-type males modify their behavior by decreasing courtship to subsequent target virgins, an effect that may last for many hours. Like wild-type males, PKA-RI mutant males are also able to distinguish between virgin and fertilized females. PKA-RI males also modify their behavior towards virgin females after prior exposure to a fertilized female, but such an effect is short-lived, suggesting a defect in memory rather than learning. We also show that under these conditions the behavior of PKA-RI males is similar to that of amnesiac, dunce and rutabaga males.

Animals↗

[Mating success and courtship ritual in strains of Drosophila melanogaster carrying mutation flamenco].

Mating success was examined in groups of Drosophila melanogaster carrying mutation flamMS (SS, MSn1-2, and MSn1-3) and in wild-type flies. The proportion of normally copulating males was significantly lower in the mutant strains. The reduction in mating efficiency was caused by changes in male behavior rather than in female attractiveness. Individual analyses showed that male mating behavior in strains carrying flamMS was qualitatively and quantitatively different from that in the wild-type strain Canton S. The proportion of males that performed consecutive courtship stages was significantly lower in the mutant strains. The sequence and duration of some courtship stages (in particular, orientation and wing vibration) in mutant flies was shown to be altered. The significance of the flamenco locus in regulation of processes occurring at the organismal level are discussed.

Animals↗

The dissonance mutation at the no-on-transient-A locus of D. melanogaster: genetic control of courtship song and visual behaviors by a protein with putative RNA-binding motifs.

Genetic and molecular results are here presented revealing that the dissonance (diss) courtship song mutation is an allele of the no-on-transient-A (nonA) locus of Drosophila melanogaster. diss (now called nonAdiss) was originally isolated as a mutant with aberrant pulse song, although it was then noted to exhibit defects in responses to visual stimuli as well. The lack of transient spikes in the electroretinogram (ERG) and optomotor blindness associated with nonAdiss are shown to be similar to the visual abnormalities caused by the original nonA mutations. nonAdiss failed to complement either the ERG or optomotor defects associated with four other nonA mutations. However, all four of these nonA mutants--which were isolated on visual criteria alone--sang a normal courtship song. nonAdiss complemented at least three of the nonA mutations with regard to the singing phenotype, as assessed by a new method for temporal analysis of the male's pulse song. Both visual and song abnormalities caused by nonAdiss were rescued by P-element-mediated transformation with overlapping 11 and 16 kilobase (kb) fragments of genomic DNA (originally cloned from the nonA locus by Jones and Rubin, 1990). Analysis of behavioral phenotypes in transformed flies carrying mutagenized versions of the 11 kb genomic fragment (in a nonAdiss genomic background) localized the rescuing DNA to a region containing an open reading frame that encodes a polypeptide (NONA) with similarity to a family of RNA-binding proteins. Immunohistochemical determination of NONA's spatial and temporal expression revealed that it is localized to the nuclei of cells in many neural and non-neural tissues, at all stages of the life cycle after very early in development. Genetic connections between the control of two quite different behaviors--reproductive and visual--are discussed, along with precedences for generally expressed gene products playing roles in specific behaviors.

Alleles↗

Sexual dimorphisms in a neuromuscular system regulating courtship in the green anole lizard: effects of season and androgen treatment.

During the breeding season, male green anoles (Anolis carolinensis) court females by extending a red throat fan called a dewlap. Motoneurons controlling this sexually dimorphic behavior are located in two portions of the brain stem: (a) the vagal portion of nucleus ambiguus (AmbX), and (b) the region containing the glossopharyngeal portion of nucleus ambiguus and the ventral motor nucleus of the facial nerve (AmbIX/VIImv). These motoneurons project to the ceratohyoideus muscle via the ramus pharyngo-laryngeus IX+X. To investigate the effects of season on and androgen regulation of neural and peripheral structures controlling dewlap extension, two experiments were conducted: (a) During the breeding and nonbreeding seasons, motoneuron number, soma size, and nucleus size were investigated in intact males and females and in castrated males treated with a testosterone propionate (TP) or a blank Silastic capsule. (b) Cross-sectional area of the nerve and muscle fiber size, number, and density were investigated in the four treatment groups during the breeding season only. No significant differences were found in motoneuron number. In the breeding season, subtle male-biased sex differences existed in both AmbX and AmbIX/VIImv soma size. Nerve cross-sectional area and muscle fiber size and number were substantially larger in males than females. Muscle fiber density was higher in females. No consistent effects due to season or androgen treatment were detected, although characteristics of motoneurons were in some cases slightly larger in the nonbreeding season. These results suggest that, while parallels to behavior exist between the sexes, morphological changes in adulthood in the dewlap motoneurons and muscle do not normally regulate courtship behavior in the male green anole.

Animals↗

Variation in courtship sounds among three geographical strains of Drosophila mercatorum.

Sound recordings of courtship in Drosophila mercatorum were analyzed with an oscilloscope. Sounds in this species consist of two kinds of pulses, referred to as the A and B sounds, respectively. These differ from each other in their oscilloscope pattern and in the behavior accompanying them. A comparative study of three strains from widely separated geographical regions (New York, El Salvador, and Hawaii) revealed significant differences among strains in the interpulse interval (ipi) of the A sound. The ipi of the B sound increased as it proceeded for New York and Hawaii males but not for males from El Salvador. These characteristics may influence mating success in interstrain crosses. The present results suggest that the first steps toward divergence in the nature of the sound are quantitative; this may affect the response threshold of the female. Genetic systems responsible for female receptivity may have been independently developed in the two sexes.

Animal Communication↗

Spectral analysis of courtship songs in behavioral mutants of Drosophila melanogaster.

Spectral analyses were applied to the courtship songs of the mutants cacophony (cac), dissonance (diss), fruitless (fru), and period (per), as well as to the double mutant cac diss. Aberrant intervals between song pulses were observed in diss, cac, cac diss, and fru songs. diss males displayed a defect in song hums manifested by an irregular sine wave, although the fundamental frequencies were normal. Sine song frequencies and intrapulse frequencies were aberrant in cac diss males. Two per mutant alleles (pero1 and pers) were associated with normal song [corrected] pulses and hums. These findings are discussed with regard to the mechanisms of song production and the role of these sounds in Drosophila reproduction.

Animals↗

The courtship disorders: a further investigation.

The senior author had put forward the hypothesis that voyeurism , exhibitionism, obscene telephone calling, toucheurism - frotteurism , and at least some cases of the preferential rape pattern all distort normal courtship behavior in basically the same manner. An earlier study (Freund et al., 1983) tested part of this hypothesis, namely that voyeurism , exhibitionism, and frotteurism - toucheurism , are closely related. However, the outcome of one of the experiments by which this test was performed could be interpreted as indicating that the relationship between these anomalous erotic inclinations consists simply in that they may be expressions of a disinclination toward intercourse. The present study was conducted to test this interpretation. It compared 16 exhibitionists with 16 sexually normal controls on penile responses to auditorily presented descriptions of the four phases of normal sexual interaction (location of a suitable partner; pretactile erotic interaction; tactile interaction short of intercourse; the situation of intercourse) and sexually neutral situations. These descriptions were narratives about the examined subject's involvement in such interaction. No differences in responding between exhibitionists and normal controls were observed. Both groups responded most to descriptions of intercourse. The second largest response was to tactile interaction (short of intercourse). The third was to situations of partner location and pretactile interaction, which were not differentiated from each other. Responses to sexually neutral situations were smaller than those to descriptions of any category of sexual interaction. This result refutes the hypothesis that exhibitionism develops on the basis of a disinclination toward intercourse. The close relationship between exhibitionism, and voyeurism and toucheurism (or frotteurism ), had been demonstrated earlier.(ABSTRACT TRUNCATED AT 250 WORDS)

Adult↗

Sexual coloration, plasma concentrations of sex steroid hormones, and responses to courtship in the female keeled earless lizard (Holbrookia propinqua).

The reproductive condition, steroid hormone concentrations in the plasma, and behavior of bright and plain female Holbrookia propinqua were examined. Bright females performed significantly more aggressive rejection of courtship than plain females. Bright females were significantly more likely than plain females to have follicles larger than 5.0 mm or oviductal eggs; females with large follicles or oviductal eggs had significantly higher concentrations of progesterone and androgen than those with small follicles and lacking oviductal eggs. Plasma progesterone, androgen, and estradiol levels in the females studied behaviorally were significantly higher for the bright females than for the plain ones. Females undergoing rapid brightening were significantly more likely to be sexually receptive and copulate than were either plain or fully brightened females. The role of sex steroid hormones in coloration and behavior and the adaptive value of chromatic signalling by females are discussed.

Aggression↗

Wings and legs are production sites for the desert locust courtship-inhibition pheromone, phenylacetonitrile.

Mature gregarious male desert locusts, Schistocerca gregaria, emit the courtship inhibition pheromone phenylacetonitrile. Wings and legs, in particular the fore wings, have been identified as the main releasing sites. Abdomen and head emit only trace amounts of this pheromone. In contrast veratrole, another typical component of male volatiles, is emitted by all body parts. Epidermal gland cells in the identified phenylacetonitrile releasing appendages are the putative sites of its biosynthesis. Incubation of these body parts in the presence of (14)C-phenylalanine results in the production of (14)C-phenylacetonitrile. Some of the phenylacetonitrile appears to be degraded to HCN and benzaldehyde presumably enhancing the repellent character of phenylacetonitrile. HCN is only detectable in volatiles of mature gregarious male desert locusts. Possible advantages of the observed distribution of the phenylacetonitrile release sites and of the cyanogenesis in relation to mating behaviour are discussed.

Acetonitriles↗

Testosterone is correlated with courtship but not aggression in the tropical buff-banded rail, Gallirallus philippensis.

Past studies into the roles of testosterone in birds have focused on species that occur in temperate regions. In such species, plasma testosterone levels are high during the establishment of territories in the spring and are associated with increased aggression. In contrast to most temperate species, tropical birds frequently defend territories year-round, during which time territoriality often occurs in a nonsexual context. The few studies that have been carried out on tropical birds show lower levels of circulating testosterone than occur in their temperate counterparts. In some year-round territorial tropical species, testosterone and aggression are dissociated, while in other species testosterone still plays a role in regulating aggression. This study examined the relationship between aggression and plasma testosterone levels in a year-round territorial, subtropical population of the buff-banded rail with characteristics typical of tropical species. Peak testosterone levels were substantially lower than those found in temperate species. Males displayed a seasonal peak in plasma testosterone level when their partners were most likely to be fertile. At other times, testosterone levels were mostly undetectable, despite year-round territoriality. We found that T levels increased with courtship behavior but showed no relationship with aggression, supporting the hypothesis that dissociation between testosterone and territoriality may be widespread among tropical avian taxa.

Aggression↗

Arginine vasotocin effects on courtship behavior in male white perch (Morone americana).

Arginine vasotocin (AVT) and its mammalian homologue, arginine vasopressin (AVP), have been shown to have widespread behavioral effects in vertebrates. AVT was evaluated for its effectiveness in stimulating an important courtship behavior termed 'attending' in male white perch, Morone americana. Attending consists of close and continuous following of the female with occasional contact in the abdominal area. We tested the behavioral effectiveness of AVT in stimulating attending when administered either intraperitoneally (IP) or intracerebroventricularly (ICV). We also tested IP injections of AVT alone and in combination with an AVP V(1) receptor antagonist (Manning compound). None of the IP injections of either AVT or Manning compound produced consistent effects on attending behavior. In contrast, ICV injections of AVT did significantly increase attending behavior and at low dosages. Circulating levels of testosterone and 11-ketotestosterone were not affected approximately 80 min following injection by any of the treatments. The strong behavioral effects observed with ICV administration support a central site of action for AVT in stimulating attending behavior. This is a complex behavior that shows similarities to behaviors mediated by AVT and AVP in other vertebrates, providing further evidence of a conserved behavioral role for these peptides.

Androgens↗

Role of X chromosomal song genes in the evolution of species-specific courtship songs in Drosophila virilis group species.

In the Drosophila virilis group the males of the virilis phylad species produce courtship song consisting of pulse trains with no pauses between successive sound pulses, whereas the males of the montana phylad species produce songs with clear pauses between the sound pulses. We obtained song data for F1 hybrids between D. virilis (representing the virilis phylad) or D. flavomontana (representing the montana phylad) females and the males of several species of the D. virilis group to study the interaction of X chromosomal and autosomal song genes affecting species differences in song. In crosses with D. virilis females, X chromosomal (or maternal) factors masked variation in pulse length despite variation in heterospecific autosomal song genes. To the contrary, in crosses with D. flavomontana females, X chromosomal genes largely determined the pause length and interacted with autosomal genes to determine the pulse length. In the montana phylad species, pulse length showed dominance toward shorter pulses and pause length toward longer pauses. The first-mentioned trait also indicated the epistatic effects of X chromosomal and autosomal components.

Analysis of Variance↗

Mating behaviour and evidence for a female released courtship pheromone in the signal crayfish Pacifastacus leniusculus.

The mating behaviour of the signal crayfish, Pacifastacus leniusculus (Dana), has been categorised into seven distinct stages: orientation; contact; seizure; turning; mounting; spermatophore deposition; and dismounting. These categories were used in the development of a bioassay to test whether mature females release a sex pheromone during the breeding season to which males respond. Water conditioned by 1) mature females (MF water), 2) immature females (IF water) and 3) freshwater (C water), were injected through standard aquarium air-stones into tanks containing mature males. The behaviour of the male was recorded on video for 15 minutes before and after injection of each test water. The time that the males spent exhibiting the three behavioural categories of 'quiescent,' 'motile' and 'handling' of the air-stone, were recorded. 'Handling' of the air-stone included the mating behaviours of seizure, mounting, and spermatophore deposition onto the surface of the air-stone. Males exposed to MF water exhibited significantly increased levels of motile activity and handling behaviour, than males exposed to C water or IF water. These results demonstrate for the first time the presence of a sex pheromone, released during the breeding season by mature females, that stimulates courtship and mating behaviour in male P. leniusculus.

Animals↗

Differential effects of courtship and mating on receptivity and brain metabolism in female red-sided garter snakes (Thamnophis sirtalis parietalis).

In the female red-sided garter snake (Thamnophis sirtalis parietalis), the loss of receptivity following intromission during mating can be prevented by injection of a local anesthetic (tetracaine) in the cloacal region prior to courtship and mating. Females that were courted and then mated had significantly higher uptake of radio-labeled [14C]2-fluoro-2-deoxyglucose (2-DG) in the preoptic area (25%) and significantly lower uptake in the ventromedial hypothalamus (-20%) compared with females that were courted but not mated. Tetracaine-treated females had accumulation patterns similar to courted but unmated females and to females exposed only to other females. These results suggest that in the female red-sided garter snake, sensory input from the cloaca during mating alters patterns of metabolism in those brain areas most often associated with female sexual behavior.

Anesthetics, Local↗

Multi-layered regulation of courtship behaviour.

The fruitless gene governs courtship in male, but not female, Drosophila, yet it is expressed and specifically spliced in the brains of both sexes. New experiments reveal that a splice-recognition site retained in the mature message in females provides the basis for sex-specific translational repression.

Alternative Splicing↗

Kin selection and cooperative courtship in wild turkeys.

In the few species of birds in which males form display partnerships to attract females, one male secures most or all of the copulations. This leads to the question of why subordinate males help in the absence of observable reproductive benefits. Hamilton's concept of kin selection, whereby individuals can benefit indirectly by helping a relative, was a crucial breakthrough for understanding apparently altruistic systems. However in the only direct test of kin selection in coordinated display partnerships, partners were unrelated, discounting kin selection as an explanation for the evolution of cooperation. Here I show, using genetic measures of relatedness and reproductive success, that kin selection can explain the evolution of cooperative courtship in wild turkeys. Subordinate (helper) males do not themselves reproduce, but their indirect fitness as calculated by Hamilton's rule more than offsets the cost of helping. This result confirms a textbook example of kin selection that until now has been controversial and also extends recent findings of male relatedness on avian leks by quantifying the kin-selected benefits gained by non-reproducing males.

Altruism↗

Overexpression of the Drosophila vesicular monoamine transporter increases motor activity and courtship but decreases the behavioral response to cocaine.

Aminergic signaling pathways have been implicated in a variety of neuropsychiatric illnesses, but the mechanisms by which these pathways influence complex behavior remain obscure. Vesicular monoamine transporters (VMATs) have been shown to regulate the amount of monoamine neurotransmitter that is stored and released from synaptic vesicles in mammalian systems, and an increase in their expression has been observed in bipolar patients. The model organism Drosophila melanogaster provides a powerful, but underutilized genetic system for studying how dopamine (DA) and serotonin (5HT) may influence behavior. We show that a Drosophila isoform of VMAT (DVMAT-A) is expressed in both dopaminergic and serotonergic neurons in the adult Drosophila brain. Overexpression of DVMAT-A in these cells potentiates stereotypic grooming behaviors and locomotion and can be reversed by reserpine, which blocks DVMAT activity, and haloperidol, a DA receptor antagonist. We also observe a prolongation of courtship behavior, a decrease in successful mating and a decrease in fertility, suggesting a role for aminergic circuits in the modulation of sexual behaviors. Finally, we find that DMVAT-A overexpression decreases the fly's sensitivity to cocaine, suggesting that the synaptic machinery responsible for this behavior may be downregulated. DVMAT transgenes may be targeted to additional neuronal pathways using standard Drosophila techniques, and our results provide a novel paradigm to study the mechanisms by which monoamines regulate complex behaviors relevant to neuropsychiatric illness.

Animals↗