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Gender identity disorder and courtship disorder.

A hypothetical connection between gender identity disorder and courtship disorder was investigated in 274 heterosexual male patients of the following types: nontransvestic fetishists for female attire; transvestic fetishists; patients with gender identity disorder of adolescence and adulthood, nontranssexual type; and transsexuals. Of these patients, 53 had also demonstrated one or more of the putative expressions of courtship disorder. The proportion each of these types contributed to this group of 53 patients with a courtship disorder was compared with the same type's proportional contribution to the group of 221 gender identity patients without a courtship disorder. The transvestic fetishists contributed a significantly larger percentage and the transsexuals a significantly smaller percentage of individuals to the group with a courtship disorder than to that without a courtship disorder. Theoretical implications of this asymmetry are discussed.

Adult↗

Mushroom body ablation impairs short-term memory and long-term memory of courtship conditioning in Drosophila melanogaster.

We have evaluated the role of the Drosophila mushroom bodies (MBs) in courtship conditioning, in which experience with mated females causes males to reduce their courtship toward virgins (Siegel and Hall, 1979). Whereas previous studies indicated that MB ablation abolished learning in an olfactory conditioning paradigm (deBelle and Heisenberg, 1994), MB-ablated males were able to learn in the courtship paradigm. They resumed courting at naive levels within 30 min after training, however, while the courtship of control males remained depressed 1 hr after training. We also describe a novel courtship conditioning paradigm that established long-term memory, lasting 9 days. In MB-ablated males, memory dissipated completely within 1 day. Our results indicate that the MBs are not required for learning and immediate recall of courtship conditioning but are required for consolidation of short-term and long-term associative memories.

Animals↗

Visual input regulates circuit configuration in courtship conditioning of Drosophila melanogaster.

Courtship and courtship conditioning are behaviors that are regulated by multiple sensory inputs, including chemosensation and vision. Globally inhibiting CaMKII activity in Drosophila disrupts courtship plasticity while leaving visual and chemosensory perception intact. Light has been shown to modulate CaMKII-dependent memory formation in this paradigm and the circuitry for the nonvisual version of this behavior has been investigated. In this paradigm, volatile and tactile pheromones provide the primary driving force for courtship, and memory formation is dependent upon intact mushroom bodies and parts of the central complex. In the present study, we use the GAL4/UAS binary expression system to define areas of the brain that require CaMKII for modulation of courtship conditioning in the presence of visual, as well as chemosensory, information. Visual input suppressed the ability of mushroom body- and central complex-specific CaMKII inhibition to disrupt memory formation, indicating that the cellular circuitry underlying this behavior can be remodeled by changing the driving sensory modality. These findings suggest that the potential for plasticity in courtship behavior is distributed among multiple biochemically and anatomically distinct cellular circuits.

Animals↗

Sex steroid communication in the ring dove brain during courtship.

This review examines possible role of progesterone receptor (PR) and androgen receptor (AR) "cross-talk" in the expression of courtship behaviour in the ring dove (Streptopelia risoria). In doves, although androgen has been mostly associated with aggressive courtship behaviour and progesterone with the initiation of incubation, progesterone administration to courting birds terminates the aggressive component of courtship whilst having no effect on nesting behaviour. Recent results in doves have identified a high density of androgen receptor and progesterone receptor immunoreactivity (AR-ir and PR-ir) in the hypothalamus of both sexes in regions known to be directly involved in courtship and incubation behaviour. Nuclear AR-ir in courting birds is widespread throughout the brain. Nuclear PR-ir is only localized in discrete regions of the preoptic hypothalamus of both sexes. In the anterior and posterior hypothalamus of courting birds an increase number of AR-ir and PR-ir neurons colocalizes (70-90%) in the nucleus preopticus anterior (POA), nucleus preopticus medialis (POM), nucleus preopticus paraventricularis magnocellularis (PPM), nucleus hypothalami lateralis posterioris (PLH), and tuberal hypothalamus (Tu). A lower percentage of colocalization is seen in birds at other stages of the breeding cycle. The high percentage of AR-ir and PR-ir colocalization in the preoptic hypothalamus of courting doves supports previous reports involving progesterone acting in these brain regions to terminate the androgen-dependent aggressive courtship behaviour in male doves. The increase in PR-ir staining intensity in AR-ir neurons in courting birds suggests that this progesterone-dependent termination of aggressive courtship display in males occurs at the receptor level and may be orchestrated by central oestrogen.

Animal Communication↗

Male-male courtship behavior induced by ectopic expression of the Drosophila white gene: role of sensory function and age.

Male-male courtship behavior was recently reported to be induced in large populations of Drosophila (e.g., 600-1500 flies) by ectopic expression of the white (w) gene. Little is known about the basis of this behavior; in male-female courtship, sensory cues are believed to play an important role. Previous data are consistent with the possibility that misexpression of w causes abnormal reception or processing of sensory information. We show here that w-induced male-male courtship occurs in isolated pairs of flies. Thus the behavior does not depend on sensory cues found only among large populations of flies, or on cues produced only by a small subset of such populations. This finding enabled quantitative analysis of mechanisms that underlie the behavior. Specifically, male-male courtship does not depend on the reception of olfactory information, nor on the reception or generation of auditory cues, as determined by surgical ablation of antennae, maxillary palps, or wings. Although the rapid onset of the behavior following w induction suggested that its basis could lie in a modulation of sensory physiology, we found visual, olfactory, and gustatory function to be normal in physiological or behavioral tests. The only sensory deprivation to produce an effect on male-male courtship was testing under dim red light; the percentage of flies courting another male was reduced to one-fourth of control values. A striking age dependence of the behavior is also documented: courtship between paired male mini-w+ flies was not observed in tests of very young (1-day-old) flies, but occurs at high levels between the ages of 1 and 4 weeks.

Aging↗

Predator-induced nest site preference: safe nests allow courtship in sticklebacks.

Reproductive activities often increase the susceptibility of individuals to predators. Individuals may be able to reduce this risk of predation, however, by their choice of breeding habitat, as the structural complexity of habitats is known to affect predator foraging success. Here we show that the presence of predators induces a preference for structurally complex nest sites over open ones in male three-spined sticklebacks, Gasterosteus aculeatus. To investigate whether this predator-induced nest site preference can decrease the known negative effect of predators on courtship activity, we recorded the reduction in courtship activity during predator presentations for males in vegetated and open nest sites. Habitat structure affected the response to a predator when males were in competition. A male in a vegetated site reduced courtship activity less than a male in an open site. Habitat had no effect on courtship when males were solitary, however. This suggests that male-male competition and the possibility of losing mating opportunities to other males affect risk taking. Females, who were unaware of the predator, preferred the male in the vegetated site, which showed less reduction in courtship, when the males were exposed to a predator, but chose randomly between the males when the predator was absent. Thus, a preference for vegetated nest sites under predation risk may be beneficial not only by increasing the probability of survival, but also by reducing the negative impact of predators on courtship activity and mating probability. Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

Female responses to isolated signals from multimodal male courtship displays in the wolf spider genus Schizocosa (Araneae: Lycosidae).

Male wolf spiders within the genus Schizocosa display considerable variation in foreleg ornamentation as well as in courtship communication. Multiple modes of male signalling have evolved in a number of species. Divergence in courtship signals among species within this genus may be directly associated with variation in the sensory sensitivities of conspecific females. We isolated the visual and vibratory courtship cues of four species of Schizocosa and recorded conspecific female receptivity to each isolated cue. We also examined female receptivity to complete multimodal courtship signals. We found that the sensory sensitivities of conspecific females were associated with the predominant modes of male courtship communication. Species in which females use mostly stridulatory cues in assessing conspecific males tended to have stridulation-based male courtship displays (S. duplex and S. uetzi) while the opposite was true for species in which females used more visual cues in male assessment (S. stridulans and S. crassipes). This study suggests coevolution between male signal design and female sensory design. We discuss possible scenarios that could be driving this coevolution, including hypotheses of sensory bias and environmental constraints. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

Males mate guard in absentia through extended effects of postcopulatory courtship in the parasitoid wasp Spalangia endius (Hymenoptera: Pteromalidae).

The proximal mechanisms leading to monandry have been little studied in most insect orders, including Hymenoptera. In the parasitoid wasp Spalangia endius, mated females are less attractive (less often mounted) than virgins and are unreceptive (unlikely to allow copulation). Which aspects of mating are responsible was tested by observing male responses toward females whose mating had been interrupted at various stages. All females were allowed to receive precopulatory courtship and to open their genital aperture to copulate. Then some were interrupted before copulation, some after copulation but before postcopulatory courtship, and some were allowed to complete postcopulatory courtship. Females that had copulated were not less attractive than females that had not. In contrast, females that had received postcopulatory courtship were clearly both less attractive and less receptive. Thus, postcopulatory courtship functions as extended mate guarding, by making the female less attractive and less receptive to subsequent males even after the original male is no longer present. The effect of postcopulatory courtship on female attractiveness was persistent but imperfect: when males were presented sequentially to mated females, most but not all males retreated without mounting, and a female could repulse more than twenty males in succession.

Animals↗

Changes in exploration from courtship to the breeding state in red-rumped parrots (Psephotus haematonotus).

Animals explore their environment to gather information about feeding areas, resting places, or suitable territories for reproduction. During the annual cycle seasonal variation in exploration occurs at least in some species. In this study, exploratory behaviour of red-rumped parrots (Psephotus haematonotus) was investigated during part of the reproductive period including courtship and incubation. On 2 test days 14 pairs in either the courtship or the breeding state were confronted with three unknown objects in their familiar aviary for 6 h on each of 2 days. Test days were separated by a 2-day pause. Exploration by the males did not change from courtship to breeding, whereas in females latency to first contact increased, and duration of exploration and the number of objects touched decreased from courtship to breeding state. Considering the pair as a unit, the total duration of exploration was shorter in breeding pairs than in pairs during courtship. Changes in exploration from the first to the second day of object exposure occurred in both sexes. Males explored longer on the second day, females contacted the objects significantly later on the second day but showed no significant changes in duration of exploration. The changes in exploration might be due to changed cost/benefit considerations during courtship and breeding.

Journal Article↗

Male calling between courtship sequences in whitethroats: a way to counter intrusions from neighbouring rivals.

In whitethroats (Sylvia communis) unmated and recently mated males often emit a specific call, the woid-call, after a sequence of courtship diving songs when the female is still present. In the experiments we tested the deterring function of the woid-calls.A simple playback experiment showed that unmated males react to woid-calls played at their territory border by initially increasing their distance to the speaker in an escape-like manner before they approached in the same way as to territorial song. In another experiment we simulated a courtship event at a neighbour by playing back sequences of courtship songs from the territory border until the territory owner started to approach the speaker. We then either continued with one sequence of courtship song or switched to woid-calls. The results showed that the woid-calls delayed approaches from the territory owner. Together, the two experiments suggest a deterring effect of woid-calls. The implications of this for their use in courtship contexts where courting males often suffer intrusions from neighbouring males are discussed. We believe that woid-calls are used both preventively and defensively to avoid intrusions when other males have obtained information about female presence and courtship.

Journal Article↗

Wing-beat frequency mutants and courtship behavior in Drosophila melanogaster males.

Flightless mutations that affect wing-beat frequency (WBF) of Drosophila melanogaster were examined for their effect on male courtship. WBFs were measured using a fixed-wire tether that completely supports the fly in an attitude similar to hovering flight. The two spontaneous mutations, one of which reduces WBF to one-half normal and the other to zero, were placed on an isogenic background and were compared to an isogenic wildtype strain and to a genetically heterogeneous wildtype strain. Time to mating under noncompetitive conditions (single pair matings) was not significantly different among the four male types in one experiment. In a second experiment, although the time to mating varied significantly among the four male types, there was no association between the WBF that was characteristic of a male type and the length of time to mating. Time to mating was not significantly correlated with WBF, wing size, or body mass in either experiment. Genetically heterogeneous wild-type females were significantly more receptive (had shorter times to mating) than inbred wild-type females toward all four male types. During the time-to-mating tests, all four male types appeared to show typical courtship behaviors. Therefore, the male types were compared for possible differences in four components of the male courtship song: sine song frequency, interpulse interval, intrapulse frequency (= carrier frequency), and wing cycles per pulse. One or another of these components showed significant differences among the four male types (e.g., genetically heterogeneous, wild-type males showed a significantly higher sine song frequency and intrapulse frequency than males of the three isogenic types). However, the average values for all four male types were within reported wild-type ranges for each courtship song component. Although the two mutations drastically reduce WBF during tethered flight, they do not have any similar major effect on courtship. Apparently they affect muscles or neuronal control mechanisms that are not common elements shared by the wing movement of flight and male courtship song.

Animals↗

Behavior and cytogenetics of fruitless in Drosophila melanogaster: different courtship defects caused by separate, closely linked lesions.

The fruitless (fru) courtship mutant was dissected into three defects of male reproductive behavior, which were separable as to their genetic etiologies by application of existing and newly induced chromosomal aberrations. fru itself is a small inversion [In(3R) 90C; 91B] on genetic and cytological criteria. Uncovering the fru distal breakpoint with deletions usually led to males with two of the fru courtship abnormalities: no copulation attempts with females (hence, behavioral sterility) and vigorous courtship among males, including the formation of "courtship chains." However, certain genetic changes involving region 91B resulted in males who formed courtship chains but who mated with females. Uncovering the fru proximal breakpoint led to males that passively elicit inappropriately high levels of courtship. This elicitation property was separable genetically from the sterility and chain formation phenotypes and provisionally mapped to the interval 89F-90F, which includes the fru proximal breakpoint. Behavioral sterility and chaining were also observed in males expressing certain abnormal genotypes, independent of the fru inversion. These included combinations of deficiencies, each with a breakpoint in 91B, and a transposon inserted in 91B.

Animals↗

Courtship anomalies caused by doublesex mutations in Drosophila melanogaster.

The role played by the sex-determining gene doublesex (dsx) and its influence on Drosophila courtship were examined. Against a background of subnormal male-like behavior that is reported to be an attribute of haplo-X flies homozygous for the original dsx mutation, and given that a sex-specific muscle is unaffected by genetic variation at this locus, analyses of several reproductive behaviors and control for genetic background effects indicated that XY dsx mutants are impaired in their willingness to court females. When they did court, certain behavioral actions were normal, including components of courtship song. However, these mutants never produced courtship humming sounds. Mature XY dsx flies elicited anomalously high levels of courtship; that this occurs merely because of a delay in imaginal development was experimentally discounted. The current analysis reconciled two ostensibly conflicting reports involving the courtship-stimulating qualities of this mutant type. Such experiments also uncovered a new behavioral anomaly: dsx mutations caused chromosomal males to court other males at abnormally high levels. These results are discussed from the perspective of doublesex's influence on internal tissues of adult Drosophila involved in the triggering and neural control of male- and female-like elements of courtship, reproductive pheromone production, or a combination of such factors.

Analysis of Variance↗

Courtship in Saccharomyces cerevisiae: an early cell-cell interaction during mating.

During conjugation in Saccharomyces cerevisiae, two cells of opposite mating type (MATa and MAT alpha) fuse to form a diploid zygote. Conjugation requires that each cell locate an appropriate mating partner. To investigate how yeast cells select a mating partner, we developed a competition mating assay in which wild-type MAT alpha cells have a choice of two MATa cell mating partners. We first demonstrated that sterile MAT alpha 1 cells (expressing no a- or alpha-specific gene products) do not compete with fertile MATa cells in the assay; hence, wild-type MATa and MAT alpha cells can efficiently locate an appropriate mating partner. Second, we showed that a MATa strain need not be fertile to compete with a fertile MATa strain in the assay. This result defines an early step in conjugation, which we term courtship. We showed that the ability to agglutinate is not necessary in MATa cells for courtship but that production of a-pheromone and response to alpha-pheromone are necessary. Thus, MATa cells must not only transmit but must also receive and then respond to information for effective courtship; hence, there is a "conversation" between the courting cells. We showed that the only alpha-pheromone-induced response necessary in MATa cells for courtship is production of a-pheromone. In all cases tested, a strain producing a higher level of a-pheromone was more proficient in courtship than one producing a lower level. We propose that during courtship, a MAT alpha cell selects the adjacent MATa cell producing the highest level of a-pheromone.

Blotting, Northern↗

Hormonal control of courtship and copulatory behavior in male Cnemidophorus inornatus, a direct sexual ancestor of a unisexual, parthenogenetic lizard.

The hormonal control of courtship and copulatory behavior in male Cnemidophorus inornatus, one of the gonochoristic ancestral species of the all-female, parthenogenetic C. uniparens, was determined. Significantly fewer castrated males courted females than intact and sham-castrated control males. Silastic capsules containing dihydrotestosterone or testosterone reinstated courtship and copulatory behavior in a significant number of castrates. While significantly greater numbers of castrates treated with androgens resumed courtship, some of the castrates receiving progesterone implants also resumed intense courtship and copulatory behavior. Exogenous progesterone also maintained courtship behaviors in a significant number of intact males at a time when control males ceased to court. The implications of these findings for our understanding of the evolution of hormone-brain-behavior relationships and sex steroid hormone mechanisms of action are discussed.

Animals↗

Hormonal independence of courtship behavior in the male garter snake.

Garter snakes exhibit a dissociated reproductive tactic in which gonadal activity is minimal at the time of mating, increasing only after the breeding season has ended. Experiments are presented demonstrating that neither short-term nor longterm castration affects courtship behavior in adult male red-sided garter snakes (Thamnophis sirtalis parietalis). So long as males have passed through a low-temperature dormancy period (hibernation), castration either shortly after emergence in the spring, or before entering winter dormancy in the fall, does not prevent the display of intense courtship behavior on emergence. Similarly, males castrated during mating activity the previous spring prior to the annual testicular growth phase actively courted females on emergence from hibernation. Males adrenalectomized and castrated during low-temperature dormancy also courted females on emergence. Hypophysectomy during or before low-temperature dormancy did not prevent males from displaying high-intensity courtship behavior on emergence from hibernation. Treatment with sex steroid hormones, as well as hypothalamic and pituitary hormones, and a variety of neural and metabolic affectors also fails to elicit courtship behavior in noncourting males during the summer. It was concluded that causal mechanisms controlling courtship behavior in the red-sided garter snake are fundamentally different, at least at the physiological level, from those mechanisms described for many laboratory and domesticated species.

Adrenal Glands↗

Role of the anterior hypothalamus-preoptic area in the regulation of courtship behavior in the male Canadian red-sided garter snake (Thamnophis sirtalis parietalis): lesion experiments.

Lesions of the medial preoptic area and/or the ventromedial hypothalamus resulted in an abrupt and immediate decline in courtship behavior in adult male Canadian red-sided garter snakes (Thamnophis sirtalis parietalis). Lesions of the more anterior portions of the preoptic area resulted in a more gradual, delayed decline in courtship behavior. Male snakes sustaining lesions dorsal, ventral, or caudal to the anterior hypothalamus-preoptic area exhibited no change in courtship behavior relative to controls. Measurements of testis size, spermatogenic stage, and circulation levels of androgens revealed no differences between any of the groups. There were marked differences in the change in hematocrit over time between the groups. These results indicate that the anterior hypothalamus-preoptic area is involved in the control of courtship behavior in the adult male red-sided garter snake. Moreover, it is suggested that the stimulus affecting the anterior hypothalamus-preoptic area to activate courtship is temperature related.

Androgens↗

Cholinergic neurons mediate CaMKII-dependent enhancement of courtship suppression.

In Drosophila, calcium/calmodulin-dependent protein kinase II (CaMKII) activity is crucial in associative courtship conditioning for both memory formation and suppression of courtship during training with a mated female. We have previously shown that increasing levels of constitutively active CaMKII, but not calcium-dependent CaMKII, in a subset of neurons can decrease the initial level of courtship and enhance the rate of suppression of courtship in response to a mated female. In this study, we demonstrate that a subpopulation of noncholinergic, nondopaminergic, non-GABAergic neurons can cause CaMKII-dependent reductions in initial courtship, but only cholinergic neurons enhance training-dependent suppression. These data suggest that processing of pheromonal signals in two subpopulations of neurons, likely antennal lobe projection neurons, is critical for behavioral plasticity.

Adaptation, Psychological↗