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Multivariate analysis of Drosophila courtship.

Courtship records of 15 pairs of Drosophila melanogaster were analyzed for temporal stationarity of courtship behaviors, behavioral diversity, behavioral intercorrelations, sequential properties, and information transmission for both sexes. Durations of one male behavior, "orient-back," and two female behaviors, "preen" and "stand still," were found to change from the first to the second half of courtship. Male diversity was greater than female diversity, and both were stationary over time. Correlation analyses failed to single out any particular male or female behaviors as being influential in controlling courtship duration. Male behavior sequences formed several multibehavior loops; female behavior consisted of only a few terminal two-tuple transitions. Transmission analysis carried out on the joint male/female transition matrix showed a higher transmission rate from males to females (12%) than from females to males (7%). Potential applications of this multivariate analysis to investigations of neurobiological and evolutionary aspects of Drosophila courtship behavior are proposed.

Journal Article↗

Behavioral and sensory basis of courtship success in Drosophila melanogaster.

In Drosophila some individuals are more successful at mating than others. Reproductive fitness is strongly dependent upon the ability to recognize and compete for members of the opposite sex. Experiments were designed to answer two questions. (i) What behavioral components are characteristic or predictive of successful courtship? and (ii) How important is the information transmitted in the different sensory channels for courtship success in each sex? These questions were approached by two experimental procedures. Flies having a sensory deficiency (olfactory, auditory, or visual) competed with wild-type flies of the same sex for mates. Males were found to rely upon sensory channels different from those used by females in order to court successfully. In addition, the courtships of pairs of various genotypes were recorded and subjected to multivariate analysis. The multivariate courtship profiles deviated most widely from those of successful wild-type pairs when the male or female was unable to receive information in the sensory channel most important for successful mating by that sex. Both sequential and quantitative courtship properties were altered when one participant was deficient in ability to receive certain sensory information.

Animals↗

Control of male reproductive behavior by the central nervous system of Drosophila: dissection of a courtship pathway by genetic mosaics.

In gynandromorphs of Drosophila, a detailed examination was made of the association between male courtship behavior and the chromosomal genotype of various parts of the central nervous system. Mosaic flies that behave as males repeatedly show a shorter courtship than normal males. If there is to be male behavior, the posterior dorsal brain must be haplo-X on at least one side for occurrence of the early courtship events. Tapping, following of females and wing extension. Licking (proboscis extension) has nearly the same focus but is submissive; that is, male tissue must be present in both left and right dorsal brain. The next courtship step, attempted copulation, has a focus (especially for actual genital contact) located in the thoracic ganglia, though apparently not in a discrete region. Attempted copulation, which can occur even in mosaics with a gravid abdomen, may be correlated with the presence of sex combs. The role of courtship foci are interpreted in terms of known sensory inputs to and functions of the major insect ganglia.

Animals↗

Neural control of homosexual courtship in Drosophila melanogaster.

Immature D. melanogaster males, like virgin females, often elicit vigorous courtship from mature males. Since males perform the same behaviors in response to attractive males and females, the question arises as to whether the foci--cells in the courting male's nervous system that must be haplo-X for the fly to perform a behavior--are the same for homosexual and heterosexual courtship. To answer this question, we analyzed the behaviors that normal males perform in response to sexually attractive flies. From these data, we calculated the probabilities that a fly with haplo-X tissue in a focus for homosexual and heterosexual courtship would perform the behavior in response to one or both of the two sex objects with which it was tested. Next, we observed the courtship behaviors that gynandromorphs (sex mosaics) performed in response to attractive males and females. Since the numbers of mosaics that performed a behavior in response to one or both of the sex objects with which they were tested were not significantly different from the predicted values, we conclude that the foci for performance of homosexual and heterosexual courtship are almost certainly identical.

Animals↗

Removal of both antennae influences the courtship and aggressive behaviors in male crickets.

To test whether insect antennae are necessary for eliciting courtship and aggression toward appropriate partners, we antennectomized adult male crickets (Gryllus bimaculatus) and observed their behavior toward other antennectomized males and intact females. At 7 days after removal of both antennae, pairs of antennectomized males were placed together; 80% displayed courtship behavior, generating courtship song by rubbing their forewings together, toward other antennectomized males, and 20% displayed aggressive behavior. Only 45% courted intact females. No intact males courted antennectomized males, and 80% displayed aggressive behavior. All intact males courted females. The results for males with one antenna removed were essentially the same as for intact males. These findings indicate that a high proportion of male crickets with both antennae removed court other males and fail to display male-male aggression, demonstrating that removal of antennae from male crickets induces male-male courtship and that an antenna is necessary for the expression of male-male aggression. Moreover, brain serotonin (5-hydroxytryptamine; 5-HT) levels in male crickets were significantly reduced at 7 days after removal of antennae. The reduction of 5-HT was detected primarily in the central body of the brain. Thus, 5-HT in the central body of the male cricket brain may be involved in the behavioral changes.

Aggression↗

Sex steroid and corticosterone levels of Adélie penguins (Pygoscelis adeliae) during courtship and incubation.

Plasma levels of sex steroids (progesterone, 17alpha-hydroxyprogesterone, total androgens, and estradiol) were measured at six stages from courtship to late incubation in Adélie penguins. The pattern of change detected in the levels of plasma total androgens (males) and estradiol (females) was consistent with that found in many birds, with elevated levels during courtship (total androgens, 4 ng/ml; estradiol, 0.5 ng/ml) declining to low, stable levels during incubation. Progesterone levels declined moderately from 1.3 to 0.75-1.0 ng/ml in females following egg laying, but levels of 0.8-1.2 ng/ml persisted in males throughout the study period. 17alpha-Hydroxyprogesterone levels were consistently low (approximately 0.2 ng/ml) in females but progressively declined in males from 0.75 during courtship to <0.3 ng/ml at egg laying and during foraging. Plasma corticosterone levels were measured over the same period and were elevated in both males and females at courtship (16-18 ng/ml) and while fasting on the nest (11-15 ng/ml), but had declined in birds returning from foraging at sea, suggesting that elevated levels are related to the metabolic demands of fasting.

17-alpha-Hydroxyprogesterone↗

Behavior and single gene substitution in Drosophila melanogaster. I. Mating and courtship differences with w, cn, and bw loci.

The effect of single allele substitutions into an isogenic background in Oregon-R inbred lines of Drosophila melanogaster on courtship and mating patterns has been studied. A comparison has been made between the white locus w, wco, we, the wild type w+, cn, bw, and cn bw to test the effect of eye pigmentation in influencing courtship and mating patterns. It was found that w, we, wco, cn, and bw females were more successful in mating than were wild-type and cn bw females, cn bw females being less successful than wild-type females. Also, w and cn bw males were equally successful in mating but less successful than wild-type males during the 20-min test period. The mutant males performed as well as the wild-type after courtship was initiated. The behavioral parameters measured were (1) courtship latency, the time from exposure of male to female until orientation; (2) mating speed, the time from beginning of orientation of male to female until successful copulation, and (3) copulation time.

Animals↗

Vasotocin treatment inhibits courtship in male zebra finches; concomitant androgen treatment inhibits this effect.

Zebra finches evolved in arid areas of Australia. Their reproduction is stimulated by water availability, which is unpredictable. Cheng (Poult. Sci. Rev. 5 (1993) 37) hypothesized that the primary mechanism controlling reproduction in species relying on unpredictable cues should be inhibitory. The onset of stimulatory environmental conditions terminates the inhibition, allowing rapid initiation of reproduction. As the primary hormone regulating water balance in birds, arginine vasotocin (AVT) appears a likely candidate to modulate reproduction in finches. Drought conditions cause sustained AVT release, which in other species inhibits androgen production. To determine whether increased AVT inhibits reproductive behavior, intact males were tested with females and divided into three groups matched for courtship behavior. Osmotic minipumps containing (a) saline, (b) 264 ng AVT, or (c) 1320 ng AVT in saline were implanted subcutaneously and males tested 48 h later. AVT-treated males socialized with females, but the high dose significantly reduced singing and courtship displays. To determine whether AVT acted by depressing androgen secretion, additional males were given subcutaneous androgen implants and divided into two groups matched for courtship behavior. Males were then implanted with minipumps containing (a) saline or (b) the high AVT dose. Males treated with AVT plus androgen showed no deficits in courtship behavior. These data suggest that AVT secretion during periods of drought may inhibit reproduction by inhibiting androgen production. Inhibition of reproductive behavior by AVT may be a more general phenomenon. Large quantities of AVT or, in mammals the closely-related peptide vasopressin (VP), are released when animals are stressed, and high levels of AVT/VP may inhibit reproductive behavior. The extremely short half-life of these peptides means that once proximal factors become more favorable, the gonads should rapidly be released from the peptides' inhibitory actions.

Androgens↗

The role of sex steroids in courtship, pairing and pairing behaviors in the socially monogamous zebra finch.

The purpose of this study was to test whether sex steroid actions are necessary for courtship and pairing in socially monogamous birds. We examined the effects of an aromatase inhibitor, 1,4,6-androstatriene-3,17-dione (ATD), combined with an anti-androgen, flutamide (F), on the behavior and pairing status of initially unpaired male and female zebra finches (Taeniopygia guttata). In the first experiment, 24 adult males were implanted with either a combination of ATD and flutamide or empty implants. Two weeks after implantation, birds were housed in aviaries containing 3 ATD + F males, 3 control males, and 3 females and allowed 2 weeks to pair, with observations 7 times during the 2-week period. A second experiment tested the effects of these same treatments in females. During the first 4 days of testing, ATD + F males were less likely to attack conspecifics than were control males. ATD + F males were also less likely to "greet," or approach, females than were control males, but other courtship behaviors, including directed singing, were unaffected. ATD + F females did not differ from control females on any courtship behavior measured. Furthermore, these treatments did not affect pairing behaviors (time spent clumping or in a nest box together) or the likelihood of pairing with a partner of the opposite sex. ATD + F treatments in females did, however, increase the likelihood of same-sex pairing. This suggests that, although sex steroids may regulate some courtship behaviors in males, they do not regulate pairing behaviors and have little effect on the likelihood that a male or female will be chosen as a mate by a bird of the opposite sex.

Aggression↗

A putative Drosophila pheromone receptor expressed in male-specific taste neurons is required for efficient courtship.

Propagation in higher animals requires the efficient and accurate display of innate mating behaviors. In Drosophila melanogaster, male courtship consists of a stereotypic sequence of behaviors involving multiple sensory modalities, such as vision, audition, and chemosensation. For example, taste bristles located in the male forelegs and the labial palps are thought to recognize nonvolatile pheromones secreted by the female. Here, we report the identification of the putative pheromone receptor GR68a, which is expressed in chemosensory neurons of about 20 male-specific gustatory bristles in the forelegs. Gr68a expression is dependent on the sex determination gene doublesex, which controls many aspects of sexual differentiation and is necessary for normal courtship behavior. Tetanus toxin-mediated inactivation of Gr68a-expressing neurons or transgene-mediated RNA interference of Gr68a RNA leads to a significant reduction in male courtship performance, suggesting that GR68a protein is an essential component of pheromone-driven courtship behavior in Drosophila.

Animals↗

Effect of genes, social experience, and their interaction on the courtship behaviour of transgenic Drosophila males.

Behaviour depends (a) on genes that specify the neural and non-neural elements involved in the perception of and responses to sensory stimuli and (b) on experience that can modulate the fine development of these elements. We exposed transgenic and control Drosophila melanogaster males, and their hybrids, to male siblings during adult development and measured the contribution of genes and of experience to their courtship behaviour. The transgene CheB42a specifically targets male gustatory sensillae and alters the perception of male inhibitory pheromones which leads to frequent male-male interactions. The age at which social experience occurred and the genotype of tester males induced a variable effect on the intensity of male homo- and heterosexual courtship. The strong interaction between the effects of genes and of social experience reveals the plasticity of the apparently stereotyped elements involved in male courtship behaviour. Finally, a high intensity of homosexual courtship was found only in males that simultaneously carried a mutation in their white gene and the CheB42a transgene.

Animals↗

Preferential rape in the theory of courtship disorder.

The theory of courtship disorder suggests voyeurism, exhibitionism, frotteurism, and preferential rape (a paraphilic preference for coercive sex) are expressions of a common underlying disturbance. Previous research has demonstrated that voyeurism, exhibitionism, and frotteurism are relatively likely to co-occur; however, their associations with rape are weaker. One explanation is that rapists are more heterogeneous and may not always be motivated by a paraphilic interest in coercive sex. Paraphilic interests can be identified using phallometric testing, but the sensitivity of the test is attenuated by voluntary control over erectile responding. Another approach is to look for a behavioral marker associated with paraphilic rape-proneness. Freund (1990) has argued that exhibitionism may be the "hub" of courtship disorder because exposing one's genitals to a stranger is rare in control subjects. We predicted that rapists who also engaged in exhibitionistic activity would be more likely than other rapists to also engage in other expressions of courtship disorder, i.e., voyeurism and frotteurism. To rule out the possibility that rapists who report exhibitionistic activity only differ in their willingness to report anomalous sexual behavior, we also predicted no difference between the groups in their self-reported involvement in other paraphilic activities such as sadism or masochism, or paraphilic target preferences such as fetishism or transvestism. Finally, we predicted no difference between groups in their willingness to admit to attempting or committing rape. The results supported our predictions, and lend credence to the notion of preferential rape as an expression of courtship disorder. Moreover, these results suggest that the presence of another paraphilia, particularly exhibitionism, may be useful as a behavioral marker for paraphilic rape-proneness.

Adult↗

The role of central parts of the brain in the control of sound production during courtship in Drosophila melanogaster.

The question of the roles of the two main parts of the insect brain, the mushroom bodies and the central complex, in controlling motor coordination and triggering a variety of behavioral programs, including sound production, remains controversial. With the aim of improving our understanding of this question, we studied the parameters of songs used by five-day-old males during courtship for fertilized wild-type females (Canton-S, C-S) over 5-min periods at 25 degrees C; males were of two wild-type Drosophila melanogaster lines (Berlin and C-S). Berlin males lacking mushroom bodies because of treatment with hydroxyurea during development (chemical removal of the mushroom bodies) were used, along with two mutants with defects in the mushroom bodies (mbm1 and mud1), two mutants with defects in the central complex (ccbKS127 and cexKS181), and mutant cxbN71 with defects in both the mushroom bodies and the central complex. The experiments reported here showed that courtship songs in males lacking mushroom bodies were virtually identical to those of wild-type males. The main parameters of pulsatile song in mutants mbm1 and mud1 (interpulse interval and train duration) were insignificantly different from those of the songs of wild-type flies, though the stability of the pulse oscillator was the same. Flies of these lines were no different from wild-type flies in terms of courtship success (percentage of copulating pairs in 10-min tests). Conversely, the songs of mutants with defects in the central complex differed from those of wild-type males. Firstly, there was degradation of the stability of the pulse oscillator and interpulse intervals were very variable. In addition, pulses were often significantly longer and appeared multicyclic, as in the well-known cacophony mutant, while the mean train duration was significantly shorter. Males of the line cexKS181 usually courted very intensely, though abnormal sounds were generally emitted. Mutants cexKS181 and ccbKS127 were significantly less successful in courtship than wild-type flies. These data show that the central complex appears to play a very important role in controlling song, while the mushroom bodies are not related to this function.

Animal Communication↗

Temperature activation of courtship behavior in the male red-sided garter snake (Thamnophis sirtalis parietalis): role of the anterior hypothalamus-preoptic area.

Adult male red-sided garter snakes (Thamnophis sirtalis parietalis) received radio-frequency lesions prior to 17 weeks of low-temperature hibernation. Animals found to have bilateral lesions of the anterior hypothalamus-preoptic area (AH-POA) failed to exhibit courtship behavior on emergence from hibernation. Those individuals in which the major portion of the destruction was centered in the anterior POA also exhibited deficits in thermoregulatory behavior. Animals that received unilateral lesions of the AH-POA initiated courtship behavior after controls and had an abbreviated period of courtship; these animals exhibited normal responses to thermal stimuli. Male snakes with lesions outside the AH-POA courted normally and demonstrated no differences in thermoregulatory behavior compared with the surgical controls. These results indicate that in male red-sided garter snakes, an intact AH-POA is critical for the integration of thermal stimuli that activate seasonal courtship behavior.

Animals↗

Genetics of divergence in male wing pigmentation and courtship behavior between Drosophila elegans and D. gunungcola.

Many sex-specific traits involved in mating consist of functionally coordinated morphologies and behaviors. How the components of these complex traits evolve and become coordinated during evolution is unknown. In order to understand how such trait complexes evolve and diversify, we must decipher the genetic underpinnings of their components. In this study, we begin to elucidate the genetic architecture underlying differences in functionally related male pigmentation and behavior between two Asian Drosophila melanogaster group species, D. elegans and D. gunungcola. D. elegans possesses a male-specific wing melanin spot and a stereotypical wing display element in male courtship, whereas D. gunungcola lacks both of these traits. Using reciprocal F1 male hybrids, we demonstrate that the X-chromosome contains a major locus or loci required for wing spot formation and that autosomal loci largely determine the male courtship display. Using phenotypic and genetic analysis of backcross progeny, we further demonstrate that both the wing spot and courtship differences between the two species are polygenic and both depend at least in small part on genetic factors on both the X and the autosomes. Finally, we find that male wing spot size and courtship wing display are highly correlated in backcross progeny, suggesting that linkage or pleiotropy may have been involved in their coordinated evolution.

Animals↗

The nonverbal basis of attraction: flirtation, courtship, and seduction.

According to a familiar phrase, the "language" of love is universal. Recent ethological studies of nonlinguistic communication in courtship using facial expression, gesture, posture, distance, paralanguage, and gaze have begun to establish that a universal, culture-free, nonverbal sign system may exist (Eibl-Eibesfeldt, 1975), which is available to all persons for negotiating sexual relationships. The nonverbal mode, more powerful than the verbal for expressing such fundamental contingencies in social relationships as liking, disliking, superiority, timidity, fear and so on, appears to be rooted firmly in man's zoological heritage (Bateson, 1966, 1968). Paralleling a vertebrate-wide plan, human courtship expressivity often relies on nonverbal signs of submissiveness (meekness, harmlessness) and affiliation (willingness to form a social bond). Adoption of a submissive-affiliative social pose enables a person to convey an engaging, nonthreatening image that tends to attract potential mates. This report explores several conspicuous nonlinguistic cues that appear to be used widely in contexts of flirtation, courtship, and seduction. The expressive units are discussed from the standpoint of their occurence in five phases of courtship, and are illustrated by four cases.

Animals↗

Bottleneck effects on genetic variance for courtship repertoire.

Bottleneck effects on evolutionary potential in mating behavior were addressed through assays of additive genetic variances and resulting phenotypic responses to drift in the courtship repertoires of six two-pair founder-flush lines and two control populations of the housefly. A simulation addressed the complication that an estimate of the genetic variance for a courtship trait (e.g., male performance vigor or the female requirement for copulation) must involve assays against the background behavior of the mating partners. The additive "environmental" effect of the mating partner's phenotype simply dilutes the net parent-offspring covariance for a trait. However, it there is an interaction with this "environmental" component, negative parent-offspring covariances can result under conditions of high incompatibility between the population's distributions for male performance and female choice requirements, despite high levels of genetic variance. All six bottlenecked lines exhibited significant differentiation from the controls in at least one measure of the parent-offspring covariance for male performance or female choice (estimated by 50 parent-son and 50 parent-daughter covariances for 10 courtship traits per line) which translated to significant phenotypic drift. However, the average effect across traits or across lines did not yield a significant net increase in genetic variance due to bottlenecks. Concerted phenotypic differentiation due to the founder-flush event provided indirect evidence of directional dominance in a subset of traits. Furthermore, indirect evidence of genotype-environment interactions (potentially producing genotype-genotype effects) was found in the negative parent-offspring covariances predicted by the male-female interaction simulation and by the association of the magnitude of phenotypic drift with the absolute value of the parent-offspring covariance. Hence, nonadditive genetic effects on mating behavior may be important in structuring genetic variance for courtship, although most of the increases in genetic variance would be expected to reflect inbreeding depression with relatively rare situations representing the facilitation of speciation by bottlenecks.

Analysis of Variance↗

Attracting female attention: the evolution of dimorphic courtship displays in the jumping spider Maevia inclemens (Araneae: Salticidae).

Males of the dimorphic jumping spider (Maevia inclemens) differ in both their morphologies and courtship displays (i.e. phase I). The tufted morph stilts and waves from an average distance of 9 cm from a female, whereas the grey morph crouches and sidles from an average distance of 3 cm from a female. The objective of this study was to determine the significance of the different courtship displays using computeranimated versions of males performing phase I courtship in a Y-maze where first male movement and then the distance of the stimulus was controlled. Females selected the first male that they orientated to at the close distance of 4 cm and at the far distance of 16cm. However, there was no preference for the first male at the intermediate distance of 8 cm or the furthest distance of 24 cm. In addition, males have morph-specific advantages regarding the time it takes to attract female attention. Grey males attracted female attention in less time than tufted males at 4 and 8 cm. However, tufted males attracted female attention in less time than grey males at 16 cm. These results suggest a mechanism for the evolution of two different courtship displays whereby each morph has an advantage at different distances from the female.

Animals↗