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The role of cuticular pheromones in courtship conditioning of Drosophila males.

Courtship conditioning is an associative learning paradigm in Drosophila melanogaster, wherein male courtship behavior is modified by experience with unreceptive, previously mated females. While the training experience with mated females involves multiple sensory and behavioral interactions, we hypothesized that female cuticular hydrocarbons function as a specific chemosensory conditioned stimulus in this learning paradigm. The effects of training with mated females were determined in courtship tests with either wild-type virgin females as courtship targets, or with target flies of different genotypes that express distinct cuticular hydrocarbon (CH) profiles. Results of tests with female targets that lacked the normal CH profile, and with male targets that expressed typically female CH profiles, indicated that components of this CH profile are both necessary and sufficient cues to elicit the effects of conditioning. Results with additional targets indicated that the female-specific 7,11-dienes, which induce naive males to court, are not essential components of the conditioned stimulus. Rather, the learned response was significantly correlated with the levels of 9-pentacosene (9-P), a compound found in both males and females of many Drosophila strains and species. Adding 9-P to target flies showed that it stimulates courting males to attempt to copulate, and confirmed its role as a component of the conditioned stimulus by demonstrating dose-dependent increases in the expression of the learned response. Thus, 9-P can contribute significantly to the conditioned suppression of male courtship toward targets that express this pheromone.

Alkenes↗

Characterization of courtship sounds of species of the subgroup fasciola (Diptera, Drosophilidae, Drosophila repleta group): interspecific and interpopulational analyses.

The aim of this work was to characterize the male courtship song pattern of various species of the fasciola subgroup and to determine the level of variation both within and among species. The parameters analyzed were intrapulse interval (PI), interpulse interval (IPI), and intrapulse frequency (IF). Six different species were analyzed: D. coroica (three populations), D. ellisoni, D. fascioloides, D. moju, D. onca, and D. rosinae (one population each). There were significant differences among the six species for these three courtship song parameters. The IPI was the most variable parameter among these species, suggesting that this parameter is important for female discrimination. Four different hypotheses could explain this variation: 1. different selection pressures with absence of flow gene; 2. intraspecific sexual selection; 3. sympatric effects on song evolution; and 4. genetic drift. The PI was the only parameter that was significantly different among the three population of D. coroica. Low variability among populations within the same species was already observed for other subgroups and could be explained by the following hypotheses: strong selection acting on the song parameters, gene flow, or recent colonization from a common source. Additional studies of the courtship song of other species of the fasciola subgroup, as well as for other subgroups of the repleta group, and studies, using molecular makers, that focus on the genetic basis of the differences among these species in courtship song would allow us to evaluate the association of courtship song and sexual isolation in these species, and would also help us to understand the evolution of these behavioural differences.

Animals↗

Initial courtship behavior and stalking: how should we draw the line?

As noted by Schaum and Parrish (1995), stalking blurs the boundaries between normal courtship and obsessive behavior. Consequently, stalking proves an elusive phenomenon to define and to study. Where does courtship end and stalking begin? To address this question, 197 women and 44 men from the University of Pittsburgh who had loved someone who did not love them in return were surveyed about their feelings and actions in response to this rejection. Factor analyses revealed six groupings of behaviors in response to unrequited courtship: approach, surveillance, intimidation, harming oneself, verbal abuse/mild physical harm, and extreme physical harm. Approach behavior was reported to be used more often by men, but there were no significant differences between men and women in the self-reported prevalence of other types of courtship behaviors. Correlations reveal that feelings of anger and depression were the most common predictors of violent behavior for both men and women. Pursuer-perceptions of what behaviors connoted "going too far" in pursuit of a relationship proved unreliable. However, when pursuers were asked whether their love interest was afraid of them, fear was a reaction perceived in response to intimidation. Although it is likely that pursuer perceptions of where to draw the line would differ from the view of the love object, these results suggest that engaging in intimidation would be an appropriate place to draw the line between courtship and stalking.

Adolescent↗

Conditioning deficits of CaM-kinase transgenic Drosophila melanogaster in a new excitatory courtship assay.

Courtship suppression is an associative conditioning procedure in Drosophila melanogaster that is ethologically based and capable of being tested on individual flies. We have expanded the range of the courtship conditioning by developing an excitatory procedure in which male flies learn to associate a novel odor with the courtship stimulating cues of virgin females. Wild-type males normally court other mature males very little, but following training, the odor alone is able to elicit increased levels of courtship towards an object male. Flies expressing an inhibitor of calcium/calmodulin-dependent protein kinase II (CaMKII) were previously shown to have no retention one hour after training in the courtship suppression task, as manifested in their persistent courting of a virgin female. A possible trivial explanation for this response is that the CaMKII-inhibited fly strains (ala1 and ala2) were merely hyperactive courters. The poor performance of these mutants in the new excitatory conditioning procedure confirms that their conditioning deficit results from a disruption of an associative mechanism per se.

Animals↗

[The fan-shaped and ellipsoid bodies of the brain central complex are involved in the control of courtship behavior and communicative sound production in Drosophila melanogaster males].

To elucidate the role of the fan-shaped and ellipsoid bodies (FB and EB) of the central complex of Drosophila melanogaster brain in the control of male courtship behavior and singing, we analyzed characteristics of the courtship behavior and parameters of the communicative sound signals accompanying it in wild type flies and in flies from 5 mutant strains with various anatomical defects in FB and EB. The following strains of flies were used for experiments: Canton S (wild type, the control), eboKS263 with defects only in EB and ebo1041, ceb849, ceb892 and cbdKS96 with both the FB and EB damaged in different manner. The data obtained suggest that the FB and EB are involved: 1) in maintenance of a high courtship activity level, 2) in the control of accuracy of male following movements when courting a female, 3) in the control of the form and stability of sound elements in courtship songs, 4) in the control of the rhythmic structure of courtship songs determined by the stability of the pacemakers, and 5) in setting up a correspondence between the current behavior and the external situation.

Animal Communication↗

Enhanced fos expression in the zebra finch (Taeniopygia guttata) brain following first courtship.

Young zebra finch males that court a female for the first time develop a stable preference for the females of that species. On the neuronal level, consolidation of the imprinted information takes place. Here we demonstrate that first courtship or being chased around in the cage leads to enhanced fos expression in forebrain areas implicated in learning and imprinting in zebra finch males compared with birds reared in isolation or in the aviary. Two of the forebrain areas highly active during first courtship (as demonstrated by the 14C-2-deoxyglucose technique), the imprinting locus latral neo/hyperstriatum ventrale (LNH) and the secondary visual area hyperstriatum accessorium/dorsale (HAD), demonstrate enhanced fos expression. Two other imprinting-related areas, the medial neo/hyperstriatum ventrale (MNH) and archistriatum/neostriatum caudale (ANC), do show c-fos induction; however, the areas are not congruous with those demarcated by the 2-DG autoradiographic studies. Additional telencephalic areas include the olfactory lobe, the information storage site lobus parolfactorius (LPO), the memory site hippocampus, the auditory caudomedial neostriatum implicated in the strength of song learning, and the caudolateral neostriatum, which is comparable to the mammalian prefrontal cortex. In addition, c-fos is induced by first courtship and chasing in neurosecretory cell groups of the preoptic area and hypothalamus associated with the repertoire of sexual behavior and stress or enhanced arousal. Enhanced fos expression is also observed in brainstem sources of specific (noradrenergic, catecholaminergic) and nonspecific (reticular formation) activating pathways with inputs to higher brain areas implicated in the imprinting process. Birds reared in isolation or alternatively in the aviary with social and sexual contact to conspecifics showed attenuated or no fos expression in most of the above-mentioned areas. First courtship and chasing both lead to enhanced uptake of 2-DG in the four imprinting areas, as well as subsequent changes in spine density-an anatomical manifestation of the imprinting process. fos expression in the imprinting and other telencephalic, preoptic, hypothalamic, and mesencephalic brain regions indicates processing of stimuli originating from exposure (like chasing) and the analysis of stimuli in a behaviorally relevant, sexually explicit context (like first courtship). c-fos induction in these brain areas indicates its involvement in the triggering of neural changes that accompany the learning process of imprinting, leading eventually to alterations in dendritic spine density in the zebra finch.

Animals↗

A test of how well the repeatability of courtship predicts its heritability.

We compared repeatability and heritability estimates for courtship traits in the housefly Musca domestica L., to test the expectation that the repeatability of a behaviour is an upper-bound estimate of its heritability. The repeatabilities across and within mating partners for two courtship traits, male wing buzzing (Buzz) and female wing extension (Wing-out), were compared to heritabilities calculated from the divergence among subpopulations due to founder-flush events or artificial selection. The male's Buzz showed highly significant repeatabilities within and across female partners, but the values ranged widely (0.3-0.5). The intensity of Buzz decreased significantly with successive courtship attempts to the same female, apparently as a male strategy to optimize his efforts to gain copulation by modulating behaviour for specific females. Repeatabilities for female Wing-out within and across male partners were not significant, but this trait yielded significant heritabilities in two assessments (based on successful courtships). We attribute this contradiction to biased sampling because only the subset of reluctant females met the necessary restriction to refuse copulation throughout four successive courtships. These results demonstrate that repeatabilities are labile within individuals and may actually underestimate heritabilities when there is a genetic predisposition not to repeat performance. We concur with related studies that repeatabilities can identify evolutionary potential, but we advise caution in interpreting low values or comparing their absolute values.Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

Geographical variation in male courtship behaviour and sexual isolation in wolf spiders of the genus Schizocosa.

We surveyed 12 populations of the wolf spider Schizocosa crassipes (Walckenaer) and S. nr. crassipes in Tennessee, Mississippi, Louisiana and Florida, in the U.S. to determine the extent of variation in male courtship behaviour when observed in standard laboratory conditions. We observed variation in both the frequency of occurrence and the sequence of the four principal male courtship behaviours and, when compared statistically, most of these behaviours differed between populations. Although there was no clinal pattern in the frequency of courtship behaviour, we observed geographically consistent patterns in the sequence in which male courtship behaviours are displayed. We conducted two subsequent studies to determine whether sexual isolation among selected populations existed. In the first study, we performed male-female reciprocal crosses of brush-legged spiders (S. nr. crassipes) between two populations from the Yazoo-Mississippi Delta physiographical region and one population from the Loess Bluff region of Mississippi and measured mating success, number of matings that produced egg sacs, and the number of egg sacs that hatched young. The results of that experiment suggest that sexual isolation occurs among populations in different physiographical regions but not between populations within regions of the state. In a second crossing study, we paired the brush-legged spiders from one of the Mississippi physiographical regions (Yazoo-Mississippi Delta) with Schizocosa ocreata (Hentz) from Ohio. The results of that cross suggest that the Mississippi Delta population is more similar to the northern S. ocreata populations than to S. crassipes. We discuss the results of the crossing experiments in terms of what was revealed about geographical patterns in male courtship behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

Behavioural isolation between two closely related Hawaiian Drosophila species: the role of courtship.

Behavioural isolation plays a critical role in several recent models of speciation. A detailed understanding of the process of speciation requires analysis of taxonomic groups that have not completed reproductive isolation. We studied D. silvestris and D. heteroneura because they are still in the process of divergence: behavioural isolation between them is incomplete, and neither postzygotic nor ecological isolation has been detected. Behavioural isolation is due to the failure of courtships between male D. silvestris and female D. heteroneura: there is no postzygotic isolation from either parental species. The F1 hybrids are as successful in courtship with parental individuals as same-species pairs, which suggests that the hybrids resemble male D. heteroneura or female D. silvestris in some behaviour patterns that are crucial to mating success. We searched for this crucial resemblance by examining courtship between F1 hybrids and the parental adults. We found that successful F1 males are somewhat more similar than unsuccessful F1 males to D. heteroneura males, but nevertheless they were intermediate between males of the two species. We also found that in both species the presence of female wings is necessary for courtship to proceed to copulation. These results reinforce an earlier report that behavioural isolation between these species is largely attributable to the decision as to whether to court at all, rather than to the details of courtship. Copyright 2000 The Association for the Study of Animal Behaviour.

Journal Article↗

Involvement of endogenous prolactin in the expression of courtship behavior in the newt, Cynops pyrrhogaster.

In order to ascertain the involvement of endogenous prolactin (PRL) in the expression of courtship behavior by the male newt, Cynops pyrrhogaster, antiserum against homologous PRL was administered to male newts, which had been exhibiting courtship behavior in the field and had been taken to the laboratory. The effect of the antiserum on the preference of terrestrial over aquatic habitats also was investigated. Within 24 hr of the first injection of antiserum, both incidence and frequency of courtship behavior declined markedly compared with those in the preimmune serum-injected newts. The anti-PRL serum-induced decline became more conspicuous thereafter. These animals came to prefer a terrestrial habitat. Cessation of antiserum injection and administration of PRL restored the expression of courtship behavior to a certain extent and altered their habitat preference from terrestrial to aquatic. The results strongly suggest the involvement of endogenous PRL in the expression of courtship behavior as well as in the preference of aquatic habitat.

Animals↗

Seasonal changes in courtship song and the medial preoptic area in male European starlings (Sturnus vulgaris).

In male starlings (Sturnus vulgaris) courtship song plays a critical role in mate attraction. During the breeding season courtship song occurs prior to copulation and appears to reflect male sexual arousal. Outside the breeding season starlings sing, but song appears unrelated to reproduction. The aromatization of testosterone (T), likely within the medial preoptic nucleus (POM), is critical for the expression of male sexual arousal. The present study was performed to determine whether seasonal changes in the POM might relate to seasonal changes in courtship singing behavior in male starlings. T concentrations, the volume of the POM, and aromatase within the POM were examined both during and outside of the breeding season in male starlings. Song was also recorded at these times both with and without a female present. The POM was largest and contained dense aromatase immunostaining only during the spring breeding season, when T concentrations were highest and males responded to a female with an increase in courtship song. Outside the breeding season the volume of the POM was small, T concentrations were low, and males displayed no changes in song expression in response to female conspecifics. Song bout length was positively related to POM volume, and males sang longer songs in spring. Only males with nestboxes in spring responded to a female, and the POM tended to be larger in these males, suggesting that nestbox possession might influence neuroplasticity within the POM. Overall, the findings suggest that T-dependent plasticity and aromatase activity within the POM might regulate courtship singing in a wild songbird.

Animals↗

Courtship latency in male Drosophila melanogaster.

Male Drosophila melanogaster differ in the age at which they reach sexual maturity following eclosion from the pupa. Courtship latency, which is the time taken by a male to initiate courtship of a conspecific female, is related to age. Young males take significantly longer than older males to begin courtship. The probability that a male will initiate courship is influenced by the physiological state of the female. Males of different genotypes readily court mature (3-day-old) virgin females, but they differ significantly in their reaction to immature (12-hr-old) and fertilized females. Genes located on the third chromosome largely control male courtship latency, but responses to immature and fertilized females have different genetic bases, suggesting that the relevant stimulus inputs governing these responses also differ. The adaptive significance of courtship directed toward immature or fertilized females, which rarely mate, probably depends on the average level of sexual responsiveness of potentially receptive mature virgin females in a given population.

Age Factors↗

Behavioral analysis of Drosophila mutants displaying abnormal male courtship.

We describe six recessive autosomal male sterile mutations in Drosophila, generated by mobilization of single P-elements, exhibiting abnormal male courtship behavior. Detailed analysis of courtship behavior elicited by virgin wild type females indicated that five of the six mutants are affected in the early steps of courtship. The sixth mutant is blocked at the step of attempted copulation which occurs later in the courtship sequence. All of the mutants have normal olfactory responses and normal locomotor activity. No defect in the visual modality has been observed for the five mutants affected in the initiation of courtship. The mutant blocked at attempted copulation lacks the 'on' and 'off' transients, but this appears to be due to genetic background rather than the mutation itself. Abnormal spermatogenesis was observed in five of the mutants. Spermatogenic defects vary and include lesions in the proliferation of the germline, in meiosis, and in the differentiation and maturation of the spermatids into motile sperm.

Animals↗

Experimental manipulation of sexual selection and the evolution of courtship song in Drosophila pseudoobscura.

Courtship song serves as a sexual signal and may contribute to reproductive isolation between closely related species. Using lines of Drosophila pseudoobscura experimentally selected under different sexual selection regimes, we tested whether increased promiscuity and enforced monogamy led to evolutionary changes in courtship song elements. In D. pseudoobscura, males produce both a low and high rate repetition song. We found that both song types diverged after selection and that the direction of changes was consistent with ordered hypotheses of predicted directions of change under the different mating system structures. In particular, latency to the initiation of song and duration of the interpulse interval (IPI) decreased in highly promiscuous lines and increased in monogamous lines. These results suggest that courtship song may rapidly evolve under different mating system structures, representing either functional evolution of more stimulatory song under conditions of strong sexual selection, or might result from increased courtship vigor, or represent correlated evolution. Some speciation theory predicts that increased sexual selection should result in increased variance of traits between allopatric populations, facilitating reproductive isolation. We also found that courtship song elements were not equally variable between replicate lines.

Animals↗

Mutations and natural genetic variation in the courtship song of Drosophila.

All Drosophila species have a courtship repertoire by which a male stimulates a female to mate with him. In many species, males vibrate their wings to produce courtship song, an element of courtship that plays an important role in female choice. Each species has a unique courtship song, with the major differences among species songs being in timing and/or structure. Analysis of genetic mutations has revealed 17 genes that affect courtship song in Drosophila melanogaster. Most of the genes were first identified as affecting another trait and were subsequently shown to affect song. Quantitative genetic studies have demonstrated a polygenetic additive genetic architecture for many song traits. Few candidate genes, identified through the classical genetic approach, coincide with the regions implicated as affecting natural variation. With many new tools in genetic analysis and the multiple Drosophila genome projects currently underway, the ability to relate mutational and quantitative analyses will improve.

Animals↗

Central DSP-4 treatment decreases norepinephrine levels and courtship behavior in male zebra finches.

In zebra finches, gonadal steroids activate male courtship, including singing, and also strongly modulate norepinephrine (NE) levels and turnover in brain areas regulating courtship behavior. In a previous study, systemic administration of DSP-4 caused significant decreases in courtship singing. These behavioral decrements were correlated with the degree of NE depletion in several vocal control nuclei. In the present study, we attempted to further decrease brain NE levels while minimizing systemic effects by infusing DSP-4 directly into the third ventricle. DSP-4 treatment significantly reduced NE levels in three of six vocal control nuclei and both hypothalamic nuclei sampled without significantly altering dopamine or serotonin levels in any areas. DSP-4-treated males took longer to begin singing and performed fewer song bouts and courtship displays. Interestingly, behavioral deficits were limited to courtship song displays, other behavior patterns, including female-directed behaviors like approach and follow, were unaffected by DSP-4 treatment. DSP-4 treatment appeared to affect singing behavior by causing deficits in initial attentiveness to females and initiation of singing rather than by affecting song structure.

Animals↗

Courtship behavior of male white perch, Morone americana: evidence for control by androgens.

Courtship behaviors are androgen-dependent in many vertebrates and castration often decreases courtship. We examined the effectiveness of castration in reducing courtship behaviors and 11-ketotestosterone (KT) and testosterone (T) in restoring them in male white perch. Castrates were given implants containing KT, T or no hormone. Sham-operated males received implants without hormone. Three weeks later, males were exposed to an ovulated female for 1 h and two courtship behaviors were quantified. Attending behavior involves close and continuous following of a female with occasional contact. Circling involves rapid transits around the female in a circular pattern or back and forth in front of her. In plasma samples taken immediately after observations, KT and T were below detectable levels in castrated males but at high physiological levels in males implanted with KT or T. Castrated males given KT attended females more than castrated males given T implants or implants containing no hormone, but not more than sham-operated males. Circling was eliminated by castration but restored by implantation with T or 11-KT to values exhibited by sham-operated males. This is one of the few demonstrations that KT can regulate courtship behavior in a non-territorial and economically important fish species.

Aggression↗

Conditional disruption of synaptic transmission induces male-male courtship behavior in Drosophila.

It is reported here that male-male courtship behavior is evoked instantaneously in the fruit fly Drosophila by conditional disruption of synaptic transmission. A temperature-sensitive allele of the Drosophila dynamin gene shibire (shi(ts1)) was expressed by using the GAL4/UAS system to disrupt synaptic transmission from GAL4-positive neurons in a temperature-dependent manner. An enhancer-trap GAL4 line C309 directing shi(ts1) expression in central and peripheral neurons (C309/UAS-shi(ts1)) initiated stereotypical precopulatory behavior toward other mature males immediately after a temperature shift from the permissive to restrictive temperature. At the restrictive temperature, C309/UAS-shi(ts1) males formed "courtship chains" and exhibited abnormally high levels of head-to-head interactions. The temperature-induced male-male courtship is attributable not to an increase in sexual attractiveness but to an increase in sexual activity of C309/UAS-shi(ts1) males. Interestingly, the temperature-induced increase in sexual activity is specific toward male partners, because C309/UAS-shi(ts1) males courted receptive virgin females less vigorously and copulated less efficiently after shifted to the restrictive temperature. Among the GAL4-positive neurons in C309, conditional disruption of certain cholinergic neurons but not the mushroom body intrinsic neurons plays a critical role in the induction of male-male courtship. These neurons may be involved in inhibitory systems that normally suppress aberrant male-male courtship. The presented strategy that can induce behavioral abnormalities by disrupting synaptic transmission in an acute and noninvasive manner will allow further exploration as to how distinct neuronal groups control sexual orientation and other aspects of reproductive behavior in Drosophila.

Alleles↗