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Distribution of dopamine-like immunoreactivity suggests a role for dopamine in the courtship display behavior of the blue crab, Callinectes sapidus.

Injection of dopamine initiates a posture in the blue crab, Callinectes sapidus, identical to courtship display behavior of the male crab. The threshold for proctolin-induced rhythmic components of courtship display is lowered in preparations when dopamine is co-applied with proctolin. To elucidate the anatomical substrate of this behavior, immunocytochemistry was used to map dopamine-immunoreactive neurons. Courtship display is sex-specific, and dependent on the hormonal, developmental, and seasonal state of the animal. We compared the distribution of dopamine-like immunoreactivity between adults and juveniles of both sexes across seasons, with hormonal alteration, and with the distribution of proctolin-like immunoreactivity. Dopamine-like immunoreactivity was found throughout the nervous system in identical patterns between the sexes and hormonal states. Differences were found between juveniles and adults that are not obviously correlated with the development of behavior. Two areas of staining were of interest: neurites that longitudinally traverse and terminate in the posterior ventral nerve cord, and a neuron in the esophageal ganglion that has projections to the pericardial organ. The results do not suggest that proctolin-like and dopamine-like immunoreactivity co-localize, but in the subesophageal ganglion there was a region of close proximity.

Animals↗

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

Testosterone (T) was administered intracranially to intact adult male Canadian red-sided garter snakes (Thamnophis sirtalis parietalis) in the fall and in the summer. Implants in the anterior hypothalamus-preoptic area (AH-POA) as well as other regions of the brain, including the medial and basal hypothalamus, thalamus, medial cortex, and third ventricle, failed to elicit courtship behavior. The spermatogenic stage was more advanced, and the circulating level of androgens was significantly higher, in animals that received implants of T in the AH-POA. These findings suggest the sex steroid-concentrating sites in the AH-POA of the garter snake are involved in feedback regulation of pituitary gonadotropin secretion and not in the control of courtship behavior. Hematocrit was found to be higher in animals that received implants of T, regardless of location, a response that may be related to changes in blood chemistry prior to hibernation. These findings support previous research indicating that in the adult male red-sided garter snake, a species exhibiting a dissociated reproductive tactic, courtship behavior is independent of testicular androgens.

Androgens↗

Breeding experience modulating androgen dependent courtship behavior in male ring doves (Streptopelia risoria).

Courtship behavior of reproductively experienced males was compared with that of age-matched inexperienced males before and after castration and during treatments with hormones (testosterone propionate and estradiol benzoate). Inexperienced males displayed significantly lower levels of nest-oriented courtship, the nest-cooing (nest-soliciting) and wingflippings; however, agonistic courtship, the bow-cooing, was not affected by breeding experience. The lower level of nest-cooing display in the inexperienced males persisted even when their circulating level of androgen as measured by RIA was no different from that of experienced males. Estrogen treatments, however, produced no group difference in levels of nest-coo display. These results were discussed in the context of hormone-behavior relationships.

Animals↗

Social condition affects the courtship behavior of male ring doves with posterior medial hypothalamic lesions.

Following bilateral lesions to the posterior medial hypothalamus (homologue of the mammalian ventromedial nucleus), adult male ring doves regain full courtship behavior and the ability to stimulate female egg-laying when housed continuously with females. Males with PMH lesions housed singly and only tested periodically with females continue to show deficits in courtship. These findings suggest that the social environment present in adulthood itself can directly influence recovery from brain lesions. They also demonstrate the importance of PMH in the mediation of male ring dove courtship behavior.

Animals↗

Prolactin acts centrally to enhance newt courtship behavior.

The effects of intracerebroventricular (ICV) and intraperitoneal (IP) injections of ovine prolactin (PRL), antiserum against newt PRL, and antibody against the newt PRL receptor on the expression of courtship behavior of male newts, Cynops pyrrhogaster, were studied to see whether PRL acts centrally or peripherally to induce the behavior. Injections of PRL by either route into gonadotropin-primed males enhanced the expression of the behavior dose-dependently. The minimum effective amount of ovine PRL administered intracerebroventricularly was 0.1 microg, whereas it was 100 microg when injected intraperitoneally. ICV injection of antiserum against newt PRL blocked the spontaneously occurring male courtship behavior when the anti-newt PRL serum was given either intracerebroventricularly or intraperitoneally. The minimum effective dose of the antiserum administered intracerebroventricularly was 0.05 microl, whereas it was 20 microl when injected intraperitoneally. Neither ICV nor IP injection of preimmune serum affected the expression of the behavior. Furthermore, ICV, but not IP, administration of 0.3 microg of anti-newt PRL receptor antibody, purified from antiserum against newt PRL receptor by use of an antigen-conjugated affinity column, blocked the spontaneously occurring courtship behavior in sexually developed males. Neither ICV nor IP injection of the same amount of normal rabbit IgG affected the expression of the behavior. The results strongly suggest that endogenous PRL enhances the behavior by acting centrally through the PRL receptors localized in the brain area.

Animals↗

Gonadotropin antagonist modulates courtship behavior in male red-sided garter snakes, Thamnophis sirtalis parietalis.

Behavioral studies were used to investigate the central effects of chicken-I GnRH, chicken-II GnRH, and D-Phe2,6,Pro3-GnRH, a GnRH antagonist, on the courtship behavior of male red-sided garter snakes, Thamnophis sirtalis parietalis. Intracerebroventricular (i.c.v.) injections of chicken-I or chicken-II GnRH had no effect on time spent courting or latency to court when experimental males were exposed to unmated females, or when experimental males were exposed to the female sex attractiveness pheromone. I.c.v. injections of D-Phe2,6,Pro3-GnRH caused a significant decrease in latency to court when experimental males were exposed to unmated females. When males injected with D-Phe2,6,Pro3-GnRH were exposed to the female sex attractiveness pheromone, it caused a significant increase in time spent courting compared to that in saline-injected controls. D-Phe2,6,Pro3-GnRH was not able to initiate courtship behavior during the nonbreeding season, indicating that courtship behavior is dependent on the interaction of multiple components. This study does demonstrate that a hormone or neuropeptide can modulate sexual behavior in garter snakes.

Animal Communication↗

The voyeurs' guide to Drosophila melanogaster courtship.

Drosophila melanogaster has a long and distinguished history as a model organism in studies of sex-specific behaviour. Courtship is relevant to a wide variety of areas of biological research, from investigations into the evolution of sex-specific behaviours, to studies of the molecular mechanisms underlying the perception and processing of sex-specific information, and the identification of genes that enable a fly to behave in a sex-specific fashion. To address any of these issues it is essential that courtship behaviour is investigated in a robust, reproducible and reliable manner. In this review I consider many of the problems that one might encounter in a study of Drosophila courtship, and how such issues may be addressed.

Journal Article↗

Neuroendocrine suppression of female courtship in a wild passerine: corticotropin-releasing factor and endogenous opioids.

During emergencies in their natural environments, vertebrates initiate coping mechanisms that redirect behavior away from nonessential activities and towards survival. Reproductive behaviors are suppressed. Evidence from field studies on passerine birds shows that this inhibition may not depend on the suppression of gonadal sex steroids, since during the breeding season they remain elevated despite activation of the stress response. We hypothesize that an alternate, central mechanism mediates the inhibition of reproductive behavior during stress in passerines. In this study, we tested the intracerebroventricular effects of endogenous opioids and corticotropin-releasing factor (CRF), neuropeptides implicated in the stress response, on courtship behavior in wild-caught female white-crowned sparrows. Beta-endorphin (beta-EN) significantly inhibited copulation solicitation, an estrogen-dependent courtship display, 30 min after treatment. Naloxone, an opioid antagonist, enhanced the behavior. CRF caused a suppression of solicitation that was reversible by pretreatment with naloxone, suggesting an intermediary role for endogenous opioids in CRF-induced suppression of courtship. The effects of beta-EN and CRF on solicitation appear to be independent of any general effects on locomotor activity. These results support our hypothesis that stress neuropeptides orchestrate coping behaviors in wild birds.

Adaptation, Psychological↗

Misexpression of the white (w) gene triggers male-male courtship in Drosophila.

We report here that the general ectopic expression of a tryptophan/guanine transmembrane transporter gene, white (w), induces male-male courtship in Drosophila. Activation of a hsp-70/miniwhite (mini-w) transgene in mature males results in a marked change in their sexual behavior such that they begin to vigorously court other mature males. In transformant populations containing equal numbers of both sexes, most males participate, thus forming male-male courtship chains, circles, and lariats. Mutations that ablate the w transgene function also abolish this inducible behavior. Female sexual behavior does not appear to be altered by ectopic w expression. By contrast, when exposed to an active homosexual courtship environment, non-transformant males alter their behavior and actively participate in the male-male chaining. These findings demonstrate that, in Drosophila, both genetic and environmental factors play a role in male sexual behavior.

ATP-Binding Cassette Transporters↗

Display lateralisation in the courtship behaviour of the alpine newt (Triturus alpestris).

Among urodelan amphibians, European newts of the genus Triturus have evolved a complex sequence of displays to the effect of stimulating the female to actively pick up the spermatophore deposited by the male. Courtship is varied, with plenty of potential for asymmetric uses of tails and of asymmetric turns. The analysis of the temporal structure of the behavioural sequences showed that both males and females significantly utilise the same side during a single courtship sequence. Moreover, if an individual prefers to exhibit one side during its lateral displays, then the duration of its displays on that side is longer than the duration of its displays on the other side. The only significantly contralateral display is creep-on. Less than 50% of the alpine newts show a significant left- or right-side preference and no lateral bias at population level was found. At the population level, male Triturus vulgaris showed a marked bias towards left turning after spermatophore deposition, in spite of the existence of both "left-turning" and "right-turning" individuals in the population (Green, 1997). The presence of population biases may be adaptive if it can influence the sensory processes and the motor responses involved in social interactions. In fact, Triturus vulgaris is characterised by a more complex courtship sequence than T. alpestris and by complex male-male interference mechanisms, but no selective advantages were shown for either left- or right-turning males (Green, 1997).

Journal Article↗

Costly but worthless gifts facilitate courtship.

What are the characteristics of a good courtship gift? We address this question by modelling courtship as a sequential game. This is structured as follows: the male offers a gift to a female; after observing the gift, the female decides whether or not to accept it; she then chooses whether or not to mate with the male. In one version of the game, based on human courtship, the female is uncertain about whether the male intends to stay or desert after mating. In a second version, there is no paternal care but the female is uncertain about the male's quality. The two versions of the game are shown to be mathematically equivalent. We find robust equilibrium solutions in which mating is predominantly facilitated by an "extravagant" gift which is costly to the male but intrinsically worthless to the female. By being costly to the male, the gift acts as a credible signal of his intentions or quality. At the same time, its lack of intrinsic value to the female serves to deter a "gold-digger", who has no intention of mating with the male, from accepting the gift. In this way, an economically inefficient gift enables mutually suitable partners to be matched.

Animal Communication↗

Courtship song recognition in the Drosophila melanogaster complex: heterospecific songs make females receptive in D. melanogaster, but not in D. sechellia.

The courtship song emitted by male wing vibration has been regarded as one of the most important signals in sexual isolation in the species of the Drosophila melanogaster complex. Inter- and intraspecific crosses were observed using males whose wings were removed (mute) or females whose aristae were removed (deaf). Females of D. melanogaster, D. simulans, and D. mauritiana mated with heterospecific males in the song-present condition (cross between normal females and winged males) more often than in the no-song condition (cross between normal females and wingless males or between aristaless females and winged males) or they showed no preference between the two conditions. It is possible that in these females heterospecific courtship songs play a role as if they were conspecific. In contrast, the females of D. sechellia mated with D. melanogaster or D. simulans males in the no-song condition more often than in the song-present condition, suggesting that they reject males with heterospecific song. Female mate recognition depending on the courtship song in D. melanogaster, D. simulans, and D. mauritiana is considered to be relatively broader and that in D. sechellia narrower.

Animals↗

Testosterone regulates androgen receptor immunoreactivity in the copulatory, but not courtship, neuromuscular system in adult male green anoles.

Androgens regulate the expression of male reproductive behaviour in diverse vertebrate species, often acting on androgen receptors (AR) to induce structural or functional changes in the nervous system and periphery. Male green anoles possess two sexually dimorphic neuromuscular systems, one controlling throat fan (dewlap) extension, which occurs during courtship, and the other mediating copulatory organ function. Although androgens are required for behavioural activation in both systems, testosterone has differential effects on the neuromuscular morphology. It increases the size of copulatory muscle fibres during the breeding season, but significant effects on dewlap muscle fibre size and motoneurone soma size in either system have not been detected. Corresponding to the lack of testosterone-induced morphological effects in the courtship system, relatively low levels of AR are expressed in the associated motoneurones. The present experiment had two goals, aiming to determine whether: (i) the other courtship and copulatory neuromuscular tissues express AR and (ii) testosterone and/or seasonal environmental changes regulate AR expression. The percentage of AR+ nuclei was evaluated in both the breeding and nonbreeding seasons in gonadally intact adult males (Experiment 1) and in castrated males treated with either testosterone or vehicle (Experiment 2). AR was extensively expressed in the dewlap and copulatory muscles, and in a high percentage of the copulatory motoneurones, but immunoreactivity did not vary across season. Testosterone increased the percentage of AR+ nuclei in the copulatory muscles of both breeding and nonbreeding males but not in the dewlap muscle or copulatory motoneurones. Finally, the target structures for both systems (cartilages and hemipenes) expressed AR in all animals. Therefore, the effects of testosterone on AR immunoreactivity suggest that up-regulation of the receptors may be important for morphological change. However, because all structures investigated in the present experiment expressed AR, the data also indicate that the receptors are involved with other functions.

Animals↗

Drosophila retained/dead ringer is necessary for neuronal pathfinding, female receptivity and repression of fruitless independent male courtship behaviors.

Mutations in the Drosophila retained/dead ringer (retn) gene lead to female behavioral defects and alter a limited set of neurons in the CNS. retn is implicated as a major repressor of male courtship behavior in the absence of the fruitless (fru) male protein. retn females show fru-independent male-like courtship of males and females, and are highly resistant to courtship by males. Males mutant for retn court with normal parameters, although feminization of retn cells in males induces bisexuality. Alternatively spliced RNAs appear in the larval and pupal CNS, but none shows sex specificity. Post-embryonically, retn RNAs are expressed in a limited set of neurons in the CNS and eyes. Neural defects of retn mutant cells include mushroom body beta-lobe fusion and pathfinding errors by photoreceptor and subesophageal neurons. We posit that some of these retn-expressing cells function to repress a male behavioral pathway activated by fruM.

Alternative Splicing↗

Mapping of the anatomical circuit of CaM kinase-dependent courtship conditioning in Drosophila.

Globally inhibiting CaM kinase activity in Drosophila, using a variety of genetic techniques, disrupts associative memory yet leaves visual and chemosensory perception intact. These studies implicate CaM kinase in the plastic processes underlying learning and memory but do not identify the neural circuitry that specifies the behavior. In this study, we use the GAL4/UAS binary expression system to define areas of the brain that require CaM kinase for modulation of courtship conditioning. The CaM kinase-dependent neurons that determine the response to the mated female during conditioning and those involved in formation and expression of memory were found to be located in distinct areas of the brain. This supports the idea that courtship conditioning results in two independent behavioral modifications: a decrement in courtship during the conditioning period and an associative memory of conditioning. This study has allowed us for the first time to genetically determine the circuit of information flow for a memory process in Drosophila. The map we have generated dissects the behavior into multiple components and will provide tools that allow both molecular and electrophysiological access to this circuit.

Animals↗

Genetics of sexual isolation based on courtship song between two sympatric species: Drosophila ananassae and D. pallidosa.

Sexual isolation has been considered one of the primary causes of speciation and its genetic study has the potential to reveal the genetics of speciation. In Drosophila, the importance of courtship songs in sexual isolation between closely related species has been well investigated, but studies analysing the genetic basis of the difference in the courtship songs associated with sexual isolation are less well documented. Drosophila ananassae and Drosophila pallidosa are useful for studies of sexual isolation, because of their sympatric distribution and absence of postmating isolation. Courtship songs are known to play a crucial role in sexual isolation between these two species, and the female discrimination behaviour against the courting male has been revealed to be controlled by a very narrow region on the second chromosome. In this study we investigated the genetic basis controlling the song differences associated with their sexual isolation, using intact and wingless males with chromosomes substituted between species. The results obtained from F1 hybrid males between these species indicate the dominance of the song characters favoured by D. pallidosa females. In addition, the results obtained from backcross F2 males indicate that chromosome 2 had a major effect on the control of the song characters associated with sexual isolation.

Acoustic Stimulation↗

[NO-synthase inhibitor prevents flight-induced activation of agressiveness and courtship in male].

Previous experience of flying enhances the aggressiveness (Hofmann and Stevenson, 2000, Nature, 403: 613) and accelerates the courtship behaviour (Dyakonova and Krushinski, 2003, DAN, 390: 709-712) of crickets Gryllus bimacultus. We present evidence that these effects may be mediated by activation of nitric oxide synthesis. The effects of flying on fighting and courtship were largely abolished in crickets who received haemocoel injections of a nonspecific NO-synthase inhibitor LNNA. Unlike this, LNNA exerted no significant effects on aggressive and courtship behaviour of nonflown males.

Aggression↗

The courtship of the paying patient.

This article argues for a prohibition on the courtship of the paying patient by individual practitioners, groups, institutions, and corporations. Our society fails to provide universal access to health care. While we await societal resolution, the private provider retains partial responsibility for addressing issues of distributive justice. All private physicians, groups, institutions, and corporations should offer a fair share of underreimbursed, nonemergency care. If economic survival or beneficent economic commitments of a provider are at risk, the provider may explicitly limit underreimbursed services by a rational system of accepting or rejecting nonemergency, indigent patients. The system should be one that, if implemented by all providers, would meet regional societal needs. One might analyze the issue of courtship of the paying patient as a problem of distributive justice. The desire of paying patients to receive personalized care or nonmedical amenities and to have the freedom to buy the best possible medical care, and the economic interests of the providers (the desire of the already affluent provider to seek further gain and growth, the need of a threatened provider to survive, and the dependency of educational institutions on patient payments) all conflict with distributive justice. The marketing interest of providers conflicts with the greater need of the poor to receive information about health, and it conflicts with public need for protection against misleading solicitation. The possibly higher per capita cost of treating the poor, along with possibly lower success rates, create a conflict between cost-effective allocation of limited resources, on the one hand, and egalitarian distributive justice, on the other. The competitive market principle may even protect, rather than defeat, the principle of justice. The author, however, writing from the point of view of the physician, has analyzed the question in terms of a conflict between the economic interest of the physician (perhaps as agent of his group, institution, or corporation) and the trustworthiness of the profession. The courtship of the paying patient represents a failure of the professional virtues of truthfulness, loyalty, and respect for persons.

Attitude to Health↗