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Hormones and midbrain mediation of courtship behavior in the male ring dove (Streptopelia risoria).

The nucleus intercollicularis (ICo) region of the midbrain has been shown to concentrate testosterone and to be involved in the neural control of avian vocal behavior. The present study investigated the possible role of the ICo region in the androgen-dependent courtship behavior of male ring doves, with the use of radio-frequency lesion and intracranial hormone-implant techniques. In the first experiment, bilateral lesions in the ICo region reduced nest-cooing but did not specifically alter other courtship behavior of males tested with stimulus females. No effect on gonadal state could be discerned. In a second experiment, unilateral 30-ga. implants containing crystalline testosterone propionate (TP) or cholesterol (C) were placed in the ICo region or in other parts of midbrain of castrated male doves. The TP implants activated nest-cooing when placed in the ICo region only. Other behaviors were not elicited or not selectively affected by TP implants. Diffusion from the implant was probably not responsible for the elicitation of nest-cooing, since the weights of the peripheral steroid-sensitive tissues of males implanted with TP and C did not differ and since many sites near, but not in, the ICo region were ineffective. These data demonstrate the importance of the ICo region in the expression of nest-cooing, vocal courtship behavior, in the male ring dove. Furthermore, the facilitation of certain vocal courtship behavior by hormonal stimulation is in line with the suggestion that the central gray/ICo/torus semicircularis regions of the vertebrate brain are analogous in their mediation of vocalizations.

Animals↗

Median bundle neurons coordinate behaviours during Drosophila male courtship.

Throughout the animal kingdom the innate nature of basic behaviour routines suggests that the underlying neuronal substrates necessary for their execution are genetically determined and developmentally programmed. Complex innate behaviours require proper timing and ordering of individual component behaviours. In Drosophila melanogaster, analyses of combinations of mutations of the fruitless (fru) gene have shown that male-specific isoforms (Fru(M)) of the Fru transcription factor are necessary for proper execution of all steps of the innate courtship ritual. Here, we eliminate Fru(M) expression in one group of about 60 neurons in the Drosophila central nervous system and observe severely contracted courtship behaviour, including rapid courtship initiation, absence of orienting and tapping, and the simultaneous occurrence of wing vibration, licking and attempted copulation. Our results identify a small group of median bundle neurons, that in wild-type Drosophila appropriately trigger the sequential execution of the component behaviours that constitute the Drosophila courtship ritual.

Animals↗

Courtship behaviour of ostriches (Struthio camelus) towards humans under farming conditions in Britain.

1. The courtship behaviour of adult male and female ostriches was observed in the presence and absence of human beings. 2. Courtship behaviours in both males and females were more prevalent in the presence of humans. 3. Exposure to a human for a short period did not stimulate courtship behaviour in the period immediately after the human had withdrawn. 4. Courtship behaviour towards humans may be important in the reproductive success of ostriches in a farming environment.

Agriculture↗

The dissonance mutant of courtship song in Drosophila melanogaster: isolation, behavior and cytogenetics.

The dissonance mutant of courtship song was induced by chemical mutagenesis. This X-chromosomal mutation causes the D. melanogaster male's acoustical output, resulting from his wing vibrations directed at a female, to include very long and loud tone "pulses." Yet, a given train of pulses starts out as normal, with the signals in all but the shortest singing bouts eventually becoming polycyclic and high-amplitude. The aberrant songs caused by diss (map position, 1-52; cytological interval, 14C1-2 to 14C4-5) were quantitatively compared to those produced by mutant cacophony males, whose pulses are much more uniformly polycyclic (due to a mutation mapping elsewhere on the X chromosome). Males or females expressing diss are normal in several "general" behaviors. Yet diss males not only sing abnormally, but they also exhibit longer-than-normal mating latencies in their courtship of females. These decrements seem to be associated, at least in part, with visually aberrant behavior of diss flies--measured with regard to male courtship per se, and also in tests of more general visual responses. Such defects were found when testing diss males or females, and the genetic etiology of the visual impairments were provisionally mapped to the same locus to which the song abnormality has been localized. Neurogenetic connections between the control of courtship singing behavior and visual system functions are discussed with respect to the new song mutation (diss) and the older one (cac)--which also turned out to be genetically related to a mutation that causes abnormalities of light-induced behavior and physiology.

Animals↗

Copulatory courtship and cryptic female choice in red flour beetles Tribolium castaneum.

Males of many animal species engage in courtship behaviours during and after copulation that appear to be solely aimed at stimulating the female. It has been suggested that these behaviours have evolved by cryptic female choice, whereby females are thought to impose biases on male postmating paternity success. Males of the red flour beetle Tribolium castaneum rub the lateral edges of the females' elytra with their tarsi during copulation. We manipulated female perception of this behaviour by tarsal ablation in males, thus preventing males from reaching the edge of the female elytra with their manipulated legs, and by subsequently performing a series of double-mating experiments where the copulatory behaviour was quantified. We found a positive relationship between the intensity of the copulatory courtship behaviour and relative fertilization success among unmanipulated males. This pattern, however, was absent in manipulated males, where female perception of male behaviour differed from that actually performed. Thus, female perception of male copulatory courtship behaviour, rather than male behaviour per se, apparently governs the fate of sperm competing over fertilizations within the female, showing that copulatory courtship is under selection by cryptic female choice.

Animals↗

Functional coupling of acoustic and chemical signals in the courtship behaviour of the male Drosophila melanogaster.

During courtship, the male Drosophila melanogaster sends signals to the female through two major sensory channels: chemical and acoustic. These signals are involved in the stimulation of the female to accept copulation. In order to determine the respective importance in the courtship of these signals, their production was controlled using genetical and surgical techniques. Males deprived of the ability to emit both signals are unable to mate, demonstrating that other (e.g. visual or tactile) signals are not sufficient to stimulate the female. If either acoustic or chemical signals are lacking, the courtship success is strongly reduced, the lack of the former having significantly more drastic effects. However, the accelerated matings of males observed with males bearing wild-type hydrocarbons compared with defective ones, whichever the modality of acoustic performance (wing vibration or playback), strongly support the role of cuticular compounds to stimulate females. We can conclude that among the possible factors involved in communication during courtship, acoustic and chemical signals may act in a synergistic way and not separately in D. melanogaster.

Animal Communication↗

The unusual inheritance pattern of the courtship songs in closely related grasshopper species of the Chorthippus albomarginatus-group (Orthoptera: Gomphocerinae).

Sibling species of the Chorthippus albomarginatus-group are exceptional among all European grasshopper species because they have an extremely elaborate courtship behaviour. Here, we present a genetic analysis of the courtship song differences between two closely related grasshopper species Ch. albomarginatus and Ch. oschei. Measurements of seven courtship song characters and one parameter of courtship visual display were compared among parentals, F(1) and F(2) hybrids, and backcrosses. Variation in one morphological character, the number of stridulatory pegs, was studied as well. The distributions of song phenotypes were more consistent with a type-III genetic architecture, which involves complementary or duplicate pairs of loci. We suggest that this type of architecture may provide the novel elements of the hybrid songs, even more complex than the parental ones, which may offer a new starting point for sexual selection. One parameter of the visual display appeared to be controlled by a single locus, but differences in other characters of visual display could be explained by a more complex inheritance pattern. Inheritance of the peg number differences was consistent with a simple polygenic additive model. Sex linkage and/or maternal effects were not detected for any trait. We found a disproportionate influence of the Ch. albomarginatus parent on most hybrid song characteristics and the visual display, and this may support a previously suggested idea that Ch. albomarginatus is an ancestral form and Ch. oschei is a derived species.

Animal Communication↗

Effects of testosterone metabolites and estrogen in the midbrain control of courtship behavior in the male ring dove (Streptopelia risoria).

Relative effectiveness of testosterone (T) and its metabolites in the midbrain to induce courtship behavior was investigated in male ring doves. T, 5 alpha-dihydrotestosterone (5 alpha-DHT), 5 beta-dihydrotestosterone, estradiol-17 beta (E), or cholesterol was implanted in or around the nucleus intercollicularis region (ICo) of the castrated male. Resulting courtship behavior was observed by pairing the treated males with stimulus females. T, 5 alpha-DHT and E activated nest-cooling behavior pattern when placed in the ICo region only. Other male courtship behaviors were not induced, induced in each brain region tested or induced by all the hormones. Hormone diffusions from the implant did not appear to be responsible for these results since many implants within 1 mm of the ICo region failed to activate nest-cooling. These data support the idea that the midbrain along with the hypothalamic region mediate male courtship behavior.

Androgens↗

The effect of substrate on the efficacy of seismic courtship signal transmission in the jumping spider Habronattus dossenus (Araneae: Salticidae).

The jumping spider Habronattus dossenus Griswold 1987 (Salticidae) communicates using seismic signals during courtship and can be found on rocks, sand and leaf litter. We examined the filtering properties of, and tested the efficacy of male courtship signals on, these natural substrates. These substrates have drastically different filtering properties at the distances at which the males court. Rocks sharply attenuated all frequencies, with considerable variability among different rocks. Desert sand showed band-pass properties, attenuating frequencies contained in the animal's signal. Leaf litter passed all frequencies and was the most favourable signalling environment. In behavioural trials, the proportion of males mating successfully was significantly higher on leaf litter than on rocks or desert sand. Males did not modify their courtship behaviour on different substrates. Therefore, the effectiveness of male courtship seismic signals appears to be strongly constrained by the available substratum resources.

Animal Communication↗

Fruitless is in the regulatory pathway by which ectopic mini-white and transformer induce bisexual courtship in Drosophila.

Bisexual courtship in male Drosophila melanogaster may be induced in some circumstances. These include ectopic expression of the transformer (tra) gene, ectopic expression of the mini-white (mw) gene, and the homozygous presence of mutant alleles of the fruitless (fru) gene. Experiments were performed to determine if ectopic mw and fru, as well as ectopic tra and fru, acted in the same pathway to control courtship. Male flies homozygous for the frusat allele court females little if at all and males at a low level. When homozygous, the frusat allele suppresses the bisexual courtship induced by both ectopic mw and ectopic tra, indicating that the fru wild-type function is necessary for expression of the ectopic mw and ectopic tra effect. This demonstrates that fru shares a pathway controlling courtship behavior with these ectopically expressed genes.

Animals↗

Social transmission of courtship behavior and mating preferences in brown-headed cowbirds, Molothrus ater.

Studies of the social learning of courtship behaviors and mating preferences of brown-headed cowbirds (Molothrus ater) are reviewed. Earlier work has suggested that cowbirds from behaviorally distinct populations mate preferentially with others from the same population. Studies are described which indicate that patterns of courtship behavior that differ by population can be socially transmitted across generations are described. Social background affects male songs, female preferences for males as mates, and courtship interactions between females and males. Thus, social traditions influence mating decisions and may, ultimately, impact reproductive success in this species. Recent work on possible social mechanisms involved in the ontogeny of courtship behavior is described. The implications of these findings are discussed, and future research directions are suggested.

Adaptation, Psychological↗

Drosophila conditioned courtship: two ways of testing memory.

In Drosophila, courtship reduction in male flies that have previous experience of courting a mated female is a result of the counterconditioning of an attractive unconditioned stimulus (US)--the aphrodisiac--which becomes an aversive conditioned stimulus (CS) after being paired with an aversive US--the antiaphrodisiac. In a retention test with a virgin female lacking the antiaphrodisiac, males retain a lower level of courtship for 3 hr after training. However, a measure of courtship suppression, the learning index (LI), decreases significantly after only 1 hr. In contrast, in the retraining test with a mated female, the LI shows no decrease for 8 hr but falls below significance 16 hr after training. These results are discussed in terms of the transfer of training. Nonspecific transfer and nonassociative behavioral modifications play little, if any, role in the transfer of training. The retraining test is recommended as a new protocol for studying conditioned courtship. According to the model proposed here, in tests with a virgin female, the duration of memory retention is limited by the retention of the direct association between the CS and the aversive motivational system or by the retention of an internal representation of the US. In retraining tests, the CS-US association seems to be the only factor involved in transfer 3 or more hours after training.

Animals↗

A gene necessary for normal male courtship, yellow, acts downstream of fruitless in the Drosophila melanogaster larval brain.

The fruitless (fru) gene is a member of the Drosophila melanogaster somatic sex determination genetic pathway. Although it has been hypothesized that the primary function of fru is to regulate a genetic hierarchy specifying development of adult male courtship behavior, genes acting downstream of fru have not yet been identified. Here we demonstrate that the yellow (y) gene is genetically downstream of fru in the 3(rd)-instar larval brain. Yellow protein is present at elevated levels in neuroblasts, which also show expression of male-specific FRU proteins, compared to control neuroblasts without FRU. A location for y downstream of fru in a genetic pathway was experimentally demonstrated by analysis of fru mutants lacking transcription of zinc-finger DNA binding domains, and of animals with temporal, spatial, or sexual mis-expression of male-specific FRU. A subset of fru and y mutants is known to reduce levels of a specific behavioral component of the male courtship ritual, wing extension, and FRU and Yellow were detected in the general region of the brain whose maleness is necessary for development of that behavior. We therefore hypothesized that ectopic expression of Yellow in the 3(rd)-instar brain, in a y null background, would rescue low levels of wing extension and male competitive mating success, and this was found to be the case. Overall, these data suggest that y is a downstream member of the fru branch of the D. melanogaster sex determination hierarchy, where it plays a currently unknown role in the development of adult male wing extension during courtship.

Animals↗

Aromatase inhibition abolishes courtship behaviours in the ring dove (Streptopelia risoria) and reduces androgen and progesterone receptors in the hypothalamus and anterior pituitary gland.

The aim of this study was to determine in the ring dove, the effects of aromatase inhibition on the expression of aggressive courtship and nest-soliciting behaviours in relation to the distribution of cells containing immunoreactive androgen (AR) and progesterone (PR) receptor in the hypothalamus and pituitary gland. Isolated sexually experienced ring doves were transferred in opposite sex pairs to individual breeding cages, and then injected with the aromatase inhibitor, fadrozole (four males and four females), or saline vehicle (four males and four females) for 3 days at 12 hourly intervals. Saline-injected control males displayed aggressive courtship behaviours (bow-cooing and hop-charging) and nest-soliciting throughout the study, and control females displayed nest-soliciting. By day 3, fadrozole treatment resulted in the disappearance of all these behaviours and in a decrease or disappearance of AR and PR in the anterior pituitary gland, and in the nucleus preopticus paraventricularis magnocellularis (PPM), nucleus preopticus medialis (POM), nucleus hypothalami lateralis posterioris (PLH), and ventral, lateral and dorsal nucleus tuberalis in the hypothalamus (VTu, LTu, DTu). In the nucleus preopticus anterior (POA), fadrozole treatment decreased AR in both sexes and decreased PR in females but not in males. Cells containing co-localized nuclear AR and PR were found in all hypothalamic areas examined, and in the anterior pituitary gland. Fadrozole is suggested to reduce the local availability of estrogen required indirectly for the induction of AR, and except in cells containing PR in the male POA, for the direct induction of PR. It is suggested that aggressive courtship behaviour is terminated by "cross talk" between aromatase-independent PR and aromatase-dependent AR co-localized in neurons in the POA. Aromatase-independent PR may increase in the male POA in response to visual cues provided by a partner. Aromatase-dependent PR in the POM, and basal hypothalamus may play a role in the facilitatory effect of progesterone on estrogen-induced nest-orientated behaviours.

Animal Communication↗

The role of the flabellar and ellipsoid bodies of the central complex of the brain of Drosophila melanogaster in the control of courtship behavior and communicative sound production in males.

The role of flabellar and ellipsoid bodies of the central complex of the brain of Drosophila melanogaster males in controlling courtship behavior and the accompanying sound production was studied by comparative analysis of the characteristics of courtship and singing in wild-type flies and individuals of five mutant lines with different anatomical defects in these parts of the brain. Investigations were performed using the following fly lines: Canton S (wild-type, controls), ebo(KS263), with defects only in the ellipsoid bodies, and ebo(1041), ceb(849), ceb(892), and cba(KS96), with different levels of abnormality in both parts of the central complex. The data obtained here indicated that the flabellar and ellipsoid bodies are involved in: 1) maintaining a high level of courtship activity; 2) regulating the precision of male movements while following females; 3) in controlling the nature and stability of the sound elements from which communicative signals are constructed; 4) in regulating the rhythmic structure of signals dependent on the stability of pacemakers function; 5) establishing the relationship between behavior and the context in which it occurs.

Animal Communication↗

Evolutionary insights into the regulation of courtship behavior in male amphibians and reptiles.

Comparative studies of species differences and similarities in the regulation of courtship behavior afford an understanding of evolutionary pressures and constraints shaping reproductive processes and the relative contributions of hormonal, genetic, and ecological factors. Here, we review species differences and similarities in the control of courtship and copulatory behaviors in male amphibians and reptiles, focusing on the role of sex steroid hormones, the neurohormone arginine vasotocin (AVT), and catecholamines. We discuss species differences in the sensory modalities used during courtship and in the neural correlates of these differences, as well as the value of particular model systems for neural evolution studies with regard to reproductive processes. For example, in some genera of amphibians (e.g., Ambystoma) and reptiles (e.g., Cnemidophorus), interspecific hybridizations occur, making it possible to compare the ancestral with the descendant species, and these systems provide a window into the process of behavioral and neural evolution as well as the effect of genome size. Though our understanding of the hormonal and neural correlates of mating behavior in a variety of amphibian and reptilian species has advanced substantially, more studies that manipulate hormone or neurotransmitter systems are required to assess the functions of these systems.

Amphibians↗

High levels of LH and testosterone in a tropical seabird with an elaborate courtship display.

For birds breeding in temperate areas, territoriality and courtship co-occur during spring and are mirrored by elevated plasma levels of luteinizing hormone (LH) and testosterone. By contrast, several tropical species show no major changes in LH and testosterone throughout the year and plasma levels are often low. This study describes changes in body condition, LH and testosterone levels throughout breeding in a tropical seabird, the magnificent frigatebird (Fregata magnificens). Although bi-parental care is the rule in frigatebirds, these seabirds are unique among seabirds in having a lek-like mating system where males perform an elaborate courtship display during which an extravagant sexual ornament (gular pouch) is inflated. Body condition peaked during the displaying stage in males and declined gradually in both sexes, being the lowest during chick-rearing. LH and testosterone titers were the highest in displaying males with a fully inflated gular pouch (means: 23.6 ng/ml for LH and 6.2 ng/ml for testosterone) and remained low (means: 3.8 ng/ml for LH and 0.3 ng/ml for testosterone) during incubation and chick-rearing in both sexes. Body condition was positively related to testosterone levels in displaying males. In male frigatebirds, high levels of testosterone are not associated with aggression but are probably required to display the inflated gular pouch and to perform an elaborate courtship display. Because males invest in incubation and chick-rearing, testosterone has to return to low levels, to permit parental care. This hormonal pattern differs greatly with that of the few tropical seabirds studied so far and is probably related to the unique reproductive strategy of frigatebirds.

Aggression↗

Genetic basis and evolution of species-specific courtship song in the Drosophila auraria complex.

The interpulse interval (IPI) of courtship song in the Drosophila auraria complex is the only parameter that is consistently species-specific among the several courtship elements examined within the complex. The genetic basis of the species-specific courtship song was examined by analysing the song of interspecific hybrids and of backcross progeny. IPI of all interspecific hybrids except two showed intermediate values, suggesting autosomal control of species-specific IPI. However, significant deviation for shorter IPI from midparent was found in thirteen out of 20 crosses. The chromosomal analysis between D. auraria and D. biauraria revealed that the two major autosomes had significantly large effects on IPI, but the sex chromosome and cytoplasm had no effect. Since no interaction was detected, it is concluded that each autosome acts additively in the determination of species-specific IPI. The common ancestors of the D. auraria complex may also have had autosomal control of IPI, which has been conserved during speciation in the complex.

Animals↗