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Effects of courtship on brain gonadotropin hormone-releasing hormone and plasma steroid concentrations in a female amphibian (Taricha granulosa).

Courtship-induced changes in plasma steroid and brain gonadotropin-releasing hormone (GnRH) concentrations in Taricha granulosa were determined with respect to changes in female sexual receptivity. Females were sacrificed at several times after courtship initiation. Concentrations of GnRH (determined by RIA) in the anterior telencephalon were high at courtship initiation (females unreceptive), but decreased by sperm transfer (females receptive). Courtship had no affect on GnRH concentrations in any other brain region examined. Furthermore, courted, receptive females had higher plasma levels of estradiol than did uncourted controls, and estradiol levels remained elevated above control levels 24 hr after courtship initiation. Courtship had no influence on plasma progesterone or corticosterone levels. To determine if the observed changes in GnRH concentrations in the telencephalon were localized to the nervus terminalis, courted females and controls were sacrificed after 5, 20, or 60 min of courtship. Nervus terminalis GnRH concentrations were higher in courted females than in uncourted controls. These results may represent the first documentation of a naturally occurring physiological change in the nervus terminalis.

Animals

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

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

Animals

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

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

Blotting, Northern

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

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

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

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

Effects of the apterous4 mutation on Drosophila melanogaster males' courtship.

The apterous4 (ap4) mutation has pleiotropic effects on the morphology and lifespan of D. melanogaster adults. The mutation also affects one aspect of male sexual behaviour. Unlike normal, sexually mature males, three day old ap4/ap4 males synthesize courtship-stimulating hydrocarbons that immature males make; as a result they elicit vigorous courtship (Jallon and Hotta, 1979; Jallon et al., 1986). Data presented in this report indicate that expression of the apterous4 mutation does not affect all aspects of sexual maturation in males. Specifically, 24-30 hour old ap4/ap4 males are capable of performing all of the courtship behaviors, including abdominal vibration, a newly described response of mutant and wild-type males to virgin females. However, the mutant males spend less time courting and are less likely to perform some of the courtship behaviours than age-matched controls. These abnormalities are probably indirect effects of the apterous4 allele's previously unreported effects on the flies' motor functions, which include partial paralysis, lack of coordination, and sluggishness.

Animals

Courtship behavior of Drosophila genetically or surgically deprived of basiconic sensilla.

The lack of basiconic antennal sensilla in the mutant lozenge was used to assess the role of these olfactory receptors in the courtship behavior of Drosophila melanogaster. Under normal light conditions, lozenge males courted virgin females much less than wild-type males did. However, when visual courtship stimuli were eliminated by studying behavior under dim red light, the two kinds of males courted individual wild-type virgin females with the same intensity, and the latency to copulation was similar. Also, no difference in courtship vigor was observed if the two kinds of males were paired in red light with a mated female. These data suggest that antennal basiconic sensilla are important for neither the perception of the attraction pheromone(s) of virgin females nor the inhibitory pheromone(s) of mated females. Similar assays with males deprived of maxillary palps make it unlikely that the basiconic-like sensilla on these appendages are needed to perceive the attraction pheromones. However, the unexpectedly high courtship activity of palp-deprived males toward mated females suggests that basiconic-like maxillary sensilla may be receptors of inhibitory female compounds.

Animals

Plasma testosterone and 17 beta-estradiol concentrations, and aromatase activity, during courtship in male Triturus carnifex.

Plasma testosterone and 17 beta-estradiol were monitored during the main phases of male Triturus carnifex courtship. "Inactive" males showed higher levels of testosterone with respect to those found during the various courtship phases. Estradiol was low in "inactive" males, and it reached the highest values at the beginning of the courtship. In addition aromatase activity was higher in the brain of the newts at the beginning of the courtship. These findings seem to support the "aromatization hypothesis" in Triturus carnifex.

Animals

Quantitative genetic analysis of courtship and reproduction in female Drosophila melanogaster.

Three hundred mother-daughter pairs were analyzed for seven attributes related to courtship and reproduction. Only the lag time from first courtship to copulation was significantly heritable; genetic correlations involving this attribute were not significant. The genetic correlation between fertility and lag time to first courtship was negative and significant. However, this genetic correlation is expected to have little impact on the retention of additive genetic variance or on response to selection because it involves two attributes with low heritabilities. The pattern of phenotypic covariation among traits is largely explained by environmental causes and is consistent with that found in a previous analysis of father-son pairs (Gromko, 1987).

Alleles

Artificial neural network classification of Drosophila courtship song mutants.

Courtship songs produced by Drosophila males--wild-type, plus the cacophony and dissonance behavioral mutants--were examined with the aid of newly developed strategies for adaptive acoustic analysis and classification. This system used several techniques involving artificial neural networks (a.k.a. parallel distributed processing), including learned vector quantization of signals and non-linear adaption (back-propagation) of data analysis. "Pulse" song from several individual wild-type and mutant males were first vector-quantized according to their frequency spectra. The accumulated quantized data of this kind, for a given song, were then used to "teach" or adapt a multiple-layered feedforward artificial neural network, which classified that song according to its original genotype. Results are presented on the performance of the final adapted system when faced with novel test data and on acoustic features the system decides upon for predicting the song-mutant genotype in question. The potential applications and extensions of this new system are discussed, including how it could be used to screen for courtship mutants, search novel behavior patterns or cause-and-effect relationships associated with reproduction, compress these kinds of data for digital storage, and analyze Drosophila behavior beyond the case of courtship song.

Algorithms

Drosophila courtship song cycles in normal and period mutant males revisited.

Courtship songs of normal males and those expressing short-period, long-period, and arrhythmic mutations at the period (per) locus of Drosophila melanogaster have been reanalyzed for rhythmic components, using spectral treatments of the fluctuating rates of tone pulse production that occur during courtship. It was concluded, as in previous studies, that such songs are strongly rhythmic, except for courtships performed by per01 males. Songs produced by males expressing this and other per alleles were compared to computer-generated "random" ones. Interpulse interval variations influenced by per01 and songs stimulated to be arrhythmic both were found to be associated with cryptic rhythmicities; several such period values, extracted by the spectral analyses, defined very short cycle durations. We discuss the implications of these findings and of some recently reported results that have challenged the existence of rhythmicity in Drosophila songs.

Animal Communication

Courtship song and mating speed in hybrids between Drosophila melanogaster and Drosophila simulans.

Courtship song and mating speed of hybrids between Drosophila melanogaster and D. simulans were investigated. The courtship song of hybrid males is identical to that of D. simulans, suggesting that X chromosome determination, known from the cross between D. pseudoobscura and D. persimilis, is also possible here. Wingbeat frequency of hybrids is intermediate between that of the two parents, demonstrating that courtship song and wingbeat frequency are inherited independently of each other. In mating test, hybrid males cout and are accepted by D. simulans females more than hybrid females (presumably because their song is more "acceptable" to the former). D. melanogaster males readily, hybrids less readily, and D. simulans least.

Animals

Inheritance of male courtship sound characteristics in Drosophila littoralis.

Males of Drosophila littoralis vibrate their wings during courtship to deliver a "love song." This consists of 25- to 50-ms-long pulses with a basic frequency of about 250-400 Hz, separated by 250- to 500-ms pauses. When recording the sounds of flies from several localities in Europe, we found that males of one strain from northern Finland displayed courtship sounds with an unusually low wing beat frequency (below 250 Hz). In a genetic analysis utilizing marker stocks, the anomalous frequency was found to be caused by genes on all major autosomes, the strongest factors being on the second chromosome. Interaction between genes on chromosome 2 and on the fused chromosome 3-4 was non-additive. In low-frequency sounds, the number of cycles in the pulse (CN) was decreased, so that the length of the sound pulse (PL) remained more or less unchanged. We suggest that the genetically and physiologically most thoroughly controlled trait in the sound of Drosophila littoralis is the length of the pulse.

Animals

Courtship behavior and control of reproductive isolation in Drosophila mojavensis: genetic analysis of population hybrids.

Drosophila mojavensis from the Sonora region and Baja California show asymmetrical sexual isolation in the laboratory: males from Sonora mate equally frequently with Sonora and Baja females, while the mating success of Baja males with Sonora females is reduced. This failure has been localized to three separate behavioral landmarks occurring during courtship. Genetic analysis was conducted using reciprocal F1 hybrids of Sonora and Baja strains to examine inheritance patterns of the responsible courtship behaviors. Mating success and propensity of F1 males were similar to Sonora males. F1 females mated with males of Sonora and Baja races equally, although mating propensity of F1 females was intermediate between that of Sonora and Baja females. Males of Baja strains presented with F1 females showed a relatively high level of failure at attempted intromission. Genes for mating behaviors are located in the autosomes, but different loci responsible for the sexual isolation appear to act in males and females.

Animals

Role of catecholamines in the courtship behavior of male ring doves.

The role of catecholamines in the expression of male courtship behavior in ring doves was examined using central administration of pharmacological agents. Males treated with 6-hydroxydopamine or U-14,624, which depleted norepinephrine (NE) levels in the preoptic-hypothalamic area, showed increased levels of bow-coo and nest-coo displays. Conversely, males treated with tyramine or desipramine, which elevated NE levels in the preoptic-hypothalamic area, showed decreased levels of bow-coo and nest-coo displays. Drug-induced changes in dopamine levels were not consistent with any changes in behavior. This suggests that in the male ring dove NE in the preoptic-hypothalamic area is important in the expression of courtship displays.

Animals