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Effects of midbrain lesions on lordosis and ultrasound production.

Previous studies suggest the control of lordosis by competing neural systems. A lordosis-inhibiting system is thought to originate in the lateral septum (LS) or preoptic area (POA) and project to midbrain sites, including the ventral tegmental area (VTA) and ventral half of the periaqueductal gray (vPAG). A lordosis-facilitating system is thought to originate in the ventromedial hypothalamus and project to the dorsal half of the PAG (dPAG). To test the generality of this model, we subjected female hamsters to control operations or lesions of the dPAG, vPAG or VTA. Subjects in hormone-induced estrus were observed for lordosis responses to males and manual stimulation and for rates of production of the "ultrasounds" used by hamsters for courtship. PAG lesions of both types depressed ultrasound rate, lordosis initiation and lordosis maintenance. VTA lesions failed to affect ultrasound rate, lordosis maintenance or lordosis responses to males. However, damage to the dorsomedial tegmentum was correlated with deficits in lordosis initiation. These results suggest the concentration in the PAG of mechanisms controlling vocalizations, including the ultrasounds used by hamsters for courtship. They confirm midbrain control of lordosis initiation and maintenance but fail to support any simple specialization of dorsal and ventral areas for these dimensions. They also fail to support a specialization of the dorsal and ventral halves of the midbrain for lordosis facilitation and inhibition, but do suggest several midbrain areas likely to be involved in the first of these functions.

Analysis of Variance↗

Social housing conditions around puberty determine later changes in plasma cortisol levels and behavior.

A recent study found that male guinea pigs raised in large, mixed age/sex groups exhibited an unexpected suppression of their cortisol response at 4 mo of age. The present study examined the effect of social experience around the time of puberty on cortisol response suppression and social behavior at 4 mo of age. Males reared in large, mixed age/sex groups were either pair-housed with a female or moved to another large colony at 55 days of age. When tested at 4 mo, pair-housed males exhibited much higher levels of courtship and sexual behavior than did colony-housed males, and a shorter latency to begin courtship when with an unfamiliar adult female. In addition, pair-housed males showed much higher levels of agonistic behavior and a shorter latency to escalated aggression with an unfamiliar adult male. Pair-housed males also had lower basal cortisol concentrations and exhibited a greater increment in cortisol levels when isolated in a novel cage than did colony-housed males. Finally, pair-housed males showed a smaller increment in cortisol levels when with the stimulus female or male than when isolated, but colony-housed males did not. The findings demonstrate that social housing conditions around the time of puberty can have pervasive effects on social behavior and hypothalamic-pituitary-adrenal (HPA) activity in 4-mo-old males. Further, these findings are consistent with the notion that changes in HPA activity contribute to social behavior development beyond the time of sexual maturity.

Adaptation, Physiological↗

Blueprints for behavior: genetic specification of neural circuitry for innate behaviors.

Innate behaviors offer a unique opportunity to use genetic analysis to dissect and characterize the neural substrates of complex behavioral programs. Courtship in Drosophila involves a complex series of stereotyped behaviors that include numerous exchanges of multimodal sensory information over time. As we will discuss in this review, recent work has demonstrated that male-specific expression of Fruitless transcription factors (Fru(M) proteins) is necessary and sufficient to confer the potential for male courtship behaviors. Fru(M) factors program neurons of the male central and peripheral nervous systems whose function is dedicated to sexual behaviors. This circuitry seems to integrate sensory information to define behavioral states and regulate conserved neural elements for sex-specific behavioral output. The principles that govern the circuitry specified by Fru(M) expression might also operate in subcortical networks that govern innate behaviors in mammals.

Animals↗

Effects of androgens on behavioral and vomeronasal responses to chemosensory cues in male terrestrial salamanders (Plethodon shermani).

Chemosensory stimuli and sex steroid hormones are both required for the full expression of social behaviors in many species. The terrestrial salamander, Plethodon shermani, is an emerging nonmammalian system for investigating the nature and evolution of pheromonal communication, yet little is known regarding the role of sex steroid hormones. We hypothesized that increased circulating androgen levels in male P. shermani enhance chemoreception through morphological, behavioral, and physiological mechanisms. Experimental elevation of plasma androgens increased development of cirri, morphological structures thought to enhance the transfer of chemosensory cues from the substrate to the vomeronasal organ (VNO). Elevated plasma androgens also increased expression of a chemo-investigatory behavior (nose tapping) and increased preference for some female-derived chemosensory cues. Male-produced courtship pheromones activated a large number of cells in the VNO as measured by the method of agmatine uptake. However, androgen levels did not affect the total number of vomeronasal cells activated by male-produced courtship pheromones. Future studies will determine whether androgens potentially modulate responsiveness of the VNO to female-derived (as opposed to male-derived) chemosensory cues.

Analysis of Variance↗

A sexual selection theory longitudinal analysis of sexual segregation and integration in early adolescence.

The three objectives in this longitudinal study were motivated by sexual selection theory. The theory specifies the role of sexually segregated groups and the effects of dominance in male groups and relational/indirect aggression in female groups for heterosexual relationships. Using a multi-method, multi-informant, longitudinal design we studied youngsters (N=138) across their first two years of middle school. First, we examined the nature of change in segregation and dating popularity across two years during early adolescence. Second, a model derived from sexual selection theory is tested to explain the ways in which boys and girls are nominated for hypothetical dates (dating popularity). Third, we examined the role of "poke and push courtship" behavior in boys' and girls' dating popularity. Results indicate that although groups did not become more integrated with time, changes in peer group sexual integration co-varied dynamically with dating popularity. Secondly, dominance-related strategies were more important for boys than girls in dating popularity whereas indirect, or relational, aggression strategies were more important for girls than boys. Third, "poke and push courtship" behaviors did not influence peer group integration or dating.

Adolescent↗

Urinary testosterone levels in the male blind mole rat (Spalax ehrenbergi) affect female preference.

This study investigated the sexual attraction of female blind mole rats to four groups of male mole rats: (a) intact males raised in captivity; (b) intact males trapped in the field; (c) captive males injected with testosterone; (d) captive castrated males. In the first part we measured blood testosterone, androstenedione, and dihydrotestosterone (DHT) levels, by radioimmunoassay; and urine testosterone levels, measured by GC-MS. The second part examined the relationship between urine testosterone levels in males and their attractiveness to females. Higher blood and urine testosterone levels were found in the field animals and in those injected with testosterone compared to captive intact or castrated animals: urine testosterone levels in the two other groups were not detectable. Blood androstenedione levels were also higher in the field animals and in those injected with testosterone compared to captive intact or castrated mole rats. Blood dihydrotestosterone levels were not detectable in all four experimental groups. Female mole rats chose to spend a longer period of time next to males with high blood and urine testosterone levels and high blood androstenedione levels than next to those with lower levels of these hormones. Because courtship and sexual behavior are influenced both by high levels of blood and urine testosterone and high levels of blood androstenedione, we suggest that the low levels of courtship and other sexual behavior in captive mole rats may be related to the lack of female attraction to these males, which display low levels of all three parameters.

Androstenedione↗

Factors affecting male song evolution in Drosophila montana.

D. montana (a species of the D. virilis group) has spread over the northern hemisphere, populations from different areas showing both genetic and phenotypic divergence. The males of this species produce an elaborate courtship song, which plays a major role both in species recognition and in intraspecific mate choice. The genetic architecture and physical constraints, as well as the importance of the signal for species recognition, set boundaries within which this signal can vary. Within these limits, courtship song parameters may change, depending on the males' physical condition and on the environment they inhabit. Females are likely to affect song evolution by exerting directional selection toward higher carrier frequencies. Given this complexity, only a comprehensive, multidisciplinary approach, starting with traditional field observation and combining controlled behavioral experiments, biometric measurements, and sophisticated molecular techniques, has the potential of shedding light on the past history and the evolution of this signal, and, eventually, adding to our understanding of the mechanisms, functions, and outcomes of sexual selection in acoustic communication systems.

Animals↗

Control of male sexual behavior and sexual orientation in Drosophila by the fruitless gene.

Sexual orientation and courtship behavior in Drosophila are regulated by fruitless (fru), the first gene in a branch of the sex-determination hierarchy functioning specifically in the central nervous system (CNS). The phenotypes of new fru mutants encompass nearly all aspects of male sexual behavior. Alternative splicing of fru transcripts produces sex-specific proteins belonging to the BTB-ZF family of transcriptional regulators. The sex-specific fru products are produced in only about 500 of the 10(5) neurons that comprise the CNS. The properties of neurons expressing these fru products suggest that fru specifies the fates or activities of neurons that carry out higher order control functions to elicit and coordinate the activities comprising male courtship behavior.

Age Factors↗

Neuronal cell death during sexual differentiation and lateralisation of vocal communication.

A rodent analogy has been established to investigate the neural mechanisms occurring during sexual differentiation and lateralization. A sexually dimorphic hypothalamic nucleus (SDApc) is closely associated with a stereotyped, courtship vocalisation in male gerbils. Stereological analysis of SDApc cytoarchitecture reveals that neuron number and nuclear volume are asymmetric in male adults. Strikingly, neuron number on the left side of the SDApc correlates significantly with the rate of the courtship call in males. Exogenous testosterone treatment in female neonates masculinises and lateralises SDApc structure and function. Neuronal programmed cell death (apoptosis), manifested in SDApcs of neonates, is more frequent in females. Significantly, apoptosis in males is lateralised, as revealed by lateral asymmetry of neuron number at postnatal day 16. It is concluded that neuroendocrine-dependent, sexual differentiation and lateralization are concurrent and influenced by apoptotic mechanisms. It is suggested that apoptosis is the result of a genetically-driven device, inherent in postmitotic, undifferentiated cells which may have recently migrated into the SDApc. The genomic mechanism inducing lateralised apoptosis is apparently activated only neonatally in males.

Animals↗

Behavioral effects of chronic exposure to low levels of lead in Drosophila melanogaster.

Through human activity lead has become a serious environmental neurotoxin, known to affect activity levels, attention and both sensory and cognitive function in children. Study of lead would be facilitated by having a model system that could be manipulated easily and quickly. We find Drosophila melanogaster ideal as such, and we have been studying effects of lead on courtship, fecundity and locomotor activity. We raised Canton-S flies from eggs to adult day 6-7 on medium made with lead acetate solution (2-100 microgram/g), or with distilled water, and we measured adult body lead burdens by means of Inductively Coupled Plasma Mass Spectrometry (ICP-MS). To measure courtship effectiveness, five virgin females and five virgin males were transferred into an empty vial and the number of females that mated within 20min was recorded. To measure fecundity, all adult offspring from eggs produced by one female within 12 days of mating were counted. To measure locomotor activity, individual flies were transferred to a grid-labeled petri dish and the number of lines crossed in 30s was counted. The number of females mating within 20min was increased significantly by exposure to 2 or 8 microgram/g lead, and was decreased significantly by exposure to 20 or 50 microgram/g lead. Fecundity was increased significantly by exposure to 2 microgram/g lead, but was unaffected by exposure to 20 microgram/g lead. Locomotor activity was consistently higher for males than for females, and was significantly reduced only by exposure to 50 microgram/g lead, and then only for males. We thus defined for Drosophila a lowest observable effect level (LOEL) of 2 microgram/g lead, which is considerably lower than the doses shown previously to affect this animal. The dose-response curve was biphasic for the number of females mating within 20min, an example of hormesis, a non-linear response that has been reported for low levels of stressors as diverse as pollutants and radiation. We hope from further studies with Drosophila to understand better how lead affects the developing nervous system, and thus ultimately its effects on children.

Animals↗

Modulation of respiratory pump rate by reproductive behaviors in freely behaving pairs of Aplysia fasciata.

Respiratory pumping in Aplysia is a spontaneously occurring behavior whose neural circuitry has been explored, but whose natural functions are incompletely understood. Respiratory pump rate was examined in freely behaving pairs of Aplysia fasciata, to determine whether it is modified by the occurrence of mating and other behaviors. The background rate of respiratory pumping was approximately 2/hour. This rate was maintained while animals were immobile, moving in place, crawling, or feeding. The rate was increased to over 8/hour during courtship and to approximately 4/hour during female mating and was reduced to approximately 1/hour during male-mating. These data suggest that respiratory pumping has a reproductive function, perhaps in dispersal of pheromones that are released during female-mating and courtship. Respiratory pumping never occurred while animals were swimming, suggesting that respiratory pumping and swimming may be mutually incompatible behaviors. Respiratory pumping was less common by night than by day.

Animals↗

Male mouse (Mus musculus) ultrasonic vocalizations to female urine: why is heterosexual experience necessary?

Previous research was consistent with the hypothesis that urinary chemosignals from female mice (Mus musculus) serve as a conditioned stimulus (CS) for the elicitation of male ultrasonic courtship vocalizations while some other unknown aspect of the female serves as an unconditioned stimulus (US). According to this hypothesis adult heterosexual experience is necessary for males to pair the urinary CS with the US. Three experiments further examined this hypothesis. Experiment 1 demonstrated that the hypothesized US was not female behavior. Experience with anesthesized males and females was just as effective as experience with awake conspecifics. Experiments 2 and 3, however, question the primacy of the Classical Conditioning hypothesis. In both experiments sexually naive males were allowed contact with either normal females or female surrogates. The female surrogates were neonatally castrated males (Experiment 2) or hypophysectomized females (Experiment 3), both of which appeared to possess the hypothesized US but not the urinary CS on the basis of previous research. While exposure to normal females caused the highest level of vocalization to urine from normal females, exposure to the two classes of female surrogates also resulted in vocalizations to the urine of normal females. Thus under some circumstances, males do not require experience with a normal female to emit ultrasounds to urine from normal females. Factors in addition to Classical Conditioning must be operating to account fully for the role of adult heterosexual experience in causing female urine to come to elicit male courtship vocalization.

Animals↗

Effect of alternating bulls as a management tool to improve the reproductive performance of suckled Zebu cows in the humid tropics of Costa Rica.

A study was undertaken to evaluate the effect of alternating bulls between a single and a multiple sire mating (MSM) program on the reproductive performance of suckled Zebu cows raised under range conditions in the humid tropics of Costa Rica. Multiparous Zebu cows (n=94) suckling calves were distributed between two experimental trials (A and B) consisting of 47 animals each. A single sire mating (SSM) system was alternated weekly with a MSM system with three bulls. This period lasted for 8 weeks. To facilitate estrous expression, four cows were strategically synchronized (estrus-stimulated) in alternate weeks. Courtship predominated over mounting under non-stimulated estrus, for each mounting performed an average of 6.0 and 6.3 courtship activities were recorded in the SSM and MSM, respectively. Under the influence of strategic synchronization corresponding values were 3.9 and 4.2 in the SSM and MSM, respectively (P>0.05). Blood samples for progesterone evaluation were taken twice weekly. All cows in trial A were in anestrus at the start of the study. By second week, 5 out of the 47 cows had initiated estrous cycles and by the third week six were pregnant. In contrast in trial B, 9 out of 47 had initiated estrous cycles before interacting with the bulls and on week 3, only two females had become pregnant and three had initiated estrous cycles. Significant differences were found in the cumulative percentage of cows pregnant between trials A and B (P<0.05). Even though these results occurred, the rotation of the bulls (one or three), or the type of cows (estrus-stimulated or not) did not influence the results in this study.

Anestrus↗

Sex differences in the densities of alpha 2-adrenergic receptors in the song control system, but not the medial preoptic nucleus in zebra finches.

In songbirds, song is regulated by a specialized group of brain nuclei known as the song system. Other aspects of courtship, such as male sexual interest in a female, are likely regulated by the medial preoptic nucleus (POM). The song control system and the POM are rich in norepinephrine, which appears to regulate courtship behaviors, including song. Zebra finches (Taeniopygia guttata) exhibit an extreme sexual dimorphism in song behavior; males sing, primarily to attract or maintain mates, and females do not. We explored possible sex differences in the distribution and density of the alpha(2)-adrenergic receptors in the song system and POM of zebra finches. Receptors were labeled with the selective ligand, [(3)H] RX821002, via autoradiographic procedures. In males, dense alpha(2)-receptors were observed in the song system (Area X, the high vocal center (HVc), the lateral portion of the magnocellular nucleus of the anterior neostriatum, and the robust nucleus of the archistriatum). In contrast, in females neither the lateral portion of the magnocellular nucleus of the anterior neostriatum nor the HVc could be identified based on alpha(2)-receptor binding. Females lack Area X and indeed differential alpha(2)-binding was not observed within the female lobus parolfactorius. The robust nucleus of the archistriatum contained less dense alpha(2)-binding in females compared to males. Alpha(2)-binding in the POM was similar in males and females. The dimorphism in alpha(2)-binding in nuclei of the song system likely relates to the dimorphism in song behavior observed in male and female zebra finches.

Animals↗

Learning without performance in PKC-deficient Drosophila.

The performance of a task is often assumed to be a prerequisite for the learning of many tasks, including the associative conditioning of courtship in the fruit fly, Drosophila melanogaster. Transgenic flies specifically inhibited for the enzyme protein kinase C dissociate the acquisition of learning and memory from performance of the task. They fail to show immediate suppression of courtship but nonetheless develop normal memory of it.

Animals↗

From behavior to development: genes for sexual behavior define the neuronal sexual switch in Drosophila.

The isolation and analysis of Drosophila mutants with altered sexual orientation lead to the identification of novel branches in the sex-determination cascade which govern the sexually dimorphic development of the nervous system. One such example is the fruitless (fru) gene, the mutation of which induces male-to-male courtship and malformation of a male-specific muscle, the muscle of Lawrence (MOL). Since the MOL is formed in wild-type flies when the innervating nerve is male, regardless of the sex of the MOL itself, the primary site of Fru function is likely to be the motoneurons controlling the MOL. The fru gene produces multiple transcripts including sex-specific ones. A female-specific mRNA from the fru locus has a putative Transformer (Tra) binding site in its 5' untranslated region, suggesting that fru is a direct target of Tra. The fru transcripts encode a set of proteins similar to the BTB (Bric à brac, Tramtrack and Broad-complex)-Zn finger family of transcription factors. Mutations in the dissatisfaction (dsf) gene result in male-to-male courtship and reduced sexual receptivity of females. The dsf mutations also give rise to poor curling of the abdomen in males during copulation and failure of egg-laying by females. The latter phenotypes are ascribable to aberrant innervation of the relevant muscles. A genetic analysis reveals that expression of the dsf phenotypes depends on Tra but not on Doublesex (Dsx) or Fru, suggesting that dsf represents another target of Tra. Taken together, these findings suggest that the sex-determination protein Tra has at least three different targets, dsx, fru and dsf, each of which represents the first gene in a branch of the sex-determination hierarchy functioning in a mutually-exclusive set of neuronal cells in the Drosophila central nervous system.

Animals↗

Genetic identification of neurons controlling a sexually dimorphic behaviour.

In the fruit fly Drosophila melanogaster, locomotor activity is sexually dimorphic: female flies constantly modulate their activity pattern whereas males show a steadier, stereotyped walking pace [1]. Here, we mapped the area of the brain controlling this behavioural dimorphism. Adult male Drosophila expressing a dominant feminising transgene in a small cluster of neurons in the pars intercerebralis exhibited a female-like pattern of locomotor activity. Genetic ablation of these neurons prevented the feminisation of the locomotor activity of transgenic males. The results suggest that this cluster of neurons modulates sex-specific activity, but is not involved in initiating fly locomotion. Nor does it control male courtship behaviour, because feminisation of courtship was not correlated with the feminisation of locomotor activity.

Animals↗

Sexual behaviour in Drosophila is irreversibly programmed during a critical period.

Sexual differentiation in Drosophila is controlled by a short cascade of regulatory genes, the expression pattern of which determines all aspects of maleness and femaleness, including complex behaviours displayed by males and females [1-3]. One sex-determining gene is transformer (tra), the activity of which is needed for female development. Flies with a female karyotype (XX) but which are mutant for tra develop and behave as males. In such flies, a female phenotype can be restored by a transgene that carries the female-specific cDNA of tra under the control of a heat-shock promoter. This transgene, called hs[trafem], also transforms XY animals into sterile females [4]. When we raised these XX and XY 'females' at 25 degreesC, however, they displayed vigorous male courtship while at the same time, as a result of their female pheromone pattern, they were attractive to males. Intriguingly, their male courtship behaviour was indiscriminately directed towards both females and males. When we forced expression of tra by heat shock, applied during a limited period around puparium formation, male behaviour was abolished and replaced by female behaviour. We conclude that sexual behaviour is irreversibly programmed during a critical period as a result of the activity or inactivity of a single control gene.

Animals↗