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Correlations between catecholamine levels and sexual behavior in male zebra finches.

In zebra finches, the combined actions of estrogens and androgens activate male courtship, including singing, and also strongly modulate norepinephrine (NE) levels and turnover in brain areas known to be involved in controlling courtship behavior. To determine whether changes in NE levels mediate changes in courtship, we administered DSP-4 to males and measured its effects on monoamine levels and reproductive behavior. DSP-4 treatment did not affect serotonin (5-HT), had small, variable effects on dopamine (DA), and caused moderate, nonsignificant reductions in NE. However, in DSP-4-treated males, NE levels in specific vocal-control nuclei showed high positive correlations with courtship singing. There were no significant correlations between NE levels in hypothalamic nuclei and any behavior or DA or 5-HT levels in any nuclei and any behavior. DSP-4-treated males took longer to begin singing and performed fewer song bouts and courtship displays, but their songs could not be differentiated from those of control males. This suggests that their behavioral deficits resulted from deficits in attention rather than an inability to sing.

Animals↗

The genetic architecture of house fly mating behavior.

This chapter summarizes several experimental approaches used to identify the effects of dominance, epistasis, and genotype-by-environment interactions in the genetic architecture of the mating behavior of the common house fly (Musca domestica L.). Quantitative genetic investigations of mating behavior hold special intrigue for unraveling the complexities of fitness traits, with applications to theory on sexual selection and speciation. Besides being well suited to large-scale quantitative genetic protocols, the house fly has a remarkably complex courtship repertoire, affording special opportunities for studies on communication, social interactions, and learning. Increased additive genetic variances for the courtship repertoire of experimentally bottlenecked populations provided evidence for the presence of dominance and/or epistasis. Negative genetic variances in these populations suggested genotype-by-environment interactions, where the environment is the mating partner. Line cross assays of populations that had been subjected to selection for divergent courtship repertoire confirmed that both dominance and epistasis have significant effects. These crosses also showed more directly that the expression of the male's genotype is dependent upon the preferences of his mating partner. Repeatability studies also detailed how males alter their courtship performances with successive encounters within and across females, such that the males learn to improve their techniques in securing copulations. A review of 41 animal behavior studies found that a wide range of traits and taxa have dominance, epistasis, and genotype-y-environment interactions, although house fly courtship may remain a unique model where learning is an intersexually selected trait. Future development of more sophisticated molecular techniques for the M. domestica genome will help unravel the underlying biochemical and developmental pathways of these quantitative genetic interactions for a more complete understanding of the processes of inbreeding depression, outbreeding depression, and pleiotropy.

Animals↗

Breeding behavior of the American black bear Ursus americanus.

By means of videotape, we observed and timed courtship, mating, and refractoriness in the American black bear while confined in a semicaptive environment. Sixty-six courtships resulted in mating. During courtship, the male detected estrus and receptivity in the female. Courtship lasted 22.5+/-2.3s (mean+/-S.E.M.). We videotaped and reviewed 61 matings that included mounts, pelvic thrusts, and flutters. Mating lasted 51.6+/-4.2s. Within these 61 matings, there were 24 pelvic thrusts and 14 flutters observed. Pelvic thrusts averaged 2.3+/-0.3 times, while flutters averaged 11.1+/-2.9 times. Of the 61 courtship-mating segments, 28 segments depicted a male refractory period, while 27 segments depicted a female refractory period. The male refractory period lasted 44.9+/-6.1s during which the male sat and licked his genital region. The female walked, ran away, or turned and attacked the male during her 54.2+/-8.3s refractory period. This is the first time a large number of paired ursid matings has been described and timed.

Animals↗

Co-adaptation of pheromone production and behavioural responses in Drosophila melanogaster males.

In Drosophila melanogaster, male courtship behaviour is genetically controlled and is influenced by sex pheromones. 7-tricosene (7-T) induces a dose-dependent inhibition of male-male courtship, whereas 7,11-dienes stimulate male courtship of females. There is a geographical quantitative variation in the production of two predominant male hydrocarbons, 7-T and 7-pentacosene (7-P). We have previously found that 7-P, the main hydrocarbon from males of West African strains, stimulates males that mainly produce 7-T. Using both 'natural' and genetically engineered strains, we find that genetic factors coding for low levels of 7-P in males have co-evolved with factor(s) coding for male responses to high levels of 7-P. These two phenotypes are coded by factors on different chromosomes: the intraspecific polymorphism for the production of 7-T and 7-P is largely controlled by chromosome 2, whereas the variation in courtship towards 7-P-rich males is largely controlled by chromosome 3. The polymorphism of male courtship towards 7-P-rich males shows no correlation with the variation in male responses to female flies.

Alkenes↗

Change in the signal-response sequence responsible for asymmetric isolation between Drosophila planitibia and Drosophila silvestris.

Drosophila planitibia and Drosophila silvestris form a species pair that is an example of species diverged through a founder event. These species exhibit asymmetric sexual isolation, courtships between D. planitibia males and D. silvestris females being more successful than courtships between D. silvestris males and D. planitibia females. When analyzing the signal-response courtship sequence in these species, we found that D. silvestris females responded to male circling by standing or preening while D. planitibia females required further signals from the male to stop walking. The main reason for the reduced mating success rate of D. silvestris males with D. planitibia females was that the females responded to male circling by walking and the males did not proceed to the head-under-wings (HUW) position of a walking female. Another critical phase in these courtships was the HUW position in D. silvestris, where males proceeded almost immediately to wing and leg vibration. The courtships between D. planitibia male and D. silvestris female proceeded in a signal-response coordination until the male went to the HUW position, where he fanned his wings for too long a period before proceeding to wing and leg vibration. Thus, it seems that the asymmetric isolation between D. planitibia (ancestral species) and D. silvestris (derived species) is mainly due to a loss of transitions in the signal-response chain of D. silvestris. A change in the behavior of the males in the HUW position has caused further isolation between the species in both directions.

Animals↗

Reduced reproductive success for a conditioning mutant in experimental populations of Drosophila melanogaster.

Male Drosophila melanogaster that have courted newly-emerged males can modify their subsequent courtship behavior to avoid further courtship with immature males for up to 6 hr (previously reported). Here, it was hypothesized that such an experience-dependent modification would afford a mating advantage to normal males over males that carried a mutation that affects learning and memory. Coisogenic lines were constructed which varied at the dunce gene (dnc+ and dncM14 alleles) in order to test this hypothesis. Whether previously experienced with immature males or not, dnc+ and dncM14 males were indistinguishable in their response and mating efficiency when individually paired with virgin females. However, courtship performance of dnc+ and dncM14 males was different if they were first experienced with immature males and were then individually tested in an artificial population of nine immature males and one virgin female. In this situation, dnc+ males spent much less time in courtship with immature males and achieved copulation in one-third the time required for dncM14 males. As a control, the behavior and mating efficiency of courtship-naive dnc+ and dncM14 males in the artificial population was indistinguishable. In competition for a single virgin female, experienced dncM14 males showed a slight mating advantage over experienced dnc+ males. But when competition by experienced males for a single virgin female took place in the presence of nine immature males, dnc+ males were the successful maters in three-fourths of the trials.

Animals↗

Extended reproductive roles of the fruitless gene in Drosophila melanogaster revealed by behavioral analysis of new fru mutants.

The fruitless mutants fru3 and fru4 were assessed for sex-specific reproductive-behavioral phenotypes and compared to the previously reported fru mutants. Among the several behavioral anomalies exhibited by males expressing these relatively new mutations, some are unique. fru3 and fru4 males are less stimulated to court females than fru1 and fru2. No courtship pulse song is generated by either fru3 or fru4 males, even though they perform brief wing extensions. fru3 and fru4 males display significantly less chaining behavior than do fru1 males. The hierarchy of courtship responses by fru males directed toward females vs. males, when presented with both sexes simultaneously, is that fru1 males perform vigorous and indiscriminant courtship directed at either sex; fru4 males are similarly indiscriminant, but courtship levels were lower than fru1; fru2 males prefer females; fru3 males show a courtship bias toward males. fru3 and fru4 males essentially lack the Muscle of Lawrence (MOL). On several reproductive criteria, there was no difference between fru-variant females and fru+. The increases in phenotypic severity measured for the new mutants are discussed in the context of the emerging molecular genetics of fru and with regard to the gene's position within the sex-determination pathway.

Animals↗

In vitro nitric oxide effects on basal and gonadotropin-releasing hormone-induced gonadotropin secretion by pituitary gland of male crested newt (Triturus carnifex) during the annual reproductive cycle.

The objective of this study was to test the possible nitric oxide (NO) involvement in pituitary gonadotropin secretion in the male crested newt, Triturus carnifex. Pituitaries were incubated in vitro with medium alone, GnRH, NO donor (NOd, sodium nitroprusside), NO synthase inhibitor (NOSi, Nomega-nitro-L-arginine methyl ester), cGMP analogue (cGMPa, 8-bromo-cGMP), soluble guanylate cyclase inhibitor (sGCi, cystamine), GnRH plus NOSi, GnRH plus sGCi, and NOd plus sGCi during the annual reproductive cycle: pre-reproduction, reproduction (noncourtship and courtship), and the refractory, recovery, and estivation periods. To determine pituitary gonadotropin secretion indirectly, newt testes were superfused in vitro with preincubated pituitaries, and androgen release was determined. NO synthase (NOS) activity and cGMP levels were assessed in the preincubated pituitaries. Medium alone- and GnRH-preincubated pituitary increased androgen secretion during pre-reproduction, noncourtship, courtship, and recovery; the GnRH-induced increase was higher than the medium alone-induced increase during pre-reproduction, noncourtship, and recovery. NOd and cGMPa increased androgens in all reproductive phases considered except courtship; the NOd- and cGMP-induced increase was higher than the medium alone-induced increase during pre-reproduction, noncourtship, and recovery. NOS activity was highest during courtship and lowest during the refractory and estivation periods. GnRH increased NOS activity during pre-reproduction, noncourtship, and recovery. Cyclic GMP levels were highest during courtship and lowest during the refractory period and estivation. GnRH increased cGMP levels during pre-reproduction, noncourtship, and recovery, while NOd did so during all reproductive phases considered. These results suggest that basal and GnRH-induced gonadotropin secretion are up-regulated by NO in the pituitary gland of the male Triturus carnifex.

Androgens↗

'Un chant d'appel amoureux': acoustic communication in moths

Tympanal sound receptors in moths evolved in response to selective pressures provided by echolocating insectivorous bats. The presence of these ultrasound detectors also set the stage for the later evolution of ultrasonic courtship signals in the tympanate moth families. Male moths have repeatedly exploited the bat-detection mechanisms in females for the purpose of finding, identifying and obtaining mates. Ultrasonic courtship has been described in several members of the moth families Arctiidae, Noctuidae and Pyralidae, and ultrasound is predicted to play a significant role in the courtship of other tympanate moths including the Sphingidae, Lymantriidae, Notodontidae and Geometridae. Ultrasonic signals are involved in species recognition, in male-male competition for mates and in female mate-choice systems. Pre-existing motor systems, including those involved in bat defence, have also been exploited for the purpose of generating high-frequency courtship signals. Sound production mechanisms in moths include thoracic tymbals, tegular tymbals, alar castanets and genital stridulatory organs. Thus, in both their sensory and motor aspects, the weapons of bat/moth warfare have frequently evolved into components of courtship systems.

Journal Article↗

Discrimination of conspecific individuals via cuticular pheromones by males of the cricket Gryllus bimaculatus.

Cuticular substances on the body surface of crickets serve as pheromones that elicit a variety of different behaviors in male crickets. Antennal contact between males and females resulted in courtship behavior, and that between two males resulted in aggressive displays. As a first step in elucidating how crickets recognize and discriminate individuals, behavioral responses of male individuals to cuticular substances of conspecific males or females were investigated. The behavioral responses of males to antennal or palpal stimulation with an isolated antenna from a male or a female were recorded. To both antennal and palpal stimulation with female antennae, the majority of males responded with courtship behavior; to stimulation with male antennae, males responded with aggressive displays. To gain insight into the chemical nature of the behaviorally relevant components, isolated antennae were washed in either n-hexane, acetone or ethanol before behavior assays. Washed antennae no longer elicited courtship or aggressive responses in males. Next, polypropylene fibers were smeared with substances from the body surface of females and used for antennal stimulation. This experiment showed that the quality and quantity of cuticular substances appear to be highly age-dependent. Significantly more males responded with courtship behavior to cuticular substances from younger females. Isolated males generally showed higher levels of aggression than males reared in groups. Grouped males also were more likely to display courtship behavior towards antennae from younger females, and aggressive behavior towards antennae from older females. These results suggest that male discrimination of mating partners depends on the nature of female cuticular substances.

Age Factors↗

Involvement of arginine vasotocin in reproductive events in the male newt Cynops pyrrhogaster.

Effects of arginine vasotocin (AVT) on reproductive events such as courtship behavior, pheromone release, and spermatophore discharge were investigated in the male newt Cynops pyrrhogaster. AVT enhanced the incidence and frequency of androgen-induced courtship behavior. In this case, AVT was likely to act centrally because the behavior was evoked with a much smaller amount of AVT when the hormone was administered intracerebroventricularly than when given intraperitoneally. Involvement of endogenous AVT in spontaneously occurring courtship behavior was also evidenced by the fact that administration of a V1 (vasopressor) receptor antagonist, [d(CH2)5(1), Tyr(Me)2, Arg8-vasopressin] suppressed the expression of the courtship behavior. The water in which AVT-treated males had been kept showed considerable female-attracting activity as compared with the water in which saline-injected males had been kept. Moreover, the content of sodefrin, a female-attracting pheromone in the abdominal gland, was decreased by the intraperitoneal injection of AVT, suggesting that the neurohypophyseal hormone stimulated the release of sodefrin from the abdominal gland into the water. AVT induced contraction of the excised abdominal gland concentration-dependently, and, again, the V1 receptor antagonist suppressed the AVT-induced contraction. Thus, we concluded that AVT induces the pheromone discharge, acting peripherally on a contractile structure of the abdominal gland. AVT was also found to induce spermatophore deposition in the male kept in the absence of the female. Administration of the V1 receptor blocker to the sexually developed males suppressed the spermatophore deposition. All these results indicate the involvement of AVT in reproductive events acting centrally and peripherally.

Animals↗

Differential effects of 11-ketotestosterone on dimorphic traits in a teleost with alternative male reproductive morphs.

Species with dimorphic males typically have one morph (type I) associated with territoriality and courtship, and another morph (type II) associated with cuckoldry. In teleost fishes, type I males generally have higher levels of the androgen 11-ketotestosterone (KT) than type II males. Is KT causal to phenotypic differences between morphs? We investigated this question in the midshipman fish (Porichthys notatus) in which type I males have detectable levels of KT during the breeding season, whereas levels in type II males are usually undetectable. Type I midshipman will either cuckold or hold territories and court females, whereas type II males are only known to cuckold. Acoustic courtship by type I's is supported by the sonic motor nucleus, which innervates a sound-producing sonic muscle. Type I males have larger sonic motor nuclei and larger sonic muscles than type II males, consistent with the more dynamic vocal repertoire of type I's. Here, we tested whether intraperitoneal KT implants in adult type II males would induce type I male-like traits in brain, sonic muscle, and behavior. Type II's treated with KT did not differ from blank-implanted type II's in sonic motor nucleus volume. Sonic muscle mass increased in KT-implanted type II's, but did not reach the relative mass naturally observed in type I's. While neither territoriality nor courtship were induced, cuckoldry behavior intensified in KT-implanted type II's. Thus, for some but not all characters, KT exaggerated the expression of already existing type II male traits rather than inducing the type I male-like traits of territoriality, courtship, and an expansive vocal motor system.

Animals↗

Aromatase activity and regulation of sexual behaviors in the green anole lizard.

Sexual behaviors in green anoles are regulated by steroid hormones. Androgens activate the display of masculine courtship and copulatory behaviors, and estradiol activates feminine receptivity. Testosterone can also facilitate receptivity in females. The present study was conducted to test the role of converting testosterone to estradiol (aromatization) in the regulation of sexual and aggressive behaviors. Adult males and females were gonadectomized and implanted with a Silastic capsule containing either testosterone propionate (T) or estradiol benzoate (E) or with an empty (blank, BL) capsule. T- and BL-treated animals were then given injections of either Fadrozole (FAD, an aromatase inhibitor) or saline (SAL). E-treated animals received saline injections. Each individual was then tested alternately with male and female stimulus animals. Overall, T stimulated masculine sexual behaviors and receptivity, but the androgen had little effect on the display of aggressive behaviors. The inhibition of aromatase activity by treatment with Fadrozole eliminated the effect of T on receptivity. In contrast, Fadrozole treatment had no effect on mounting behavior or the frequency of courtship bouts. The inhibition of aromatase activity did increase the number of dewlap extensions (the display of a red throat fan) during courtship. These results suggest that the metabolism of testosterone to estradiol is a mechanism through which androgens can facilitate receptivity, and that such aromatization of testosterone is not required for the display of masculine sexual behaviors. In addition, females performed courtship and mounting behaviors less frequently than males, suggesting that an organizational component to steroid hormone regulation of sexual behaviors may exist in the anole.

Aggression↗

The genetics of mating recognition between Drosophila simulans and D. sechellia.

During courtship, visual and chemical signals are often exchanged between the sexes. The proper exchange of such signals ensures intraspecific recognition. We have examined the genetic basis of interspecific differences in male mating behaviour and pheromone concentration between Drosophila simulans and D. sechellia by using Drosophila simulans/D. sechellia introgression lines. Our results show a majority of quantitative trait loci (QTLs) explaining variation in both male mating behaviour and pheromone concentration to be located on the third chromosome. One QTL found on the third chromosome explains variation in time needed to start courtship and copulation as well as time spent courting. The position of such QTL (approximately 84A-88B) with effects on courtship and copulation aspects of mating includes the candidate sex determination gene doublesex (84E5-6) and Voila (86E1-2), a gene that affects male courtship in D. melanogaster. One additional third chromosome QTL explained variation in 7-tricosene pheromone concentrations among males. The interval mapping position of this QTL (approximately 68E-76E) did not overlap with the position detected for differences in mating behaviour and the intervals did not include candidate genes previously identified as having an effect on D. melanogaster cuticular hydrocarbon production. We did not detect any directionality of the effect of Drosophila sechellia allele introgressions in male mating recognition.

Alkenes↗

A Drosophila DEG/ENaC channel subunit is required for male response to female pheromones.

Odorants and pheromones as well as sweet- and bitter-tasting small molecules are perceived through activation of G protein-coupled chemosensory receptors. In contrast, gustatory detection of salty and sour tastes may involve direct gating of sodium channels of the DEG/ENaC family by sodium and hydrogen ions, respectively. We have found that ppk25, a Drosophila melanogaster gene encoding a DEG/ENaC channel subunit, is expressed at highest levels in the male appendages responsible for gustatory and olfactory detection of female pheromones: the legs, wings, and antennae. Mutations in the ppk25 gene reduce or even abolish male courtship response to females in the dark, conditions under which detection of female pheromones is an essential courtship-activating sensory input. In contrast, the same mutations have no effect on other behaviors tested. Importantly, ppk25 mutant males that show no response to females in the dark execute all of the normal steps of courtship behavior in the presence of visible light, suggesting that ppk25 is required for activation of courtship behavior by chemosensory perception of female pheromones. Finally, a ppk25 mutant allele predicted to encode a truncated protein has dominant-negative properties, suggesting that the normal Ppk25 protein acts as part of a multiprotein complex. Together, these results indicate that ppk25 is necessary for response to female pheromones by D. melanogaster males, and suggest that members of the DEG/ENaC family of genes play a wider role in chemical senses than previously suspected.

Aging↗

Regulation of behavioral and pheromonal aspects of sex determination in Drosophila melanogaster by the Sex-lethal gene.

We have shown that the Sex-lethal (Sxl) gene, which controls morphological aspects of sex determination in Drosophila melanogaster, also regulates sexual behavior. Chromosomal males that are hemizygous for a deletion of the entire Sxl locus perform normal courtship and synthesize the two courtship-inhibiting pheromones that normal males make. However, ectopic expression of female-specific Sex-lethal gene products drastically alters chromosomal males' ability to perform and elicit courtship and increases the probability that they will synthesize a courtship-stimulating pheromone or fail to synthesize one of the inhibitory pheromones. These observations suggest that male sexual behavior is a consequence of the Sxl gene's being functionally inactive in haplo-X flies.

Animals↗

Role of gonadotrophin releasing hormones in the reproductive behaviour of female ring doves (Streptopelia risoria).

This work was carried out to determine whether gonadotrophin releasing hormone(s) (GnRH) plays a role in the courtship behaviour of female ring doves. In particular, it tests the hypothesis that a high dose of oestrogen suppresses the synergistic action of GnRH with oestrogen which is required to elicit courtship behaviour. The hypothesis was formulated to provide an explanation for a finding in a previous study that ovariectomized ring doves treated with a high dose of oestrogen, paradoxically, failed to show courtship behaviour. In the present study, treatment with synthetic LH releasing hormone (RH) enabled such birds to show typical female behaviour, thus supporting the hypothesis. This LH-RH effect was not apparently mediated by LH or progesterone since treatment with LH or progesterone failed to induce courtship behaviour. In addition, LH-RH can synergise with sub-threshold doses of oestrogen to induce both nest-soliciting and squatting behaviour. This portion of the LH-RH effect complements similar results reported for the rat. Finally, the results of thyrotrophin releasing hormone point to the specificity of LH-RH and the results of an anti-ovulation LH-RH analogue agree with the LH-RH effect.

Animals↗

The neural mechanisms of mate choice: a hypothesis.

Scientists have described many physical and behavioral traits in avian and mammalian species that evolved to attract mates. But the brain mechanisms by which conspecifics become attracted to these traits is unknown. This paper maintains that two aspects of mate choice evolved in tandem: 1) traits that evolved in the "display producer" to attract mates and, 2) corresponding neural mechanisms in the "display chooser" that enable them to become attracted to these display traits. Then it discusses our (in-progress) fMRI brain scanning project on human romantic attraction, what we believe is a developed form of "courtship attraction" common to avian and mammalian species as well as the primary neural mechanism underlying avian and mammalian mate choice. The paper hypothesizes that courtship attraction is associated with elevated levels of central dopamine and norepinephrine and decreased levels of central serotonin in reward pathways of the brain. It also proposes that courtship attraction is part of a triune brain system for mating, reproduction and parenting. 1)The sex drive evolved to motivate birds and mammals to court any conspecifics. 2) The attraction system evolved to enable individuals to discriminate among potential mating partners and focus courtship activities on particular individuals, thereby conserving mating time and energy. 3) The neural circuitry for attachment evolved to enable individuals to complete species-specific parental duties.

Brain Mapping↗