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Vaginal scent marking: effects on ultrasonic calling and attraction of male golden hamsters.

Male hamsters were tested for their responses to areas that had been scent marked by intact or vaginectomized females to determine the effects of naturally deposited vaginal secretions on male behavior. In the first experiment males produced more ultrasonic courtship calls when investigating areas marked by intact females than areas scented by vaginectomized females, demonstrating that vaginal marks facilitate such calling. In a wind-tunnel preference test situation in which scent-marked alleys and clean alleys served as sources of odor, males approached the scented alley first if it had been freshly marked by intact females but not if it had been scented by vaginectomized females or other males. Thus, the odors of vaginal marks are sufficient to attract males over short distances. After males entered these alleys they showed a preference for odors of both intact and vaginectomized females over no odors, but still spent significantly more time investigating the odors of intact females than those of vaginectomized females. These experiments indicate that vaginal secretions are one of the primary cues that elicit male courtship calling, and the small quantities of vaginal secretions deposited by females in vaginal marks are sufficient to elicit ultrasonic calling and attract males over short distances. Thus it is likely that vaginal scent marking and ultrasonic calling by females interact to facilitate attraction and location of mates during courtship.

Animals↗

Motivational control of sexual behavior in Aplysia fasciata: sequencing and modulation by sexual deprivation and by addition of partners.

Variables similar to those affecting feeding behavior also modulate sexual behavior in Aplysia fasciata, indicating that mating is under motivational control. Motivated behaviors are often patterned into appetitive and consummatory components. Courtship, the appetitive phase of male sexual behavior, was temporally related to subsequent mating. Although many bouts of courtship failed to lead to mating, most incidents of mating were preceded by courtship. Motivation is also characterized by satiation after the goal of the behavior is achieved. We found an increase in likelihood to mate following a period of sexual isolation. Motivated behaviors are also modulated by environmental stimuli that induce arousal. Time spent mating was found to be a function of the number of potential mates accessible for mating. Number of A. fascinata participating in a mating group was also found to be a function of number of potential mates available.

Animals↗

Songs of Holcostethus strictus (Fabricius): a different repertoire among landbugs (Heteroptera: Pentatomidae).

Songs emitted during mating by male and female Holcostethus strictus were recorded as substrate vibrations. Spectra of the vibrational signals have a dominant frequency peak between 100 and 260 Hz and in this respect reflect the general characteristic of the family Pentatomidae. Songs of H. strictus differ from the song repertoire of the southern green stink bug Nezara viridula (L.) (Pentatomidae) in many respects. The female calling and courtship songs differ in echeme and phrase duration. The male calling song is composed of spectrally different subunits. The male courtship song is characterised by three types of spectrally and temporally different echemes. The male copulatory song is composed of echemes of two types, which constitute a phrase of less regular temporal structure. In H. strictus, males start to sing first and female songs are less complex than in N. viridula. The female calling song is evoked by male calling and does not trigger male response. The female and male courtship song phrases are superimposed on one another and we have not observed any obvious regularity in their exchange. The possible role of different songs in H. strictus is discussed and compared with that in other pentatomide landbug species.

Journal Article↗

Sexual behaviour of Zebu bulls in the humid tropics of Costa Rica: single versus multiple-sire groups.

This study compares the courtship and mounting behaviour of Bos indicus bulls in single- and multiple-sire groups in Costa Rica (latitude 10 degrees 25'N, longitude 84 degrees 32'W, annual precipitation of 3096mm, temperature of 24 degrees C and humidity of 85.3%). Four, 3-4 year-old Brahman bulls with previous sexual experience were used to sire a group of 120 multiparous cows (average of 128 days post-partum and a body condition score of 2.5) allocated to two groups of 60 each: (1) single-sire mating group (SSM) and (2) multiple-sire mating group of three bulls (MSM). Bulls were rotated among groups every 7 days for 28 days. The frequency, type and duration of sexual activities (mounting and mounting attempts) and courtship activities (smelling and licking genital area, butting, supporting the head over a female and the sign of Flehmen) were calculated for each mating group. Descriptive and non-parametric statistics (Wilcoxon, Mann-Withney tests) were used to calculate differences between mating programmes. Sexual activities tended to be more frequent in the SSM group than the MSM group (267 versus 124, P>0.05). Differences in the ratios of sexual to courtship activities between both groups were significant (1:3 in SSM and 1:6 in MSM, P>0.05). Pregnancy rate averages were 28 and 37%, respectively, (P>0.05). It is concluded that under these conditions multiple-sire mating and single-sire mating achieved similar pregnancy rates.

Animals↗

Distribution of aromatase immunoreactivity in the forebrain of red-sided garter snakes at the beginning of the winter dormancy.

Until recently, it has been difficult to identify the exact location of aromatase containing cells in the brain. The development of new antibodies has provided a sensitive tool to analyze the distribution of aromatase immunoreactive (ARO-ir) material at a cellular level of resolution. In the present study we examined, for the first time, the distribution of ARO-ir cells in the brain of a reptile, the red-sided garter snake, at the beginning of the winter dormancy. ARO-ir cells were found at all rostro-caudal levels in the red-sided garter snake brain. Although weakly stained cells were distributed throughout the brain, more intensely immunoreactive cells were primarily concentrated in the preoptic area, anterior hypothalamus, septum and nucleus sphericus. Although androgens are elevated upon emergence from hibernation in the male red-sided garter snake, initiation of courtship behavior appears to be independent of direct androgen control. To date, the only known stimulus found to initiate courtship is a period of low temperature dormancy followed by exposure to warm temperatures. Circumstantial data, however, suggest an indirect role in the activation of male copulatory behavior for estrogenic metabolites of testosterone produced in the brain by aromatization during the winter dormancy. This study provides the first documentation of the distribution of ARO-ir cells in a reptilian species and demonstrates that while the aromatase enzyme occurs in most regions of the brain, the ARO-ir cells that appear to contain the highest concentration of enzyme are clustered in brain areas classically associated with the control of courtship behavior and mating in vertebrates. These data are consistent with the idea that estrogens locally produced in the brain may participate in some way to the activation of sexual behavior in this species also. This notion should now be experimentally tested by analyzing annual changes in aromatase activity and immunoreactivity and assessing the effects of pharmacological blockade of the enzyme activity at different times of the year.

Animals↗

Overexpression of a CREB repressor isoform enhances the female sexual receptivity in Drosophila.

The cAMP responsive transcriptional factor, CREB, is highly conserved among animal species, and its activity affects their behavior. In Drosophila melanogaster, one of alternatively spliced products of the CREB gene, dCREB2-a, is a transcriptional activator, while another isoform, dCREB2-b, is a repressor of dCREB2-a. Here, we demonstrate that overexpression of dCREB2-b in virgin females enhances their sexual receptivity. We studied the role of dCREB2 in female mating behavior using two transgenic lines, hs-dCREB2-a and hs-dCREB2-b, which overexpress respective products with heat-shock treatment. Wild-type males started their courtship behavior and mated more quickly with heat-shocked hs-dCREB2-b females than with non-heat-shocked hs-dCREB2-b females. Overexpression of dCREB2-a in females affected neither their courtship behavior nor mating frequency. The effects of overexpressed dCREB2-b were not due to elevated locomotor activities of heat-shocked females nor due to more vigorous courtship behavior of paired wild-type males. CREB might be involved in female sexual behavior of animals.

Animals↗

Social malleability in cowbirds: new measures reveal new evidence of plasticity in the eastern subspecies (Molothrus ater ater).

The social experiences of young Molothrus ater ater cowbirds were manipulated in a 2-year study. In the 1st year, males were housed with pairs of canaries. The males were tested in 3 social contexts. Also, vocal repertoires were recorded and played back to females. In contrast to a previous study of the M. a. artemisiae subspecies, the males did not vocalize to the canaries in courtship tests (T. M. Freeberg, A. P. King, & M. J. West, 1995) but showed incompetent courtship of female cowbirds. In their 2nd year, half of the males were housed with older males and female cowbirds, and half were housed with only females. Those exposed to older males courted much more successfully than did those deprived of such experience. All males developed new repertoires, and song potencies did not correlate across years. The data reveal intraspecific variation in the ontogeny of mate recognition but intraspecific dependence on social learning to acquire courtship skills.

Animals↗

Period-homologous sequence polymorphisms in subterranean mammals of the Spalax ehrenbergi superspecies in Israel.

Restriction fragment length polymorphism (RFLPs) of the mouse period-homologous sequence were studied in 13 populations of the four chromosomal species (2n = 52, 54, 58 and 60) of the mole rat, Spalax ehrenbergi superspecies in Israel. The period locus of Drosophila melanogaster is implicated in controlling the circadian rhythm as well as the male courtship song rhythm. Multiple DNA homologies exist in the mole rat and correspond to more than 10 loci. The level of polymorphism is very high, with a large number of alleles per locus, increasing from the northern to the southern species along a gradient of increasing aridity. Variation was also found in an isolated desert population, with a unique fragment specific to this population. Fragment variation allows distinction between chromosomal species, and confirms earlier evidence that gene flow does not occur between them. A correlation was found between some allelic fragments and the number of apparent harmonics of the courtship calls. This finding suggests an interesting testable hypothesis that the existence of a locus (homology) is responsible for the courtship call parameters.

Animals↗

Molecular evolution of the cacophony IVS6 region in sandflies.

A number of insects produce acoustic signals during courtship. Genes involved in the control of the courtship song are particularly interesting from an evolutionary viewpoint because interspecific variation in this signal is potentially important as a reproductive isolation mechanism and, as a consequence, in the speciation process. The cacophony gene was identified by a mutation affecting the "lovesong" in Drosophila melanogaster. Phlebotomine sandflies (Diptera: Psychodidae) also produce acoustic stimuli during courtship and therefore cacophony can be used as an interesting molecular marker in evolutionary studies in these important disease vectors. In this paper we have studied the molecular evolution of the IVS6 region of cacophony in sandflies. We compared the level of divergence in the exon sequences encoding this conserved domain in Drosophila and Phlebotomines. We also analysed the high level of variation in an intron that is present in sandflies but that was lost in Drosophila during evolution. The available cacophony sequences were also used for a phylogenetic analysis of some species of the Neotropical genus Lutzomyia.

Amino Acid Sequence↗

Signal efficacy and the evolution of male dimorphism in the jumping spider, Maevia inclemens.

An extreme form of male dimorphism occurs in the jumping spider, Maevia inclemens, where two male morphs differ dramatically in both morphology and behavior. We examined the visual signals transmitted to females by the two different morphs during courtship display. Even though the display of each male is different, as a function of behavior and courtship distance, the males present females with images that are almost identical in height and visual target area. By substituting live males with computer-animated sequences of male courtship behavior, the male morphs were superimposed upon each other, and females were determined to have two distinct male-recognition templates. We hypothesize that the dimorphic males of this species evolved through sensory exploitation and selection for alternative male strategies that stimulate different fields of the female visual range.

Journal Article↗

Philopatry of male marine turtles inferred from mitochondrial DNA markers.

Recent studies of mitochondrial DNA (mtDNA) variation among marine turtle populations are consistent with the hypothesis that females return to beaches in their natal region to nest as adults. In contrast, less is known about breeding migrations of male marine turtles and whether they too are philopatric to natal regions. Studies of geographic structuring of restriction fragment and microsatellite polymorphisms at anonymous nuclear loci in green turtle (Chelonia mydas) populations indicate that nuclear gene flow is higher than estimates from mtDNA analyses. Regional populations from the northern and southern Great Barrier Reef were distinct for mtDNA but indistinguishable at nuclear loci, whereas the Gulf of Carpentaria (northern Australia) population was distinct for both types of marker. To assess whether this result was due to reduced philopatry of males across the Great Barrier Reef, we determined the mtDNA haplotypes of breeding males at courtship areas for comparison with breeding females from the same three locations. We used a PCR-restriction fragment length polymorphism approach to determine control region haplotypes and designed mismatch primers for the identification of specific haplotypes. The mtDNA haplotype frequencies were not significantly different between males and females at any of the three areas and estimates of Fst among the regions were similar for males and females (Fst = 0.78 and 0.73, respectively). We conclude that breeding males, like females, are philopatric to courtship areas within their natal region. Nuclear gene flow between populations is most likely occurring through matings during migrations of both males and females through nonnatal courtship areas.

Animals↗

dissatisfaction, a gene involved in sex-specific behavior and neural development of Drosophila melanogaster.

Few mutations link well defined behaviors with individual neurons and the activity of specific genes. In Drosophila, recent evidence indicates the presence of a doublesex-independent pathway controlling sexual behavior and neuronal differentiation. We have identified a gene, dissatisfaction (dsf), that affects sex-specific courtship behaviors and neural differentiation in both sexes without an associated general behavioral debilitation. Male and female mutant animals exhibit abnormalities in courtship behaviors, suggesting a requirement for dsf in the brain. Virgin dsf females resist males during courtship and copulation and fail to lay mature eggs. dsf males actively court and attempt copulation with both mature males and females but are slow to copulate because of maladroit abdominal curling. Structural abnormalities in specific neurons indicate a role for dsf in the differentiation of sex-specific abdominal neurons. The egg-laying defect in females correlates with the absence of motor neuronal innervation on uterine muscles, and the reduced abdominal curling in males correlates with alteration in motor neuronal innervation of male ventral abdominal muscles. Epistasis experiments show that dsf acts in a tra-dependent and dsx-independent manner, placing dsf in the dsx-independent portion of the sex determination cascade.

Abdomen↗

Cryptic forcible insemination: male snakes exploit female physiology, anatomy, and behavior to obtain coercive matings.

Whether males can inseminate uncooperative females is a central determinant of mating system evolution that profoundly affects the interpretation of phenomena such as multiple mating by females, mate choice, reproductive seasonality, and courtship tactics. Forcible insemination is usually inferred from direct physical battles between the sexes and has been dismissed on intuitive grounds for many kinds of animals. For example, snakes have elongate flexible bodies (making it difficult for a male to restrain a female physically), males are typically smaller than females, and copulation requires female cloacal gaping to enable intromission. Male garter snakes (Thamnophis sirtalis) do not display any overt aggression during courtship and simply lie over the female and exhibit rhythmic pulsating caudocephalic waves of muscular contraction; previous studies have interpreted this behavior as a mechanism for eliciting female receptivity. In contrast, we show that male garter snakes forcibly inseminate females. They do so by taking advantage of specific features of snake physiology, respiratory anatomy, and antipredator behavior. The snake lung extends along most of the body, with the large posterior section (the saccular lung) lacking any respiratory exchange surface. Rhythmic caudocephalic waves by courting male garter snakes push anoxic air from the saccular lung forward and across the respiratory surfaces such that females cannot obtain oxygen. Their stress response involves cloacal gaping, which functions in other contexts to repel predators by extruding feces and musk but in this situation permits male intromission. Thus, superficially benign courtship behaviors may involve cryptic coercion even in species for which intuition dismisses any possibility of forcible insemination.

Animals↗

The effect of transformer, doublesex and intersex mutations on the sexual behavior of Drosophila melanogaster.

We have identified the effects of genes that regulate sex determination on female-specific tissues in the abdomen that produce sex pheromones and parts of the central nervous system that function when a male performs courtship. To do this, we monitored the sexual behaviors of flies with mutations in the transformer (tra), doublesex (dsx) and intersex (ix) genes. Except for tra, which transforms diplo-X flies so that they look and function like normal males, these mutations do not have the same effect on pheromone-producing tissues and the central nervous system as they do on the appearance of the fly. The dsx and ix mutations, which make diplo-X-flies look like intersexes, do not transform the flies so that they can perform courtship, suggesting that these genes do not regulate the development of sex-specific parts of the central nervous system. Conversely, the ix mutation, which has no effect on the appearance of haplo-X flies, makes the flies sexually attractive and impairs their ability to perform courtship, which implies that the ix gene is active in internal tissues of males.

Animals↗

The nonA gene in Drosophila conveys species-specific behavioral characteristics.

The molecular basis of species-specific differences in courtship behavior, a critical factor in preserving species boundaries, is poorly understood. Genetic analysis of all but the most closely related species is usually impossible, given the inviability of hybrids. We have therefore applied interspecific transformation of a single candidate behavioral locus, no-on-transient A (nonA), between Drosophila virilis and D. melanogaster, to investigate whether nonA, like the period gene, might encode species-specific behavioral information. Mutations in nonA can disrupt both visual behavior and the courtship song in D. melanogaster. The lovesong of nonA(diss) mutant males superficially resembles that of D. virilis, a species that diverged from D. melanogaster 40-60 mya. Transformation of the cloned D. virilis nonA gene into D. melanogaster hosts carrying a synthetic deletion of the nonA locus restored normal visual function (the phenotype most sensitive to nonA mutation). However, the courtship song of transformant males showed several features characteristic of the corresponding D. virilis signal, indicating that nonA can act as a reservoir for species-specific information. This candidate gene approach, together with interspecific transformation, can therefore provide a direct avenue to explore potential speciation genes in genetically and molecularly tractable organisms such as Drosophila.

Animals↗

Male coercion and convenience polyandry in a calopterygid damselfly.

Copulation in odonates requires female cooperation because females must raise their abdomen to allow intromission. Nevertheless in Calopteryx haemorrhoidalis haemorrhoidalis (Odonata) males commonly grasp ovipositing females and apparently force copulations. This has been interpreted as a consequence of extreme population density and male-male competition. We studied this behavior at two sites on a river that had different densities over three years. As predicted, at high densities most matings were forced (i.e. not preceded by courtship), but at low density most were preceded by courtship. Courtship matings were shorter at high density, but density did not affect the duration of forced matings. Females cooperated in forced matings even if they had very few mature eggs. Furthermore, females mated more times if they experienced higher male harassment during oviposition, and at low density second and subsequent matings were more likely to be forced. We interpret these results to mean that females engage in "convenience polyandry", because they gain more by accepting copulation than by resisting males. The results also suggest that females might trade copulations for male protection, because under extreme population density harassment by males is so intense that they can impede oviposition.

Abdomen↗

Sexual selection and mating advantages in the giant sperm species, Drosophila bifurca.

Mate choice may be exercised by either sex; however, females are generally choosier than males because they invest more in their gametes. Female choice is often based on direct benefits, such as better reproductive output, whereas male choice appears to be based on differences in female fecundity. However, when gamete production is limited, sexual selection theories predict that mate choice will be decisive for reproductive success in both sexes. Here, we investigate how mating advantage is achieved in Drosophila bifurca, a giant sperm species in which both sexes produce only a few gametes. Our initial expectations were as follows: (1) females would discriminate against sperm-depleted males to avoid fertility cost; and (2) males would discriminate against inseminated females to reduce sperm competition and increase the assurance of paternity of individual gametes. Differences in courtship behaviors were analyzed with regard to the sexual maturity, which is reached after 22 days in males at 21 degrees C, and the reproductive history of both sexes (inseminated versus virgin for females, and sperm-depleted versus sperm-loaded for males). Our results show that: (1) sexual immaturity precludes mating in both sexes; (2) virgin females do not discriminate between sperm-loaded and sperm-depleted males, and (3) males mate preferentially with virgin females, because inseminated females fend off the male, which tended to bring male courtship to an end. Female remating was limited, but increased significantly when the first male was sperm-depleted. Contrary to our initial expectations, these findings suggest that male sperm depletion does not affect female mating preference, whereas the success of male courtship is driven by female behavior. The possibility that female remating was only promoted in response to low sperm transfer is discussed in relation to the gametic system of this species.

Animals↗

Evolution of female remating behaviour following experimental removal of sexual selection.

The relatively small number of ova produced by a female can be fertilized by a single ejaculate in most species. Why females of many species mate with multiple males is therefore enigmatic, especially given that costs associated with remating have been well documented. Recently, it has been argued that females may remate at a maladaptive rate as an outcome of sexually antagonistic coevolution: the evolutionary tug-of-war between manipulation by one sex and resistance to being manipulated by the other sex. We tested this hypothesis experimentally for the evolution of the female remating interval in a naturally promiscuous species, Drosophila melanogaster. In two replicate populations, sexual selection was removed through enforced monogamous mating with random mate assignment, or retained in polyandrous controls. Monogamy constrains the reproductive success of mates to be identical, thereby converting prior conflicts between mates into opportunities for mutualism. Under these experimental conditions, the sexually antagonistic coevolution hypothesis generates explicit predictions regarding the direction of evolutionary change in female remating behaviour. These predictions are contingent upon the mechanism of male manipulation, which may be mediated biochemically by seminal fluids or behaviourally by courtship. Levels of divergence in female remating interval across lines, and in male ejaculatory and courtship effects on female remating, were quantified after 84 generations of selection. Data refute the hypothesis that the evolutionary change in female remating behaviour was due to sexually antagonistic coevolution of courtship signal and receiver traits. The data were, however, consistent with a hypothesis of sexual conflict mediated through ejaculate manipulation. Monogamy-line males evolved ejaculates that were less effective in inducing female non-receptivity and monogamy-line females evolved to remate less frequently, symptomatic of lowered resistance to ejaculate manipulation. The consistency of the results with alternative hypotheses to explain female promiscuity are discussed.

Animals↗