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Schistosoma mansoni: a chemoattractive factor released by males and its receptor in females.

Chemoattraction between developing (23- to 28-day-old) and adult (10- to 12-week-old) Schistosoma mansoni males and females has been described previously. We report here on in vitro attraction of mature worms to released products of worms of the opposite sex. In the absence of a stimulus worms migrated randomly and did not show any preference in their orientation. Males were not significantly attracted to released products of females. Females exhibited greatest attraction to released products of males, and some attraction to the lipid fraction of released products of males. The aqueous fraction of male-released products slightly repelled females. Prior exposure to released products of males rendered females unresponsive to subsequent exposure, suggesting receptor saturation. These findings have important potential implications for the control of schistosomiasis by intercepting attraction and mating between males and females.

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Incipient speciation by sexual isolation in Drosophila: concurrent evolution at multiple loci.

Drosophila melanogaster from Zimbabwe and nearby regions shows strong but asymmetric sexual isolation from its cosmopolitan counterparts. By creating stable chromosome-substitution lines, earlier studies were able to show that the two major autosomes have very large effects on both male mating success and female mating preference. In this study, we genetically dissect this sexual isolation by recombination analysis between a whole-chromosome substitution line (which carries a Zimbabwe-derived third chromosome) and a strain with seven visible markers on that chromosome. Four loci are responsible for male mating success and three others are found to control female mating preference. Because male and female traits are not closely linked, their strong association among isofemale lines is most likely a reflection of sexual selection in nature. The results suggest that a large number of behavioral loci may evolve concurrently in the incipient stage of speciation before other aspects of reproductive isolation (such as hybrid sterility) have become evident. The results shed light on the population genetic processes underlying the formation of nascent species, as well as modes of speciation.

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What initiates speciation in passion-vine butterflies?

Studies of the continuum between geographic races and species provide the clearest insights into the causes of speciation. Here we report on mate choice and hybrid viability experiments in a pair of warningly colored butterflies, Heliconius erato and Heliconius himera, that maintain their genetic integrity in the face of hybridization. Hybrid sterility and inviability have been unimportant in the early stages of speciation of these two Heliconius. We find no evidence of reduced fecundity, egg hatch, or larval survival nor increases in developmental time in three generations of hybrid crosses. Instead, speciation in this pair appears to have been catalyzed by the association of strong mating preferences with divergence in warning coloration and ecology. In mate choice experiments, matings between the two species are a tenth as likely as matings within species. F1 hybrids of both sexes mate frequently with both pure forms. However, male F1 progeny from crosses between H. himera mothers and H. erato fathers have somewhat reduced mating success. The strong barrier to gene flow provided by divergence in mate preference is probably enhanced by frequency-dependent predation against hybrids similar to the type known to occur across interracial hybrid zones of H. erato. In addition, the transition between this pair falls at the boundary between wet and dry forest, and rare hybrids may also be selected against because they are poorly adapted to either biotope. These results add to a growing body of evidence that challenge the importance of genomic incompatibilities in the earliest stages of speciation.

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Distribution of putative male sex pheromones among Lutzomyia sandflies (Diptera: Psychodidae).

Male Lutzomyia longipalpis produce terpene sex pheromones in glandular tissue underlying the cuticle. The pheromones are transmitted to the surface via cuticle-lined ducts (measuring 0.25 microm in diameter), each of which reaches the surface in the centre of a papule (measuring 3-3.5 microm in diameter). Similar papules, in a range of shapes but all characterized by the presence of a central pore and absence of macroserae, occur in some other species of sandfly. The aim of the present study was to determine the distribution of sex pheromones in sandflies of the genus Lutzomyia that do and do not have the papules. The results indicate that sex pheromones are not widely distributed amongst male Lutzomyia spp. Male members of the genus can be subdivided into three groups: those that produce terpenes and have cuticular papules; those that do not produce terpenes but still have the associated papules; and those that have neither terpenes nor papules. The papules seen in the species that do not synthesise sex pheromones are presumably vestigial, non-functional structures. Such species may have stopped producing pheromone as the result of changes in the way in which the females found and selected mates or changing feeding preferences. A similar event has occurred in the Lepidoptera, where vestigial pheromone-secreting structures remain in some species which no longer produce pheromone. Lutzomyia lenti collected in southern Brazil produced a novel diterpene whereas male L. lenti from north-eastern Brazil did not, supporting suggestions by others that L. lenti is, like L. longipalpis, a species complex.

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Overt female mate competition and preference for central males in a lekking antelope.

In mammals, there exists only scant evidence of female mate choice in species mating on arenas, so-called leks. This has led to hypotheses of lek evolution that are based on benefits to females from reduced harassment by males, low predation risk, or improved availability of scarce nutrients. Here I report that female topi antelopes (Damaliscus lunatus) compete aggressively for matings with preferred males on central lek territories. Females fight at higher rates and more likely disrupt mating attempts of others in the lek center than elsewhere. Contrary to the predictions of the alternative hypotheses, food resources were insignificant, and harassment levels and estimated predation risk were higher on than off lek. These results clearly demonstrate female competition for mates in a lekking mammal in which a female chooses between males for the sole purpose of mating. The finding suggests that the forces leading to lek evolution in mammals and birds may be more similar than previously acknowledged.

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Gender effects on aggression, dominance rank, and affiliative behaviors in a flock of captive adult cockatiels (Nymphicus hollandicus).

Agonistic and affiliative behaviors and spatial positioning were studied in a small psittacine species, Nymphicus hollandicus. Subjects studied were flock-housed breeders including five hens and seven cocks. Fifteen-minute focal animal samples were collected for the entire flock in a randomly distributed order during mate selection and the onset of the breeding season. All agonistic behaviors were recorded, including the winner and loser of each interaction, along with allopreening and copulation behavior. Point samples were recorded every 60s to determine social spacing. Rates of aggression were significantly higher for male cockatiels than female cockatiels. Results based on dyadic agonistic interactions showed males to rank significantly higher in the social hierarchy than females. Associations within the flock were not random. Individual birds associated more with specific birds than would be predicted by chance. Analysis of spatial data revealed that there were both same-sex and opposite-sex preferred associations within the flock. Male cockatiels engaged in allopreening behavior with females significantly more than with other males. Observations of breeding and nesting behaviors revealed pairs, a triad, and extra-pair mating.

Journal Article↗

The evolution of female mate choice by sexual conflict.

Although empirical evidence has shown that many male traits have evolved via sexual selection by female mate choice, our understanding of the adaptive value of female mating preferences is still very incomplete. It has recently been suggested that female mate choice may result from females evolving resistance rather than attraction to males, but this has been disputed. Here, we develop a quantitative genetic model showing that sexual conflict over mating indeed results in the joint evolution of costly female mate choice and exaggerated male traits under a wide range of circumstances. In contrast to tradition explanations of costly female mate choice, which rely on indirect genetic benefits, our model shows that mate choice can be generated as a side-effect of females evolving to reduce the direct costs of mating.

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Effect on growth and reproduction of hormone immersed and masculinized fighting fish Betta splendens.

To produce all-male progenies in the fighting fish, Betta splendens, six groups of fry were subjected to discrete immersion treatment at different 17alpha-methyltestosterone (MT) doses (viz. 100, 200, 500, 700, 900, and 1,000 microg/l) for a constant duration (3 hr/day) and frequency (second, fifth, and eighth day after hatching). The treatment at 900 microg/l led to 98% masculinization and 71% survival at sexual maturity. Treated groups, which showed significant deviation from the 1:1 sex ratio, were classified into two different series: S1 and S2. The groups that showed nearly cent-percent masculinization were classified as S1, and the other groups were classified as S2. The S1 males showed remarkably slower growth and attained 3.5 cm total length compared to 6.0 cm attained by a normal male. The S2 males attained 5.4 cm total length. Apart from these morphological defects, both S1 and S2 males suffered functional (decreased sperm count and sperm motility) and behavioral defects (incomplete embracing during mating) in their reproductive ability, leading to approximately 50% and 30% reduction in fecundity per mating, respectively. The cumulative fecundity loss suffered by the S1 male during its active reproductive phase is discussed. When normal and sex-reversed males were presented, a female preferred the former. Progeny testing of the sex-reversed males showed the occurrence of 12.75% males, indicating the possible role of autosomal genes in the sex determination mechanism of this species. Discrete immersion treatment at optimal/super-optimal doses ensured not only a higher percentage of masculinization, but also a higher frequency of homogametic males (XX).

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Ventral tegmental area involvement in pair bonding in male prairie voles.

Dopamine is known to play a critical role in social attachment in monogamous voles. However, little is known about the neurochemical regulation of central dopamine release during pair bond formation. Here we examine the effects on partner preference formation in male prairie voles of neurochemical manipulations in the ventral tegmental area (VTA), a major source of dopamine to brain regions implicated in pair bonding. Administration of NBQX, an AMPA receptor antagonist, or bicuculline, a GABA receptor antagonist, into the VTA induced partner preferences within 6 h in the absence of mating. We also found that, after unilateral administration of NBQX into the VTA, neuronal activation, as indicated by the expression of the immediate early gene c-fos, was decreased in the nucleus accumbens, prefrontal cortex, and medial amygdala, but was unchanged in the lateral septum and in a control region, the arcuate nucleus. These results confirm a role for the VTA in partner preference formation in monogamous voles and extend the list of neurochemicals important in pair bonding to include glutamate and GABA.

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Effects of vomeronasal organ removal on olfactory sex discrimination and odor preferences of female ferrets.

Previous research suggests that body odorants, including anal scents and urinary odors, contribute to sex discrimination and mate identification in European ferrets of both sexes. We assessed the possible role of the vomeronasal organ (VNO) in these functions by surgically removing the organ bilaterally in sexually experienced female ferrets. Lesioned (VNOx) and sham-operated control (VNOi) females reliably discriminated between male- and female-derived anal scent gland as well as fresh urinary odors in habituation/dishabituation tests. However, VNOi females spent significantly more time than VNOx subjects investigating male urinary odors in these tests. Also, VNOi females, but not VNOx subjects, preferred to investigate day-old male versus female urine spots as well as wooden blocks that had previously been soiled by male versus female ferrets. Both groups of female ferrets preferred to approach volatile odors from a breeding male instead of an estrous female in Y-maze tests and both groups showed similar levels of receptive sexual behavior in response to a male's neck grip. The VNO is apparently not required for olfactory sex discrimination or mate recognition in this carnivore, but instead may play a role in promoting continued contact with nonvolatile body odors previously deposited by opposite-sex conspecifics during territorial scent marking.

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Degree of male ornamentation affects female preference for conspecific versus heterospecific males.

Several studies have shown female preference for conspecific males with the attached artificial ornaments of more elaborate heterospecifics. However, preference for heterospecifics under natural conditions is relatively rare. We tested what factors affect behavioural mechanisms of species isolation using three species of estrildid finch (genus Uraeginthus) that occur in both sympatry and allopatry. These finches differ in degree of sexual dimorphism; male ornamentation; behavioural and morphological similarity; and phylogenetic distance. Paired mate-choice trials were used in which females were presented with a conspecific and heterospecific male to test which of the above between-species differences best predicted the degree of premating isolation. The three species differed in the degree of species-specific mate preference shown. Females from the brighter two species discriminated against dull males, independently of sympatry-allopatry, similarity and phylogenetic distance. Females from the dull species reacted to conspecific males and brighter heterospecific males equally strongly, independently of similarity and phylogenetic distance. In contrast to previous studies, an equal preference for heterospecific and conspecific males was found under natural conditions. It is suggested that differences between closely related species in male ornamentation affect the likelihood that premating isolation will occur due to the fact that sexual selection tends to drive preferences for exaggerated ornamentation.

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The effects of sex, reproductive condition and context on discrimination of conspecific odours by giant pandas.

To elucidate some of the functions of chemical communication in giant pandas, Ailuropoda melanoleuca, we systematically exposed captive males, oestrous and nonoestrous females to each others' odours by providing access to one another's temporarily unoccupied enclosures. In comparison with control observations, pandas from all reproductive categories displayed a dramatic increase in several measures of chemosensory responsiveness, both when visiting another panda's enclosure and when encountering scent deposited in their home enclosure. Evidence for discrimination between odours encountered as a resident versus a visitor was minimal, but male pandas' response patterns were suggestive of a territorial function. We also found evidence for discrimination of sex and reproductive condition via chemical cues. Males showed a marked preference for female odours, investigating, licking, scent marking and vocalizing more in response to female than male odours. Males also vocalized more in response to oestrous than nonoestrous female odours. Nonoestrous females licked more and oestrous females vocalized more when encountering male than female odours. Our data on vocalizations suggest a potential role for odours in the activation of sexual motivation. We argue that the low reproductive success observed in captivity may be attributed in part to failure to provide sufficient opportunities for chemical communication, and encourage the judicious management of social odours to promote natural mating (e.g. using intersexual odour exposure to increase libido and decrease aggressiveness prior to mating introductions). Copyright 2000 The Association for the Study of Animal Behaviour.

Journal Article↗

Sexual incentive motivation, olfactory preference, and activation of the vomeronasal projection pathway by sexually relevant cues in non-copulating and naive male rats.

There are some apparently healthy male rats that fail to mate after repeated testing with receptive females. We have previously shown that these "non-copulator (NC)" males show no partner preference for a receptive female when given the opportunity to physically interact with a sexually receptive female or a sexually active male. We also demonstrated that although NC males prefer odors from estrous females to odors from anestrous females, this preference is significantly reduced in comparison to the preference displayed by copulating (C) males. The aim of the present study was to evaluate in NC males sexual incentive motivation, that is, the approach behavior of male rats to either a sexually receptive female or a sexually active male in a test where the subjects can smell, hear, and see the stimulus animal but prevents their physical interaction. In addition, we determined whether NC rats have alterations in their ability to detect odors from conspecifics or odors related to food. In the detection of odors from conspecifics, we determined if these NC males are sexually attracted toward odors from receptive females or sexually active males. For food-related odors, we quantified the time it took the subjects to locate a hidden a piece of apple. Finally, using the induction of Fos-immunoreactivity (Fos-IR) as an index of neuronal activation, we compared the response of the vomeronasal projection pathway (VN pathway) of C and NC male rats exposed to estrous bedding. Males without sexual experience (WSE) were included in all experiments to determine the importance of previous heterosexual experience in the different behavioral tests and in the activity of the VN pathway. In the sexual incentive motivation test, we found that C and WSE male rats have a clear preference for estrous females over sexually active males, whereas NC male rats showed no preference. In odor tests, our results showed that C males had a clear preference for odors from estrous females as opposed to odors from sexually active males. Although NC and WSE male rats showed a preference for estrous female odors, this preference was significantly reduced compared to that shown by C males. No differences were found between WSE, C, and NC males in the detection of stimuli associated with food-related odors. A significant increase in Fos-IR was observed in the mitral cell layer of the accessory olfactory bulb in all groups when exposed to estrous bedding. However, only the C male rats exposed to estrous female bedding showed an increase Fos-IR in all structures of the VN pathway. An increase in Fos-IR was observed in the medial preoptic area (MPOA) of WSE males exposed to estrous bedding. No increases in Fos-IR were detected along the VN pathway in NC male rats. We proposed that NC male rats do not display sexual behavior due to a reduced sexual motivation that could be caused by alterations in the neuronal activity of the VN pathway during the processing of estrous odors.

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The evolution of song repertoires and immune defence in birds.

Song repertoires (the number of different song types sung by a male) in birds provide males with an advantage in sexual selection because females prefer males with large repertoires, and females may benefit because offspring sired by preferred males have high viability. Furthermore, males with large repertoires suffer less from malarial parasites, indicating that a large repertoire may reflect health status. We hypothesize that sexual selection may cause a coevolutionary increase in parasite virulence and host immune defence because sexual selection increases the risk of multiple infections that select for high virulence. Alternatively, a female mate preference for healthy males will affect the coevolutionary dynamics of host-parasite interactions by selecting for increased virulence and hence high investment by hosts in immune function. In a comparative study of birds, repertoire size and relative size of the spleen, which is an important immune defence organ, were strongly, positively correlated accounting for almost half of the variance. This finding suggests that host-parasite interactions have played an important role in the evolution of song repertoires in birds.

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Reproductive isolation in natural populations of Drosophila melanogaster from Brazzaville (Congo).

The aim of this work is to analyze the homogamy previously detected between two natural populations of Drosophila melanogaster from Brazzaville. It is shown that mating isolation was still maintained under laboratory conditions 10 years after the populations samples were trapped. Isolation seemed to be due mainly to premating isolation and we checked for any suggestion of post-mating mortality of hybrids. Pre-mating isolation was not symmetrical, and significant chi2 values were found in 3/4 possible 3-way mating choice experiments. The only exception involved a male from the countryside and two females (one from each population) for which no significant mating preference was detected. Mortality of hybrids was intermediate between those of the parental strains showing a clear maternal effect and the existence of partial dominance. Major differences in the cuticular hydrocarbons were also found and they could account for the isolation. These findings in populations from African breweries indicate that they are closely related to European ones, suggesting that this phenomenon is not a case of sympatric speciation, but probably attributable to the reintroduction of an allopatric population.

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Cues for eavesdroppers: do frog calls indicate prey density and quality?

Predators and parasites that eavesdrop on the mating signals of their prey often preferentially select individuals within a prey/host species that produce specific cues. Mechanisms driving such signal preferences are poorly understood. In the tungara frog Physalaemus pustulosus, conspecific females, frog-eating bats, and blood-sucking flies all prefer complex to simple mating calls. In this study we assess the natural signal variation in choruses in the wild and test two hypotheses for why eavesdroppers prefer complex calls: (1) prey quality: complex calls indicate better quality of prey/host, and (2) prey density: complex calls indicate higher prey/host density. Call complexity is not correlated with frog length, mass, or body condition, but it does signal higher abundance of prey/host. Thus, increased effectiveness of attack may have played a role favoring the preference for complex calls in eavesdropping heterospecifics.

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Courtship song recognition in the Drosophila melanogaster complex: heterospecific songs make females receptive in D. melanogaster, but not in D. sechellia.

The courtship song emitted by male wing vibration has been regarded as one of the most important signals in sexual isolation in the species of the Drosophila melanogaster complex. Inter- and intraspecific crosses were observed using males whose wings were removed (mute) or females whose aristae were removed (deaf). Females of D. melanogaster, D. simulans, and D. mauritiana mated with heterospecific males in the song-present condition (cross between normal females and winged males) more often than in the no-song condition (cross between normal females and wingless males or between aristaless females and winged males) or they showed no preference between the two conditions. It is possible that in these females heterospecific courtship songs play a role as if they were conspecific. In contrast, the females of D. sechellia mated with D. melanogaster or D. simulans males in the no-song condition more often than in the song-present condition, suggesting that they reject males with heterospecific song. Female mate recognition depending on the courtship song in D. melanogaster, D. simulans, and D. mauritiana is considered to be relatively broader and that in D. sechellia narrower.

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Frequency-dependent sexual selection.

Sexual selection by female choice is expected to give rise to a frequency-dependent sexual advantage in favour of preferred male phenotypes: the rarer the preferred phenotypes, the more often they are chosen as mates. This 'rare-male advantage' can maintain a polymorphism when two or more phenotypes are mated preferentially: each phenotype gains an advantage when it is rarer than the others; no preferred phenotype can then be lost from the population. Expression of preference may be complete or partial. In models of complete preference, females with a preference always mate preferentially. Models of partial preference are more realistic: in these models, the probability that a female mates preferentially depends on the frequency with which she encounters the males she prefers. Two different 'encounter models' of partial preference have been derived: the O'Donald model and the Charlesworth model. The encounter models contain the complete preference model as a limiting case. In this paper, the Charlesworth model is generalized to allow for female preference of more than one male phenotype. Levels of frequency dependence can then be compared in the O'Donald and Charlesworth models. The complete preference model and both encounter models are formulated in the same genetical terms of preferences for dominant and recessive male phenotypes. Polymorphic equilibria and conditions for stability are derived for each of the three models. The models are then fitted to data of frequencies of matings observed in experiments with the two-spot ladybird. The complete preference model gives as good a fit as the encounter models to the data of these and other experiments. The O'Donald and Charlesworth encounter models are shown to produce a very similar frequency-dependent relation. Generally, as females become less choosy, they express their preference with more dependence on male frequency, whereas the resulting selection of the males becomes less frequency dependent. More choosy females are more constant in expressing their preference, producing greater frequency dependence in the selection of the males.

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