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Mechanisms underlying the sperm quality advantage in Drosophila melanogaster.

Contrary to early predictions of sperm competition theory, postcopulatory sexual selection favoring increased investment per sperm (e.g., sperm size, sperm quality) has been demonstrated in numerous organisms. We empirically demonstrate for Drosophila melanogaster that both sperm quality and sperm quantity independently contribute to competitive male fertilization success. In addition to these independent effects, there was a significant interaction between sperm quality and quantity that suggests an internal positive reinforcement on selection for sperm quality, with selection predicted to intensify as investment per sperm increases and the number of sperm competing declines. The mechanism underlying the sperm quality advantage is elucidated through examination of the relationship between female sperm-storage organ morphology and the differential organization of different length sperm within the organ. Our results exemplify that primary sex cells can bear secondary sexual straits.

Animals↗

Female choice in Drosophila: evidence from Hawaii and implications for evolutionary biology.

Details of female choice of mate in Drosophila silvestris of Hawaii strikingly parallels epigamic behavioral systems in many other animals and may be common in other species of Drosophilidae. Females respond selectively to male circling, wing displays, songs and tactile stimulation with foreleg cilia, a quantitative character that is highly variable in some populations. I hypothesize that the female can exert choice based on these cues from individual males that differ genetically by quantitative trait loci. Laboratory tests show that one third of courting males are repeatedly rejected in favor of a minority of 'alpha' males. This result imposes non-random mating at the local population level. Past multiple-choice lab tests, widely used to measure 'isolation' between pairs of populations or species of Drosophila may be flawed, since random mating has been assumed in the interpretation of results. Pre-mating sexual selection is clearly a powerful intrapopulation force in population biology. This view creates difficulties for discerning any proposed simultaneous interpopulation selective events in the presence of strong female choice. The long-held theory assuming that there is significant selection for pre-mating isolation between groups is questionable.

Animals↗

Captivity masks inbreeding effects on male mating success in butterflies.

Small isolated populations are frequently genetically less diverse than core populations, resulting in higher homozygosity that can hamper their long-term survival. The decrease in fitness of organisms owing to matings between relatives is well known from captive and laboratory animals. Such inbreeding can have strongly deleterious effects on life-history traits and survival, and can be critical to the success of population conservation. Because pedigrees are hard to follow in the wild, most field studies have used marker loci to establish that fitness declines with increasing homozygosity. Very few have experimentally explored the effects of inbreeding in the wild, or compared observations in the laboratory with field conditions. Here, using a technique involving the transfer of marker dusts during copulation, we show that a small decrease in mating success of captive inbred male butterflies in cages is greatly accentuated in conditions with unconstrained flight. Our results have important implications for conservation and for studies of sexual selection because they show that the behaviours underlying patterns of mating can be profoundly influenced by a history of inbreeding or by any restraining experimental conditions.

Animals↗

Incipient speciation by sexual isolation in Drosophila melanogaster: extensive genetic divergence without reinforcement.

The collection of Drosophila melanogaster from Zimbabwe and nearby regions (the Z-type) yield females who would not mate with the cosmopolitan D. melanogaster males (the M-type). To dissect the genetic basis of this sexual isolation, we constructed 16 whole-chromosome substitution lines between two standard Z- and M-lines. The results were as follows: (1) All substitution lines appear normal in viability and fertility in both sexes, indicating no strong postmating isolation. (2) The genes for the behaviors are mapped to all three major chromosomes with the same ranking and comparable magnitude of effects for both sexes: III > II >> X > or = 0 (III, II and X designate the effects of the three chromosomes). The results suggest less evolution on the X than on autosomes at loci of sexual behavior. (3) The genes for "Z-maleness" are many and somewhat redundant. Whole-chromosome effects for Z-maleness appear nearly additive and show little dominance. (4) In contrast, "Z-femaleness" has less redundancy as partial genotypes never exhibit full phenotypic effects. Epistatic interactions and incomplete dominance can sometimes be detected. (5) The extensive genetic divergence underlying sexual isolation has evolved in the absence of detectable reduction in hybrid fitnesses. Sexual selection has apparently been a driving force of multiple facets of speciation at the nascent stage without reinforcement.

Animals↗

Sequence variation in the coding region of the melanocortin-1 receptor gene (MC1R) is not associated with plumage variation in the blue-crowned manakin (Lepidothrix coronata).

Avian plumage traits are the targets of both natural and sexual selection. Consequently, genetic changes resulting in plumage variation among closely related taxa might represent important evolutionary events. The molecular basis of such differences, however, is unknown in most cases. Sequence variation in the melanocortin-1 receptor gene (MC1R) is associated with melanistic phenotypes in many vertebrate taxa, including several avian species. The blue-crowned manakin (Lepidothrix coronata), a widespread, sexually dichromatic passerine, exhibits striking geographic variation in male plumage colour across its range in southern Central America and western Amazonia. Northern males are black with brilliant blue crowns whereas southern males are green with lighter blue crowns. We sequenced 810 bp of the MC1R coding region in 23 individuals spanning the range of male plumage variation. The only variable sites we detected among L. coronata sequences were four synonymous substitutions, none of which were strictly associated with either plumage type. Similarly, comparative analyses showed that L. coronata sequences were monomorphic at the three amino acid sites hypothesized to be functionally important in other birds. These results demonstrate that genes other than MC1R underlie melanic plumage polymorphism in blue-crowned manakins.

Animals↗

Evolution of ejaculates: patterns of phenotypic and genotypic variation and condition dependence in sperm competition traits.

Sperm competition is widely recognized as a potent force in evolution, influencing male behavior, morphology, and physiology. Recent game theory analyses have examined how sperm competition can influence the evolution of ejaculate expenditure by males and the morphology of sperm contained within ejaculates. Theoretical analyses rest on the assumption that there is sufficient genetic variance in traits important in sperm competition to allow evolving populations to move to the evolutionarily stable equilibrium. Moreover, patterns of genotypic variation can provide valuable insight into the nature of selection currently acting on traits. However, our knowledge of genetic variance underlying traits important in sperm competition is limited. Here we examine patterns of phenotypic and genotypic variation in four sperm competition traits in the dung beetle Onthophagus taurus. Testis weight, ejaculate volume, and copula duration were found to have high coefficients of additive genetic variation (CV(A)S), which is characteristic of fitness traits and traits subject to sexual selection. Heritabilities were high, and there was some evidence for Y-linked inheritance in testis weight. In contrast, sperm length had a low CV(A), which is characteristic of traits subject to stabilizing selection. Nevertheless, there was little residual variance so that the heritability of sperm length exceeded 1.0. Such a pattern is consistent with Y-linked inheritance in sperm length. Interestingly, we found that testis weight and sperm length were genetically correlated with heritable male condition. This finding holds important implications for potential indirect benefits associated with the evolution of polyandry.

Animals↗

Disturbed sexual characteristics in male mosquitofish (Gambusia holbrooki) from a lake contaminated with endocrine disruptors.

Previous laboratory studies have demonstrated that estrogenic and antiandrogenic chemicals can alter several sexual characteristics in male poeciliid fishes. Whether similar disturbances occur under field conditions remains to be confirmed. Lake Apopka, Florida, is contaminated with numerous chemicals, some of which possess endocrine-disrupting activity. Male mosquitofish (Gambusia holbrooki) were collected monthly from December 2000 through May 2001 from Lake Apopka and two nearby reference lakes, Orange Lake and Lake Woodruff National Wildlife Refuge. Selected sexual characteristics were compared temporally and among lakes during the collection period. Male fish from Lake Apopka had slightly shorter gonopodia and on average 32 and 47% fewer sperm cells per milligram testis, when compared with the fish collected from Orange Lake and Lake Woodruff, respectively. The testes weights increased markedly during spring, with significantly smaller testes in fish from Lake Apopka than from Orange Lake, but surprisingly, the smallest testes occurred in males obtained from the Lake Woodruff population. The highest liver weights were found in the Lake Apopka population. Whole-body concentrations of testosterone and estradiol varied among months; the peak testosterone concentration occurred in January and was significantly lower in male fish from Lake Apopka compared with Orange Lake. The intensity of male courtship behavior was highly correlated to body testosterone concentration, but no statistically significant differences in sexual behavior among the lakes were found. We conclude that sexual characteristics of relevance to male reproductive capacity are altered in the Lake Apopka mosquitofish population, and we discuss the presence of chemicals with antiandrogenic effects in Lake Apopka as a possible cause of the observed alterations.

Androgen Antagonists↗

Developmental isolation and subsequent adult behavior of Drosophila pseudoobscura.

When isolated from all contact with other flies until the opportunity to exercise mating choices, Drosophila pseudoobscura displays genetically-mediated compensatory behavior: heterokaryotypic females are more vulnerable than homokaryotes to wholly restricted contact with conspecifics. They are less able to demonstrate normal frequency-dependent sexual selection after isolation than their own homokaryotypes. However, the later the isolation in this holometabolous insect--eggs, first instars, second instars, third instars, pupae, imagoes--the fewer the behavioral effects of this isolation.

Animals↗

Plumage colour in nestling blue tits: sexual dichromatism, condition dependence and genetic effects.

Sexual-selection theory assumes that there are costs associated with ornamental plumage coloration. While pigment-based ornaments have repeatedly been shown to be condition dependent, this has been more difficult to demonstrate for structural colours. We present evidence for condition dependence of both types of plumage colour in nestling blue tits (Parus caeruleus). Using reflectance spectrometry, we show that blue tit nestlings are sexually dichromatic, with males having more chromatic (more 'saturated') and ultraviolet (UV)-shifted tail coloration and more chromatic yellow breast coloration. The sexual dimorphism in nestling tail coloration is qualitatively similar to that of chick-feeding adults from the same population. By contrast, the breast plumage of adult birds is not sexually dichromatic in terms of chroma. In nestlings, the chroma of both tail and breast feathers is positively associated with condition (body mass on day 14). The UV/blue hue of the tail feathers is influenced by paternally inherited genes, as indicated by a maternal half-sibling comparison. We conclude that the expression of both carotenoid-based and structural coloration seems to be condition dependent in blue tit nestlings, and that there are additional genetic effects on the hue of the UV/blue tail feathers. The signalling or other functions of sexual dichromatism in nestlings remain obscure. Our study shows that nestling blue tits are suitable model organisms for the study of ontogenetic costs and heritability of both carotenoid-based and structural colour in birds.

Aging↗

Animal communication: complex call production in the túngara frog.

Animals' sound-producing organs often act as an integrated whole--particular vocal structure are not directly associated with the creation of discrete syllables. But here we show that the 'chuck' of the 'whine-chuck' mating call of the túngara frog, Physalaemus pustulosus, is caused by a fibrous mass attached to the vocal folds; the chuck is eliminated by removal of this structure, although the frog still tries to produce the sound. Sexual selection affects the acoustic complexity of the frog's call, so evolution may have shaped this unusual vocalization, which is akin to the two-voiced song of songbirds.

Animals↗

A model of the interaction between 'good genes' and direct benefits in courtship-feeding animals: when do males of high genetic quality invest less?

Conflict between mates over the amount of parental investment by each partner is probably the rule except in rare cases of genetic monogamy. In systems with parental care, males may frequently benefit by providing smaller investments than are optimal for individual female partners. Females are therefore expected to choose males that will provide the largest amounts of parental investment. In some species, however, the preferred males provide less care than their rivals. Focusing on species in which males invest by feeding their mates, I use a simple model to demonstrate the conditions under which males preferred by females may have optimal donations that are smaller than those of less-preferred rivals. Pre-mating female choice may sufficiently bias the perception of mate availability of preferred males relative to their rivals such that preferred males gain by conserving resources for future matings. Similarly, 'cryptic' biases in favour of high-quality ejaculates by females can compensate for smaller than average donations received from preferred males. However, post-fertilization cryptic choice should not change the optimal donations of preferred males relative to their rivals. I discuss the implications of this work for understanding sexual selection in courtship-feeding animals, and the relevance of these systems to understanding patterns of investment for animals in general.

Animals↗

Accelerated protein evolution and origins of human-specific features: Foxp2 as an example.

Genes responsible for human-specific phenotypes may have been under altered selective pressures in human evolution and thus exhibit changes in substitution rate and pattern at the protein sequence level. Using comparative analysis of human, chimpanzee, and mouse protein sequences, we identified two genes (PRM2 and FOXP2) with significantly enhanced evolutionary rates in the hominid lineage. PRM2 is a histone-like protein essential to spermatogenesis and was previously reported to be a likely target of sexual selection in humans and chimpanzees. FOXP2 is a transcription factor involved in speech and language development. Human FOXP2 experienced a >60-fold increase in substitution rate and incorporated two fixed amino acid changes in a broadly defined transcription suppression domain. A survey of a diverse group of placental mammals reveals the uniqueness of the human FOXP2 sequence and a population genetic analysis indicates possible adaptive selection behind the accelerated evolution. Taken together, our results suggest an important role that FOXP2 may have played in the origin of human speech and demonstrate a strategy for identifying candidate genes underlying the emergences of human-specific features.

Alleles↗

The population genetics of sperm displacement.

This article reports the results of some sperm displacement experiments, as well as the results of a theoretical study of selection arising from genetic differences in displacing ability. The experimental work involved the use of three genetic marker stocks in double and triple matings. The speed of displacement following the matings was determined by scoring the progeny of each female daily. There were clear differences between strains in their displacing ability. It is shown how new information concerning the displacement process results when three markers are used; however, no new light is shed by these experiments on the mechanism of displacement. The theoretical study of selection resulting from displacement uses a one-locus, two-allele model in which three diploid male genotypes confer different displacing abilities. The results indicate stable equilibria if (1) there is heterosis, and (2) there are certain nontransitive relationships in displacing ability among the different kinds of double matings. Some evolutionary consequences are discussed in which sperm displacement is regarded as a form of sexual selection.

Alleles↗

Quantitative genetics of ultrasonic advertisement signalling in the lesser waxmoth Achroia grisella (Lepidoptera: pyralidae).

Males of the lesser waxmoth Achroia grisella (Lepidoptera: Pyralidae) produce ultrasonic advertisement signals attractive to females within several metres. Previous studies showed that females prefer male signals that are louder, delivered at a faster rate, and have a greater asynchrony between pulses produced by the right and left wings. These three signal characters vary considerably within populations but are repeatable within individuals. Breeding experiments employing half-sib designs were conducted on both collectively and individually reared moths to determine genetic variance within and covariance among these signal characters. Heritabilities of all signal characters were significant among collectively reared moths. Heritabilities for signal rate and right-left wing asynchrony interval were not significant, however, among individually reared moths, suggesting the presence of significant nonadditive genetic variance or common environmental variation. Development time was also significantly heritable, but only under individual rearing. The only significant genetic correlation was between signal rate and length of the right-left wing asynchrony and this was negative. Our findings on heritability of signal characters are consistent with a coevolutionary sexual selection mechanism, but the absence of signal x development genetic correlation fails to support specifically a good-genes mechanism. The variation in heritability among conditions suggests that environmental variance may be high, and may render selection on signal characters by female choice ineffective. Thus, additive genetic variance for these characters may be maintained in the presence of directional female choice.

Animals↗

Mating behavior of a flower-visiting longhorn beetle Zorion guttigerum (Westwood) (Coleoptera: Cerambycidae: Cerambycinae).

Long-range sex pheromones have been demonstrated for several cerambycid beetle species. Our field study on the mating behavior of Zorion guttigerum, on the basis of its temporal and spatial distributions on mating and feeding sites (flowers), and longevity, however, suggests that such pheromones are not used by this species. Plant characteristics rather than long-range sex pheromones may play an important role in bringing both sexes together. Adult activities on flowers occur exclusively during the day with two peaks, one around midday and the other in the late afternoon. Overall operational sex ratio is male-biased ( approximately 1 female symbol:1.5 male symbol) but it becomes very highly male-biased ( approximately 1 female symbol:9 male symbol) when mating and feeding activities decrease to the minimum in mid-afternoon, suggesting that females leave flowers to oviposit during that period of time. For cerambycid species whose females oviposit alone, and in which mating and oviposition occur on different plants or different plant parts, the operational sex ratio appears to vary significantly over time on the mating sites. The number and duration of pair-bondings also vary over time for Z. guttigerum. Fewer and shorter pair-bondings in the morning may suggest a strong sexual selection process. After approximately 2 h of selection, both sexes tend to engage in longer pair-bondings and mate more times before females leave the mating sites in mid-afternoon. Details of the mating behavior are described here.

Animal Feed↗

Sex in Drosophila mauritiana: a very high level of amino acid polymorphism in a male reproductive protein gene, Acp26Aa.

Many genes pertaining to male reproductive functions have been shown to evolve rapidly between species, and evidence increasingly suggest the influence of positive Darwinian selection. The accessory gland protein gene (Acp26Aa) of Drosophila is one such example. In order to understand the mechanism of selection, it is often helpful to examine the pattern of polymorphism. We report here that the level of amino acid polymorphism in the N-terminal quarter of Acp26Aa is high in Drosophila melanogaster and is unprecedented in its sibling species Drosophila mauritiana. We postulate that (1) this N-terminal segment may play a role in sperm competition, and (2) D. mauritiana may have been under much more intense sexual selection than other species. Both postulates have important ramifications and deserve to be tested rigorously.

Amino Acid Sequence↗

History of U.S. military contributions to the study of sexually transmitted diseases.

Sexually transmitted diseases have posed a threat to military service members throughout history. Among these diseases, syphilis, gonorrhea, and human immunodeficiency virus infections have accounted for the most significant morbidity and mortality rates in the U.S. military. In response, military researchers have made significant contributions to the treatment and prevention of these diseases. We review the impact of these diseases through the history of the U.S. Armed Forces and review selected sexually transmitted disease-oriented publications of U.S. military researchers.

Biomedical Research↗

Selective advantage for sexual reproduction.

This paper develops a simplified model for sexual reproduction within the quasispecies formalism. The model assumes a diploid genome consisting of two chromosomes, where the fitness is determined by the number of chromosomes that are identical to a given master sequence. We also assume that there is a cost to sexual reproduction, given by a characteristic time tau(seek) during which haploid cells seek out a mate with which to recombine. If the mating strategy is such that only viable haploids can mate, then when tau(seek) = 0, it is possible to show that sexual reproduction will always out compete asexual reproduction. However, as tau(seek) increases, sexual reproduction only becomes advantageous at progressively higher mutation rates. Once the time cost for sex reaches a critical threshold, the selective advantage for sexual reproduction disappears entirely. The results of this paper suggest that sexual reproduction is not advantageous in small populations per se, but rather in populations with low replication rates. In this regime, the cost for sex is sufficiently low that the selective advantage obtained through recombination leads to the dominance of the strategy. In fact, at a given replication rate and for a fixed environment volume, sexual reproduction is selected for in high populations because of the reduced time spent finding a reproductive partner.

Biological Evolution↗