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PubMed2002

Discrimination of MHC-derived odors by untrained mice is consistent with divergence in peptide-binding region residues.

Genes of the major histocompatibility complex (MHC) play a central role in immune recognition, yet they also influence the odor of individuals. Mice can be trained to distinguish odors mediated by classical MHC loci; however, training can introduce confounding behavioral artifacts. This study demonstrates that mice can distinguish some, but not all, naturally occurring allelic variants at classical MHC loci without prior training. This result suggests that MHC-disassortative mating preferences might operate by means of small MHC-based odor differences, and could therefore contribute to diversifying selection acting on MHC loci. Here we show that odors of two MHC mutant mouse strains (bm1 and bm3) can be distinguished, even after genetic background is controlled by intercrossing strains. These two strains differ by five amino acids, three of which are predicted to chemically contact peptides bound to the peptide-binding region (PBR), the site of antigen presentation for T cell recognition. However, the odors of neither bm1 nor bm3 were distinguished from their parental B6 haplotype after randomizing genomic background, despite discrimination of pure-bred B6 and bm1 strain odors. These combined results suggest that (i) there may be an MHC odor discrimination threshold based on divergence in PBR residues, providing a more logical pattern of MHC-based odor discrimination than found in previous training studies, where discrimination ability was not correlated with PBR divergence; and (ii) additional (non-MHC) mutations that influence odor have accumulated in these strains during the 100 generations of divergence between pure B6 and bm1 strains.

Alleles↗

PubMed2003

Neural images of pursuit targets in the photoreceptor arrays of male and female houseflies Musca domestica.

Male houseflies use a sex-specific frontal eye region, the lovespot, to detect and pursue mates. We recorded the electrical responses of photoreceptors to optical stimuli that simulate the signals received by a male or female photoreceptor as a conspecific passes through its field of view. We analysed the ability of male and female frontal photoreceptors to code conspecifics over the range of speeds and distances encountered during pursuit, and reconstructed the neural images of these targets in photoreceptor arrays. A male's lovespot photoreceptor detects a conspecific at twice the distance of a female photoreceptor, largely through better optics. This detection distance greatly exceeds those reported in previous behavioural studies. Lovespot photoreceptors respond more strongly than female photoreceptors to targets tracked during pursuit, with amplitudes reaching 25 mV. The male photoreceptor also has a faster response, exhibits a unique preference for stimuli of 20-30 ms duration that selects for conspecifics and deblurs moving images with response transients. White-noise analysis substantially underestimates these improvements. We conclude that in the lovespot, both optics and phototransduction are specialised to enhance and deblur the neural images of moving targets, and propose that analogous mechanisms may sharpen the neural image still further as it is transferred to visual interneurones.

Animals↗

PubMed2005

Male reproductive physiology as a sexually selected handicap? Erectile dysfunction is correlated with general health and health prognosis and may have evolved as a marker of poor phenotypic quality.

Many extravagant physical traits are selected because they are used as cues for mate choice (sexual selection). Why is mate choice driven by costly ornaments? A theory of sexual selection posits that extravagant traits are preferred because are reliable indicators of superior (heritable) phenotypic quality. In particular, the preferred traits can be expressed only in individuals with superior conditions because are handicaps which impose a high cost to the carrier. The human penis achieves its reproductive function by the complex neuro-vascular mechanisms that controls erection. Surprisingly, erectile dysfunction and infertility, two condition which nearly annihilate fitness, are widespread medical conditions which affect millions of people of any age worldwide. The very high incidence of erectile dysfunction appears as an evolutionary paradox. Impotence is associated with all major systemic diseases as well depression and stress. Stress is also one of the causes of infertility. Therefore, male reproduction appears to be extremely sensitive to internal and external stressors. Moreover, erectile dysfunction is a predictor of myocardial infarction and stroke, whereas men with regular sexual activity have lower risk of death due to coronary disease. This large body of medical literature makes erection the best case for a fitness indicator in the human male. We suggest that the fragility of male sexual physiology observed in modern society is the specific consequence of an evolutionary process driven by the handicap principle.

Animals↗

PubMed2002

Microsatellite markers associated with quantitative trait loci controlling antibody response to Escherichia coli and Salmonella enteritidis in young broilers.

A unique resource population was produced to facilitate detection of microsatellite markers associated with quantitative trait loci controlling antibody (Ab) response in broiler chickens. Three F1 males were produced by mating two lines divergently selected on Ab response to Escherichia coli vaccination. Each F1 male was mated with females from four genetic backgrounds: F1, high-Ab line (HH), low-Ab line and commercial line, producing three resource families, each with four progeny types. About 1700 chicks were immunized with E. coli and Salmonella enteritidis vaccines. Selective genotyping was conducted on the individuals with highest or lowest average Ab to E. coli and S. enteritidis within each progeny type in each sire family. Twelve markers were significantly associated with Ab to E. coli and six of them were also associated with Ab to S. enteritidis, mostly exhibiting a similar low effect (approximately 0.35 phenotypic SD) in all progeny types. Four markers exhibited a highly significant and much larger effect (approximately 1.7 SD), but only in progeny of females from the HH, suggesting that a backcross to the high parental line should be preferred over the commonly used F2 population. Results from two markers suggested a quantitative trait locus on chromosome 2 around 400 cM. The marker MCW0083, significant in two sire families, is closely linked to the bone morphogenetic protein 2 (BMP2) gene, known to be associated with the control of T-cell transformation in humans.

Animals↗

PubMed2004

Chemical and visual communication during mate searching in rock shrimp.

Mate searching in crustaceans depends on different communicational cues, of which chemical and visual cues are most important. Herein we examined the role of chemical and visual communication during mate searching and assessment in the rock shrimp Rhynchocinetes typus. Adult male rock shrimp experience major ontogenetic changes. The terminal molt stages (named "robustus") are dominant and capable of monopolizing females during the mating process. Previous studies had shown that most females preferably mate with robustus males, but how these dominant males and receptive females find each other is uncertain, and is the question we examined herein. In a Y-maze designed to test for the importance of waterborne chemical cues, we observed that females approached the robustus male significantly more often than the typus male. Robustus males, however, were unable to locate receptive females via chemical signals. Using an experimental set-up that allowed testing for the importance of visual cues, we demonstrated that receptive females do not use visual cues to select robustus males, but robustus males use visual cues to find receptive females. Visual cues used by the robustus males were the tumults created by agitated aggregations of subordinate typus males around the receptive females. These results indicate a strong link between sexual communication and the mating system of rock shrimp in which dominant males monopolize receptive females. We found that females and males use different (sex-specific) communicational cues during mate searching and assessment, and that the sexual communication of rock shrimp is similar to that of the American lobster, where females are first attracted to the dominant males by chemical cues emitted by these males. A brief comparison between these two species shows that female behaviors during sexual communication contribute strongly to the outcome of mate searching and assessment.

Analysis of Variance↗

PubMed2003

Behavioral genetics of Drosophila ananassae.

Drosophila ananassae has a unique status among Drosophila species because of certain peculiarities in its genetic behavior. The most unusual feature of this species is its relatively high frequency of spontaneous male recombination. The results of studies on non-sexual behavior, such as phototactic responses, eclosion rhythm, and preferences for oviposition and pupation sites, lead us to suggest that this behavior is under polygenic control, with a substantial amount of additive genetic variation. Sexual isolation has been reported in D. ananassae with the degree of such isolation being stronger in isofemale lines than in natural populations. The significant variations seen in the mating propensity of several isofemale strains, inversion karyotypes and wild type strains, the diminishing effects of certain mutations on the sexual activity of males, and the positive responses to selection for high and low mating propensity point to a genetic control of sexual behavior in D. ananassae. Males contribute more to variation and thus are more subject to intrasexual selection than females. There is a positive correlation between sternopleural bristle number, mating propensity and fertility in D. ananassae. This correlation between morphometric traits and mating success suggests that larger flies are more successful in mating than smaller ones. There is also evidence for adaptive plasticity and a trade-off between longevity and productivity in D. ananassae. Rare, specific courtship song parameters that provide males with a mating advantage have also been reported in different geographic strains of D. ananassae. The remating behavior of males and females, sperm displacement, and the bi-directional selection for female remating speed indicate that post-mating behavior in this species may also be under genetic control. The occurrence of size assortative mating further indicates that there is size-dependent sexual selection in D. ananassae.

Animals↗

PubMed2007

Role of (3Z,6Z,8E)-dodecatrien-1-ol in trail following, feeding, and mating behavior of Reticulitermes hesperus.

Trail pheromones mediate communication among western subterranean termites, Reticulitermes hesperus Banks. Repetitive passages of >or=28 termites were required to establish a pheromone trail and trails needed to be reinforced because they lasted <48 hr. The minimal threshold concentration for inducing responses from termite workers and secondary reproductives was between 0.01 and 0.1 fg/cm of (3Z,6Z,8E)-dodecatrien-1-ol (henceforth, dodecatrienol). Workers showed optimal trail-following behavior to dodecatrienol at a concentration of 10 fg/cm. Trails with concentrations >10 pg/cm were repellent to workers. Workers did not detect pheromone gradients, responding equally to increasing or decreasing gradients of dodecatrienol, and termite workers were not able to differentiate between different concentrations of dodecatrienol. Termites preferred dodecatrienol trails to 2-phenoxyethanol trails. Antennae played a key role in trail pheromone perception. Dodecatrienol acted as an arrestant for worker termites (10 fg/cm2) and male alates (5 ng/cm2), whereas sternal gland extracts from females attracted male alates. Workers and alates, upon contact with filter paper disks treated with higher doses (10 fg/cm2 and 5 ng/cm2, respectively) of dodecatrienol, were highly excited (increased antennation and palpation) and repeatedly returned to the treated disks. Dodecatrienol did not act as a phagostimulant when offered on a paper towel disk. Reticulitermes hesperus is highly responsive to dodecatrienol, and it may play an important role in orientation of workers and alates.

Animals↗

PubMed2004

Can too strong female choice deteriorate male ornamentation?

Competition for limited resources can have fundamental implications for population dynamics. However, the effects of resource depletion have rarely been discussed in the context of sexual selection, even though mate choice typically favours males who outperform others in securing access to some limited resource. Here, we develop a model to investigate the question of resource competition as a form of male-male competition in the context of male sexual displays. We phrase our model in terms of male bowerbirds either searching for or stealing resources (ornamental objects) valued by females, and compare the model findings with published studies of time allocation to various activities in different species of bowerbirds. The basic idea of the model, however, extends to cases where the resource is used less directly for the development of sexual ornamentation, such as males excluding others' access to food. We show that if males compete for resources used in sexual displays, intense female preference for high-quality displays can lead to poorer prospects for efficient choice by females. This is because males benefit from excluding others' access to resources used in displays, damaging the overall efficiency of resource use in the population, and the accuracy with which females can judge male ability to gain such resources. The evolution of female choice may therefore have a self-limiting nature when it poses a selection pressure on male resource acquisition.

Animals↗

PubMed1983

Differences in the fatty acid profile and beta-oxidation by heart homogenates of rats fed cis and trans octadecenoic acids.

Female Wistar rats were fed a fat-free diet containing either 5% partially hydrogenated corn oil (52.2% elaidate) or 5% oleic acid (67% oleate) with 8.6% linoleate providing 1% of calories 2 weeks before mating and were maintained on this diet throughout pregnancy and lactation. Fatty acid analysis of the developing organs as well as beta-oxidation by heart homogenates with [1-14C]palmitate, [1-14C]elaidate and [1-14C]oleate of the developing male and female progeny were determined and compared with age-matched controls on a stock diet. Results show that irrespective of the cis and trans 18:1 in the diet, the maternal plasma at term contained mostly cis 18:1, with 5% trans for the rats on the trans diet. The placenta and fetal liver contained 40 and 60% less trans, respectively, than did the maternal plasma. trans 18:1 was not detected in fetal brain or heart. Regardless of diet or sex, the order of preference for the heart was palmitate greater than elaidate greater than oleate. There was an increase in the rate of beta-oxidation of all the substrates, especially in the females on the trans diet, suggesting a stimulation of one or more of the enzymes involved. Above all, the myocardium showed a unique capacity to retain n-6 and n-3 fatty acids when the levels of these decreased in the serum.

Animals↗

PubMed2005

Escalation, retreat, and female indifference as alternative outcomes of sexually antagonistic coevolution.

Verbal and quantitative genetic models of sexually antagonistic coevolution suggest that coevolutionary arms races should be common. Sexual selection favors exaggeration of male persistence traits that are costly to females, and females, in turn, are selected to resist these traits. The heightened resistance by females is thought to then favor further exaggeration in the male trait, leading to an escalating coevolutionary arms race between persistence and resistance traits. Much of this theory, however, is based on an (implicit) assumption that there are tight constraints on how female resistance can evolve. We develop a theory that identifies and relaxes these constraints, allowing female resistance to evolve in a fashion that better reflects known empirical patterns in the evolution of female preference functions (the resistance trait). Our results suggest that evolutionary arms races that lead to the exaggeration of persistence and resistance will be much less common than formerly predicted. Females sometimes evolve indifference to male traits rather than resistance and can even evolve to discriminate against these traits. These alternative outcomes depend on the existence of genetic variance in the components of the female sensory system underlying female resistance and on the strength of natural selection acting on these components. Female indifference tends to evolve when natural selection on the sensory system is weak, and under these conditions, sexually antagonistic coevolution tends not to reduce female fitness significantly at equilibrium. When natural selection on the female sensory system is strong, however, then arms races are more likely, and female fitness is then sometimes significantly depressed at equilibrium. Sexually antagonistic coevolution is thus likely to have strong deleterious effects on population fitness only when female sensory traits are under strong natural selection to perform functions in addition to those involved with mating. Together, these results suggest that identifying the nature of genetic variation in and the strength of natural selection on female resistance should be a central goal of future studies of sexual conflict.

Animal Communication↗

PubMed2000

Nestling growth and song repertoire size in great reed warblers: evidence for song learning as an indicator mechanism in mate choice.

Females of many songbird species show a preference for mating with males that have larger song repertoires, but the advantages associated with this preference are uncertain. We tested the hypothesis that song complexity can serve as an indicator of male quality because the development of the brain regions underlying song learning and production occurs when young birds typically face nutritional and other stresses, so that song reflects how well a male fared during post-hatch development. A key prediction of this hypothesis is that variation in nestling condition should correspond to variation in the adult song repertoires of individuals. We used data from a long-term study of the great reed warbler (Acrocephalus arundinaceus) to test this prediction, correlating two measures of nestling development with subsequent repertoire size of males. We found that the length of the innermost primary feather, a standard measure of development, significantly predicted first-year repertoire size. The relationship between repertoire size and body mass was nearly significant, in spite of the large variance inherent in this measure. These data support the idea that song may provide females with information about a male's response to developmental stress, which in turn is expected to correlate with indirect or direct benefits she might receive.

Animals↗

PubMed2003

Evolution of cooperation: cooperation defeats defection in the cornfield model.

"Cooperation" defines any behavior that enhances the fitness of a group (e.g. a community or species), but which, by its nature, can be exploited by selfish individuals, meaning, firstly, that selfish individuals derive an advantage from exploitation which is greater than the average advantage that accrues to unselfish individuals. Secondly, exploitation has no intrinsic fitness value except in the presence of the "cooperative behavior". The mathematics is described by the simple Prisoner's Dilemma Game (PDG). It has previously been shown that koinophilia (the avoidance of sexual mates displaying unusual or atypical phenotypic features, such as mutations) stabilizes any inherited strategy in the simple or iterated PDG, meaning that it cannot be displaced by rare forms of alternative behavior which arise through mutation or occasional migration. In the present model equal numbers of cooperators and defectors (in the simple PDG) were randomly spread in a two-dimensional "cornfield" with uniformly distributed resources. Every individual was koinophilic, and interacted (sexually and in the PDG tournaments) only with individuals from within its immediate neighborhood. This model therefore tested whether cooperation can outcompete defection or selfishness in a straight, initially equally matched, evolutionary battle. The results show that in the absence of koinophilia cooperation was rapidly driven to extinction. With koinophilia there was a very rapid loss of cooperators in the first few generations, but thereafter cooperation slowly spread, ultimately eliminating defection completely. This result was critically dependent on sampling effects of neighborhoods. Small samples (resulting from low population densities or small neighborhood sizes) increase the probability that a chance neighborhood comes to consist predominantly of cooperators. A sexual preference for the most common phenotype in the neighborhood then makes that phenotype more common still. Once this occurs cooperation's spread becomes almost inevitable.

Animals↗

PubMed2006

Progestin concentrations are increased following paced mating in midbrain, hippocampus, diencephalon, and cortex of rats in behavioral estrus, but only in midbrain of diestrous rats.

BACKGROUND: The progesterone (P(4)) metabolite, 5alpha-pregnan-3alpha-ol-20-one (3alpha,5alpha-THP), acts in the midbrain ventral tegmental area (VTA) to modulate the intensity and duration of lordosis. 3alpha,5alpha-THP can also have anti-anxiety and anti-stress effects in part through actions in the hippocampus. Separate reports indicate that manipulating 3alpha,5alpha-THP levels in the VTA or hippocampus respectively can influence lordosis and affective behavior. 3alpha,5alpha-THP levels can also be altered by behavioral experiences, such as mating or swim stress. Whether endogenous levels of 3alpha,5alpha-THP modulate and/or are increased in response to affective and/or reproductively-relevant behaviors was investigated. METHODS: In Experiment 1, rats in behavioral estrus or diestrus were individually tested sequentially in the open field, elevated plus maze, partner preference, social interaction, and paced mating tasks and levels of 17beta-estradiol (E(2)), P(4), dihydroprogesterone (DHP), and 3alpha,5alpha-THP in serum, midbrain, hippocampus, diencephalon, and cortex were examined. In Experiments 2 and 3, rats in behavioral estrus or diestrus, were individually tested in the battery indicated above, with, or without, paced mating and tissues were collected immediately after testing for later assessment of endocrine measures. RESULTS: In Experiment 1, behavioral estrous, compared to diestrous, rats demonstrated more exploratory, anti-anxiety, social, and reproductive behaviors, and had higher levels of E(2) and progestins in serum, midbrain, hippocampus, diencephalon, and cortex. In Experiment 2, in midbrain and hippocampus, levels of 3alpha,5alpha-THP and its precursor DHP were increased among rats in behavioral estrus that were mated. In diencephalon, and cortex, DHP levels were increased by mating. In Experiment 3, in midbrain, levels of 3alpha,5alpha-THP and its precursor DHP were increased among diestrous rats that were tested in the behavioral battery with mating as compared to those tested in the behavioral battery without mating. CONCLUSIONS: Increased levels of 3alpha,5alpha-THP in behavioral estrus versus diestrous rats are associated with enhanced exploratory, anti-anxiety, social, and reproductive behaviors. Rats in behavioral estrus that are mated have further increases in 3alpha,5alpha-THP and/or DHP levels in midbrain, hippocampus, diencephalon, and cortex than do non-mated rats in behavioral estrus, whereas diestrous rats only show 3alpha,5alpha-THP increases in midbrain in response to behavioral testing that included mating.

Animals↗

PubMed2004

Song function and the evolution of female preferences: why birds sing, why brains matter.

Analyzing the function of song and its evolution as a communication signal provides an essential backdrop for understanding the physiological and neural mechanisms responsible for song learning, perception, and production. The reverse also is true-understanding the mechanisms underlying song learning provides insight into how song has evolved as a communication signal. Song has two primary functions: to repel other males from a defended space and to attract females and stimulate their courtship. The developmental stress hypothesis we present here builds on studies of the development of the song system to suggest how learned features of song, including complexity and local dialect structure, can serve as indicators of male quality useful to females in mate choice. The link between song and male quality depends on the fact that brain structures underlying song learning largely develop during the first few months post-hatching and that during this same period, songbirds are likely to be subject to nutritional and other developmental stresses. Individuals faring well in the face of stress are able to invest more resources to brain development and are expected to be correspondingly better at song learning. Learned features of song thus become reliable indicators of male quality, with reliability maintained by the developmental costs of song. Data from both field and laboratory studies are now beginning to provide broad support for the developmental stress hypothesis, illustrating the utility of connecting mechanistic and evolutionary analyses of song learning.

Animal Communication↗

PubMed2005

Colour vision and speciation in Lake Victoria cichlids of the genus Pundamilia.

Lake Victoria cichlids are one of the most speciose groups of vertebrates. Selection on coloration is likely playing an important role in their rapid speciation. To test the hypothesis that sensory biases could explain species differences in mating preferences and nuptial coloration, we studied seven populations of four closely related species of the genus Pundamilia that differ in visual environment and male nuptial colour. Microspectrophotometry determined that the wavelength of maximum absorption (lambdamax) of the rod pigment and three cone pigments were similar in all four species. Only the long wavelength sensitive (LWS) pigment varied among species, with 3-4 nm shifts in lambdamax that correlated with differences in the LWS opsin sequence. These subtle shifts in lambdamax coincided with large shifts in male body colour, with red species having longer LWS pigments than blue species. Furthermore, we observed within and between species a correlation between water transparency and the proportion of red/red vs. red/green double cones. Individuals from turbid water had more red/red double cones than individuals from clear water. The variation in LWS lambdamax and in the proportion of red/red double cones could lead to differences in perceived brightness that may explain the evolution of variation in male coloration. However, other factors, such as chromophore shifts and higher order neural processing, should also be investigated to fully understand the physiological basis of differential responses to male mating hues in cichlid fish.

Africa, Eastern↗

PubMed1995

Contribution of stress and gender to exploratory preferences for familiar versus unfamiliar conspecifics.

An apparatus for measuring the exploratory preferences of rats for familiar and unfamiliar conspecifics in a novel environment was designed. The exploratory behavior of males and females was compared and contrasted to that elicited in response to an acute aversive event. Sprague-Dawley male and female rats were exposed to restraint and 60, 1 s, 1 mA tailshocks and returned to their home cage. Either 2 or 24 h later, they were placed in a novel environment with a familiar cage-mate and an unfamiliar conspecific of the same sex. Relative to unstressed controls and females, males stressed 2 h previously decreased the exploration of the unfamiliar conspecific, exhibiting a rapid decrease over the course of the trial. In response to the stressor, however both sexes, however, decreased the exploration of the familiar conspecific, decreased their overall activity, and returned preferentially to their starting quadrant. None of these stress-induced effects were evident 24 h later upon the first or second exposure to the apparatus. Thus, exposure to the stressor transiently increased perseveration and decreased activity in males and females, but only decreased the exploration of novel conspecifics in males. These results indicate that a number of behavioral responses to stressors are conserved across gender, but those relating to novelty are more pronounced in males.

Animals↗

PubMed1979

Recognition among mice. Evidence from the use of a Y-maze differentially scented by congenic mice of different major histocompatibility types.

Previous studies of mating preference signified that mice can sense one another's major histocompatibility complex (MHC) types, probably by olfaction. This conclusion has now been substantiated by the use of a Y-maze whose two arms were differentially scented with currents of air conducted through boxes occupied by B6 (H-2b) males and by B6-H-2k congenic males. Four B6 mice, two males and two females, were successfully trained, by water deprivation and reward, to enter the arm scented by B6 or B6-H-2k males. One of the males and one of the females were trained to select the B6-scented arm; the other male and female were trained to select the B6-H-2k-scented arm. Untrained mice showed no MHC discrimination in the maze. The performance of the trained mice in distinguishing between MHC congenic homozygous F2 segregants derived from a cross of B6-H-2k with B6 was as good as their performance in distinguishing the respective inbred strains, thus essentially eliminating alternative and significant additional explanations of MHC-associated sensory discrimination. The data further indicate that chemosensory discrimination of MHC types can be entirely dissociated from sex differences and from the circumstances of mating.

Animal Communication↗

PubMed2004

The role of male age, sperm age and mating history on fecundity and fertilization success in the hide beetle.

Models of age-related mate choice predict female preference for older males as they have proven survival ability. However, these models rarely address differences in sperm age and male mating history when evaluating the potential benefits to females from older partners. We used a novel experimental design to assess simultaneously the relative importance of these three parameters in the hide beetle, Dermestes maculatus. In a two-part experiment we first explored age-related male mating success and subsequently examined the consequences of male age, sperm age and male mating history on female fecundity and fertilization success. In a competitive mating environment, intermediate-age males gained significantly higher mating success than younger or older males. To test the consequences for females of aged-related male mating success, a second set of females were mated to males varying in age (young, intermediate-age and old), in numbers of matings and in timing of the most recent mating. We found that male age had a significant impact on female fecundity and fertilization success. Females mated to intermediate-age males laid more eggs and attained consistently higher levels of fertilization success than females with young and old mates. A male's previous mating history determined his current reproductive effort; virgin males spent longer in copula than males with prior mating opportunities. However, differences in copulation duration did not translate into increased fecundity or fertilization success. There was also little evidence to suggest that fertilization success was dependent on the age of a male's sperm. The experiment highlights the potential direct benefits accrued by females through mating with particular aged males. Such benefits are largely ignored by traditional viability models of age-related male mating success.

Age Factors↗