Sperm: seminal fluid interactions and the adjustment of sperm quality in relation to female attractiveness

An important predictor of male fitness is the fertilizing efficiency of their ejaculates. Ejaculates are costly to produce and males are predicted to devote greater resources to copulations with reproductively superior females. It is well established that males allocate different numbers of sperm to ejaculates. However, less is known about how males adjust their sperm quality, which has important implications for our understanding of fertilization and the evolution of sexual strategies. Here we test in the fowl, Gallus gallus, whether males adjust their sperm velocity by differentially allocating seminal fluid to copulations with attractive and unattractive females. To disentangle the contributions of sperm and seminal fluid to sperm velocity, we separated and remixed sperm and seminal fluid from ejaculates allocated to females of different attractiveness. We show that dominant males increase the velocity of the sperm they invest in more attractive females by allocating larger ejaculates that contain seminal fluid that increases sperm velocity. Furthermore, we find weak evidence that males also allocate sperm with higher velocity, irrespective of seminal fluid, to more attractive females.     

Co-evolution of male and female reproductive characters across the Scathophagidae (Diptera)

Sperm morphometry is extremely variable across species, but a general adaptive explanation for this diversity is lacking. As sperm must function within the female, variation in sperm form may be associated with variation in female reproductive tract morphology. We investigated this and other potential evolutionary associations between male and female reproductive characters across the Scathophagidae. Sperm length was positively associated with the length of the spermathecal (sperm store) ducts, indicating correlated evolution between the two. No association was found between sperm length and spermathecal size. However, the size of the spermathecae was positively associated with testis size indicating co-evolution between male investment in sperm production and female sperm storage capacity. Furthermore, species with a higher degree of polyandry (larger testes) had longer spermathecal ducts. However, no associations between sperm length or length variation and testis size were found which suggests greater sperm competition sensu stricto does not select for longer sperm.     

Sex allocation and sexual conflict in simultaneously hermaphroditic animals

Links between sex allocation (SA) and sexual conflict in simultaneous hermaphrodites have been evident since Charnov''s landmark paper published 30 years ago. We discuss two links, namely the potential for sexual conflict over SA between sperm donor and recipient, and the importance of post-copulatory sexual selection and the resulting sexual conflict for the evolution of SA. We cover the little empirical and theoretical work exploring these links, and present an experimental test of one theoretical prediction. The link between SA and sexual conflict is an interesting field for future empirical and theoretical research.     

Effects of Sexual Selection Upon Sperm Morphology and Sexual Skin Morphology in Primates

We review possible effects of sexual selection upon sperm morphology, and sexual skin morphology, in primates. Comparative morphometric studies, involving 31 species representing 21 primate genera, revealed a positive relationship between volume of the sperm midpiece, occurrences of multiple partner matings by females, and large relative testes sizes, which indicate sperm competition. The midpiece houses the mitochondria required to power sperm motility. Hence, sperm competition may have influenced the evolution of increased mitochondrial loading in species where females mate with multiple partners during the fertile period. Females of some Old World monkey species and female chimpanzees exhibit large estrogen-dependent sexual skin swellings during the follicular phase of the menstrual cycle. Studies of mandrills support the conclusion that swellings act primarily as sexually attractive, graded signals and that swelling size may indicate current reproductive quality. Measurements of the genitalia in chimpanzees indicate a secondary function for female swellings. The swelling increases the operating depth of the female's vagina by 50% during the fertile phase of her cycle. Males have evolved long, filiform penes capable of placing sperm close to the os cervix during competitive multipartner matings. This may exemplify how morphologic specializations in females can influence the coevolution of advantageous genitalic specializations in males: the phenomenon that Eberhard (1985) dubbed cryptic female choice.     

Male–female relatedness and patterns of male reproductive investment in guppies

Inbreeding can cause reductions in fitness, driving the evolution of pre- and postcopulatory inbreeding avoidance mechanisms. There is now considerable evidence for such processes in females, but few studies have focused on males, particularly in the context of postcopulatory inbreeding avoidance. Here, we address this topic by exposing male guppies (Poecilia reticulata) to either full-sibling or unrelated females and determining whether they adjust investment in courtship and ejaculates. Our results revealed that males reduce their courtship but concomitantly exhibit short-term increases in ejaculate quality when paired with siblings. In conjunction with prior work reporting cryptic female preferences for unrelated sperm, our present findings reveal possible sexually antagonistic counter-adaptations that may offset postcopulatory inbreeding avoidance by females.     

Sexual selection in hermit crabs: a review and outlines of future research

The information currently available on sexual selection in hermit crabs is reviewed to identify the role of males and females before, during and after mating. According to this information, possible mechanisms of male–male competition, female choice and/or sexual conflict are suggested. Important male components that may affect mating success include dragging the female shell, rotations of the female's shell and male cheliped palpations, and male size and/or shell characteristics (species and size). Possible female determinants of male mating/fertilization success include size (as an indicator of egg production capacity), signalling of sexual receptivity to males, delay from mate guarding to copulation and mating duration. Avenues for deeper exploration in males include the role of the number and morphometry of male sexual tubes during sperm transfer, and whether ejaculate size and sperm number can be adjusted with variable situations of sperm competition intensity and risk. In females it would be interesting to investigate the chemical and behavioural mechanisms affecting spermatophore breakage for sperm release and the variable duration from sperm transfer to spawning. Given these possibilities, and that sperm is externally deposited on the female's body but inside her shell (except for those species that do not use shells, e.g. Birgus , or species where shells are rather small and do not cover the body totally, e.g. Parapagurus ), we conclude that hermit crabs are unique subjects for separating male and female effects, particularly with respect to the applicability of current ideas in sexual selection such as female choice and sexual conflict. Some practical ideas are provided to disentangle both hypotheses using these animals.     

PATTERNS OF PATERNITY SKEW AMONG POLYANDROUS SOCIAL INSECTS: WHAT CAN THEY TELL US ABOUT THE POTENTIAL FOR SEXUAL SELECTION?

Monogamy results in high genetic relatedness among offspring and thus it is generally assumed to be favored by kin selection. Female multiple mating (polyandry) has nevertheless evolved several times in the social Hymenoptera (ants, bees, and wasps), and a substantial amount of work has been conducted to understand its costs and benefits. Relatedness and inclusive fitness benefits are, however, not only influenced by queen mating frequency but also by paternity skew, which is a quantitative measure of paternity biases among the offspring of polyandrous females. We performed a large‐scale phylogenetic analysis of paternity skew across polyandrous social Hymenoptera. We found a general and significant negative association between paternity frequency and paternity skew. High paternity skew, which increases relatedness among colony members and thus maximizes inclusive fitness gains, characterized species with low paternity frequency. However, species with highly polyandrous queens had low paternity skew, with paternity equalized among potential sires. Equal paternity shares among fathers are expected to maximize fitness benefits derived from genetic diversity among offspring. We discuss the potential for postcopulatory sexual selection to influence patterns of paternity in social insects, and suggest that sexual selection may have played a key, yet overlooked role in social evolution.     

Sexuelle Selektion und die Evolution von Kopulationen au?erhalb des Paarbundes: Spielregeln der Weibchen

Zusammenfassung Seit Darwins grundlegendem Werk von 1871 über sexuelle Selektion ist es kein Geheimnis mehr, daß der Pfau sein Rad schlagen kann, weil die Weibchen es so wollen. Anerkanntermaßen ist der Pfau polygam; dasjenige Männchen mit dem prächtigsten Gefieder hat die meisten Weibchen und den größten Reproduktionserfolg. Durch die Zuchtwahl der Weibchen (female choice) wurden die Männchen so schön, wie sie es heute sind. In monogamen Paarungssystemen konnte man auffällig gefärbte Männchen auf die gleiche Weise allerdings nicht erklären. Dies wurde erst möglich, seit sich die Erkenntnis durchgesetzt hat, daß auch bei monogamen Arten gewisse Männchen durch Fremdkopulationen mehr Nachkommen haben als andere. Erst seit wenigen Jahren ist bekannt, daß es häufig die Weibchen sind, die die aktive Rolle beim Vollzug von Seitensprüngen spielen. Welche selektiven Vorteile können Weibchen durch dieses Verhalten erlangen?Die basale Theorie der sexuellen Selektion wird auf der Grundlage klassischer Arbeiten besprochen. Darauf fußend wird ein überblick über die aktuelle Diskussion von Fremdkopulationen (extra-pair copulations [EPCs]) gegeben. Die für Weibchen möglichen Fitness-Gewinne, die zur Evolution von EPCs geführt haben können, werden zusammengefaßt. Ein Zusammenhang wird hergestellt unter dem Gesichtspunkt, wie sich EPCs jeweils auf den Gesamtfortpflanzungserfolg (total reproductive value) der Weibchen auswirken. Die Notwendigkeit wird erläutert, die theoretischen Ansätze aus einem übergeordneten Blickwinkel zu betrachten, ohne von vornherein die eine oder andere Erklärung auszuschließen.Als gelungenes Beispiel wird u. a. die Hypothese der eingeschränkten Weibchen (constrained female hypothesis) von Gowaty (1996) besprochen. Diese besagt, daß Weibchen zur Erlangung eines beliebigen zusätzlichen Fitness-Gewinns auf EPCs angewiesen sein können, da sie in ihrer Wahl eines passenden Partners oft behindert sind (z. B. durch die Männchen selber). Schließlich werden mögliche Ansatzpunkte zur Überprüfung der Hypothesen in Feldstudien aufgezeigt; zukünftig sollte vermehrt auch die Qualität der Weibchen als Grundlage für deren Verhaltensentscheidungen betrachtet werden.

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Sexual selection and the evolution of extra-pair copulation: rules of the game from the females' point of view

Summary In view of Darwin's fundamental thesis on sexual selection, which appeared in 1871, it is no longer a secret that a peacock can spread his tail because the females want it that way. It is generally acknowledged that the peacock is polygamous; whichever male has the most attractive plumage wins the most females and therefore achieves the greatest reproductive success. It is due to selective breeding by the females, i. e. female choice that the males have become what they are today. In monogamous mating systems, however, the source of striking colouring in males could not be explained the same way until evidence for the common occurrence of extra-pair copulation became more accepted. This mating behaviour offers an explanation of how certain males, even in monogamous species, can produce more offspring than others. Only in recent years was it recognised that it is often the females which play the active role in the initiation of extra-pair copulation. What fitness gains can the females expect to achieve through this behaviour?This review commences with an introduction to fundamental theories of sexual selection. Progressing from this, the current discussion of extra-pair copulation (EPC) is reviewed. Conceivable fitness gains for the females, which may have resulted in the evolution of EPC, are summarised. A connection is noted between the various possible fitness gains in their effect on the total reproductive value of the females. The necessity of considering all of these theories from a more general perspective, without having to dismiss any explanations from the outset, is made clear. Gowaty's constrained female hypothesis (1996) is one example in which this has been achieved. This hypothesis proposes that females can be obliged to engage in EPC in order to obtain any kind of extra fitness gain, since they are often constrained in their choice of a partner (e.g. by the males themselves). In conclusion, possible directions are suggested for the testing of these hypotheses in field studies; in future more emphasis should be put on intrinsic quality differences between the females while investigating their mating behaviour.