Operational sex ratio influences the opportunity for sexual selection in guppies

The opportunity for sexual selection was greater when the operational sex ratio (OSR) in guppies Poecilia reticulata was biased towards males. This could be due to an increase in both male-male competition and female mate choice under male-biased OSR.     

Offspring sex ratio in the Common Tern Sterna hirundo, a species with negligible sexual size dimorphism

In many vertebrates, male offspring are affected more than female offspring by adverse conditions during growth, resulting in facultative adjustment of offspring sex ratio by parents in response to social and environmental conditions during breeding. The greater vulnerability of male offspring is generally attributed to their higher energy requirements associated with their larger size, although greater sensitivity to adverse conditions could be related to other factors such as negative effects of androgens on male physiology. To control for sexual differences in body size, we examined variation in offspring sex ratio in the Common Tern Sterna hirundo , a species with negligible sexual size dimorphism. In this species, the last-laid egg (termed the c-egg) is smaller than the first two and hatches last, so that the chick obtains relatively little food and hence has a low probability of survival to fledging. This species thus provides a powerful model for examining sex-linked mortality and sex ratio variation under natural conditions. We found that the sex ratio of c-eggs, but not of earlier laid eggs, was significantly biased in favour of females. Chicks hatched from c-eggs (termed c-chicks) had low survival but female c-chicks had significantly higher survival than male c-chicks. These data provide strong evidence that factors other than sexual size dimorphism are responsible for producing greater vulnerability of male offspring to adverse conditions during growth.     

Patterns of fluctuating asymmetry in sexual ornaments predict female choice

Extravagant secondary sexual characters are assumed to have arisen and be maintained by sexual selection. While traits like horns, antlers and spurs can be ascribed to intrasexual competition, other traits such as extravagant feather ornaments, displays and pheromones have to be ascribed to mate choice. A number of studies have tested whether females exert selection on the size of male ornaments, but only some of these have recorded female preferences for the most extravagantly ornamented males. Here I demonstrate that female choice can be directly predicted from the relationship between the degree of fluctuating asymmetry and the size of a secondary sexual character. Fluctuating asymmetry is an epigenetic measure of the ability of individuals to cope with stress, and it occurs when an individual is unable to undergo identical development of an otherwise bilaterally symmetric trait on both sides of its body. There is a negative relationship between the degree of fluctuating asymmetry and the absolute size of an ornament in those bird species with a female preference for the largest male sex trait, while there is a flat or U-shaped relationship among species without a female preference. These results suggest that females prefer exaggerated secondary sexual characters if they reliably demonstrate the ability of males to cope with genetic and environmental stress. Some species may demonstrate a flat or U-shaped relationship between the degree of fluctuating asymmetry and the absolute size of an ornament because (i) the genetic variance in viability signalled by the secondary sex trait has been depleted; (ii) the secondary sex trait is not particularly costly and therefore does not demonstrate condition dependence; or because (iii) the sex traits can be considered arbitrary traits rather than characters reflecting good genes.     

Directional postcopulatory sexual selection is associated with female sperm storage in Trinidadian guppies

Female sperm storage (FSS) is taxonomically widespread and often associated with intense sperm competition, yet its consequences on postcopulatory sexual selection (PCSS) are poorly known. Theory predicts that FSS will reduce the strength of PCSS, because sperm characteristics favored before and after FSS may be traded‐off, and opportunities for nondirectional PCSS should increase. We explored these questions in the guppy (Poecilia reticulata), by allowing females to mate multiply and by comparing the paternity pattern in two successive broods. Contrary to predictions, the variance in male fertilization success increased after FSS, driven by a change in male paternity share across broods. This change was positively associated with sperm velocity (measured before FSS) but not with the duration of FSS, indirectly suggesting that faster sperm were better in entering female storage organs, rather than in persisting within them. Other male traits, such as male size and orange color, heterozygosity, and relatedness to the female, did not influence paternity after FSS. These results indicate that processes associated with FSS tend to reinforce the strength of PCSS in guppies, rather than weaken it. Further work is necessary to test whether this pattern changes in case of more prolonged FSS.     

Control of sex ratio by female spider mites

Control of sex ratio in the progenies of mated female spider mites was investigated in two laboratory experiments. In Experiment I, a single strain of Tetranychus urticae Koch was reared in four different environments. In Experiment II, 4 different geographic strains of T. urticae were reared in a single environment.Results indicated that spider mite females were able to control the sex ratio of their progeny at two levels: an initial control of mean population ratio according to ovipositional environment. In addition, a secondary control resulted in an approximately uniform daily ratio. Although each parental female produced a varying number of eggs from day to day, a more or less constant fraction of those eggs were fertilized every day. It is concluded that both of these characteristics are probably important to the colonizing ability of the species.

---

Résumé L'étude de la régulation du sexe de la descendance de femelles inséminées d'acariens est basée sur deux expériences de laboratoire. Dans l'expérience I, une lignée de Tetranychus urticae a été élevée dans deux conditions différentes de qualité des feuilles pendant le développement des parents, d'une part, pendant leur ponte et le développement de la descendance d'autre part. Dans l'expérience II, quatre souches géographiques différentes de T. urticae ont été élevées dans les mêmes conditions.Il est apparu nettement que les femelles dans des conditions différentes ont donné des descendances avec des taux sexuels variant dans certaines limites. Ces différences étaient statistiquement significatives dans les duex lots de données expérimentales. Les taux sexuels des populations d'acariens variaient aussi avec les conditions écologiques. Dans les bonnes conditions, la proportion de femelles dans la descendance était la plus élevée. Les taux sexuels des populations variaient aussi entre les quatre souches géographiques.Les analyses ont aussi révélé que les femelles d'acariens sont capables de contrôler le taux sexuel à deux niveaux: une régulation initiale du taux de la population en fonction des conditions écologiques ou de la lignée géographique et un contrôle secondaire qui s'est traduit par un taux quotidien approximativement uniforme.En utilisant un argument mathématique simple incorporant la fécondité caractéristique des acariens, nous avons conclu que ces deux caractéristiques de la rulation sont probablement d'une importance capitale pour le pouvoir colonisateur de cette espèce.

     

Offspring sex ratio skew in the sexually monomorphic house martin Delichon urbicum

Sex ratio at conception may be under selection pressure, given that male and female offspring differ in the cost of production or generate different fitness returns under specific conditions. We studied adjustments in the primary, secondary and tertiary sex ratio in house martin Delichon urbicum, which is a sexually monomorphic, socially monogamous, colonial bird. Males of this species engage in extra‐pair copulations with heavy males acquiring the highest fertilization success. We analyzed variation in the sex ratio in relation to clutch size and parental characteristics including body condition, wing length, as well as length and pigmentation of the white rump patch during three breeding seasons. The only variable which significantly explained the variation in the sex ratio was wing length of the social father and mother. The proportion of sons among offspring was positively correlated to wing length of the social father and negatively correlated to mother wing length. Social father wing length positively correlated with mean brood body mass at fledging, which may suggest that females that mated with long‐winged males produced sons, which acquired the highest total fertilization success. Consequently, our results indicate that house martin females may adaptively adjust offspring sex composition at egg laying in relation to the characteristics of their social mate.     

Non-seasonal changes in the intensity of female mate preference and offspring sex ratio in the wild guppy Poecilia reticulata

It has been demonstrated that the exaggeration of male sexual ornaments and the intensity of female mate preferences of a wild guppy population change over a period of several months. However, the factors that determine the short-term changes in male ornaments and female preferences remained unclear. In this study, we examined the effect of season on these short-term changes by measuring these traits in the same seasons of different years for a wild guppy population in Okinawa, Japan. We also compared the characteristics of the offspring in each collection term, as female guppies are known to have the ability to control offspring characteristics, such as brood size and sex ratios, depending on their mates' attractiveness. Results showed that the total lengths of the males changed seasonally; males in the summer were larger than those in the spring. In contrast, the size of orange spots in males and the intensity of female mating preferences differed in the same seasons of different years. Brood size and offspring body size in each term showed seasonal changes. However, offspring sex ratios exhibited different patterns in the same seasons of different years. Females produced female-biased broods when attractive males with large orange spots were rare. These results suggest that short-term changes in some traits of adult male and female guppies as well as offspring sex ratios may be not determined by seasonal factors, and that these traits may be interrelated.     

Operational sex ratio, sexual conflict and the intensity of sexual selection

Modern sexual selection theory indicates that reproductive costs rather than the operational sex ratio predict the intensity of sexual selection. We investigated sexual selection in the polygynandrous common lizard Lacerta vivipara . This species shows male aggression, causing high mating costs for females when adult sex ratios (ASR) are male-biased. We manipulated ASR in 12 experimental populations and quantified the intensity of sexual selection based on the relationship between reproductive success and body size. In sharp contrast to classical sexual selection theory predictions, positive directional sexual selection on male size was stronger and positive directional selection on female size weaker in female-biased populations than in male-biased populations. Thus, consistent with modern theory, directional sexual selection on male size was weaker in populations with higher female mating costs. This suggests that the costs of breeding, but not the operational sex ratio, correctly predicted the strength of sexual selection.     

Perspective: female remating,operational sex ratio,and the arena of sexual selection in Drosophila species

Abstract.— As commonly observed among closely related species within a variety of taxa, Drosophila species differ considerably in whether they exhibit sexual dimorphism in coloration or morphology. Those Drosophila species in which male external sexual characters are minimal or absent tend, instead, to have exaggerated ejaculate traits such as sperm gigantism or seminal nutrient donations. Underlying explanations for the interspecific differences in the presence of external morphological sexual dimorphism versus exaggerated ejaculate traits are addressed here by examining the opportunity for sexual selection on males to occur before versus after mating in 21 species of Drosophila . Female remating frequency, an important component of the operational sex ratio, differs widely among Drosophila species and appears to dictate whether the arena of sexual selection is prior to, as opposed to after, copulation. Infrequent female mating results in fewer mating opportunities for males and thus stronger competition for receptive females that favors the evolution of male characters that maximize mating success. On the other hand, rapid female remating results in overlapping ejaculates in the female reproductive tract, such that ejaculate traits which enhance fertilization success are favored. The strong association between female remating frequency in a given species and the presence of sexually selected external versus internal male characters indicates that the relationship be examined in other taxa as well.     

Manipulation of offspring sex ratio by second-mated female house wrens

In 1973, Trivers and Willard proposed that offspring sex ratio should be associated with the quality of parental care likely to be provided to the offspring. We tested this hypothesis by comparing fledgling sex ratios in nests of first- and second-mated female house wrens (Troglodytes aedon). In our Wyoming population, second-mated females typically receive little or no male parental assistance and fledge fewer and lower-quality young compared with first-mated females. Assuming that being of lower quality has stronger negative effects on the future reproductive success of males than that of females in this polygynous population, we predicted that fledgling sex ratios in the nests of second-mated females would be female-biased compared with the fledgling sex ratios of first-mated females. Additionally, we asked whether any sex bias at fledging could have resulted from male-biased nestling mortality caused by sex-biased parental provisioning. As predicted, mean fledgling sex ratios in nests of second-mated females were more female-biased than fledgling sex ratios in nests of first-mated females. However, we found no evidence of either sex-biased nestling mortality or sex-biased parental provisioning. These findings suggest that females are responding to their status as second-mated females and to the associated low-quality parental care that their young are likely to receive by producing female-biased clutches rather than manipulating the offspring sex ratio through sex-biased nestling mortality.     

Offspring allocation in externally ovipositing fig wasps with varying clutch size and sex ratio

The simultaneous optimization of clutch size and sex ratio isa tricky problem. Unless parameters such as host size or fecundityexist to pin down the optimal clutch size, this problem remainselusive to analytical analysis. This is because the fitnesslandscape with respect to clutch size and sex ratio does nothave one single evolutionarily stable peak toward which thepopulation can evolve. To solve this problem, I used a computeremulation to optimize both clutch size and sex ratio using externallyovipositing fig wasps as a model taxon. The simulation approachallows the use of integer numbers of eggs rather than assumingthat females can produce any sex ratio between 0 and 1. Whenfemales have no information about the patches on which theyoviposit, they produce either large clutches with a strong femalebias or clutches of a single male egg. When females have completeknowledge of their oviposition site, a set of conditional substrategiesis evolutionarily stable. Again, these substrategies are eitherlarge clutches with a female bias or dutches consisting of asingle male egg. This dichotomous oviposition pattern resultsin unrelated males sharing a fig, a condition conducive to theevolution of fatal fighting. Selection on female ovipositionstrategies may therefore be an important driving force behindhigh levels of fighting observed between male fig wasps.     

Offspring sex ratio in the sequentially polygamous Penduline Tit Remiz pendulinus

Despite the growing literature on facultative sex-ratio adjustment in chromosomal sex-determining vertebrate taxa (birds, mammals), the consistency of results is often low between studies and species. Here, we investigate the primary and secondary offspring sex ratio of a small passerine bird, the Eurasian Penduline Tit (Remiz pendulinus) in three consecutive years. This species has a uniquely diverse breeding system, in which the male (and/or the female) abandons the nest during egg-laying, and starts a new breeding attempt. This allowed us to test (1) whether patterns of parental care, i.e., male-only care, female-only care or biparental desertion, influence offspring sex ratio, and (2) whether the offspring sex ratio is repeatable between successive clutches of males and females. Using molecular markers to sex 497 offspring in 176 broods, we show that (1) offspring sex ratio does not depend on which parent provides care, and (2) the offspring sex ratio is not repeatable between clutches of a given individual. The overall primary and secondary offspring sex ratio at a population level is not different from parity (54 ± 6% males, and 50 ± 3% (mean ± SE), respectively). We suggest that ecological and phenotypic factors, rather than individual traits of parents, may influence offspring’s sex, and conclude that there is currently no evidence for a facultative adjustment of offspring sex ratio in the Penduline Tit.     

Offspring and sex ratio are independent of sperm management in Eupelmus orientalis females

Sperm stocks in both males and females of the parthenogenetic wasp Eupelmus orientalis were investigated at various points during reproduction and compared to the progeny of females in controlled conditions. One day-old virgin males had approximately 5500 sperm, and from a total of about 1697 sperm transferred per copulation, 21% are stored in the spermathecae by females 24 hours after mating. At the end of the egg-laying period (at least 42 days), 2/5 of the initial amount of sperm remained in this storage organ. This decrease (from approximately 350 to 150) occurred essentially during the first 21 days of egg-laying activity, indicating that the majority of sperm stored were used during this period. Between 21 days and the end of fertile life, the number of sperm remained constant. The mean offspring production throughout reproductive life after one mating was 153, with 56.5% of the daughters laid at the beginning of the laying activity. Sex ratio was entirely female biased during the first 15 days (mean=0.65), then it decreased and became nearly equal after 20 days. Present results propose that females maximize the production of daughters i.e. of inseminated eggs until the 20th day and after this time lay as many daughters and sons despite their still having stored sperm. Physiological constraints due to ageing are proposed to explain this phenomenon.     

The strategic use of sex in wild female western gorillas

Human females, unlike most mammals, are sexually active outside of fertile periods. This decoupling of sexual behavior from its conceptive function has had an enormous impact on human social relationships, and yet we know little about why there was selection for nonconceptive mating. Here we examine one form of nonconceptive mating, the mating that occurs during pregnancy or post‐conceptive (PC) mating, in wild western gorillas (Gorilla gorilla). Using a near complete mating record for five females during gestation, we show that pregnant females varied in the timing and frequency of mating, and used PC mating conditionally, synchronizing copulations to occur on days when other females mated, and refraining from mating for lengthy periods when no other females mated. As pregnant females mated exclusively with the same male before and after conception, and mated in response to group female (and not male) behavior, we conclude that western gorillas used PC mating as a form of female competition, and not to confuse paternity or to obtain immediate benefits from the male, as suggested earlier. The male initiated copulations preferentially with females of high rank, rather than distinguishing between pregnant and cycling females. Therefore, PC mating appears to be a strategy by which high‐ranking pregnant females attempt to minimize male interest in other females, while reinforcing their own status and potentially delaying conception in others. These findings indicate that female‐mating competition is more important than considered earlier, and may be a factor in the evolution of nonconceptive mating in humans. Am. J. Primatol. 71:1011–1020, 2009. © 2009 Wiley‐Liss, Inc.     

The effects of sex ratio on sexual competition in the European lobster

During the breeding season an individual's access to mates may be affected by operational sex ratios, causing strong variation in mating success. We manipulated adult sex ratios of the European lobster, Homarus gammarus, to test the predictions of models that relate sexual competition to (1) the sex ratio, (2) the time that an individual is not available to mate and (3) 'collateral investment', whereby two males contribute to a single clutch. The model predictions proved to be relatively insensitive to collateral investment. Male-male competition predominated in the male-biased but not in the female-biased sex ratio. This matches the predictions of one model that incorporates an extended period of female receptivity because the time that a male was unavailable to mate was small compared to the time spent by females in cohabitation and parental care. Although females increased their competitiveness when males were in the minority, male competition remained high. The insensitivity of male-male competition to sex ratios may be due to an upper limit to the costs that males can afford when there is a serious risk of injury, preventing males from increasing their aggression when females are in short supply. Copyright 1999 The Association for the Study of Animal Behaviour.     

Isolation of sex pheromone mimic of the American cockroach by monitoring with male/female ratio in electroantennogram

The electroantennogram (EAG) technique was applied as a monitoring tool to isolate germacrene-D, a sex pheromone mimic of the American cockroach. Large values of the male/female ratio, and its index derived from EAG responses of adult males and females to a standard, indicated significant amounts of germacrene-D in plant fractions. Monitoring with behavioural assays and gas chromatographic analysis during the isolation confirmed that the EAG technique using the male/female index was a good indicator in the isolation of sexually active components.     

Regulation of sexual odor preference by sex steroids in the posterodorsal medial amygdala in female rats

Our previous study in male rats demonstrated that bilateral administration of flutamide, an androgen receptor (AR) antagonist, into the posterodorsal medial amygdala (MePD) increased the time sniffing male odors to as high as that sniffing estrous odors, eliminating the preference for estrous odors over male odors. This made us speculate that under blockade of AR in the MePD, testosterone-derived estrogen acting on the same brain region arouses interest in male odors which is otherwise suppressed by concomitant action of androgen. In cyclic female rats, endogenous androgen has been thought to be involved in inhibitory regulation of estrogen-activated sexual behavior. Thus, in the present study, we investigated the possibility that in female rats the arousal of interest in male odors is also normally regulated by both estrogen and androgen acting on the MePD, as predicted by our previous study in male rats. Implantation of either the estrogen receptor blocker tamoxifen (TX) or a non-aromatizable androgen 5α-dihydrotestosterone (DHT) into the MePD of ovariectomized, estrogen-primed female rats eliminated preference for male odors over estrous odors by significantly decreasing the time sniffing male odors to as low as that sniffing estrous odors. The subsequent odor discrimination tests confirmed that the DHT and TX administration did not impair the ability to discriminate between male and estrous odors. These results suggest that in estrous female rats estrogen action in the MePD plays critical roles in the expression of the preference for male odors while androgen action in the same brain region interferes with the estrogen action.