No intra-locus sexual conflict over reproductive fitness or ageing in field crickets

Differences in the ways in which males and females maximize evolutionary fitness can lead to intra-locus sexual conflict in which genes delivering fitness benefits to one sex are costly when expressed in the other. Trade-offs between current reproductive effort and future reproduction and survival are fundamental to the evolutionary biology of ageing. This leads to the prediction that sex differences in the optimization of age-dependent reproductive effort may generate intra-locus sexual conflict over ageing rates. Here we test for intra-locus sexual conflict over age-dependent reproductive effort and longevity in the black field cricket, Teleogryllus commodus. Using a half-sib breeding design, we show that the most important components of male and female reproductive effort (male calling effort and the number of eggs laid by females) were positively genetically correlated, especially in early adulthood. However, the genetic relationships between longevity and reproductive effort were different for males and females, leading to low genetic covariation between male and female longevity. The apparent absence of intra-locus sexual conflict over ageing suggests that male and female longevity can evolve largely independently of one another.     

Influence of sex-specific stimuli on ovarian maturation in primary and secondary reproductives of the dampwood termite Zootermopsis angusticollis

Abstract. It was hypothesized that female primary and secondary reproductives of the termite Zootermopsis angusticollis Hagen require sex-specific stimuli from a reproductive male to trigger ovarian maturation. To test this hypothesis, experimental laboratory colonies were established that contained female primary or secondary reproductives nesting: alone; with a male; with a female; and with three third- to fourth-instar larvae. Following colony initiation, we measured changes in body mass, ovariole number and fecundity over 60 days. Results indicate that the reproductive maturation of female primaries was promoted by contact with a male but inhibited by the presence of another female. Secondary females were not affected by the presence of another reproductive, regardless of sex, but the development of reproductive competency of primary females appeared to depend on male-specific stimulation. Reliance on male–female interaction to induce female reproductive development may ensure that the resources of newly dealate females are not wasted on producing larvae that would have a poor chance of surviving in the absence of a male. By contrast, secondary females maturing within established colonies are likely to have a mate and immediate assistance with non-reproductive tasks, and therefore do not delay ovarian maturation and reproduction until they perceive male-specific stimuli. These results demonstrate that male-specific stimuli affect only the reproductive development of primary females, and suggest that the degree to which primary and secondary females depend on mate assistance may have shaped their physiological responses to the presence of a reproductive male.     

Male parental care, female reproductive success, and extrapair paternity

Birds differ considerably in the degree of male parental care,and it has been suggested that interspecific variation in extrapairpaternity is determined by the relative importance of benefitsto females from male parental care and good genes from extrapairsires. I estimated the relationship between extrapair paternityand the importance of male parental care for female reproductivesuccess mainly based on male removal studies, using a comparativeapproach. The reduction in female reproductive success causedby the absence of a male mate was positively correlated withthe male contribution to feeding offspring. The frequency ofextrapair paternity was negatively related to the reductionin female reproductive success caused by the absence of a mate.This was also the case when potentially confounding variablessuch as developmental mode of offspring and sexual dichromatismwere considered. A high frequency of extrapair paternity occursparticularly in bird species in which males play a minor rolein offspring provisioning and in which attractive males providerelatively little parental care. Bird species with frequentextrapair paternity thus appear to be those in which direct fitness benefits from male care are small, females can readilycompensate for the absence of male care, and indirect fitnessbenefits from extrapair sires are important.     

Dietary self-selection and discrimination threshold in wild Anastrepha obliqua females (Diptera,Tephritidae)

Experiments were performed on adult wild Anastrepha obliqua females to (I) determine whether there is a critical period during the reproductive phase when the absence of protein in the diet impairs egg production; (II) determine whether the females are able to self-select an optimal diet; and (III) determine the discrimination threshold for protein in newly emerged females as well as in 10-day old females that had either been deprived of protein or not deprived. The results showed that protein ingestion is essential during the preovipositing phase to allow egg production, that females are self-selective and that this behavior changes with age according to the reproductive status, and that protein deprivation strongly alters the discrimination threshold for protein. Newly emerged females have discriminatory thresholds similar to protein-deprived insects.     

Early growth,dominance acquisition and lifetime reproductive success in male and female cooperative meerkats

In polygynous species, variance in reproductive success is higher in males than females. There is consequently stronger selection for competitive traits in males and early growth can have a greater influence on later fitness in males than in females. As yet, little is known about sex differences in the effect of early growth on subsequent breeding success in species where variance in reproductive success is higher in females than males, and competitive traits are under stronger selection in females. Greater variance in reproductive success has been documented in several singular cooperative breeders. Here, we investigated consequences of early growth for later reproductive success in wild meerkats. We found that, despite the absence of dimorphism, females who exhibited faster growth until nutritional independence were more likely to become dominant, whereas early growth did not affect dominance acquisition in males. Among those individuals who attained dominance, there was no further influence of early growth on dominance tenure or lifetime reproductive success in males or females. These findings suggest that early growth effects on competitive abilities and fitness may reflect the intensity of intrasexual competition even in sexually monomorphic species.     

Hormonal and behavioral changes at puberty in the squirrel monkey

Developmental changes in the reproductive behavior and physiology of 9 male and 15 female juvenile squirrel monkeys were evaluated in a 20-month study. Plasma levels of gonadal steroids remained relatively low for this species until most animals reached puberty between 2.5 and 3 years of age. Longitudinal assessment of plasma progesterone levels indicated that the onset of ovarian cycles tended to be synchronized between females although the 5 heaviest females began to cycle earlier than the rest. The heavier females reached puberty at a time which was appropriate to their birth in the wild, whereas most of the remaining females conceived 6 months later during a second period of reproductive activity that coincided with the laboratory mating season. Pubescent males underwent their first seasonal elevation in plasma testosterone levels during the second period and its onset was synchronized across all males. Thus, even in the absence of adults, pubertal processes in the squirrel monkey were strongly influenced by the seasonal breeding pattern. In addition, behavioral observations revealed that social maturation closely parallels reproductive ability in females, whereas males enter a protracted subadult stage after puberty.     

Reproductive cycle of the blacknose shark Carcharhinus acronotus in the Gulf of Mexico

The reproductive periodicity of blacknose sharks Carcharhinus acronotus in the northern Gulf of Mexico was determined by examining reproductive tissues from specimens collected monthly from 2002 to 2005. Monthly changes in shell gland mass, right ovary mass and ovarian follicle diameter were assessed for 74 mature females. Temporal changes in testes mass, testes width and proportion of mature spermatocysts were examined for 64 mature males. Trends in female reproductive tissues suggested an annual peak in reproductive activity during June and July, while trends in male variables suggested an annual reproductive peak during May and June. Although male and female reproductive activity peaked in different months, a strong synchronicity existed between the proportion of mature spermatocysts and the diameter of the largest ovarian follicle. Based on these results, the mating season of blacknose sharks lasts from mid-May to July in the Gulf of Mexico. Maximum embryo sizes were observed in May, which suggested that partition occurs during late May or early June. Results indicate that blacknose sharks have a clearly defined annual cycle in the Gulf of Mexico. This conclusion is further supported by the complete absence of gravid females without vitellogenic ovarian follicles among all mature females examined.     

The Effects of Perceived Mating Opportunities on Patterns of Reproductive Investment by Male Guppies

Males pay considerable reproductive costs in acquiring mates (precopulatory sexual selection) and in producing ejaculates that are effective at fertilising eggs in the presence of competing ejaculates (postcopulatory sexual selection). Given these costs, males must balance their reproductive investment in a given mating to optimise their future reproductive potential. Males are therefore expected to invest in reproduction prudently according to the likelihood of obtaining future matings. In this study we tested this prediction by determining whether male reproductive investment varies with expected future mating opportunities, which were experimentally manipulated by visually exposing male guppies (Poecilia reticulata) to high or low numbers of females in the absence of competing males. Our experiment did not reveal consistent effects of perceived future mating opportunity on either precopulatory (male mate choice and mating behaviour) or postcopulatory (sperm quality and quantity) investment. However, we did find that male size and female availability interacted to influence mating behaviour; large males visually deprived of females during the treatment phase became more choosy and showed greater interest in their preferred female than those given continuous visual access to females. Overall, our results suggest males tailor pre- rather than postcopulatory traits according to local female availability, but critically, these effects depend on male size.     

Sociosexual behavior, male mating tactics, and the reproductive cycle of giraffe Giraffa camelopardalis

Female distribution exerts a major impact on male mating tactics. Giraffe cows have a reproductive cycle, and a social system, that should favor a male roaming reproductive tactic. We conducted a 2-year study of female Rothschild's giraffe (G. c. rothschildi) reproductive endocrinology in order to characterize attributes of the reproductive cycle and investigate how female endocrine and behavioral cues influence mating activity. We used non-invasive fecal steroid methods to determine reproductive state among females residing in a herd in a large outdoor enclosure. We found that females had an estrous cycle of 14.7 days and that they regularly had multiple ovarian cycles prior to conception. Adult males were more likely to associate with, and sexually investigate, females when they were cycling than when they were either pregnant or acyclic. During the estrous cycle, male-female proximity and sociosexual behavior were more pronounced during the probable fertile phase than the rest of the cycle. Sexual activity between giraffe coincided with the periovulatory period, with male interest in females peaking during the fertile window in the absence of proceptive behavior by females. We conclude that males detect reliable cues revealing female reproductive status and partition their reproductive effort in response to such cues. We propose that male giraffe adopt a roaming reproductive strategy with their large size, enabling them to search for and mate guard fertile females while minimizing metabolic costs.     

Reproductive success in relation to dominance rank in the absence of prime-age males in Barbary macaques

In some primate species dominance rank of males is correlated with reproductive success, whereas in other species this relationship is inconsistent. Barbary macaques (Macaca sylvanus) live in a promiscuous mating system in which males are ranked in a dominance hierarchy that influences their access to females. High-ranking males usually monopolize fertile females during their estrous period and show increased mating activities. Subadult males generally rank below adult males. For Barbary macaque females in the Gibraltar colony, there was no correlation between dominance status and reproductive success. Paternity data for 31 offspring collected over four consecutive breeding seasons were used to test whether male social rank was associated with reproductive success and whether reproductive success was mainly confined to a small number of males. Genetic variation was assessed using 14 microsatellite markers for a dataset of 127 individuals sampled in all five social groups of the Gibraltar colony. Paternity analysis was conducted for offspring in one social group only, where all in-group males were sampled. Eighty-three percent of the offspring could be assigned to an in-group candidate father; none of the extra-group males appeared to have sired an infant. Male dominance rank was not found to contribute to the observed variation in male reproductive output. Fifty-nine percent of the offspring was sired by two low-ranking males, whereas the two top-ranking males sired one-fifth. A highly significant correlation was found for male age and dominance rank. Reproductive success of subadult males might be explained by the gap in the age distribution of male group members. These missing prime males are usually regarded as serious competitors for older males. Subadult males may have gained easier access to females in their absence. In addition, the presence of inbreeding avoidance mechanisms, which might also have overpowered possible rank effects, cannot be excluded.     

Manipulating reproductive effort leads to changes in female reproductive scheduling but not oxidative stress

The trade‐off between reproductive investment and lifespan is the single most important concept in life‐history theory. A variety of sources of evidence support the existence of this trade‐off, but the physiological costs of reproduction that underlie this relationship remain poorly understood. The Free Radical Theory of Ageing suggests that oxidative stress, which occurs when there is an imbalance between the production of damaging Reactive Oxygen Species (ROS) and protective antioxidants, may be an important mediator of this trade‐off. We sought to test this theory by manipulating the reproductive investment of female mice (Mus musculus domesticus) and measuring the effects on a number of life history and oxidative stress variables. Females with a greater reproductive load showed no consistent increase in oxidative damage above females who had a smaller reproductive load. The groups differed, however, in their food consumption, reproductive scheduling and mean offspring mass. Of particular note, females with a very high reproductive load delayed blastocyst implantation of their second litter, potentially mitigating the costs of energetically costly reproductive periods. Our results highlight that females use strategies to offset particularly costly periods of reproduction and illustrate the absence of a simple relationship between oxidative stress and reproduction.     

Characterization of the estrous cycle in Octodon degus

We characterized the reproductive cycle of Octodon degus to determine whether reproductive maturation is spontaneous in juveniles and if ovarian cyclicity and luteal function are spontaneous in adults. Laboratory-reared prepubertal and adult females were monitored for vaginal patency and increased wheel-running. Sexual receptivity was assessed by pairing adult females with a male 1) continuously, 2) at the time of vaginal patency, or 3) following estradiol treatment. Blood samples were assayed for estradiol and progesterone concentrations on Days 1, 4, 8, and 16 relative to vaginal opening. Ovarian tissues were collected 6 and 16 days after behavioral estrus and 6 days after copulation for histology. In juveniles, the onset of cyclic vaginal patency and increased wheel-running activity was spontaneous, occurred in the absence of proximal male cues, and appeared at regular intervals (17.5 ± 1.4 days). In adults, vaginal patency and increased wheel-running occurred cyclically (21.2 ± 0.6 days) in the absence of proximal male cues, and these traits predicted the time of sexual receptivity. Corpora lutea develop spontaneously and are maintained for 12-14 days. The ovaries had well-developed corpora lutea 6 days after mating and 6 days after estrus without mating. Progesterone concentrations were highest in the second half of the cycle when corpora lutea were present and estradiol concentrations peaked on the day of estrus. Thus, female degus appear to exhibit a spontaneous reproductive cycle consistent with other Hystricognathi rodents. Octodon degus is a novel model with which to examine the mechanisms underlying different reproductive cycles.     

Circulating LH levels and the response to exogenous GnRH in the common mole-rat: implications for reproductive regulation in this social, seasonal breeding species

The effects of breeding season and reproductive status on male and female reproduction were investigated in the common mole-rat, Cryptomys hottentotus hottentotus, a cooperatively breeding rodent which exhibits a unique combination of seasonal breeding and a reproductive division of labor. Pituitary function was examined by measuring the luteinizing hormone (LH) responses to single doses of 2 microg exogenous gonadotrophin-releasing hormone (GnRH) and physiological saline in 69 males and 58 females from 35 wild caught colonies. Neither males nor females exhibited any apparent manifestation of season on basal LH concentrations or on pituitary sensitivity to stimulation by exogenous GnRH. The continuance of reproductive function during the nonbreeding period is essential in common mole-rat males and females, as this period coincides with the period of maximal dispersal opportunity in the winter rainfall area they inhabit. Normal circulating levels of reproductive hormones in dispersing animals may aid intersexual recognition, assist pairbond formation, and thus prime animals for independent reproduction. Circulating basal concentrations of LH as well as LH levels measured in response to a single exogenous GnRH challenge were not significantly different between the reproductive and non-reproductive groups of either sex, suggest the absence of a physiologically well-defined suppression of reproduction in subordinate common mole-rats.     

Betacellulin overexpression in the mouse ovary leads to MAPK3/MAPK1 hyperactivation and reduces litter size by impairing fertilization

The epidermal growth factor receptor (EGFR) and its ligands are emerging as key molecules in regulating female reproduction. Here, we used a transgenic mouse model to evaluate whether and at which level of the reproduction cascade higher-than-normal levels of the EGFR ligand betacellulin (BTC) in the reproductive organs affect fertility. Western blots and immunohistochemistry revealed increased BTC levels in uterus and ovaries from transgenic females, particularly evident in granulosa cells of antral follicles. Onset of puberty, estrous cyclicity, and the anatomy and histology of reproductive organs at puberty were not altered as compared to control females. Fertility tests revealed a reduction (~50%) in litter size as the major reproductive deficit of transgenic females. Embryo implantation was delayed in transgenic females, but this was not the reason for the reduced litter size. Transgenic females produced a normal number of oocytes after natural ovulation. The in vivo fertilization rate was significantly reduced in untreated transgenic females but returned to normal levels after superovulation. Impaired oocyte fertilization in the absence of superovulation treatment was associated with MAPK3/MAPK1 hyperactivation in BTC transgenic ovaries, whereas similar levels of MAPK3/MAPK1 activation were detected in transgenic and control ovaries after superovulation treatment. Thus, tight regulation of MAPK3/MAPK1 activity appears to be essential for appropriate granulosa cell function during oocyte maturation. Our study identified hitherto unknown effects of BTC overabundance in reproduction and suggests BTC as a novel candidate protein for the modulation of fertility.     

Subsociality and female reproductive success in a mycophagous thrips: An observational and experimental analysis

Oviparous females of the haplodiploid, facultatively viviparous thrips Elaphrothrips tuberculatus(Thysanoptera: Phlaeothripidae) guard their eggs against female conspecifics and other egg predators. The intensity of maternal defense increases with clutch size. Field and laboratory observations indicate that cannibalism by females is an important selective pressure favoring maternal care. Experimental removals of guarding females showed that egg guarding substantially increases egg survivorship and that the survivorship of undefended eggs is higher in the absence of nonguarding female conspecifics than in their presence. The fecundity of viviparous females increases with the number of eggs cannibalized. The reproductive success of oviparous females increases with body size and local food density and decreases with local density of breeding females. Social behavior may not have advanced beyond maternal care in Elaphrothrips tuberculatusbecause, relative to Hymenoptera, capabilities for helping relatives are few or nonexistent, and the causes of variation in female reproductive success are not influenced easily by cooperation among females.     

Determinants of reproductive success in female Columbian ground squirrels

Summary We studied the reproductive success of female Columbian ground squirrels (Spermophilus columbianus) in southwestern Alberta for nine years. We defined reproductive success as the number of offspring surviving their first hibernation, classified as yearlings. The number of weaned juveniles explained one third of the variance in number of yearlings at emergence from their first hibernation the following spring, and much of the variance in individual reproductive success originated after weaning. Weight of adult females at emergence from hibernation was correlated with annual reproductive success. The mother's survival beyond weaning and the subsequent winter's snow accumulation had positive effects on annual reproductive success, whereas population density and summer temperature had negative effects. We found no effects on annual reproductive success of date of litter emergence, weight at emergence as a yearling, presence or absence of adult kin, distance from the natal site, location within the study area, winter temperature or summer precipitation. Age of first breeding did not affect lifetime reproductive success, which ranged from 0 to 19 yearlings produced over a lifetime. The greatest source of variation in lifetime reproductive success for females surviving to breeding age was offspring survival, followed by reproductive lifespan.     

Mating modifies female life history in a haplodiploid spider mite

Mating usually modifies females' resource allocation pattern, often as a result of conflicts between male and female partners. Can such a switch occur even in the absence of sexual conflicts? We addressed this issue in the haplodiploid spider mite Tetranychus urticae, whose biology and population structure considerably reduce conflicts between males and females over reproductive decisions. Comparing virgin and mated females, we tested the hypothesis that mated females modify their allocation pattern so as to maximize their probability of producing daughters. Mated females produced fewer but larger eggs, resulting in an overall similar reproductive effort but an increased probability of producing daughters, since in this species larger eggs are more likely to be fertilized and thus to become female. Moreover, mated females concentrated their reproduction early in life. Again, this might be a way to produce more daughters, since sperm is more abundant early in life. For virgins, spreading reproductive investment might be a way to save resources to extend life span, thus increasing their probability of encountering a sexual partner. Females with multiple opportunities for mating produced fewer eggs and a less female-biased sex ratio than once-mated females, raising the question of why multiple mating often occurs in this species.     

Impacts of male and food density on female performance in the brackish cladoceran Daphniopsis australis

The roles of male and food density in regulating female performance were investigated in the brackish cladoceran, Daphniopsis australis. Parthenogenetic females and ephippial females were tested using a 2 × 4 factorial experiment involving the presence and the absence of a male cross-classified with nil, low, medium and high food densities. For parthenogenetic females, the male presence and food density failed to trigger the switch from asexual to sexual reproduction, but the presence of male negatively affected parthenogenesis through egg abortion. Food density affected the animal longevity but depended on the male presence. The reproductive output was favoured by increasing food densities, but the male presence increased egg abortion, suggesting male being an added stress factor to parthenogenetic females. For ephippial females, food densities affected the frequency of switch from sexual to asexual modes in the absence and the presence of a male. However, the male enhanced switch frequency under low and high food densities. Longevity was increased with the male presence but was unaffected by food density. The ephippial females successfully produced diapausing eggs with the male presence. Although, ephippial females could switch to parthenogenesis but the reproductive output of switched ephippial females was inferior to that of parthenogenetic females since birth. The results reveal that the male presence and food density can impact the performance of female D. australis. Hence, this study provides an insight into the understanding of the reproductive biology of cladocerans and a possible alternative explanation for population dynamic of this species and other cladocerans in the field.     

Aggregation of whelks, Buccinum undatum, near feeding predators: the role of reproductive requirements

In the Mingan Islands, northern Gulf of St Lawrence (eastern Canada), the whelk Buccinum undatum displays a strong escape response to its predator, the asteroid Leptasterias polaris, nevertheless large sexually mature individuals occasionally approach feeding L. polaris to obtain food. In this study, we investigated the hypothesis that reproductive requirements increase the tendency of sexually mature whelks to approach feeding asteroids. Prior to egg laying, females (which invest more energy than males into the production of reproductive structures) represented 72% of the adult whelks that approached feeding L. polaris, but only 36% of the adults randomly collected from the study area. Furthermore, females that were attracted to feeding asteroids had smaller reproductive organs (after accounting for body size) than females randomly collected from the study area. Similarly, prior to egg laying, females fed longer and ingested more food than males when tested in the presence of L. polaris in the laboratory. After egg laying, however, females and males displayed a similar tendency to feed in the presence of a predator, both in the field and in the laboratory. Predator-impact indices, computed by contrasting the feeding activity of whelks in the absence and presence of a predator, indicated that females (but not males) responded more boldly to predators prior to than after egg laying, despite a general decrease in feeding activity at that period. Taken together, our observations indicate that the tendency of adult whelks to approach feeding predators is influenced by potential reproductive gains. Because such gains are presumably more directly linked to a given feeding opportunity in sexually active individuals, whelks may be selected to display increased levels of boldness towards predators with the onset of sexual maturity. Thus, potential reproductive benefits may partly explain the size-dependent tendency of whelks to approach feeding asteroids. Copyright 2001 The Association for the Study of Animal Behaviour.