Alternative life histories in the Atlantic salmon: genetic covariances within the sneaker sexual tactic in males

Alternative reproductive tactics are ubiquitous in many species. Tactic expression often depends on whether an individual's condition surpasses thresholds that are responsible for activating particular developmental pathways. Two central goals in understanding the evolution of reproductive tactics are quantifying the extent to which thresholds are explained by additive genetic effects, and describing their covariation with condition-related traits. We monitored the development of early sexual maturation that leads to the sneaker reproductive tactic in Atlantic salmon (Salmo salar L.). We found evidence for additive genetic variance in the timing of sexual maturity (which is a measure of the surpassing of threshold values) and body-size traits. This suggests that selection can affect the patterns of sexual development by changing the timing of this event and/or body size. Significant levels of covariation between these traits also occurred, implying a potential for correlated responses to selection. Closer examination of genetic covariances suggests that the detected genetic variation is distributed along at least five directions of phenotypic variation. Our results show that the potential for evolution of the life-history traits constituting this reproductive phenotype is greatly influenced by their patterns of genetic covariance.     

Loss of Wnt5a disrupts primordial germ cell migration and male sexual development in mice

Disruptions in the regulatory pathways controlling sex determination and differentiation can cause disorders of sex development, often compromising reproductive function. Although extensive efforts have been channeled into elucidating the regulatory mechanisms controlling the many aspects of sexual differentiation, the majority of disorders of sex development phenotypes are still unexplained at the molecular level. In this study, we have analyzed the potential involvement of Wnt5a in sexual development and show in mice that Wnt5a is male-specifically upregulated within testicular interstitial cells at the onset of gonad differentiation. Homozygous deletion of Wnt5a affected sexual development in male mice, causing testicular hypoplasia and bilateral cryptorchidism despite the Leydig cells producing factors such as Hsd3b1 and Insl3. Additionally, Wnt5a-null embryos of both sexes showed a significant reduction in gonadal germ cell numbers, which was caused by aberrant primordial germ cell migration along the hindgut endoderm prior to gonadal colonization. Our results indicate multiple roles for Wnt5a during mammalian reproductive development and help to clarify further the etiology of Robinow syndrome (OMIM 268310), a disease previously linked to the WNT5A pathway.     

Short-day increases in aggression are independent of circulating gonadal steroids in female Siberian hamsters (Phodopus sungorus)

Among the suite of adaptations displayed by seasonally-breeding rodents, individuals of most species display reproductive regression and concomitant decreases in gonadal steroids during the winter. In addition, some species display increased aggression in short "winter-like" days compared with long "summer-like" day lengths. For example, male Syrian and Siberian hamsters held in short days express heightened levels of aggression that are independent of gonadal steroids. Virtually nothing is known, however, regarding seasonal aggression in female Siberian hamsters (Phodopus sungorus). Studies were undertaken to determine female levels of aggression in long and short days as well as the role of gonadal steroids in mediating this behavior. In Experiment 1, females were housed in long or short days for 10 weeks and resident-intruder aggression was assessed. Prior to testing, estrous cycle stages were determined by vaginal cytology and females were tested during both Diestrus I and Proestrus. In Experiment 2, hormone levels were experimentally manipulated; long-day females were ovariectomized (OVx) or given sham surgeries whereas short-day females were implanted with capsules containing 17beta-estradiol (E(2)) or Progesterone (P). In Experiment 3, both long- and short-day females were ovariectomized and implanted with either an exogenous E(2) or blank capsule, or given a sham surgery. Short-day hamsters displayed increased aggression relative to long-day females. Aggression was not affected by estrous stage. There was no difference in aggression between long-day OVx and sham animals. Furthermore, neither exogenous E(2) nor P had any significant effect on aggression. These results support previous findings of increased non-breeding aggression and suggest that short-day aggression is not likely mediated by circulating levels of gonadal steroids. These results also suggest that the endocrine regulation of seasonal aggression may be similar between the sexes.     

植物有性生殖对温度胁迫反应的研究进展

开花植物的有性生殖阶段对温度胁迫高度敏感,高温热害和低温冷害都会对这一过程造成严重影响。本文全面总结了温度胁迫对作物有性生殖的影响,明确花粉发育过程是有性生殖过程中对温度胁迫最敏感的时期;转录组和蛋白质组的研究结果表明,蛋白激酶、热激转录因子、热休克蛋白等可能参与花粉发育期对热胁迫的信号转导。理解植物在有性生殖发育阶段如何适应温度胁迫的机理,为遗传育种实践中筛选对温度耐受的作物品种提供指导,也为基因工程选育对温度耐受的品种提供可能。     

Maturation of social reward in adult male Syrian hamsters does not depend on organizational effects of pubertal testosterone

The rewarding value of female sexual stimuli develops across puberty, as sexually-naïve adult, but not prepubertal, male hamsters show a conditioned place preference (CPP) for both vaginal secretions and a receptive female. Similarly, only adults show an endogenous testosterone surge when they encounter vaginal secretions. Testosterone by itself can condition a place preference in male rodents. Therefore, Experiment 1 assessed whether the endogenous testosterone surge elicited by vaginal secretions is necessary to show a CPP. Both gonad-intact and gonadectomized, testosterone-treated adult males showed a CPP for vaginal secretions, indicating that the rewarding value of this social cue is independent of an endogenous testosterone surge. However, organizational effects of pubertal testosterone could be necessary for adolescent development of social reward, as pubertal testosterone organizes adult-typical expression of sexual behavior. To investigate this possibility, in Experiment 2, sexually-naïve prepubertal and adult male hamsters were gonadectomized and received testosterone-filled capsules four weeks later. Testing began after two weeks of testosterone replacement. Adult males showed a CPP for both vaginal secretions and a receptive female, whether or not they experienced pubertal testosterone. Thus, the acquisition of positive valence of sexual stimuli is not organized by pubertal testosterone. Taken together, the ability of female sexual stimuli to serve as an unconditioned reward to adult male hamsters is independent of the chemosensory-induced endogenous testosterone surge and also organizational effects of pubertal testosterone. Instead, sexual reward may be dependent either on activational effects of testosterone or gonadal hormone-independent mechanisms.     

Old males reduce melanin‐pigmented traits and increase reproductive outcome under worse environmental conditions in common kestrels

Secondary sexual traits displayed by males and females may have evolved as a signal of individual quality. However, both individual quality and investment on producing or maintaining enhanced sexual traits change as individuals age. At the same time, the costs associated to produce sexual traits might be attenuated or increased if environmental conditions are benign or worse respectively. Accordingly, environmental conditions are expected to shape the association between the expression of sexual traits and their reproductive outcome as individuals age. Nonetheless, little is known about the environmental influence on the co‐variation between sexual traits and reproductive outcome throughout the life of individuals. We studied the age‐dependency of the number and size of back spots, a melanin‐based and sexual trait in adults of common kestrels (Falco tinnunculus). We analysed the age‐dependence of reproductive traits and the environmental influence, defined as vole abundance, using a 10‐year individual‐based dataset. We broke down age‐related changes of reproductive traits into within‐ and between‐individual variation to assess their contribution to population‐level patterns. Our results showed a within‐individual decrease in the number, but not the size, of back spots in males. The size of back spots was positively correlated with food availability in males. Reproductive performance of males increased as they aged, in agreement with the life‐history theory but depending of vole abundance. Remarkably, we found that having fewer back spots was positively associated with clutch size only for old individuals under low‐food conditions. We suggest that environmental variation may shape the association between the expression of a sexual signal and reproductive outcome. We speculate that the reliability of sexual traits is higher when environmental conditions are poor only for old individuals. Within an evolutionary context, we suggest that the expression of sexual traits might be constrained by environmental conditions at later stages of life.     

Lack of Support for the Association between Facial Shape and Aggression: A Reappraisal Based on a Worldwide Population Genetics Perspective

Antisocial and criminal behaviors are multifactorial traits whose interpretation relies on multiple disciplines. Since these interpretations may have social, moral and legal implications, a constant review of the evidence is necessary before any scientific claim is considered as truth. A recent study proposed that men with wider faces relative to facial height (fWHR) are more likely to develop unethical behaviour mediated by a psychological sense of power. This research was based on reports suggesting that sexual dimorphism and selection would be responsible for a correlation between fWHR and aggression. Here we show that 4,960 individuals from 94 modern human populations belonging to a vast array of genetic and cultural contexts do not display significant amounts of fWHR sexual dimorphism. Further analyses using populations with associated ethnographical records as well as samples of male prisoners of the Mexico City Federal Penitentiary condemned by crimes of variable level of inter-personal aggression (homicide, robbery, and minor faults) did not show significant evidence, suggesting that populations/individuals with higher levels of bellicosity, aggressive behaviour, or power-mediated behaviour display greater fWHR. Finally, a regression analysis of fWHR on individual''s fitness showed no significant correlation between this facial trait and reproductive success. Overall, our results suggest that facial attributes are poor predictors of aggressive behaviour, or at least, that sexual selection was weak enough to leave a signal on patterns of between- and within-sex and population facial variation.     

Plasticity in reproductive phenotypes reveals status-specific correlations between behavioral, morphological, and physiological sexual traits

Reproductive success is determined by a complex interplay between multiple sexual traits that promote mate acquisition and, following copulation, provide control over paternity. The intensity of sexual competition that individuals experience often fluctuates, and here we investigate how this influences the expression of reproductive traits and their relationships. We show in the fowl, Gallus gallus, that males of different social status, which experience different intensities of sexual competition, before and after copulation, have different reproductive phenotypes. Dominant males are more vigilant, feed less, and have larger sexual ornaments than subordinate males. Experimentally manipulating social status revealed that these differences were phenotypically plastic, indicating multiple sexual traits were dependent on the social environment. We integrated these data with previous published findings on changes in sperm numbers and velocity to show that relationships between traits were different for males when they were dominant and when they were subordinate. Furthermore, when males switched status a complex array of negative and positive correlations between the degree traits changed was observed. Our results suggest that variation in the intensity of sexual competition generates reversible plasticity in reproductive phenotypes and that relationships between sexual traits may be variable and influence the evolution of reproductive strategies.     

Reproduction and sexual dimorphism in two populations of the polymorphic spiny lizard Sceloporus minor from Hidalgo,México

Male‐biased sexual dimorphism is extensive in New World spiny lizards (Sceloporus: Phrynosomatidae) and is particularly prominent in the polymorphic minor lizard Sceloporus minor. However, the possible relationship between patterns of sexual dimorphism and gonadal development is little known for most species. In this study, we explored aspects of sexual dimorphism in noncolor morphological traits in S. minor and characterized the gonadal cycle in males and females from each of two sites (El Enzuelado and La Manzana) in Hidalgo, México, differing in key ecological aspects linked to life‐history trait expression in other lizards. Males were generally larger than females in each population and expressed larger forms of several other morphological traits, although not all comparisons attained statistical significance. Both sexes attained reproductive maturity at a larger size in El Enzuelado relative to La Manzana, and females from El Enzuelado had larger litters. Gonadal cycles differed substantially between the two populations and suggest that reproductive activity of males and females was synchronous at El Enzuelado and asynchronous at La Manzana, an unusual pattern. Geographic variation in sex‐specific responses to environmental variables may be at least partly responsible for the exceptional species diversity of spiny lizards in México.     

Gonadal hormones during puberty organize environment-related social interaction in the male rat

This study examined the role of gonadal androgens during puberty on the development of environment-related social interaction (SI) in male rats. SI in an unfamiliar environment versus SI in a familiar environment was evaluated in young adult rats as a function of sex and gonadal status. Intact male rats at 60 days of age exhibited a differential response to the two environments, whereas SI in intact female rats at 60 days was equivalent in the two environments. Furthermore, male rats castrated as juveniles and tested for SI at 60 days displayed a pattern of environment-related SI similar to SI in intact adult female rats. This effect of juvenile castration on SI in male rats was prevented by chronic exposure to testosterone propionate (TP) over Days 30 through 60. SI in male rats castrated in adulthood, on the other hand, was not altered either 2 or 4 weeks postcastration. The results from this study indicate that pubertal secretions of gonadal androgen(s) are necessary for the development of environment-related SI in male rats. In contrast, secretions of gonadal androgens in adulthood do not appear to be critical for the continued expression of environment-related SI, as suggested by the observation that environment-related SI in male rats remains unchanged by castration in adulthood.     

Evidence that female preferences have shaped male signal evolution in a clade of specialized plant-feeding insects

Mate choice is considered an important influence in the evolution of mating signals and other sexual traits, and--since divergence in sexual traits causes reproductive isolation--it can be an agent of population divergence. The importance of mate choice in signal evolution can be evaluated by comparing male signal traits with female preference functions, taking into account the shape and strength of preferences. Specifically, when preferences are closed (favouring intermediate values), there should be a correlation between the preferred values and the trait means, and stronger preferences should be associated with greater preference-signal correspondence and lower signal variability. When preferences are open (favouring extreme values), signal traits are not only expected to be more variable, but should also be shifted towards the preferred values. We tested the role of female preferences in signal evolution in the Enchenopa binotata species complex of treehoppers, a clade of plant-feeding insects hypothesized to have speciated in sympatry. We found the expected relationship between signals and preferences, implicating mate choice as an agent of signal evolution. Because differences in sexual communication systems lead to reproductive isolation, the factors that promote divergence in female preferences--and, consequently, in male signals--may have an important role in the process of speciation.     

Extreme aggression in male squid induced by a β-MSP-like pheromone

Male-male aggression is widespread in the animal kingdom and subserves many functions related to the acquisition or retention of resources such as shelter, food, and mates. These functions have been studied widely in the context of sexual selection, yet the proximate mechanisms that trigger or strengthen aggression are not well known for many taxa. Various external sensory cues (visual, audio, chemical) acting alone or in combination stimulate the complex behavioral interactions of fighting behaviors. Here we report the discovery of a 10 kDa protein, termed Loligo β-microseminoprotein (Loligo β-MSP), that immediately and dramatically changes the behavior of male squid from calm swimming and schooling to extreme fighting, even in the absence of females. Females synthesize Loligo β-MSP in their reproductive exocrine glands and embed the protein in the outer tunic of egg capsules, which are deposited on the open sea floor. Males are attracted to the eggs visually, but upon touching them and contacting Loligo β-MSP, they immediately escalate into intense physical fighting with any nearby males. Loligo β-MSP is a distant member of the chordate β-microseminoprotein family found in mammalian reproductive secretions, suggesting that this gene family may have taxonomically widespread roles in sexual competition.     

Adaptive advantages of patterns of growth and asexual reproduction of the sea anemone Metridium senile (L.) in intertidal and submerged populations

Individual size, rate of growth, and mode and frequency of asexual reproduction are life-history traits of primary importance for sea anemones. These traits determine sexual reproductive output, affect an individual's probability of survival, and are crucial in adapting an individual to its environmental surroundings. The sea anemone Metridium senile (L.) is highly variable in ecological distribution and life history, including rate of growth, individual size, and rate of asexual reproduction. Gonad size (measured as cross-sectional area of gonadal tissue) increases with body weight, so individuals should grow as large and as rapidly as possible to maximize individual sexual reproductive output. Cessation of growth and small body size in intertidal populations suggest that growth is constrained by genetic or environmental conditions. The growth of intertidal individuals transplanted to harbor-float panels demonstrated that growth limits are imposed by environmental factors, most probably limited food and feeding time and damage from wave exposure (which stimulates fragmentation). Individuals in harbor-float populations, which are continuously immersed, grow much larger, and large individuals comprise a greater proportion of the population than in the intertidal zone. The highest rate of fragmentation observed was on harbor-float panels. Patterns of growth and asexual reproduction provide adaptive advantages for M. senile. For harborfloat individuals, large individual size increases gamete production and may increase feeding efficiency. For intertidal individuals, asexual reproduction allows growth despite individual size constraints and rapid population growth, with specific advantages resulting from clone formation.     

Repair of reproductive deficits by neural transplantation

Transplantation of brain tissue has been used to ameliorate the genetic lesion of the hypogonadal mutant mouse. This animal does not synthesize gonadotropin-releasing hormone (GnRH) and so has an infantile reproductive system. Implantation of normal fetal or neonatal preoptic area containing GnRH neurons reverses many aspects of the reproductive deficiency. Pituitary and plasma levels of gonadotropins rise, followed by growth of the gonads and sexual organs. Pituitary release of gonadotropins is episodic, suggesting that the grafted tissue is integrated into the "pulse generator." The vast majority of grafted animals do not show castration-induced elevations of luteinizing hormone (LH) nor respond to exogenous steroids with a depression in circulating LH. Negative feedback of gonadal steroids seems to be inoperative. In contrast, some females can show ovulatory surges of LH in response to mating (reflex ovulation), after administration of exogenous steroid (progesterone), and, on rare occasion, ovulation cycles occur spontaneously. Anatomical studies demonstrate that reproductive recovery is dependent on the outgrowth of GnRH axons to the host median eminence. Some but not all of the GnRH neurons within the grafts contribute to this innervation. GnRH axons exit into the host along well-defined pathways, recapitulating in part the paths taken by normal axons. How the graft and host are integrated to produce the panoply of reproductive responses is the subject of current study.     

Nutrient restriction induces failure of reproductive function and molecular changes in hypothalamus–pituitary–gonadal axis in postpubertal gilts

People on a diet to lose weight may be at risk of reproductive failure. To investigate the effects of nutrient restriction on reproductive function and the underlying mechanism, changes of reproductive traits, hormone secretions and gene expressions in hypothalamus–pituitary–gonadal axis were examined in postpubertal gilts at anestrus induced by nutrient restriction. Gilts having experienced two estrus cycles were fed a normal (CON, 2.86 kg/d) or nutrient restricted (NR, 1 kg/d) food regimens to expect anestrus. NR gilts experienced another three estrus cycles, but did not express estrus symptoms at the anticipated fourth estrus. Blood samples were collected at 5 days’ interval for consecutive three times for measurement of hormone concentrations at the 23th day of the fourth estrus cycle. Individual progesterone concentrations of NR gilts from three consecutive blood samples were below 1.0 ng/mL versus 2.0 ng/mL in CON gilts, which was considered anestrus. NR gilts had impaired development of reproductive tract characterized by absence of large follicles (diameter ≥ 6 mm), decreased number of corepus lutea and atrophy of uterus and ovary tissues. Circulating concentrations of IGF-I, kisspeptin, estradiol, progesterone and leptin were significantly lower in NR gilts than that in CON gilts. Nutrient restriction down-regulated gene expressions of kiss-1, G-protein coupled protein 54, gonadotropin-releasing hormone, estrogen receptor α, progesterone receptor, leptin receptor, follicle-stimulating hormone and luteinizing hormone and insulin-like growth factor I in hypothalamus–pituitary–gonadal axis of gilts. Collectively, nutrient restriction resulted in impairment of reproductive function and changes of hormone secretions and gene expressions in hypothalamus–pituitary–gonadal axis, which shed light on the underlying mechanism by which nutrient restriction influenced reproductive function.     

Sexual maturation and related changes in aspartate transcarbamylase activity of gonad tissues in the soft shell clam (Mya arenaria)

The information on the relation between the gonadal aspartate transcarbamylase (ACTase) activity and the sexual maturation in molluscs is very fragmentary and is still absent in Mya arenaria. The determination of ATCase activity, energy reserves levels and maturation stages were done in gonads of clams. Results showed that the seasonal cycle of storage and utilization of energy reserves in gonads of clams are linked to the bimodal reproduction well known in this bivalve. ATCase activity was high in clams at development and ripe stages, while this activity was low for individuals found in indifferent, spawning or spent stages. This difference can be explained by the fact that during gonad development, gonadal synthetic activity increased following the mitotic events associated to the reproductive cycle. The results presented in this paper have confirmed the link between ATCase activity and gametogenesis in M. arenaria. Further work should be realized in order to assess if ATCase activity could be considered as a potential biomarker to evaluate the disruption of sexual maturation in clams collected in sites such as the St. Lawrence estuary.     

Relationships among male sexually selected traits in the bean bug,Riptortus pedestris (Heteroptera: Alydidae)

Current concepts of sexual selection suggest that male reproductive success is determined by multiple sexual traits. As expression and production of multiple sexual traits are frequently associated with each other, positive or negative correlations among multiple sexual traits ensue. These relationships among traits associated with male reproductive success may be crucial in the evolution of male reproductive strategies. Here, we investigate phenotypic relationships among sexually selected traits in the armed bean bug Riptortus pedestris. In this insect, males with a larger body and weapon are more likely to win male–male competitions, and males with a larger weapon or higher courtship rate are more attractive to females. There was a significant positive correlation between body size and weapon size, whereas the courtship rate had significant negative correlations with body size and weapon size. Our results suggested that there was a phenotypic trade‐off between courtship rate and male morphology. In this insect, smaller males may make more effort in courtship behavior as an alternative mating tactic.     

Sexual isolation promotes divergence between parapatric lake and stream stickleback

Speciation can be initiated by adaptive divergence between populations in ecologically different habitats, but how sexually based reproductive barriers contribute to this process is less well understood. We here test for sexual isolation between ecotypes of threespine stickleback fish residing in adjacent lake and stream habitats in the Lake Constance basin, Central Europe. Mating trials exposing females to pairings of territorial lake and stream males in outdoor mesocosms allowing for natural reproductive behaviour reveal that mating occurs preferentially between partners of the same ecotype. Compared to random mating, this sexual barrier reduces gene flow between the ecotypes by some 36%. This relatively modest strength of sexual isolation is surprising because comparing the males between the two ecotypes shows striking differentiation in traits generally considered relevant to reproductive behaviour (body size, breeding coloration, nest size). Analysing size differences among the individuals in the mating trials further indicates that assortative mating is not related to ecotype differences in body size. Overall, we demonstrate that sexually based reproductive isolation promotes divergence in lake–stream stickleback along with other known reproductive barriers, but we also caution against inferring strong sexual isolation from the observation of strong population divergence in sexually relevant traits.     

Gonochorism and seasonal variations in the gonads of the labrid Symphodus (Crenilabrus) ocellatus (Forsskål)

A histological study of gonadal organization of 354 Symphodus ocellatus caught in the Bay of Naples from June 1985 to December 1986 demonstrated the absence of sexual inversion in this species. Four phases of gonadal development, related to the seasonal cycle, are described. During the reproductive period the ovaries showed the asynchrony in oocyte development that is typical of serial spawners. The testes of three types of males present in this species (territorial, satellite and sneaker males) were examined: the only difference found was in the gonadal growth pattern during the reproductive period.     

Environmental heterogeneity,genotype-by-environment interactions and the reliability of sexual traits as indicators of mate quality

Exaggerated sexual displays are often supposed to indicate the indirect benefits females may receive from sexual reproduction with displaying males, but empirical evidence for positive relationships between the genetic quality and sexual trait quality is scant. The explanation for this might lie in the fact that mixing of reproductive individuals whose development has been influenced by genotype-by-environment interactions (GEIs) can blur the relationship between the individual male genetic quality and phenotype as perceived by females. Strong GEIs can generate an ecological crossover, where different genotypes are superior in environments that are separated either in space or time. Here, we use a stochastic simulation model to show that even a weak GEI, which does not generate an obvious ecological crossover, can neutralize or even reverse the relationship between genetic quality and sexual trait size in the presence of environmental heterogeneity during development. Our model highlights the importance of developmental selection in evolution of traits and allows us to predict the situations in which sexual displays might not be reliable indicators of genetic quality.