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1.
Disentangling the relationship between age and reproduction is central to understand life‐history evolution, and recent evidence shows that considering condition‐dependent mortality is a crucial piece of this puzzle. For example, nonrandom mortality of ‘low‐condition’ individuals can lead to an increase in average lifespan. However, selective disappearance of such low‐condition individuals may also affect reproductive senescence at the population level due to trade‐offs between physiological functions related to survival/lifespan and the maintenance of reproductive functions. Here, we address the idea that condition‐dependent extrinsic mortality (i.e. simulated predation) may increase the age‐related decline in male reproductive success and with it the potential for sexual conflict, by comparing reproductive ageing in Drosophila melanogaster male/female cohorts exposed (or not) to condition‐dependent simulated predation across time. Although female reproductive senescence was not affected by predation, male reproductive senescence was considerably higher under predation, due mainly to an accelerated decline in offspring viability of ‘surviving’ males with age. This sex‐specific effect suggests that condition‐dependent extrinsic mortality can exacerbate survival‐reproduction trade‐offs in males, which are typically under stronger condition‐dependent selection than females. Interestingly, condition‐dependent extrinsic mortality did not affect mating success, hinting that accelerated reproductive senescence is due to a decrease in male post‐copulatory fitness components. Our results support the recent proposal that male ageing can be an important source of sexual conflict, further suggesting this effect could be exacerbated under more natural conditions.  相似文献   

2.
The trade‐off between the allocation of resources toward somatic maintenance or reproduction is one of the fundamentals of life history theory and predicts that females invest in offspring at the expense of their longevity or vice versa. Mate quality may also affect life history trade‐offs through mechanisms of sexual conflict; however, few studies have examined the interaction between mate quality and age at first mating in reproductive decisions. Using house crickets (Acheta domesticus), this study examines how survival and reproductive trade‐offs change based on females’ age at first reproduction and exposure to males of varying size. Females were exposed to either a large (presumably high‐quality) or small male at an early (young), middle (intermediate), or advanced (old) age, and longevity and reproductive investment were subsequently tracked. Females mated at a young age had the largest number of eggs but the shortest total lifespans while females mated at older ages produced fewer eggs but had longer total lifespans. The trade‐off between age at first mating and eggs laid appears to be mediated through higher egg‐laying rates and shorter postmating lifespans in females mated later in life. Exposure to small males resulted in shorter lifespans and higher egg‐laying rates for all females indicating that male manipulation of females, presumably through spermatophore contents, varies with male size in this species. Together, these data strongly support a trade‐off between age at first reproduction and lifespan and support the role of sexual conflict in shaping patterns of reproduction.  相似文献   

3.
Although the reasons why organisms age and die are generally well understood, it has recently been suggested that an optimal life span has evolved not only as the result of trade‐offs between reproductive performances early and late in life, but also that a balance between the costs and benefits of the number of mating has also played an important role in the evolution of ageing in both sexes. By using four seed beetle (Acanthoscelides obtectus) lines selected for different life history traits, but which have also inadvertently created monoandrous and polyandrous conditions, we showed that males evolved to affect the mortality patterns of females in a way consistent to the postmating sexual selection generated by sexually antagonistic co‐evolution theory. Monoandrous males, irrespectively of body weight and other life history traits specific to their lines, evolved to increase the longevity of control females kept under starvation and suppressed fecundity, compared with males that originated in the lines with effectively polyandrous conditions. When females were allowed to lay eggs, the effects of males from different lines and mating type history on the senescence of females were substantially weaker. We found that males in the line that was evolved to decelerate senescence and polyandrous conditions stimulate the earlier onset of females’ oviposition, relative to males stemmed from the line with accelerated senescence and monoandrous conditions. This fact may explain the absence of difference in the mean longevities between the control females mated to these males and highlight the importance of sexual selection in the evolution of ageing.  相似文献   

4.
Understanding trade‐offs in wild populations is difficult, but important if we are to understand the evolution of life histories and the impact of ecological variables upon them. Markers that reflect physiological state and predict future survival would be of considerable benefit to unraveling such trade‐offs and could provide insight into individual variation in senescence. However, currently used markers often yield inconsistent results. One underutilized measure is hematocrit, the proportion of blood comprising erythrocytes, which relates to the blood's oxygen‐carrying capacity and viscosity, and to individual endurance. Hematocrit has been shown to decline with age in cross‐sectional studies (which may be confounded by selective appearance/disappearance). However, few studies have tested whether hematocrit declines within individuals or whether low hematocrit impacts survival in wild taxa. Using longitudinal data from the Seychelles warbler (Acrocephalus sechellensis), we demonstrated that hematocrit increases with age in young individuals (<1.5 years) but decreases with age in older individuals (1.5–13 years). In breeders, hematocrit was higher in males than females and varied relative to breeding stage. High hematocrit was associated with lower survival in young individuals, but not older individuals. Thus, while we did not find support for hematocrit as a marker of senescence, high hematocrit is indicative of poor condition in younger individuals. Possible explanations are that these individuals were experiencing dehydration and/or high endurance demands prior to capture, which warrants further investigation. Our study demonstrates that hematocrit can be an informative metric for life‐history studies investigating trade‐offs between survival, longevity, and reproduction.  相似文献   

5.
Allocation decisions depend on an organism's condition which can change with age. Two opposite changes in life‐history traits are predicted in the presence of senescence: either an increase in breeding performance in late age associated with terminal investment or a decrease due to either life‐history trade‐offs between current breeding and future survival or decreased efficiency at old age. Age variation in several life‐history traits has been detected in a number of species, and demographic performances of individuals in a given year are influenced by their reproductive state the previous year. Few studies have, however, examined state‐dependent variation in life‐history traits with aging, and they focused mainly on a dichotomy of successful versus failed breeding and non‐breeding birds. Using a 50‐year dataset on the long‐lived quasi‐biennial breeding wandering albatross, we investigated variations in life‐history traits with aging according to a gradient of states corresponding to potential costs of reproduction the previous year (in ascending order): non‐breeding birds staying at sea or present at breeding grounds, breeding birds that failed early, late or were successful. We used multistate models to study survival and decompose reproduction into four components (probabilities of return, breeding, hatching, and fledging), while accounting for imperfect detection. Our results suggest the possible existence of two strategies in the population: strict biennial breeders that exhibited almost no reproductive senescence and quasi‐biennial breeders that showed an increased breeding frequency with a strong and moderate senescence on hatching and fledging probabilities, respectively. The patterns observed on survival were contrary to our predictions, suggesting an influence of individual quality rather than trade‐offs between reproduction and survival at late ages. This work represents a step further into understanding the evolutionary ecology of senescence and its relationship with costs of reproduction at the population level. It paves the way for individual‐based studies that could show the importance of intra‐population heterogeneity in those processes.  相似文献   

6.
Telomere dynamics in natural populations have been linked to survival, reproduction, and energetic investment. Given their putative role in mediating life‐history trade‐offs, telomeres are also a likely candidate for maintaining honesty in sexually selected signals; few studies to date, however, have demonstrated a correlation between sexual signals and telomere dynamics. Here, we show that plumage coloration in male common yellowthroats (Geothlypis trichas) is correlated with both relative telomere length and with the rate of telomere loss between years. Elevated antioxidant capacity is also associated with reduced telomere loss, but only among older males. Previous work in this population has demonstrated that males with brighter plumage are in better condition, have higher reproductive success, and are more likely to survive over winter. Thus, the signal attribute associated with mate choice in this system also conveys reliable information about telomere dynamics. At present, it is unclear whether telomere maintenance plays a causal role in maintaining signal honesty or whether the correlation arises due to underlying variation in individual resources or genotypes. We suggest that subsequent work should consider the possibility that fundamental trade‐offs between signal investment and cell‐level processes that influence aging and reproductive senescence may provide a foundation for understanding the maintenance of sexual signal honesty.  相似文献   

7.
Hypotheses exploring the influence of dietary conditions on the life‐history trade‐off between survival and reproductive success are extensively tested in female insects but only rarely explored in males. The present study examines the impact of dietary quality and female access on age‐specific reproduction and survival of the male Mexican fruit fly Anastrepha ludens Loew (Diptera: Tephritidae). There is a clear cost of female access for males with access to dietary protein, measurable as a decrease in life expectancy, which is further influenced by the age when females are introduced. A protein deficient diet reduces the lifespan benefit of virginity and masks the detrimental effect of female access on male life expectancy. Dietary protein is not necessary for reproductive success, although access to protein at eclosion improves the lifetime reproductive success of males compared to when it is delayed. Overall, reproductive success diminishes as the male flies age, regardless of the dietary conditions, providing evidence for reproductive senescence in males. Delaying the males' access to a protein source fails to influence the negative effect of age on reproductive ability. Because age‐specific reproductive rates decline with age, regardless of diet, male fitness does not benefit from lifespan extension. Therefore, males can be expected to allocate available resources towards reproductive effort in favour of an extended lifespan, regardless of mate and protein availability.  相似文献   

8.
Oxidative stress (i.e., more oxidants than antioxidants) has been proposed as a proximate currency in life‐history trade‐offs, which if studied in an ecological setting allow a more realistic perspective on the origin and evolution of trade‐offs. Therefore, the aim here was to investigate the impact of ecological and individual factors for variation in markers of oxidative stress using both experimental and correlational data. Total glutathione (tGSH), oxidized glutathione (GSSG), plasma antioxidant capacity (OXY), and plasma‐reactive oxygen metabolites (ROM) were measured in more than 700 breeding great tits (Parus major). The main results revealed a pronounced sex difference, with females having lower ROM and OXY, but higher tGSH compared with males. In addition, birds breeding in the evergreen areas had higher tGSH compared with those in the deciduous habitat, but the experimentally manipulated breeding density had no significant effect on any of the redox markers. Independent of the sex differences, the larger the reproductive investment the lower the ROM of both males and females. Taken together, the extracellular markers – ROM and OXY – revealed similar results and were highly correlated. Interestingly, the direction of their effects was in the opposite direction to the endogenously synthesized tGSH and GSSG. This highlights the need to combine extracellular markers with endogenously synthesized antioxidants to understand its implications for the origin and evolution of trade‐offs in an ecological setting.  相似文献   

9.
Telomeres in human fibroblasts shorten progressively during in vitro culturing and trigger replicative senescence. Furthermore, shortened telomeres can be used as biomarkers of disease. These observations have led to the suggestion that telomere dynamics may also be associated with viability and selection for life history variation in non‐human taxa. Model systems to examine this suggestion would particularly benefit from the coexistence of multiple phenotypes within the same species with different life history trade‐offs, since those could be compared in terms of telomere characteristics. This scenario also provokes the classic question of why one morph does not have marginally higher fitness and replaces the others. One explanation is that different morphs have different reproductive tactics with equal relative fitness. In Australian painted dragons (Ctenophorus pictus), males differ in head color, the presence or absence of a gular bib, and reproductive expenditure. Red males out‐compete yellow males in dominance contests, while yellow males copulate quickly and have higher success in sperm competition than red males. Males with bibs better defend partners against rival matings, at the cost of loss of body condition. We show that yellow‐headed and bib‐less males have longer telomeres than red, blue and bibbed males, suggesting that telomere length is positively associated with higher investment into self‐maintenance and less reproductive expenditure.  相似文献   

10.
Fitness depends on both the resources that individuals acquire and the allocation of those resources to traits that influence survival and reproduction. Optimal resource allocation differs between females and males as a consequence of their fundamentally different reproductive strategies. However, because most traits have a common genetic basis between the sexes, conflicting selection between the sexes over resource allocation can constrain the evolution of optimal allocation within each sex, and generate trade‐offs for fitness between them (i.e. ‘sexual antagonism’ or ‘intralocus sexual conflict’). The theory of resource acquisition and allocation provides an influential framework for linking genetic variation in acquisition and allocation to empirical evidence of trade‐offs between distinct life‐history traits. However, these models have not considered the emergence of trade‐offs within the context of sexual dimorphism, where they are expected to be particularly common. Here, we extend acquisition–allocation theory and develop a quantitative genetic framework for predicting genetically based trade‐offs between life‐history traits within sexes and between female and male fitness. Our models demonstrate that empirically measurable evidence of sexually antagonistic fitness variation should depend upon three interacting factors that may vary between populations: (1) the genetic variances and between‐sex covariances for resource acquisition and allocation traits, (2) condition‐dependent expression of resource allocation traits and (3) sex differences in selection on the allocation of resource to different fitness components.  相似文献   

11.
A central objective of evolutionary biology is understanding variation in life‐history trajectories and the rate of aging, or senescence. Senescence can be affected by trade‐offs and behavioural strategies in adults but may also be affected by developmental stress. Developmental stress can accelerate telomere degradation, with long‐term longevity and fitness consequences. Little is known regarding whether variation in developmental stress and telomere dynamics contributes to patterns of senescence during adulthood. We investigated this question in the dimorphic white‐throated sparrow (Zonotrichia albicollis), a species in which adults of the two morphs exhibit established differences in behavioural strategy and patterns of senescence, and also evaluated the relationship between oxidative stress and telomere length. Tan morph females, which exhibit high levels of unassisted parental care, display faster reproductive senescence than white females, and faster actuarial senescence than all of the other morph–sex classes. We hypothesized that high oxidative stress and telomere attrition in tan female nestlings could contribute to this pattern, since tan females are small and potentially at a competitive disadvantage even as nestlings. Nestlings that were smaller than nest mates had higher oxidative stress, and nestlings with high oxidative stress and fast growth rates displayed shorter telomeres. However, we found no consistent morph–sex differences in oxidative stress or telomere length. Results suggest that oxidative stress and fast growth contribute to developmental telomere attrition, with potential ramifications for adults, but that developmental oxidative stress and telomere dynamics do not account for morph–sex differences in senescence during adulthood.  相似文献   

12.
The reproductive effort that a male directs to a familiar female declines over time, suggesting decreasing marginal returns. But is this diminishing returns a function of increasing reproductive costs or decreasing benefits of sustained effort? Here, we use the restoration of male reproductive effort with unfamiliar females to differentiate the role of diminishing returns and lifetime costs of increased reproductive effort of male guppies. We kept males with familiar or unfamiliar females throughout their lives and manipulated their ability to either court or mate with females. We found that increased male reproductive effort with novel mates lead to an immediate trade‐off in the form of reduced foraging effort. Further, males able to mate with a series of unfamiliar females had lower lifetime growth, indicating the primary cost of male reproductive effort in guppies arises from copulation rather than courtship. The lifetime growth trade‐offs were significant only when males mated with unfamiliar mates, suggesting that male reproductive effort with familiar females declines before it is restricted by physical exhaustion. These findings provide some of the first evidence of longitudinal costs of increased male reproductive effort in a vertebrate.  相似文献   

13.
Josh R. Auld  Anne Charmantier 《Oikos》2011,120(8):1129-1138
Reproductive senescence, an intra‐individual decline in reproductive function with age, is widespread, but proximate factors determining its rate remain largely unknown. Most studies of reproductive senescence focus on females, leaving senescence in male function and its implications for female function largely understudied. We constructed linear mixed models to explore the interactive effects of paternal and maternal age and a life‐history trait (i.e. age at first reproduction) on four fitness components (i.e. laying date, clutch size, number of fledglings and number of recruits) measured in a wild, breeding population of blue tits Cyanistes caeruleus ogliastrae where individual breeding success has been followed for over 30 years (our dataset spanned 29 years). Previous studies have shown that, across female lifespan, laying date decreases and subsequently increases; earlier laying dates result in higher fitness because hatchlings have greater access to a seasonal food source. Our analyses reveal that females that initiate reproduction early in life show a greater delay in laying date with old age. In addition to delayed laying dates, older females lay smaller clutches. However, the magnitude of female age effects was influenced by the age at first reproduction of their breeding partners. Senescence of laying date and clutch size was reduced when females mated with males that reproduced early in life compared to males that delayed reproduction. We confirmed that both laying date and clutch size were significantly correlated with reproductive fitness suggesting that these dynamics early in the breeding cycle can have long‐term consequences. These complex phenotypic interactions shed light on the proximate mechanisms underlying reproductive senescence in nature and highlight the potential importance of cross‐sex age by life‐history interactions.  相似文献   

14.
Actuarial senescence is widespread in age‐structured populations. In growing populations, the progressive decline of Hamiltonian forces of selection with age leads to decreasing survival. As actuarial senescence is overcompensated by a high fertility, actuarial senescence should be more intense in species with high reproductive effort, a theoretical prediction that has not been yet explicitly tested across species. Wild boar (Sus scrofa) females have an unusual life‐history strategy among large mammals by associating both early and high reproductive effort with potentially long lifespan. Therefore, wild boar females should show stronger actuarial senescence than similar‐sized related mammals. Moreover, being polygynous and much larger than females, males should display higher senescence rates than females. Using a long‐term monitoring (18 years) of a wild boar population, we tested these predictions. We provided clear evidence of actuarial senescence in both sexes. Wild boar females had earlier but not stronger actuarial senescence than similar‐sized ungulates. Both sexes displayed similar senescence rates. Our study indicates that the timing of senescence, not the rate, is associated with the magnitude of fertility in ungulates. This demonstrates the importance of including the timing of senescence in addition to its rate to understand variation in senescence patterns in wild populations.  相似文献   

15.
In the socially polymorphic spider Anelosimus studiosus, males mature early in the reproductive season and recruit to the webs of juvenile females and guard them until they mature. During the period before females mature, males and females engage in repeated bouts of non‐conceptive (play) sexual behavior, where the pair courts and engages in mock copulation; both males and females gain performance‐enhancing experience via these encounters. In this study, we examined the factors that underlie individual variation in the tendency to engage in non‐conceptive mating and determine whether it impacts male–male competition for females. We found that docile females, being less resistant to mating in general, are more likely to accept male courtship and non‐conceptive copulation as juveniles. Personality type influenced the exhibition of non‐conceptive sexual behavior in males as well. High body condition males of the aggressive phenotype were more likely to engage in non‐conceptive sexual behavior than males with lower body condition. Body condition did not influence docile males’ propensity to engage in non‐conceptive sexual behavior, but female size did. Docile males engaged in more non‐conceptive sexual displays with larger females. Engaging in non‐conceptive sexual displays negatively impacted male performance in staged male–male contests for access to females. This cost was greatest for males of the aggressive phenotype, which are otherwise favored in male–male contests. Our findings indicate expression of non‐conceptive sexual displays is linked to personality and results in reproductive performance trade‐offs for male A. studiosus.  相似文献   

16.
Extra‐pair paternity (EPP) is often linked to male age in socially monogamous vertebrates; that is, older males are more likely to gain EPP and less likely to be cuckolded. However, whether this occurs because males improve at gaining paternity as they grow older, or because “higher quality” males that live longer are preferred by females, has rarely been tested, despite being central to our understanding of the evolutionary drivers of female infidelity. Moreover, how extra‐pair reproduction changes with age within females has received even less attention. Using 18 years of longitudinal data from an individually marked population of Seychelles warblers (Acrocephalus sechellensis), we found considerable within‐individual changes in extra‐pair reproduction in both sexes: an early‐life increase and a late‐life decline. Furthermore, males were cuckolded less as they aged. Our results indicate that in this species age‐related patterns of extra‐pair reproduction are determined by within‐individual changes with age, rather than differences among individuals in longevity. These results challenge the hypothesis—based on longevity reflecting intrinsic quality—that the association between male age and EPP is due to females seeking high‐quality paternal genes for offspring. Importantly, EPP accounted for up to half of male reproductive success, emphasizing the male fitness benefits of this reproductive strategy. Finally, the occurrence of post‐peak declines in extra‐pair reproduction provides explicit evidence of senescence in infidelity in both males and females.  相似文献   

17.
Individual variation in resource acquisition should have consequences for life‐history traits and trade‐offs between them because such variation determines how many resources can be allocated to different life‐history functions, such as growth, survival and reproduction. Since resource acquisition can vary across an individual's life cycle, the consequences for life‐history traits and trade‐offs may depend on when during the life cycle resources are limited. We tested for differential and/or interactive effects of variation in resource acquisition in the burying beetle Nicrophorus vespilloides. We designed an experiment in which individuals acquired high or low amounts of resources across three stages of the life cycle: larval development, prior to breeding and the onset of breeding in a fully crossed design. Resource acquisition during larval development and prior to breeding affected egg size and offspring survival, respectively. Meanwhile, resource acquisition at the onset of breeding affected size and number of both eggs and offspring. In addition, there were interactive effects between resource acquisition at different stages on egg size and offspring survival. However, only when females acquired few resources at the onset of breeding was there evidence for a trade‐off between offspring size and number. Our results demonstrate that individual variation in resource acquisition during different stages of the life cycle has important consequences for life‐history traits but limited effects on trade‐offs. This suggests that in species that acquire a fixed‐sized resource at the onset of breeding, the size of this resource has larger effects on life‐history trade‐offs than resources acquired at earlier stages.  相似文献   

18.
Correlated responses to bi‐directional selection on thorax length, examined on several life‐history traits and chromosome inversion polymorphisms, have revealed apparent novel trade‐offs in Drosophila ananassae. We provide evidence of trade‐offs between hatching time and pupal period, pupal period and egg‐pupa development time, and pupal period and larval development time (LDT). Body size shows positive correlations with ovariole number, LDT and DT (egg–fly). We provide evidence of sexual dimorphism for trade‐offs between longevity and body size and starvation and longevity in females only. Trade‐offs between wing/thorax (W/T) ratio and longevity, W/T ratio and starvation, and DT (egg‐ fly) and longevity are evident in males only. Sexual dimorphism is also evident for inversion polymorphism with body size and longevity. A longevity assay suggests that low line females outlived high line females whereas high line males outlived low line males. The mean longevity in males is negatively correlated with the 2L‐ST and 3R‐ST arrangement frequencies whereas the 3L‐ST arrangement frequency is positively correlated with the mean longevity in males but opposite arrangements are found in females. Absolute starvation resistance is negatively correlated with 2L‐ST and 3R‐ST chromosome arrangements and results in a trade‐off between longevity and absolute starvation resistance in females. Analyses of fecundity, hatchability, and viabilities based on age intervals in both G10 and G13 suggest that the early reproduction is favoured in D. ananassae. The productivity percentage is highest in the high line and there is no effect of late reproduction on it. Overall, we provide some unravelled trade‐offs and striking sex differences, which may help in understanding the life‐history evolution of the species. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 90 , 669–685.  相似文献   

19.
To invest in energetically demanding life history stages, individuals require a substantial amount of resources. Physiological traits, particularly those related to energetics, can be useful for examining variation in life history decisions and trade‐offs because they result from individual responses to environmental variation. Leptin is a protein hormone found in mammals that is proportional to the amount of endogenous fat stores within an individual. Recently, researchers have confirmed that a mammalian leptin analogue (MLA), based on the mammalian sequence of leptin, is present with associated receptors and proteins in avian species, with an inhibitory effect on foraging and body mass gain at high circulating levels. While MLA has been both quantified and manipulated in avian species, little is currently known regarding whether plasma MLA in wild‐living species and individuals is associated with key reproductive decisions. We quantified plasma MLA in wild, Arctic‐nesting female common eiders (Somateria mollissima) at arrival on the breeding grounds and followed them to determine subsequent breeding propensity, and reproductive phenology, investment, and success. Common eiders are capital‐income breeding birds that require the accumulation of substantial fat stores to initiate laying and successfully complete incubation. We found that females with lower plasma MLA initiated breeding earlier and in a shorter period of time. However, we found no links between plasma MLA levels and breeding propensity, clutch size, or reproductive success. Although little is still known about plasma MLA, based on these results and its role in influencing foraging behaviors and condition gain, plasma MLA appears to be closely linked to reproductive timing and is therefore likely to underlie trade‐offs surrounding life history decisions.  相似文献   

20.
The Free Radical Theory of Ageing (FRTA) predicts that oxidative stress, induced when levels of reactive oxygen species exceed the capacity of antioxidant defenses, causes ageing. Recently, it has also been argued that oxidative damage may mediate important life‐history trade‐offs. Here, we use inbred lines of the decorated cricket, Gryllodes sigillatus, to estimate the genetic (co)variance between age‐dependent reproductive effort, life span, ageing, oxidative damage, and total antioxidant capacity within and between the sexes. The FRTA predicts that oxidative damage should accumulate with age and negatively correlate with life span. We find that protein oxidation is greater in the shorter lived sex (females) and negatively genetically correlated with life span in both sexes. However, oxidative damage did not accumulate with age in either sex. Previously we have shown antagonistic pleiotropy between the genes for early‐life reproductive effort and ageing rate in both sexes, although this was stronger in females. In females, we find that elevated fecundity early in life is associated with greater protein oxidation later in life, which is in turn positively correlated with the rate of ageing. Our results provide mixed support for the FRTA but suggest that oxidative stress may mediate sex‐specific life‐history strategies in G. sigillatus.  相似文献   

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