Variation in fecundity and other reproductive traits in freshwater mussels

1. Life histories of the highly diverse and endangered North American freshwater mussel fauna are poorly known. We investigated reproductive traits of eight riverine mussel species in Alabama and Mississippi, U.S.A.: Amblema plicata, Elliptio arca, Fusconaia cerina, Lampsilis ornata, Obliquaria reflexa, Pleurobema decisum, Quadrula asperata and Q. pustulosa, and compare our results with existing life history information for other species. 2. These eight species had reproductive traits characteristic of large, outcrossing populations: hermaphrodites were rare, we found no evidence of protandry, and sex ratios were even or slightly male‐biased. 3. Age at sexual maturity varied among species, ranging from <1 to 2 years for L. ornata to 3–9 years for Q. asperata. In all species, most mature females participated in reproduction and fertilisation success was high. 5. Fecundity was related positively to both length and age, but length was the best predictor. In six species, fecundity increased exponentially with increasing size; in two species the rate of increase in fecundity declined in larger animals. In four species, fecundity declined in older animals. These latter results indicate weak reproductive senescence; however, in all species, older individuals continued to produce large numbers of offspring. Mean annual fecundity differed widely among species ranging from 9647 to 325 709. Within‐species differences in fecundity were found among rivers and among populations within a river. 6. The wide variation in reproductive traits among species indicates the existence of widely divergent life history strategies in freshwater mussels.     

Food availability as a major driver in the evolution of life‐history strategies of sibling species

Life‐history theory predicts trade‐offs between reproductive and survival traits such that different strategies or environmental constraints may yield comparable lifetime reproductive success among conspecifics. Food availability is one of the most important environmental factors shaping developmental processes. It notably affects key life‐history components such as reproduction and survival prospect. We investigated whether food resource availability could also operate as an ultimate driver of life‐history strategy variation between species. During 13 years, we marked and recaptured young and adult sibling mouse‐eared bats (Myotis myotis and Myotis blythii) at sympatric colonial sites. We tested whether distinct, species‐specific trophic niches and food availability patterns may drive interspecific differences in key life‐history components such as age at first reproduction and survival. We took advantage of a quasi‐experimental setting in which prey availability for the two species varies between years (pulse vs. nonpulse resource years), modeling mark‐recapture data for demographic comparisons. Prey availability dictated both adult survival and age at first reproduction. The bat species facing a more abundant and predictable food supply early in the season started its reproductive life earlier and showed a lower adult survival probability than the species subjected to more limited and less predictable food supply, while lifetime reproductive success was comparable in both species. The observed life‐history trade‐off indicates that temporal patterns in food availability can drive evolutionary divergence in life‐history strategies among sympatric sibling species.     

Evaluating the life‐history trade‐off between dispersal capability and reproduction in wing dimorphic insects: a meta‐analysis

A life‐history trade‐off exists between flight capability and reproduction in many wing dimorphic insects: a long‐winged morph is flight‐capable at the expense of reproduction, while a short‐winged morph cannot fly, is less mobile, but has greater reproductive output. Using meta‐analyses, I investigated specific questions regarding this trade‐off. The trade‐off in females was expressed primarily as a later onset of egg production and lower fecundity in long‐winged females relative to short‐winged females. Although considerably less work has been done with males, the trade‐off exists for males among traits primarily related to mate acquisition. The trade‐off can potentially be mitigated in males, as long‐winged individuals possess an advantage in traits that can offset the costs of flight capability such as a shorter development time. The strength and direction of trends differed significantly among insect orders, and there was a relationship between the strength and direction of trends with the relative flight capabilities between the morphs. I discuss how the trade‐off might be both under‐ and overestimated in the literature, especially in light of work that has examined two relevant aspects of wing dimorphic species: (1) the effect of flight‐muscle histolysis on reproductive investment; and (2) the performance of actual flight by flight‐capable individuals.     

Trade‐offs between life‐history traits at range‐edge and central locations

The allocation of resources to different life‐history traits should represent the best compromise in fitness investment for organisms in their local environment. When resources are limiting, the investment in a specific trait must carry a cost that is expressed in trade‐offs with other traits. In this study, the relative investment in the fitness‐related traits, growth, reproduction and defence were compared at central and range‐edge locations, using the seaweed Ascophyllum nodosum as a model system. Individual growth rates were similar at both sites, whereas edge populations showed a higher relative investment in reproduction (demonstrated by a higher reproductive allocation and extended reproductive periods) when compared to central populations that invested more in defence. These results show the capability of A. nodosum to differentially allocate resources for different traits under different habitat conditions, suggesting that reproduction and defence have different fitness values under the specific living conditions experienced at edge and central locations. However, ongoing climate change may threaten edge populations by increasing the selective pressure on specific traits, forcing these populations to lower the investment in other traits that are also potentially important for population fitness.     

An experimental test for alternative reproductive strategies underlying a female-limited polymorphism

Polymorphism often corresponds to alternative mating tactics in males, but much less is known about this relationship in females. However, recent work suggests that selection for alternative reproductive strategies in females can maintain genetic variation in important life-history traits. Brown anole lizards (Anolis sagrei) exhibit a genetically based polymorphism in dorsal pattern that is expressed only by females, which occur in bar (B), diamond (D) and intermediate diamond-bar (DB) morphs. Here, we use a combination of natural history data, captive breeding studies and phenotypic manipulations of reproductive investment to test the hypothesis that this polymorphism corresponds to morph-specific patterns of reproductive investment. Three years of data from wild females and two generations of captive breeding revealed no differences among morphs in the frequency of egg production or in the number, frequency, size or sex ratio of offspring. Manipulations of reproductive investment via surgical ovariectomy revealed significant costs of reproduction with respect to survival, growth, immune function and haematocrit, but the magnitudes of these costs did not differ among morphs. Collectively, our results refute the hypothesis that this sex-limited polymorphism is maintained by selection for alternative reproductive strategies. We compare this finding to other systems in which polymorphic females exhibit alternative reproductive tactics and discuss other selective factors that could maintain polymorphism in anoles.     

Inbreeding alters context‐dependent reproductive effort and immunity in male crickets

Infection can cause hosts to drastically alter their investment in key life‐history traits of reproduction and defence. Infected individuals are expected to increase investment in defence (e.g., by increasing immune function) and, due to trade‐offs, investment in other traits (e.g., current reproduction) should decrease. However, the terminal investment hypothesis postulates that decreased lifespan due to infection and the associated reduction in the expectation for future offspring will favour increased investment towards current reproduction. Variation in intrinsic condition will likely influence shifts in reproductive investment post‐infection, but this is often not considered in such assessments. For example, the extent of inbreeding can significantly impact an individual's lifetime fitness and may influence its reproductive behaviour following a threat of infection. Here, we investigated the effects of inbreeding status on an individual's reproductive investment upon infection, including the propensity to terminally invest. Male crickets (Gryllodes sigillatus) from four genetically distinct inbred lines and one outbred line were subjected to a treatment from an increasing spectrum of simulated infection cue intensities, using heat‐killed bacteria. We then measured reproductive effort (calling effort), survival and immune function (antibacterial activity, circulating haemocytes and haemocyte microaggregations). Inbred and outbred males diverged in how they responded to a low‐dose infection cue: relative to unmanipulated males, outbred males decreased calling effort, whereas inbred males increased calling effort. Moreover, we found that inbred males exhibited higher antibacterial activity and numbers of circulating haemocytes compared with outbred males. These results suggest that an individual's inbreeding status may have consequences for context‐dependent shifts in reproductive strategies, such as those triggered by infection.     

Geographical variation in reproductive ageing patterns and life‐history strategy of a short‐lived passerine bird

We investigated differences in ageing patterns in three measures of breeding performance in populations of barn swallows Hirundo rustica L. from Spain and Denmark differing in breeding latitude and hence migration distance and duration of the breeding season. We found differences in ageing patterns between populations. Generally, young (i.e. yearling) and old females (i.e. ≥ 5 years of age) laid their first eggs later and produced smaller clutches than middle‐aged females (i.e. 2–4 years of age) in both populations. The southernmost population (i.e. Spanish) showing the shorter migratory distance experienced a greater within‐individual increase in timing of breeding and clutch size in early life and a greater within‐individual decrease in laying date but not in clutch size during senescence compared with the northernmost population (i.e. Danish). We also found that the number of fledglings produced annually was related to the age of the two members of the breeding pairs with pairs composed of young and old females performing less well than breeding pairs composed of middle‐aged females. We did not find reproductive senescence for the age of the male while controlling for the age of the female on the number of fledglings produced annually by the breeding pair. Differential survival between individuals did not explain age effects on laying date or annual clutch size in neither population. However, the increase in the number of fledglings produced annually with age was partly explained by the disappearance of poor‐quality members of the pairs, mainly poor‐quality males. Age‐related breeding success (i.e. number of fledglings) was similar for barn swallows from Spain and Denmark. Therefore, the study of ageing patterns and life‐history strategies in free‐ranging animals from more than a single population can throw new light on life‐history theory, population dynamics and evolutionary studies of senescence.     

The early‐life environment and individual plasticity in life‐history traits

We tested whether the early‐life environment can influence the extent of individual plasticity in a life‐history trait. We asked: can the early‐life environment explain why, in response to the same adult environmental cue, some individuals invest more than others in current reproduction? Moreover, can it additionally explain why investment in current reproduction trades off against survival in some individuals, but is positively correlated with survival in others? We addressed these questions using the burying beetle, which breeds on small carcasses and sometimes carries phoretic mites. These mites breed alongside the beetle, on the same resource, and are a key component of the beetle's early‐life environment. We exposed female beetles to mites twice during their lives: during their development as larvae and again as adults during their first reproductive event. We measured investment in current reproduction by quantifying average larval mass and recorded the female's life span after breeding to quantify survival. We found no effect of either developing or breeding alongside mites on female reproductive investment, nor on her life span, nor did developing alongside mites influence her size. In post hoc analyses, where we considered the effect of mite number (rather than their mere presence/absence) during the female's adult breeding event, we found that females invested more in current reproduction when exposed to greater mite densities during reproduction, but only if they had been exposed to mites during development as well. Otherwise, they invested less in larvae at greater mite densities. Furthermore, females that had developed with mites exhibited a trade‐off between investment in current reproduction and future survival, whereas these traits were positively correlated in females that had developed without mites. The early‐life environment thus generates individual variation in life‐history plasticity. We discuss whether this is because mites influence the resources available to developing young or serve as important environmental cues.     

Trait‐mediated effects of predation across life‐history stages in container mosquitoes

1. It was determined if the predatory midge Corethrella appendiculata Grabham imposes a fitness cost in a native mosquito, Ochlerotatus triseriatus Say, and an invasive mosquito, Aedes albopictus Skuse. The hypothesis that decreased activity of immature prey in the presence of predator cues is associated with life history costs through all life cycle stages was tested. 2. In experiment 1, individual larvae of O. triseriatus or A. albopictus were raised in the presence or absence of predation cues at two resource levels. Prey were video recorded to detect behavioural responses and to measure development time, size at emergence, and adult longevity. In experiment 2, prey populations were reared in similar environments and the frequency of predator cue additions was varied. 3. Only O. triseriatus reduced its activity in the presence of predation cues. Predation cues were associated with longer immature development times and shorter adult life spans in O. triseriatus, whereas in A. albopictus, the cues were associated with a larger size of emerging adults. 4. In the present study, it was found that behavioural modifications during the larval stage can affect mosquitoes through multiple stages of their complex life cycle. The species‐specific behavioural differences are probably attributable to the longer evolutionary history O. triseriatus has with predators, relative to the invasive A. albopictus.     

Hot and not‐so‐hot females: reproductive state and thermal preferences of female Arizona Bark Scorpions (Centruroides sculpturatus)

For ectotherms, environmental temperatures influence numerous life history characteristics, and the body temperatures (T_b) selected by individuals can affect offspring fitness and parental survival. Reproductive trade‐offs may therefore ensue for gravid females, because temperatures conducive to embryonic development may compromise females' body condition. We tested whether reproduction influenced thermoregulation in female Arizona Bark Scorpions (Centruroides sculpturatus). We predicted that gravid females select higher T_b and thermoregulate more precisely than nonreproductive females. Gravid C. sculpturatus gain body mass throughout gestation, which exposes larger portions of their pleural membrane, possibly increasing their rates of transcuticular water loss in arid environments. Accordingly, we tested whether gravid C. sculpturatus lose water faster than nonreproductive females. We determined the preferred T_b of female scorpions in a thermal gradient and measured water loss rates using flow‐through respirometry. Gravid females preferred significantly higher T_b than nonreproductive females, suggesting that gravid C. sculpturatus alter their thermoregulatory behaviour to promote offspring fitness. However, all scorpions thermoregulated with equal precision, perhaps because arid conditions create selective pressure on all females to thermoregulate effectively. Gravid females lost water faster than nonreproductive animals, indicating that greater exposure of the pleural membrane during gestation enhances the desiccation risk of reproductive females. Our findings suggest that gravid C. sculpturatus experience a trade‐off, whereby selection of higher T_b and increased mass during gestation increase females' susceptibility to water loss, and thus their mortality risk. Elucidating the mechanisms that influence thermal preferences may reveal how reproductive trade‐offs shape the life history of ectotherms in arid environments.     

Living on the edge: life‐history of olive baboons at Gashaka‐Gumti National Park,Nigeria

Baboons are the most successful and ubiquitous African primates, renowned for their behavioral and reproductive flexibility, which enable them to inhabit a wide variety of habitat types. Owing to a number of long‐term field studies, comparative behavioral, developmental, demographic, and life‐history data are available from several populations, but study sites show a heavy bias toward South and East African savannahs, with little research in West or Central Africa. Life‐history data from such areas are important if we are fully to understand the nature of the environmental factors that limit baboon distribution. Here, we present demographic data for olive baboons at Gashaka‐Gumti National Park (GGNP), Nigeria, collected from December 2000–February 2006, and use these data to test comparative models of baboon life‐history. The GGNP habitat, which includes large areas of rainforest, is an environment in which baboons are little studied, and rainfall is much higher than at previous study sites. GGNP troop size data are presented from censuses, as well as life‐history data for two troops, one of which is within the park and wild‐feeding (Kwano troop), whereas the other dwells at the park edge, and supplements its diet by crop‐raiding (Gamgam troop). Troop sizes at GGNP are small compared with other field sites, but fit within previously suggested ranges for baboons under these climatic conditions. Inter‐birth intervals in Kwano troop were long compared with most studied populations, and values were not as predicted by comparative models. Consistent with known effects of food enhancement, Gamgam troop experienced shorter inter‐birth intervals and lower infant mortality than Kwano troop. We indicate some possible factors that exclude baboons from true rainforest, and suggest that the clearing of forests in Central and West Africa for agricultural land may allow baboons to extend their range into regions from which they are currently excluded. Am. J. Primatol. 71:293–304, 2009. © 2009 Wiley‐Liss, Inc.     

Estimation of fitness from energetics and life‐history data: An example using mussels

Changing environments have the potential to alter the fitness of organisms through effects on components of fitness such as energy acquisition, metabolic cost, growth rate, survivorship, and reproductive output. Organisms, on the other hand, can alter aspects of their physiology and life histories through phenotypic plasticity as well as through genetic change in populations (selection). Researchers examining the effects of environmental variables frequently concentrate on individual components of fitness, although methods exist to combine these into a population level estimate of average fitness, as the per capita rate of population growth for a set of identical individuals with a particular set of traits. Recent advances in energetic modeling have provided excellent data on energy intake and costs leading to growth, reproduction, and other life‐history parameters; these in turn have consequences for survivorship at all life‐history stages, and thus for fitness. Components of fitness alone (performance measures) are useful in determining organism response to changing conditions, but are often not good predictors of fitness; they can differ in both form and magnitude, as demonstrated in our model. Here, we combine an energetics model for growth and allocation with a matrix model that calculates population growth rate for a group of individuals with a particular set of traits. We use intertidal mussels as an example, because data exist for some of the important energetic and life‐history parameters, and because there is a hypothesized energetic trade‐off between byssus production (affecting survivorship), and energy used for growth and reproduction. The model shows exactly how strong this trade‐off is in terms of overall fitness, and it illustrates conditions where fitness components are good predictors of actual fitness, and cases where they are not. In addition, the model is used to examine the effects of environmental change on this trade‐off and on both fitness and on individual fitness components.     

Macroevolutionary evidence suggests trait‐dependent coevolution between behavior and life‐history

Species with fast life‐histories typically prioritize current over future reproductive events, compared to species with slow life‐histories. These species therefore require greater energetic input into reproduction, and also likely have less time to realize their reproductive potential. Hence, behaviors that increase access to both resources and mating opportunities, at a cost of increased mortality risk, could coevolve with the pace of life‐history. However, whether this prediction holds across species, remains untested under standardized conditions. Here, we test how risky behaviors, which facilitate access to resources and mating opportunities (i.e., activity, boldness, and aggression), along with metabolic rate, coevolve with the pace of life‐history across 20 species of killifish that present remarkable divergences in the pace of life‐history. We found a positive association between the pace of life‐history and aggression, but interestingly not with other behavioral traits or metabolic rate. Aggression is linked to interference competition, and in killifishes is often employed to secure mates, while activity and boldness are more relevant for exploiting energetic resources. Our results suggest that the trade‐off between current and future reproduction plays a more prominent role in shaping mating behavior, while behaviors related to energy acquisition may be influenced by ecological factors.     

Heat shock proteins mediate trade‐offs between early‐life reproduction and late survival in Drosophila melanogaster

Ageing and the resulting increased likelihood mortality are the inescapable fate of organisms because selection pressures on genes that exert their function late in life is weak, promoting the evolution of genes that enhance early‐life reproductive performance at the same time as sacrificing late survival. Heat shock proteins (HSP) are known to buffer various environmental stresses and are also involved in protein homeostasis and longevity. The characteristics of genes for HSPs (hsp) imply that they affect various life‐history traits, which in turn affect longevity; however, little is known about the effects of hsp genes on life‐history traits and their interaction with longevity. In the present study, the effects of hsp genes on multiple fitness traits, such as locomotor activity, total fecundity, early fecundity and survival time, are investigated in Drosophila melanogaster Meigen using RNA interference (RNAi). In egg‐laying females, RNAi knockdown of six hsp genes (hsp22, hsp23, hsp67Ba, hsp67Bb, hsp67Bc and hsp27‐like) does not shorten survival but rather increases it. Knockdown of five of those genes on an individual basis reduces early‐life reproduction, suggesting that several hsp genes mediate the trade‐off between early reproduction and late survival. The data indicate a positive effect of hsp genes on early reproduction and also negative effects on survival time, supporting the antagonistic pleiotropic effects predicted by the optimality theory of ageing.     

Recovery and immune priming modulate the evolutionary trajectory of infection‐induced reproductive strategies

In response to parasite exposure, organisms from a variety of taxa undergo a shift in reproductive investment that may trade off with other life‐history traits including survival and immunity. By suppressing reproduction in favour of somatic and immunological maintenance, hosts can enhance the probability of survival and recovery from infection. By plastically enhancing reproduction through terminal investment, on the other hand, hosts under the threat of disease‐induced mortality could enhance their lifetime reproductive fitness through reproduction rather than survival. However, we know little about the evolution of these strategies, particularly when hosts can recover and even bequeath protection to their offspring. In this study, we develop a stochastic agent‐based model that competes somatic maintenance and terminal investment strategies as they trade off differentially with lifespan, parasite resistance, recovery and transgenerational immune priming. Our results suggest that a trade‐off between reproduction and recovery can drive directional selection for either terminal investment or somatic maintenance, depending on the cost of reproduction to lifespan. However, some conditions, such as low virulence with a high cost of reproduction to lifespan, can favour diversifying selection for the coexistence of both strategies. The introduction of transgenerational priming into the model favours terminal investment when all strategies are equally likely to produce primed offspring, but favours somatic maintenance if it confers even a slight priming advantage over terminal investment. Our results suggest that both immune priming and recovery may modulate the evolution of reproductive shift diversity and magnitude upon exposure to parasites.     

The demographic and life‐history costs of fear: Trait‐mediated effects of threat of predation on Aedes triseriatus

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Time spent in distinct life history stages has sex‐specific effects on reproductive fitness in wild Atlantic salmon

In species with complex life cycles, life history theory predicts that fitness is affected by conditions encountered in previous life history stages. Here, we use a 4‐year pedigree to investigate if time spent in two distinct life history stages has sex‐specific reproductive fitness consequences in anadromous Atlantic salmon (Salmo salar). We determined the amount of years spent in fresh water as juveniles (freshwater age, FW, measured in years), and years spent in the marine environment as adults (sea age, SW, measured in sea winters) on 264 sexually mature adults collected on a river spawning ground. We then estimated reproductive fitness as the number of offspring (reproductive success) and the number of mates (mating success) using genetic parentage analysis (>5,000 offspring). Sea age is significantly and positively correlated with reproductive and mating success of both sexes whereby older and larger individuals gained the highest reproductive fitness benefits (females: 62.2% increase in offspring/SW and 34.8% increase in mate number/SW; males: 201.9% offspring/SW and 60.3% mates/SW). Younger freshwater age was significantly related to older sea age and thus increased reproductive fitness, but only among females (females: ?33.9% offspring/FW and ?32.4% mates/FW). This result implies that females can obtain higher reproductive fitness by transitioning to the marine environment earlier. In contrast, male mating and reproductive success was unaffected by freshwater age and more males returned at a younger age than females despite the reproductive fitness advantage of later sea age maturation. Our results show that the timing of transitions between juvenile and adult phases has a sex‐specific consequence on female reproductive fitness, demonstrating a life history trade‐off between maturation and reproduction in wild Atlantic salmon.     

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.