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PubMed2005

Are reproductive and somatic senescence coupled in humans? Late, but not early, reproduction correlated with longevity in historical Sami women.

Evolutionary theory of senescence emphasizes the importance of intense selection on early reproduction owing to the declining force of natural selection with age that constrains lifespan. In humans, recent studies have, however, suggested that late-life mortality might be more closely related to late rather than early reproduction, although the role of late reproduction on fitness remains unclear. We examined the association between early and late reproduction with longevity in historical post-reproductive Sami women. We also estimated the strength of natural selection on early and late reproduction using path analysis, and the effect of reproductive timing on offspring survival to adulthood and maternal risk of dying at childbirth. We found that natural selection favoured both earlier start and later cessation of reproduction, and higher total fecundity. Maternal age at childbirth was not related to offspring or maternal survival. Interestingly, females who produced their last offspring at advanced age also lived longest, while age at first reproduction and total fecundity were unrelated to female longevity. Our results thus suggest that reproductive and somatic senescence may have been coupled in these human populations, and that selection could have favoured late reproduction. We discuss alternative hypotheses for the mechanisms which might have promoted the association between late reproduction and longevity.

Aging↗
PubMed2005

Cost of reproduction in a long-lived bird: incubation effort reduces immune function and future reproduction.

Life-history theory predicts that increased current reproductive effort should lead to a fitness cost. This cost of reproduction may be observed as reduced survival or future reproduction, and may be caused by temporal suppression of immune function in stressed or hard-working individuals. In birds, consideration of the costs of incubating eggs has largely been neglected in favour of the costs of brood rearing. We manipulated incubation demand in two breeding seasons (2000 and 2001) in female common eiders (Somateria mollissima) by creating clutches of three and six eggs (natural range 3-6 eggs). The common eider is a long-lived sea-duck where females do not eat during the incubation period. Mass loss increased and immune function (lymphocyte levels and specific antibody response to the non-pathogenic antigens diphtheria and tetanus toxoid) was reduced in females incubating large clutches. The increased incubation effort among females assigned to large incubation demand did not lead to adverse effects on current reproduction or return rate in the next breeding season. However, large incubation demand resulted in long-term fitness costs through reduced fecundity the year after manipulation. Our data show that in eiders, a long-lived species, the cost of high incubation demand is paid in the currency of reduced future fecundity, possibly mediated by reduced immune function.

Analysis of Variance↗
PubMed2005

Does habitat fragmentation affect temperature-related life-history traits? A laboratory test with a woodland butterfly.

Habitat fragmentation may change local climatic conditions leading to altered selection regimes for life-history traits in small ectotherms, including several insects. We investigated temperature-related performance in terms of fitness among populations of the woodland butterfly Pararge aegeria (L.) originating from populations of a closed, continuous woodland landscape versus populations of an open, highly fragmented agricultural landscape in central Belgium. Female fecundity and longevity were evaluated in a temperature-gradient experiment. As predicted, females of woodland landscape origin reached higher maximum daily fecundity and lifetime number of eggs than did agricultural landscape females at low ambient temperatures, but this reversed at high ambient temperature. Egg weight decreased with temperature, and eggs of woodland butterflies were smaller. Contrary to what is generally assumed, remaining thorax mass was a better predictor of lifetime reproductive output than was abdomen mass. Since we used the F2 generation from wild-caught females reared under common garden conditions, the observed effects are likely to rely on intrinsic, heritable variation. Our results suggest that differential selection regimes associated with different landscapes intervene by intraspecific variation in the response of a butterfly to variation in ambient temperature, and may thus be helpful when making predictions of future impacts on how wild populations respond to environmental conditions under a global change scenario, with increasing temperatures and fragmented landscapes.

Adaptation, Physiological↗
PubMed2005

Changes in maternal investment in eggs can affect population dynamics.

The way that mothers provision their offspring can have important consequences for their offspring's performance throughout life. Models suggest that maternally induced variation in life histories may have large population dynamical effects, even perhaps driving cycles such as those seen in forest Lepidoptera. The evidence for large maternal influences on population dynamics is unconvincing, principally because of the difficulty of conducting experiments at both the individual and population level. In the soil mite, Sancassania berlesei, we show that there is a trade-off between a female's fecundity and the per-egg provisioning of protein. The mother's position on this trade-off depends on her current food availability and her age. Populations initiated with 250 eggs of different mean sizes showed significant differences in the population dynamics, converging only after three generations. Differences in the growth, maturation and fecundity of the initial cohort caused differences in the competitive environment for the next generation, which, in turn, created differences in their growth and reproduction. Maternal effects in one generation can therefore lead to population dynamical consequences over many generations. Where animals live in environments that are temporally variable, we conjecture that maternal effects could result in long-term dynamical effects.

Animal Nutritional Physiological Phenomena↗
PubMed2005

Biology of extinction risk in marine fishes.

We review interactions between extrinsic threats to marine fishes and intrinsic aspects of their biology that determine how populations and species respond to those threats. Information is available on the status of less than 5% of the world's approximately 15500 marine fish species, most of which are of commercial importance. By 2001, based on data from 98 North Atlantic and northeast Pacific populations, marine fishes had declined by a median 65% in breeding biomass from known historic levels; 28 populations had declined by more than 80%. Most of these declines would be sufficient to warrant a status of threatened with extinction under international threat criteria. However, this interpretation is highly controversial, in part because of a perception that marine fishes have a suite of life history characteristics, including high fecundity and large geographical ranges, which might confer greater resilience than that shown by terrestrial vertebrates. We review 15 comparative analyses that have tested for these and other life history correlates of vulnerability in marine fishes. The empirical evidence suggests that large body size and late maturity are the best predictors of vulnerability to fishing, regardless of whether differences among taxa in fishing mortality are controlled; there is no evidence that high fecundity confers increased resilience. The evidence reviewed here is of direct relevance to the diverse criteria used at global and national levels by various bodies to assess threat status of fishes. Simple life history traits can be incorporated directly into quantitative assessment criteria, or used to modify the conclusions of quantitative assessments, or used as preliminary screening criteria for assessment of the approximately 95% of marine fish species whose status has yet to be evaluated either by conservationists or fisheries scientists.

Animals↗
PubMed2006

Evidence for adaptive male mate choice in the fruit fly Drosophila melanogaster.

Theory predicts that males will benefit when they bias their mating effort towards females of higher reproductive potential, and that this discrimination will increase as males become more resource limited. We conducted a series of experiments to test these predictions in a laboratory population of the fruitfly, Drosophila melanogaster. In this species, courtship and copulation have significant costs to males, and females vary greatly in fecundity, which is positively associated with body size. When given a simultaneous choice between small and large virgin females, males preferentially mated with larger, more fecund, females. Moreover, after males had recently mated they showed a stronger preference for larger females. These results suggest that male D. melanogaster adaptively allocate their mating effort in response to variation in female quality and provide some of the first support for the theoretical prediction that male stringency in mate choice increases as resources become more limiting.

Adaptation, Physiological↗
PubMed2006

Disparate maturation adaptations to size-dependent mortality.

Body size is an important determinant of resource use, fecundity and mortality risk. Evolution of maturation size in response to size-dependent selection is thus a fundamental part of life-history theory. Increased mortality among small individuals has previously been predicted to cause larger maturation size, whereas increased mortality among large individuals is expected to have the opposite effect. Here we use a continuously size-structured model to demonstrate that, contrary to these widespread expectations, increased mortality among small individuals can have three alternative effects: maturation size may increase, decrease or become evolutionarily bistable. We show that such complex responses must be reckoned with whenever mortality is size-dependent, growth is indeterminate, reproduction impairs growth and fecundity increases with size. Predicting adaptive responses to altered size-dependent mortality is thus inherently difficult, since, as demonstrated here, such mortality cannot only reverse the direction of adaptation, but also cause abrupt shifts in evolutionarily stable maturation sizes.

Adaptation, Biological↗
PubMed2002

Male-only care and classical polyandry in birds: phylogeny, ecology and sex differences in remating opportunities.

It has been argued recently that the combination of male-only parental care and classical polyandry in birds is the most interesting and yet the least understood of all avian breeding systems. Despite a huge number of hypotheses, careful comparative analyses have repeatedly failed to identify consistent ecological differences between species showing male-only care and closely related species showing other patterns of care. This has led to the suggestion that such analyses fail because the crucial differences are between ancient lineages rather than between closely related species. Here, therefore, I use comparisons between families to test three well-known hypotheses: that male-only care is associated with: (i) a low rate of fecundity; (ii) large egg size relative to female size; or (iii) female-biased opportunities for remating. Families showing male-only care do not differ from families showing female-only care with respect to rate of fecundity or relative egg size. There is, however, a significant difference between these two groups of families with respect to an index of remating opportunities, nesting density. Families showing female-only care nest at high density, while those showing male-only care nest at very low density. This is one of the first times a consistent ecological correlate has been identified for male-only care in birds. It suggests that female-only care arises (or persists) in families where remating opportunities are abundant for both sexes, whereas male-only care arises (or persists) in families where remating opportunities are rare for both sexes and particularly scarce for males. This in turn suggests that sex differences in remating opportunities are the key ecological factor in determining male-only care and classical polyandry in birds.

Animals↗
PubMed2002

Life-history correlates of the evolution of live bearing in fishes.

Selection for live bearing is thought to occur when the benefits of increasing offspring survival exceed the costs of reduced fecundity, mobility and the increased metabolic demands of carrying offspring throughout development. We present evidence that live bearing has evolved from egg laying 12 times in teleost (bony) fishes, bringing the total number of transitions to 21 to 22 times in all fishes, including elasmobranchs (sharks and rays). Live bearers produce larger offspring than egg layers in all of 13 independent comparisons for which data were available. However, contrary to our expectation there has not been a consistent reduction in fecundity; live bearers have fewer offspring in seven out of the 11 available comparisons. It was predicted that live bearers would have a larger body size, as this facilitates accommodation of developing offspring. This prediction was upheld in 14 out of 20 comparisons. However, this trend was driven by elasmobranchs, with large live bearers in seven out of eight comparisons. Thus, while the evolution of live bearing in elasmobranchs is correlated with predicted increases in offspring size and adult size, teleost live bearers do not have such a consistent suite of life-history correlates. This suggests that constraints or selection pressures on associated life histories may differ in live-bearing elasmobranchs and teleost fishes.

Animals↗
PubMed2002

Resistance of cultivated tomato to cell content-feeding herbivores is regulated by the octadecanoid-signaling pathway.

The octadecanoid signaling pathway has been shown to play an important role in plant defense against various chewing insects and some pathogenic fungi. Here, we examined the interaction of a cell-content feeding arachnid herbivore, the two-spotted spider mite (Tetranychus urticae Koch), with cultivated tomato (Lycopersicon esculentum) and an isogenic mutant line (defenseless-1 [def-1]) that is deficient in the biosynthesis of the octadecanoid pathway-derived signal, jasmonic acid (JA). Spider mite feeding and fecundity on def-1 plants was significantly greater than on wild-type plants. Decreased resistance of def-1 plants was correlated with reduced JA accumulation and expression of defensive proteinase inhibitor (PI) genes, which were induced in mite-damaged wild-type leaves. Treatment of def-1 plants with methyl-JA restored resistance to spider mite feeding and reduced the fecundity of female mites. Plants expressing a 35S::prosystemin transgene that constitutively activates the octadecanoid pathway in a Def-1-dependent manner were highly resistant to attack by spider mites and western flower thrips (Frankliniella occidentalis), another cell-content feeder of economic importance. These findings indicate that activation of the octadecanoid signaling pathway promotes resistance of tomato to a broad spectrum of herbivores. The techniques of amplified fragment length polymorphism (AFLP) and bulk segregant analysis were used to map the Def-1 gene to a region on the long arm of chromosome 3 that is genetically separable from the map position of known JA biosynthetic genes. Tight linkage of Def-1 to a T-DNA insertion harboring the maize (Zea mays) Dissociation transposable element suggests a strategy for directed transposon tagging of the gene.

Acetates↗
PubMed2000

Evolution in stressful environments. I. Phenotypic variability, phenotypic selection, and response to selection in five distinct environmental stresses.

Considerable debate has accompanied efforts to integrate the selective impacts of environmental stresses into models of life-history evolution. This study was designed to determine if different environmental stresses have consistent phenotypic effects on life-history characters and whether selection under different stresses leads to consistent evolutionary responses. We created lineages of a wild mustard (Sinapis arvensis) that were selected for three generations under five stress regimes (high boron, high salt, low light, low water, or low nutrients) or under near-optimal conditions (control). Full-sibling families from the six selection histories were divided among the same six experimental treatments. In that test generation, lifetime plant fecundity and six phenotypic traits were measured for each plant. Throughout this greenhouse study, plants were grown individually and stresses were applied from the early seedling stage through senescence. Although all stresses consistently reduced lifetime fecundity and most size- and growth-related traits, different stresses had contrasting effects on flowering time. On average, stress delayed flowering compared to favorable conditions, although plants experiencing low nutrient stress flowered earliest and those experiencing low light flowered latest. Contrary to expectations of Grime's triangle model of life-history evolution, this ruderal species does not respond phenotypically to poor environments by flowering earlier. Most stresses enhanced the evolutionary potential of the study population. Compared with near-optimal conditions, stresses tended to increase the opportunity for selection as well as phenotypic variance, although both of these quantities were reduced in some stresses. Rather than favoring traits characteristic of stress tolerance, such as slow growth and delayed reproduction, phenotypic selection favored stress-avoidance traits: earlier flowering in all five stress regimes and faster seedling height growth in three stresses. Phenotypic correlations reinforced direct selection on these traits under stress, leading to predicted phenotypic change under stress, but no significant selection in the control environment. As a result of these factors, selection under stress resulted in an evolutionary shift toward earlier flowering. Environmental stresses may drive populations of ruderal plant species like S. arvensis toward a stress-avoidance strategy, rather than toward stress tolerance. Further studies will be needed to determine when selection in stressful environments leads to these alternative life-history strategies.

Biological Evolution↗
PubMed2000

Reproductive effort in viscous populations.

Here I study a kin selection model of reproductive effort, the allocation of resources to fecundity versus survival, in a patch-structured population. Breeding females remain in the same patch for life. Offspring have costly, partial long-distance dispersal and compete for breeding sites, which become vacant upon the death of previous occupants. The main result is that the evolutionarily stable reproductive effort decreases as offspring dispersal rate increases. The result can be understood as follows: In a well-mixed population with global competition, neither adults nor juveniles compete with relatives, but in a patch-structured population with dispersal restricted to the juvenile phase, juveniles experience relatively less competition with relatives than adults, thus making juveniles relatively more valuable. Because this asymmetry between adults and juveniles decreases with the dispersal rate, so does the evolutionarily stable level of allocation to fecundity.

Animals↗
PubMed2001

An unusual sex-determination system in South American field mice (Genus Akodon): the role of mutation, selection, and meiotic drive in maintaining XY females.

The mechanism of sex determination in mammals appears highly conserved: the presence of a Y chromosome triggers the male developmental pathway, whereas the absence of a Y chromosome results in a default female phenotype. However, if the Y chromosome fails to initiate the male pathway (referred to as Y*), XY* females can result, as is the case in several species of South American field mice (genus Akodon). The breeding genetics in this system inherently select against the Y* chromosome such that the frequency of XY* females should decrease rapidly to very low frequencies. However, in natural populations of Akodon, XY* females persist at substantial frequencies; for example, 10% of females are XY* in A. azarae and 30% in A. boliviensis. We develop a mathematical model that considers the potential roles of three evolutionary forces in maintaining XY* females: Y-to-Y* chromosome transitions (mutation), chromosome segregation distortion (meiotic drive), and differential fecundity (selection). We then test the predictions of our model using data from breeding colonies of A. azarae. We conclude that any single force is inadequate to maintain XY* females. However, a combination of segregation bias of the male and female Y chromosomes during spermatogenesis/oogenesis and increased fecundity in XY* females could account for the observed frequencies of XY* females.

Animals↗
PubMed2001

The contribution of phenotypic plasticity to adaptation in Lacerta vivipara.

Correlation between intraspecific phenotypic variability and variation of environmental conditions could reflect adaptation. Different phenotypes may result from differential expression of a genotype in different environments (phenotypic plasticity) or from expression of different genotypes (genetic diversity). Populations of Lacerta vivipara exhibit larger adult body length, lower age at maturity, higher fecundity, and smaller neonatal size in humid habitats compared to dry habitats. We conducted reciprocal transplants of juvenile L. vivipara to test for the genetic or plastic origin of this variation. We captured gravid females from four populations that differed in the relative humidity of their habitats, and during the last 2 to 4 weeks of gestation, we manipulated heat and water availability under laboratory conditions. Juveniles were released into the different populations and families were divided to compare growth rate and survival of half-sibs in two environments. Growth rate and survival were assessed using capture-recapture techniques. Growth rate was plastic in response to postnatal conditions and did not differ between populations of origin. Survival differed between populations of origin, partially because of differences in neonatal body length. The response of juvenile body length and body condition to selection in the different habitats was affected by the population of origin. This result cannot be simply interpreted in terms of adaptation; however, phenotypic plasticity of fecundity or juvenile size most probably resulted in adaptive reproductive strategies. Adaptation to the habitat by means of genetic specialization was not detected. Further investigation is needed to discriminate between genetic and long-term maternal effects.

Adaptation, Physiological↗
PubMed2002

The evolution of trade-offs: testing predictions on response to selection and environmental variation.

The concept of phenotypic trade-offs is a central element in evolutionary theory. In general, phenotypic models assume a fixed trade-off function, whereas quantitative genetic theory predicts that the trade-off function will change as a result of selection. For a linear trade-off function selection will readily change the intercept but will have to be relatively stronger to change the slope. We test these predictions by examining the trade-off between fecundity and flight capability, as measured by dorso-longitudinal muscle mass, in four different populations of the sand cricket, Gryllus firmus. Three populations were recently derived from the wild, and the fourth had been in the laboratory for 19 years. We hypothesized that the laboratory population had most likely undergone more and different selection from the three wild populations and therefore should differ from these in respect to both slope and intercept. Because of geographic variation in selection, we predicted a general difference in intercept among the four populations. We further tested the hypothesis that this intercept will be correlated with proportion macropterous and that this relationship will itself vary with environmental conditions experienced during both the nymphal and adult period. Observed variation in the phenotypic trade-off was consistent with the predictions of the quantitative genetic model. These results point to the importance of modeling trade-offs as dynamic rather than static relationships. We discuss how phenotypic models can incorporate such variation. The phenotypic trade-off between fecundity and dorso-longitudinal muscle mass is determined in part by variation in body size, illustrating the necessity of considering trade-offs to be multi factorial rather than simply bivariate relationships.

Animals↗
PubMed2002

Evolutionary optimality applied to Drosophila experiments: hypothesis of constrained reproductive efficiency.

The general purpose of the paper is to test evolutionary optimality theories with experimental data on reproduction, energy consumption, and longevity in a particular Drosophila genotype. We describe the resource allocation in Drosophila females in terms of the oxygen consumption rates devoted to reproduction and to maintenance. The maximum ratio of the component spent on reproduction to the total rate of oxygen consumption, which can be realized by the female reproductive machinery, is called metabolic reproductive efficiency (MRE). We regard MRE as an evolutionary constraint. We demonstrate that MRE may be evaluated for a particular Drosophila phenotype given the fecundity pattern, the age-related pattern of oxygen consumption rate, and the longevity. We use a homeostatic model of aging to simulate a life history of a representative female fly, which describes the control strain in the long-term experiments with the Wayne State Drosophila genotype. We evaluate the theoretically optimal trade-offs in this genotype. Then we apply the Van Noordwijk-de Jong resource acquisition and allocation model, Kirkwood's disposable soma theory. and the Partridge-Barton optimality approach to test if the experimentally observed trade-offs may be regarded as close to the theoretically optimal ones. We demonstrate that the two approaches by Partridge-Barton and Kirkwood allow a positive answer to the question, whereas the Van Noordwijk-de Jong approach may be used to illustrate the optimality. We discuss the prospects of applying the proposed technique to various Drosophila experiments, in particular those including manipulations affecting fecundity.

Aging↗
PubMed2002

Hormonal manipulation of offspring number: maternal effort and reproductive costs.

We used exogenous gonadotropin hormones to physiologically enlarge litter size in the bank vole (Clethrionomys glareolus). This method allowed the study design to include possible production costs of reproduction and a trade-off between offspring number and body size at birth. Furthermore, progeny rearing and survival and postpartum survival of the females took place in outdoor enclosures to capture salient naturalistic effects that might be present during the fall and early winter. The aim of the study was to assess the effects of the manipulation on the growth and survival of the offspring and on the reproductive effort, survival, and future fecundity of the mothers. Mean offspring body size was smaller in enlarged litters compared to control litters at weaning, but the differences disappeared by the winter. Differences in litter sizes disappeared before weaning age due to higher mortality in enlarged litters. In addition to the effects of the litter size, offspring performance was probably also influenced by the ability of the mother to support the litter. Experimental females had higher reproductive effort at birth, and they also tended to have higher mortality during nursing. Combined effects of high reproductive effort at birth and high investment in nursing the litter entailed costs for the experimental females in terms of decreased probability of producing a second litter and a decreased body mass gain. Thus, enlarged litter size had both survival and fecundity costs for the mothers. Our results suggest that the evolution of litter size and reproductive effort is determined by reproductive costs for the mothers as well as by a trade-off between offspring number and quality.

Animals↗
PubMed2003

Genetic architecture of differences between populations of cowpea weevil (Callosobruchus maculatus) evolved in the same environment.

We investigated the genetic architecture underlying differentiation in fitness-related traits between two pairs of populations of the seed beetle Callosobruchus maculatus (Coleoptera: Bruchidae). These populations had geographically distant (> 2000 km) origins but evolved in a uniform laboratory environment for 120 generations. For each pair of populations (Nigeria x Yemen and Cameroon x Uganda) we estimated the means of five fitness-related characters and a measure of fitness (net reproductive rate R0) in each of the parental populations and 12 types of hybrids (two F1 and two F2 lines and eight backcrosses). Models containing up to nine composite genetic parameters were fitted to the means of the 14 lines. The patterns of line means for all traits in the Nigeria x Yemen cross and for four traits (larval survival, developmental rate, female body weight, and fecundity) in the Cameroon x Uganda cross were best explained by models including additive, dominance, and maternal effects, but excluding epistasis. We did not find any evidence for outbreeding depression for any trait. An epistatic component of divergence was detected for egg hatching success and R0 in the Cameroon x Uganda cross, but its sign was opposite to that expected under outbreeding depression, that is, additive x additive epistasis had a positive effect on the performance of F2 hybrids. All traits except fecundity showed a pattern of heterosis. A large difference of egg-hatching success between the two reciprocal F1 lines in that cross was best explained as fertilization incompatibility between Cameroon females and sperm carrying Uganda genes. The results suggest that these populations have not converged to the same life-history phenotype and genetic architecture, despite 120 generations of uniform natural selection. However, the absence of outbreeding depression implies that they did not evolve toward different adaptive peaks.

Animals↗