The short-term and long-term effects of parental age in the bean weevil (Acanthoscelides obtectus)

We examined the influence of parental age on life history traits of their offspring in the lines of bean weevil that have evolved different rates of senescence. Measurements included preadult traits (egg size, embryonic developmental time, total preadult developmental time, preadult viability) and adult traits (body weight, total realized fecundity of females, first day of egg laying, early fecundity, late fecundity and longevity). The negative parental age effects were observed for all traits except for the early and total realized fecundity. We did not detect statistically significant line×parental age interactions for either preadult- or adult-survival, so offspring survival did not change with parental age after selection for early vs. late reproduction. It seems that selection acting on the quality of offspring produced by parents of different ages has not been responsible for the evolution of senescence in bean weevil. This revised version was published online in August 2006 with corrections to the Cover Date.     

Phenotypic selection and regulation of reproduction in different environments in wild barley

Plasticity of the phenotypic architecture of wild barley, Hordeum spontaneum, was studied in response to water and nutrient stress. Direct and indirect selection on several vegetative and reproductive traits was estimated and path analysis used to reveal how regulating pathways via maternal investment differed between environments. Vegetative traits displayed differential regulating effect on fitness across experimental environments: (1) increase in size was selected for under optimal conditions and under water stress, but not under nutrient stress; (2) allocation to root biomass was selected for under optimal conditions, but it had no effect under nutrient stress and was strongly selected against when water was limiting; (3) delayed onset of reproduction was selected under nutrient limitation whereas earlier onset was selected under water stress. The regulating effect of reproductive traits on final reproductive output also differed across treatments, operating either at the 'early' stage of plant development through varying the number of initiated spikelets per spike (no stress and water stress treatment) or at the 'late' developmental stage adjusting the fertile spikelet weight (no stress and nutrient stress treatment). Reproductive output was regulated via seed abortion under no stress and water stress treatments. Although the underlying mechanism of the regulation through abortion has yet to be discovered, the specific mechanism of abortion under water stress appears to be different from that under optimal conditions. Our results demonstrate that not only is the character architecture in wild barley plastic and sensitive to changing availability of water and nutrients, but the regulating mechanism of maternal investment is also environmentally sensitive.     

Experimental evidence of early costs of reproduction in conspecific viviparous and oviparous lizards

Reproduction entails costs, and disentangling the relative importance of each stage of the reproductive cycle may be important to assess the costs and benefits of different reproductive strategies. We studied the early costs of reproduction in oviparous and viviparous lizard females of the bimodal reproductive species Zootoca vivipara. Egg retention time in oviparous females is approximately one-third of the time in viviparous females. We compared the vitellogenesis and egg retention stages that are common to both reproductive modes. Precisely, we monitored the thermoregulatory behaviour, the weight gain and the immunocompetence of the females. Moreover, we injected an antigen in half of the females (immune challenge) to study the trade-offs between reproductive mode and immune performance and between different components of the immune system. Finally, we experimentally induced parturition in viviparous females at the time of egg laying in oviparous females. Oviparous and viviparous females did not show strong differences in response to the immune challenge. However, viviparous females spent more time thermoregulating while partially hidden and gained more weight than oviparous females. The greater weight gain indicates that the initial period of egg retention is less costly for viviparous than for oviparous females or that viviparous females are able to save and accumulate energy at this period. This energy may be used by viviparous females to cope with the subsequent costs of the last two-third of the gestation. Such an ability to compensate the higher costs of a longer egg retention period may account for the frequent evolution of viviparity in squamate reptiles.     

The effects of reproduction on courtship, fertility and longevity within and between alternative male mating tactics of the horned beetle, Onthophagus binodis

Life history theory provides a powerful tool to study an organism's biology within an evolutionary framework. The notion that males face a longevity cost of competing for and displaying to females lies at the core of sexual selection theory. Likewise, recent game theory models of the evolution of ejaculation strategies assume that males face a trade-off between expenditure on the ejaculate and expenditure on gaining additional matings. Males of the dung beetle Onthophagus binodis adopt alternative reproductive tactics in which major males fight for and help provision females, and minor males sneak copulations with females that are guarded by major males. Minor males are always subject to sperm competition, and consistent with theoretical expectation, minor males have a greater expenditure on their ejaculate than major males. We used this model system to seek evidence that mating comes at a cost for future fertility and/or male expenditure on courtship and attractiveness, and to establish whether these traits vary between alternative mating tactics. We monitored the lifespan of males exposed to females and nonmating populations, and sampled males throughout their lives to assess their fertility and courtship behaviour. We found a significant longevity cost of reproduction, but no fertility cost. On average, males from mating populations had a lower courtship rate than those from nonmating populations. This small effect, although statistically nonsignificant, was associated with significant increases in the time males required to achieve mating. Minor males had lower courtship rates than major males, and took longer to achieve mating. Although we did not measure ejaculate expenditure in this study, the correlation between lower courtship rate and longer mating speed of minor males documented here with their greater expenditure on the ejaculate found in previous studies, is consistent with game theory models of ejaculate expenditure which assume that males trade expenditure on gaining matings for expenditure on gaining fertilizations.     

One big, and many small reasons that direct selection on offspring number is still open for discussion

In a recent paper, I proposed that natural selection should act to increase offspring number when diversification bet hedging is favoured. The simple underlying reasoning is that a target diversification strategy is more reliably generated with increasing sample size. The intention of opening a discussion has been realized; recent criticisms of the idea argue that selection does not act to increase offspring number when population size is large or infinite. Here I agree that criticisms have merit; indeed they are largely confined to the caveats discussed in my original paper. The critique, however, implies a verdict of outright rejection of the idea of selection on offspring number, which would be erroneous. Contrary to the assertions of the criticism, then, the importance of selection acting directly on offspring number remains an open question.     

Parent–offspring conflict and selection on egg size in turtles

The trade‐off between offspring size and number can present a conflict between parents and their offspring. Because egg size is constrained by clutch size, the optimal egg size for offspring fitness may not always be equivalent to that which maximizes parental fitness. We evaluated selection on egg size in three turtle species (Apalone mutica, Chelydra serpentina and Chrysemys picta) to determine if optimal egg sizes differ between offspring and their mothers. Although hatching success was generally greater for larger eggs, the strength and form of selection varied. In most cases, the egg size that maximized offspring fitness was greater than that which maximized maternal fitness. Consistent with optimality theory, mean egg sizes in the populations were more similar to the egg sizes that maximized maternal fitness, rather than offspring fitness. These results provide evidence that selection has maximized maternal fitness to achieve an optimal balance between egg size and number.     

Horn growth rate and longevity: implications for natural and artificial selection in thinhorn sheep (Ovis dalli)

We used horn measurements from natural and hunted mortalities of male thinhorn sheep Ovis dalli from Yukon Territory, Canada, to examine the relationship between rapid growth early in life and longevity. We found that rapid growth was associated with reduced longevity for sheep aged 5 years and older for both the hunted and natural mortality data sets. The negative relationship between growth rate and longevity in hunted sheep can at least partially be explained by morphologically biased hunting regulations. The same trend was evident from natural mortalities from populations that were not hunted or underwent very limited hunting, suggesting a naturally imposed mortality cost directly or indirectly associated with rapid growth. Age and growth rate were both positively associated with horn size at death for both data sets, however of the two growth rate appeared to be a better predictor. Large horn size can be achieved both by individuals that grow horns rapidly and by those that have greater longevity, and the trade-off between growth rate and longevity could limit horn size evolution in this species. The similarity in the relationship between growth rate and longevity for hunted and natural mortalities suggests that horn growth rate should not respond to artificial selection. Our study highlights the need for the existence and study of protected populations to properly assess the impacts of selective harvesting.     

Effects of aging and reproduction on protein quality control in soma and gametes of Drosophila melanogaster

In organisms with a soma-germ demarcation, the germline must be 'preserved' such that harmful damage is not transmitted to the offspring. Keeping the progeny free of damage may be achieved by gametes enjoying elevated, and/or more functional, homeostatic maintenance systems. This possibility was approached here by testing whether the soma and maturating oocytes (eggs) dissected from female Drosophila melanogaster in reproductive ages display differential capacities for protein quality control and whether these capacities change during aging and mating. Eggs exhibited a high capacity to prevent protein aggregation, strong capacity for 26S proteasome-dependent degradation and reduced levels of oxidatively damaged (carbonylated) proteins compared to the soma. The capacity to prevent protein aggregation was not affected in either soma or eggs by age and/or mating, while the 26S proteasome capacity declined in the soma but was maintained in the eggs of aged females. However, the levels of carbonylated proteins increased with age in both soma and eggs, and this increase was more pronounced in females allowed to mate continuously. Furthermore, the levels of carbonylated proteins in the eggs of mated flies correlated negatively with the propensity of the eggs to develop into an adult fly. In young flies, mating caused a decrease in 26S proteasome capacity and an increase in protein carbonylation in the soma, but not in the eggs. These results are in line with trade-off theories of aging where aging is considered a consequence of investment in reproduction over somatic maintenance.     

Effects of organ removal on trade‐offs between sexual and clonal reproduction in the stoloniferous herb Duchesnea indica

Resource allocation patterns and trade‐off between sexual and clonal reproduction in clonal plants have been extensively studied, but little is known about effects of organ removal on the trade‐offs in clonal plants. To examine the effects, we conducted an experiment with the stoloniferous herb Duchesnea indica in which we removed plant organs like roots, flowers, or fruits. Removing roots significantly increased number of ramets and biomass allocation to stolons, but decreased number of fruits. Removing flowers or fruits greatly increased number of ramets and biomass allocation to stolons and roots, but decreased spacer length, number of fruits, and fruits set. Onset and median date of flowering phenology of D. indica shifted after flowers, fruits, or roots were removed. These results may indicate that removing organs can affect trade‐off between sexual and clonal reproduction of D. indica.     

Life History theory hypotheses on child growth: Potential implications for short and long‐term child growth,development and health

Life history theory integrates ecological, physiological, and molecular layers within an evolutionary framework to understand organisms’ strategies to optimize survival and reproduction. Two life history hypotheses and their implications for child growth, development, and health (illustrated in the South African context) are reviewed here. One hypothesis suggests that there is an energy trade‐off between linear growth and brain growth. Undernutrition in infancy and childhood may trigger adaptive physiological mechanisms prioritizing the brain at the expense of body growth. Another hypothesis is that the period from conception to infancy is a critical window of developmental plasticity of linear growth, the duration of which may vary between and within populations. The transition from infancy to childhood may mark the end of a critical window of opportunity for improving child growth. Both hypotheses emphasize the developmental plasticity of linear growth and the potential determinants of growth variability (including the role of parent–offspring conflict in maternal resources allocation). Implications of these hypotheses in populations with high burdens of undernutrition and infections are discussed. In South Africa, HIV/AIDS during pregnancy (associated with adverse birth outcomes, short duration of breastfeeding, and social consequences) may lead to a shortened window of developmental plasticity of growth. Furthermore, undernutrition and infectious diseases in children living in South Africa, a country undergoing a rapid nutrition transition, may have adverse consequences on individuals’ cognitive abilities and risks of cardio‐metabolic diseases. Studies are needed to identify physiological mechanisms underlying energy allocation between biological functions and their potential impacts on health.     

The ecological advantage of sexual reproduction in multicellular long-lived organisms

We present a model for the advantage of sexual reproduction in multicellular long-lived species in a world of structured resources in short supply. The model combines features of the Tangled Bank and the Red Queen hypothesis of sexual reproduction and is of broad applicability. The model is ecologically explicit with the dynamics of resources and consumers being modelled by differential equations. The life history of consumers is shaped by body mass-dependent rates as implemented in the metabolic theory of ecology. We find that over a broad range of parameters, sexual reproduction wins despite the two-fold cost of producing males, due to the advantage of producing offspring that can exploit underutilized resources. The advantage is largest when maturation and production of offspring set in before the resources of the parents become depleted, but not too early, due to the cost of producing males. The model thus leads to the dominance of sexual reproduction in multicellular animals living in complex environments, with resource availability being the most important factor affecting survival and reproduction.     

The effects of rat selection and reproduction on learning ability in the offspring

We studied the effect of selective reproduction on rats purchased from a breeder, so as to detect learning impairments in their offspring. The parent rats selected for mating were categorized as bright or dull in terms of learning ability on the basis of a learning test in a water maze or passive avoidance response (PAR). Offspring were tested in the same way as the selected rats, with their responses being compared for each generation. 1. The matings, consisting of bright X bright, bright X dull, dull X bright, dull X dull, resulted in a significant increase in the number of dull offspring of dull X dull from parents selected by water maze or PAR learning test. 2. Bright or dull mothers selected by PAR learning were treated with chlorpromazine 6 mg/kg (2 times/day/s.c.) on the 13th day of gestation. Afterwards, a slight increase in dull offspring of dull parents was seen. 3. These results indicate that in order to evaluate the learning ability of F1 in a reproductive test, it is first necessary to check the reaction of the parents, purchased F0, and to classify them as bright or dull.     

The genetics of primary and secondary sexual character trade-offs in a horned beetle

When structures compete for shared resources, this may lead to acquisition and allocation trade-offs so that the enlargement of one structure occurs at the expense of another. Among the studies of morphological trade-offs, their importance has been demonstrated primarily through experimental manipulations and comparative analyses. Relatively, a few studies have investigated the underlying genetic basis of phenotypic patterns. Here, we use a half-sibling breeding design to determine the genetic underpinnings of the phenotypic trade-off between head horns and the male copulatory organ or aedeagus that has been found in the dung beetle Onthophagus taurus. Instead of the predicted negative genetic covariance among characters that trade-off, we find positive genetic covariance between absolute horn and aedeagus length and zero genetic covariance between relative horn and aedeagus length. Therefore, although the genetic covariance between absolute horn and aedeagus length would constrain the independent evolution of primary and secondary sexual characters in this population, there was no evidence of a trade-off. We discuss alternative hypotheses for the observed patterns of genetic correlation between traits that compete for resources and the implications that these have for selection and the evolution of such traits.     

Effects of early resource limitation and compensatory growth on lifetime fitness in the ladybird beetle (Harmonia axyridis)

Acceleration of growth following a period of diet restriction may result in either complete or partial catch-up in size. The existence of such compensatory growth indicates that organisms commonly grow at rates below their physiological maxima and this implies a cost for accelerated growth. We examined patterns of accelerated growth in response to temporary resource limitation, and assayed both short and long-term costs of this growth in the ladybird beetle Harmonia axyridis. Subsequent to the period of food restriction, accelerated growth resulted in complete compensation for body sizes, although we observed greater larval mortality during the period of compensation. There were no effects on female fecundity or survivorship within 3 months of maturation. Females did not discriminate against males that had undergone compensatory growth, nor did we observe effects on male mating behaviour. However, individuals that underwent compensatory growth died significantly sooner when deprived of food late in adult life, suggesting that longer-term costs of compensatory growth may be quite mild and detectable only under stressful conditions.     

Reproductive ageing and sexual selection on male body size in a wild population of antler flies (Protopiophila litigata)

Little is known about the importance of trade-offs between ageing and other life history traits, or the effects of ageing on sexual selection, particularly in wild populations suffering high extrinsic mortality rates. Life history theory suggests that trade-offs between reproduction and somatic maintenance may constrain individuals with higher initial reproductive rates to deteriorate more rapidly, resulting in reduced sexual selection strength. However, this trade-off may be masked by increased condition dependence of reproductive effort in older individuals. We tested for this trade-off in males in a wild population of antler flies (Protopiophila litigata). High mating rate was associated with reduced longevity, as a result of increased short-term mortality risk or accelerated ageing in traits affecting viability. In contrast, large body size was associated with accelerated ageing in traits affecting mating success, resulting in reduced sexual selection for large body size. Thus, ageing can affect sexual selection and evolution in wild populations.     

Delayed effects of larval predation risk and food quality on anuran juvenile performance

Metamorphosis can disrupt the correlation structure between juvenile and adult traits, thus allowing relatively independent evolution in contrasting environments. We used a multiple experimental approach to investigate how diet quality and larval predation risk affected the rates of growth and development in painted frogs (Discoglossus galganoi), and how these changes influence post-metamorphic performance. A high-energy diet entailed growth advantages only if predation risk did not constrain energy acquisition, whereas diet quality affected primarily the extension of the larval period. Predation risk influenced juvenile shape, most likely via the effects on growth and differentiation rates. Juvenile frogs emerging from predator environments had shorter legs and longer bodies than those from the nonpredator tanks. However, these morphological changes did not translate into differences in relative jumping performance. Neither size-adjusted lipid storage nor fluctuating asymmetry was significantly influenced by food quality or predation risk. Our data suggest that the post-metamorphic costs of predator avoidance during the larval phase are mostly a consequence of small size at metamorphosis.     

Evolution of sex-biased maternal effects in birds: II. Contrasting sex-specific oocyte clustering in native and recently established populations

In species that produce broods of multiple offspring, parents need to partition resources among simultaneously growing neonates that often differ in growth requirements. In birds, multiple ovarian follicles develop inside the female at the same time, resulting in a trade-off of resources among them and potentially limiting maternal ability for sex-specific allocation. We compared resource acquisition among oocytes in relation to their future sex and ovulation order in two populations of house finches with contrasting sex-biased maternal strategies. In a native Arizona population, where mothers do not bias offspring sex in relation to ovulation order, the male and female oocytes did not show sex-specific trade-offs of resources during growth and there was no evidence for spatial or temporal segregation of male and female oocytes in the ovary. In contrast, in a recently established Montana population where mothers strongly bias offspring sex in relation to ovulation order, we found evidence for both intra-sexual trade-offs among male and female oocytes and sex-specific clustering of oocytes in the ovary. We discuss the importance of sex-specific resource competition among offspring for the evolution of sex-ratio adjustment and sex-specific maternal resource allocation.