Total Reproductive Allocation Research Articles

Influences of climatic and social environment on variable maternal allocation among offspring in Alpine marmots.

In an environment with limited resources, parents may trade-off the number of offspring produced against offspring mass. To maximize fitness under unpredictable environments, females must not only maximize mean annual reproductive success but also minimize between-year variation in reproductive success. Thus, preferred strategies of maternal allocation might be to maximize the mass of their offspring or to produce a number of offspring of variable body masses. Many social species have evolved in variable and unpredictable environments where only the social environment can be predicted. If mothers seem to alter their total reproductive allocation to offspring depending on their social environment, how the total expenditure is allocated between the different offspring is still unknown. Here, we analysed how climatic and social environments influence strategies of maternal allocation and how these strategies impact pup first-year survival in a wild population of Alpine marmots monitored between 1990 and 2016. We found that females acted as income breeders using resources immediately available for reproduction. Our results showed that the proportion of maternal mass allocated to offspring varied mainly with litter size. However, how maternal allocation is shared between pups depended on climatic and social environments. In general, mothers tended to have litters of greater average mass and small variability in favourable social environments or when resources are abundant and lighter average pup mass but high variability in unfavourable social environments or when resources are scarce. This variable allocation could correspond to dynamic bet-hedging such that mothers influence the variance of pup mass within the litter in response to poor current environmental conditions. Our analysis of first-year survival showed that females should maximize the body mass of their young whatever the conditions will be because pups of higher mass have higher survival, regardless of environmental conditions. When resources are scarce, this strategy might not be achievable for all pups so that mothers produced variable pups. In large litters, this strategy increased first-year survival. Because pup variability affects parental fitness, differential allocation between pups of the same litter could have large consequences on fitness and thus on reproductive strategies of social species.

Accessory costs of seed production

Accessory costs of reproduction are those that are necessary to mature a seed, but that do not involve the direct cost of provisioning the seed itself. This study aims to quantify accessory costs in a range of species, and test whether they decrease as a proportion of total reproductive expenditure with increasing seed mass, as might be expected if economies of scale came into play at larger seed sizes. We also test whether accessory costs varied with growth form, pollination mode, and dispersal mode, with the expectation that biotic pollination and dispersal modes should incur greater costs. Reproductive allocation (dry biomass) over one season, was calculated for 14 diclinous angiosperm species. Accessory costs averaged 73% of total reproductive allocation, with the majority spent on packaging and dispersal. Total accessory costs, packaging and dispersal costs, and costs incurred prior to pollination were proportional to direct costs of reproduction in major axis regressions. However, larger seeded species incurred significantly greater costs associated with aborted seeds and fruits, and matured a smaller proportion of ovules. This is consistent with larger seeded species being more selective of the ovules/embryos matured than small-seeded species. Total accessory costs, and proportion of ovules aborted, were also significantly greater for biotically dispersed species, but only due to an association with larger seed masses. Costs associated with abortions were lower for biotically pollinated species, due to a general trend of more ovules per ovary, resulting in greater cost sharing. This study demonstrates that expenditure on items other than seeds accounts for the majority of reproductive allocation in flowering plants. Yet, far more literature exists on seed mass variation than on investment in accessory structures. We found a proportional relationship between accessory costs and seed mass that warrants further investigation within the context of selection on margin returns on investment.

Does body-volume constrain reproductive output in lizards?

The numbers and sizes of eggs produced by adult females ultimately determine the viability of populations, as well as the evolutionary fitness of the females themselves. Despite an enormous amount of literature on the adaptive significance of fecundity variation within and among populations, simpler questions--such as the proximate mechanisms by which a female determines her clutch size--have attracted less attention. Our surgical manipulations show that the amount of space available to hold eggs within a female's abdomen influences her total reproductive allocation, enabling her to flexibly modify her reproductive output as she grows larger.

Resource Allocation and the Evolution of Self-Fertilization in Plants.

This article develops a simple evolutionarily stable strategy (ESS) model of resource allocation in partially selfing plants, which incorporates reproductive and sex allocation into a single framework. The analysis shows that, if female fitness gain increases linearly with resource investment, total reproductive allocation is not affected by sex allocation, defined as the fraction of reproductive resources allocated to male function. All else being equal, the ESS total reproductive allocation increases with increasing selfing rate if the fitness of selfed progeny is more than half that of outcrossed progeny, while the ESS sex allocation is always a decreasing function of the selfing rate. Self-fertilization is much more common in annual than in perennial plants, and this association has been commonly interpreted in terms of an effect of life history on mating system. The model in this article shows that self-fertilization can itself cause the evolution of the annual habit. Incorporating the effects of pollen discounting may not have any influence on total reproductive allocation if female fitness gain is a linear function of resource investment, although the evolutionarily stable sex allocation is altered. Evolution of the selfing rate is found to be independent of reproductive and sex allocation under the mass-action assumption that self- and outcross pollen are deposited simultaneously on receptive stigmas and compete for access to ovules.

Resource Allocation and the Evolution of Self-Fertilization in Plants

This article develops a simple evolutionarily stable strategy (ESS) model of resource allocation in partially selfing plants, which incorporates reproductive and sex allocation into a single framework. The analysis shows that, if female fitness gain increases linearly with resource investment, total reproductive allocation is not affected by sex allocation, defined as the fraction of reproductive resources allocated to male function. All else being equal, the ESS total reproductive allocation increases with increasing selfing rate if the fitness of selfed progeny is more than half that of outcrossed progeny, while the ESS sex allocation is always a decreasing function of the selfing rate. Self‐fertilization is much more common in annual than in perennial plants, and this association has been commonly interpreted in terms of an effect of life history on mating system. The model in this article shows that self‐fertilization can itself cause the evolution of the annual habit. Incorporating the effects of pollen discounting may not have any influence on total reproductive allocation if female fitness gain is a linear function of resource investment, although the evolutionarily stable sex allocation is altered. Evolution of the selfing rate is found to be independent of reproductive and sex allocation under the mass‐action assumption that self‐ and outcross pollen are deposited simultaneously on receptive stigmas and compete for access to ovules.

Life-History Consequences of Divergent Selection on Egg Size in Drosophila melanogaster.

Life histories are generally assumed to evolve via antagonistic pleiotropy (negative genetic correlations) among traits, and trade-offs between life-history traits are typically studied using either phenotypic manipulations or selection experiments. We investigated the trade-off between egg size and fecundity in Drosophila melanogaster by examining both the phenotypic and genetic relationships between these traits after artificial selection for large and small eggs, relative to female body size. Egg size responded strongly to selection in both directions, increasing in the large-egg selected lines and decreasing in the small-egg selected lines. Phenotypic correlations between egg size and fecundity in the large-egg selected lines were negative, but no relationship between these traits occurred in either the control or small-egg selected lines. There was no negative genetic correlation between egg size and fecundity. Total reproductive allocation decreased in the small-egg selected lines but did not increase in the large-egg lines. Our results have three implications. First, our selection procedure may have forced females selected for large eggs into a physiological trade-off not reflected in a negative genetic correlation between these traits. Second, the lack of a negative genetic correlation between egg size and number suggests that the phenotypic trade-off frequently observed between egg size and number in other organisms may not evolve over the short term via a direct genetic trade-off whereby increases in egg size are automatically accompanied by decreased fecundity. Finally, total reproductive allocation may not evolve independently of egg size as commonly assumed.

Colony-level single-sex alate production inPheidole ceres andPheidole pilifera (Formicidae: Myrmicinae)

Patterns of colony-level reproductive investment were estimated inPheidole ceres andP. pilifera. Both species exhibited strong degrees of colony-level, single-sex alate production, but differed in the extent to which colonies produced only one sex. The proportion of reproductive investment allocated to males was negatively correlated to the total reproductive allocation in both species.

SEX ALLOCATION IN THE MONOECIOUS HERB BEGONIA SEMIOVATA.

Sex-allocation models predict that the evolution of self-fertilization should result in a reduced allocation to male function and pollinator attraction in plants. The evolution of sex allocation may be constrained by both functional and genetic factors, however. We studied sex allocation and genetic variation for floral sex ratio and other reproductive traits in a Costa Rica population of the monoecious, highly selfing annual Begonia semiovata. Data on biomass of floral structures, flower sex ratios, and fruit set in the source population were used to calculate the average proportion of reproductive allocation invested in male function. Genetic variation and genetic correlations for floral sex ratio and for floral traits related to male and female function were estimated from the greenhouse-grown progeny of field-collected maternal families. The proportion of reproductive biomass invested in male function was low (0.34 at flowering, and 0.07 for total reproductive allocation). Significant among-family variation was detected in the size (mass) of individual male and female flowers, in the proportion of male flowers produced, and in the proportion of total flower mass invested in male flowers. Significant among-family variation was also found in flower number per inflorescence, petal length of male and female flowers, and petal number of female flowers. Except for female petal length, we found no difference in the mean value of these characters between selfed and outcrossed progeny, indicating that, with the possible exception of female petal length, the among-family variation detected was not the result of variation among families in the level of inbreeding. Significant positive phenotypic and broad-sense genetic correlations were detected between the mass of individual male and female flowers, between male and female petal length, and between number of male and number of female flowers per inflorescence. The ratio of stamen-to-pistil mass (0.33) was low compared to published data for autogamous species with hermaphroditic flowers, suggesting that highly efficient selfing mechanisms may evolve in monoecious species. Our results indicate that the study population harbors substantial genetic variation for reproductive characters. The positive genetic correlation between investment in male and female flowers may reflect selection for maximum pollination efficiency, because in this self-pollinating species, each female flower requires a neighboring male flower to provide pollen.

Seed size, pollination costs and angiosperm success

Seed plants capture pollen before seeds are dispersed and abort unpollinated ovules. As a result, each seed is associated with an accessory cost that represents the costs of pollen capture and the costs of aborted ovules. Accessory costs may explain the minimum seed size among species, because these costs are likely to comprise a greater proportion of total reproductive allocation in species with smaller seeds. For very small propagules, the costs of pollination may not be worth the benefits, perhaps explaining the persistence of pteridophytic reproduction at small propagule sizes. The smallest angiosperm seeds are much smaller than the smallest gymnosperm seeds, both in the fossil record and in the modern flora. This suggests that angiosperms can produce pollinated ovules more cheaply than gymnosperms. Pollination becomes less efficient as a species decreases in abundance, and this loss of efficiency is greater for species with a higher accessory cost per seed. We propose that the greater reproductive efficiency of angiosperms when rare can explain why angiosperm-dominated floras were more speciose than the gymnosperm-dominated floras they replaced.

Reproductive effort in cotton grass tussock tundra

Eriophorum vaginatum ssp. spissum is a dominant plant species of undisturbed cotton grass tussock tundra in Alaska. It also quickly invades and dominates recently disturbed sites. The hypothesis tested in this research was that the success of E. vaginatum on disturbed sites might be achieved through a higher allocation of biomass to reproductive structures relative to other tundra species. Reproductive allocation of tundra plants in general also was compared with plants of the temperate zone. The results indicate that E. vaginatum is about average among the common tundra species in terms of total reproductive allocation, allocation to seeds, and the proportion of total reproductive allocation that is accounted for by viable seeds. Tundra species, on a relative basis, allocate less biomass to all reproductive structures than temperate species but not necessarily less biomass to the output of viable seeds.