Investment In Female Function Research Articles

Hermaphrodites and parasitism: size-specific female reproduction drives infection by an ephemeral parasitic castrator

Sex can influence patterns of parasitism because males and females can differ in encounter with, and susceptibility to, parasites. We investigate an isopod parasite (Hemioniscus balani) that consumes ovarian fluid, blocking female function of its barnacle host, a simultaneous hermaphrodite. As a hermaphrodite, sex is fluid, and individuals may allocate energy differentially to male versus female reproduction. We predicted the relationship between barnacle size and female reproductive function influences the distribution of parasites within barnacle populations. We surveyed 12 populations spanning ~400 km of coastline of southern California and found intermediate-sized barnacles where most likely to be actively functioning as females. While it is unclear why larger individuals are less likely to be actively reproducing as females, we suggest this reduced likelihood is driven by increased investment in male reproductive effort at larger sizes. The female function-size relationship was mirrored by the relationship between size and parasitism. We suggest parasitism by Hemioniscus balani imposes a cost to female function, reinforcing the lack of investment in female function by the largest individuals. Within the subset of suitable (=female) hosts, infection probability increased with size. Hence, the distribution of female function, combined with selection for larger hosts, primarily dictated patterns of infection.

Effect of water availability and genetic diversity on flowering phenology, synchrony and reproductive investment in summer squash

SUMMARYCurrent agricultural practices rely on crops with developmental phenologies adapted to local climate, photoperiods and soils; however, global climate change will alter some abiotic factors (e.g. temperature and precipitation). Previously adapted varieties may be poorly prepared for these changing conditions, if such conditions induce mismatched phenologies. Crops that depend on cross-pollination and synchronous flowering may be most susceptible, e.g. monoecious plants have separate male and female flowers, and changes in flowering synchrony may alter yield. Using genetically diverse (open-pollinated (OP)) and genetically homogeneous (hybrid) varieties of a monoecious crop, courgette, also known as zuchinni (Cucurbita pepo), phenological responses to experimentally manipulated moisture conditions were explored in an agricultural context. Under drier and wetter conditions, the hybrid courgette plants shifted towards a male-biased floral sex ratio due to the reduced production of female flowers. However, flowering synchrony and fruit production were unaffected by moisture treatment in both varieties. The hybrid and OP varieties differed in many traits related to floral sex ratios, phenology, synchrony and fruit production. Further, the OP variety displayed more phenotypic variation than the hybrid in many traits. Being in a population context rather than relying on self-pollination increased the availability of potential mates for a given female flower in both the hybrid and, particularly, the OP variety. Thus, the increased genetic diversity found in OP v. hybrid varieties may buffer the possible environmental effects on flowering synchrony within a cropping context. Finally, the likelihood of female flowers setting fruit increased with the number of male flowers within a population, and the rate of increase was higher in the hybrid variety. In summary, climate change is predicted to reduce investment in female function in some monoecious crops and genetically diverse varieties may play an important role in maintaining reproductive synchrony in altered environments.

Trade‐offs between clonal and sexual reproduction in Sagittaria latifolia (Alismataceae) scale up to affect the fitness of entire clones

Many plants combine sexual reproduction with vegetative propagation, but how trade-offs between these reproductive modes affect fitness is poorly understood. Although such trade-offs have been demonstrated at the level of individual shoots (ramets), there is little evidence that they scale up to affect genet fitness. For hermaphrodites, reproductive investment is further divided between female and male sexual functions. Female function should generally incur greater carbon costs than male function, which might involve greater nitrogen (N) costs. Using a common garden experiment with diclinous, clonal Sagittaria latifolia we manipulated investment in reproduction through female and male sex functions of 412 plants from monoecious and dioecious populations. We detected a 1:1 trade-off between biomass investment in female function and clonal reproduction. For male function, there was no apparent trade-off between clonal and sexual reproduction in terms of biomass investment. Instead, male function incurred a substantially higher N cost. Our results indicate that: trade-offs between investment in clonal propagation and sexual reproduction occur at the genet level in S. latifolia; and sexual reproduction interferes with clonal expansion, with investment in female function limiting the quantity of clonal propagules produced, and investment in male function limiting the nutrient content of clonal propagules.

FLOWERING PHENOLOGY AND SEXUAL ALLOCATION IN SINGLE-MUTATION LINEAGES OF ARABIDOPSIS THALIANA

We investigated the relationship between flowering time and sexual allocation in wild-type Arabidopsis thaliana and in genetically similar lineages with single-locus mutations of floral induction genes. We examined whether the mechanisms of growth and development that govern resource investment would permit the independent evolution of reproductive phenology and sexual allocation, or whether constraints, manifested as pleiotropic effects of the single mutations, would link these two life-history traits. Flowering times differed significantly among genotypes, and, as expected, later flowering times were associated with larger vegetative size. Later flowering genotypes produced heavier floral parts (larger petals, in particular), and allocated a significantly lower proportion of biomass to androecia, especially in final allocations that included fruit biomass. At least part of this pleiotropic covariation of flowering time and sexual allocation is likely to be mediated by vegetative size and the rate of resource supply to growing reproductive tissues, because the larger fruits of late-flowering genotypes required the same time, or proportionately less time than the difference in biomass, to mature. Because fruit mass is considered an investment in female function, sexual allocation measured at the end of a growing season tends to be highly female biased in angiosperms. We consider the implications of the pleiotropic association of flowering time, vegetative size, and sexual investment for the theory of sex allocation, and suggest that the idiosyncratic phenology of sexual investment in flowering plants creates a departure from a central assumption of Fisher's seminal sex allocation argument.

Sexual selection in a simultaneous hermaphrodite with hypodermic insemination: body size, allocation to sexual roles and paternity

Theory predicts that variation in body size within a population of simultaneous hermaphrodites should affect sex allocation, leading to individual differences in mating strategies and increased investment in female function with size. Small animals with fewer resources should invest proportionally more of their resources in male function than large animals, resulting in sperm displacement and paternity patterns that are independent of body size. This study investigated the effect of body size on mating patterns, egg production (an indirect measure of sperm transfer) and paternity in Alderia modesta, a sperm-storing, hermaphroditic sea slug with hypodermic insemination. The relative sizes of two hermaphrodites affected the probability and duration of inseminations; smaller animals inseminated larger mates for longer than vice versa. Sperm transfer began at a smaller size and age than egg production, and estimates of both sperm transfer and egg production increased with body size. Paternity patterns varied widely; in this species, unpredictable sperm precedence patterns may be a consequence of hypodermic insemination and the lack of a well-defined sperm storage organ. Hypodermic injections across all sections of the body successfully transferred sperm and fertilized eggs. The function and consequences of hypodermic insemination are discussed. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.

Multiple mating, paternity, and body size in a simultaneous hermaphrodite, Aplysia californica

Sperm displacement and sperm competition prove difficult to measure, but are crucial elements in predicting sex allocation strategies of sperm-storing hermaphrodites. Body size is predicted to affect sex allocation so that within a population, large animals invest a greater proportion of resources in female function than do small animals. These mating strategies depend on sperm displacement abilities and lead to similar levels of paternity across body sizes despite differences in resource level. The present study investigated mating patterns, multiple paternity, and sperm competition in a field population of a simultaneously hermaphroditic sea slug, Aplysia californica (California sea hare). Animals mating in the female role were larger than the mean for the population, indirectly supporting theoretical predictions for increased investment in female function with body size. However, contrary to predictions, animals mating in the male role were not different in size from the population mean or the animals they inseminated. Individual tagging revealed that sea slugs are capable of moving across distances that allow for the sampling of many potential mates, and that they mate repeatedly in both sexual roles. Microsatellite paternity analysis demonstrated that multiple mating in the field leads to multiple paternity, and last-sperm donors achieve high levels of paternity. There was no effect of body size on paternity. Further paternity studies are needed to reveal the mechanisms of sperm precedence patterns in A. californica. Copyright 2003.

Size‐dependent sex allocation within flowers of the annual herb Clarkia unguiculata (Onagraceae): ontogenetic and among‐plant variation

The relative allocation of resources to male and female functions may vary among flowers within and among individual plants for many reasons. Several theoretical models of sex allocation in plants predict a positive correlation between the resource status of a flower or individual and the proportion of reproductive resources allocated to female function. These models assume that, independent of resource status, a negative correlation exists between male and female investment. Focusing on the allocation of resources within flowers, we tested these theoretical predictions and this assumption using the annual Clarkia unguiculata (Onagraceae). We also sought preliminary evidence for a genetic component to these relationships. From 116 greenhouse-cultivated plants representing 30 field-collected maternal families, multiple flowers and fruits per plant were sampled for gamete production, pollen : ovule ratio, seed number, ovule abortion, seed biomass/fruit, mean individual seed mass, and petal area. If sex allocation changes as predicted, then (1) assuming that flowers produced early have access to more resources than those produced later, basal flowers should exhibit a higher absolute and proportional investment in female function than distal flowers and (2) plants of high resource status (large plants) should produce flowers with a higher proportional investment in female function than those of low resource status. Within plants, variation in floral traits conformed to the first prediction. Among plants and families, no significant effects of plant size (dry stem biomass) on intrafloral proportional sex allocation were observed. We detected no evidence for a negative genetic correlation between male and female investment per flower, even when controlling for plant size.

Variation in floral sex allocation and reproductive success within inflorescences of Corydalis ambigua (Fumariaceae): pollination efficiency or resource limitation?

Summary The variation of floral sex allocation with flower position within inflorescences was investigated in the spring ephemeral, Corydalis ambigua. Investment in female function (pistil), attraction (corolla) and nectar production decreased from bottom to top flowers, whereas male investment (stamen) did not differ. This self‐incompatible species appears to set seeds as a result of visitation by nectar robbing bumblebee queens. The tendency of bees to visit lower flowers first and then move upwards within an inflorescence should result in directional pollen flow from bottom to top flowers. Naturally pollinated upper flowers set fewer seeds than intermediate and lower flowers due to pollen limitation. The lack of differences in seed set and seed mass per pod following artificial outcrossing indicated that resource limitation did not explain the variation in seed production of flowers in different positions. Pollen viability also did not differ significantly between flower positions. A model of pollination was developed that incorporated the visitation pattern of bumblebees and observed variations in nectar distribution between flower positions. This predicted that receipt of outcross pollen would decrease from bottom to top flowers, but that pollen export to other plants would not differ between flower positions provided that the pollen exchange rate of pollinators was either small or positively correlated with nectar content of each flower position. The observed pattern of floral sex allocation would then be parallel to relative success of pollen export and import between flower positions within inflorescences.