Plasticity Of Sex Differentiation Research Articles

Conservative seed fertility maintains hermaphrodites in gynodioecious Daphne jezoensis: a test of the sex-differential plasticity hypothesis.

Gynodioecy is assumed to be an evolutionary transition from hermaphroditism to dioecy. However, if hermaphrodites can better flexibly regulate seed production depending on resource availability than females, i.e. sex-differential plasticity (SDP), gynodioecy can be a stable state. In the gynodioecious shrub Daphne jezoensis, hermaphrodites generally exhibit low seed fertility and largely act as males. We examined the SDP hypothesis and the cost of fruit production to clarify why D. jezoensis did not evolve into unisexual morphs. We evaluated the size and resource dependency of reproduction in field experiments by manipulating soil nutrient and light conditions. We compared the plant size and pollen production among females, fruiting hermaphrodites and non-fruiting hermaphrodites. We then analysed the effect of current fruit production on subsequent flower production, i.e. the cost of fruit production. The fruiting ability was independent of plant size and resource availability in both sexual phenotypes, indicating the absence of SDP in D. jezoensis. Hermaphrodites produced larger-sized pollen and allocated more resources to pollen production in the non-fruiting year than in the fruiting year. In contrast, the cost of fruit production was not revealed for either sexual phenotype, even in the absence of pollen limitation, and even when fruit production was maximized. SDP could not explain the maintenance of hermaphrodites in D. jezoensis. Alternatively, the lower cost of fruit production in hermaphrodites due to their potentially low fruiting ability may hinder the evolutionary shift to dioecy.

Absence of sex differential plasticity to light availability during seed maturation in Geranium sylvaticum.

Sex-differential plasticity (SDP) hypothesis suggests that since hermaphrodites gain fitness through both pollen and seed production they may have evolved a higher degree of plasticity in their reproductive strategy compared to females which achieve fitness only through seed production. SDP may explain the difference in seed production observed between sexes in gynodioecious species in response to resource (nutrients or water) availability. In harsh environments, hermaphrodites decrease seed production whereas females keep it relatively similar regardless of the environmental conditions. Light availability can be also a limiting resource and thus could theoretically affect differently female and hermaphrodite seed output even though this ecological factor has been largely overlooked. We tested whether the two sexes in the gynodioecious species Geranium sylvaticum differ in their tolerance to light limitation during seed maturation in the field. We used a fully factorial block experiment exposing female and hermaphrodite plants to two different light environments (control and shade) after their peak flowering period. Specifically, we measured fruit and seed production in response to decreased light availability and compared it between the sexes. Shading reduced the number of fruits and seeds produced, but the decrease was similar between the sexes. Furthermore, shading delayed seed production by three days in both sexes, but did not affect seed mass, seed P content, or the probability of re-flowering the following year. Our results give no evidence for reproductive SDP in response to light during seed maturation.

Gender plasticity and sexual system stability in Wurmbea

Sexually dimorphic populations are often located in drier habitats than cosexual populations. Gender plasticity (GP), whereby hermaphrodites alter female and male functions depending on resources, and sex-differential plasticity (SDP) between hermaphrodites and unisexuals are predicted to affect sexual system stability. Here, GP and SDP are evaluated in cosexual and gynodioecious Wurmbea biglandulosa and sub-dioecious and dioecious W. dioica. GP was evaluated under two resource conditions, compared among sexual systems and assessed as to whether (1) males produced perfect flowers and (2) hermaphrodites altered investment in perfect (female function) and total (male function) flowers. SDP was assessed within sexual systems as differences between sex functions of hermaphrodites vs. unisexuals. Males and hermaphrodites were compared to assess whether size thresholds for female function differed among sexual systems. Plasticity costs were evaluated using correlations between female function and male traits in hermaphrodites, and in W. dioica by comparing hermaphrodite and male regressions between plant size and pollen production. In dioecious W. dioica no males exhibited GP, whereas 100 % did in gynodioecious and cosexual W. biglandulosa. In sub-dioecious W. dioica, resources affected GP (high, 66 %; low, 42 %). Hermaphrodites in all sexual systems reduced perfect but not total flowers under low resources. Unisexuals were unaffected, demonstrating SDP for female function only. Thresholds for female function were greater in sub-dioecious W. dioica than in W. biglandulosa. Plasticity costs were detected only in sub-dioecious W. dioica. SDP for female function could assist female establishment in cosexual populations and maintain females in gynodioecious and sub-dioecious populations. Although the absence of male SDP should stabilize sub-dioecy, plasticity costs would render sub-dioecy unstable, favouring canalized males over hermaphrodites. This study highlights the importance of interactions between environmental conditions and hermaphrodite sex expression for the stability of dimorphic sexual systems.

Sex ratio and subdioecy inFragaria virginiana: the roles of plasticity and gene flow examined

Here we examined the roles of sex-differential plasticity (SDP) and gene flow in sex ratio evolution of subdioecious Fragaria virginiana. We assessed whether female frequency varied with resource availability in 17 natural populations and then characterized plasticity and mean investment in allocation to female function at flower and plant levels in the sex morphs in the glasshouse. We estimated patterns of population divergence using five microsatellite markers. We reveal SDP in fruit production substantial enough to translate into a higher equilibrium female frequency at low resources. Thus SDP can account, in part, for the strong negative relationship between female frequency and resources found in the field. Pollen-bearing morphs varied in plasticity across populations, and the degree of plasticity in fruit number was positively correlated with in situ variation in nitrogen (N) availability, suggesting an adaptive component to sex-allocation plasticity. Low neutral genetic differentiation, indicating high gene flow or recent divergence, may contribute to the absence of population differentiation in fruit-setting ability of pollen-bearing morphs despite considerable sex ratio variation. We consider how these processes, in addition to other features of this system, may work in concert to influence sex ratios and to hinder the evolution of dioecy in F. virginiana.

Darwinism renewed: contemporary studies of plant adaptation

Darwinism renewed: contemporary studies of plant adaptation

Effects of environmental heterogeneity on the distribution of sexes within and among populations in a gynodioecious species, Geranium maculatum

Populations containing both females and hermaphrodites (dimorphic) are generally found in drier sites than those with only hermaphrodites (monomorphic). The sex-differential plasticity hypothesis (SDP) suggests that this is caused by hermaphrodites reducing allocation to seeds in harsh environments, allowing female establishment. We proposed that a similar process could explain sex distribution within populations. We compared light availability and soil moisture between sites of three monomorphic and three dimorphic populations of Geranium maculatum and between microsites occupied by females and hermaphrodites within populations. We also correlated seed production in dimorphic populations with environmental measures. We found that dimorphic and monomorphic populations occurred in sites with similar soil moisture but within two dimorphic populations females occurred in drier microsites than hermaphrodites, as predicted by the SDP hypothesis. Contrary to the predictions, hermaphrodites' seed production was not influenced by the environment. Rather, females' seed production was correlated with environmental conditions in two populations, although the direction of the correlation differed between populations. Our results suggest that in this species, the SDP hypothesis does not explain sex distribution among or within populations. However, microsite environments may influence the distribution of sexes within a population and potentially aid in maintaining gynodioecy.

Reproductive functions of wild fish as bioindicators of reproductive toxicants in the aquatic environment

Impacts on the reproductive health of wild fish are thought to be suitable early-warning tools indicating contamination of surface waters with endocrine-disrupting compounds. Ecotoxicological assessment of these field observations depends on the availability of reliable biomarkers to enable a discrimination of natural variations of reproductive functions from anthropogenic impacts. Roach and perch were caught at eight sampling sites by electrofishing twice a year in summer (July-September) and late autumn/winter (November-December) over a 2-year period. The sites are characterized by different degrees of anthropogenic impact and are situated within the greater Upper Rhine catchment. Age growths, parasitization and gonadal histology of more than 3,000 fish were examined. The two dominant fish species in German surface waters perch (Perca fluviatilis L.) and roach (Rutilus rutilus L.) differ considerably regarding their suitability for biomonitoring. Even in pristine habitats, perch show several variants of sex differentiation in terms of (1) the time of first sexual maturation, (2) the course of seasonal gonadal recrudescence, and (3) the occurrence of heterologous germ cells (testes ova). A statistically significant elevated proportion of males were observed in fish obtained from a TBT-contaminated marina and suppression of gonadal ripening was observed in females caught in a sewage-contaminated brook. Both effects appear to be due to chemical contamination. The only "natural" alteration of sex differentiation in roach was related to parasitization with Ligula intestinalis (Eucestoda, Pseudophyllidea). Other deviations from the normal pattern of sex differentiation were (1) suppression of ovarian ripening and (2) asynchronic seasonal gonadal recrudescence. These are strong indicators of an anthropogenically induced impact on reproductive health. Feminization phenomena were not observed at either the individual or the population level. Interpretation of field monitoring results concerning reproductive health requires large numbers of samples and detailed knowledge of the natural plasticity of sex differentiation in the species under investigation. A better understanding of the mechanisms underlying the plasticity of sex differentiation in perch is indispensable to enable perch to be used as a bioindicator. Deviation from the strict and probably endogenous control of sex differentiation in roach is a strong and unequivocal warning signal. The subject of fish monitoring should be addressed in the context of a broader spectrum of potential risks. Seasonal and ontogenetic integrity of gonadal development and recrudescence are potent biomarkers, provided the natural process is well documented for the species under investigation.

PHENOTYPIC PLASTICITY OF HERMAPHRODITE SEX ALLOCATION PROMOTES THE EVOLUTION OF SEPARATE SEXES: AN EXPERIMENTAL TEST OF THE SEX-DIFFERENTIAL PLASTICITY HYPOTHESIS USINGSAGITTARIA LATIFOLIA(ALISMATACEAE)

Separate sexes can evolve under nuclear inheritance when unisexuals have more than twice the reproductive fitness of hermaphrodites through one sex function (e.g., when females have more than twice the seed fertility of hermaphrodites). Because separate sexes are thought to evolve most commonly via a gynodioecious intermediate (i.e., populations in which females and hermaphrodites cooccur), the conditions under which females can become established in populations of hermaphrodites are of considerable interest. It has been proposed that resource-poor conditions could promote the establishment of females if hermaphrodites are plastic in their sex allocation and allocate fewer resources to seed production under these conditions. If this occurs, the seed fertility of females could exceed the doubling required for the evolution of unisexuality under low-, but not high-resource conditions (the sex-differential plasticity hypothesis). We tested this hypothesis using replicate experimental arrays of the aquatic herb Sagittaria latifolia grown under two fertilizer treatments. The results supported the sex-differential plasticity hypothesis, with females having more than twice the seed fertility of hermaphrodites under low-, but not high-fertilizer conditions. Our findings are consistent with the idea that separate sexes are more likely to evolve under unfavorable conditions.