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Stinging nettle, Urtica dioica L., is a widely distributed subdioecious long‐rhizomatous plant. Within a single population growing in the territory of the Zvenigorod Biological Station of Lomonosov Moscow State University, eight plots were selected in stinging nettle thickets to study the variability of the sexual reproductive part in U. dioica depending on internal and external factors. As monoecious ramets are relatively common in the studied population, four gender forms were distinguished based on the ratio of male and female flowers. On 156 ramets of different gender forms and 36 genets, it was shown that predominantly male plants are shorter, while predominantly female plants have larger and more widely spread inflorescences compared to other gender forms. The pubescence type (spreading or appressed unicellular hairs) shows the greatest repeatability within the merigenet. True monoecy in stinging nettle is widespread: male and female flowers of different gender forms were found in 10 out of 36 genets. The most common inflorescence architecture types in the studied population were proxigynous and basigynous. Both normal pistillate and normal staminate flowers, as well as flowers with morphological abnormalities formed based on flowers of different sexes can be found on the same ramet, which suggests the presence of paradioecious lability. [ABSTRACT FROM AUTHOR]

RNA-sequencing was used to develop 16 microsatellite markers for the pearly everlasting, Anaphalis margaritacea var. yedoensis (Franch. et Sav.) Ohwi (Asteraceae), which inhabits gravel bars throughout the Japanese archipelago. The mean number of alleles for these 16 markers in two populations in the Hokkaido and Shizuoka Prefectures, was 3.5 and 4.0, respectively, while the mean expected heterozygosity was 0.525 and 0.560, respectively, with a significant genetic differentiation between the two populations. All markers could also be amplified in two conspecific taxa, A. margaritacea var. margaritacea and var. angustifolia, whereas 11 of the 16 markers were amplifiable in two congeneric species, A. sinica and A. alpicola. These newly developed microsatellite markers will support understanding of population genetics and mating systems in A. margaritacea var. yedoensis, and several will potentially be of use in similar studies in other Anaphalis species. [ABSTRACT FROM AUTHOR]

Idesia polycarpa is a dioecious tree; in field surveys, there are rare sex conversions in I. polycarpa individuals with bisexual flowers. To identify the molecular mechanisms underlying sex conversion in this species, an integrative analysis of the proteome and transcriptome profiles of I. polycarpa male and bisexual flowers at key developmental stages was conducted in this study using isobaric tags for relative and absolute quantification and RNA-seq technology. A total of 15,003 proteins were identified; the differentially expressed proteins (DEPs) were enriched in metabolic pathways, biosynthesis of secondary metabolites, and flavonoid metabolism pathways in all comparison groups. A total of 290,442 unigenes were obtained; these were compared with seven databases, revealing 196,366 annotated unigenes. In general, the expression of proteins and genes tended to be positively correlated, with Spearman correlation coefficients in the ranges of 0.152–0.262 (all genes and all proteins) and 0.497–0.778 (DEPs and DEGs). The integrative analysis of DEPs and DEGs between male and bisexual flowers revealed that the most significantly enriched pathways were flavonoid pathways, metabolic pathways, and the biosynthesis of secondary metabolites. Finally, four co-expressed proteins and transcripts and one gene associated with the flavonoid biosynthesis pathway were screened out. The proteins identified were p-coumaroyl shikimate 3′-hydroxylase, and shikimate/quinate hydroxycinnamoyl transferase, and the gene was caffeoyl-CoA O-methyltransferase. The analysis has revealed key potential proteins and genes involved in sex conversion at the molecular level and has provided a basis for future investigations of artificial regulation of sex differentiation in I. polycarpa. [ABSTRACT FROM AUTHOR]

When the sexual system of a rare plant species is complex, characterizing floral development, plant morphology, and subpopulation structure is essential for assessing reproductive potential. Availability of pollinator rewards and positioning of sexual morphs within inflorescences relates to outcrossing potential. We studied the complex sexual system of the recently discovered Euphorbia rosescens E.L. Bridges & Orzell, a rare species occurring in a single Florida county. Euphorbia rosescens has both monomorphic subpopulations of female plants and mixed subpopulations with male and andromonoecious plants. Most data on cyathia sex, inflorescence architecture, floral visitors, and fruit set came from one female subpopulation and one mixed subpopulation. Cyathia longevity, flowering phase, nectar secretion, pollen production, and viability were measured in female, male, and hermaphrodite cyathia. Female plants had fewer inflorescence levels and cyathia and significantly smaller cyathia than male and andromonoecious plants. Andromonoecious plants were larger and produced more cyathia relative to male plants. Within mixed populations, sex switching occurred in both directions between male and andromonoecious plants. Female plants did not switch sexes. Proximal to distal inflorescence levels transitioned from hermaphrodite to male cyathia on andromonoecious plants. Nectar secretion corresponded to initiation of staminate flowering, but in female cyathia stigmas were receptive approximately a day before nectar secretion. Pollen production and viability were similar in male and hermaphrodite cyathia. Fruit maturation was minimal, primarily observed on andromonoecious plants. Nectar availability, pollen rewards, trimorphic inequalities in inflorescence display, and low fruit set suggest high geitonogamy, thus limiting outcrossing potential for this species. [ABSTRACT FROM AUTHOR]

Revealing the environmental pressures determining the frequency of females amongst populations of sexually dimorphic plants is a key research question. Analyses of sex ratio variation have been mainly done in dioecious plants, which misses key plant sexual systems that might represent intermediate stages in the evolution of dioecy from hermaphroditism. • We investigated female frequency across populations of sexually dimorphic plant species in relation to environmental stressors (temperature, precipitation), totaling 342 species, 2011 populations, representing 40 orders and three different sexual systems (dioecy, gynodioecy and subdioecy). We also included the biome where the population was located to test how female frequency may vary more broadly with climate conditions. • After correcting for phylogeny, our results for gynodioecious systems showed a positive relationship between female frequency and increased environmental stress, with the main effects being temperature-related. Subdioecious systems also showed strong positive relationships with temperature, and positive and negative relationships related to precipitation, while no significant effects on sex ratio in dioecious plants were detected. • Combined, we show that female frequencies in an intermediate sexual system on the pathway from hermaphroditism to dioecy respond strongly to environmental stressors and have different selective agents driving female frequency. [ABSTRACT FROM AUTHOR]

Key message: Cosexual Populus ×canescens plants are inconstant females with life course plasticity of sex phenotype and can reproduce by selfing. Populus species are dioecious, but deviations from dioecy are reported in some cases. The objectives of this study were to investigate the phenotypic expression and the inheritance of subdioecy in a Populus ×canescens pedigree. The F1 progeny was monitored for sex during 14 years. Thirty per cent of individuals expressed deviations from dioecy and long-term plasticity of sex. Some plants started flowering as male, then became cosexual, and finally turned female. Two cosexual individuals were self-pollinated and generated a selfed progeny markedly impaired by inbreeding depression, but able to reproduce by outcrossing. Sex segregation of the F1 progeny statistically fitted the expected ratio 1:2:1 (female:male:cosexual). By analysis of DNA markers, the cosexual individuals were genetically clustered with the females. The segregation ratio and the genetic profile indicated that cosexual plants were female with altered sex phenotype. Linkage analysis identified a putative sex-determining region with suppressed recombination on chromosome 19 of the male Populus tremula parent. The male sex trait was linked to the pericentromeric region of the P. tremula chromosome 19, whereas the cosexual trait was linked to chromosome 19 of the female Populus alba parent. A genetic model is proposed to explain inheritance and phenotypic expression of sex. [ABSTRACT FROM AUTHOR]

In trioecious plant populations, the role of hermaphroditism is often uncertain. We investigated the advantages of hermaphroditism in the dioecious shrub Salix myrsinifolia. The sex ratio of 30 S. myrsinifolia populations in northeastern Poland (secondary range) and Lithuania (primary range) was investigated in 2010–2011. Measures of reproductive (number of catkins, number of flowers in catkins) and vegetative traits (height, diameter, number of shoots, vitality) were taken and compared among sexual morphs. In two populations, measurements collected 14 years prior on marked individuals were used to determine the rate of changes in height, crown diameter and survivorship rates. We found trioecy mostly in the secondary part of the range with an average share of hermaphrodites reaching 21% in the trioecious populations. The sex ratio varied between populations, but tended mostly towards female domination. Several traits differed significantly among sexes. The characteristics of hermaphrodites were often intermediate between males and females and provided no evidence for the higher competitive abilities of hermaphrodites. We concluded that the possible gain of hermaphroditism in the colonization process is restricted to reproduction. We consider hermaphroditism in S. myrsinifolia as an equilibrium between the allocation of resources for growth and reproduction in unstable conditions on the margins of the range.

RNA-sequencing was used to develop 16 microsatellite markers for the pearly everlasting, Anaphalis margaritacea var. yedoensis (Franch. et Sav.) Ohwi (Asteraceae), which inhabits gravel bars throughout the Japanese archipelago. The mean number of alleles for these 16 markers in two populations in the Hokkaido and Shizuoka Prefectures, was 3.5 and 4.0, respectively, while the mean expected heterozygosity was 0.525 and 0.560, respectively, with a significant genetic differentiation between the two populations. All markers could also be amplified in two conspecific taxa, A. margaritacea var. margaritacea and var. angustifolia, whereas 11 of the 16 markers were amplifiable in two congeneric species, A. sinica and A. alpicola. These newly developed microsatellite markers will support understanding of population genetics and mating systems in A. margaritacea var. yedoensis, and several will potentially be of use in similar studies in other Anaphalis species.

Abstract: Most dioecious plants are perennial and subject to trade‐offs between sexual reproduction and vegetative performance. However, these broader life‐history trade‐offs have not usually been incorporated into theoretical analyses of the evolution of separate sexes. One such analysis has indicated that hermaphroditism is favoured over unisexuality when female and male sex functions involve the allocation of nonoverlapping types of resources to each sex function (e.g. allocations of carbon to female function vs. allocations of nitrogen to male function). However, some dioecious plants appear to conform to this pattern of resource allocation, with different resource types allocated to female vs. male sex functions. Using an evolutionarily stable strategy approach, we show that life‐history trade‐offs between sexual reproduction and vegetative performance enable the evolution of unisexual phenotypes even when there are no direct resource‐based trade‐offs between female and male sex functions. This result might help explain the preponderance of perennial life histories among dioecious plants and why many dioecious plants with annual life histories have indeterminate growth with ongoing trade‐offs between sexual reproduction and vegetative growth. [ABSTRACT FROM AUTHOR]

Gynodioecy is the coexistence of hermaphrodites and females in a population. It is supposed to be an intermediate stage in the evolutionary pathway from hermaphroditism to dioecy in angiosperm. Hermaphrodites gain fitness through both seed and pollen production whereas females gain fitness only through seed production. As females spread in a gynodioecious population, sexual selection prompts hermaphrodites to invest in male function and male-biased hermaphrodites prevail. In the gynodioecious shrub Daphne jezoensis (Thymelaeaceae), female frequency is stably around 50% in most populations, and fruit-set rate of hermaphrodites is commonly low. Therefore, D. jezoensis is likely at a later stage in the evolutionary pathway. Female function of hermaphrodites (fruit-set rate, selfing rate, seed size, and germination rate) was assessed in three populations under natural conditions. In order to evaluate the potential seed fertility and inbreeding depression by selfing in hermaphrodites, hand pollination treatments were also performed. Over a 2-year period under natural conditions, 18–29% of hermaphrodites and 69–81% of females set fruit. Across all three populations, the mean fruit-set rate ranged 9.5–49.2% in females and only 3.9–10.2% in hermaphrodites. Even with artificial outcross-pollination, 59–91% of hermaphrodites failed to set any fruit. When self-pollination was performed in hermaphrodites, both of fruit-set and germination rates were decreased, indicating early-acting inbreeding depression. In addition, more than half of the hermaphrodite seeds were produced by selfing under natural pollination, but pollinator service was still required. Totally, hermaphrodites performed poorly as seed producers because of the intrinsically-low fruiting ability and a combination of autogamous selfing and strong inbreeding depression, indicating the absence of reproductive assurance. These results indicate that the mating system of D. jezoensis is functionally close to dioecy. [ABSTRACT FROM AUTHOR]

• In sexually dimorphic species, hermaphrodite flowers in gynodioecious species or male flowers in dioecious species are often larger and produce more nectar than their conspecific female flowers. As a consequence, hermaphrodite or male flowers frequently receive more pollinator visits. • Sex ratio, flower size, floral display, nectar production and floral visits were evaluated in two natural populations of Fuchsia thymifolia, a morphologically gynodioecious but functionally subdioecious insect-pollinated shrub. • Sex ratio did not differ from the expected 1:1 in the two studied populations. As expected, hermaphrodite flowers were larger than female flowers, but in contrast to the general pattern, hermaphrodite flowers did not produce nectar or produced much less than female flowers. Flower visitors were flies (68%) and bumblebees (24%), both of which showed a preference for female flowers. No sex difference was detected in either flower longevity or floral display across the flowering season. • Higher nectar production by females may attract more pollinators, and may be a strategy to enhance female reproductive success in this species. Finally, floral dimorphism and insect preferences did not seem to hamper the maintenance of sub-dioecy or prevent the evolution of dioecy in F. thymifolia. [ABSTRACT FROM AUTHOR]

Sex change affects the sex ratios of plant populations and may play an essential role in the evolutionary shift of sexual systems. Sex change can be a strategy for increasing fitness over the lifetime of a plant, and plant size, environmental factors, and growth rate may affect sex change. We described frequent, repeated sex changes following various patterns in a subdioecious Eurya japonica population over five successive years. Of the individuals, 27.5% changed their sex at least once, and these changes were unidirectional or bidirectional. The sex ratio (females/males/all hermaphrodite types) did not fluctuate over the 5 years. In our study plots, although the current sex ratio among the sexes appears to be stable, the change in sex ratio may be slowly progressing toward increasing females and decreasing males. Sex was more likely to change with higher growth rates and more exposure to light throughout the year. Among individuals that changed sex, those that were less exposed to light in the leafy season and had less diameter growth tended to shift from hermaphrodite to a single sex. Therefore, sex change in E. japonica seemed to be explained by a response to the internal physiological condition of an individual mediated by intrinsic and abiotic environmental factors. [ABSTRACT FROM AUTHOR]

Gene flow in plant populations is heavily affected by species sexual systems. In order to study the effect of sexual systems on genetic structure, we examined plastid and nuclear DNA of 12 dioecious (males and females) and 18 trioecious (males, females and hermaphrodites) populations of Salix myrsinifolia-a boreal shrub with slow range expansion. Populations were located along latitudinal gradients across submarginal and marginal parts of the range. Individuals of each sex morph were all hexaploid. We identified 10 chloroplast DNA haplotypes and scored 205 polymorphic bands with amplified fragment length polymorphism. We found dioecious populations that differed from trioecious populations via the presence of four unique haplotypes and significant difference in Nei's gene diversity index (0.119 vs. 0.116) and down-weighed marker value (1.17 vs. 1.02). The latter parameter, together with haplotype and nucleotide diversity, significantly decreased with latitude similar to the expansion front. Also, we found that 89% of hermaphrodite individuals belong to one distinct in tree parsimony network haplotype. This frequency significantly decreased with latitude towards the expansion front. We suspect that the presence of hermaphrodites in trioecious populations may represent a trade-off between the possibility of producing progeny by single hermaphrodites and genetic variability loss through autogamy. S. myrsinifolia benefits from trioecious sexual systems under colonization events. This phenomenon is no longer a gain closer to the core of the species range. [ABSTRACT FROM AUTHOR]

Most Salix species are characterized by a complex sex structure with populations that contain different ratios of male, female (dioecious populations) and even hermaphroditic individuals (trioecious populations). However, our knowledge of the temporal and spatial variability of such populations is rudimentary. I studied how sexual structure in trioecious populations of Salix myrsinifolia changes over time (from 1995 to 2012) and space in 32 populations across NE Poland and Lithuania. Additionally, the sex of marked individuals was monitored in a population over eight seasons. A number of land cover variables, latitude and population size were investigated to determine what possible mechanisms/factors affect the occurrence of hermaphroditism in this species. Sex structure was found to differ between the northern and southern parts of the study area (primary and secondary ranges), with trioecious populations occurring significantly more often in the secondary part of the range. Females dominated in northern dioecious populations, while sex ratios were balanced in the southern part of the range and in trioecious populations. The mean frequency of hermaphrodites reached 14 % and was decreasing toward the north. The sex structure remained stable over the study period, with an increasing frequency of hermaphrodites (by nearly 6 %). A detailed study of marked individuals revealed that sex is labile in this species. Over 12 % of individuals changed sex from one to three times. Most changes (67 %) were to hermaphroditism, which frequently occurred (34 %) just before death or when entering the post-reproductive period. [ABSTRACT FROM AUTHOR]

Flowering plants exhibit a diverse array of sex expression patterns that become of particular interest when considered in the context of coexisting native-invasive congeners. This study presents findings on sex expression and sex ratio variation in two congeneric tree species, the native Morus rubra and the exotic M. alba, in the Flint Hills region of the Great Plains in the United States. Both species exhibited a subdioecious breeding system (with male, female and hermaphrodite individuals co-occurring within populations), and significantly male-biased sex ratios (i.e., males are more numerous than females). Cumulative sex ratio deviation was higher in the native M. rubra populations than in the exotic M. alba. Within-species sex ratio did not vary significantly among populations. In one focus study area surveyed over three years, most individuals were consistent in sex expression, but approximately 10% switched their sex at least once; the vast majority of these changes were between unisexual and hermaphrodite morphs. There was no size dependence of sex expression based on current sampling. Our results document subdioecy as the breeding system in these Morus species, and advance the group as an interesting study system for its reproductive biology. [ABSTRACT FROM AUTHOR]

In trioecious plant populations, the role of hermaphroditism is often uncertain. We investigated the advantages of hermaphroditism in the dioecious shrub Salix myrsinifolia. The sex ratio of 30 S. myrsinifolia populations in northeastern Poland (secondary range) and Lithuania (primary range) was investigated in 2010-2011. Measures of reproductive (number of catkins, number of flowers in catkins) and vegetative traits (height, diameter, number of shoots, vitality) were taken and compared among sexual morphs. In two populations, measurements collected 14 years prior on marked individuals were used to determine the rate of changes in height, crown diameter and survivorship rates. We found trioecy mostly in the secondary part of the range with an average share of hermaphrodites reaching 21% in the trioecious populations. The sex ratio varied between populations, but tended mostly towards female domination. Several traits differed significantly among sexes. The characteristics of hermaphrodites were often intermediate between males and females and provided no evidence for the higher competitive abilities of hermaphrodites. We concluded that the possible gain of hermaphroditism in the colonization process is restricted to reproduction. We consider hermaphroditism in S. myrsinifolia as an equilibrium between the allocation of resources for growth and reproduction in unstable conditions on the margins of the range. [ABSTRACT FROM AUTHOR]

Subdioecy is thought to occupy a transitional position in the gynodioecy-dioecy pathway, explaining one of the evolutionary routes from hermaphroditism to dioecy. Quantifying any female reproductive advantage of females versus hermaphrodites is fundamental to examining the spectrum between subdioecy and dioecy; however, this is challenging, as multiple interacting factors, such as pollen limitation and resource availability, affect plant reproduction. We compared the female reproductive success of females and hermaphrodites via a field experiment in which we hand-pollinated individuals of the subdioecious shrub Eurya japonica of similar size growing under similar light conditions. Effects of pollen limitation and seed quality were also evaluated through comparing the results of hand- and natural-pollination treatments and performing additional laboratory and greenhouse experiments. Overall, females had higher fruit set and produced heavier fruit and more seeds than hermaphrodites, and these results were more pronounced for hand-pollinated than for natural-pollinated plants of both sexes. We also found that seeds naturally produced by females had a higher mean germination rate. These results indicate that females had a pronounced advantage in female reproductive success under conditions of no pollen limitation. The sexual difference in the degree of pollen limitation suggests a pollinator-mediated interaction, whereas the higher female reproductive success of females even under natural conditions implies that E. japonica is a good model species for elucidating the later stages of the gynodioecy-dioecy pathway. [ABSTRACT FROM AUTHOR]

In subdioecious populations, functional female, male and hermaphrodite individuals coexist. Subdioecy may be a transitional state towards dioecy or a breakdown of dioecy, although lability in sex expression may maintain subdioecy as a stable condition. To better understand the ecological aspects involved in sex ratio dynamics and breeding system evolution, we studied the pollination and female fitness components of female and hermaphrodite individuals of the subdioecious shrub Fuchsia microphylla. In two natural populations at the Trans-Mexican Volcanic Belt we estimated female frequency and several reproductive components of female and hermaphrodite plants under natural pollination and experimental pollination treatments. Average female frequency was 42%, and on average, 42.5% of hermaphrodites produced fruits. Female plants showed a 17-fold female fertility advantage over hermaphrodites through increased fruit production, as the number of seeds and germination rates did not differ between morphs. Hermaphrodite flowers were larger, with similar nectar production and concentration to female flowers, and pollinators did not show consistent morph preferences. Some hermaphrodites produced fruits autonomously, and female flowers excluded from pollinators produced fruits putatively by apomixis. Fruit production in hermaphrodites, but not in females, was related to height, suggesting increased investment of hermaphrodites in the female function at higher resource status. For sex ratios to be at equilibrium, the female fertility advantage should be reduced about eightfold. However, it may be that hermaphrodites are maintained by producing fruits at no cost to the male function at higher resource status, as the gender plasticity hypothesis proposes. [ABSTRACT FROM AUTHOR]

Bisexualism (and its derivatives) is a rare exception in species which already acquired the dioecious breeding system. Although it occurs in some tree species, it is still rather uncommon, except for willows, in which bisexualism occurs in at least 18 species. In most willows such unusual individuals are subdioecious, hermaphroditic or monoecious, or produce intersexes (staminate flowers transform into pistillate flowers or the other way round). The frequency of non-dioecy can vary from a single individual to whole populations, but is in principle rare. Its possible causes are both environmental and genetic. In Salix sex lability is known to be affected by parasites and anthropogenic habitat change. In interspecific willow crosses there are numerous exceptions from dioecy that are mostly caused by polyploidization. Since sex determination in willows is genetic, environmental factors are expected to determine exclusively phenotypic sex traits. [ABSTRACT FROM AUTHOR]

Different sexual morphs of dimorphic and trimorphic populations of plants have different demands for resources correlating with phenotypic and ecological dissimilarities among morphs. Few studies have investigated the variations in mycorrhizal symbiosis across the sexual morphs of trimorphic plant populations. Here, we report seasonal variations in the mycorrhizal colonization of the roots of female, male and hermaphroditic trees of Jacaratia mexicana under field conditions in central Mexico. Mycorrhizal colonization was low in the rainy season, when J. mexicana bears only leaves, but high in the dry season when it flowers and fruits. In the dry season, the mycorrhizal colonization of female and male trees was over twofold higher than that of the hermaphroditic trees, and all three morphs had low mycorrhizal colonization in the rainy season. The lack of seasonal variation in the mycorrhizal colonization of hermaphroditic trees, together with other available evidence, supports the idea that hermaphroditic trees of J. mexicana might have some physiological disadvantages compared with male and female trees. [ABSTRACT FROM AUTHOR]

Background The ‘gynodioecy–dioecy pathway’ is considered to be one of the most important evolutionary routes from hermaphroditism to separate sexes (dioecy). Despite a large accumulation of evidence for female seed fertility advantages in gynodioecious species (females and hermaphrodites coexist) in support of the first step in the gynodioecy–dioecy pathway, we still have very little evidence for the second step, i.e. the transition from gynodioecy to dioecy. Scope We review the literature to evaluate whether basic predictions by theory are supported. To establish whether females' seed fertility advantage and frequencies are sufficient to favour the invasion of males, we review these for species along the gynodioecy–dioecy pathway published in the last 5 years. We then review the empirical evidence for predictions deriving from the second step, i.e. hermaphrodites' male fertility increases with female frequency, selection favours greater male fertility in hermaphrodites in gynodioecious species, and, where males and hermaphrodites coexist with females (subdioecy), males have greater male fertility than hermaphrodites. We review how genetic control and certain ecological features (pollen limitation, selfing, plasticity in sex expression and antagonists) influence the trajectory of a population along the gynodioecy–dioecy pathway. Conclusions Females tend to have greater seed fertility advantages over hermaphrodites where the two coexist, and this advantage is positively correlated with female frequency across species, as predicted by theory. A limited number of studies in subdioecious species have demonstrated that males have an advantage over hermaphrodites, as also predicted by theory. However, less evidence exists for phenotypic selection to increase male traits of hermaphrodites or for increasing male function of hermaphrodites in populations with high female frequency. A few key case studies underline the importance of examining multiple components of male fertility and the roles of pollen limitation, selfing and plasticity, when evaluating advantages. We conclude that we do not yet have a full understanding of the transition from gynodioecy to dioecy. [ABSTRACT FROM AUTHOR]

Sex-allocation plasticity is thought to play an important role in the evolution of separate sexes in plants. Accordingly, much attention has been paid to environmentally induced variation in fruit and seed production in sexually dimorphic species, but we know little about whether this variation arises as a direct response to environmental variation or is instead an indirect consequence of changes in plant size. In this study, we characterize sex-allocation plasticity across a resource gradient for several reproductive traits in hermaphrodites of gyno(sub)dioecious Fragaria virginiana Duch. We find significant plasticity, on average, for flower number, proportion fruit set, ovule number, proportion seed set, and runner number in response to resource variation. Plasticity of most traits examined tended to be at least partially independent of variation in plant size, suggesting that it is not simply an indirect consequence of plant allometry. Moreover, we find genetic variation for plasticity of key reproductive traits. Comparisons of relative plasticities among traits reveal that F. virginiana hermaphrodites are more likely to adjust female investment via changes in fruit and seed set than ovule number, and most likely to adjust male investment via flower number rather than anther number or pollen per anther, although there is genotypic variation for plasticity in pollen per anther. Evidence of within-population variation can logically be extended to suggest that variation in hermaphrodite sex-expression seen among natural populations of F. virginiana may be due, at least in part, to sex-allocation plasticity. [ABSTRACT FROM AUTHOR]

The gender of dimorphic plant species is often affected by ecophysiological variables. Differences have been interpreted as a response of the sexes to meet specific resource demands associated with reproduction. This study investigated whether sex-specific variations in ecophysiological traits in response to water availability determine the performance of each sex in different habitats, and therefore promote extreme spatial segregation of the sexes in the subdioecious plant, Honckenya peploides. Twenty-seven plants of each sex were individually potted in dune sand and assigned randomly to one of three water treatments. Well-watered plants were watered daily to field capacity, whereas plants in the moderate and high-water stress treatments received 40% and 20%, respectively, of the water given to well-watered plants. Photochemical efficiency, leaf spectral properties and components of relative growth rate (leaf area ratio and net assimilation rate) were measured. Photochemical efficiencies integrated over time were higher in male than in female plants. Water deficit decreased maximum quantum yield in female plants more rapidly than in male plants, but female plants (unlike male plants) had recovered to initial values by the end of the experiment. Maximum quantum yield in male plants was more affected by water stress than in female plants, indicating that male plants were more susceptible to photoinhibition. The two sexes did not differ in growth rate, but male plants invested a higher proportion of their biomass in leaves, had a higher leaf area per unit biomass and lower net assimilation rate relative to female plants. Female plants had a higher water content and succulence than male plants. Differences in stomatal density between the sexes depended on water availability. The results suggest that the two sexes of H. peploides have different strategies for coping with water stress. The study also provides evidence of sex differences in allocation traits. We conclude that between-sex differences in ecophysiological and allocation traits may contribute to explain habitat-related between-sex differences in performance and, therefore, the spatial segregation of the sexes. [ABSTRACT FROM AUTHOR]

Using phylogenies to make sound inferences about character evolution depends on a variety of factors, including tree uncertainty, taxon sampling, and the degree of evolutionary lability in the character of interest. We explore the effect of these and other sources of ambiguity for maximum likelihood (ML)-based inferences of sexual-system evolution in Wurmbea, a small genus of geophytic monocots from the Southern Hemisphere. We reconstructed Wurmbea phylogeny using four noncontiguous regions (ca. 5.5 kb) of the plastid genome across a broad sampling of taxa, and we confirm that the genus is divided into two well-supported clades, each defined by its geography (Africa vs. Australasia) and variation in sexual system (i.e., uniformly monomorphic vs. sexually variable, respectively). We demonstrate that the predominantly Australian clade includes the sexually monomorphic species Iphigenia novae-zelandiae. We observe treewide uncertainty in the state of all ancestral nodes, and therefore all state transitions, when all taxa in Wurmbea are considered. We demonstrate that this is primarily a consequence of interspersion of terminals with gender dimorphism vs. monomorphism throughout the Australasian clade, rather than tree uncertainty or the presence of very short internal branches. We accounted for tree uncertainty by randomly sampling alternative resolutions of branches that are poorly supported by ML bootstrap analysis, effectively interpreting these as soft polytomies. Under the assumption that well-supported aspects of our gene tree accurately depict organismal phylogeny, there is a marked evolutionary lability in the sexual systems of Australasian Wurmbea. A more problematic issue is that our results contradict the monophyly of two sexually polymorphic Australian species, Wurmbea dioica and Wurmbea biglandulosa. If this reflects paraphyly at the species level, lateral gene transfer, or failed coalescence, then the interpretations of character transitions will need to be adjusted. Our analysis provides an example of the impediments to linking macroevolutionary pattern with microevolutionary processes for evolutionarily labile traits in recently evolved plant groups that possess a high degree of variation in sexual characters. [ABSTRACT FROM AUTHOR]

The maintenance of females in gender dimorphic populations requires that they have a fitness advantage to compensate for their loss of male reproductive function. We assess whether inbreeding avoidance provides this advantage in two subdioecious Wurmbea dioica populations by estimating seed production, outcrossing rates and inbreeding depression. Fruiting males produced less than half as many seeds as females, owing to low outcrossing rates and early acting inbreeding depression. Inbreeding coefficients of fruiting males demonstrated that progeny were more inbred than their parents, implying that few selfed progeny reach maturity, as confirmed by inbreeding depression estimates that exceeded 0.85. In a glasshouse experiment, open-pollinated females exhibited a fitness advantage of 3.7 relative to fruiting males, but when we increased fruiting male outcrossing rate, female advantage was only 1.4. This reduced advantage is insufficient to maintain females if nuclear genes control sex. Thus, inbreeding avoidance could maintain females at high frequencies, although this is contingent upon high frequencies of fruiting males, which can be altered by environmentally determined gender plasticity. [ABSTRACT FROM AUTHOR]

A functional view of gender helps evolutionary biologists evaluate the mechanisms underlying breeding-system evolution. Evolutionary pathways from hermaphroditism to dioecy include the intermediate breeding systems of gynodioecy and androdioecy. These pathways start with the invasion of unisexual mutants, females or males, respectively, followed by alteration of the hermaphrodites to allocate more to the sexual function that the unisexuals lack. Eventually, hermaphrodites become unisexual and dioecy has evolved. Some species evolving along these pathways stop short of completing this second step, or even revert back from dioecy. We evaluate the hypothesis that gender plasticity is involved in these transitions to and from dioecy. Evidence from studies of subdioecious species that have evolved along the gynodioecy pathway suggests that gender plasticity occurs and stabilizes subdioecy by lowering the cost of producing seed. Factors influencing species evolving toward androdioecy, or reverting to androdioecy from dioecy, appear to be more varied and include reproductive assurance, herbivory and gender plasticity. In general, gender specialization appears to be favored in resource-poor environments regardless of which pathway is taken to dioecy.New Phytologist(2005)doi: 10.1111/j.1469-8137.2005.01339.x© New Phytologist(2005) [ABSTRACT FROM AUTHOR]

The distribution of male and female inflorescences among stems of Myrica gale L. is strongly bimodal and indicates a genetical dimorphism in gender. Predominantly or strictly male stems are about twenty times as frequent as predominantly or strictly female stems. There is considerable variation among male and female stems in their precise gender. Quantitative values of the phenotypic sex (the observed gender based on the relative proportions of male and female inflorescences) and functional sex (the relative probabilities of transmitting genes to descendants through male and female gametes) of samples of stems are estimated. [ABSTRACT FROM AUTHOR]

The genus Cirsium comprises both gynodioecious and dioecious species. The observation of microsporogenesis in female plants of C. montanum, C. oleraceum, C. palustre and C. spinosissimum shows that the male sterility is due to a degeneration of the tapetum. This degeneration occurs more or less early according to the species and, in the light of these results, a scheme of evolution in the male sterility mechanism is proposed. Furthermore, the male sterility mechanism in C. montanum is very similar to that previously found in female plants of the dioecious species C. arvense. This fact enhances the possibility of evolution of the dioecy of C. arvense from the gynodioecy found in other species. According to these results, a general scheme of evolution of sexes in the genus Cirsium is proposed. [ABSTRACT FROM AUTHOR]

Abstract: The determination of sex-ratio variation is a useful guide for identifying models of sex-ratio evolution in gynodioecious species. This study counted the numbers of female and hermaphrodite plants in five populations of the gynodioecious shrub Gnidia wikstroemiana distributed over 2.3° of latitude in dry, highland shrublands and grasslands of South Africa. Population sizes ranged from 71 to 435 plants with populations occurring on diverse rocky terrains along a 422-m altitudinal gradient. Female frequencies varied from 35 to 47% but sex-ratios did not differ significantly among the five populations. However, sex-ratios in three populations occurring on ridge slopes deviated significantly from the 1:1 ratio while those populations located on the crests of mountains did not. These results suggest that both nuclear gynodioecy and subdioecy are involved in the sex-ratio evolution of this species. [Copyright &y& Elsevier]

Female reproductive success in females versus hermaphrodites has been well documented. However, documenting a potential advantage in male fertility of male versus hermaphrodite individuals in subdioecious species is also essential for understanding the evolutionary pathway toward dioecy from hermaphroditism via gynodioecy. Siring success in terms of fruit set, fruit mass, number of seeds and mean seed mass was compared by hand-pollinated crosses in the subdioecious shrub Eurya japonica The pollen was from male and hermaphrodite individuals, and the pollen recipients were females and hermaphrodites. Seed quality was also evaluated in terms of seed germination rate, seed germination day and seedling survival. Overall, pollen from males sired more fruits of larger size and more seeds than did pollen from hermaphrodites. The male advantage was observed when pollen recipients were females, whereas no effect was found in hermaphrodite recipients. Pollen from males also produced better quality seeds with higher germination rate and sooner germination day. Although these results could also be explained by a higher pollen load for crosses with male pollen donors, we took care to saturate the stigma regardless of the pollen donor. Therefore, these results suggest that male individuals of E. japonica have advantages in male fertility in terms of both quantity and quality. Our previous studies indicated that females exhibit higher female reproductive success compared with hermaphrodites. Thus, both the female and male functions of hermaphrodites are outperformed by females and males, respectively, raising the possibility that the subdioecious E. japonica at this study site is entering the transitional phase to dioecy along the gynodioecy-dioecy pathway., (© The Authors 2016. Published by Oxford University Press on behalf of the Annals of Botany Company.)

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 fruitsetting 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. [ABSTRACT FROM AUTHOR]

Genetic diversity of allozymes, genetic identity based on allozyme variability, and phylogenetic relationships were studied with respect to breeding system diversity, population size, and island age in 20 of the 29 species of Schiedea and Alsinidendron (Caryophyllaceae: Alsinoideae), a monophyletic lineage endemic to the Hawaiian Islands. Average levels of genetic variability in Schiedea and Alsinidendron were comparable to or higher than those found in other Hawaiian lineages for which equivalent data are available [Bidens, Tetramolopium, and the silversword alliance (Asteraceae: Madiinae)] and similar to average values for species of dicots. Allozyme variability was strongly dependent on breeding system, which varies widely in the Hawaiian Alsinoideae. Species with autogamous breeding systems showed very low variability, measured as the number of alleles per locus, percent polymorphic loci, and mean heterozygosity per locus. Outcrossing hermaphroditic and dimorphic species (those with gynodioecious, subdioecious, and dioecious breeding systems) showed significantly higher genetic variability. Small population size was associated with lower values for all measures of genetic variability. Nearly half of the species occurring in small populations are also autogamous; thus, both factors may have influenced levels of genetic variability in these species. Founder effect was apparent in one species (Schiedea adamantis), which occurs in a single large population, has a gynodioecious breeding system but a very low genetic variability. Island age appeared to have little effect on genetic variability. Slightly lower values of genetic variability for species occurring on Kaua'i and O'ahu result primarily from the occurrence of autogamous Alsinidendron species on those islands. Values for Nei's genetic identity for different species pairs were 0.201-0.942, a far greater range than in Bidens, the silversword alliance, and Tetramolopium. Using UPGMA clustering, there was only moderate support for relationships detected through cladistic analysis. Nei's unbiased genetic identity (I) was greatest among species with outcrossing breeding systems, which for the most part clustered together. Nei's genetic identities for self-fertilizing species were low, indicating that these species are less similar to one another and to outcrossing species, regardless of their affinities based on cladistic analysis. Parsimony analysis of allele frequency data supported two clades also found in phylogenetic analyses using morphological and molecular data. Clades recognized in parsimony analysis of allele frequencies were those lineages containing selfing species, indicating that conditions favoring fixation of alleles occurred in ancestral species. In contrast, maintenance of high genetic diversity in outcrossing species interferes with recognition of phylogenetic relationships using allozyme variability., (© 1996 The Society for the Study of Evolution.)

Population-genetic models of nucleo-cytoplasmic gynodioecy are shown to allow invasion of males and conversion to dioecy in a single cytotype. Pleiotropic effects of restorer alleles on fertility through male or female function can maintain a cytoplasmic polymorphism in a population that prevents evolution to dioecy regardless of the pollen fertility of males. However, a cytoplasmic polymorphism has little effect on, and may even reduce, the minimum pollen fertility required for the spread of males into an equilibrium gynodioecious population. Where the thresholds for dioecy are similar, the presence of males during a transient preequilibrium high frequency of females can accelerate evolution to dioecy by more than 50 times relative to nuclear male sterility. However, the appearance of a nonrestorable male-sterile cytotype generally eliminates males from both subdioecious and dioecious populations, converting them to purely cytoplasmic gynodioecy. These models contradict the previously suggested notion that nucleo-cytoplasmic gynodioecy represents a "stable" intermediate breeding system and instead show that such gynodioecy can generally evolve to subdioecy, and often to dioecy, as easily as nuclear gynodioecy., (© 1994 The Society for the Study of Evolution.)