Gynodioecious Populations Research Articles

Characterization of gynodioecious Pune selection lines of papaya

Among major global threats to papaya cultivation, papaya ringspot virus (PRSV) is the most challenging one. In the absence of any PRSV resistant commercial papaya cultivar, PRSV management is restricted to minimizing yield losses. ICAR-Indian Agricultural Research Institute, Regional Station, Pune has developed PRSV tolerant dioecious papaya lines, Pune Selection (PS)-1, PS-2, PS-3 and PS-5. Being dioecious these lines have limited acceptability among farmers. Gynodioecious population from these lines were developed and characterized. They are numbered PS-1-1, PS-2-1, PS-3-1 and PS-5-1. These lines were characterized against prevailing commercial gynodioecious cultivar, Red Lady, for five generations. The average plant height of PS-2-1 and PS-5-1 (183 cm) was more than Red Lady (158 cm), however, stem girth of all lines was lesser than Red Lady. The fruiting height of all lines was less than Red Lady (87 cm). Length of the fruiting column of all lines was more than Red Lady (37 cm), except in PS-1-1. Fruit yield of all lines was more than Red Lady (16 kg/plant). Intensity of PRSV infection in Red Lady (48%) was considerably more than all lines. These lines can be used for developing PRSV tolerant gynodioecious papaya variety.

Geranium sylvaticum increases pollination probability by sexually dimorphic flowers.

Sexual dimorphism is expressed as different morphologies between the sexes of a species. Dimorphism is pronounced in gynodioecious populations which consist of female and hermaphrodite individuals. The small size of female flowers in gynodioecious species is often explained by resource re-allocation to seed production instead of large flowers. However, pollinator attraction is critical to female fitness, and factors other than resource savings are needed to explain the small size of female flowers. We hypothesized that the floral size dimorphism in the perennial gynodioecious Geranium sylvaticum (L.) is adaptive in terms of pollination. To test this "pollination hypothesis," we video recorded the small female and large hermaphrodite G.sylvaticum flowers. We parameterized floral visitor behavior when visiting a flower and calculated pollination probabilities by a floral visitor as the probability of touching anther and stigma with the same body part. Pollination probability differed in terms of flower sex and pollinator species. Bumblebees had the highest pollination probability. The small female flowers were more likely to receive pollen via several pollinator groups than the large hermaphrodite flowers. The pollen display of hermaphrodites matched poorly with the stigma display of hermaphrodites, but well with that of females. Although the small size of female flowers is commonly explained by resource re-allocation, we show that sexual dimorphism in flower size may increase the main reproductive functions of the females and hermaphrodites. Dimorphism increases pollination probability in females and fathering probability of the hermaphrodites likely driving G.sylvaticum populations towards dioecy.

Extreme mitochondrial DNA divergence underlies genetic conflict over sex determination

Cytoplasmic male sterility (CMS) is a form of genetic conflict over sex determination that results from differences in modes of inheritance between genomic compartments.1-3 Indeed, maternally transmitted (usually mitochondrial) genes sometimes enhance their transmission by suppressing the male function in a hermaphroditic organism to the detriment of biparentally inherited nuclear genes. Therefore, these hermaphrodites become functionally female and may coexist with regular hermaphrodites in so-called gynodioecious populations.3 CMS has been known in plants since Darwin's times4 but is previously unknown in the animal kingdom.5-8 We relate the first observation of CMS in animals. It occurs in a freshwater snail population, where some individuals appear unable to sire offspring in controlled crosses and show anatomical, physiological, and behavioral characters consistent with a suppression of the male function. Male sterility is associated with a mitochondrial lineage that underwent a spectacular acceleration of DNA substitution rates, affecting the entire mitochondrial genome-this acceleration concerns both synonymous and non-synonymous substitutions and therefore results from increased mitogenome mutation rates. Consequently, mitochondrial haplotype divergence within the population is exceptionally high, matching that observed between snail taxa that diverged 475 million years ago. This result is reminiscent of similar accelerations in mitogenome evolution observed in plant clades where gynodioecy is frequent,9,10 both being consistent with arms-race evolution of genome regions implicated in CMS.11,12 Our study shows that genomic conflicts can trigger independent evolution of similar sex-determination systems in plants and animals and dramatically accelerate molecular evolution.

Do Genetic Drift and Gene Flow Affect the Geographic Distribution of Female Plants in Gynodioecious Lobelia siphilitica?

Variation in population sex ratio is particularly pronounced in gynodioecious angiosperms. Extremely high female frequencies in gynodioecious populations cannot be readily explained by selective forces alone. To assess the contributions of drift and gene flow to extreme sex-ratio variation, we documented sex ratio and population size in 92 populations of Lobelia siphilitica across its range and genotyped plants using plastid and nuclear genetic markers. Similarity in spatial patterns of genetic and demographic variables may suggest that drift and/or gene flow have contributed to population sex-ratio variation in L. siphilitica. We found strong spatial structuring of extremely high female frequencies: populations with >50% female plants are restricted to the south–central portion of the range. However, we did not detect any spatial structuring in population size nor metrics of genetic diversity, suggesting that extreme variation in female frequency is not strongly affected by drift or gene flow. Extreme sex-ratio variation is frequently observed in gynodioecious plants, but its causes are difficult to identify. Further investigation into mechanisms that create or maintain the spatial structure of sex ratios in gynodioecious species will provide much needed insight.

Female advantage in gynodioecious plants: A meta-analysis focused on seed quality.

In gynodioecious systems, female plants must counteract the selective disadvantage of not passing genes via pollen production, as hermaphrodites can. Theory predicts that females must produce more or better-quality seeds than hermaphrodites in order to be maintained within the same population. This female advantage has been widely measured and reported for seed number, but whether female advantage is gained through the production of better seeds remains relatively under-studied. Here, a meta-analysis approach was used to investigate whether females in gynodioecious species produce seeds of better quality than hermaphrodites (measured as seed mass, seed nutrient content, seed germinability and seedling survival and performance) in addition to achieving a larger seed production. In total, 50 studies were included, reporting traits for 34 gynodioecious species in 17 different families. Female advantage was significant for seed number and seed germination, but was not detected for seed mass, seed nutrient content or seedling performance. A female advantage in seed number was corroborated in this meta-analysis, which together with better seed germination, may explain maintenance of female plants within gynodioecious populations.

Structural variations in papaya genomes

BackgroundStructural variations (SVs) are a type of mutations that have not been widely detected in plant genomes and studies in animals have shown their role in the process of domestication. An in-depth study of SVs will help us to further understand the impact of SVs on the phenotype and environmental adaptability during papaya domestication and provide genomic resources for the development of molecular markers.ResultsWe detected a total of 8083 SVs, including 5260 deletions, 552 tandem duplications and 2271 insertions with deletion being the predominant, indicating the universality of deletion in the evolution of papaya genome. The distribution of these SVs is non-random in each chromosome. A total of 1794 genes overlaps with SV, of which 1350 genes are expressed in at least one tissue. The weighted correlation network analysis (WGCNA) of these expressed genes reveals co-expression relationship between SVs-genes and different tissues, and functional enrichment analysis shows their role in biological growth and environmental responses. We also identified some domesticated SVs genes related to environmental adaptability, sexual reproduction, and important agronomic traits during the domestication of papaya. Analysis of artificially selected copy number variant genes (CNV-genes) also revealed genes associated with plant growth and environmental stress.ConclusionsSVs played an indispensable role in the process of papaya domestication, especially in the reproduction traits of hermaphrodite plants. The detection of genome-wide SVs and CNV-genes between cultivated gynodioecious populations and wild dioecious populations provides a reference for further understanding of the evolution process from male to hermaphrodite in papaya.

Sexual conflict in protandrous flowers and the evolution of gynodioecy.

Sexual interference between male and female function in hermaphrodite plants is reduced by protandry. In environments with insufficient pollinator service, prolongation of male function owing to limited pollen removal could restrict the duration of female function and lower seed production. We provide evidence that this form of sexual conflict has played a role in the spread of females in gynodioecious populations of Cyananthus delavayi in the pollen-limited environments in which this subalpine species occurs. Using field experiments involving artificial pollen removal from the strongly protandrous flowers of hermaphrodites, we demonstrated a trade-off between male- and female-phase duration with no influence on overall floral longevity. Pollen removal at the beginning of anthesis resulted in hermaphrodite seed production matching that of females. In contrast, restricted pollen removal increased the duration of male function at the expense of female function lowering maternal fertility compared to females. This pattern was evident in five populations with females experiencing a twofold average seed fertility advantage compared to hermaphrodites. Gynodioecy often appears to evolve from protandrous ancestors and pollen limitation is widespread in flowering plants suggesting that sexual conflict may play an unappreciated role in the evolution of this form of sexual dimorphism.

Seed germination of gynodioecious species: theoretical considerations and a comparison of females and hermaphrodites.

Better seed germination of females than of hermaphrodites is not a major contributor to the greater geometric lifetime fitness that females require to be maintained in a gynodioecious population. Gynodioecy is a sexually dimorphic breeding system in which females (F, male sterile) and hermaphrodites (H) coexist in the same population. For plants with nuclear (biparental) inheritance of male sterility, theory predicts that except when the product of selfing rate (s) and inbreeding depression (δ) in H is high (sδ > 0.50), F must compensate (female advantage) for the loss of gene transmission via pollen production by producing more or higher-quality offspring than H to be maintained in the population. For species with cytoplasmic (maternal) inheritance of male sterility, the female requires only a small compensation in seed production or some other offspring fitness trait to persist. Reallocation to seeds of resources saved by loss of pollen production is expected to increase the quantity (number) and/or quality (mass, germinability) of seeds produced by F, thus compensating for the lack of pollen production. The primary aim of our study was to compare seed germination of F and H via a literature review. Based on theoretical considerations, we hypothesized that seeds of F should germinate better or equally as well as those of H. We found that of 235 case studies for 47 species Fgerm > Hgerm in 48.1%, Fgerm = Hgerm in 38.3% and Fgerm < Hgerm in 13.6%. Our results are very similar to those of a previously published meta-analysis that included germination of F and H for 12 species. For 162 cases on seed size, F > H in 29.0%, F = H in 63.6% and F < H in 7.4%. Since [(Fgerm > Hgerm) < (Fgerm ≤ Hgerm)] and [(Fseedsize > Hseedsize) < (Fseedsize ≤ Hseedsize)], these results suggest that seed quality is not a major fitness component of female advantage.

Flowering sex ratios and costs of reproduction in gynodioeciousOcotea oblonga(Lauraceae)

AbstractIn gynodioecious plant species, both female and hermaphrodite individuals produce fruit, but only hermaphrodites produce pollen. Such sex-specific differences in reproductive investment may contribute to dimorphism, but the magnitude and ecological effects are still unclear, especially for gynodioecious tropical trees where collecting flowers and determining the sex is complex. We documented flowering and fruiting over three years in a natural population of Ocotea oblonga (Lauraceae) trees in a tropical moist forest, Panama. We determined sex from freshly collected flowers, counted and measured fruit, and used long-term growth data for each individual. We confirmed that O. oblonga is gynodioecious. No tree switched sex or had flowers of both sexes. The population was hermaphrodite-biased. We found no ecological differences in reproductive investment (seed, fruit, or tree size, or growth rate) between the sexes, indicating that the sex differential in the cost of reproduction is much smaller in woody gynodioecious taxa than in dioecious taxa. Females produced more fruit than hermaphrodites, which may contribute to their persistence in the population. Accordingly, and contrary to most studies of temperate gynodioecious populations, our study of a tropical tree shows no differential cost of reproduction in a hermaphrodite-biased population. Consequently, other factors such as seed fertility or herbivory could drive the biased sex ratio in this population.

Impact of prairie fragment size on proportion of females and reproductive success of Lobelia spicata Lam., a gynodioecious species.

Due to ongoing human impacts, plant species increasingly occur in landscapes that are highly fragmented, with remaining natural habitats occupying small areas, resulting in populations that are smaller and more isolated than in previous time periods. This changed metapopulation structure is expected to have negative impacts on seed production. For example, the proportion of female plants within gynodioecious populations may be more volatile due to genetic drift in small populations associated with small habitat fragments, with concomitant impacts on seed production. My aims were to determine: (i) if variation in proportion of females is larger in smaller fragments; and (ii) if such changes in female frequency in small fragments result in reduced seed production. Thirty-two populations of Lobelia spicata Lam., a gynodioecious species, were surveyed in 2000, 2001 and 2009 in the tallgrass prairie region of Midwestern North America (Illinois and Indiana, USA). Data were collected for: proportion of female plants, total number of flowering plants (measure of population size), seed set per plant and prairie fragment size (another measure of population size). The proportion of females is more variable in smaller prairie fragments. Seed number per fruit decreases as the proportion of females increases in a population, but only significantly for female plants. The number of flowering plants is positively associated with fruit production for both genders. Populations within larger prairie fragments have higher seed production. The reproductive consequences of habitat fragmentation depend on the plant breeding system. While both sexes were negatively impacted, females were more adversely affected.

Beyond balancing selection: frequent mitochondrial recombination contributes to high-female frequencies in gynodioecious Lobelia siphilitica (Campanulaceae).

Gynodioecy is a sexual system in which females and hermaphrodites co-occur. In most gynodioecious angiosperms, sex is determined by an interaction between mitochondrial male-sterility genes (CMS) that arise via recombination and nuclear restorer alleles that evolve to suppress them. In theory, gynodioecy occurs when multiple CMS types are maintained at equilibrium frequencies by balancing selection. However, some gynodioecious populations contain very high frequencies of females. Highfemale frequencies are not expected under balancing selection, but could be explained by the repeated introduction of novel CMS types. To test for balancing selection and/or the repeated introduction of novel CMS, we characterised cytoplasmic haplotypes from 61 populations of Lobelia siphilitica that vary widely in female frequency. We confirmed that mitotype diversity and female frequency were positively correlated across populations, consistent with balancing selection. However, while low-female populations hosted mostly common mitotypes, high-female populations and female plants hosted mostly rare, recombinant mitotypes likely to carry novel CMS types. Our results suggest that balancing selection maintains established CMS types across this species, but extreme female frequencies result from frequent invasion by novel CMS types. We conclude that balancing selection alone cannot account for extreme population sex-ratio variation within a gynodioecious species.

In search of an identity for Salvia bertolonii (Lamiaceae)

Classical and geometric morphometric analyses of floral characters of Salvia pratensis agg. were performed on populations from the E Adriatic area. Both types of analyses yielded congruent results, and three groups of populations, attributed to S. verbenaca, which served as an outgroup, S. saccardiana and S. pratensis (incl. S. bertolonii), were unambiguously identified. Results confirmed floral size dimorphism, with hermaphroditic flowers being significantly bigger than female ones, while we newly identified floral shape dimorphism in S. pratensis, too. No gynodioecious populations were observed in morphologically otherwise very uniform S. saccardiana. Being a SE Alpine endemic, the latter was for the first time recorded in Croatia. While S. saccardiana merits species rank, we found no support for the segregation of S. bertolonii and S. pratensis. Clinal variation in floral characters of S. pratensis, generally detected in populations of various taxa along the E Adriatic coast, lacks the spatial consistency and geographic pattern that would allow for a clear delimitation of S. pratensis and S. bertolonii.

Sexually Antagonistic Variation and the Evolution of Dimorphic Sexual Systems.

Multicellular Eukaryotes use a broad spectrum of sexual reproduction strategies, ranging from simultaneous hermaphroditism to complete dioecy (separate sexes). The evolutionary pathway from hermaphroditism to dioecy involves the spread of sterility alleles that eliminate female or male reproductive functions, producing unisexual individuals. Classical theory predicts that evolutionary transitions to dioecy are feasible when female and male sex functions genetically trade off with one another (allocation to sex functions is sexually antagonistic) and rates of self-fertilization and inbreeding depression are high within the ancestral hermaphrodite population. We show that genetic linkage between sterility alleles and loci under sexually antagonistic selection significantly alters these classical predictions. We identify three specific consequences of linkage for the evolution of dimorphic sexual systems. First, linkage broadens conditions for the invasion of unisexual sterility alleles, facilitating transitions to sexual systems that are intermediate between hermaphroditism and dioecy (androdioecy and gynodioecy). Second, linkage elevates the equilibrium frequencies of unisexual individuals within androdioecious and gynodioecious populations, which promotes subsequent transitions to full dioecy. Third, linkage dampens the role of inbreeding during transitions to androdioecy and gynodioecy, making these transitions feasible in outbred populations. We discuss implications of these results for the evolution of dimorphic reproductive systems and sex chromosomes.

Restricted female function of hermaphrodites in a gynodioecious shrub, Daphne jezoensis (Thymelaeaceae).

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.

Sex allocation in gynodioecious Cyananthus delavayi differs between gender morphs and soil quality.

Sex allocation in Cyananthus delavayi. Gynodioecy, where females and hermaphrodites coexist in the same natural population, is particularly suitable for predicting the ecological pressures that drive the stability of gender polymorphism. Since females have a disadvantage in that they only contribute to the next generation via ovules, they should gain an advantage via other means, of which resource allocation is an important component. Thus, to study their sex allocation is very helpful to understand how the dimorphic sexual system is maintained in natural systems. We studied the sex allocation patterns and reproductive output of the gynodioecious Cyananthus delavayi in three populations with different soil qualities (organic matter, N, P and K). The hermaphroditic flowers and pistils were much larger than those of female individuals. Although both gender morphs invested similar biomass in the pistils, females allocated more of their resource pool to the seed production, while hermaphrodites allocated more to pollinator advertisement. The pollen production of hermaphrodites did not differ between populations, suggesting that pollen production by hermaphrodites was not limited by soil nutrients. Fruit set of females, but not hermaphrodites, decreased with declining soil quality, whereas seeds per fruit of both females and hermaphrodites were highest in poor soils. Overall, this study shows that females achieve greater reproductive success by allocating more of their resource pool to enhancing seed production, which should favor their presence in gynodioecious populations. The hermaphrodites achieve reproductive success from both pollen and seed production, and unnecessarily reduce their allocation to pollen production. Soil quality should explain, at least partially, the sexual allocation patterns. Furthermore, some of our findings contradict previous hypotheses, thus adding a new example to the body of research on plant sex allocation and the development of future theories.

Three psychophilous Asteraceae species with distinct reproductive mechanisms in southeastern Brazil

ABSTRACTThe Asteraceae have a high diversity of reproductive mechanisms, as shown by studies conducted mainly with species from temperate regions. Aiming to contribute knowledge on the reproduction of tropical species, we compared the floral biology and breeding systems of the co-occurring species Adenostemma brasilianum, Bidens segetum and Grazielia intermedia, and additionally identified their pollinators. The study was conducted in a seasonal semideciduous forest fragment in southeastern Brazil. Florets opened in early morning and were protandrous. In the first day of anthesis, secondary pollen presentation occurred during the staminate phase, mainly in the morning. In the same day, the pistillate phase began, being long in A. brasilianum and B. segetum (up to 4 days) and short in G. intermedia (1 day). Pollen viability was high in all three species, except for two G. intermedia individuals which showed male sterility, thus characterising a gynodioecious population. Self-compatibility was observed in A. brasilianum and self-incompatibility in B. segetum. In G. intermedia, besides sexual reproduction, apomixis was also observed. This is the first report of apomixis in the genus Grazielia. Although the studied species show distinct reproductive mechanisms, all of them benefitted from cross-pollination, which was mostly done by butterflies.

Microsatellite genetic diversity and mating systems in the columnar cactus Pachycereus pringlei (Cactaceae)

Mating systems and long-term climatic fluctuations have large impact on genetic diversity and population structure of plants; however, the genetic consequences have been poorly documented in species with geographic variation in mating systems. Here we used genetic variation from microsatellite markers to investigate the biogeographic pattern of Pachycereus pringlei, a columnar cactus with exceptional assortment of mating systems and wide range of distribution in the Sonoran Desert. Bayesian and multivariate analyses distinguished five genetic populations associated with differences in mating systems. Two hermaphroditic populations restricted to Cerralvo and Catalana islands were highly differentiated (ΦPT=0.26 and 0.21, respectively). Cerralvo Island showed the lowest genetic diversity (HO=0.26). Two other populations showed wide geographical distribution: a mostly gynodioecious population in the North comprising mostly hermaphrodite and female individuals, and a trioecious population in the South comprising unisexual and hermaphrodite individuals. The latter population exhibited the lowest level of genetic differentiation (ΦPT=0.14) and highest genetic diversity (HO=0.45). Cabo San Lucas, another trioecious population restricted to the southern end of the Baja California Peninsula, exhibits prevalence of unisexual individuals (0.76) but highly differentiated (ΦPT=0.19) and relatively moderate genetic diversity (HO=0.38). Latitudinal decrease of genetic diversity and isolation by distance supports northwards range expansion and southern BCP refugia, while Ecological Niche Models supported suitable climatic conditions during the Last Glacial Maximum in this region. These results suggest that the current biogeographic pattern of P. pringlei has been driven by longstanding climatic fluctuations associated with differential colonization abilities of genders and geographic variation of selfing and outcrossing rates. In conclusion, P. pringlei is not a panmictic assemblage and the interaction of historical vicariance and dispersal processes with mating systems may well explain present-day patterns of genetic variation.

To be or not to be better pollinated: Differences between sex morphs in marginal gynodioecious populations.

Changes in the pollinator communities of marginal plant populations can affect their pollination quantity or quality. Geographic variation in pollination success can alter the reproductive advantage that female plants require to persist within gynodioecious populations. Particularly valuable is determining the pollination success at the prezygotic stage in self-compatible gynodioecious species whose females do not exhibit enhanced seed production. In core and marginal populations of Daphne laureola, we analyzed the differences between hermaphrodites and females in the proportion of flowers visited, the stigma pollen loads, and the quantity of pollen tubes in styles. We also examined the relationship between the number of pollen tubes in styles vs. the number of pollen grains on stigmas using piecewise regression and binomial generalized linear mixed models. Pollinators deposited larger pollen loads on flowers in marginal populations. In marginal populations, female flowers received more pollinator visits and more pollen grains on their stigmas, and they had more pollen tubes in their styles than did female flowers in core populations. Both piecewise regression and binomial GLMM analyses showed that females in marginal populations had a lower proportion of grains that developed tubes than females in the core populations, which suggests decreased pollination quality. More efficient pollination services in marginal populations decreased the overall differences in the prezygotic pollination success between the sex morphs. Our results also suggest that pollination quality is lower in females of marginal populations, which could be counteracting the increased pollination in females in marginal populations.

The effect of disease on the evolution of females and the genetic basis of sex in populations with cytoplasmic male sterility.

The evolution of separate males and females is an important evolutionary transition that has occurred multiple times in flowering plants. While empirical studies have stressed the potential importance of natural enemies and organismal interactions in the evolution of separate sexes, there has been no treatment of natural enemies in the theoretical literature. We investigated the effects of disease on the evolution of females in gynodioecious populations composed of females and hermaphrodites, where sex is determined by the interaction of cytoplasmic male sterility (CMS) and nuclear restorer genes. When females are significantly more resistant than hermaphrodites, disease drives an increase in the frequency of females and sex determination becomes nuclear, creating the pre-conditions for the evolution of separate males and females. However, when females are only moderately more resistant, disease drives changes in the frequency of CMS and restorer alleles, but has little effect on the frequency of females. We discuss our results in the context of the evolution of mating systems and cyto-nuclear epistasis.

Sexual allocation in the gynodioecious species Cyananthus macrocalyx (Campanulaceae) at high elevations in the Sino-Himalaya Mountains

Sexual allocation pattern is considered highly relevant for the maintenance of females in the process of evolution from hermaphrodites to dioecy. Theoretical and empirical studies predict that gynodioecious plants should invest more resources in male function under harsher environments and/or when female frequency is higher; and that there are trade-offs between male and female function. We studied sexual allocation pattern in the gynodioecious species Cyananthus macrocalyx in two populations in SW China. The results showed that although the total flower mass in hermaphrodites was significantly higher than that in females, females and hermaphrodites allocated similar biomass to female function (pistil biomass). As a consequence, females allocated relatively more resource to female function, while hermaphrodites allocated relatively more resource to pollinator attractiveness. There was no difference in total flower mass, pistil mass, and pollen production in hermaphrodites between the two populations. These results suggest that the females compensate for the disadvantage of lacking male function by allocating relatively more resource to female function, while hermaphrodites gain fitness through both male and female functions. This study supports the idea that if females are to be maintained in a gynodioecious population, they must allocate more resource to female functions than is necessary for hermaphrodites. However, our results were in contrast to those of previous studies, in that trade-offs between male and female functions and male-biased allocation under high female frequency were not observed. This study adds to the body of research on plant sexual allocation, and for that it is, at least at some aspects, contrary to previous statements, it could be helpful for advancing the theoretical predictions in the future.