Search

Premise: Angiosperms range in sexual system from hermaphroditism through gynodioecy and androdioecy to dioecy. Trioecy, where females and males coexist with hermaphrodites, is rare. Recently, trioecy was documented in hexaploid populations of the wind‐pollinated herb Mercurialis annua in Spain. Methods: We surveyed the frequency of males, hermaphrodites, and females in M. annua across its distribution in the Iberian Peninsula, tracked sex‐ratio variation in several populations over consecutive generations, and assessed evidence for pollen limitation. In a common garden, we estimated male, female, and hermaphroditic fitness. We used controlled crosses to infer the genetic basis of male sterility. Finally, we compared predictions of a deterministic model with the distribution of observed sex ratios in the field based on our fitness estimates and the inferred genetics of sex determination. Results: Trioecy is widespread in Spanish and Portuguese populations of M. annua. Males are determined by a dominant (Y‐linked) allele, and female expression results from the interaction between cytoplasmic male sterility and multiple nuclear male sterility restorers partially linked to the male determiner. Male pollen production is approximately 12 times that of hermaphrodites, while female seed production is less than 1.12 times the observed hermaphroditic levels. The distribution of sex ratios in natural populations conforms with predictions of our deterministic simulations. Conclusions: Our study documents and accounts for a clear case of trioecy in which sex is determined by both maternally and biparentally inherited genes. [ABSTRACT FROM AUTHOR]

Life history is defined by traits that reflect key components of fitness, especially those relating to reproduction and survival. Research in life history seeks to unravel the relationships among these traits and understand how life history strategies evolve to maximize fitness. As such, life history research integrates the study of the genetic and developmental mechanisms underlying trait determination with the evolutionary and ecological context of Darwinian fitness. As a leading model organism for molecular and developmental genetics, Caenorhabditis elegans is unmatched in the characterization of life history-related processes, including developmental timing and plasticity, reproductive behaviors, sex determination, stress tolerance, and aging. Building on recent studies of natural populations and ecology, the combination of C. elegans ' historical research strengths with new insights into trait variation now positions it as a uniquely valuable model for life history research. In this review, we summarize the contributions of C. elegans and related species to life history and its evolution. We begin by reviewing the key characteristics of C. elegans life history, with an emphasis on its distinctive reproductive strategies and notable life cycle plasticity. Next, we explore intraspecific variation in life history traits and its underlying genetic architecture. Finally, we provide an overview of how C. elegans has guided research on major life history transitions both within the genus Caenorhabditis and across the broader phylum Nematoda. While C. elegans is relatively new to life history research, significant progress has been made by leveraging its distinctive biological traits, establishing it as a highly cross-disciplinary system for life history studies. [ABSTRACT FROM AUTHOR]

Abstract Background The MADS-box gene family is widely distributed in the plant kingdom, and its members typically encoding transcription factors to regulate various aspects of plant growth and development. In particular, the MIKC-type MADS-box genes play a crucial role in the determination of floral organ development and identity recognition. As a type of androdioecy plant, Chionanthus retusus have unique gender differentiation. Manifested as male individuals with only male flowers and female individuals with only bisexual flowers. However, due to the lack of reference genome information, the characteristics of MIKC-type MADS-box genes in C. retusus and its role in gender differentiation of C. retusus remain largely unknown. Therefore, it is necessary to identify and characterize the MADS-box gene family within the genome of the C. retusus. Results In this study, we performed a genome-wide identification and analysis of MIKC-type MADS-box genes in C. retusus (2n = 2x = 46), utilizing the latest reference genome, and studied its expression pattern in individuals of different genders. As a result, we identified a total of 61 MIKC-type MADS-box genes in C. retusus. 61 MIKC-type MADS-box genes can be divided into 12 subfamilies and distributed on 18 chromosomes. Genome collinearity analysis revealed their conservation in evolution, while gene structure, domains and motif analysis indicated their conservation in structure. Finally, based on their expression patterns in floral organs of different sexes, we have identified that CrMADS45 and CrMADS60 may potentially be involved in the gender differentiation of C. retusus. Conclusions Our studies have provided a general understanding of the conservation and characteristics of the MIKC-type MADS-box genes family in C. retusus. And it has been demonstrated that members of the AG subfamily, CrMADS45 and CrMADS60, may play important roles in the gender differentiation of C. retusus. This provides a reference for future breeding efforts to improve flower types in C. retusus and further investigate the role of MIKC-type MADS-box genes in gender differentiation.

Background: The MADS-box gene family is widely distributed in the plant kingdom, and its members typically encoding transcription factors to regulate various aspects of plant growth and development. In particular, the MIKC-type MADS-box genes play a crucial role in the determination of floral organ development and identity recognition. As a type of androdioecy plant, Chionanthus retusus have unique gender differentiation. Manifested as male individuals with only male flowers and female individuals with only bisexual flowers. However, due to the lack of reference genome information, the characteristics of MIKC-type MADS-box genes in C. retusus and its role in gender differentiation of C. retusus remain largely unknown. Therefore, it is necessary to identify and characterize the MADS-box gene family within the genome of the C. retusus. Results: In this study, we performed a genome-wide identification and analysis of MIKC-type MADS-box genes in C. retusus (2n = 2x = 46), utilizing the latest reference genome, and studied its expression pattern in individuals of different genders. As a result, we identified a total of 61 MIKC-type MADS-box genes in C. retusus. 61 MIKC-type MADS-box genes can be divided into 12 subfamilies and distributed on 18 chromosomes. Genome collinearity analysis revealed their conservation in evolution, while gene structure, domains and motif analysis indicated their conservation in structure. Finally, based on their expression patterns in floral organs of different sexes, we have identified that CrMADS45 and CrMADS60 may potentially be involved in the gender differentiation of C. retusus. Conclusions: Our studies have provided a general understanding of the conservation and characteristics of the MIKC-type MADS-box genes family in C. retusus. And it has been demonstrated that members of the AG subfamily, CrMADS45 and CrMADS60, may play important roles in the gender differentiation of C. retusus. This provides a reference for future breeding efforts to improve flower types in C. retusus and further investigate the role of MIKC-type MADS-box genes in gender differentiation. [ABSTRACT FROM AUTHOR]

In angiosperms cytoplasmic DNA is typically passed on maternally through ovules. Genes in the mtDNA may cause male sterility. When male-sterile (female) cytotypes produce more seeds than cosexuals, they pass on more copies of their mtDNA and will co-occur with cosexuals with a neutral cytotype. Cytoplasmic gynodioecy is a well-known phenomenon in angiosperms, both in wild and crop plants. In some conifer families (e.g. Pinaceae) mitochondria are also maternally inherited. However in some other families (e.g. Taxaceae and Cupressaceae) mtDNA is paternally inherited through the pollen. With paternal mtDNA inheritance, male cytotypes that produce more pollen than cosexuals are expected to co-occur with cosexuals. This is uncharted territory. An ESS model shows that the presence of male cytotypes selects for more female allocation in the cosexual, i.e. for sexual specialisation. An allele that switches sex from male to female can then invade. This leads to rapid loss of the neutral cytotype of the cosexual, fixation of the male cytotype and dioecy with 50% males and 50% females. The models suggest that paternal inheritance of mtDNA facilitates the evolution dioecy. Consistent with this hypothesis the Pinaceae are 100% monoecious, while dioecy is common in the Taxaceae family and in the genus Juniperus (Cupressaceae). However, no reliable data are yet available on both mode of inheritance of mtDNA and gender variation of the same species. When cosexuals benefit from reproductive assurance (high selfing rate, low inbreeding depression, low fertilisation) they maintain themselves next to males and females. This predicted pattern with three sex types present in the same population is observed in conifers in nature. [ABSTRACT FROM AUTHOR]

Background and Aims Androdioecy, the co-occurrence of males and hermaphrodites, is a rare reproductive system. Males can be maintained if they benefit from a higher male fitness than hermaphrodites, referred to as male advantage. Male advantage can emerge from increased fertility owing to resource reallocation. However, empirical studies usually compare sexual phenotypes over a single flowering season, thus ignoring potential cumulative effects over successive seasons in perennials. In this study, we quantify various components of male fertility advantage, both within and between seasons, in the long-lived perennial shrub Phillyrea angustifolia (Oleaceae). Although, owing to a peculiar diallelic self-incompatibility system and female sterility mutation strictly associated with a breakdown of incompatibility, males do not need fertility advantage to persist in this species, this advantage remains an important determinant of their equilibrium frequency. Methods A survey of >1000 full-sib plants allowed us to compare males and hermaphrodites for several components of male fertility. Individuals were characterized for proxies of pollen production and vegetative growth. By analysing maternal progeny, we compared the siring success of males and hermaphrodites. Finally, using a multistate capture–recapture model we assessed, for each sexual morph, how the intensity of flowering in one year impacts next-year growth and reproduction. Key Results Males benefitted from a greater vegetative growth and flowering intensity. Within one season, males sired twice as many seeds as equidistant, compatible hermaphroditic competitors. In addition, males more often maintained intense flowering over successive years. Finally, investment in male reproductive function appeared to differ between the two incompatibility groups of hermaphrodites. Conclusion Males, by sparing the cost of female reproduction, have a higher flowering frequency and vegetative growth, both of which contribute to male advantage over an individual lifetime. This suggests that studies analysing sexual phenotypes during only single reproductive periods are likely to provide inadequate estimates of male advantage in perennials. [ABSTRACT FROM AUTHOR]

In dioecious populations, males and females may evolve different trait values to increase fitness through their respective sexual functions. Because hermaphrodites express both sexual functions, resolving sexual conflict is potentially more difficult for them. Here, we show that hermaphrodite plants can partially resolve sexual conflict by expressing different trait values in different male and female modules (e.g. different flowers, inflorescences, branches etc.). We analysed the flowering phenology, sex allocation and selection gradients on floral traits of flowers of the andromonoecious plant Pulsatilla alpina, which produces both bisexual and male flowers. Our results indicate that strong protogyny prevents early bisexual flowers from profiting from high siring opportunities early in the reproductive season at a time when male flowers could achieve high siring success. The production of unisexual male flowers thus resolves this sexual conflict because it allows the flowers to express their male function without waiting until after the female function has been performed. Our study illustrates the resolution of sexual conflict arising from phenological constraints via modular divergence in sex allocation. We discuss the extent to which modular variation in sex allocation in the context of other sexual systems may be similarly explained. [ABSTRACT FROM AUTHOR]

Sweet osmanthus is an androdioecious plant; however, the mechanism underlying pistil sterility in male plants is still unclear. Scanning electron microscopy showed that the structure of pollen grains in the stamens does not differ between the sterile cultivar 'Chenghong Dangui' and the fertile cultivar 'Huangchuan Jingui'. Triphenyltetrazolium chloride and fluorescein diacetate staining as well as in vitro culture experiments revealed that pollen grains were active in both cultivars, indicating that the stamens in both 'Chenghong Dangui' and 'Huangchuan Jingui' could develop normally. When the pistils of the fertile cultivar 'Huangchuan Jingui' differentiated, two protrusions formed on the inner side of the stamen primordium, and these gradually developed and fused together to form the ovary, style, and stigma. The pistil of the sterile cultivar 'Chenghong Dangui' also formed two protrusions on the inner side of the stamen during differentiation; however, instead of fusing, two fronds were formed. These results suggest that male sweet osmanthus are formed due to the abortion of pistils during the development of floral organs. Transcriptome sequencing revealed that the expression levels of carpel development gene CRC, AG, and AGL11 were significantly lower in 'Chenghong Dangui' compared with 'Huangchuan Jingui' at different flowering stages, which provide new insight in the molecular mechanism of pistil abortion in 'Chenghong Dangui'. CRC and AG may regulate each other to promote carpel development.

Several examples of androdioecy appear to have evolved from dioecy and have low male frequency (< 0.5). However, the evolutionary pathway to androdioecy in Oleaceae may come from hermaphroditism. Osmanthus fragrans L. has a 1:1 sex ratio in nature populations. Significant differences are observed not only in flowering phenology but also in some floral traits between males and hermaphrodites. The protandry in the same population and the protogyny in the same plant may promote the xenogamy between genders. The majority of flower traits related with the pollen production are different between males and hermaphrodites. Males bear more flowering nodes, and more flowers per node, and larger anther in all three populations. This characteristic demonstrated that males have more male advantage than hermaphrodites. Population genetic structure of O. fragrans is genetically homogeneous at the species level, and most variations exist within a population. The percentage of variation among populations (13%) and between males and hermaphrodites (0%) is low. Moreover, genetic differentiation was very low between genders not only among populations but also in the same population. This genetic variation could be attributed to the occurrence of high levels of xenogamy between genders. Therefore, high male frequency and more male fitness advantage in males are the essential conditions for this mating system, which plays an important role during population reproduction and regeneration. The 1:1 sex ratio could be the result of integrative effects of sexual system, mating system, and reproductive success.

Abstract Background The fragrant flower plant Osmanthus fragrans has an extremely rare androdioecious breeding system displaying the occurrence of males and hermaphrodites in a single population, which occupies a crucial intermediate stage in the evolutionary transition between hermaphroditism and dioecy. However, the molecular mechanism of androdioecy plant is very limited and still largely unknown. Results Here, we used SWATH-MS-based quantitative approach to study the proteome changes between male and hermaphroditic O. fragrans pistils. A total of 428 proteins of diverse functions were determined to show significant abundance changes including 210 up-regulated and 218 down-regulated proteins in male compared to hermaphroditic pistils. Functional categorization revealed that the differentially expressed proteins (DEPs) primarily distributed in the carbohydrate metabolism, secondary metabolism as well as signaling cascades. Further experimental analysis showed the substantial carbohydrates accumulation associated with promoted net photosynthetic rate and water use efficiency were observed in purplish red pedicel of hermaphroditic flower compared with green pedicel of male flower, implicating glucose metabolism serves as nutritional modulator for the differentiation of male and hermaphroditic flower. Meanwhile, the entire upregulation of secondary metabolism including flavonoids, isoprenoids and lignins seem to protect and maintain the male function in male flowers, well explaining important feature of androdioecy that aborted pistil of a male flower still has a male function. Furthermore, nine selected DEPs were validated via gene expression analysis, suggesting an extra layer of post-transcriptional regulation occurs during O. fragrans floral development. Conclusion Taken together, our findings represent the first SWATH-MS-based proteomic report in androdioecy plant O. fragrans, which reveal carbohydrate metabolism, secondary metabolism and post-transcriptional regulation contributing to the androdioecy breeding system and ultimately extend our understanding on genetic basis as well as the industrialization development of O. fragrans.

Androdioecy is the crucial transition state in the evolutionary direction of hermaphroditism to dioecy, however, the molecular mechanisms underlying the formation of this sex system remain unclear. While popular in China for its ornamental and cultural value, Osmanthus fragrans has an extremely rare androdioecy breeding system, meaning that there are both male and hermaphroditic plants in a population. To unravel the mechanisms underlying the formation of androdioecy, we performed small RNA sequencing studies on male and hermaphroditic O. fragrans. A total of 334 miRNAs were identified, of which 59 were differentially expressed. Functional categorization revealed that the target genes of differentially expressed miRNAs were mainly involved in the biological processes of reproductive development and the hormone signal transduction pathway. We speculated that the miRNA160, miRNA167, miRNA393 and miRNA396 families may influence the sex differentiation in O. fragrans. Overall, our study is the first exploration of miRNAs in the growth and development process of O. fragrans, and is also the first study of androdioecious plants from the miRNA sequencing perspective. The analysis of miRNAs and target genes that may be involved in the sex differentiation process lay a foundation for the ultimate discovery of the androdioecious molecular mechanism in O. fragrans. [ABSTRACT FROM AUTHOR]

Intraspecific structural variations in pollen grains are widespread in angiosperms. The aims of the present review are to track down intraspecific variability in various pollen morphological features, and to evaluate their taxonomic significance in defining species boundaries. Shape of pollen grains, sculptures, aperture types and numbers, muri textures, presence/absence of opercula, folded ectexines and pollen corpus (in bisaccate pollen grains) were encountered to exhibit intraspecific variations in the various taxa examined. These structural characteristics were recognized to be associated with variations of evolutionary effects, in the spatial arrangements and structural diversity of the styles (heterostyly, enantiostyly, stigma-height dimorphism, inverostyly), anthers (heteranthery), stigma textures, and the reproductive (breeding) systems (androdioecy, dioecy, andromonoecy) of the flowers. Other internal (polyploidy) and external factors (environmental) linked with structural intraspecific pollen variations were also considered. In order to characterize intraspecific boundaries and to test the credibility of these characters, various statistical tools were surveyed to identify characters of diagnostic taxonomic values. • Various pollen morphological features witnessed intraspecific variations. • Floral/sexual diversity represented main stimulants of intraspecific pollen variations. • Pollinator-mediated selection, polyploidy, environment, are main causes of variations. • Intraspecific pollen variation restricts their perception as of taxonomic value. • Appropriate statistical tools to explore taxonomic relationships should be endorsed. [ABSTRACT FROM AUTHOR]

The plant genus Mercurialis includes dioecious, monoecious and androdioecious species (where males coexist with hermaphrodites). Its diversification involved reticulate evolution via hybridization and polyploidization. The Y chromosome of the diploid species Mercurialis annua shows only mild signs of degeneration. We used sequence variation at a Y-linked locus in several species and at multiple autosomal and pseudoautosomal loci to investigate the origin and evolution of the Y chromosome across the genus. Our study provides evidence for further cases of allopolyploid speciation. It also reveals that all lineages with separate sexes (with one possible exception) share the same ancestral Y chromosome. Surprisingly, males in androdioecious populations of hexaploid M. annua carry a Y chromosome that is not derived from either of its two putative progenitor lineages but from a more distantly related perennial dioecious lineage via introgression. These results throw new light on the evolution of sexual systems and polyploidy in Mercurialis and secure it as a promising model for further study of plant sex chromosomes. This article is part of the theme issue 'Sex determination and sex chromosome evolution in land plants'. [ABSTRACT FROM AUTHOR]

Androdioecy is a rare mating system in which males and self‐compatible hermaphrodites coexist. Such a system is proposed to be a temporary evolutionary transition between separate sexes and total hermaphroditism. Sexual conflict over occurrence of mating has been proposed as one crucial precondition for a behavioral mechanism that can maintain stable androdioecy. This work tests this hypothesis using Eulimnadia dahli, a species of clam shrimp that has maintained androdioecy for millions of years. Monitoring the behaviors of mating pairs, we examined the differences in antagonistic behaviors and propensity to mate between different types of hermaphrodites and their male counterparts. Contrary to predictions, inbred hermaphrodites from male‐less populations did not have higher resistance to mating with males, were not found to be less attractive to males, and showed no increased tendency to self‐fertilize than hermaphrodites from mixed populations. [ABSTRACT FROM AUTHOR]

Androdioecy is a rare reproductive system. Fraxinus platypoda, a woody canopy species in Japan's mountainous riparian zones, is described as a morphologically androdioecious species. In this study, we tried to detect whether F. platypoda is also functionally androdioecious. We analyzed its sexual expression, seed development, pollen morphology and germination ability, pollination systems, and mast flowering behavior. We found that the hermaphrodite trees are andromonoecious, with inflorescences bearing male and hermaphroditic flowers, whereas male individuals had only male flowers. Pollen morphology was identical in male flowers, in hermaphrodite flowers of an andromonoecious individual, and in male flowers of male individuals. Pollen from both types of individuals was capable of germination both ex vivo (on nutrient medium) and in vivo in pollination experiments. However, compared with pollen from andromonoecious trees, pollen from male trees showed a higher germination rate. The self-pollination rate of bagged hermaphroditic flowers was almost zero. The fruit set rate following cross-pollination with male pollen from a male tree was higher than that following natural pollination, whereas the rate with hermaphroditic pollen was the same. The flowering and fruiting of F. platypoda have fluctuated over 17 years; the flowering of the two types of sexual individuals exhibited clear synchronization during this period. The frequency of male individuals within the populations is 50%. The maintenance of such a proportion of males in populations of the self-incompatible F. platypoda is either indicative of a true androdioecious species with a diallelic self-incompatibility system or a cryptic-dioecious species. This alternative is discussed here. [ABSTRACT FROM AUTHOR]

Androdioecy (the coexistence of males and hermaphrodites) is a rare mating system for which the evolutionary dynamics are poorly understood. Here, we investigate the cottony cushion scale, Icerya purchasi, one of only three reported cases of androdioecy in insects. In this species, female‐like hermaphrodites have been shown to produce sperm and self‐fertilize. However, males are ocassionally observed as well. In a large genetic analysis, we show for the first time that, although self‐fertilization appears to be the primary mode of reproduction, rare outbreeding events do occur in natural populations, supporting the hypothesis that hermaphrodites mate with males and hence androdioecy is the mating system of I. purchasi. Thus, this globally invasive pest insect appears to enjoy the colonization advantages of a selfing organism while also benefitting from periodic reintroduction of genetic variation through outbreeding with males. [ABSTRACT FROM AUTHOR]

Background: The fragrant flower plant Osmanthus fragrans has an extremely rare androdioecious breeding system displaying the occurrence of males and hermaphrodites in a single population, which occupies a crucial intermediate stage in the evolutionary transition between hermaphroditism and dioecy. However, the molecular mechanism of androdioecy plant is very limited and still largely unknown. Results: Here, we used SWATH-MS-based quantitative approach to study the proteome changes between male and hermaphroditic O. fragrans pistils. A total of 428 proteins of diverse functions were determined to show significant abundance changes including 210 up-regulated and 218 down-regulated proteins in male compared to hermaphroditic pistils. Functional categorization revealed that the differentially expressed proteins (DEPs) primarily distributed in the carbohydrate metabolism, secondary metabolism as well as signaling cascades. Further experimental analysis showed the substantial carbohydrates accumulation associated with promoted net photosynthetic rate and water use efficiency were observed in purplish red pedicel of hermaphroditic flower compared with green pedicel of male flower, implicating glucose metabolism serves as nutritional modulator for the differentiation of male and hermaphroditic flower. Meanwhile, the entire upregulation of secondary metabolism including flavonoids, isoprenoids and lignins seem to protect and maintain the male function in male flowers, well explaining important feature of androdioecy that aborted pistil of a male flower still has a male function. Furthermore, nine selected DEPs were validated via gene expression analysis, suggesting an extra layer of post-transcriptional regulation occurs during O. fragrans floral development. Conclusion: Taken together, our findings represent the first SWATH-MS-based proteomic report in androdioecy plant O. fragrans, which reveal carbohydrate metabolism, secondary metabolism and post-transcriptional regulation contributing to the androdioecy breeding system and ultimately extend our understanding on genetic basis as well as the industrialization development of O. fragrans. [ABSTRACT FROM AUTHOR]

In dioecious plants, males and females frequently show 'leaky' sex expression, with individuals occasionally producing flowers of the opposite sex. This leaky sex expression may have enabled the colonization of oceanic islands by dioecious plant species, and it is likely to represent the sort of variation upon which selection acts to bring about evolutionary transitions from dioecy to hermaphroditism. Although leakiness is commonly reported for dioecious species, it is not known whether it has plastic component. The question is interesting because males or females with an ability to enhance their leakiness plastically in the absence of mates would have an advantage of being able to produce progeny by self‐fertilization. Here, we demonstrate that leaky sex expression in the wind‐pollinated dioecious herb Mercurialis annua is plastically responsive to its mating context. We compared experimental populations of females growing either with or without males. Females growing in the absence of males were leakier in their sex expression than controls growing with males, producing more than twice as many male flowers. Our results thus provide a striking instance of plasticity in the reproductive behaviour of plants that is likely adaptive. We consider how females might sense their mating environment as a function of pollen availability, and we discuss possible constraints on the evolution of plasticity in sex expression when the environmental signals that individuals receive are unreliable. [ABSTRACT FROM AUTHOR]

Fertilization assurance (Baker's contention) and multilocus coadaptation (Allard's argument) are two distinct hypotheses for the adaptive significance of self-fertilization in hermaphroditic taxa, and both scenarios have been invoked to rationalize isogenicity via incest in various plants and invertebrate animals with predominant selfing. Here we contrast Allard's argument and Baker's contention as applied to the world's only known vertebrate that routinely self-fertilizes. We pay special attention to frequencies of locally most common multilocus genotypes in Floridian populations of the Mangrove Rivulus (Kryptolebias marmoratus). Isogenicity patterns in this fish appear inconsistent with Allard's argument, thus leaving Baker's contention as the more plausible scenario (a result also supported by natural history information for this species). These results contrast with the isogenicity patterns and conclusions previously drawn from several self-fertilizing plants and invertebrate animal species. Thus, the adaptive significance of selfing apparently varies across hermaphroditic taxa.

Androdioecy is a rare reproductive system. Fraxinus platypoda, a woody canopy species in Japan’s mountainous riparian zones, is described as a morphologically androdioecious species. In this study, we tried to detect whether F. platypoda is also functionally androdioecious. We analyzed its sexual expression, seed development, pollen morphology and germination ability, pollination systems, and mast flowering behavior. We found that the hermaphrodite trees are andromonoecious, with inflorescences bearing male and hermaphroditic flowers, whereas male individuals had only male flowers. Pollen morphology was identical in male flowers, in hermaphrodite flowers of an andromonoecious individual, and in male flowers of male individuals. Pollen from both types of individuals was capable of germination both ex vivo (on nutrient medium) and in vivo in pollination experiments. However, compared with pollen from andromonoecious trees, pollen from male trees showed a higher germination rate. The self-pollination rate of bagged hermaphroditic flowers was almost zero. The fruit set rate following cross-pollination with male pollen from a male tree was higher than that following natural pollination, whereas the rate with hermaphroditic pollen was the same. The flowering and fruiting of F. platypoda have fluctuated over 17 years; the flowering of the two types of sexual individuals exhibited clear synchronization during this period. The frequency of male individuals within the populations is 50%. The maintenance of such a proportion of males in populations of the self-incompatible F. platypoda is either indicative of a true androdioecious species with a diallelic self-incompatibility system or a cryptic-dioecious species. This alternative is discussed here.

Individual plants can produce leaves that differ substantially in size, morphology and many other traits. However, leaves that play a specific role in reproduction have rarely been reported. Here, we report leaves specialized to enclose fruit clusters and enhance seed production in an annual vine, Schizopepon bryoniifolius. Enclosure leaves were produced at the end of the growing season in late autumn. They were different in greenness and structure from other leaves. Under solar radiation, the ambient temperature inside an intact enclosure was up to 4.6°C higher than that near a fruit cluster whose enclosure leaves had been removed. We found that enclosures were thicker at colder sites. Removal of enclosing leaves negatively affected fruit survival and/or growth, but we could not identify the exact mechanism. The results suggested that enclosures allow the plant to produce seeds under the cold weather the plant encounters at the end of its life. Vegetative and reproductive traits of plants have usually been studied separately. This study indicates how they can dynamically interact, as shown by an examination of associations among leaf and reproductive trait changes according to life stages. [ABSTRACT FROM AUTHOR]

The gooseneck barnacle Octolasmis warwickii has a rare sexual system called androdioecy, in which hermaphrodites and dwarf males co-occur. It has been hypothesized that dwarf males can coexist with conspecific hermaphrodites when dwarf males are capable of leaving more offspring than hermaphrodites via male reproduction. This hypothesis of reproductive superiority of dwarf males can be validated by comparing the reproductive success between dwarf males and hermaphrodites through DNA marker-based parentage testing. In the present study, we developed microsatellite DNA markers for O. warwickii, and evaluated the power of these markers to infer parentage based on simulation analysis. Using next generation sequencing, we obtained 344 microsatellite sequences suitable for designing primer sets for amplification in polymerase chain reaction (PCR). Of these, we examined the PCR amplification efficiency of 54 primer sets, of which 11 passed our primer screening in a population sample (n = 35). The developed markers exhibited moderate to high levels of polymorphisms, and met Hardy–Weinberg equilibrium with little evidence of significant allelic association to each other. Our simulated paternity inference suggested that the combinational use of the markers allows a high resolution of parentage (success rate of > 99.9%) if all candidate fathers are available. [ABSTRACT FROM AUTHOR]

We found functionally male individuals in an otherwise hermaphroditic population of Tulipa pumila (Liliaceae) located in Tuscany (central Italy). We investigated the sex ratio of this population, followed by morphometric analyses of the scape, leaves and flowers, and tests on pollen germinability and seed number and mass, in order to infer which sexual strategy produced the observed co‐occurrence of male and hermaphrodite individuals. We found that sex ratio deviated from 1:1, and functionally male individuals showed a smaller plant size and a reduced pollen fitness (germinability and siring ability) compared to hermaphrodites. These findings point to a resource‐dependent sexual allocation strategy, probably associated with gender diphasy. [ABSTRACT FROM AUTHOR]

Among vertebrate animals, only the mangrove rivulus (Kryptolebias marmoratus) was known to self-fertilize. Here, we use microsatellite analyses to document a high selfing rate (97%) in a related nominal species, Kryptolebias ocellatus, which likewise is androdioecious (populations consist of males and hermaphrodites). In contrast, we find no evidence of self-fertilization in Kryptolebias caudomarginatus (an androdioecious species closely related to the marmoratus-ocellatus clade) or in Kryptolebias brasiliensis (a dioecious outgroup). These findings indicate that the initiation of self-fertilization predated the origin of the marmoratus-ocellatus clade. From mitochondrial DNA sequences and microsatellite data, we document a substantial genetic distance between Kryptolebias marmoratus and K. ocellatus, implying that the selfing capacity has persisted in these fishes for at least several hundred thousand years.

The diversity in sperm shape and size represents a powerful paradigm to understand how selection drives the evolutionary diversification of cell morphology. Experimental work on the sperm biology of the male-hermaphrodite nematode Caenorhabditis elegans has elucidated diverse factors important for sperm fertilization success, including the competitive superiority of larger sperm. Yet despite extensive research, the molecular mechanisms regulating C. elegans sperm size and the genetic basis underlying natural variation in sperm size remain unknown. To address these questions, we quantified male sperm size variation of a worldwide panel of 97 genetically distinct C. elegans strains, allowing us to uncover significant genetic variation in male sperm size. Aiming to characterize the molecular genetic basis of C. elegans male sperm size variation using a genome-wide association study, we did not detect any significant quantitative trait loci. We therefore focused on the genetic analysis of pronounced sperm size differences observed between recently diverged laboratory strains (N2 vs. LSJ1/2). Using mutants and quantitative complementation tests, we demonstrate that variation in the gene nurf-1 underlies the evolution of small sperm in the LSJ lineage. Given the previous discovery that this same nurf-1 variation was central for hermaphrodite laboratory adaptation, the evolution of reduced male sperm size in LSJ strains likely reflects a pleiotropic consequence. Together, our results provide a comprehensive quantification of natural variation in C. elegans sperm size and first insights into the genetic determinants of Caenorhabditis sperm size, pointing at an involvement of the NURF chromatin remodeling complex. [ABSTRACT FROM AUTHOR]

Themaintenance of males at intermediate frequencies is an important evolutionary problem. Several species of Caenorhabditis nematodes have evolved a mating system in which selfing hermaphrodites and males coexist. While selfing produces XX hermaphrodites, crossfertilization produces 50% XO male progeny. Thus, male mating success dictates the sex ratio. Here, we focus on the contribution of the male secreted short (mss) gene family to male mating success, sex ratio, and population growth. The mss family is essential for sperm competitiveness in gonochoristic species, but has been lost in parallel in androdioecious species. Using a transgene to restore mss function to the androdioecious Caenorhabditis briggsae, we examined how mating system and population subdivision influence the fitness of the mss+ genotype. Consistent with theoretical expectations, when mss+ and mss-null (i.e., wild type) genotypes compete, mss+ is positively selected in both mixed-mating and strictly outcrossing situations, though more strongly in the latter. Thus, while sexual mode alone affects the fitness of mss+, it is insufficient to explain its parallel loss. However, in genetically homogenous androdioecious populations, mss+ both increases male frequency and depresses population growth. We propose that the lack of inbreeding depression and the strong subdivision that characterize natural Caenorhabditis populations impose selection on sex ratio that makes loss of mss adaptive after self-fertility evolves. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Ephedranthus S. Moore is a Neotropical genus of Annonaceae distributed from Colombia and the Guianas to Bolivia and southeastern Brazil, comprised of seven species: E. amazonicus R. E. Fr., E. boliviensis Chatrou & Pirie, E. colombianus Maas & Setten, E. dimerus J. C. Lopes, Chatrou & Mello-Silva, E. guianensis R. E. Fr., E. parviflorus S. Moore, and E. pisocarpus R. E. Fr. The genus is recognized by the combination of leaves with impressed adaxial veins, short pedicels, two to five bracts, androdioecy (i.e., some plants with only staminate flowers, other plants with only bisexual flowers), and 1-seeded monocarps. This revision updates the knowledge about the genus, providing a morphological phylogenetic analysis of Ephedranthus species, with a key, descriptions, distribution maps, and illustrations of the diagnostic characters of the species. The phylogenetic analysis shows Ephedranthus defined by having staminate flowers and basal placentation, and the seven species related as ((E. colombianus, E. boliviensis) ((E. guianensis, E. amazonicus) (E. dimerus (E. parviflorus, E. pisocarpus)))); based on this, the current subdivision of Ephedranthus is not monophyletic, and the sections are here synonymized. [ABSTRACT FROM AUTHOR]

Summary: A new species of a predator aphelenchoidid, Seinura caverna n. sp., is described and illustrated. The new species was isolated from bat guano collected from a calcareous cave in Japan. The new species is characterised by its three-lined lateral field, secretory-excretory pore at the level of the posterior two-thirds of the metacorpus, a long post-uterine sac, hermaphrodite tail shape elongate conoid with a filiform tip, and an androdioecious reproductive mode. The new species is typologically and biologically close to S. steineri , but is distinguished by its longer post-uterine sac, slightly longer stylet and slightly larger median bulb. The comparisons with other morphologically similar species, i.e. , S. chertkovi , S. tenuicaudata and S. steineri , are discussed. A molecular phylogenetic analysis based on two small and large subunit ribosomal RNA genes revealed that the new species is located at the basal position of clade 3 of the Aphelenchoididae, clearly separate from S. demani , its congener with unclear rectum and anus, which is located at the derived position, suggesting that further generic revision is necessary for the genus. Aphelenchoides lii n. comb. (= Seinura lii) is proposed. [ABSTRACT FROM AUTHOR]

Androdioecy (males and hermaphrodites) is a rare breeding system in multicellular organisms, found mostly in barnacles and branchiopod crustaceans. The most speciose and longest-lived androdioecious clade is the genus Eulimnadia Packard, 1874 (Branchiopoda, Spinicaudata), the clam shrimps, consisting of over 50 species that have maintained androdioecy for an estimated 24–180 million years. Many populations of Eulimnadia nevertheless comprise entirely "monogenic" hermaphrodites. Hypotheses proposed to explain the relative stability of androdioecy (sexual conflict, overdominance, and metapopulation model) differ in their predictions of the resistance of existing all-hermaphrodite populations to invasion of males and hermaphrodites. We tested whether all-hermaphroditic populations of Eulimnadia texana Packard, 1871 may be resistant to male invasion by adding males and "amphigenic" hermaphrodites to all-hermaphrodite, monogenic populations that have been inbred for eight generations. All-hermaphrodite populations of E. texana that have been selfing for multiple generations are easily invaded by males, both directly and indirectly. The addition of males also increased the productivity of these experimental treatments, suggesting a selective benefit to outcrossing and thus to males. These results do not align with the sexual conflict nor the overdominance models, but are consistent with the metapopulation model of the maintenance of androdioecy. [ABSTRACT FROM AUTHOR]

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]

The exotic mangrove species Laguncularia racemosa (Combretaceae) is fast-growing and was used for forestation in recent years along the southeast coast of China. The breeding system of L. racemosa is variable among populations, such as hermaphroditism, gynodioecy, and androdioecy. To determine whether androdioecy is widespread in L. racemosa, 19 planted populations were surveyed along the southeast coast of China. To determine whether local environmental factors could affect the sex ratio in androdioecious populations, the observed male frequency of different populations was compared to local average annual temperature, rainfall, and salinity. The results showed that the 19 L. racemosa populations along the southeast coast of China were androdioecious. The male frequencies of these populations varied from 31.0 to  88.9%. Partial correlation analysis showed that average annual salinity explained 74.7% of the male frequency (p = 0.001). It is reasonable to note that the male frequency followed a general trend, presenting peak that coincided with the low salinity. The average annual rainfall explained only 30.4% of the male frequency (p = 0.403). And the average annual temperature explained only 20.2% of the male frequency (p = 0.206). The variable male frequency in different androdioecious L. racemosa populations may presumably be caused by ecological or genetical processes; these hypotheses will be tested in future studies. [ABSTRACT FROM AUTHOR]

Recent studies have shown that the number of individuals that can mate with each other (the mating group size, MGS) is a major driving force in the evolution of sexual systems in animals. Androdioecy is an important yet little-studied sexual system linking dioecy and hermaphroditism. Here, the individual sex allocation of hermaphrodites carrying dwarf males in the androdioecious barnacle Scalpellum scalpellum was studied. In accord with theoretical models, it was predicted that solitarily settled hermaphrodites have (1) a lower allocation of own male functions (estimated from testis size and penis length) but (2) instead a higher allocation of own female functions (estimated from embryo numbers) with (3) more dwarf males attached as a result of their lower MGS. As predicted, solitary hermaphrodites had relatively smaller testes and shorter penises than gregarious hermaphrodites of a similar size. This indicates that the solitary hermaphrodites can adaptively adjust sex allocation. Contrary to our prediction, these did not have larger brood sizes or carried significantly more males than gregarious hermaphrodites, although the probability of having any dwarf males present was significantly higher. We relate our results to sex allocation theory and the evolutionary maintenance of androdioecy in the species. [ABSTRACT FROM AUTHOR]

Mating behaviour between a dwarf male and its hermaphrodite partner was observed for the first time in cirripedes using the androdioecious barnacle Scalpellum scalpellum. Mating between hermaphrodites was also observed. The dwarf males are located on the rim of the mantle cavity of the hermaphrodite partner. When mating, the male extends the penis, which is four times longer than its body. The penis first assumes a straight stance where it is waved around in a searching mode. Upon touching the cirri of the hermaphrodite, the penis and the cirri engage in prolonged contact during which hermaphrodite feeding is suspended. Thereafter the penis assumes a U-bend to reach into the brood chamber, where after the mantle valves are closed tightly around the penis. The nearly transparent penis is a tube of very thin cuticle, equipped with pairs of side branches but not containing any visible tissue. The penis enables the minute male, situated outside the brood chamber, to securely deposit sperm into its partner. Adjacently situated hermaphrodites interact socially in between feeding sessions by reorienting themselves on the peduncle to touch each other with their cirri. This can be followed by precopulatory behaviour, where one or both individuals extend the penis to touch their partner, leading again to actual copulation where the penis of one individual is inserted into the other. We discuss the results in the context of the diverse reproductive strategies found in cirripede barnacles. [ABSTRACT FROM AUTHOR]

Androdioecy (co-existence of hermaphrodites and dwarf males) is a fascinating yet poorly understood phenomenon. The pedunculated barnacle Scalpellum scalpellum is an emerging model species for the system. In S. scalpellum, dwarf males and hermaphrodites are very different in adult morphology (e.g., in feeding structures and reproductive organs), but they share the same larval development with nauplii followed by cypris larvae. Recently, it was found that S. scalpellum cypris larvae display both genetic and environmental sex determination, but no detailed morphological study has yet investigated how the settled cypris larvae differ subsequent to settlement. This study investigates the morphological aspects of the onset of sex determination in the cyprids of S. scalpellum by examining their metamorphosis into either dwarf males or hermaphrodites under laboratory conditions. This study emphasizes morphological differences, such as size and shape of primordial shell plates, development of a flexible peduncle and of thoracopods. It was shown that the cypris larvae start to differ already one day after settlement on either a hydroid (leading to hermaphrodites) or an adult hermaphrodite (leading to dwarf males). Dwarf males gradually developed an ovoid body shape and two pairs of circular scutal and tergal primordia. Such cyprids developed neither a carina nor any peduncle or cirri for feeding. The study concludes that the dwarf males of S. scalpellum are not just hermaphrodites arrested early in development. This entails that dwarf males constitute their own separate developmental pathways and points to S. scalpellum dwarf males being more specialized than previously stated. Finally, the study compares differences in dwarf male morphology between S. scalpellum with two other androdioecious species with less specialized dwarf males and use this to discuss evolutionary implications for the adaptive evolution of dwarf males across the Cirripedia. [ABSTRACT FROM AUTHOR]

Androdioecy is the rarest sexual system among plants. The majority of androdioecious species are herbaceous plants that have evolved from dioecious ancestors. Nevertheless, some woody and androdioecious plants have hermaphrodite ancestors, as in the Annonaceae, where androdioecious genera have arisen several times in different lineages. The majority of androdioecious species of Annonaceae belong to the Neotropical tribe Malmeeae. In addition to these species, Pseudoxandra spiritus-sancti was recently confirmed to be androdioecious. Here, we describe the morphology of male and bisexual flowers of Pseudoxandra spiritus-sancti , and investigate the evolution of androdioecy in Malmeeae. The phylogeny of tribe Malmeeae was reconstructed using Bayesian inference, maximum parsimony and maximum likelihood of 32 taxa, using DNA sequences of 66 molecular markers of the chloroplast genome, sequenced by next generation sequencing. The reconstruction of ancestral states was performed for characters associated with sexual systems and floral morphology. The phylogenetic analyses reconstructed three main groups in Malmeeae, ( Malmea ( Cremastosperma , Pseudoxandra )) sister to the rest of the tribe, and ( Unonopsis ( Bocageopsis , Onychopetalum )) sister to ( Mosannona, Ephedranthus , Klarobelia , Oxandra , Pseudephedranthus fragrans , Pseudomalmea, Ruizodendron ovale ). Hermaphroditism is plesiomorphic in the tribe, with four independent evolutions of androdieocy, which represents a synapomorphy of two groups, one that includes three genera and 14 species, the other with a single genus of seven species. Male flowers are unisexual from inception and bisexual flowers possess staminodes and functional stamens. Pseudoxandra spiritus-sancti is structurally androdioecious. [ABSTRACT FROM AUTHOR]

We address the evolution of effective number of individuals under androdioecy and gynodioecy. We analyze dynamic models of autosomal modifiers of weak effect on sex expression. In our zygote control models, the sex expressed by a zygote depends on its own genotype, while in our maternal control models, it depends on the genotype of its maternal parent. Our analysis unifies full multi-dimensional local stability analysis with the Li-Price equation, which for all its heuristic appeal, describes evolutionary change over a single generation. We define a point in the neighborhood of a fixation state from which a single-generation step indicates the asymptotic behavior of the frequency of a modifier allele initiated at an arbitrary point near the fixation state. A concept of heritability appropriate for the evolutionary modification of sex emerges from the Li-Priceframework. We incorporate our theoretical analysis into our previously-developed Bayesian inference framework to develop a new method for inferring the viability of gonochores (males or females) relative to hermaphrodites. Applying this approach to microsatellite data derived from natural populations of the gynodioecious plant Schiedea salicaria and the androdioecious killifish Kryptolebias marmoratus , we find that while female and hermaphrodite S. salicaria appear to have similar viabilities, male K. marmoratus appear to survive to reproductive age at less than half the rate of hermaphrodites. [ABSTRACT FROM AUTHOR]

Plant demography is known to depend on both spatial dynamics and life history, but how these two factors interact is poorly understood. We conducted a longitudinal study of the wind-pollinated annual plant Mercurialis annua that varies geographically in its sexual system to investigate this interaction., Metapopulation demographic models predict that regular population turnover should be a more common feature of monomorphic than dimorphic populations because males and females cannot found new populations by selfing but hermaphrodites can. We tested the prediction that rates of population turnover would be higher in monomorphic compared to dimorphic regions., We surveyed 356 populations of M. annua along five regional transects in Morocco and the Iberian Peninsula over a 3-year period to examine their demography and persistence. Each transect crossed a transition in the sexual system, from a monomorphic region where almost all populations were hermaphroditic to a dimorphic one in which most populations had separate sexes (males with females or hermaphrodites)., As predicted, rates of local apparent extinctions (i.e., the disappearance of adult plants) were nearly 50% higher in monomorphic compared to dimorphic regions. Local extinctions appeared to be driven by changes in vegetation cover, with extinctions tending to occur in sites in which perennial cover also declined. This suggests that disturbance is a primary agent of local extinctions., We further examined the influence of regional dynamics on local demographic properties by investigating patterns of spatial autocorrelation in population density across years. We found positive spatial autocorrelations in plant densities within regions for both sexual systems. However, these positive autocorrelations extended over shorter distances in monomorphic regions, perhaps as a result of greater population flux in these regions., Synthesis. Our study shows that population dynamics may be influenced by processes acting at a range of spatial scales: within patches, across patches within sites, and across sites within regions, as well as by life-history variation. In Mercurialis annua, regional variation in apparent extinction rates is affected by life history and implicated in regulating the geographical distribution of populations with different sexual systems. [ABSTRACT FROM AUTHOR]

Background: The nematode species Pristionchus pacificus has an androdioecious mating system in which populations consist of self-fertilizing hermaphrodites and relatively few males. The prevalence of males in such a system is likely to depend on the relative pros and cons of outcrossing. While outcrossing generates novel allelic combinations and can therefore increase adaptive potential, it may also disrupt the potentially beneficial consequences of repeated generations of selfing. These include purging of deleterious alleles, inheritance of co-adapted allele complexes, improved hermaphrodite fitness and increased population growth. Here we use experimental and population genetic approaches to test hypotheses relating to male production and outcrossing in laboratory and natural populations of P. pacificus sampled from the volcanic island of La Réunion. Results: We find a significant interaction between sampling locality and temperature treatment influencing rates of spontaneous male production in the laboratory. While strains isolated at higher altitude, cooler localities produce a higher proportion of male offspring at 25 °C relative to 20 or 15 °C, the reverse pattern is seen in strains isolated from warmer, low altitude localities. Linkage disequilibrium extends across long physical distances, but fails to approach levels reported for the partially selfing nematode species Caenorhabditis elegans. Finally, we find evidence for admixture between divergent genetic lineages. Conclusions: Elevated rates of laboratory male generation appear to occur under environmental conditions which differ from those experienced by populations in nature. Such elevated male generation may result in higher outcrossing rates, hence driving increased effective recombination and the creation of potentially adaptive novel allelic combinations. Patterns of linkage disequilibrium decay support selfing as the predominant reproductive strategy in P. pacificus. Finally, despite the potential for outcrossing depression, our results suggest admixture has occurred between distinct genetic lineages since their independent colonization of the island, suggesting outcrossing depression may not be uniform in this species. [ABSTRACT FROM AUTHOR]