Self-compatible Species Research Articles

The genetic control of herkogamy.

Herkogamy is the spatial separation of anthers and stigmas within complete flowers, and is a key floral trait that promotes outcrossing in many angiosperms. The degree of separation between pollen-producing anthers and receptive stigmas has been shown to influence rates of self-pollination amongst plants, with a reduction in herkogamy increasing rates of successful selfing in self-compatible species. Self-pollination is becoming a critical issue in horticultural crops grown in environments where biotic pollinators are limited, absent, or difficult to utilise. In these cases, poor pollination results in reduced yield and misshapen fruit. Whilst there is a growing body of work elucidating the genetic basis of floral organ development, the genetic and environmental control points regulating herkogamy are poorly understood. A better understanding of the developmental and regulatory pathways involved in establishing varying degrees of herkogamy is needed to provide insights into the production of flowers more adept at selfing to produce consistent, high-quality fruit. This review presents our current understanding of herkogamy from a genetics and hormonal perspective.

An experimental field study of inbreeding depression in an outcrossing invasive plant.

Inbreeding depression is likely to play an important role during biological invasion. But relatively few studies have investigated the fitness of selfed and outcrossed offspring in self-incompatible invasive plants in natural environments in their introduced range. Moreover, the majority of studies on inbreeding depression have investigated self-compatible species with mixed mating, and less is known about the intensity of inbreeding depression in outcrossing self-incompatible species. Here, we address these questions experimentally by comparing selfed and outcrossed progeny of purple loosestrife (Lythrum salicaria) over four growing seasons, including three under field conditions in a freshwater marsh in southern Ontario, Canada, a region where L. salicaria is highly invasive. The tristylous mating system of L. salicaria involves disassortative mating among floral morphs enforced by trimorphic incompatibility. However, owing to partial incompatibility, self-fertilized seed can be obtained by manual self-pollination thus facilitating comparisons of selfed and outcrossed progeny. We compared progeny with and without intraspecific competition from selfed or outcrossed neighbours and examined the influence of breeding treatment and competition on fitness correlates by measuring a range of life-history traits including: proportion of seeds germinating, days to germination, survival, proportion of plants flowering, time to flowering, vegetative mass, and inflorescence number and mass. We analysed data for each trait using functions from time series estimates of growth and two multiplicative estimates of fitness. We detected varying intensities of inbreeding depression for several traits in three of the four years of the experiment, including inflorescence mass and reproductive output. Cumulative inbreeding depression over four years averaged δ = 0.48 and 0.68, depending on the method used to estimate multiplicative fitness. The competition treatments did not significantly affect plant performance and the magnitude of inbreeding depression. Given the primarily outcrossing mating system of L. salicaria populations, the detection of inbreeding depression for several key life-history traits was as predicted by theory. Our results suggests that biparental inbreeding and low selfing in colonizing populations may have significant effects on demographic parameters such as population growth.

De novo transcriptome of a bast fibre crop Crotalaria juncea reveals T2 ribonuclease genes to investigate late-acting self-rejection of pollens

The biology of gametophytic late-acting self-rejection of pollen in pre-zygotic ovules is relatively unknown in the Fabaceae family of plants. Our understanding of the genetic basis of late-acting self-incompatibility (LSI) is limited due to a lack of sequence data and candidate genes. Crotalaria juncea, a Fabaceae family member that produces commercially important phloem fiber, possesses LSI. Due to a lack of selfed seeds, it is difficult to maintain genetic purity in germplasm and develop breeding lines for improved fiber quality and yield. To investigate candidate genes for LSI in C. juncea, a high-quality de novo transcriptome was generated. It facilitated the identification of genes from the self-incompatibility-related ribonuclease (RNase) family, specifically a Class III T2/RNase gene with sequence properties similar to S-RNase proteins. Based on conserved amino acid motifs, histidine residues, and a high isoelectric point (pI ≥9.0), Cjun_RNS3.1 was identified as an S-RNase homologue (non-S-RNase) or relic S-RNase gene. Unlike typical S-RNases found in known SI plant systems, evolutionary analysis of the Cjun_RNS3.1 protein revealed its ancestral origin. The expression of Cjun_RNS3.1 and other T2/RNase genes revealed differential expression patterns in the pistils of LSI and self-compatible Crotalaria species. The upregulation of the Cjun_RNS3.1 gene during the different stages of pollen tube development indicates that it may be involved in the LSI. In summary, the sunn hemp transcriptome is the first genomic resource reported from a Fabaceae family plant with the LSI trait. A non-S-RNase gene with an ancestral evolutionary origin and a diverse function was found, which could be related to gametophytic LSI. This could be used as a model system to investigate the molecular basis of LSI in the Fabaceae family and help develop C. juncea breeding lines for fibre improvement.

Beyond pollen:ovule ratios: Evolutionary consequences of pollinator dependence and pollination efficiency for pollen and ovule production in angiosperms.

The relative per-flower production of ovules and pollen varies broadly with angiosperm mating systems, with outcrossing types commonly producing more pollen grains per ovule than selfing types. The evolutionary causes of this variation are contentious, especially the relevance of pollination risk. Resolution of this debate may have been hampered by its focus on pollen:ovule (P:O) ratios rather than on the evolution of pollen and ovule numbers per se. Using published mean ovule and pollen counts, we analyzed associations with the proportion of removed pollen that reaches stigmas (pollen-transfer efficiency) and differences between pollinator-dependent and autogamous forms within and among species. Analyses involved Bayesian methods that simultaneously considered variation in pollen and ovule numbers and accounted for phylogenetic relatedness. We also assessed the utility of P:O ratios as mating-system proxies and their association with female outcrossing rates. Median pollen number declined consistently with pollen-transfer efficiency among species, whereas median ovule number did not. Similarly, in both intraspecific and interspecific analyses, pollinator-dependent plants produced more pollen than autogamous plants, whereas ovule production did not differ statistically. Distributions of P:O ratios overlapped extensively for self-incompatible and self-compatible species and for different mating-system classes, and P:O ratios correlated weakly with outcrossing rate. Our findings demonstrate that pollinator dependence and pollination efficiency commonly influence the evolution of pollen number per flower but have more limited effects on ovule number. P:O ratios provide ambiguous, possibly misleading, information about mating systems, especially when compared among clades.

Expanded male sex-determining region conserved during the evolution of homothallism in the green alga Volvox

Male and female genotypes in heterothallic (self-incompatible) species of haploid organisms, such as algae and bryophytes, are generally determined by male and female sex-determining regions (SDRs) in the sex chromosomes. To resolve the molecular genetic basis for the evolution of homothallic (bisexual and self-compatible) species from a heterothallic ancestor, we compared whole-genome data from Thai and Japanese genotypes within the homothallic green alga Volvox africanus. The Thai and Japanese algae harbored expanded ancestral male and female SDRs of ∼1 Mbp each, representing a direct heterothallic ancestor. Therefore, the expanded male and female ancestral SDRs may originate from the ancient (∼75 mya) heterothallic ancestor, and either might havebeen conserved during the evolution of each homothallic genotype. An expanded SDR-like region seems essential for homothallic sexual reproduction in V.africanus, irrespective of male or female origin. Our study stimulates future studies to elucidate the biological significance of such expanded genomic regions.

CAGEE: computational analysis of gene expression evolution.

Despite the increasing abundance of whole transcriptome data, few methods are available to analyze global gene expression across phylogenies. Here, we present a new software package (CAGEE) for inferring patterns of increases and decreases in gene expression across a phylogenetic tree, as well as the rate at which these changes occur. In contrast to previous methods that treat each gene independently, CAGEE can calculate genome-wide rates of gene expression, along with ancestral states for each gene. The statistical approach developed here makes it possible to infer lineage-specific shifts in rates of evolution across the genome, in addition to possible differences in rates among multiple tissues sampled from the same species. We demonstrate the accuracy and robustness of our method on simulated data, and apply it to a dataset of ovule gene expression collected from multiple self-compatible and self-incompatible species in the genus Solanum to test hypotheses about the evolutionary forces acting during mating system shifts. These comparisons allow us to highlight the power of CAGEE, demonstrating its utility for use in any empirical system and for the analysis of most morphological traits. Our software is available at https://github.com/hahnlab/CAGEE/.

Reduced pollination in bilateral flowers could reflect selfing avoidance

The multiple evolutionary transitions from radial to bilateral flower symmetry recorded across the angiosperms have been related to pollinator specialization and the promotion of outcross pollen transfer. In particular, the limited number of physical orientations associated with bilaterality results in a tighter morphological match between a flower and its specialized pollinators, which can enhance the transfer of incoming pollen while preventing autonomous, or even pollinator-mediated self-pollination when associated with dichogamy. In contrast, radial flowers have various planes of symmetry, which may allow pollination by a wider range of pollinators at the cost of more promiscuous pollen transfer. Here we used data from the literature and our own on pollen production, ovule production, and stigmatic pollen load compiled for 171 angiosperm species distributed worldwide to assess differences in pollen transfer in relation to flower symmetry within a phylogenetic comparative framework. We did not find evidence that bilateral and radial flowers differ in average pollen and ovule production. However, bilateral flowers tended to receive less pollen on their stigmas. This decrease in pollen receipt was most apparent among self-compatible species after accounting for the covarying effects of flower size and ovule production. Therefore, our findings indicate that the evolution of flower bilaterality, a trait we found is strongly associated with self-compatibility, comes at a quantitative pollination cost. However, they support the view that a bilateral flower morphology may provide the benefit of limiting self-pollen deposition, particularly in the absence of an alternative mechanism of selfing avoidance.

Fine-Mapping and Comparative Genomic Analysis Reveal the Gene Composition at the S and Z Self-incompatibility Loci in Grasses.

Self-incompatibility (SI) is a genetic mechanism of hermaphroditic plants to prevent inbreeding after self-pollination. Allogamous Poaceae species exhibit a unique gametophytic SI system controlled by two multi-allelic and independent loci, S and Z. Despite intense research efforts in the last decades, the genes that determine the initial recognition mechanism are yet to be identified. Here, we report the fine-mapping of the Z-locus in perennial ryegrass (Lolium perenne L.) and provide evidence that the pollen and stigma components are determined by two genes encoding DUF247 domain proteins (ZDUF247-I and ZDUF247-II) and the gene sZ, respectively. The pollen and stigma determinants are located side-by-side and were genetically linked in 10,245 individuals of two independent mapping populations segregating for Z. Moreover, they exhibited high allelic diversity as well as tissue-specific gene expression, matching the expected characteristics of SI determinants known from other systems. Revisiting the S-locus using the latest high-quality whole-genome assemblies revealed a similar gene composition and structure as found for Z, supporting the hypothesis of a duplicated origin of the two-locus SI system of grasses. Ultimately, comparative genomic analyses across a wide range of self-compatible and self-incompatible Poaceae species revealed that the absence of a functional copy of at least one of the six putative SI determinants is accompanied by a self-compatible phenotype. Our study provides new insights into the origin and evolution of the unique gametophytic SI system in one of the largest and economically most important plant families.

Stamen curvature and temporal flower closure assure reproductive success in an early-spring-flowering perennial in the cold desert of Middle Asia.

Floral organ movements that ensure autonomous selfing are likely to occur in species that grow in habitats with pollinator scarcity and/or an unpredictable environment. Stamen curvature and temporal flower closure are two important floral behaviors that can influence plant pollination mode and reproductive success. However, both behaviors are rarely reported within a species, and little is known about how these two movements of floral organs ensure reproductive success in an unpredictable early spring environment with few pollinators. The aim of this study was to assess whether stamen curvature and temporal flower closure ensure successful reproduction of Leontice incerta in its cold desert habitat. Flowering phenology, floral traits, stamen curvature patterns and flower visitors were surveyed. The breeding system, capacity and timing for autonomous selfing were estimated by pollination manipulations. The timing of floral opening and closure, and benefits of temporal flower closure were determined. We found that flowering of L. incerta began in late March to early April in two populations in two years, and the yellow flowers had neither nectar nor scent. Floral visitation occurred very rarely, but bees (Colletes sp.) were potential pollinators. Fruit and seed set of open and bagged flowers did not differ significantly from that of self-pollinated or cross-pollinated flowers. However, removal of stamens significantly decreased seed set. Self-pollination occurs when the stamens curve and anthers touch the stigma autonomously, suggesting autonomous selfing assurance of seed production in this self-compatible species. Both fruit and seed set of flowers that were prevented from closing were significantly lower than those of control flowers and closed flowers treated with simulated rain treatment. Therefore, stamen curvature and temporal floral closure can ensure successful sexual reproduction of L. incerta in early spring in the cold desert, where lack of pollinators otherwise may lead to pollination failure.

Identification of the genes at S and Z reveals the molecular basis and evolution of grass self-incompatibility.

Self-incompatibility (SI) is a feature of many flowering plants, whereby self-pollen is recognized and rejected by the stigma. In grasses (Poaceae), the genes controlling this phenomenon have not been fully elucidated. Grasses have a unique two-locus system, in which two independent genetic loci (S and Z) control self-recognition. S and Z are thought to have arisen from an ancient duplication, common to all grasses. With new chromosome-scale genome data, we examined the genes present at S- and Z-loci, firstly in ryegrass (Lolium perenne), and subsequently in ~20 other grass species. We found that two DUF247 genes and a short unstructured protein (SP/ZP) were present at both S- and Z- in all SI species, while in self-compatible species these genes were often lost or mutated. Expression data suggested that DUF247 genes acted as the male components and SP/ZP were the female components. Consistent with their role in distinguishing self- from non-self, all genes were hypervariable, although key secondary structure features were conserved, including the predicted N-terminal cleavage site of SP/ZP. The evolutionary history of these genes was probed, revealing that specificity groups at the Z-locus arose before the advent of various grass subfamilies/species, while specificity groups at the S-locus arose after the split of Panicoideae, Chloridoideae, Oryzoideae and Pooideae. Finally, we propose a model explaining how the proteins encoded at the S and Z loci might function to specify self-incompatibility.

Temporal changes in the Swiss flora: implications for flower-visiting insects

BackgroundLocal floristic diversity has massively decreased during the twentieth century in Central Europe even though in the 1990s diversity began increasing again in several regions. However, little is known whether this increase is equally distributed among plant groups with different reproductive traits.MethodsOur study is based on data of the Swiss Biodiversity Monitoring Program. In this program, plant species occurrence is recorded since 2001 in 450 regularly distributed 1 km2 study sites. For all 1774 plant species registered in the study, we researched data on flower/pseudanthium type and colour, reproductive system, and groups of flower visitors. We then tested whether temporal changes in species frequency were equally distributed among species with different trait states.ResultsSpecies richness and functional richness significantly increased in the study sites while functional evenness decreased. The frequency of wind-pollinated species increased more strongly than that of insect-pollinated species. Further, the frequency of species with simple, open insect-pollinated flowers and pseudanthia visited by generalist groups of insects increased slightly more strongly than the frequency of species with complex flowers visited by more specialized groups of flower visitors. Additionally, the frequency of self-compatible species increased significantly more than that of self-incompatible species. Thus, the overall increase in local plant species richness in Switzerland is mostly driven by wind- and generalist insect-pollinated, self-compatible species. In contrast, species with complex flowers, which are essential for specialized groups of flower visitors and species with self-incompatible reproductive systems profited less.ConclusionsOur study thus emphasizes the need to consider functional traits in the planning and monitoring of conservation activities, and calls for a special focus on plant species with specialized reproductive traits.

Rapid evolution of T2/S-RNase genes in Fragaria linked to multiple transitions from self-incompatibility to self-compatibility

The T2/RNase gene family is widespread in eukaryotes, and particular members of this family play critical roles in the gametophytic self-incompatibility (GSI) system in plants. Wild diploid strawberry (Fragaria) species have diversified their sexual systems via self-incompatible and self-compatible traits, yet how these traits evolved in Fragaria remains elusive. By integrating the published and de novo assembled genomes and the newly generated RNA-seq data, members of the RNase T2 gene family were systematically identified in six Fragaria species, including three self-incompatible species (Fragaria nipponica, Fragaria nubicola, and Fragaria viridis) and three self-compatible species (Fragaria nilgerrensis, Fragaria vesca, and Fragaria iinumae). In total, 115 RNase T2 genes were identified in the six Fragaria genomes and can be classified into three classes (I–III) according to phylogenetic analysis. The identified RNase T2 genes could be divided into 22 homologous gene sets according to amino acid sequence similarity and phylogenetic and syntenic relationships. We found that extensive gene loss and pseudogenization coupled with small-scale duplications mainly accounted for variations in the RNase T2 gene numbers in Fragaria. Multiple copies of homologous genes were mainly generated from tandem and segmental duplication events. Furthermore, we newly identified five S-RNase genes in three self-incompatible Fragaria genomes, including two in F. nipponica, two in F. viridis, and one in F. nubicola, which fit for typical features of a pistil determinant, including highly pistil-specific expression, highly polymorphic proteins and alkaline isoelectric point (pI), while no S-RNase genes were found in all three self-compatible Fragaria species. Surprisingly, these T2/S-RNase genes contain at least one large intron (>10 kb). This study revealed that the rapid evolution of T2/S-RNase genes within the Fragaria genus could be associated with its sexual mode, and repeated evolution of the self-compatible traits in Fragaria was convergent via losses of S-RNase.

Benefits of Insect Pollination in Brassicaceae: A Meta-Analysis of Self-Compatible and Self-Incompatible Crop Species

This paper reviewed the effects of insect pollination on the yield parameters of plants from the family Brassicaceae presenting different breeding systems. Meta-analysis indicates that in both self-compatible and self-incompatible crop species, meta-analysis indicates that seed yield (Y), silique set (SQS), number of siliquae/plant (NSQ), and the number of seeds/silique (NSSQ) increase when plants are insect-pollinated compared to when there is no insect pollination. The weight of seeds (WS), however, increased in self-incompatible species but not in self-compatible ones as a result of insect pollination. Overall, the percentage of studies showing a positive effect of insect pollination on yield parameters was higher in self-incompatible than in self-compatible species. It was shown that the ability of self-compatible species to reproduce does not fully compensate for the loss of yield benefits in the absence of insect pollination. Cultivated Brassicaceae attract a wide variety of pollinators, with honeybees (Apis spp.) such as A. mellifera L., A. cerana F., A. dorsata F., and A. florea F. (Hymenoptera: Apidae); other Apidae, such as bumblebees (Bombus spp.) (Hymenoptera: Apidae); mining bees (Hymenoptera: Andrenidae); sweat bees (Hymenoptera: Halictidae); and hoverflies (Diptera: Syrphidae) constituting the most common ones. The benefits of insect pollination imply that pollinator conservation programs play a key role in maximizing yield in cruciferous crops.

Breeding System and Response of the Pollinator to Floral Larceny and Florivory Define the Reproductive Success in Aerides odorata.

Consumption of pollination reward by felonious means in a plant species can influence the foraging behavior of its pollinator and eventually the reproductive success. So far, studies on this aspect are largely confined to interaction involving plant-pollinators and nectar robbers or thieves. However, a foraging guild in such interactions may also include floral herbivores or florivores. There is a paucity of information on the extent to which nectar larcenists may influence the foraging behavior of the pollinator and reproductive fitness of plants in the presence of a florivore. We investigated various forms of larceny in the natural populations of Aerides odorata, a pollinator-dependent and nectar-rewarding orchid. These populations differed in types of foraging guild, the extent of larceny (thieving/robbing), which can occur with or without florivory, and natural fruit-set pattern. The nectariferous spur of the flower serves as an organ of interest among the foraging insects. While florivory marked by excision of nectary dissuades the pollinator, nectar thieving and robbing significantly enhance visits of the pollinator and fruit-set. Experimental pollinations showed that the species is a preferential outbreeder and experiences inbreeding depression from selfing. Reproductive fitness of the orchid species varies significantly with the extent of floral larceny. Although nectar thieving or robbing is beneficial in this self-compatible species, the negative effects of florivory were stronger. Our findings suggest that net reproductive fitness in the affected plant species is determined by the overarching effect of its breeding system on the overall interacting framework of the foraging guild.

Interactions between breeding system and ploidy affect niche breadth in Solanum.

Understanding the factors driving ecological and evolutionary interactions of economically important plant species is important for agricultural sustainability. The geography of crop wild relatives, including wild potatoes (Solanum section Petota), have received attention; however, such information has not been analysed in combination with phylogenetic histories, genomic composition and reproductive systems to identify potential species for use in breeding for abiotic stress tolerance. We used a combination of ordinary least-squares (OLS) and phylogenetic generalized least-squares (PGLM) analyses to identify the discrete climate classes that make up the climate niche that wild potato species inhabit in the context of breeding system and ploidy. Self-incompatible diploid or self-compatible polyploid species significantly increase the number of discrete climate classes within a climate niche inhabited. This result was sustained when correcting for phylogenetic non-independence in the linear model. Our results support the idea that specific breeding system and ploidy combinations increase niche breadth through the decoupling of geographical range and niche diversity, and therefore, these species may be of particular interest for crop adaptation to a changing climate.

Female self-incompatibility type in heterostylous Primula is determined by the brassinosteroid-inactivating cytochrome P450 CYP734A50

Most flowering plants are hermaphrodites, with flowers having both male and female reproductive organs. One widespread adaptation to limit self-fertilization is self-incompatibility (SI), where self-pollen fails to fertilize ovules.1,2 In homomorphic SI, many morphologically indistinguishable mating types are found, although in heteromorphic SI, the two or three mating types are associated with different floral morphologies.3-6 In heterostylous Primula, a hemizygous supergene determines a short-styled S-morph and a long-styled L-morph, corresponding to two different mating types, and full seed set only results from intermorph crosses.7-9 Style length is controlled by the brassinosteroid (BR)-inactivating cytochrome P450 CYP734A50,10 yet it remains unclear what defines the male and female incompatibility types. Here, we show that CYP734A50 also determines the female incompatibility type. Inactivating CYP734A50 converts short S-morph styles into long styles with the same incompatibility behavior as L-morph styles, and this effect can be mimicked by exogenous BR treatment. Invitro responses of S- and L-morph pollen grains and pollen tubes to increasing BR levels could only partly explain their different invivo behavior, suggesting both direct and indirect effects of the different BR levels in S- versus L-morph stigmas and styles in controlling pollen performance. This BR-mediated SI provides a novel mechanism for preventing self-fertilization. The joint control of morphology and SI by CYP734A50 has important implications for the evolutionary buildup of the heterostylous syndrome and provides a straightforward explanation for why essentially all of the derived self-compatible homostylous Primula species are long homostyles.11.

Evolution of seed mass associated with mating systems in multiple plant families.

In flowering plants, the evolution of self‐fertilization (selfing) from obligate outcrossing is regarded as one of the most prevalent evolutionary transitions. The evolution of selfing is often accompanied by various changes in genomic, physiological and morphological properties. In particular, a set of reproductive traits observed typically in selfing species is called the “selfing syndrome”. A mathematical model based on the kinship theory of genetic imprinting predicted that seed mass should become smaller in selfing species compared with outcrossing congeners, as a consequence of the reduced conflict between maternally and paternally derived alleles in selfing plants. Here, we test this prediction by examining the association between mating system and seed mass across a wide range of taxa (642 species), considering potential confounding factors: phylogenetic relationships and growth form. We focused on three plant families—Solanaceae, Brassicaceae and Asteraceae—where information on mating systems is abundant, and the analysis was performed for each family separately. When phylogenetic relationships were controlled, we consistently observed that selfers (represented by self‐compatible species) tended to have a smaller seed mass compared with outcrossers (represented by self‐incompatible species) in these families. In summary, our analysis suggests that small seeds should also be considered a hallmark of the selfing syndrome, although we note that mating systems have relatively small effects on seed mass variation.

Pistil Mating Type and Morphology Are Mediated by the Brassinosteroid Inactivating Activity of the S-Locus Gene BAHD in Heterostylous Turnera Species.

Heterostyly is a breeding system that promotes outbreeding through a combination of morphological and physiological floral traits. In Turnera these traits are governed by a single, hemizygous S-locus containing just three genes. We report that the S-locus gene, BAHD, is mutated and encodes a severely truncated protein in a self-compatible long homostyle species. Further, a self-compatible long homostyle mutant possesses a T. krapovickasii BAHD allele with a point mutation in a highly conserved domain of BAHD acyl transferases. Wild type and mutant TkBAHD alleles were expressed in Arabidopsis to assay for brassinosteroid (BR) inactivating activity. The wild type but not mutant allele caused dwarfism, consistent with the wild type possessing, but the mutant allele having lost, BR inactivating activity. To investigate whether BRs act directly in self-incompatibility, BRs were added to in vitro pollen cultures of the two mating types. A small morph specific stimulatory effect on pollen tube growth was found with 5 µM brassinolide, but no genotype specific inhibition was observed. These results suggest that BAHD acts pleiotropically to mediate pistil length and physiological mating type through BR inactivation, and that in regard to self-incompatibility, BR acts by differentially regulating gene expression in pistils, rather than directly on pollen.

Identification and chromosome doubling of Fragaria mandshurica and F. nilgerrensis

Genetic factors that control various excellent agronomic traits are present in wild strawberries; however, barriers such as ploidy differences and interspecific incompatibility prevent the transmission of these genetic factors to cultivated strawberries. Using the colchicine method to induce doubling, the autotetraploids of Fragaria mandshurica and F. nilgerrensis were obtained, and the biological characteristics of the autotetraploids and the compatibility status distribution in different cross-pollination combinations relating to the two species were further examined. The autotetraploids showed thicker and stronger morphological characteristics than those of their corresponding diploids, especially the F. mandshurica species. The change in ploidy can cause the appearance of new excellent agronomic traits: the autotetraploid of F. nilgerrensis is more resistant to anthracnose disease, caused by Colletotrichum gloeosporioides, than the diploid, and the flowering period is significantly longer, with an increase in flower quantity. There is unilateral incompatibility between the two self-compatible species. The autotetraploids are similar to the corresponding diploid in pollination compatibility; however, due to the decline in pollen viability, the related hybridisation combinations show slightly lower compatibility. The closer the ploidy is, the easier it is to obtain hybrid offspring; the tetraploids of both species crossed with cultivated strawberries, showing possible increases in compatibility after chromosome doubling.

Inter-morph pollen flow and reproductive success in a self-compatible species with stigma-height dimorphism: the influence of herkogamy and reciprocity.

Inter-morph pollen transfer and its dependence on herkogamy and reciprocity are not completely understood in species with stigma-height dimorphism. We asked whether total stigmatic pollen loads, inter-morph fraction of pollen load and reproductive success differed between morphs in Jasminum malabaricum, a species exhibiting stigma-height dimorphism. We tested whether higher herkogamy and reciprocity resulted in higher inter-morph pollen deposition and reproductive success. We quantified individual-level estimates of herkogamy, reciprocity, total stigmatic pollen load, inter-morph stigmatic pollen fraction and fruit set for both morphs in naturally occurring populations of J. malabaricum. Total pollen load was higher in the long-styled morph, inter-morph pollen fraction was higher in the short-styled morph, but fruit set did not differ between morphs. Higher herkogamy resulted in a higher inter-morph fraction of pollen load and fruit set in the long-styled morph of one population. In the other population, only reciprocity was found to be related to inter-morph pollen deposition. This study is the first to quantify and report natural inter-morph stigmatic pollen load in a species with stigma-height dimorphism. Morph-specific differences in pollen load were similar to patterns commonly observed in heterostylous species. The results highlight the importance of both herkogamy and reciprocity in facilitating inter-morph pollen transfer. Population-specific patterns indicate that local environmental factors determine the relative functional importance of herkogamy and reciprocity.