High Selfing Rates Research Articles

Sibship and parentage reconstruction as genetic tool for designing and monitoring plant translocations

Estimating contemporary effective population size (Ne), sibship and parentage can be useful for designing and monitoring plant translocations. Contemporary Ne is an important indicator of population viability, as it considers several demographic factors, e.g. sex ratio, mate compatibility, inbreeding, nonrandom mating, and family size. Sibship and parentage analyses allow not only estimating genetic relatedness of transplants and post‐translocation generations, but also selfing rates, parent contributions to offspring and contemporary seed and pollen dispersal. Sibship assignment method was used to reconstruct offspring pedigree and parentage and to estimate contemporary Ne, selfing rate, and contemporary seed and pollen dispersal in wild (used as seed sources for translocation) and translocated populations of the self‐compatible, insect‐pollinated Dianthus deltoides (Caryophyllaceae). Source populations show very small Ne estimates (5–21) and high selfing rates (28.6–68.9%), despite high census sizes. The first post‐translocation generation (recruits and transplant seed progeny) showed small contemporary Ne estimates (19–53) and substantial selfing rates (8.4–22.3%), despite high flowering sizes, overall genetic diversity and multiple paternity. Parentage assignments revealed high levels of admixture between sources, restricted seed dispersal and successful—often restricted—pollen dispersal in the post‐translocation generation. Pollinator service, crosses between selfed or related transplants and floral display, may have contributed to selfing and pollen dispersal patterns in translocated populations. The present findings support the interest of sibship and parentage reconstruction as genetic tool for designing and monitoring plant translocations, emphasizing the importance of assessing the genetic quality of the transplants, and contributing to a more precise evaluation of population extinction risk.

Phenotypic Selection in Halenia elliptica D. Don (Gentianaceae), an Alpine Biennial with Mixed Mating System.

The transition from outcrossing to selfing is a common evolutionary trend in flowering plants, and floral traits change significantly with the evolution of selfing. Whether or not plant traits are subjected to selection remains an open question in species with mixed mating systems. We examined phenotypic selection in two populations of Halenia elliptica with different selfing rates. We found that the pollen–ovule ratio, seed size, plant height, spur length, and pollinator visitation rate in the population with the higher selfing rate were lower than those in the population with the lower selfing rate. Selfing provides reproductive assurance for populations when pollinator service is low, and the floral traits that are associated with selfing syndrome are evident in populations with a higher selfing rate but are subjected to weak selection in each of the two populations with different selfing rates. Directional selection for an early flowering time indicated that late blooming flowers could experience a risk of seed development in alpine environments, and for large plants, selection indicated that seed production could be limited by the available resources. The floral traits that are associated with pollinator attraction and specialization could be subjected to weak selection at the plant level as selfing evolves, and the selective pressures that are independent of pollinators might not change significantly; highlighting the selective biotic and abiotic pressures that shape the morphological traits of plant species and their independence from the mating system.

Evolutionary breakdown of distyly to homostyly is accompanied by reductions of floral scent inPrimula oreodoxa

AbstractReproductive traits that function in pollinator attraction may be reduced or lost during evolutionary transitions from outcrossing to selfing. Although floral scent plays an important role in attracting pollinators in outcrossing species, few studies have investigated associations between floral scent variation and intraspecific mating system transitions. The breakdown of distyly to homostyly represents a classic example of a shift from outcrossing to selfing and provides an opportunity to test whether floral fragrances have become reduced and/or changed in composition with increased selfing. Here, we evaluate this hypothesis by quantifying floral volatiles using gas chromatography‐mass spectrometry in two distylous and four homostylous populations ofPrimula oreodoxaFranchet, a perennial herb from SW China. Our analysis revealed significant variation of volatile organic compounds (VOCs) among populations ofP. oreodoxa. Although there was no difference in VOCs between floral morphs in distylous populations as predicted, we detected a substantial reduction in VOC emissions and the average number of scent compounds in homostylous compared with distylous populations. A total of 12 compounds, mainly monoterpenoids and sesquiterpenoids, distinguished homostylous and distylous morphs; of these, (E)‐β‐ocimene was the most important in contributing to the difference in volatiles, with significantly lower emissions in homostyles. Our findings support the hypothesis that the transition from outcrossing to selfing is accompanied by the loss of floral volatiles. The modification to floral fragrances inP. oreodoxaassociated with mating system change might occur because high selfing rates in homostylous populations result in relaxed selection for floral attractiveness.

Outcrossing rates in an experimentally admixed population of self-compatible and self-incompatible Arabidopsis lyrata

The transition to self-compatibility from self-incompatibility is often associated with high rates of self-fertilization, which can restrict gene flow among populations and cause reproductive isolation of self-compatible (SC) lineages. Secondary contact between SC and self-incompatible (SI) lineages might re-establish gene flow if SC lineages remain capable of outcrossing. By contrast, intrinsic features of SC plants that reinforce high rates of self-fertilization could maintain evolutionary divergence between lineages. Arabidopsis lyrata subsp. lyrata is characterized by multiple origins of self-compatibility and high rates of self-fertilization in SC-dominated populations. It is unclear whether these high rates of selfing by SC plants have intrinsic or extrinsic causes. We estimated outcrossing rates and examined patterns of pollinator movement for 38 SC and 40 SI maternal parents sampled from an admixed array of 1509 plants sourced from six SC and six SI populations grown under uniform density. Although plants from SI populations had higher outcrossing rates (mean tm = 0.78 ± 0.05 SE) than plants from SC populations (mean tm = 0.56 ± 0.06 SE), outcrossing rates among SC plants were substantially higher than previous estimates from natural populations. Patterns of pollinator movement appeared to contribute to lower outcrossing rates for SC plants; we estimated that 40% of floral visits were geitonogamous (between flowers of the same plant). The relatively high rates of outcrossing for SC plants under standardized conditions indicate that selfing rates in natural SC populations of A. lyrata are facultative and driven by extrinsic features of A. lyrata, including patterns of pollinator movement.

Cloning and Selfing Affect Population Genetic Variation in Simulations of Outcrossing, Sexual Sea Stars

AbstractMany sea stars are well known for facultative or obligate asexual reproduction in both the adult and larval life-cycle stages. Some species and lineages are also capable of facultative or obligate hermaphroditic reproduction with self-fertilization. However, models of population genetic variation and empirical analyses of genetic data typically assume only sexual reproduction and outcrossing. A recent reanalysis of previously published empirical data (microsatellite genotypes) from two studies of one of the most well-known sea star species (the crown-of-thorns sea star; Acanthaster sp.) concluded that cloning and self-fertilization in that species are rare and contribute little to patterns of population genetic variation. Here we reconsider that conclusion by simulating the contribution of cloning and selfing to genetic variation in a series of models of sea star demography. Simulated variation in two simple models (analogous to previous analyses of empirical data) was consistent with high rates of cloning or selfing or both. More realistic scenarios that characterize population flux in sea stars of ecological significance, including outbreaks of crown-of-thorns sea stars that devastate coral reefs, invasions by Asterias amurensis, and epizootics of sea star wasting disease that kill Pisaster ochraceus, also showed significant but smaller effects of cloning and selfing on variation within subpopulations and differentiation between subpopulations. Future models or analyses of genetic variation in similar study systems might benefit from simulation modeling to characterize possible contributions of cloning or selfing to genetic variation in population samples or to understand the limits on inferring the effects of cloning or selfing in nature.

Why are deleterious mutations maintained in selfing populations? An analysis of the effects of early- and late-acting mutations by a two-locus two-allele model

Frequencies of deleterious mutations are higher than expected in many plants. Here, by developing a two-locus two-allele model, I examine the effects of differential timing of the expression of deleterious mutations (two-stage effects) on the maintenance of mutations. I assume early- and late-acting loci to distinguish whether maintenance of mutations in populations with high selfing rates is explained better by two-stage effects of single mutations, or by separate mutations in both early- and late-acting loci. I found that, when ovules are overproduced, the stable frequency of early-acting mutations is higher if mutations also occur in a late-acting locus than if a late-acting mutation is lacking. The stable frequency of late-acting mutations is higher if mutations also occur in an early-acting locus than if an early-acting mutation is lacking. Selective interference does not account for these results because analyses in which the number of loci subject to mutations is equalized are included. Overproduction of ovules has little effect on maintenance if either early- or late-acting mutations are lacking, whereas when ovules are not overproduced, the two-stage effect does not enhance the maintenance of mutations. Hence, mutations occurring in both loci coupled with overproduction of ovules enhances the maintenance of mutations in populations with high selfing rates. The detailed mechanisms underlying the two-stage effect were also analyzed.

Pollination efficiency and pollen limitation in bistaminate vs tetrastaminate Lamiaceae

AbstractThe evolution of bilabiate flowers in Lamiaceae has several times led to a reduction in stamen number. It is generally assumed that stamen reduction goes along with decreasing pollen production. One should, therefore, expect that bistaminate flowers have either a more efficient pollen transfer mechanism or a higher selfíng rate than tetrastaminate species to adapt to the stamen reduction. We tested this hypothesis in a community of Lamiaceae including three bistaminate and three tetrastaminate species. The results of selfing experiments, extra‐pollination, counting pollen number, and stigmatic pollen load analysis were used to calculate the autonomous selfing index, the pollen limitation index, and the pollination efficiency index. The results indicate a significantly lower pollen number in bistaminate flowers than in tetrastaminate flowers of similar anther size. Bistaminate species shared a higher selfing rate which was related to their structural adaptation to delayed selfing. In Ziziphora clinopodioides, the delayed selfing was accompanied by low pollination efficiency, and plants suffered from pollen limitation at the location studied. However, in Salvia sclarea and Salvia hypoleuca, which benefited from a relatively high pollination efficiency, no pollen limitation was observed. Our results reveal a close association between adaptive mechanisms such as increasing precision of pollen transfer and evolution of delayed selfing with stamen reduction in bistaminate Lamiaceae.

Faster Rates of Molecular Sequence Evolution in Reproduction-Related Genes and in Species with Hypodermic Sperm Morphologies.

Sexual selection drives the evolution of many striking behaviors and morphologies and should leave signatures of selection at loci underlying these phenotypes. However, although loci thought to be under sexual selection often evolve rapidly, few studies have contrasted rates of molecular sequence evolution at such loci across lineages with different sexual selection contexts. Furthermore, work has focused on separate sexed animals, neglecting alternative sexual systems. We investigate rates of molecular sequence evolution in hermaphroditic flatworms of the genus Macrostomum. Specifically, we compare species that exhibit contrasting sperm morphologies, strongly associated with multiple convergent shifts in the mating strategy, reflecting different sexual selection contexts. Species donating and receiving sperm in every mating have sperm with bristles, likely to prevent sperm removal. Meanwhile, species that hypodermically inject sperm lack bristles, potentially as an adaptation to the environment experienced by hypodermic sperm. Combining functional annotations from the model, Macrostomum lignano, with transcriptomes from 93 congeners, we find genus-wide faster sequence evolution in reproduction-related versus ubiquitously expressed genes, consistent with stronger sexual selection on the former. Additionally, species with hypodermic sperm morphologies had elevated molecular sequence evolution, regardless of a gene’s functional annotation. These genome-wide patterns suggest reduced selection efficiency following shifts to hypodermic mating, possibly due to higher selfing rates in these species. Moreover, we find little evidence for convergent amino acid changes across species. Our work not only shows that reproduction-related genes evolve rapidly also in hermaphroditic animals, but also that well-replicated contrasts of different sexual selection contexts can reveal underappreciated genome-wide effects.

Self-Fertilization, Inbreeding, and Yield in Alfalfa Seed Production.

Selfing (self-pollination) is the ultimate form of inbreeding, or mating among close relatives. Selfing can create yield loss when inbreeding depression, defined as a lower survival and reproduction of inbred relative to outbred progeny, is present. To determine the impact of selfing in alfalfa (Medicago sativa L.), we quantified the selfing rate of 32 alfalfa seed production fields located in three regions, namely, the Pacific Northwest (PNW), the Central Valley of California (CEV), and the Imperial Valley of California (IMP). Selfing rates (the proportion of selfed seeds) varied between 5.3 and 30% with an average of 12.2% over the 32 seed production fields. In both the parents and their progeny, we observed an excess of heterozygotes relative to Hardy–Weinberg expectations. We detected notable levels of inbreeding in parents (0.231 ± 0.007 parental inbreeding coefficient) and progeny (0.229 ± 0.005). There were a 15% decrease in the number of seeds per stem (seed set) and a 13% decline in the number of seeds per pod in selfed relative to outcrossed stems, but negligible inbreeding depression for pods per raceme and seed weight. The number of racemes on selfed stems increased significantly in fields with greater selfing rates, supporting the presence of geitonogamous or among flower selfing. Despite the significant level of inbreeding depression, seed set did not decrease in fields with higher selfing rates, where the greater number of racemes on the selfed stems increased the seed set. The effects of the field selfing rate on the seed yield metrics were mostly indirect with direct effects of the number of racemes per stem. Available data indicate that the majority of selfing in alfalfa is pollinator-mediated, and thus, eliminating selfing in alfalfa seed production would require the selection of self-incompatible varieties, which, by eliminating inbreeding depression, would provide a 15% potential increase in seed yield and an increase in future hay yield.

New genomic resources and comparative analyses reveal differences in floral gene expression in selfing and outcrossing Collinsia sister species.

The evolutionary transition from outcross- to self-fertilization is one of the most common in angiosperms and is often associated with a parallel shift in floral morphological and developmental traits, such as reduced flower size and pollen to ovule ratios, known as the “selfing syndrome.” How these convergent phenotypes arise, the extent to which they are shaped by selection, and the nature of their underlying genetic basis are unsettled questions in evolutionary biology. The genus Collinsia (Plantaginaceae) includes seven independent transitions from outcrossing or mixed mating to high selfing rates accompanied by selfing syndrome traits. Accordingly, Collinsia represents an ideal system for investigating this parallelism, but requires genomic resource development. We present a high quality de novo genome assembly for the highly selfing species Collinsia rattanii. To begin addressing the basis of selfing syndrome developmental shifts, we evaluate and contrast patterns of gene expression from floral transcriptomes across three stages of bud development for C. rattanii and its outcrossing sister species Collinsia linearis. Relative to C. linearis, total gene expression is less variable among individuals and bud stages in C. rattanii. In addition, there is a common pattern among differentially expressed genes: lower expression levels that are more constant across bud development in C. rattanii relative to C. linearis. Transcriptional regulation of enzymes involved in pollen formation specifically in early bud development may influence floral traits that distinguish selfing and outcrossing Collinsia species through pleiotropic functions. Future work will include additional Collinsia outcrossing-selfing species pairs to identify genomic signatures of parallel evolution.

Evolution of apetaly in the cosmopolitan genus Stellaria

Apetaly is widespread across distantly related lineages of flowering plants and is associated with abiotic (or self-) pollination. It is particularly prevalent in the carnation family, and the cosmopolitan genus Stellaria contains many lineages that are hypothesized to have lost petals from showy petalous ancestors. But the pollination biology of apetalous species of Stellaria remains unclear. Using a substantial species-level sampling (~92% of known taxonomic diversity), we describe the pattern of petal evolution within Stellaria using ancestral character state reconstructions. To help shed light on the reproductive biology of apetalous Stellaria, we conducted a field experiment at an alpine tundra site in the southern Rocky Mountains to test whether an apetalous species (S. irrigua) exhibits higher levels of selfing than a sympatric, showy petalous congener (S. longipes). Analyses indicated that the ancestor of Stellaria was likely showy petalous and that repeated, parallel reductions of petals occurred in clades across much of the world, with uncommon reversal back to showy petals. Field experiments supported high rates of selfing in the apetalous species and high rates of outcrossing in the petalous species. Petal loss is rampant across major clades of Stellaria and is potentially linked with self-pollination worldwide. Self-pollination occurs within the buds in S. irrigua, and high propensities for this and other forms of selfing known in many other taxa of arctic-alpine habitats may reflect erratic availability of pollinators.

Mating system of Datura inoxia: association between selfing rates and herkogamy within populations.

Plant mating system determines, to a great extent, the demographic and genetic properties of populations, hence their potential for adaptive evolution. Variation in plant mating system has been documented between phylogenetically related species as well between populations of a species. A common evolutionary transition, from outcrossing to selfing, is likely to occur under environmental spatial variation in the service of pollinators. Here, we studied two phenotypically (in floral traits) and genetically (in neutral molecular markers) differentiated populations of the annual, insect-pollinated, plant Datura inoxia in Mexico, that differ in the service of pollinators (Mapimí and Cañada Moreno). First, we determined the populations’ parameters of phenotypic in herkogamy, outcrossing and selfing rates with microsatellite loci, and assessed between generation (adults and seedlings) inbreeding, and inbreeding depression. Second, we compared the relationships between parameters in each population. Results point strong differences between populations: plants in Mapimí have, on average, approach herkogamy, higher outcrossing rate (tm = 0.68), lower primary selfing rate (r = 0.35), and lower inbreeding at equilibrium (Fe = 0.24) and higher inbreeding depression (δ = 0.25), than the populations of Cañada. Outcrossing seems to be favored in Mapimí while selfing in Cañada. The relationship between r and Fe were negatively associated with herkogamy in Mapimí; here, progenies derived from plants with no herkogamy or reverse herkogamy had higher selfing rate and inbreeding coefficient than plants with approach herkogamy. The difference Fe–F is positively related to primary selfing rate (r) only in Cañada Moreno which suggests inbreeding depression in selfing individuals and then genetic purging. In conclusion, mating system evolution may occur differentially among maternal lineages within populations of Datura inoxia, in which approach herkogamy favors higher outcrossing rates and low levels of inbreeding and inbreeding depression, while no herkogamy or reverse herkogamy lead to the evolution of the “selfing syndrome” following the purge of deleterious alleles despite high inbreeding among individuals.

Seasonal change of flower sex ratio and pollinator dynamics in three reproductive ecotypes of protandrous plant

AbstractCimicifuga simplex has three genetically and ecologically distinct pollination morphs with different flowering phenology, flower sex expressions, and selfing rates. A previous study showed that strong protandry in hermaphroditic flowers of C. simplex causes there to be seasonal minority sexes; for example, unisexual female ramets are advantageous in the first half of the flowering season and bloom then (minority sex hypothesis). That study, however, did not distinguish among the three pollination morphs of C. simplex. We investigated seasonal sex ratio changes and pollinator environments of the three morphs to verify and expand the minority sex hypothesis. We investigated flowering phenology, pollinator quantity and quality, seasonal population sex ratio dynamics, and stamen/ovule ratios in hermaphroditic flowers. We also examined the seasonal female reproductive success of hermaphroditic flowers, and the effect of male flower excision on fruit set by morph II hermaphroditic flowers. Morph I (mainly hermaphroditic and female ramets) had high pollinator quality and quantity. Fruit set of hermaphroditic ramets was high throughout the flowering season. Morph II (hermaphroditic and andromonoecious ramets) had low pollinator quality, and few pollinators visited near the beginning and end of the season. Removal of male flowers led to a pollen limitation. Morph III (hermaphroditic ramets and a high selfing rate) had very low pollinator abundance throughout the flowering season, and male and female phases largely overlapped. The scarcity of male flowers in morph I is likely a result of the high pollinator quantity and quality. The high fruit set suggests that no pollen limitation existed. The low pollinator quality and quantity of the morph II population caused it to suffer from pollen limitation and may explain the absence of unisexual female ramets in this morph. The high selfing rate of morph III may be due to the extremely low pollinator abundance. Theoretical studies have indicated that the optimal investment allocation to male and female functions depends on whether a pollen limitation exists. In C. simplex, the observed relationships between pollinator environment and reproductive systems are consistent with these theoretical models.

The opportunity for outcrossing varies across the geographic range of the primarily selfing Clarkia xantiana ssp. parviflora.

The timing of self-fertilization has potentially important consequences for the trajectory of mating system evolution, the opportunity for outcrossing, and the maintenance of genetic variation in populations. For primarily selfing taxa, it remains poorly understood as to how floral variation influences the opportunity for outcrossing and whether those attributes vary among populations across geographic ranges. We examined variation in floral traits (herkogamy, protandry, flower size, stigma stage at anthesis, timing of stigma receptivity) in seven populations of Clarkia xantiana ssp. parviflora, a primarily selfing taxon, spanning from the western to eastern margins of its distribution. We also performed experimental emasculations and pollinations (followed by stigma severing) to quantify the extent of opportunities for outcrossing across flower development. There was marked among-population variation in all floral traits, particularly between far eastern and western populations. Emasculation experiments showed that the eastern populations had minimal autonomous selfing, but western populations had high rates of selfing within 24 h after anthesis. Population variation in autofertility was significantly predicted by floral trait variation, especially protandry and petal size. Greater protandry both extended the time over which outcrossing could potentially occur and reduced the probability of autonomous selfing, suggesting that there may be a tradeoff that results in fitness loss when pollinator visits are not common. The east-west pattern of differentiation in some floral traits parallels that of postglacial range expansion, suggesting that selection on the mating system may have been strong in the process of range expansion.

Hidden genetic variance contributes to increase the short-term adaptive potential of selfing populations.

Standing genetic variation is considered a major contributor to the adaptive potential of species. The low heritable genetic variation observed in self-fertilizing populations has led to the hypothesis that species with this mating system would be less likely to adapt. However, a non-negligible amount of cryptic genetic variation for polygenic traits, accumulated through negative linkage disequilibrium, could prove to be an important source of standing variation in self-fertilizing species. To test this hypothesis, we simulated populations under stabilizing selection subjected to an environmental change. We demonstrate that, when the mutation rate is high (but realistic), selfing populations are better able to store genetic variance than outcrossing populations through genetic associations, notably due to the reduced effective recombination rate associated with predominant selfing. Following an environmental shift, this diversity can be partially remobilized, which increases the additive variance and adaptive potential of predominantly (but not completely) selfing populations. In such conditions, despite initially lower observed genetic variance, selfing populations adapt as readily as outcrossing ones within a few generations. For low mutation rates, purifying selection impedes the storage of diversity through genetic associations, in which case, as previously predicted, the lower genetic variance of selfing populations results in lower adaptability compared to their outcrossing counterparts. The population size and the mutation rate are the main parameters to consider, as they are the best predictors of the amount of stored diversity in selfing populations. Our results and their impact on our knowledge of adaptation under high selfing rates are discussed.

Maintenance of high inbreeding depression in selfing populations: Two-stage effect of early- and late-acting mutations

High estimates of inbreeding depression have been obtained in many plant populations with high selfing rates. However, deleterious mutations might be purged from such populations as a result of selfing. I developed a simulation model assuming the presence of mutations at two sets of loci, namely, early- and late-acting loci, and the selective abortion of embryos coupled with ovule overproduction. In the model, early-acting loci are expressed during embryo initiation, and less vigorous embryos are aborted. Late-acting loci are expressed after selective abortion ends; the surviving embryos (seeds) compete, and some of them form the next generation. If mutations are allowed to occur in both early- and late-acting loci, both types increase in frequency in populations with high selfing rates. However, this phenomenon does not occur if mutations occur only in the early- or only in the late-acting loci. Consistent results are observed even if the total number of loci in which mutations are allowed to occur is the same for simulations with both early- and late-acting loci, only early-acting loci, or only late-acting loci, indicating that the presence of both types of loci is the causal factor. Thus, the two-stage effect, or occurrence of both early- and late-acting mutations, promotes the maintenance of these mutations in populations with high selfing rates.

Management Practices and Breeding History of Varieties Strongly Determine the Fine Genetic Structure of Crop Populations: A Case Study Based on European Wheat Populations

As the effects of climate change begin to be felt on yield stability, it is becoming essential to promote the use of genetic diversity in farmers’ fields. The presence of genetic variability in variety could fulfil this purpose. Indeed, the level of intra-varietal genetic diversity influences the spatio-temporal stability of yields and the disease susceptibility of crop species. Breeding history of varieties and their management practices are two factors that should influence intra-varietal genetic diversity. This paper describes the genetic diversity of eight wheat samples covering a gradient from modern single varieties to on-farm mixtures of landraces. This gradient discriminates between landrace, historical and modern varieties, considering the breeding history of varieties, between single-varieties and mixtures of varieties, and between ex situ and in situ de facto strategy in terms of management practices. Genetic diversity of these samples was analyzed with the help of 41 single nucleotide polymorphism markers located in neutral regions, through computing genetic indices at three different levels: Allelic, haplotypic and genetic group level. Population structure and kinship were depicted using discriminant analysis and kinship network analysis. Results revealed an increase in the complexity of the genetic structure as we move on the gradient of variety types (from modern single variety to in situ on-farm mixtures of landraces). For the landraces, the highest levels of genetic diversity have been observed for a landrace (Solina d’Abruzzo) continuously grown on-farm in the region of Abruzzo, in Italy, for many decades. This landrace showed an excess of haplotypic diversity compared to landraces or the historical variety that were stored in genebanks (ex situ conservation). Genetic analyses of the mixtures revealed that, despite a very high selfing rate in wheat, growing in evolutionary mixtures promotes recombination between different genetic components of the mixture, a second way to increase the level of haplotype diversity. When management practices such as growing in mixture and on-farm management are combined, they substantially increase the different levels of genetic diversity of the populations (allelic, haplotypic, genetic group diversity), and consequently promote their adaptability. Our results confirm the need to develop and manage evolving diversified large populations on-farm. These results invite crop diversity managers such as genebank curators, community seed bank managers and farmers’ organizations to adapt their management strategies to the type of variety they wish to manage, because we have shown that their choices have a strong influence on the genetic composition of the crop populations.

Multiple founder events explain the genetic diversity and structure of the model allopolyploid grass Brachypodium hybridum in the Iberian Peninsula hotspot.

It is accepted that contemporary allopolyploid species have originated recurrently, but very few cases have been documented using multiple natural formations of the same species. To extend our knowledge, we have investigated the multiple origins, genetic variation and structure of the allotetraploid grass Brachypodium hybridum with respect to its progenitor diploid species B. distachyon (D genome) and B. stacei (S genome). For this, our primary focus is the Iberian Peninsula, an evolutionary hotspot for the genus Brachypodium. We analysed 342 B. hybridum individuals from 36 populations using ten nuclear SSR loci and two plastid loci. The B. hybridum genetic profiles were compared with those previously reported for B. stacei and B. distachyon. In addition, phylogenetic analysis of the plastid data was performed for a reduced subset of individuals. The nuclear simple sequence repeat (SSR) genetic analysis detected medium to high genetic diversity, with a strong south-to-north genetic structure cline, and a high selfing rate in B. hybridum. Comparative genetic analysis showed a close relatedness of current B. hybridum D allelic profiles with those of B. distachyon, but a lack of similarity with those of B. stacei, suggesting another B. stacei source for the B. hybridum S alleles. Plastid analysis detected three different bidirectional allopolyploidization events: two involved distinct B. distachyon-like ancestors and one involved a B. stacei-like ancestor. The south-eastern Iberian Peninsula B. hybridum populations were more genetically diverse and could have originated from at least two hybridization events whereas north-eastern/north-western Iberian Peninsula B. hybridum populations were less diverse and may have derived from at least one hybridization event. The genetic and evolutionary evidence supports the plausible in situ origin of the south-eastern and northern Iberian Peninsula B. hybridum allopolyploids from their respective local B. distachyon and unknown B. stacei ancestors. The untapped multiple origins and genetic variation detected in these B. hybridum populations opens the way to future evolutionary analysis of allopolyploid formation and genomic dominance and expression in the B. hybridum-B. distachyon-B. stacei grass model complex.

Effect of partial selfing and polygenic selection on establishment in a new habitat

This article analyzes how partial selfing in a large source population influences its ability to colonize a new habitat via the introduction of a few founder individuals. Founders experience inbreeding depression due to partially recessive deleterious alleles as well as maladaptation to the new environment due to selection on a large number of additive loci. I first introduce a simplified version of the inbreeding history model to characterize mutation‐selection balance in a large, partially selfing source population under selection involving multiple nonidentical loci. I then use individual‐based simulations to study the eco‐evolutionary dynamics of founders establishing in the new habitat under a model of hard selection. The study explores how selfing rate shapes establishment probabilities of founders via effects on both inbreeding depression and adaptability to the new environment, and also distinguishes the effects of selfing on the initial fitness of founders from its effects on the long‐term adaptive response of the populations they found. A high rate of (but not complete) selfing is found to aid establishment over a wide range of parameters, even in the absence of mate limitation. The sensitivity of the results to assumptions about the nature of polygenic selection is discussed.

Marker‐assisted parentage analysis reveals high individual selfing rates in tetraploid red clover genotypes selected for seed yield

AbstractSeed yield is a major breeding target in tetraploid red clover. We investigated if marker‐assisted parentage analysis can identify progeny plants with two high seed‐yielding parents in tetraploid red clover and if this technique is more advantageous than traditional half‐sib selection. Parentage analysis was successfully performed on the progeny from the 10% highest seed‐yielding genotypes from a second‐cycle family selection trial: 16.0% of progeny were identified with a high seed‐yielding father. However, progeny plants with two high seed‐yielding parents did not produce more seeds than traditionally selected progeny (27.3 g vs. 30.7 g/plant, respectively). The 10% highest seed‐yielding genotypes displayed on average 2% self‐fertilization. Four genotypes were self‐fertile with individual selfing rates up to 20%. Our results discourage the use of marker‐assisted parentage analysis to improve seed yield in tetraploid red clover when the material has been preselected for seed yield. Breeders should be aware that intensive selection for seed yield in tetraploid red clover may inadvertently lead to selection for increased self‐fertility, which may increase inbreeding in the long term.