Rate Of Self-fertilization Research Articles

High but variable outcrossing rates in the invasive Geranium carolinianum (Geraniaceae).

• Premise of the study: Mating system plays an important role in population establishment and persistence, maintenance of genetic variation, and adaptive ability, especially for invasive species that colonize new environments to which they may be poorly adapted. In mixed-mating species, population differences in self-fertilization rates often arise due to variation in local ecological conditions (e.g., pollinator or mate availability) or genetic variation in traits promoting selfing or outcrossing. Knowledge of how and why selfing rates vary can help us understand how populations respond to different environments, how this affects patterns of genetic variation, and the role of mating systems in biological invasions.• Methods: We determined outcrossing rates in invasive (China) and native (US) populations of the weedy annual Geranium carolinianum in natural populations and an open-pollinated common garden to answer the following questions: To what extent do populations vary in mating system? Do invasive populations differ from native populations? Is interpopulation variation in mating system under genetic control?• Key results: Despite having many characteristics of selfing species, we found high variation in outcrossing rates (tm from 0.1 to 1.0) resulting from both environmental and genetic variation. Outcrossing rates were generally high and plastic in the Chinese populations.• Conclusions: A delayed selfing mechanism provides reproductive assurance while allowing facultative outcrossing when mates are not limiting. In invasive populations, high outcrossing rates were facilitated by large admixed founding populations, promoting local adaptation and the maintenance of genetic diversity.

Extensive pollen immigration and no evidence of disrupted mating patterns or reproduction in a highly fragmented holm oak stand

Aims Forest fragmentation and reduced tree population densities can potentially have negative impacts on mating patterns, offspring genetic diversity and reproductive performance. The aim of the present study is to test these hypotheses comparing an extremely fragmented, low tree density (~0.02 trees/ha) holm oak (Quercus ilex L.) stand from Central Spain with a nearby high tree density stand (~50 trees/ha). Methods We genotyped adult trees and seeds from the low-density stand (436 seeds from 15 families) and the high-density stand (404 seeds from 11 families) using nine microsatellite markers. With these data, we performed paternity analyses, determined pollen flow, mating patterns and pollen pool structure, and estimated progeny genetic diversity in both stands. We also studied seed set and production and performed a pollen supplementation experiment to determine whether reduced tree density has limited foreign pollen availability. Important Findings We have found extensive pollen immigration (>75%) into the low tree density stand and Monte Carlo simulations revealed that pollen moves larger distances than expected from null models of random dispersal. Mating patterns and differentiation of pollen pools were similar in the high-density stand and the low-density stand but we found higher inter-annual differentiation of pollen pools in the for mer. Progeny genetic diversity and self-fertilization rates did not differ between the low-density stand and the high-density stand. Seed set rates were significantly lower in the low-density stand than in the high-density stand and experimental cross-pollen supplementation evidenced that foreign pollen availability is indeed a limiting factor in the former. However, seed crops did not differ between the low-density stand and the high-density stand, indicating that limitation of foreign pollen is not likely to be of great concern in terms of reduced seed production and potential recruitment. Poor forest regeneration due to other ecological and human factors is probably a more important threat for the persistence of fragmented and low tree density stands than reduced pollen flow and only extremely small and isolated tree populations would be expected to suffer severe loss of genetic diversity in the long term.

Genetic and experimental evidence for a mixed-age, mixed-origin bank of kelp microscopic stages in southern California.

Laboratory studies have demonstrated that the microscopic stages of kelps can rapidly resume development from a delayed state. Like terrestrial seeds or aquatic resting eggs, banks of delayed kelp stages may supplement population recovery after periods of stress, playing an important role for kelp populations that experience adult sporophyte absences due to seasonal or interannual disturbances. We found that removing the microscopic stages from natural rock substratum could prevent the appearance of juvenile kelp sporophytes for three months and the establishment of a diverse kelp assemblage for over four months within a southern California kelp forest. Juveniles were observed within one month in plots where microscopic stages were left intact, which may confer an advantage for the resulting sporophytes as they attain larger sizes before later recruiting neighbors. Microsatellite diversity was high (expected heterozygosity HE approximately 0.9) for juveniles and adults within our sites. Using a microsatellite-based parentage analysis for the dominant kelp, Macrocystis pyrifera, we estimated that a portion of the new M. pyrifera sporophyte recruits had originated from their parents at least seven months after their parents had disappeared. Similar delay durations have been demonstrated in recent laboratory studies. Additionally, our results suggest that zoospore dispersal distances > 50 m may be supported by including additional microsatellite loci in the analysis. We propose a mixed-age and, potentially, a mixed-origin bank of M. pyrifera gametophytes promotes maximal genetic diversity in recovering populations and reduces population genetic subdivision and self-fertilization rates for intact populations by promoting the survival of zoospores dispersed > 10 m and during inhospitable environmental conditions.

Paternity testing in an autotetraploid alfalfa breeding polycross

Determining unknown parentage in autotetraploid alfalfa (Medicago sativa L.) (2n = 4x = 32) can improve breeding gains. An exclusion analysis-based paternity testing SAS code, amenable to genotyping errors, is presented for autotetraploid species utilizing co-dominant molecular markers with ambiguous dosage. To demonstrate the paternity testing SAS code, 19 SSR loci were genotyped and analyzed on 1,107 progeny from a commercial, isolated, clonally replicated, 16-parent alfalfa breeding polycross which was pollinated by leafcutter bees (Megachile rotundata F.). Paternal assignment success rate was over 90 %. Among typed progeny, 45 % were the result of self-fertilization. Significant differences were detected between the 15 parents that produced seed and were observed as fathers for (1) total fertilizing pollen contribution (% deviation from expectation), (2) self-fertilization rates (%), and (3) outcrossing fertilizing pollen contribution (% deviation from expectation). Physical within-cage distance between parental plants was correlated with outcrossing fertilizing pollen frequency (negative power function). Parental seed yield was positively correlated with total fertilizing pollen contribution, particularly with self-fertilization rates (42 % self-fertilization and 17 % outcrossing). These correlations suggest that selecting for increased seed yield may result in indirect selection for increased self-fertilization rates. Parental total fertilizing pollen contribution was 62 % determined by outcrossing and 35 % by self-fertilization. This study cautions alfalfa breeders that heretofore unconsidered sources of inbreeding could be present in some breeding materials. This study also provides cost effective and easy to use molecular genetic tools for detecting, managing, and/or selecting against (through breeding) those sources of inbreeding.

Drought alters the expression of mating system traits in two species of Clarkia

Variation in mating system traits can have important consequences for plant populations by affecting reproductive assurance, the expression of inbreeding depression, and the colonization of and persistence in new or altered habitats. Environmental stressors, such as drought, have been hypothesized to induce higher rates of self-fertilization, yet this hypothesis has rarely been tested. Here we measure the response of two sister species of self-compatible annual herbs from contrasting habitats, Clarkia breweri and C. concinna, to an experimentally imposed greenhouse drought treatment. We find that the species differ in their baseline per-flower autogamy rates and the degree of spatial and temporal separation of male and female function within their flowers. Both species show a reduction in temporal separation of anthesis and stigma receptivity with the drought treatment. However, the species from the more mesic habitat, C. concinna, increases its low autogamy rate under drought conditions, whereas the species from the more xeric habitat, C. breweri, decreases its high autogamy rate under drought conditions. Neither species showed a response to drought in flower size or anther-stigma distance. Our results demonstrate that the induction of selfing under environmental stress cannot be assumed and that, in this case, the developmental timing of flower maturation is more plastic than floral morphology.

Detecting selection along environmental gradients: analysis of eight methods and their effectiveness for outbreeding and selfing populations

Thanks to genome-scale diversity data, present-day studies can provide a detailed view of how natural and cultivated species adapt to their environment and particularly to environmental gradients. However, due to their sensitivity, up-to-date studies might be more sensitive to undocumented demographic effects such as the pattern of migration and the reproduction regime. In this study, we provide guidelines for the use of popular or recently developed statistical methods to detect footprints of selection. We simulated 100 populations along a selective gradient and explored different migration models, sampling schemes and rates of self-fertilization. We investigated the power and robustness of eight methods to detect loci potentially under selection: three designed to detect genotype-environment correlations and five designed to detect adaptive differentiation (based on F(ST) or similar measures). We show that genotype-environment correlation methods have substantially more power to detect selection than differentiation-based methods but that they generally suffer from high rates of false positives. This effect is exacerbated whenever allele frequencies are correlated, either between populations or within populations. Our results suggest that, when the underlying genetic structure of the data is unknown, a number of robust methods are preferable. Moreover, in the simulated scenario we used, sampling many populations led to better results than sampling many individuals per population. Finally, care should be taken when using methods to identify genotype-environment correlations without correcting for allele frequency autocorrelation because of the risk of spurious signals due to allele frequency correlations between populations.

Microsatellite markers for the native Texas perennial grass, Panicum hallii (Poaceae)

We developed microsatellites for Panicum hallii for studies of gene flow, population structure, breeding experiments, and genetic mapping. Next-generation (454) genomic sequence data were used to design markers. Eighteen robust markers were discovered, 15 of which were polymorphic across six accessions of P. hallii var. hallii. Fourteen of the markers cross-amplified in a P. capillare accession. For the 15 polymorphic markers, the total number of alleles per locus ranged from two to 26 (mean: 11.0) across six populations (11-19 individuals per population). Observed heterozygosity (mean: 0.031) was 13.7 times lower than the expected heterozygosity (mean: 0.426). The deficit of heterozygous individuals is consistent with P. hallii having a high rate of self-fertilization. These markers will be useful for studies in P. hallii and related species.

Mating Systems and Reproductive Success in Hermaphroditic Bay Scallop, Argopecten irradians irradians (Lamarck 1819), Inferred by Microsatellite‐based Parentage Analysis

In this study, we used microsatellite‐based parentage analysis to assess the mating system of a mass spawning bay scallop, Argopecten irradians irradians. Parental assignment was performed on 237 offspring and 14 candidate parents. Overall, 95% of all offspring were unambiguously allocated to their putative parents based on information from six highly polymorphic loci. Offspring from both the self‐fertilization and cross‐fertilization style were observed, the large variation of self‐fertilization rate among individuals displays a mixed mode of mating. The high variances in reproductive success resulted in a decline in effective breeding numbers (Nb/N = 0.71). The self‐fertilization rate was 32.89% for this mass spawning and the coefficient of inbreeding was estimated at around 16.5% per generation. When selfed larvae were removed, the relationship between relatedness and individual reproductive success was proved insignificant between half‐sibings and unrelated, which suggest that selection pressures other than avoidance of inbreeding may be responsible for the evolution and maintenance of the self‐compatible mating system of A. i. irradians. These results lend insight to A. i. irradians cultivation in present China and provide a genetic basis for the development of more effective spawning techniques.

Mating system shifts on the trailing edge.

The trailing edges of species ranges are becoming a subject of increasing interest as the environment changes due to global warming. Trailing edge populations are likely to face extinction because of a decline in numbers and an inability to evolve new adaptations with sufficient speed. Discussions of character change in the trailing edge have focused on physiological, exomorphic and phenological traits. The mating pattern within populations has not been part of the discourse, in spite of the fact that the mating pattern may affect the ability of populations to respond to environmental change and to maintain their sizes. In this paper, the case is made that a substantial increase in self-fertilization rates may occur via plastic responses to stress. Small populations on the trailing edge are especially vulnerable to environmental change because of inadequate levels of cross-fertilization. Evidence is presented that a deficiency of cross-seed production is due to inadequate pollinator services and a paucity of self-incompatibility alleles within populations. Evidence also is presented that if plants are self-compatible, self-fertilization may compensate in part for this deficiency through a stress-induced increase in levels of self-compatibility and stress-induced alterations in floral morphology that elevate self-pollination. Whereas increased self-fertility may afford populations the time to adapt to their changing environments, it can be concluded that increased selfing is not a panacea for the ills of environmental change, because it will lead to substantial reductions in genetic diversity, which may render adaptation unlikely.

Effects of floral display size on male and female reproductive success in Mimulus ringens

The number of flowers blooming simultaneously on a plant may have profound consequences for reproductive success. Large floral displays often attract more pollinator visits, increasing outcross pollen receipt. However, pollinators frequently probe more flowers in sequence on large displays, potentially increasing self-pollination and reducing pollen export per flower. To better understand how floral display size influences male and female fitness, we manipulated display phenotypes and then used paternity analysis to quantify siring success and selfing rates. To facilitate unambiguous assignment of paternity, we established four replicate (cloned) arrays of Mimulus ringens, each consisting of genets with unique combinations of homozygous marker genotypes. In each array, we trimmed displays to two, four, eight or 16 flowers. When fruits ripened, we counted the number of seeds per fruit and assigned paternity to 1935 progeny. Siring success per flower declined sharply with increasing display size, while female success per flower did not vary with display. The rate of self-fertilization increased for large floral displays, but siring losses due to geitonogamous pollen discounting were much greater than siring gains through increased self-fertilization. As display size increased, each additional seed sired through geitonogamous self-pollination was associated with a loss of 9·7 seeds sired through outcrossing. Although total fitness increased with floral display size, the marginal return on each additional flower declined steadily as display size increased. Therefore, a plant could maximize fitness by producing small displays over a long flowering period, rather than large displays over a brief flowering period.

BEST OF BOTH WORLDS? ASSOCIATION BETWEEN OUTCROSSING AND PARASITE LOADS IN A SELFING FISH

Mixed-mating strategies (i.e., intermediate levels of self-fertilization and outcrossing in hermaphrodites) are relatively common in plants and animals, but why self-fertilization (selfing) rates vary so much in nature has proved difficult to explain. We tested the hypothesis that parasites help maintain mixed-mating using a partially selfing fish (Kryptolebias marmoratus) as a model. We show that outcrossed progeny in the wild are genetically more diverse and less susceptible to multiple parasite infections than their selfed counterparts. Given that outcrossing in K. marmoratus can only be attained by male-hermaphrodite matings, our data provide an explanation for the coexistence of males and hermaphrodites in androdioecious species where hermaphrodites are unable to outcross among themselves. Moreover, our study provides evidence that parasites contribute to maintaining mixed-mating in a natural animal population.

Flower Development and the Evolution of Self-fertilization in Amsinckia: The Role of Heterochrony

We studied the development of 26 flower traits under natural conditions in three clades of the genus Amsinckia (Boraginaceae). Each clade contained both a derived highly self-fertilizing taxon and an ancestral more highly outcrossing taxon. The more outcrossing taxa contained two flower morphs—pins and thrums—with opposite positioning of the sex organs (heterostyly). The highly selfing taxa had smaller flowers with sex organs in close proximity (homostyly). Growth trajectories were quantified over the entire or nearly the entire period from primordium initiation to flower opening. These trajectories were compared in the heterochronic framework and, in contrast with previous studies, character size was tracked over time rather than relative to another character. We focused on three hypotheses: (1) The distinct developmental trajectories leading to pins and thrums should be similar in all clades, while the trajectories leading to homostylous flowers might differ among clades. This was supported. Specifically, contrasting growth rates of stamen and pistil heights in heterostylous flowers caused pin and thrum flowers to have the reciprocal arrangement of anther and stigma heights. From the viewpoint of heterochrony, the decreased size (paedomorphosis) of the homostylous morph, compared to pins and thrums, resulted from decreased growth rate (neoteny) and earlier offset (progenesis) in all clades. Nevertheless, multiple heterochronic processes were involved in the mosaic development and evolution of homostylous flowers. (2) We tested the hypothesis that small, self-fertilizing flowers have reduced development times, one of the proposed selective advantages of increased self-fertilization rates. We found in contrast that developmental duration of homostylous flowers was either the same (two clades) or longer (one clade) compared to duration of pins and thrums. (3) Finally, we tested von Baer’s Law, which proposes that developmental differences among closely related taxa should arise later in development than differences among more distantly related taxa. Von Baer’s Law was supported strongly among homostyles, moderately among thrums and weakly among pins.

Reconstruction of pedigrees in clonal plant populations

We present a Bayesian method for the reconstruction of pedigrees in clonal populations using co-dominant genomic markers such as microsatellites and single nucleotide polymorphisms (SNPs). The accuracy of the algorithm is demonstrated for simulated data. We show that the joint estimation of parameters of interest such as the rate of self-fertilization is possible with high accuracy even with marker panels of moderate power. Classical methods can only assign a very limited number of statistically significant parentages in this case and would therefore fail. Statistical confidence is estimated by Markov Chain Monte Carlo (MCMC) sampling. The method is implemented in a fast and easy to use open source software that scales to large datasets with many thousand individuals.

Fifteen microsatellite loci for the decollate snail, Rumina decollata

We characterized 15 microsatellite loci from the decollate snail, Rumina decollata. Loci were screened in 21 individuals and several individuals of the congener Rumina saharica. There was ample allelic diversity (6–12 alleles per locus) but observed heterozygosity values were extremely low (0–0.421). This was expected given the high self-fertilization rate in this species. Ten of the 15 loci were successfully amplified in R. saharica. These loci provide tools for examining the population genetics and taxomomic boundaries in R. decollata and its allies.

Indel-Associated Mutation Rate Varies with Mating System in Flowering Plants

A recently proposed mutational mechanism, indel-associated mutation (IDAM), posits that heterozygous insertions/deletions (indels) increase the point mutation rate at nearby nucleotides due to errors during meiosis. This mechanism could have especially dynamic consequences for the evolution of plant genomes, because the high degree of variation in the rate of self-fertilization among plant species causes differences in the heterozygosity of alleles, including indel alleles, segregating in plant species. In this study, we investigated the consequences of IDAM for species differing in mating system using both forward population genetic simulations and genomewide DNA resequencing data from Arabidopsis thaliana, Oryza sativa, and Oryza rufipogon. Simulations of different levels of selfing suggest that the effect of IDAM on surrounding nucleotide diversity should decrease with increasing selfing rate. Further simulations incorporating selfing rates and the time of onset of selfing suggest that the time since the switch to selfing also affects patterns of nucleotide diversity due to IDAM. Population genetic analyses of A. thaliana and Oryza DNA sequence data sets empirically confirmed our simulation results, revealing the strongest effect of IDAM in the outcrossing O. rufipogon, a weaker effect in the recently evolved selfer O. sativa, and the weakest effect in the relatively ancient selfer A. thaliana. These results support the novel idea that differences in life history, such as the level of selfing, can affect the per-individual mutation rate among species.

Mating patterns in an indoor miniature Cryptomeria japonica seed orchard as revealed by microsatellite markers

To evaluate the practical application of an indoor seed orchard, we compared the seed production, final germination rates and mating patterns of potted Cryptomeria japonica D. Don clones in two miniature seed orchards: one in a glasshouse and the other outdoors. There was no statistically significant difference in total seed production between the orchards (P = 0.275, ANOVA). However, the final germination rate of seeds produced in the indoor orchard was significantly lower than that of seeds produced in the outdoor orchard (P < 0.01, Wilcoxon test). The average self-fertilization rate was higher in the indoor orchard (27.2%) than in the outdoor orchard (5.6%), and this parameter was strongly associated with the reduced final germination rates of the seeds harvested in the glasshouse. Pollen contamination was considerably lower in the indoor seed orchard (48.9% in the outdoor seed orchard and 4.4% in the indoor seed orchard). χ2 tests indicated that the paternal contributions of each constituent clone differed significantly in both orchards (P < 0.001 in both cases). The strong variations in paternal contributions among indoor orchard clones was significantly correlated with their respective pollen production capacities (Spearman ρ = 0.693, P < 0.001). To reduce the high self-fertilization rates, we recommend the use of orchard clones with similar pollen production capacity in indoor seed orchards.

Blooming phenology and self-incompatibility of some commercial cherry ( Prunus avium L.) cultivars in Iran

Self-sterility in cherry ( Prunus avium L.) is one of the most important problems in commercial cherry orchards in Iran and many other countries. The main objective of this research was to determine self-fertility rate and blooming phenology of 25 commercial cherry ( P. avium L.) cultivars in Iran. This study was conducted during 2006–2007. Results revealed that the stage of bud swell in all cultivars began late second decade of March, which lasts 8–11 days. In all cultivars the duration end bloom to petal fall stage took 5 days. Cherry cultivars were classified into three groups considering their overlapping pollination. The cultivars each group had a great overlapped pollination toward each other. The results also showed that there was a significant difference among cultivars in respect of the fruit set ( P < 1%). The fruit set of open pollination treatment among the cultivars varied in range of 7.36–41.58%. However fruit set was not observed in isolated pollination (artificial and natural self-pollination). In open pollination, the cultivars were also classified into four groups, i.e. weak fertility (Moreau and Ferracida), mean fertility (Siah Mashhad, Napelon, Proteva, Haj usefy, Lambert, Roshun, Sefid Ghermeze Baghno, Arak, Belamarka, Siah Shabestar, Ghermeze Rezaeyeh, Abardeh, Soraty Lavisan and Victoria), good fertility (Peshrase Mashhad, Dovoumras Mashhad, Bing, Shesheiy Mashhad, Siah Ghazvin and Sefid Rezaeyeh), very good fertility (Zarde Daneshkadeh, Dirrase Italia and No. 1 karaj). All cultivars were found to need cross-pollination and included in self-sterile group.

The Functional Transfer of Genes From the Mitochondria to the Nucleus: The Effects of Selection, Mutation, Population Size and Rate of Self-Fertilization

The transfer of mitochondrial genes to the nucleus is a recurrent and consistent feature of eukaryotic genome evolution. Although many theories have been proposed to explain such transfers, little relevant data exist. The observation that clonal and self-fertilizing plants transfer more mitochondrial genes to their nuclei than do outcrossing plants contradicts predictions of major theories based on nuclear recombination and leaves a gap in our conceptual understanding how the observed pattern of gene transfer could arise. Here, with a series of deterministic and stochastic simulations, we show how epistatic selection and relative mutation rates of mitochondrial and nuclear genes influence mitochondrial-to-nuclear gene transfer. Specifically, we show that when there is a benefit to having a mitochondrial gene present in the nucleus, but absent in the mitochondria, self-fertilization dramatically increases both the rate and the probability of gene transfer. However, absent such a benefit, when mitochondrial mutation rates exceed those of the nucleus, self-fertilization decreases the rate and probability of transfer. This latter effect, however, is much weaker than the former. Our results are relevant to understanding the probabilities of fixation when loci in different genomes interact.

EXPERIMENTAL EVIDENCE FOR FREQUENCY DEPENDENT SELF-FERTILIZATION IN THE GYNODIOECIOUS PLANT,SILENE VULGARIS

After over a half century of empirical and theoretical research regarding the evolution and maintenance of gynodioecy in plants, unexplored factors influencing the relative fitnesses of females and hermaphrodites remain. Theoretical studies suggest that hermaphrodite self-fertilization (selfing) rate influences the maintenance of gynodioecy and we hypothesized that population sex ratio may influence hermaphrodite selfing rate. An experimental test for frequency-dependent self-fertilization was conducted using replicated populations constructed with different sex ratios of the gynodioecious plant Silene vulgaris. We found that hermaphrodite selfing increased with decreased hermaphrodite frequency, whereas evidence for increased inbreeding depression was equivocal. We argue that incorporation of context dependent inbreeding into future models of the evolution of gynodioecy is likely to yield novel insights into sex ratio evolution.

Density‐Dependent Regulation of the Sex Ratio in an Annual Plant

Sex ratios are subject to strong frequency-dependent selection regulated by the mating system and the relative male versus female investment. In androdioecious plant populations, where males co-occur with hermaphrodites, the sex ratio depends on the rate of self-fertilization by hermaphrodites and on the relative pollen production of males versus hermaphrodites. Here, we report evolutionary changes in the sex ratio from experimental mating arrays of the androdioecious plant Mercurialis annua. We found that the progeny sex ratio depended strongly on density, with fewer males in the progeny of plants grown under low density. This occurred in part because of a plastic adjustment in pollen production by hermaphrodites, which produced more pollen when grown at low density than at high density. Our results provide support for the prediction that environmental conditions govern sex ratios through their effects on the relative fertility of unisexual versus hermaphrodite individuals.