Postcopulatory sexual selection reduces genetic diversity in experimental populations of Caenorhabditis elegans

Postcopulatory sexual selection affects the evolution of numerous features ranging from mating behavior to seminal fluid toxicity to the size of gametes. In an earlier study of the effect of sperm competition risk on sperm size evolution, experimental populations of the nematode Caenorhabditis elegans were maintained either by outcrossing (sperm competition present) or by selfing (no sperm competition), and after 60 generations, significantly larger sperm had evolved in the outcrossing populations. To determine the effects of this selection on population genetic variation, we assessed genetic diversity in a large number of loci using random amplification of polymorphic DNA-PCR. Nearly 80% of the alleles present in parental strain populations persisted in the 6 experimental populations after the 60 generations and, despite a 2.2-fold difference in expected heterozygosity, the resulting levels of genetic variation were equivalent between the outcrossing and selfing experimental populations. By inference, we conclude that genetic hitchhiking due to sexual selection in the experimental populations dramatically reduced genetic diversity. We use the levels of variation in the selfing populations as a control for the effects of drift, and estimate the strength of sexual selection to be strong in obligatorily outcrossing populations. Although sequential hermaphrodites like C. elegans probably experience little sexual selection in nature, these data suggest that sexual selection can profoundly affect diversity in outcrossing taxa.     

普通野生稻小种群的交配系统与遗传多样性

小种群的遗传动态是保育遗传学关注的核心问题之一,而种群遗传动态又与交配系统密切相关.普通野生稻(Oryza rufipogon Griff.)是具有重要经济价值的濒危物种,目前其种群规模都较小,研究其小种群交配系统与遗传变异性对普通野生稻的保护具有重要意义.运用7对SSR引物,对采自江西东乡普通野生稻小种群的36份种茎和其中20个家系共计601份子代进行了分析.结果显示:该种群的表观异交率为0.318,多位点法估计(MLTR)的多位点异交率为0.481;50%以上的子代共享亲本,非随机交配明显;东乡普通野生稻种群交配系统属于混合交配类型.比较亲本和子代种群的遗传变异性显示:子代种群比亲本种群遗传变异性更丰富;子代种群的杂合子不足与种群变小自交比例上升有关;而亲本种群杂合子过剩可能与杂合基因型的选择优势有关.这些结果说明创造条件扩大种群规模对普通野生稻的原生境保护显得尤为重要.     

Mating system and the evolution of quantitative traits: an experimental study of Mimulus guttatus

Abstract The mating system of a population profoundly influences its evolution. Inbreeding alters the balance of evolutionary forces that determine the amount of genetic variation within a population. It redistributes that variation among individuals, altering heritabilities and genetic correlations. Inbreeding even changes the basic relationships between these genetic statistics and response to selection. If populations differing only in mating system are exposed to the same selection pressures, will they respond in qualitatively different ways? Here, we address this question by imposing selection on an index of two negatively correlated traits (flower size and development rate) within experimental populations that reproduce entirely by outcrossing, entirely by self‐fertilizing, or by a mixture of outcrossing and selfing. Entirely selfing populations responded mainly by evolving larger flowers whereas outcrossing populations also evolved more rapid development. Divergence occurred despite an equivalent selection regime and no direct effect of mating system on fitness. The study provides an experimental demonstration of how the interaction of selection, genetic drift, and mating system can produce dramatic short‐term changes in trait means, variances, and covariances.     

MUTATIONAL MELTDOWN IN SELFING ARABIDOPSIS LYRATA

The majority of plant species and many animals are hermaphrodites, with individuals expressing both female and male function. Although hermaphrodites can potentially reproduce by self‐fertilization, they have a high prevalence of outcrossing. The genetic advantages of outcrossing are described by two hypotheses: avoidance of inbreeding depression because selfing leads to immediate expression of recessive deleterious mutations, and release from drift load because self‐fertilization leads to long‐term accumulation of deleterious mutations due to genetic drift and, eventually, to extinction. I tested both hypotheses by experimentally crossing Arabidopsis lyrata plants (self‐pollinated, cross‐pollinated within the population, or cross‐pollinated between populations) and measuring offspring performance over 3 years. There were 18 source populations, each of which was either predominantly outcrossing, mixed mating, or predominantly selfing. Contrary to predictions, outcrossing populations had low inbreeding depression, which equaled that of selfing populations, challenging the central role of inbreeding depression in mating system shifts. However, plants from selfing populations showed the greatest increase in fitness when crossed with plants from other populations, reflecting higher drift load. The results support the hypothesis that extinction by mutational meltdown is why selfing hermaphroditic taxa are rare, despite their frequent appearance over evolutionary time.     

Consequences of multiple mating‐system shifts for population and range‐wide genetic structure in a coastal dune plant

Evolutionary transitions from outcrossing to selfing can strongly affect the genetic diversity and structure of species at multiple spatial scales. We investigated the genetic consequences of mating‐system shifts in the North American, Pacific coast dune endemic plant Camissoniopsis cheiranthifolia (Onagraceae) by assaying variation at 13 nuclear (n) and six chloroplast (cp) microsatellite (SSR) loci for 38 populations across the species range. As predicted from the expected reduction in effective population size (N_e) caused by selfing, small‐flowered, predominantly selfing (SF) populations had much lower nSSR diversity (but not cpSSR) than large‐flowered, predominantly outcrossing (LF) populations. The reduction in nSSR diversity was greater than expected from the effects of selfing on N_e alone, but could not be accounted for by indirect effects of selfing on population density. Although selfing should reduce gene flow, SF populations were not more genetically differentiated than LF populations. We detected five clusters of nSSR genotypes and three groups of cpSSR haplotypes across the species range consisting of parapatric groups of populations that usually (but not always) differed in mating system, suggesting that selfing may often initiate ecogeographic isolation. However, lineage‐wide genetic variation was not lower for selfing clusters, failing to support the hypothesis that selection for reproductive assurance spurred the evolution of selfing in this species. Within three populations where LF and SF plants coexist, we detected genetic differentiation among diverged floral phenotypes suggesting that reproductive isolation (probably postzygotic) may help maintain the striking mating‐system differentiation observed across the range of this species.     

The influence of persistent individual differences and age at maturity on effective population size

Ratios of effective populations size, N(e), to census population size, N, are used as a measure of genetic drift in populations. Several life-history parameters have been shown to affect these ratios, including mating system and age at sexual maturation. Using a stochastic matrix model, we examine how different levels of persistent individual differences in mating success among males may affect N(e)/N, and how this relates to generation time. Individual differences of this type are shown to cause a lower N(e)/N ratio than would be expected when mating is independent among seasons. Examining the way in which age at maturity affects N(e)/N, we find that both the direction and magnitude of the effect depends on the survival rate of juveniles in the population. In particular, when maturation is delayed, lowered juvenile survival causes higher levels of genetic drift. In addition, predicted shifts in N(e)/N with changing age at maturity are shown to be dependent on which of the commonly used definitions of census population size, N, is employed. Our results demonstrate that patterns of mating success, as well as juvenile survival probabilities, have substantial effects on rates of genetic drift.     

Small reductions in corolla size and pollen: ovule ratio,but no changes in flower shape in selfing populations of the North American <Emphasis Type="Italic">Arabidopsis lyrata</Emphasis>

The shift from outcrossing to selfing is often accompanied by striking changes in floral morphology towards a “selfing syndrome”, which is characterized by flowers with reduction in size, pollen: ovule (P/O) ratio, and herkogamy. This study aims to test whether such changes have occurred in the North American Arabidopsis lyrata, which is of particular interest because of the relatively recent transitions to selfing in this system. Flower size, flower shape, herkogamy levels, P/O ratio, and floral integration of six self-incompatible (outcrossing) and six self-compatible (selfing) populations of A. lyrata were measured in a common environment using conventional and geometric morphometrics methods. Although selfers had on average 9.2% smaller corollas, 8.4% longer pistils, and 21.5% lower P/O ratios than outcrossers, there were no differences in shape, floral integration, and herkogamy between outcrossing and selfing populations. Moreover, most variation in floral traits was explained by population genetic background rather than by mating system. We conclude that selfing populations in A. lyrata have not evolved a selfing syndrome.     

Patterns of genetic diversity in outcrossing and selfing populations of Arabidopsis lyrata

Arabidopsis lyrata is normally considered an obligately outcrossing species with a strong self-incompatibility system, but a shift in mating system towards inbreeding has been found in some North American populations (subspecies A. lyrata ssp. lyrata). This study provides a survey of the Great Lakes region of Canada to determine the extent of this mating system variation and how outcrossing rates are related to current population density, geographical distribution, and genetic diversity. Based on variation at microsatellite markers (progeny arrays to estimate multilocus outcrossing rates and population samples to estimate diversity measures) and controlled greenhouse pollinations, populations can be divided into two groups: (i) group A, consisting of individuals capable of setting selfed seed (including autogamous fruit set in the absence of pollinators), showing depressed outcrossing rates (T(m) = 0.2-0.6), heterozygosity (H(O) = 0.02-0.06) and genetic diversity (H(E) = 0.08-0.10); and (ii) group B, consisting of individuals that are predominantly self-incompatible (T(m) > 0.8), require pollinators for seeds set, and showing higher levels of heterozygosity (H(O) = 0.13-0.31) and diversity (H(E) = 0.19-0.410). Current population density is not related to the shift in mating system but does vary with latitude. Restricted gene flow among populations was evident among all but two populations (F(ST) = 0.11-0.8). Group A populations were more differentiated from one another (F(ST) = 0.78) than they were from group B populations (F(ST) = 0.59), with 41% of the variation partitioned within populations, 47% between populations, and 12% between groups. No significant relationship was found between genetic and geographical distance. Results are discussed in the context of possible postglacial expansion scenarios in relation to loss of self-incompatibility.     

High selfing and high inbreeding depression in peripheral populations of Juncus atratus

The mating system of a plant is the prime determinant of its population genetic structure. However, mating system effects may be modified by postzygotic mechanisms like inbreeding depression. Furthermore, historical as well as contemporary ecological factors and population characteristics, like the location within the species range can contribute to genetic variability. Using microsatellite markers we assessed the population genetic structure of the wind-pollinated Juncus atratus in 16 populations from peripheral and nearly central areas of the distribution range and studied the mating system of the species. In three peripheral populations, outcrossing rates at seeds stage were low (mean t(m) = 5.6%), suggesting a highly autogamous mating system. Despite this fact, on adult stage both individual heterozygosity (mean H(O) = 0.48) and gene diversity (mean H(E) = 0.58) were high even in small populations. Inbreeding coefficients were consistently low among all populations (mean F(IS) = 0.15). Within the three peripheral populations indirect estimates of lifetime inbreeding depression were surprisingly high (delta(eq) = 0.96) and inbreeding depression could be shown to act mostly on early seedling establishment. Similar conditions of autogamy combined with high inbreeding depression are typical for plants with a large lifetime genomic mutation rate that cannot avoid selfing by geitonogamy. However, the results presented here are unexpected for small-statured, herbaceous plants. Substantial genetic differentiation among all populations was found (mean F(ST) = 0.24). An isolation-by-distance pattern was apparent on large scale but not on local scale suggesting that the overall pattern was largely influenced by historical factors, e.g. colonization, whereas locally genetic drift was of greater importance than gene flow. Peripheral populations exhibited lower genetic diversity and higher inbreeding coefficients when compared with subcentral populations.     

EFFECTIVE POPULATION SIZE AND GENETIC DRIFT IN TRISTYLOUS EICHHORNIA PANICULATA (PONTEDERIACEAE)

Populations of the tristylous, annual Eichhornia paniculata are markedly differentiated with respect to frequency of mating types. This variation is associated with evolutionary changes in mating system, from predominant outcrossing to high self-fertilization. To assess the potential influence of genetic drift acting on this variation, we estimated effective population size in 10 populations from northeastern Brazil using genetic and demographic methods. Effective size (N_e) was inferred from temporal changes in allele frequency at two to eight isozyme loci and also calculated using five demographic variables: 1) the number of flowering individuals (N); 2) temporal fluctuations in N; 3) variance in flower number; 4) frequency of mating types; and 5) selfing rate. Average N_e based on isozyme data was 15.8, range 3.4–70.6, and represented a fraction (mean N_e/N = 0.106) of the census number of individuals (mean N = 762.8; range: 30.5–5,040). Temporal variation in N and variance in flower number each reduced N_e to about a half of N whereas mating type frequencies and selfing rate caused only small reductions in N_e relative to N. All estimates of N_e based on demographic variables were considerably larger than those obtained from genetic data. The two kinds of estimates were in general agreement, however, when all demographic variables were combined into a single measure. Monte Carlo simulations indicated that effective size must be fewer than about 40 for drift to overcome the frequency-dependent selection that maintains the polymorphism for mating type. Applying the average N_e/N value to 167 populations censused in northeastern Brazil indicated that 72% had effective sizes below this number. This suggests that genetic drift is likely to play a dominant role in natural populations of E. paniculata.     

Breeding systems in flowering plants and the control of variability

Plants have three basic means of reproduction, by outcrossing, by selfing, and asexually. In most plant populations, at least two and often all three of these options are everpresent, so that individuals adopt mixed mating strategies at evolutionarily stable strategy (ESS) threshholds. Because mating systems are genetically controlled and affect genotype structure, they are liable to feedback. Productive habitats with a large standing crop are more likely to favour outcrossing, while unproductive habitats may favour asexuality or selfing, so that mating systems may change through seral development, even within the same species. Outcrossing tends to break up linkage disequilibria, but may also favour the creation of adaptive linkage groups. Mechanisms whereby male sexual selection, small population size and selfing can influence the genetic structure of populations are examined.     

RECENT EVOLUTION AND DIVERGENCE AMONG POPULATIONS OF A RARE MEXICAN ENDEMIC,CHIHUAHUA SPRUCE,FOLLOWING HOLOCENE CLIMATIC WARMING

Fragmentation and reduction in population size are expected to reduce genetic diversity. However, examples from natural populations of forest trees are scarce. The range of Chihuahua spruce retreated northward and fragmented coincident with the warming climate that marked the early Holocene. The isolated populations vary from 15 to 2441 trees, which provided an opportunity to test whether census number is a good predictor of genetic diversity. Mean expected heterozygosity, H_e, based on 24 loci in 16 enzyme systems, was 0.093 for 10 sampled populations, which is within the range reported for conifers. However, estimates varied more than twofold among populations and H_e was closely related to the logarithm of the number of mature trees in the population (r_He,N = 0.93). Diversity among populations, F_ST, was 24.8% of the total diversity, which is higher than that observed in almost all conifer species studied. Nei's genetic distance, D, was not related to geographic distance between populations, and D? was 0.033, which is higher than estimates for most wide-ranging species. Most populations had excess homozygosity and the fixation index, F_IS, was higher than that reported for all but one species of conifer. Nm, the number of migrants per generation, was 0.43 to 0.76, depending on estimation procedure, and is the smallest observed in conifers. The data suggest that populations of Chihuahua spruce have differentiated by drift and that they are effectively isolated. The results illustrate how a combination of paleontological observation and molecular markers can be used to illuminate recent evolutionary events. Multilocus estimates of outcrossing for two small populations were zero (complete selfing) and 0.153, respectively, which are in striking contrast to the near complete outcrossing observed in most conifers. The high fixation index and a high proportion of empty seeds (45%) suggest that inbreeding may be a serious problem for conservation of Chihuahua spruce.     

Mixed mating in natural populations of the chestnut blight fungus, Cryphonectria parasitica

As in plants, fungi exhibit wide variation in reproductive strategies and mating systems. Although most sexually reproducing fungi are either predominantly outcrossing or predominantly selfing, there are some notable exceptions. The haploid, ascomycete chestnut blight pathogen, Cryphonectria parasitica, has previously been shown to have a mixed mating system in one population in USA. In this report, we show that both selfing and outcrossing occur in 10 additional populations of C. parasitica sampled from Japan, Italy, Switzerland and USA. Progeny arrays from each population were assayed for segregation at vegetative incompatibility (vic) and DNA fingerprinting loci. Outcrossing rates (t(m)) were estimated as the proportion of progeny arrays showing segregation at one or more loci, corrected by the probability of nondetection of outcrossing (alpha). Estimates of t(m) varied from 0.74 to 0.97, with the lowest rates consistently detected in USA populations (0.74-0.78). Five populations (four in USA and one in Italy) had t(m) significantly less than 1, supporting the conclusion that these populations exhibit mixed mating. The underlying causes of variation in outcrossing rates among populations of C. parasitica are not known, but we speculate that--as in plants--outcrossing is a function of ecological, demographic and genetic factors.     

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.     

Limiting inbreeding in disjunct and isolated populations of a woody shrub

Pollen movements and mating patterns are key features that influence population genetic structure. When gene flow is low, small populations are prone to increased genetic drift and inbreeding, but naturally disjunct species may have features that reduce inbreeding and contribute to their persistence despite genetic isolation. Using microsatellite loci, we investigated outcrossing levels, family mating parameters, pollen dispersal, and spatial genetic structure in three populations of Hakea oldfieldii, a fire‐sensitive shrub with naturally disjunct, isolated populations prone to reduction in size and extinction following fires. We mapped and genotyped a sample of 102 plants from a large population, and all plants from two smaller populations (28 and 20 individuals), and genotyped 158–210 progeny from each population. We found high outcrossing despite the possibility of geitonogamous pollination, small amounts of biparental inbreeding, a limited number of successful pollen parents within populations, and significant correlated paternity. The number of pollen parents for each seed parent was moderate. There was low but significant spatial genetic structure up to 10 m around plants, but the majority of successful pollen came from outside this area including substantial proportions from distant plants within populations. Seed production varied among seven populations investigated but was not correlated with census population size. We suggest there may be a mechanism to prevent self‐pollination in H. oldfieldii and that high outcrossing and pollen dispersal within populations would promote genetic diversity among the relatively small amount of seed stored in the canopy. These features of the mating system would contribute to the persistence of genetically isolated populations prone to fluctuations in size.     

Ecological context of the evolution of self-pollination in Clarkia xantiana: population size, plant communities, and reproductive assurance

The repeated evolutionary transition from outcrossing to self-pollination in flowering plants has been suggested to occur because selfing provides reproductive assurance. Reports from biogeographical and ecological surveys indicate that selfing taxa are often associated with stressful and ephemeral environments, situations in which plant abundance is low (e.g., Baker's law) and with novel plant communities, however experimental tests of ecological hypotheses are few. In this study, we examined the ecological context of selection on mating system traits (herkogamy and protandry) in a California annual, Clarkia xantiana, where natural selfing populations differ from outcrossing populations in that they are often of small size or low density and occur mainly outside the range of pollinator-sharing congeners. We constructed artificial populations of plants with broad genetic variation in floral traits and manipulated two ecological factors, plant population size, and the presence versus absence of pollinator-sharing congeners, in the center of the geographic range of outcrossing populations. We found evidence for context-dependent selection on herkogamy and protandry via female fitness in which reduced traits, which promote autonomous selfing, were favored in small populations isolated from congeners whereas selection was comparatively weak in large populations or when congeners were present. In small, isolated populations, the fertility of plants with low herkogamy or protandry was elevated by 66% and 58%, respectively, compared to those with high herkogamy or protandry. The presence of pollinator-sharing congeners augmented bee visitation rates to C. xantiana flowers by 47% for all bees and by 93% for pollen specialists. By facilitating pollinator visitation, congeners mitigated selection on mating system traits in small populations, where outcross mating success is often low (the Allee effect). We also found support for the hypothesis that pollinator availability directly influenced variation in the strength of selection on herkogamy among populations. The striking parallels between our experimental results and patterns of variation in ecological factors across the geographic range of outcrossing and selfing populations suggest that reproductive assurance may play a central role in directing mating system evolution in C. xantiana.     

Reproductive success and effective population size in woodrats (Neotoma macrotis)

Discrepancies between the census size and the genetically effective size of populations (N(e)) can be caused by a number of behavioural and demographic factors operating within populations. Specifically, strong skew in male reproductive success, as would be expected in a polygynous mating system, could cause a substantial decrease in N(e) relative to census size. Because the mating system of Neotoma macrotis had previously been described as one nearing harem polygyny, I examined the distribution of reproductive success and genetic variation within a population of this species. Combining genetic data and three years of field observations, I show that variance in reproductive success does not deviate from poisson expectations within either sex and variance in success is similar between the sexes. Furthermore, both males and females had multiple partners across litters in addition to some evidence of multiple paternity within litters. Despite a lack of strong skew in reproductive success, an estimate of N(e) based on a number of demographic parameters suggests that the ratio of N(e)/N in this population is 0.48. Although the ratio of N(e)/N suggests that the population is experiencing higher rates of genetic drift than would be expected based on census size alone, the population maintains high levels of genetic diversity. Estimates of neighbourhood size and patterns of recruitment to the study site suggest that immigration plays an important role in this population and may contribute to the maintenance of high levels of genetic diversity.     

Variation in Outcrossing Rates in Eichhornia paniculata: Temporal Changes in Populations of Contrasting Style Morph Structure

Abstract Surveys of mating-system parameters in populations of the annual, self-compatible, tristylous, emergent aquatic, Eichhornia paniculata (Pontederiaceae) from N.E. Brazil and Jamaica have indicated that the species exhibits a wide range of outcrossing rates. To investigate whether temporal variation in outcrossing rate was also a feature of populations, open-pollinated families were sampled from five populations of contrasting style morph structure from N.E. Brazil over three consecutive years (1987–1989). Multilocus estimates of outcrossing rate ( t ) were obtained from assays of isozyme polymorphisms using starch gel electrophoresis. There was significant variation both among populations and between years in the frequency of outcrossing. Outcrossing in three tristylous populations was high ( t > 0.80), with relatively small fluctuations occurring over the three-year sampling period. In contrast, in a dimorphic and monomorphic population considerable self-fertilization occurred and the frequency of outcrossing declined significantly from 1987 to 1989 in both populations. In the dimorphic population, increased selfing was associated with a marked reduction in population size and an increase in the frequency of selfing variants of the mid-styled morph. The significance of temporal variation in outcrossing frequency in plant populations is discussed in relation to its effect on population genetic structure and recent models of mating-system evolution.     

Sex ratio, reproductive mode and genetic diversity in Triops cancriformis

1. Aquatic invertebrates display a wide array of alternative reproductive modes from apomixis to hermaphroditism and cyclical parthenogenesis. These have important effects on genetic diversity and population structure. Populations of the 'living fossil' Triops cancriformis display a range of sex ratios, and various reproductive modes are thought to underlie this variation. Using sex ratio information and histological analyses European populations have been inferred to be gonochoric (with separate males and females), selfing hermaphroditic and androdioecious, a rare reproductive mode in which selfing hermaphrodites coexist with variable proportions of males. In addition, some populations have been described as meiotic parthenogens.
2. Here we use population genetic analysis using microsatellite loci in populations with a range of sex ratios including a gonochoric population, and marker segregation patterns in heterozygote individuals reared in isolation, to clarify the reproductive mode in this species.
3. Our data show that populations in general have very low levels of genetic diversity. Non-gonochoric populations show lower genetic diversity, more heterozygote deficiencies, higher inbreeding coefficients and stronger linkage disequilibria than the gonochoric population. The maintenance of some heterozygosity in populations is consistent with some male influence in T. cancriformis populations, as would be expected from an androdioecious reproductive system. Results of marker segregation in eggs produced in isolation from non-gonochoric populations indicate that meiosis occurs and are consistent with two reproductive modes: selfing hermaphroditism and a type of ameiotic parthenogenesis.
4. Overall, our data indicate that androdioecy and selfing hermaphroditism are the most likely reproductive modes of non-gonochoric European Triops populations. Triops populations are strongly structured, suggesting high genetic drift and low levels of gene flow.     

Mitosis, stature and evolution of plant mating systems: low-Phi and high-Phi plants

There is a long-recognized association in plants between small stature and selfing, and large stature and outcrossing. Inbreeding depression is central to several hypotheses for this association, but differences in the evolutionary dynamics of inbreeding depression associated with differences in stature are rarely considered. Here, we propose and test the Phi model of plant mating system evolution, which assumes that the per-generation mutation rate of a plant is a function of the number of mitoses (Phi) that occur from zygote to gamete, and predicts fundamental differences between low-Phi (small-statured) and high-Phi (large-statured) plants in the outcomes of the joint evolution of outcrossing rate and inbreeding depression. Using a large dataset of published population genetic studies of angiosperms and conifers, we compute fitted values of inbreeding depression and deleterious mutation rates for small- and large-statured plants. Consistent with our Phi model, we find that populations of small-statured plants exhibit a range of mating systems, significantly lower mutation rates, and intermediate inbreeding depression, while large-statured plants exhibit very high mutation rates and the maximum inbreeding depression of unity. These results indicate that (i) inbred progeny typically observed in large-statured plant populations are completely lost prior to maturity in nearly all populations; (ii) evolutionary shifts from outcrossing to selfing are generally not possible in large-statured species, rather, large-statured species are more likely to evolve mating systems that avoid selfing such as self-incompatibility and dioecy; (iii) destabilization of the mating system-high selfing rate with high-inbreeding depression-might be a common occurrence in large-statured species; and (iv) large-statured species in fragmented populations might be at higher risk of extinction than previously thought. Our results help to unify and simplify a large and diverse field of research, and serve to emphasize the importance that developmental and genetic constraints play in the evolution of plant mating systems.