Does inbreeding avoidance maintain gender dimorphism in Wurmbea dioica (Colchicaceae)?

The maintenance of females in gender dimorphic populations requires that they have a fitness advantage to compensate for their loss of male reproductive function. We assess whether inbreeding avoidance provides this advantage in two subdioecious Wurmbea dioica populations by estimating seed production, outcrossing rates and inbreeding depression. Fruiting males produced less than half as many seeds as females, owing to low outcrossing rates and early acting inbreeding depression. Inbreeding coefficients of fruiting males demonstrated that progeny were more inbred than their parents, implying that few selfed progeny reach maturity, as confirmed by inbreeding depression estimates that exceeded 0.85. In a glasshouse experiment, open-pollinated females exhibited a fitness advantage of 3.7 relative to fruiting males, but when we increased fruiting male outcrossing rate, female advantage was only 1.4. This reduced advantage is insufficient to maintain females if nuclear genes control sex. Thus, inbreeding avoidance could maintain females at high frequencies, although this is contingent upon high frequencies of fruiting males, which can be altered by environmentally determined gender plasticity.     

INBREEDING EFFECTS IN CLARKIA TEMBLORIENSIS (ONAGRACEAE) POPULATIONS WITH DIFFERENT NATURAL OUTCROSSING RATES

Inbreeding depression is commonly observed in natural populations. The deleterious effects of forced inbreeding are often thought to be less pronounced in populations with self-pollinating mating systems than in primarily outcrossing populations. We tested this hypothesis by comparing the performance of plants produced by artificial self- and cross-pollination from three populations whose outcrossing rate estimates were 0.03, 0.26, and 0.58. Outcrossing rates and inbreeding coefficients were estimated using isozyme polymorphisms as genetic markers. Analysis of F statistics suggests that biparental inbreeding as well as self-fertilization contribute to the level of homozygosity in the seed crop. Biparental inbreeding will reduce the heterozygosity of progeny produced by outcrossing, relative to random outcrossing expectations, and hence will reduce the effects of outcrossing versus self-fertilization. Heterotic selection may increase the average heterozygosity during the life history. Selfed and outcrossed seeds from all three populations were equally likely to germinate and survive to reproduce. However, inbreeding depression was observed in fecundity traits of plants surviving to reproduction in all three populations. Even in the population whose natural self-fertilization rate was 97%, plants grown from seed produced by self-pollination produced fewer fruits and less total seed weight than plants grown from outcrossed seed. There was no detectable inbreeding depression in estimated lifetime fitness. Inbreeding effects for all reproductive yield characters were most severe in the accession from the most outcrossing population and least severe in the accession from the most self-fertilizing population.     

The cost of selfing inEncyclia cochleata (Orchidaceae)

To determine whether inbreeding depression accounts for the maintenance of outcrossing in populations of the self-compatible orchidEncyclia cochleata, the estimated selective advantage of selfing was compared to a measure of inbreeding depression. Individuals from three populations ofE. cochleata and some of their progeny were phenotyped using isozyme analysis. The electrophoretic data were used to estimate the outcrossing rate and the theoretical cost of outcrossing. Inbreeding depression was estimated by comparing the fitness of the progeny resulting from both types of pollinations. The seeds from outcrossed and selfed hand-pollinations and naturally pollinated seeds from a population of the triandrous form ofE. cochleata were grown aseptically on culture media, and their development over the next three years recorded. Inbreeding was common, particularly in one population (outcrossing rate 40%). However, the level of inbreeding depression was only 1–2%, considerably less inbreeding depression than expected.     

Patterns of self compatibility, inbreeding depression, outcrossing, and sex allocation in a marine bryozoan suggest the predominating influence of sperm competition

Sex allocation by simultaneous hermaphrodites is theoretically influenced by selfing rate, which is in turn influenced by the benefits of enhanced genomic transmission and reproductive assurance relative to the cost of inbreeding depression. The experimental investigation of these influences in seed plants has a rich pedigree, yet although such an approach is equally relevant to colonial invertebrates, which globally dominate subtidal communities on firm substrata, such studies have been scarce. We reared self‐compatible genets of the marine bryozoan Celleporella hyalina s.l. in the presence and absence of allosperm, and used molecular genetic markers for paternity analysis of progeny to test theoretical predictions that: (1) genets from focal populations with high selfing rates show less inbreeding depression than from focal populations with low selfing rates; (2) genets whose selfed progeny show inbreeding depression prefer outcross sperm (allosperm); and (3) genets bias sex allocation toward female function when reared in reproductive isolation. Offspring survivorship and paternity analysis were used to estimate levels of inbreeding depression and preference for outcrossing or selfing. Sex allocation was assessed by counting male and female zooids. As predicted, inbreeding depression was severe in selfed progeny of genets derived from the populations with low self‐compatibility rates, but, with one exception, was not detected in selfed progeny of genets derived from the populations with higher self‐compatibility rates. Also, as predicted, genets whose selfed progeny showed inbreeding depression preferred outcrossing, and a genet whose selfed progeny did not show inbreeding depression preferred selfing. Contrary to prediction, sex allocation in the majority of genets was not influenced by reproductive isolation. Lack of economy of male function may reflect the over‐riding influence of allosperm‐competition in typically dense breeding populations offering good opportunity for outcrossing. We suggest that hermaphroditism may be a plesiomorphic character of the crown group Bryozoa, prevented by phylogenetic constraint from being replaced by gonochorism and therefore not necessarily adaptive in all extant clades. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 519–531.     

GENETIC CONSEQUENCES OF OUTCROSSING IN THE CLEISTOGAMOUS ANNUAL,IMPATIENS CAPENSIS. II. OUTCROSSING RATES AND GENOTYPIC CORRELATIONS

The genetic consequences of a plant's mating system depend on both the degree of outcrossing and the genetic relationship between mates. We examined the electrophoretic genotypes of seeds derived from cleistogamous (CL) and chasmogamous (CH) flowers in six populations of the facultatively cleistogamous annual, Impatiens capensis. Multilocus estimates of the outcrossing rates for the strongly protandrous CH flowers ranged from 0.29 to 0.71 and were higher than estimates based on single-locus data. Such results suggest that the CH flowers experience variable levels of both geitonogamous self-fertilization and biparental inbreeding. A new and generally applicable technique based on the relative level of inbreeding within progeny groups provided direct estimates of the correlation between the genotypic values of outcrossed mates. These correlations varied widely among populations and contributed up to half of the inbreeding observed among the CH progeny. Such biparental inbreeding biases estimates of the outcrossing rate based on the mixed-mating model downward and influences mating-system evolution by decreasing the “cost of meiosis.”     

Increased inbreeding but not homozygosity in small populations of Sabatia angularis (Gentianaceae)

Understanding how the mating system varies with population size in plant populations is critical for understanding their genetic and demographic fates. We examined how the mating system, characterized by outcrossing rate, biparental inbreeding rate, and inbreeding coefficient, and genetic diversity varied with population size in natural populations of the biennial Sabatia angularis. We found a significant, positive relationship between outcrossing and population size. Selfing was as high as 40% in one small population but was only 7% in the largest population. Despite this pattern, observed heterozygosity did not vary with population size, and we suggest that selection against inbred individuals maintains observed heterozygosity in small populations. Consistent with this hypothesis, we found a trend of lower inbreeding coefficients in the maternal than progeny generation in all of the populations, and half of the populations exhibited significant excesses of adult heterozygosity. Moreover, genetic diversity was not related to population size and was similar across all populations examined. Our results suggest that the consequences of increased selfing for population fitness in S. angularis, a species that experiences significant inbreeding depression, will depend on the relative magnitude and consistency of inbreeding depression and the demographic cost of selection for outcrossed progeny in small populations.     

Relatively weak inbreeding depression in selfing but also in outcrossing populations of North American Arabidopsis lyrata

Hermaphroditic plants can potentially self‐fertilize, but most possess adaptations that promote outcrossing. However, evolutionary transitions to higher selfing rates are frequent. Selfing comes with a transmission advantage over outcrossing, but self‐progeny may suffer from inbreeding depression, which forms the main barrier to the evolution of higher selfing rates. Here, we assessed inbreeding depression in the North American herb Arabidopsis lyrata, which is normally self‐incompatible, with a low frequency of self‐compatible plants. However, a few populations have become fixed for self‐compatibility and have high selfing rates. Under greenhouse conditions, we estimated mean inbreeding depression per seed (based on cumulative vegetative performance calculated as the product of germination, survival and aboveground biomass) to be 0.34 for six outcrossing populations, and 0.26 for five selfing populations. Exposing plants to drought and inducing defences with jasmonic acid did not magnify these estimates. For outcrossing populations, however, inbreeding depression per seed may underestimate true levels of inbreeding depression, because self‐incompatible plants showed strong reductions in seed set after (enforced) selfing. Inbreeding‐depression estimates incorporating seed set averaged 0.63 for outcrossing populations (compared to 0.30 for selfing populations). However, this is likely an overestimate because exposing plants to 5% CO₂ to circumvent self‐incompatibility to produce selfed seed might leave residual effects of self‐incompatibility that contribute to reduced seed set. Nevertheless, our estimates of inbreeding depression were clearly lower than previous estimates based on the same performance traits in outcrossing European populations of A. lyrata, which may help explain why selfing could evolve in North American A. lyrata.     

Mating system, sex ratio, and persistence of females in the gynodioecious shrub Daphne laureola L. (Thymelaeaceae)

Although in gynodioecious populations male steriles require a fecundity advantage to compensate for their gametic disadvantage, southern Spanish populations of the long-lived shrub Daphne laureola do not show any fecundity advantage over hermaphrodites in terms of seed production and early seedling establishment. By using allozyme markers, we assess the mating system of this species in five populations differing in sex ratio, and infer levels of inbreeding depression over the whole life cycle by comparing the inbreeding coefficients at the seed and adult plant stages. Extremely low outcrossing rates (0.001相似文献   

Effect of inbreeding depression on outcrossing rates among populations of a tropical pine

Inbreeding depression is common among plants and may distort mating system estimates. Mating system studies traditionally ignore this effect, nonetheless an assessment of inbreeding depression that may have occurred before progeny evaluation could be necessary. In the neotropical Pinus chiapensis inbreeding depression was evaluated using regression analysis relating progeny F-values with seed germinability, the mating system was analysed in three populations with contrasting size, using isozymes, obtained a corrected outcrossing rate. Selfing decreased seed viability by 19%, relative to an outcrossed plant. Multilocus outcrossing rates, t(m), varied widely among populations. In the two smallest populations t(m) congruent with 1. Therefore, inbreeding depression did not affect the estimates, but overestimated t(m) by 10% in the third population, which has a true mixed mating system (selfing was the major source of inbreeding), and an unusually low t(m) for pines (t(m) = 0.54, uncorrected, t(m) = 0.49, corrected). Inbreeding depression may be an uneven source of bias for outcrossing estimates even at the infraspecific level. Accuracy [corrected] but not precision [corrected] may be gained by including inbreeding depression in outcrossing estimates. Therefore, caution should be taken when comparing t(m) among species or even populations within the same species.     

Mating patterns and demography in the tristylous daffodil Narcissus triandrus

Mating patterns in plant populations are influenced by interactions between reproductive traits and ecological conditions, both factors that are likely to vary geographically. Narcissus triandrus, a wide-ranging heterostylous herb, exhibits populations with either two (dimorphic) or three (trimorphic) style morphs and displays substantial geographical variation in demographic attributes and floral morphology. Here, we investigate this variation to determine if demography, morphology, and mating system differ between the two sexual systems. Our surveys in Portugal and NW Spain indicated that dimorphic populations were less dense, of smaller size, and had larger plants and flowers compared to trimorphic populations. Outcrossing rates estimated using allozyme markers revealed similar outcrossing rates in dimorphic and trimorphic populations (t(m) dimorphic=0.759; t(m) trimorphic=0.710). All populations experienced significant inbreeding in progeny (mean F=0.143). In contrast, parental estimates of inbreeding were not significantly different from zero (mean F=0.062), implying that few inbred offspring survive to reproductive maturity due to inbreeding depression. Although the majority of inbreeding results from selfing, significant levels of biparental inbreeding were also detected in eight of the nine populations (mean s(s)-s(m)=0.081). Density was negatively associated with levels of selfing but positively associated with biparental inbreeding. Population size was positively associated with outcrossing but not biparental inbreeding. There were no consistent differences among the style morphs in outcrossing or biparental inbreeding indicating that the maintenance of trimorphism vs dimorphism is unlikely to be associated with inbreeding of maternal parents.     

Inbreeding depression in self-incompatible and self-compatible populations of Leavenworthia alabamica

Inbreeding depression is one of the leading factors preventing the evolution of self-fertilization in plants. In populations where self-fertilization evolves, theory suggests that natural selection against partially recessive deleterious alleles will reduce inbreeding depression. The purpose of this study was to evaluate this hypothesis by comparing the magnitude of inbreeding depression in self-incompatible and self-compatible populations of Leavenworthia alabamica. Within-population crosses were conducted to compare the quantity and quality of offspring produced by outcrossing and self-fertilization. These progeny were grown in a common greenhouse and inbreeding depression was measured in germination, survival, biomass, transition rate to flowering, flower number, petal length, pollen grains/anther, pollen viability, and ovule number. In comparison to outcrossing, self-fertilization led to the production of fewer and smaller seeds within self-incompatible populations. Moreover, inbreeding depression was observed in eight of 11 offspring traits within self-incompatible populations of L. alabamica. In contrast, there was significant inbreeding depression only in flower number within self-compatible populations. The results of this study are consistent with the idea that self-fertilization selectively removes partially recessive deleterious alleles causing inbreeding depression in natural plant populations. However, in plant species such as L. alabamica where self-compatibility may evolve in small populations following long-distance dispersal, declines in inbreeding depression may also be facilitated by genetic drift.     

Mating system variation along a successional gradient in the allogamous and colonizing plant Crepis sancta (Asteraceae)

We analysed mating system in an annual and colonizing plant, Crepis sancta, that occupies different successional stages in the French Mediterranean region. Based on a previous experiment, we hypothesized that low inbreeding depression measured in young successional stages should select for selfing whereas higher inbreeding depression in old stages should select for outcrossing. Nine populations of C. sancta (Asteraceae) from contrasting successional stages were used to analyse (1) Seed set after autonomous and enforced selfing in controlled conditions and (2) outcrossing rates in natural conditions using allozymes (progeny array analysis). We found that C. sancta possesses a pseudo‐self‐incompatibility system and that mating system varies among populations. Allozymes revealed that the population multilocus outcrossing rates vary from 0.77 to 0.99. The lowest outcrossing rates occur in the youngest successional stages and complete outcrossing is found in old stages. The data partially agree with the predictions we made and the results are more generally discussed in the light of factors changing during succession. We did not find any evidence of reproductive assurance in the nine populations, contrary to what is often assumed as a major factor governing mating system evolution in colonizing species. We propose that mating system variation can be interpreted as the result of the balance between the cost of outcrossing and inbreeding depression in a metapopulation context.     

Kinship between parents reduces offspring fitness in a natural population of Rhododendron brachycarpum

Background and Aims

A reduction in offspring fitness resulting from mating between neighbours is interpreted as biparental inbreeding depression. However, little is known about the relationship between the parents'' genetic relatedness and biparental inbreeding depression in their progeny in natural populations. This study assesses the effect of kinship between parents on the fitness of their progeny and the extent of spatial genetic structure in a natural population of Rhododendron brachycarpum.

Methods

Kinship coefficients between 11 858 pairs of plants among a natural population of 154 R. brachycarpum plants were estimated a priori using six microsatellite markers. Plants were genotyped, and pairs were selected from among 60 plants to vary the kinship from full-sib to unrelated. After a hand-pollination experiment among the 60 plants, offspring fitness was measured at the stages of seed maturation (i.e. ripening) under natural conditions, and seed germination and seedling survival under greenhouse conditions. In addition, spatial autocorrelation was used to assess the population''s genetic structure.

Key Results

Offspring fitness decreased significantly with increasing kinship between parents. However, the magnitude and timing of this effect differed among the life-cycle stages. Measures of inbreeding depression were 0·891 at seed maturation, 0·122 (but not significant) at seed germination and 0·506 at seedling survival. The local population spatial structure was significant, and the physical distance between parents mediated the level of inbreeding between them.

Conclusions

The level of inbreeding between individuals determines offspring fitness in R. brachycarpum, especially during seed maturation. Genetic relatedness between parents caused inbreeding depression in their progeny. Therefore, biparental inbreeding contributes little to reproduction and instead acts as a selection force that promotes outcrossing, as offspring of more distant (less related) parents survive better.     

Inbreeding depression and mixed mating in Leptosiphon jepsonii: a comparison of three populations

BACKGROUND AND AIMS: Inbreeding depression is thought to play a central role in the evolution and maintenance of cross-fertilization. Theory indicates that inbreeding depression can be purged with self-fertilization, resulting in positive feedback for the selection of selfing. Variation among populations of Leptosiphon jepsonii in the timing and rate of self-fertilization provides an opportunity to study the evolution of inbreeding depression and mating systems. In addition, the hypothesis that differences in inbreeding depression for male and female fitness can stabilize mixed mating in L. jepsonii is tested. METHODS: In a growth room experiment, inbreeding depression was measured in three populations with mean outcrossing rates ranging from 0.06 to 0.69. The performance of selfed and outcrossed progeny is compared at five life history stages. To distinguish between self-incompatibility and early inbreeding depression, aborted seeds and unfertilized ovules were counted in selfed and outcrossed fruits. In one population, pollen and ovule production was quantified to estimate inbreeding depression for male and female fitness. KEY RESULTS: Both prezygotic barriers and inbreeding depression limited self seed set in the most outcrossing population. Cumulative inbreeding depression ranged from 0.297 to 0.501, with the lowest value found in the most selfing population. Significant inbreeding depression for early life stages was found only in the more outcrossing populations. Inbreeding depression was not significant for pollen or ovule production. CONCLUSIONS: The results provide modest support for the hypothesized relationship between inbreeding depression and mating systems. The absence of early inbreeding depression in the more selfing populations is consistent with theory on purging. Differences in male and female expression of inbreeding depression do not appear to stabilize mixed mating in L. jepsonii. The current estimates of inbreeding depression for L. jepsonii differ from those of previous studies, underscoring the effects of environmental variation on its expression.     

Fine-scale genetic structure enhances biparental inbreeding by promoting mating events between more related individuals in wild soybean (Glycine soja; Fabaceae) populations

Outcrossing between genetically distant individuals in a plant population enhances allelic heterozygosity-an important source for genetic diversity and adaptive evolution. Fine-scale spatial genetic structure (FSGS) can interfere with outcrossing by promoting mating between more related individuals. To test the influence of FSGS on outcrossing, FSGS and outcrossing rates were analyzed with four wild soybean (Glycine soja) populations from different habitats, using simple sequence repeat (SSR) fingerprints. Spatial autocorrelation analysis indicated variable FSGS (15.44-25.87 m) in all four populations. Multilocus mixed-mating analysis of 1605 progeny indicated substantial variation in single-locus outcrossing (T(s) = 6.3-12.6%) although the total outcrossing rates as estimated by multilocus outcrossing (T(m) = 12.8-17%) did not vary significantly among populations. The comparison between FSGS and outcrossing rates demonstrated that strong FSGS with large genetic patch size can enhance biparental inbreeding by promoting mating between more related individuals in a population. The results suggest that patch size management can aid in situ conservation by avoiding formation of strong FSGS and encouraging true outcrossing among individuals.     

Inbreeding depression and outbreeding depression in Digitalis purpurea: optimal outcrossing distance in a tetraploid

An optimal crossing distance exists within plant populations if inbreeding and outbreeding depression operate simultaneously. In a population of tetraploid Digitalis purpurea, maternal plants were pollinated with donors at four distances: 0 (self-pollination), 1, 6 and 30 m. Lifetime fitness of F1 progeny was investigated in greenhouse experiments, and significant inbreeding and outbreeding depression were detected at five vs. three life history traits. Inbreeding depression increased at later life stages, whereas outbreeding depression was relatively constant. The existence of within-population outbreeding depression suggests substantial genetic structuring at moderate distances in D. purpurea, and corroborates recent findings of significant outbreeding depression in F1 progeny in polyploids. The moderate inbreeding depression found in this predominately outcrossing population supports the notion that effects of inbreeding are less severe in polyploids than in diploids.     

FLORAL CORRELATES AND FITNESS CONSEQUENCES OF MATING-SYSTEM VARIATION IN TURNERA ULMIFOLIA

Outcrossing rates varied from 0% to 69% among Jamaican populations of Turnera ulmifolia. A correlation between increasing herkogamy and outcrossing rate occurred among populations. Predictions from sex-allocation theory were tested by estimating allocation to reproductive functions. Significant differences in allocation patterns occurred among populations, but they were not correlated with outcrossing rates. The fitness consequences of inbreeding were assessed in high- and low-density greenhouse experiments for nine populations with variable outcrossing rates. No evidence for inbreeding depression occurred in early portions of the life history, but multiplicative fitness functions provide evidence for inbreeding depression. We tested the prediction that selfing populations have lower levels of inbreeding depression than outcrossing populations but found no significant correlation.     

Anther-stigma separation is associated with inbreeding depression in Datura stramonium,a predominantly self-fertilizing annual

Abstract.— Genetically based variation in outcrossing rate generates lineages within populations that differ in their history of inbreeding. According to some models, mating-system modifiers in such populations will demonstrate both linkage and identity disequilibrium with fitness loci, resulting in lineage-specific inbreeding depression. Other models assert that differences among families in levels of inbreeding depression are mainly attributable to random accumulation of genetic load, unrelated to variation at mating-system loci. We measured female reproductive success of selfed and outcrossed progeny from naturally occurring lineages of Datura stramonium , a predominantly self-fertilizing annual weed that has heritable variation in stigma-anther separation, a trait that influences selfing rates. Progeny from inbred lineages (as identified by high degree of anther-stigma overlap) showed equal levels of seed production, regardless of cross type. Progeny from mixed lineages (as identified by relatively high separation between anthers and stigma) showed moderate levels of inbreeding depression. We found a significant correlation between anther-stigma separation and relative fitness of selfed and outcrossed progeny, suggesting that family-level inbreeding depression may be related to differences among lineages in inbreeding history in this population. Negative inbreeding depression in putatively inbred lineages may be due in part to additive effects or to epistatic interactions among loci.     

Pollination biology and outcrossing rates in hermaphroditic Schiedea lydgatei (Caryophyllaceae)

The influence of outcrossing and pollination biology on the maintenance of hermaphroditism was studied for Schiedea lydgatei (Caryophyllaceae: Alsinoideae), a species endemic to Moloka`i in the Hawaiian Islands. Schiedea lydgatei is the only hermaphroditic species in an otherwise dimorphic clade and hermaphroditism is likely the result of a reversal from a gynodioecious ancestor. Both wind and native moths in the family Pyralidae are responsible for pollination in S. lydgatei. Outcrossing rates were generally high (0.80), especially in years when the greatest number of plants were flowering. The combination of high outcrossing rates and substantial inbreeding depression indicates that at present females would not be favored in the population. Pollination by both wind and insects is consistent with the hypothesis that hermaphroditism is the result of a relatively recent reversal, as the ancestor of S. lydgatei was probably wind pollinated and gynodioecious with few females in the populations. A shift from wind to predominately insect pollination on Moloka`i may have resulted in increased outcrossing rates and prevented the expression of high inbreeding depression among progeny of hermaphrodites, a condition that would select against females and favor a reversal to hermaphroditism. Because few females were likely to have been present in ancestral populations that colonized Moloka`i, founder effect is another potential explanation for loss of females. In either case, current high levels of outcrossing prevent re-establishment of females in populations of S. lydgatei.