INBREEDING DEPRESSION IN GYNODIOECIOUS LOBELIA SIPHILITICA: AMONG-FAMILY DIFFERENCES OVERRIDE BETWEEN-MORPH DIFFERENCES

If inbreeding depression is caused by deleterious recessive alleles, as suggested by the partial dominance hypothesis, a negative correlation between inbreeding and inbreeding depression is predicted. This hypothesis has been tested several times by comparisons of closely related species or comparisons of populations of the same species with different histories of inbreeding. However, if one is interested in whether this relationship contributes to mating-system evolution, which occurs within populations, comparisons among families within a population are needed; that is, inbreeding depression among individuals with genetically based differences in their rate of selfing should be compared. In gynodioecious species with self-compatible hermaphrodites, hermaphrodites will have a greater history of potential inbreeding via both selfing and biparental inbreeding as compared to females and may therefore express a lower level of inbreeding depression. We estimated the inbreeding depression of female and hermaphrodite lineages in gynodioecious Lobelia siphilitica in a greenhouse experiment by comparing the performance of selfed and outcrossed progeny, as well as sibling crosses and crosses among subpopulations. We did not find support for lower inbreeding depression in hermaphrodite lineages. Multiplicative inbreeding depression (based on seed germination, juvenile survival, survival to flowering, and flower production in the first growing season) was not significantly different between hermaphrodite lineages (δ = 0.30 ± 0.08) and female lineages (δ = 0.15 ± 0.18), although the trend was for higher inbreeding depression in the hermaphrodite lineages. The population-level estimate of inbreeding depression was relatively low for a gynodioecious species (δ = 0.25) and there was no significant inbreeding depression following biparental inbreeding (δ = 0.01). All measured traits showed significant variation among families, and there was a significant interaction between family and pollination treatment for four traits (germination date, date of first flowering, number of flowers, and aboveground biomass). Our results suggest that the families responded differently to selfing and outcrossing: Some families exhibited lower fitness following selfing whereas others seemed to benefit from selfing as compared to outcrossing. Our results support recent simulation results in that prior inbreeding of the lineages did not determine the level of inbreeding depression. These results also emphasize the importance of determining family-level estimates of inbreeding depression, relative to population-level estimates, for studies of mating-system evolution.     

EFFECTS OF DIFFERENT LEVELS OF INBREEDING ON PROGENY FITNESS IN PLANTAGO CORONOPUS

Inbreeding depression (δ) is a major selective force favoring outcrossing in flowering plants. Many phenotypic and genetic models of the evolution of selfing conclude that complete outcrossing should evolve whenever inbreeding depression is greater than one-half, otherwise selfing should evolve. Recent theoretical work, however, has challenged this view and emphasized (1) the importance of variation in inbreeding depression among individuals within a population; and (2) the nature of gene action between deleterious mutations at different loci (epistasis) as important determinants for the evolution of plant mating systems. The focus of this study was to examine the maintenance of inbreeding depression and the relationship between inbreeding level and inbreeding depression at both the population and the individual level in one population of the partially self-fertilizing plant Plantago coronopus (L.). Maternal plants, randomly selected from an area of about 50 m² in a natural population, were used to establish lines with expected inbreeding coefficients (f) of 0, 0.25, 0.50, 0.75, and 0.875. Inbreeding depression was estimated both in the greenhouse and at the site of origin of the maternal plants by comparing growth, survival, flowering, and seed production of the progeny with different inbreeding coefficients. No significant inbreeding depression for these fitness traits was detected in the greenhouse after 16 weeks. This was in strong contrast to the field, where the traits all displayed significant inbreeding depression and declined with increased inbreeding. The results were consistent with the view that mutation to mildly deleterious alleles is the primary cause of inbreeding depression. At the family level, significantly different maternal line responses (maternal parent × inbreeding level interaction) provide a mechanism for the invasion of a selfing variant into the population through any maternal line exhibiting purging of its genetic load. At the population level, evidence for synergistic epistasis was detected for the probability of flowering, but not for total seed production. At the family level, however, a significant interaction between inbreeding level and maternal families for both traits was observed, indicating that epistasis could play a role in the expression of inbreeding depression among maternal lines.     

No correlation between inbreeding depression and delayed selfing in the freshwater snail Physa acuta

Inbreeding depression, one of the main factors driving mating system evolution, can itself evolve as a function of the mating system (the genetic purging hypothesis). Classical models of coevolution between mating system and inbreeding depression predict negative associations between inbreeding depression and selfing rate, but more recent approaches suggest that negative correlations should usually be too weak or transient to be detected within populations. Empirical results remain unclear and restricted to plants. Here, we evaluate, for the first time, the within-population genetic correlation between inbreeding depression and a trait that controls the amount of self-fertilization (the waiting time) in a self-fertile hermaphroditic animal, the freshwater snail Physa acuta. Using a large quantitative-genetic design (36 grand-families and 348 families), we observe abundant within-population family-level genetic variation for both inbreeding depression (estimated for survival, fecundity, and size) and the degree of behavioral selfing avoidance. However, we detected no correlation between waiting time and inbreeding depression across families. In agreement with recent models, this result shows that mutational variance rather than differential purging accounts for most of the genetic variance in inbreeding depression within a population.     

Fitness Traits and Dispersal Ability in the Herb Tragopogon pratensis (Asteraceae): Decoupling the Role of Inbreeding Depression and Maternal Effects

Abstract: Inbreeding depression can decrease several fitness traits and maternal effects can strongly influence the amount of inbreeding depression. Understanding the effects of inbreeding depression on plant fitness is especially important in the context of habitat fragmentation, where plant populations become smaller and more isolated, exhibiting increasing levels of inbreeding depression. We examined the joint influence of inbreeding depression and maternal effects on life cycle traits and dispersal ability in the herb Tragopogon pratensis that grows in fragmented populations in Europe. We conducted experimental crosses to obtain selfed and outcrossed progeny in two contrasted environments. In particular, we produced a first generation of seeds and plants that were self-pollinated again to produce a second generation of seeds. Individual seeds were weighed and their pappuses measured to estimate the dispersal potential. Pollination treatment only had a significant effect on seed mass and dispersal ability. Coefficients of inbreeding depression did not differ between selfed and outcrossed plants. Seed mass had a significant effect on germination date. Environment had a significant effect on mass of the second generation of seeds and the interaction between pollination treatment and family was significant for six traits, indicating the existence of strong maternal effects in T. pratensis. Results suggest population differentiation. Overall, T. pratensis populations exhibited a good performance under selfing, in terms of life cycle traits and dispersal ability, which would allow the species to cope with problems associated with fragmentation.     

Problems in measuring among-family variation in inbreeding depression

Understanding the sources of variation in inbreeding depression within populations is important for understanding the evolution of selfing rates. At the population level, inbreeding depression is due to decreased heterozygosity caused by inbreeding, which decreases overdominance and increases the frequency of expression of recessive deleterious alleles. However, within individual families inbreeding has two distinct consequences: it reduces heterozygosity and it restricts the alleles present in offspring to those present in the parent. Outcrossing both increases heterozygosity and brings new alleles into a family (compared to the alleles present if the plant is self-pollinated). Both consequences of inbreeding affect offspring fitness, but the most common experimental design used to measure among-family variation in inbreeding depression cannot distinguish them. The result is that variance in inbreeding depression among families is confounded by genetic variation in the traits being measured. Also, correlations (among families) between measures of inbreeding depression or between inbreeding depression and mean trait values are confounded by genetic variation in the traits being measured. I conclude that more complex crossing designs that allow estimation of breeding values for individual families are required to accurately detect and measure among-family variation in inbreeding depression.     

Within and between population variation in inbreeding depression in the locally threatened perennial Scabiosa columbaria

Inbreeding depression plays a central role within the conservation genetics paradigm. Until now inbreeding depression is incorporated into models of population viability as a mean value (e.g. number of lethal equivalents) for all traits in a population. In this study of the locally threatened perennial plant species Scabiosa columbaria we investigated both the mean and the variance among families of inbreeding depression in eight life history traits for five natural populations varying in size from 300 to more than 120,000 individuals. Significant inbreeding depression was found in all populations and all traits. The mean inbreeding depression value per trait was never correlated to population size. Within each population, highly significant variation in inbreeding depression between families (VIFLID) was found. Per trait, families with inbreeding depression next to families with outbreeding depression were often found within the same population. Inbreeding depression at the family level was in many cases not correlated among traits and independent of correlations between traits themselves. VIFLID was negatively correlated with population size: in two traits these correlations were significant. The results underline that inbreeding depression is a complex, highly dynamic phenomenon. Models of viability should incorporate inbreeding depression distributions, with a trait specific mean and variance. Moreover, models of metapopulation dynamics should incorporate genotype quality as factor in colonization success.     

Spawning, copulation and inbreeding coefficients in marine invertebrates

Patterns of population genetic variation have frequently been understood as consequences of life history covariates such as dispersal ability and breeding systems (e.g. selfing). For example, marine invertebrates show enormous variation in life history traits that are correlated with the extent of gene flow between populations and the magnitude of differentiation among populations at neutral genetic markers (FST). Here we document an unexpected correlation between marine invertebrate life histories and deviation from Hardy-Weinberg equilibrium (non-zero values of FIS, the inbreeding coefficient). FIS values were significantly higher in studies of species with free-spawned planktonic sperm than in studies of species that copulate or have some form of direct sperm transfer to females or benthic egg masses. This result was robust to several different analytical approaches. We note several mechanisms that might contribute to this pattern, and appeal for more studies and ideas that might help to explain our observations.     

Maintenance of Inbreeding Depression in a Highly Self-Fertilizing Tree, Magnolia obovata Thunb

Inbreeding depression is a major selective force that maintains outcrossing in flowering plants. If the long life and large mature size of trees cause high inbreeding depression via mitotic mutations and half-sib competition, these characteristics may increase inbreeding depression sufficiently to maintain traits that facilitate outcrossing even with high primary selfing rates (proportion of selfed ovules). Here, I report the maintenance of inbreeding depression in a population of a tree (Magnolia obovata Thunb.) with primary selfing rates greater than 0.8 resulting from geitonogamy. The progenies exhibited inbreeding depression for germination, seedling survival, and seedling mass (δ = 0.29–0.38), but no significant difference between crossing type in seedling height. Cumulative inbreeding depression for early survival (from zygote to 2-year-old stage) estimated from these results and from prior data on embryonic survival was high (δ_e = 0.91). The fixation index at maturity based on six allozyme loci was low (F_is = 0.08), indicating that significant inbreeding depression for late survival results in a low level of inbreeding with respect to gene transmission to the next generation. From these results, I estimated that inbreeding depression for late and lifetime survival equaled 0.69 and 0.97, respectively. These results suggest that M. obovata trees maintain high inbreeding depression at both early and late life stages, resulting in a low level of inbreeding despite a high primary selfing rate. The high inbreeding depression can be explained by previous theories and is consistent with the predicted maintenance of inbreeding depression in highly self-fertilizing tree populations. The inbreeding load due to the high primary selfing rate represents a cost of this tree’s pollination system for outcrossing, which is based on automimicry and mass flowering. Co-ordinating editor: S.-M. Chang     

Evidence for an epigenetic role in inbreeding depression

Inbreeding depression (i.e. negative fitness effects of inbreeding) is central in evolutionary biology, affecting numerous aspects of population dynamics and demography, such as the evolution of mating systems, dispersal behaviour and the genetics of quantitative traits. Inbreeding depression is commonly observed in animals and plants. Here, we demonstrate that, in addition to genetic processes, epigenetic processes may play an important role in causing inbreeding effects. We compared epigenetic markers of outbred and inbred offspring of the perennial plant Scabiosa columbaria and found that inbreeding increases DNA methylation. Moreover, we found that inbreeding depression disappears when epigenetic variation is modified by treatment with a demethylation agent, linking inbreeding depression firmly to epigenetic variation. Our results suggest an as yet unknown mechanism for inbreeding effects and demonstrate the importance of evaluating the role of epigenetic processes in inbreeding depression.     

Individual Variation in Inbreeding Depression: The Roles of Inbreeding History and Mutation

We use mutation-selection recursion models to evaluate the relative contributions of mutation and inbreeding history to variation among individuals in inbreeding depression and the ability of experiments to detect associations between individual inbreeding depression and mating system genotypes within populations. Poisson mutation to deleterious additive or recessive alleles generally produces far more variation among individuals in inbreeding depression than variation in history of inbreeding, regardless of selfing rate. Moreover, variation in inbreeding depression can be higher in a completely outcrossing or selfing population than in a mixed-mating population. In an initially random mating population, the spread of a dominant selfing modifier with no pleiotropic effects on male outcross success causes a measurable increase in inbreeding depression variation if its selfing rate is large and inbreeding depression is caused by recessive lethals. This increase is observable during a short period as the modifier spreads rapidly to fixation. If the modifier alters selfing rate only slightly, it fails to spread or causes no measurable increase in inbreeding depression variance. These results suggest that genetic associations between mating loci and inbreeding depression loci could be difficult to demonstrate within populations and observable only transiently during rapid evolution to a substantially new selfing rate.     

The inbreeding depression cost of selfing: Importance of flower size and population size in Collinsia parviflora (Veronicaceae)

Inbreeding depression should evolve with selfing rate when frequent inbreeding results in exposure of and selection against deleterious alleles. The selfing rate may be modified by plant traits such as flower size, or by population characteristics such as census size that can affect the probability of biparental inbreeding. Here we quantify inbreeding depression (δ) among different population sizes of Collinsia parviflora, a wildflower with interpopulation variation in flower size, by comparing fitness components and multiplicative fitness of experimentally produced selfed and outcrossed offspring. Selfed offspring had reduced multiplicative fitness compared to outcrossed offspring, but inbreeding depression was low in all combinations of population size and flower size (δ ≤ 0.05) except in large populations of large-flowered plants (δ = 0.45). The decrement to multiplicative fitness with inbreeding was not affected by population size nested within flower size, but differed between small- and large-flowered plants: small-flowered populations had lower overall inbreeding depression (δ = 0.04) compared to large-flowered populations (δ = 0.25). The difference in load with flower size suggests that either selection has removed deleterious recessive alleles or these alleles have become fixed in small-flowered, potentially more selfing populations, but that purging has not occurred to the same extent in presumably outcrossing large-flowered populations.     

Variability of individual genetic load: consequences for the detection of inbreeding depression

Inbreeding depression is a key factor affecting the persistence of natural populations, particularly when they are fragmented. In species with mixed mating systems, inbreeding depression can be estimated at the population level by regressing the average progeny fitness by the selfing rate of their mothers. We applied this method using simulated populations to investigate how population genetic parameters can affect the detection power of inbreeding depression. We simulated individual selfing rates and genetic loads from which we computed fitness values. The regression method yielded high statistical power, inbreeding depression being detected as significant (5?% level) in 92?% of the simulations. High individual variation in selfing rate and high mean genetic load led to better detection of inbreeding depression while high among-individual variation in genetic load made it more difficult to detect inbreeding depression. For a constant sampling effort, increasing the number of progenies while decreasing the number of individuals per progeny enhanced the detection power of inbreeding depression. We discuss the implication of among-mother variability of genetic load and selfing rate on inbreeding depression studies.     

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.     

EVOLUTION OF GYNODIOECY AND MAINTENANCE OF FEMALES: THE ROLE OF INBREEDING DEPRESSION,OUTCROSSING RATES,AND RESOURCE ALLOCATION IN SCHIEDEA ADAMANTIS (CARYOPHYLLACEAE)

Levels of inbreeding depression, outcrossing rates, and phenotypic patterns of resource allocation were studied to examine their relative importance in the maintenance of high numbers of females in gynodioecious Schiedea adamantis (Caryophyllaceae), an endemic Hawaiian shrub found in a single population on Diamond Head Crater, Oahu. In studies of inbreeding depression in two greenhouse environments, families of hermaphrodites exhibited significant inbreeding depression (δ = 0.60), based on a multiplicative fitness function using seeds per capsule, germination, survival, and the inflorescence biomass of progeny. Differences between inbred and outcrossed progeny were smallest at the early stage of seeds per capsule and more pronounced at the later stages of survival and inflorescence production. These results are consistent with inbreeding depression caused by many mutations of small effect. Using allozyme analyses, the inbreeding coefficient of adult plants in the field was not significantly different from zero, implying that δ in nature may be equal to one. The single locus estimate of the outcrossing rate for hermaphrodites was 0.50 based on progeny that survived to flowering; corrected for the disproportionate loss before flowering of progeny from selfing, the adjusted outcrossing rate at the zygote stage was 0.32, suggesting that considerable selfing occurs in hermaphrodites. Females were totally outcrossed. When females and hermaphrodites were compared for reproductive output in the field, females produced over twice as many seeds per plant as hermaphrodites, primarily because females had far more capsules per inflorescence than hermaphrodites. Females had greater mass per seed than hermaphrodites in the field, either because of greater provisioning or reduced inbreeding depression. There was no significant differential mortality with respect to sex over a seven year period. The higher number of seeds per plant of females, combined with substantial inbreeding depression and relatively high selfing rates for hermaphrodites, are probably responsible for the maintenance of females in this population. The predicted frequency of females based on data for seed production, the adjusted selfing rate, and inbreeding depression is 42%, remarkably close to the observed frequency of 39%. High levels of inbreeding depression suggest that considerable quantitative genetic variation is present for traits affecting fitness in this population, despite low allozyme variability and a presumed founder effect.     

Inbreeding depression in perennial Lychnis viscaria (Caryophyllaceae): effects of population mating history and nutrient availability

We studied inbreeding depression in a perennial plant, Lychnis viscaria, in three populations differing in their inbreeding history and population size by measuring several traits at two nutrient levels over the plant's life cycle. The observed levels of inbreeding depression (cumulative inbreeding depression, from -0.057 to 0.629) were high for a plant with a mixed mating system. As expected, the population with a low level of isozyme variation expressed the least inbreeding depression for seed germination. Highest inbreeding depression for germination was found in the largest and genetically most variable population. No clear differences between populations in expression of inbreeding depression in the later life stages were found. The population level inbreeding depression varied with the nutrient conditions and among populations and life stages, but we found no evidence that inbreeding depression increased with lower nutrient availability. These results emphasize the importance of measuring inbreeding depression under several environmental conditions and over life stages.     

Patterns of inbreeding depression in a population of Brassica cretica (Brassicaceae): evidence from family-level analyses

In this investigation, we have collected family-structured data from a partly self-compatible, outcrossing population of Brassica cretica to estimate and compare the effects of one-generation selfing on different types of characters. Inbreeding not only depressed characters that should be positively correlated with fitness irrespective of habitat, e.g. germinability, leaf number and inflorescence size, but also resulted in later flowering, smaller and more asymmetric flowers, and an increased production of basal branches. Population-level estimates of inbreeding depression were similar in magnitude to estimates reported in other wild plant species. There was a tendency for direct components of fitness to exhibit a stronger response to inbreeding than other characters, but only when the differences between selfed and outbred offspring were measured in standard deviation units. Family-level estimates of inbreeding depression were weakly correlated across characters. Given these and other observations, we hypothesize that the genetic basis of inbreeding depression varies across the life cycle and that changes in local inbreeding will lead to shifts in the mean phenotypes of B. cretica populations. However, judging from data on current levels of population divergence, quite large changes in inbreeding will be required to influence large-scale patterns of variation in this species.  © 2002 The Linnean Society of London. Biological Journal of the Linnean Society , 2002, 76 , 317–325.     

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.     

THE RELATIONSHIP BETWEEN MATING-SYSTEM CHARACTERS AND INBREEDING DEPRESSION IN MIMULUS GUTTATUS

Several recent theoretical considerations of mating-system evolution predict within-population covariation between levels of inbreeding depression and genetically controlled mating-system characters. If inbreeding depression is caused by deleterious recessive alleles, families with characters that promote self-pollination should show lower levels of inbreeding depression than families with characters that promote outcrossing. The converse is expected if inbreeding depression is due to overdominant allelic interactions. Whether these associations between mating-system and viability loci evolve will have important consequences for mating-system evolution. The evolution of selfing within the genus Mimulus is associated with a reduction in stigma-anther separation (i.e., a loss of herkogamy) and high autogamous seed set. In this study we compared families from two M. guttatus populations that differed genetically in their degree of stigma-anther separation. In one of these populations we also compared families that differed genetically in the degree to which they autogamously set seed in a pollinator-free greenhouse. Dams often differed significantly in levels of inbreeding depression for aboveground biomass and flower production, but variation in inbreeding depression was never explained by herkogamy class or autogamy class. Several factors might account for why families with traits associated with selfing did not show lower inbreeding depression, and these are discussed. Our study also demonstrated significant variation among self progeny from a given female likely due to differences in pollination date and position of fruit maturation. The detection of significant dam × sire interactions suggests biparental inbreeding or differences in combining ability for specific pairs of parents.     

Purging of inbreeding depression within a population of Oxalis alpina (Oxalidaceae)

? Premise of the study: Variation among individuals in levels of inbreeding depression associated with selfing levels could influence mating system evolution by purging deleterious alleles, but empirical evidence for this association is limited. ? Methods: We investigated the association of family-level inbreeding depression and presumed inbreeding history in a tristylous population of Oxalis alpina (Oxalidaceae). ? Key results: Mid-styled individuals possessed the greatest degree of self-compatibility (SC) and produced more autogamous capsules than short- or long-styled individuals. Offspring of highly self-compatible mid-styled individuals showed reduced inbreeding depression. Mid-styled plants that produced capsules autogamously exhibited reduced stigma-anther separation compared to mid-styled plants that produced no capsules autogamously. Reduced inbreeding depression was not correlated with stigma-anther separation, suggesting that self-compatibility and autogamy evolve before morphological changes in stigma-anther separation. ? Conclusions: Purging of inbreeding depression occurred in SC mid-styled maternal families. Low inbreeding depression in SC mid-styled plants may lead to retention of the mid-styled morph in populations, despite the occurrence of higher selfing rates in mid-styled relative to short- or long-styled morphs. Variation among individuals in levels of self-fertilization within populations may lead to associations between inbreeding lineages and lower levels of inbreeding depression, influencing the evolution of mating systems.     

Pollination ecology and inbreeding depression control individual flowering phenologies and mixed mating

We analyze evolution of individual flowering phenologies by combining an ecological model of pollinator behavior with a genetic model of inbreeding depression for plant viability. The flowering phenology of a plant genotype determines its expected daily floral display which, together with pollinator behavior, governs the population rate of geitonogamous selfing (fertilization among flowers on the same plant). Pollinators select plant phenologies in two ways: they are more likely to visit plants displaying more flowers per day, and they influence geitonogamous selfing and consequent inbreeding depression via their abundance, foraging behavior, and pollen carry‐over among flowers on a plant. Our model predicts two types of equilibria at stable intermediate selfing rates for a wide range of pollinator behaviors and pollen transfer parameters. Edge equilibria occur at maximal or minimal selfing rates and are constrained by pollinators. Internal equilibria occur between edge equilibria and are determined by a trade‐off between pollinator attraction to large floral displays and avoidance of inbreeding depression due to selfing. We conclude that unavoidable geitonogamous selfing generated by pollinator behavior can contribute to the common occurrence of stable mixed mating in plants.