Developmental stability in Brassica cretica: the effect of crossing distance on fluctuating asymmetry in cotyledon morphology

In the present investigation of Brassica cretica, a wild relative of the cultivated cabbage, B. oleracea, we performed an extensive crossing experiment, involving self-pollinations, random outcrosses within populations and hybridizations between populations or species, to evaluate the relationship between crossing distance and developmental stability, estimated as the absolute difference between the right and left lobe of the cotyledons. The frequency distribution of the right-minus-left scores had a narrower peak than expected for normally-distributed data, but there was no directional asymmetry or antisymmetry. Despite evidence for inbreeding depression in seedling biomass and cotyledon size, the type of cross had negligible influence on cotyledon asymmetry. Separate analyses of between-population hybrids revealed differences among progenies from different pairs of populations and a tendency for the F1 hybrid means to decrease with the geographic distance separating the parent populations, but only for the two size variables. Based on these and other observations, we propose that the degree of cotyledon asymmetry is unrelated to genome-wide characteristics, such as the level of heterozygosity and genomic co-adaptation, and that cotyledon asymmetry is unrelated to the level of genetic stress experienced by each individual. Hence, there is no reason to consider measures of asymmetry as more sensitive indicators of genetic health than conventional fitness variables.     

THE SIGNIFICANCE OF GENETIC EROSION IN THE PROCESS OF EXTINCTION. IV. INBREEDING DEPRESSION AND HETEROSIS EFFECTS CAUSED BY SELFING AND OUTCROSSING IN SCABIOSA COLUMBARIA

The effects of self-fertilization, within-population crosses (WPC) and between-population crosses (BPC) on progeny fitness were investigated in the greenhouse for Scabiosa columbaria populations of varying size. Plants grown from field collected seeds were hand pollinated to produce selfed, WPC, and BPC progeny. The performance of these progenies was examined throughout the entire life cycle. The different pollination treatments did not significantly affect germination, seedling-to-adult survival, flowering percentage and the number of flower heads. But severe inbreeding depression was demonstrated for biomass production, root development, adult survival, and seed set. Additionally, multiplicative fitness functions were calculated to compare relative fitnesses for progeny. On average, WPC progeny showed a more than 4-fold, and BPC progeny an almost 10-fold, advantage over selfed progeny, indicating that S. columbaria is highly susceptible to inbreeding. No clear relationship was found between population size and level of inbreeding depression, suggesting that the genetic load has not yet been reduced substantially in the small populations. A significant positive correlation was found between plant dry weight and total fitness. In two out of six populations, the differences between the effects of the pollination treatments on dry weight increased significantly when seedlings were grown under competitive conditions. This result is interpreted as an enhancement of inbreeding depression under these conditions. It is argued that improvement of the genetic exchange between populations may lower the probability of population extinction.     

Offspring fitness in relation to population size and genetic variation in the rare perennial plant species Gentiana pneumonanthe (Gentianaceae)

Seeds were sampled from 19 populations of the rare Gentiana pneumonanthe, ranging in size from 5 to more than 50,000 flowering plants. An analysis was made of variation in a number of life-history characters in relation to population size and offspring heterozygosity (based on seven polymorphic isozyme loci). Life-his-tory characters included seed weight, germination rate, proportion of seeds germinating, seedling mortality, seedling weight, adult weight, flower production per plant and proportion of plants flowering per family. Principal component analysis (PCA) reduced the dataset to three main fitness components. The first component was highly correlated with adult weight and flowering performance, the second with germination performance and the third component with seed and seedling weight and seedling mortality. The latter two components were considered as being maternally influenced, since these comprised life-history traits that were significantly correlated with seed weight. Multiple regression analysis showed that variation in the first fitness component was mainly associated with heterozygosity and not with population size, while the third fitness component was only correlated with population size and not with heterozygosity. The latter relationship appeared to be non-linear, which suggests a stronger loss of fitness in the smallest populations. The second (germination) component was neither correlated with population size nor with genetic variation. There was only a weak association between population size, heterozygosity and the population coefficients of variation for each life history character. Most correlation coefficients were negative, however, which suggests that there is more variation among progeny from smaller populations. We conclude that progeny from small populations of Gentiana pneumonanthe show reduced fitness and may be phenotypically more variable. One of the possible causes of the loss of fitness is a combination of unfavourable environmental circumstances for maternal plants in small populations and increased inbreeding. The higher phenotypic variation in small populations may also be a result of inbreeding, which can lead to deviation of individuals from the average phenotype through a loss of developmental stability.     

THE SIGNIFICANCE OF GENETIC EROSION IN THE PROCESS OF EXTINCTION. IV. INBREEDING LOAD AND HETEROSIS IN RELATION TO POPULATION SIZE IN THE MINT SALVIA PRATENSIS

If, because of genetic erosion, the level of homozygosity in small populations is high, additional selfing will result in small reductions of fitness. In addition, in small populations with a long inbreeding history selection may have purged the population of its genetic load. Therefore, a positive relationship between population size (or level of genetic variation) and level of additional inbreeding depression, here referred to as inbreeding load, may be expected. In a previous study on the rare and threatened perennial Salvia pratensis, a positive correlation between population size and level of allozyme variation has been demonstrated. In the present study, the inbreeding load in six populations of varying size and allozyme variation was investigated. In the greenhouse, significant inbreeding load in mean seed weight, proportion of germination, plant size, regenerative capacity, and survival was demonstrated. In a field experiment with the two largest and the two smallest populations, survival of selfed progeny was 16% to 63% lower than survival of outcrossed progeny. In addition, survival of outcrossed progeny was, with the exception of the largest population, lower (16% to 37%) than of hybrid progeny, resulting from crosses between populations. Effects on plant size were qualitatively similar to the effects on survival, but these effects were variable in time because of differential survival of larger individuals. In all populations the total inbreeding load, that is, the effects on size and survival multiplicated, increased in time. It was demonstrated that inbreeding load in different characters may be independent. At no time and for no character was inbreeding load or the heterosis effect correlated to the mean number of alleles per locus, indicating that allozyme variation is not representative for variation at fitness loci in these populations. Combined with results of previous investigations, these results suggest that the small populations are in an early phase of the genetic erosion process. In this phase, allozyme variation, which is supposed to be (nearly) neutral, has been affected by genetic erosion but the selectively nonneutral variation is only slightly affected. These results stress the need for detailed information about the inbreeding history of small populations. The relative performance of selfed progeny was lowest in all populations, in the greenhouse as well as in the field, and inbreeding depression could still influence the extinction probabilities of the small populations.     

Inbreeding depression and low between-population heterosis in recently diverged experimental populations of a selfing species

In fragmented populations, genetic drift and selection reduce genetic diversity, which in turn results in a loss of fitness or in a loss of evolvability. Genetic rescue, that is, controlled input of diversity from distant populations, may restore evolutionary potential, whereas outbreeding depression might counteract the positive effect of this strategy. We carried out self-pollination and crosses within and between populations in an experimental subdivided population of a selfing species, Triticum aestivum L., to estimate the magnitude of these two phenomena. Surprisingly, for a self-fertilizing species, we found significant inbreeding depression within each population for four of the six traits studied, indicating that mildly deleterious mutations were still segregating in these populations. The progeny of within- and between-population crosses was very similar, indicating low between-population heterosis and little outbreeding depression. We conclude that relatively large population effective sizes prevented fixation of a high genetic load and that local adaptation was limited in these recently diverged populations. The kinship coefficient estimated between the parents using 20 neutral markers was a poor predictor of the progeny phenotypic values, indicating that there was a weak link between neutral diversity and genes controlling fitness-related traits. These results show that when assessing the viability of natural populations and the need for genetic rescue, the use of neutral markers should be complemented with information about the presence of local adaptation in the subdivided population.     

Effects of crossing distance on offspring fitness and developmental stability in Dalechampia scandens (Euphorbiaceae)

Crosses between genetically close and distant populations of Dalechampia scandens (Euphorbiaceae) were made to test whether the responses of various fitness components and measurements of developmental stability were affected by the outcrossing distance (level of outbreeding). Two fecundity traits, seed set and seed mass, decreased consistently with increasing level of outbreeding, and hybrids between the most divergent populations were sterile. Effects of the genetic distance between parental populations on viability traits, survival and vigor at 1 month of age, were highly idiosyncratic. Hybrids of one long-distance combination had no reduction in survival and vigor, while both traits were greatly reduced in the other long-distance combination. The expression of outbreeding depression on fecundity traits differed between reciprocal crosses in some hybrids but not others; thus, hybrid breakdown may have been due to cytoplasmic-by-nuclear gene interactions, reduced endosperm formation, or an interaction between progeny and maternal genotype. None of the measures of developmental stability had a consistent relationship with either genetic distance between parental populations or seedling vigor. These results suggest that fecundity and viability traits may be differentially affected by hybridization, probably due to differences in genetic architecture among populations. This study also confirms that developmental stability, as measured by the level of fluctuating asymmetry, is not a reliable index of genetic stress.     

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.     

Developmental instability in Brassica campestris (Cruciferae): fluctuating asymmetry of foliar and floral traits

The degree of fluctuating asymmetry of bilateral traits provides a measure of developmental instability, which can be influenced by genetic as well as environmental stress. We studied genetic variation between and within two populations of the mustard Brassica campestris for asymmetry of foliar (cotyledon width) and floral (petal length and width) traits as well as for phenological (germination and flowering) and performance (biomass and flowering) traits. The two populations differed in mean expression of most traits, including asymmetry. However, within-population estimates of genetic variability tended to be lower for asymmetry than other traits. Asymmetry was greater in the population that had lower biomass accumulation and flower production, which supports the idea that population-level asymmetry may be indicative of population-level performance. However, within each population, evidence that performance was negatively correlated with asymmetry was equivocal. Within populations there was little or no concordance among estimates of asymmetry based on different structures, i.e., plants that had highly asymmetrical cotyledons did not tend to have highly asymmetrical petals. The lack of a general buffering capacity at the individual level may be explained by developmental processes (e.g., action of different genes or morphogens) as well as evolutionary processes (e.g., selection on asymmetry of different traits).     

THE RELATIONSHIP BETWEEN INBREEDING DEPRESSION AND PRIOR INBREEDING AMONG POPULATIONS OF FOUR MIMULUS TAXA

When recessive mutations are the primary cause of inbreeding depression, a negative relationship between the levels of prior inbreeding and inbreeding depression is expected. We tested this prediction using 15 populations chosen a priori to represent a wide range of prior inbreeding among four closely related taxa of the Mimulus guttatus species complex. Artificially selfed and outcrossed progeny were grown under controlled growth-chamber conditions, and inbreeding depression was estimated for each population as one minus the ratio of the fitness of selfed to outcrossed progeny. Estimates of inbreeding depression varied from 0% to 68% among populations. Inbreeding coefficients, estimated from electrophoretic assay of field-collected progenies, ranged from 0.02 to 0.76. All five fitness traits displayed a negative association between inbreeding depression and the inbreeding coefficient, but only height showed a statistically significant correlation. Inbreeding depression was also not correlated with the level of genetic variability. In addition, populations with similar levels of prior inbreeding showed significant differences of inbreeding depression, whereas populations with different levels of prior inbreeding showed similar inbreeding depression. Within populations, inbreeding depression did not differ between progeny selfed one versus two generations. Our results are weakly consistent with the recessive mutation model of inbreeding depression, but suggest that additional factors, including genotype-by-environment interaction and complex modes of inheritance, may influence the expression of inbreeding depression.     

BIPARENTAL INBREEDING AND INTERREMNANT MATING IN A PERENNIAL PRAIRIE PLANT: FITNESS CONSEQUENCES FOR PROGENY IN THEIR FIRST EIGHT YEARS

Despite fundamental importance to population dynamics, mating system evolution, and conservation management, the fitness consequences of breeding patterns in natural settings are rarely directly and rigorously evaluated. We experimentally crossed Echinacea angustifolia, a widespread, perennial prairie plant undergoing radical changes in distribution and abundance due to habitat fragmentation. We quantified the effects of both biparental inbreeding and crossing between remnant populations on progeny survival and reproduction in the field over the first eight years. Lifetime fitness is notoriously difficult to assess particularly for iteroparous species because of the long sequence and episodic nature of selection events. Even with fitness data in hand, analysis is typically plagued by nonnormal distributions of overall fitness that violate the assumptions of the usual parametric statistical approaches. We applied aster modeling, which integrates the measurements of separate, sequential, nonnormally distributed annual fitness components, and estimated current biparental inbreeding depression at 68% in progeny of sibling‐mating. The effect of between‐remnant crossing on fitness was negligible. Given that relatedness among individuals in remnant populations is already high and dispersal very limited, inbreeding depression may profoundly affect future dynamics and persistence of these populations, as well as their genetic composition.     

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.     

Fluctuating asymmetry as an indicator of fitness: can we bridge the gap between studies?

There is growing evidence from both experimental and non-experimental studies that fluctuating asymmetry does not consistently index stress or fitness. The widely held--yet poorly substantiated--belief that fluctuating asymmetry can act as a universal measure of developmental stability and predictor of stress-mediated changes in fitness, therefore staggers. Yet attempts to understand why the reported relationships between fluctuating asymmetry, stress and fitness are so heterogeneous--i.e. whether the associations are truly weak or non-existent or whether they become confounded during different stages of the analytical pathways remain surprisingly scarce. Hence, we attempt to disentangle these causes, by reviewing the various statistical and conceptual factors that are suspected to confound potential relationships between fluctuating asymmetry, stress and fitness. Two main categories of factors are discerned: those associated with the estimation of developmental stability through fluctuating asymmetry and those associated with the effects of genotype and environment on developmental stability. Next, we describe a series of statistical tools that have recently been developed to help reduce this noise. We argue that the current lack of a theoretical framework that predicts if and when relationships with developmental stability can be expected, urges for further theoretical and empirical research, such as on the genetic architecture of developmental stability in stressed populations. If the underlying developmental mechanisms are better understood, statistical patterns of asymmetry variation may become a biologically meaningful tool.     

Inbreeding depression in two species of Mimulus (Scrophulariaceae) with contrasting mating systems

We examined the effect of self- and cross-pollination on germination success, flowering probability, pollen and ovule production, survivorship, and adult aboveground biomass in two species of Mimulus with contrasting mating systems: the highly seifing M. micranthus and an outcrossing population of M. guttatus. Cross-pollinations were performed both within and between populations in order to examine the scale at which the genetic load is distributed. We found significant inbreeding depression in M. guttatus in four of the six traits, with the highest inbreeding depression observed in biomass (68% and 69% based on within- and between-population crosses, respectively) and lowest in ovule production (21% based on between-population crosses only). M. micranthus displayed significant inbreeding depression in only two of the six traits examined. Again, we observed the highest inbreeding depression in biomass (47–60% based on within- and between-population crosses, respectively), but both traits showing significant differences between self and outcross progeny expressed lower inbreeding depression than in M. guttatus. We detected no significant inbreeding depression for either pollen or ovule production in M. micranthus. An estimate of total inbreeding depression based on the multiplicative effects of all traits was also lower in M. micanthus than∗∗∗ in M. guttatus. Our results are consistent with the expected purging of genetic load in populations with high selfing rates. The absence of inbreeding depression in M. micranthus pollen and ovule production, two traits with strong links to fitness in a selfing annual, further suggests the important role of directional selection in determining the population's genetic load. Comparison of cross-pollinations made within and between populations revealed little evidence of divergence of genetic load among the M. micranthus and M. guttatus populations examined.     

Evolution of Adaptation and Mate Choice: Parental Relatedness Affects Expression of Phenotypic Variance in a Natural Population

Mating between relatives generally results in reduced offspring viability or quality, suggesting that selection should favor behaviors that minimize inbreeding. However, in natural populations where searching is costly or variation among potential mates is limited, inbreeding is often common and may have important consequences for both offspring fitness and phenotypic variation. In particular, offspring morphological variation often increases with greater parental relatedness, yet the source of this variation, and thus its evolutionary significance, are poorly understood. One proposed explanation is that inbreeding influences a developing organism’s sensitivity to its environment and therefore the increased phenotypic variation observed in inbred progeny is due to greater inputs from environmental and maternal sources. Alternatively, changes in phenotypic variation with inbreeding may be due to additive genetic effects alone when heterozygotes are phenotypically intermediate to homozygotes, or effects of inbreeding depression on condition, which can itself affect sensitivity to environmental variation. Here we examine the effect of parental relatedness (as inferred from neutral genetic markers) on heritable and nonheritable components of developmental variation in a wild bird population in which mate choice is often constrained, thereby leading to inbreeding. We found greater morphological variation and distinct contributions of variance components in offspring from highly related parents: inbred offspring tended to have greater environmental and lesser additive genetic variance compared to outbred progeny. The magnitude of this difference was greatest in late-maturing traits, implicating the accumulation of environmental variation as the underlying mechanism. Further, parental relatedness influenced the effect of an important maternal trait (egg size) on offspring development. These results support the hypothesis that inbreeding leads to greater sensitivity of development to environmental variation and maternal effects, suggesting that the evolutionary response to selection will depend strongly on mate choice patterns and population structure.     

Exploring the genetic basis and proximate causes of female fertility advantage in gynodioecious Thymus vulgaris

In many gynodioecous species, females produce more viable seeds than hermaphrodites. Knowledge of the relative contribution of inbreeding depression in hermaphrodites and maternal sex effects to the female fertility advantage and the genetic basis of variation in female fertility advantage is central to our understanding of the evolution of gender specialization. In this study we examine the relative contribution of inbreeding and maternal sex to the female fertility advantage in gynodioecious Thymus vulgaris and quantify whether there is genetically based variation in female fertility advantage for plants from four populations. Following controlled self and outcross (sib, within-population, and between-population) pollination, females had a more than twofold fertility advantage (based on the number of germinating seeds per fruit), regardless of the population of origin and the type of pollination. Inbreeding depression on viable seed production by hermaphrodites occurred in two populations, where inbreeding had been previously detected. Biparental inbreeding depression on viable seed production occurred in three of four populations for females, but in only one population for hermaphrodites. Whereas the maternal sex effect may consistently enhance female fertility advantage, inbreeding effects may be limited to particular population contexts where inbreeding may occur. A significant family x maternal sex interaction effect on viable seed production was observed, illustrating that the extent of female fertility advantage varies significantly among families. This result is due to greater variation in hermaphrodite (relative to female) seed fertility between families. Despite this genetic variation in female fertility advantage and the highly female biased sex ratios in populations of T. vulgaris, gynodioecy is a stable polymorphism, suggesting that strong genetic and/or ecological constraints influence the stability of this polymorphism.     

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.     

The effect of inbreeding on fluctuating asymmetry in Scabiosa canescens (Dipsacaceae)

Developmental instability and fluctuating asymmetry (FA) describe the inability of organisms to correct for random accidents under development and has become a major but controversial topic in evolutionary biology. Theoretical models predict that the level of FA should increase as a result of inbreeding, but empirical results are ambiguous. Moreover, the relationship between fitness and FA is still debated. In the current study, plants from a population of Scabiosa canescens, a locally rare species in southern Sweden, were raised under uniform growth conditions to examine the effects of one-generation of selfing and outcrossing on FA in flower morphology. The level of flower FA was significantly higher (p = 0.038) for inbred progeny than for offspring derived from outcross pollinations. Given that earlier studies of this species have found no negative relation between heterozygosity and FA, the results support the conclusion that expression of deleterious recessive alleles are responsible for the increase of FA. There was no correlation between FA and estimates of five fitness-related traits when estimated at the individual level. However, a companion study found significant inbreeding depression for all fitness traits, and a negative association between FA and fitness could therefore be asserted at the treatment level (inbred/outbred progeny). Hence, FA seems to be useful to predict inbreeding depression in S. canescens, but specific individuals with high fitness cannot be identified based on their FA levels.     

Inbreeding alters resistance to insect herbivory and host plant quality in Mimulus guttatus (Scrophulariaceae)

Previous studies have demonstrated genetic variation for resistance to insect herbivores and host plant quality. The effect of plant mating system, an important determinant of the distribution of genetic variation, on host plant characteristics has received almost no attention. This study used a controlled greenhouse experiment to examine the effect of self- and cross-pollination in Mimulus guttatus (Scrophulariaceae) on resistance to and host plant quality for the xylem-feeding spittlebug Philaenus spumarius (Homoptera: Cercopidae). Spittlebugs were found to have a negative effect on two important fitness components in M. guttatus, flower production and above ground biomass. One of two M. guttatus populations examined showed a significant interaction between the pollination and herbivore treatments. In this case, the detrimental effects of herbivores on biomass and flower production were much more pronounced in inbred (self) plants. The presence of spittlebug nymphs increased inbreeding depression by as much as three times. Pollination treatments also had significant effects on important components of herbivore fitness, but these effects were in opposite directions in our two host plant populations. Spittlebug nymphs maturing on self plants emerged as significantly larger adults in one of our host plant populations, indicating that inbreeding increased host plant quality. In our second host plant population, spittlebug nymphs took significantly longer to develop to adulthood on self plants, indicating that inbreeding decreased host plant quality. Taken together these results suggest that the degree of inbreeding in host plant populations can have important and perhaps complex effects on the dynamics of plant-herbivore interactions and on mating-system evolution in the host.     

A quantitative model of the relationship between phenotypic variance and heterozygosity at marker loci under partial selfing.

Negative relationships between allozyme heterozygosity and morphological variance have often been observed and interpreted as evidence for increased developmental stability in heterozygotes. However, inbreeding can also generate such relationships by decreasing heterozygosity at neutral loci and redistributing genetic variance at the same time. I here provide a quantitative genetic model of this process by analogy with heterozygosity-fitness relationships. Inbreeding generates negative heterozygosity-variance relationships irrespective of the genetic architecture of the trait. This holds for fitness traits as well as neutral traits, the effect being stronger for fitness traits under directional dominance or overdominance. The order of magnitude of heterozygosity-variance regressions is compatible with empirical data even with very low inbreeding. Although developmental stability effects cannot be excluded, inbreeding is a parsimonious explanation that should be seriously considered to explain correlations between heterozygosity and both mean and variance of phenotypes in natural populations.     

The effects of inbreeding, genetic dissimilarity and phenotype on male reproductive success in a dioecious plant

Pollen fate can strongly affect the genetic structure of populations with restricted gene flow and significant inbreeding risk. We established an experimental population of inbred and outbred Silene latifolia plants to evaluate the effects of (i) inbreeding depression, (ii) phenotypic variation and (iii) relatedness between mates on male fitness under natural pollination. Paternity analysis revealed that outbred males sired significantly more offspring than inbred males. Independently of the effects of inbreeding, male fitness depended on several male traits, including a sexually dimorphic (flower number) and a gametophytic trait (in vitro pollen germination rate). In addition, full-sib matings were less frequent than randomly expected. Thus, inbreeding, phenotype and genetic dissimilarity simultaneously affect male fitness in this animal-pollinated plant. While inbreeding depression might threaten population persistence, the deficiency of effective matings between sibs and the higher fitness of outbred males will reduce its occurrence and counter genetic erosion.