EVOLUTION OF THE MAGNITUDE AND TIMING OF INBREEDING DEPRESSION IN PLANTS

Estimates of inbreeding depression obtained from the literature were used to evaluate the association between inbreeding depression and the degree of self-fertilization in natural plant populations. Theoretical models predict that the magnitude of inbreeding depression will decrease with inbreeding as deleterious recessive alleles are expressed and purged through selection. If selection acts differentially among life history stages and deleterious effects are uncorrelated among stages, then the timing of inbreeding depression may also evolve with inbreeding. Estimates of cumulative inbreeding depression and stage-specific inbreeding depression (four stages: seed production of parent, germination, juvenile survival, and growth/reproduction) were compiled for 79 populations (using means of replicates, N = 62) comprising 54 species from 23 families of vascular plants. Where available, data on the mating system also were collected and used as a measure of inbreeding history. A significant negative correlation was found between cumulative inbreeding depression and the primary selfing rate for the combined sample of angiosperms (N = 35) and gymnosperms (N = 9); the correlation was significant for angiosperms but not gymnosperms examined separately. The average inbreeding depression in predominantly selfing species (δ = 0.23) was significantly less (43%) than that in predominantly outcrossing species (δ = 0.53). These results support the theoretical prediction that selfing reduces the magnitude of inbreeding depression. Most self-fertilizing species expressed the majority of their inbreeding depression late in the life cycle, at the stage of growth/reproduction (14 of 18 species), whereas outcrossing species expressed much of their inbreeding depression either early, at seed production (17 of 40 species), or late (19 species). For species with four life stages examined, selfing and outcrossing species differed in the magnitude of inbreeding depression at the stage of seed production (selfing δ = 0.05, N = 11; outcrossing δ = 0.32, N = 31), germination (selfing δ = 0.02, outcrossing δ = 0.12), and survival to reproduction (selfing δ = 0.04, outcrossing δ = 0.15), but not at growth and reproduction (selfing δ = 0.21, outcrossing δ = 0.27); inbreeding depression in selfers relative to outcrossers increased from early to late life stages. These results support the hypothesis that most early acting inbreeding depression is due to recessive lethals and can be purged through inbreeding, whereas much of the late-acting inbreeding depression is due to weakly deleterious mutations and is very difficult to purge, even under extreme inbreeding.     

Little to no inbreeding depression in a tapeworm with mixed mating

Meta‐studies on hermaphrodites have found a negative relationship between primary selfing rates and levels of inbreeding depression (ID) and, thus, generally support purging in inbred systems. However, in plants, high among‐taxa variance in ID results in no difference in the mean ID between outcrossing and mixed‐mating taxa. Selective interference likely explains high ID among mixed‐mating taxa, whereas low levels of ID among mixed‐mating taxa are not as stressed. Among animal hermaphrodites, primarily molluscs, there are little data on mixed‐mating systems. To fill a taxonomic and mating system gap, we tested for ID in a mixed‐mating tapeworm, Oochoristica javaensis. We provide a direct estimate of ID across infection of an intermediate host by comparing selfing rates at two life history stages. We found little to no evidence for ID, and the level of ID falls in line with what is reported for highly selfing species even though O. javaensis has mixed mating. We discuss this result within the context of kin mating in O. javaensis. Our results emphasize that primary selfing rates alone may be insufficient to classify the inbreeding history in all species when testing for a relationship to ID. Mixed‐mating taxa, and possibly some outcrossing taxa, may exhibit low levels of ID if biparental inbreeding is also driving purging. We advocate that ID studies report estimates of inbreeding history (e.g. F_IS or identity disequilibrium) from nature‐derived adult samples to provide context rather than relying on primary selfing rates alone.     

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     

Inbreeding depression is high in a self‐incompatible perennial herb population but absent in a self‐compatible population showing mixed mating

High inbreeding depression is thought to be one of the major factors preventing evolutionary transitions in hermaphroditic plants from self‐incompatibility (SI) and outcrossing toward self‐compatibility (SC) and selfing. However, when selfing does evolve, inbreeding depression can be quickly purged, allowing the evolution of complete self‐fertilization. In contrast, populations that show intermediate selfing rates (a mixed‐mating system) typically show levels of inbreeding depression similar to those in outcrossing species, suggesting that selection against inbreeding might be responsible for preventing the transition toward complete self‐fertilization. By implication, crosses among populations should reveal patterns of heterosis for mixed‐mating populations that are similar to those expected for outcrossing populations. Using hand‐pollination crosses, we compared levels of inbreeding depression and heterosis between populations of Linaria cavanillesii (Plantaginaceae), a perennial herb showing contrasting mating systems. The SI population showed high inbreeding depression, whereas the SC population displaying mixed mating showed no inbreeding depression. In contrast, we found that heterosis based on between‐population crosses was similar for SI and SC populations. Our results are consistent with the rapid purging of inbreeding depression in the derived SC population, despite the persistence of mixed mating. However, the maintenance of outcrossing after a transition to SC is inconsistent with the prediction that populations that have purged their inbreeding depression should evolve toward complete selfing, suggesting that the transition to SC in L. cavanillesii has been recent. SC in L. cavanillesii thus exemplifies a situation in which the mating system is likely not at an equilibrium with inbreeding depression.     

Population genetics and breeding system of Tupistra pingbianensis (Liliaceae), a naturally rare plant endemic to SW China

The levels and partitioning of genetic diversity and inbreeding depression were investigated in Tupistra pingbianensis, a narrow endemic of South-east Yunnan, China, characterized by a naturally fragmented distribution due to extreme specialization on a rare habitat type. Here genetic diversity and patterns of genetic variation within and among 11 populations were analyzed using AFLP markers with 97 individuals across its whole geographical range. High levels of genetic variation were revealed both at the species level (P₉₉ = 96.012%; H_t = 0.302) and at the population level (P₉₉ = 51.41%; H_s = 0.224). Strong genetic differentiation among populations was also detected (F_ST = 0.2961; ⍬^嘦= 0.281), which corresponded to results reported for typical animal-pollinated, mixed selfing and outcrossing plant species. This result was consistent with mating patterns detected by our pollination experiments. The indirect estimate of gene flow based on ⍬^嘦 was low (N_m = 0.64). Special habitat and its life history traits may play an important role in shaping the genetic diversity and the genetic structure of this species. A pollination experiment also fail to detect significant inbreeding depression upon F₁ fruit set, seed weight and germinate rate fitness-traits. Since naturally rare species T. pingbianensis is not seriously genetically impoverished and likely to have adapted to tolerating a high level of inbreeding early in its history, we propose this species need only periodic monitoring to ensure their continued persistence but not intervention to remain viable.     

Effects of inbreeding on the magnitude of inbreeding depression in a highly self-fertilizing tree, Magnolia obovata

Inbreeding depression is one of the major selective forces driving the evolution of mating systems. Previous theories predict that long-lived plants will show a negative correlation between inbreeding depression and the level of inbreeding (as determined by an inbreeding coefficient) at maturity, but the extent of this correlation may vary among life stages because of variation in the genetic basis for inbreeding depression at different stages. To test this prediction, I used electrophoretic allozyme analysis and pollination experiments to examine the fixation index (F _is) at maturity and inbreeding depression in the early and late life stages of two populations with different outcrossing rates of a highly self-fertilizing tree, Magnolia obovata. The magnitude of inbreeding depression for early survival (δ _e) in an outcrossing population (t _m = 0.51; F _is = −0.015) was higher (δ _e = 0.97) than that in an inbreeding population (t _m = 0.18; F _is = 0.15; δ _e = 0.38). From these results, I estimated that both populations exhibited high inbreeding depression for late survival (δ _l) (0.94 in the outcrossing population and 0.93 in the inbreeding one) and lifetime survival (δ _t) (0.99 and 0.96, respectively). My results and previously published data demonstrate the predicted relationship between inbreeding depression and the level of inbreeding for early survival, but not for late survival. This suggests that there is a differential genetic basis for inbreeding depression at different life stages. The inbreeding depression for late survival appears to play a central role in the maintenance of reproductive traits that promote outcrossing in M. obovata.     

Mating Systems of Psychotria tenuinervis (Rubiaceae): Distance from Anthropogenic and Natural Edges of Atlantic Forest Fragment

The aim of this study was to determine and compare the mating systems among Psychotria tenuinervis populations at anthropogenic edges, natural edges, and the forest interior using allozyme electrophoresis of naturally pollinated progeny arrays. P. tenuinervis showed low outcrossing rates, varying from 37% to 50% of the mating attributable to outcrossing and 50% to 63% attributable to self-fertilization, in the three habitats. The forest interior had the highest outcrossing rate (t _m = 0.50 and t _s = 0.43) among the three habitats. However, there were no differences in either multilocus or single-locus rates among the three habitats, indicating that the contribution of biparental inbreeding to the apparent selfing rate in these populations was very low. The multilocus (t _m) and single-locus (t _s) outcrossing rates for the P. tenuinervis in the sample plots within each habitat showed great heterogeneity. In conclusion, edge creation seems not to influence its mating systems. Additionally, although P. tenuinervis is a distylous species, the population’s inbreeding can be attributed almost entirely to self-fertilization.     

Mating system and ploidy influence levels of inbreeding depression in Clarkia (Onagraceae)

Inbreeding depression is the reduction in offspring fitness associated with inbreeding and is thought to be one of the primary forces selecting against the evolution of self-fertilization. Studies suggest that most inbreeding depression is caused by the expression of recessive deleterious alleles in homozygotes whose frequency increases as a result of self-fertilization or mating among relatives. This process leads to the selective elimination of deleterious alleles such that highly selfing species may show remarkably little inbreeding depression. Genome duplication (polyploidy) has also been hypothesized to influence levels of inbreeding depression, with polyploids expected to exhibit less inbreeding depression than diploids. We studied levels of inbreeding depression in allotetraploid and diploid species of Clarkia (Onagraceae) that vary in mating system (each cytotype was represented by an outcrossing and a selfing species). The outcrossing species exhibited more inbreeding depression than the selfing species for most fitness components and for two different measures of cumulative fitness. In contrast, though inbreeding depression was generally lower for the polyploid species than for the diploid species, the difference was statistically significant only for flower number and one of the two measures of cumulative fitness. Further, we detected no significant interaction between mating system and ploidy in determining inbreeding depression. In sum, our results suggest that a taxon's current mating system is more important than ploidy in influencing levels of inbreeding depression in natural populations of these annual plants.     

Inbreeding depression and selfing rates in a self-compatible,hermaphroditic species,Schiedea membranacea (Caryophyllaceae)

Inbreeding depression and selfing rates were investigated in Schiedea membranacea (Caryophyllaceae), a hermaphroditic species endemic to the Hawaiian Islands. Most theoretical models predict high inbreeding depression in outcrossing hermaphroditic species and low inbreeding depression in inbreeding species. Although high outcrossing rates and high levels of inbreeding depression are characteristic of many species of Schiedea, self- fertilization is common among relatives of hermaphroditic S. membranacea, and high selfing rates and low levels of inbreeding depression were predicted in this species. Sixteen individuals grown in the greenhouse were used to produce selfed and outcrossed progeny. Inbreeding depression, which was evident throughout the stages measured (percentage viable seeds per capsule, mean seed mass, percentage seed germination, percentage seedling survival, and biomass after 8 mo), averaged 0.70. Inbreeding depression among maternal families varied significantly for all measured traits and ranged from −0.12 to 0.97. Using isozyme analysis, the multilocus selfing rate varied from 0.13 to 0.38 over 4 yr. Contrary to the initial prediction of high selfing and low inbreeding depression based on phylogenetic relationships within Schiedea, low selfing rates and high levels of inbreeding depression were found in S. membranacea. These results indicate that outcrossing is stable in this species and maintained by high levels of inbreeding depression.     

Plant traits correlated with generation time directly affect inbreeding depression and mating system and indirectly genetic structure

Background  

Understanding the mechanisms that control species genetic structure has always been a major objective in evolutionary studies. The association between genetic structure and species attributes has received special attention. As species attributes are highly taxonomically constrained, phylogenetically controlled methods are necessary to infer causal relationships. In plants, a previous study controlling for phylogenetic signal has demonstrated that Wright's F _ST, a measure of genetic differentiation among populations, is best predicted by the mating system (outcrossing, mixed-mating or selfing) and that plant traits such as perenniality and growth form have only an indirect influence on F _ST via their association with the mating system. The objective of this study is to further outline the determinants of plant genetic structure by distinguishing the effects of mating system on gene flow and on genetic drift. The association of biparental inbreeding and inbreeding depression with population genetic structure, mating system and plant traits are also investigated.     

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 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.     

The cost of fluctuating inbreeding depression

We present a phenotypic model for the evolution of self-fertilization in an infinite population of annual hermaphrodites for the case in which fitness and inbreeding depression vary among generations (e.g., due to fluctuations in the environment from year to year). Conditions for the evolution of selfing, mixed mating, and outcrossing are derived and are compared with results from numerical calculations that assume a normal distribution of inbreeding depression. In contrast to the situation in which inbreeding depression does not vary, when inbreeding depression fluctuates in a stochastic manner among generations with a mean less than 0.5, selfing is not necessarily selected. Thus, fluctuating inbreeding depression can be viewed as an additional cost of selfing that may stabilize mixed mating systems. These results emphasize the need to take into account fluctuating inbreeding depression in empirical studies aimed at understanding mating system evolution in annuals.     

OUTCROSSING RATE AND INBREEDING DEPRESSION IN TWO ANNUAL MONOECIOUS HERBS,BEGONIA HIRSUTA AND B. SEMIOVATA

Most models of mating-system evolution predict inbreeding depression to be low in inbred populations due to the purging of deleterious recessive alleles. This paper presents estimates of outcrossing rates and inbreeding depression for two highly selfing, monoecious annuals Begonia hirsuta and B. semiovata. Outcrossing rates were estimated using isozyme polymorphisms, and the magnitude of inbreeding depression was quantified by growing progeny in the greenhouse produced through controlled selfing and outcrossing. The estimated single-locus outcrossing rate was 0.03 ± 0.01 (SE) for B. hirsuta and 0.05 ± 0.02 for B. semiovata. In both species, the seed production of selfed flowers was on average 12% lower than that of outcrossed flowers (B. hirsuta P = 0.07, B. semiovata P < 0.05, mixed model ANOVAs). There was no significant effect of crosstype on germination rate or survival, but selfed offspring had a lower dry mass than outcrossed offspring 18 weeks after planting in both species (on average 18% lower in B. hirsuta and 31% lower in B. semiovata). Plants that were the products of selfing began flowering later than plants produced through outcrossing in B. semiovata, but not in B. hirsuta. The effects of crosstype on seed production (B. semiovata), days to first flower and offspring dry mass (both species) varied among maternal parents, as indicated by significant crosstype x maternal parent interactions for these characters. Both species showed significant inbreeding depression for total fitness (estimated as the product of seed production, germination rate, survival and dry mass at 18 weeks). In B. hirsuta, the average total inbreeding depression was 22% (range -57%-98%; N = 23 maternal parents), and in B. semiovata, it was 42% (-11%-84%; N = 21). This study demonstrates that highly selfing populations can harbor substantial inbreeding depression. Our findings are consistent with the hypothesis that a high mutation rate to mildly deleterious alleles contributes to the maintenance of inbreeding depression in selfing populations.     

INBREEDING DEPRESSION IN TWO MIMULUS TAXA MEASURED BY MULTIGENERATIONAL CHANGES IN THE INBREEDING COEFFICIENT

In mixed-mating plant populations, one can estimate the relative fitness of selfed progeny w by measuring the inbreeding coefficient F and selfing rate s of adults of one generation, together with F of adults in the following generation (after selection). In the first application of this multigenerational method, we estimated F and s for adults over three consecutive generations in adjacent populations of two annual Mimulus taxa: the outbreeding M. guttatus and the inbreeding M. platycalyx. This gave estimates of w for the last two generations. Although average multilocus selfing rates were high in both taxa (0.63 in M. guttatus; 0.84 in M. platycalyx), the relative fitness of selfed progeny averaged only 0.19 in M. guttatus and 0.32 in M. platycalyx. An alternative estimator for w that incorporates biparental inbreeding gave even lower estimates of w. These values are significantly below the 0.5 threshold thought to favor selfing, and show that partially selfing populations can harbor substantial genetic load. In accordance with the purging hypothesis, the more highly selfing M. platycalyx showed marginally lower inbreeding depression than M. guttatus in both years (P = 0.08). Inbreeding depression and selfing rates also varied among years in concert among taxa. Several sources of bias are discussed, but computer simulations indicate it is unlikely that w is biased downwards by linkage of marker loci to load loci.     

Reduced mate availability leads to evolution of self‐fertilization and purging of inbreeding depression in a hermaphrodite

Basic models of mating‐system evolution predict that hermaphroditic organisms should mostly either cross‐fertilize, or self‐fertilize, due to self‐reinforcing coevolution of inbreeding depression and outcrossing rates. However transitions between mating systems occur. A plausible scenario for such transitions assumes that a decrease in pollinator or mate availability temporarily constrains outcrossing populations to self‐fertilize as a reproductive assurance strategy. This should trigger a purge of inbreeding depression, which in turn encourages individuals to self‐fertilize more often and finally to reduce male allocation. We tested the predictions of this scenario using the freshwater snail Physa acuta, a self‐compatible hermaphrodite that preferentially outcrosses and exhibits high inbreeding depression in natural populations. From an outbred population, we built two types of experimental evolution lines, controls (outcrossing every generation) and constrained lines (in which mates were often unavailable, forcing individuals to self‐fertilize). After ca. 20 generations, individuals from constrained lines initiated self‐fertilization earlier in life and had purged most of their inbreeding depression compared to controls. However, their male allocation remained unchanged. Our study suggests that the mating system can rapidly evolve as a response to reduced mating opportunities, supporting the reproductive assurance scenario of transitions from outcrossing to selfing.     

RECONSTRUCTING ORIGINS OF LOSS OF SELF‐INCOMPATIBILITY AND SELFING IN NORTH AMERICAN ARABIDOPSIS LYRATA: A POPULATION GENETIC CONTEXT

Theoretical and empirical comparisons of molecular diversity in selfing and outcrossing plants have primarily focused on long‐term consequences of differences in mating system (between species). However, improving our understanding of the causes of mating system evolution requires ecological and genetic studies of the early stages of mating system transition. Here, we examine nuclear and chloroplast DNA sequences and microsatellite variation in a large sample of populations of Arabidopsis lyrata from the Great Lakes region of Eastern North American that show intra‐ and interpopulation variation in the degree of self‐incompatibility and realized outcrossing rates. Populations show strong geographic clustering irrespective of mating system, suggesting that selfing either evolved multiple times or has spread to multiple genetic backgrounds. Diversity is reduced in selfing populations, but not to the extent of the severe loss of variation expected if selfing evolved due to selection for reproductive assurance in connection with strong founder events. The spread of self‐compatibility in this region may have been favored as colonization bottlenecks following glaciation or migration from Europe reduced standing levels of inbreeding depression. However, our results do not suggest a single transition to selfing in this system, as has been suggested for some other species in the Brassicaceae.     

MUTATIONAL MELTDOWN IN SELFING ARABIDOPSIS LYRATA

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

Inbreeding and inbreeding depression in a threatened endemic plant,the African violet (Saintpaulia ionantha ssp. grotei), of the East Usambara Mountains,Tanzania

Mating among closely‐related individuals in small and isolated plant populations may result in reduced vigour of the inbred offspring, i.e. inbreeding depression, especially in naturally outbreeding plants. Occurrence of inbreeding and inbreeding depression was studied in Saintpaulia ionantha ssp. grotei, a threatened endemic plant species with a narrow ecological amplitude from the East Usambara Mountains. The level of inbreeding (measured as the fixation index, F) was investigated in twelve populations by analyzing variation at one microsatellite marker locus. The effect of one generation of selfing and outcrossing on the progeny fitness was studied by controlled crosses in two small patches that differ in the level isolation. The fixation index (F) across the populations was on the average 0.21 and varied among the populations from substantial inbreeding (F = 0.58) to surplus heterozygosity (F = −0.29). High inbreeding depression (δ) was observed at early and late stages of the life‐cycle. The isolated patch exhibited lower inbreeding depression than did the non‐isolated patch. The results of this study suggest that inbreeding and subsequent inbreeding depression are potential threats to the survival of Saintpaulia populations.     

STRONG INBREEDING DEPRESSION IN TWO SCANDINAVIAN POPULATIONS OF THE SELF‐INCOMPATIBLE PERENNIAL HERB ARABIDOPSIS LYRATA

Inbreeding depression is a key factor influencing mating system evolution in plants, but current understanding of its relationship with selfing rate is limited by a sampling bias with few estimates for self‐incompatible species. We quantified inbreeding depression (δ) over two growing seasons in two populations of the self‐incompatible perennial herb Arabidopsis lyrata ssp. petraea in Scandinavia. Inbreeding depression was strong and of similar magnitude in both populations. Inbreeding depression for overall fitness across two seasons (the product of number of seeds, offspring viability, and offspring biomass) was 81% and 78% in the two populations. Chlorophyll deficiency accounted for 81% of seedling mortality in the selfing treatment, and was not observed among offspring resulting from outcrossing. The strong reduction in both early viability and late quantitative traits suggests that inbreeding depression is due to deleterious alleles of both large and small effect, and that both populations experience strong selection against the loss of self‐incompatibility. A review of available estimates suggested that inbreeding depression tends to be stronger in self‐incompatible than in self‐compatible highly outcrossing species, implying that undersampling of self‐incompatible taxa may bias estimates of the relationship between mating system and inbreeding depression.