Inbreeding depression in Campanula rapunculoides L. I. A comparison of inbreeding depression in plants derived from strong and weak self-incompatibility phenotypes

The evolution of selfing taxa from outcrossing ancestors has occurred repeatedly and is the subject of many theoretical models, yet few empirical studies have examined the immediate consequences of inbreeding in a population with variable expression of self-incompatibility. Because self-incompatibility breaks down with floral age in Campanula rapunculoides, we were able to mate outbred and selfed maternal plants in a crossing design which produced progeny with inbreeding coefficients of 0, 0.25, 0.50 and 0.75. Cumulative inbreeding depression in plants that were selfed for one generation was very high in families derived from strongly self-incompatible plants (average δ = 0.98), and somewhat lower in families derived from plants with weaker expression of self-incompatibility (average δ = 0.90). Relative to outbred progeny, inbred progeny produced fewer seeds, had lower rates of germination, less vegetative growth and fewer flowers per plant. Inbred progeny also took longer to germinate, and longer to produce a first leaf and to flower. Interestingly, inbred plants also produced 40% fewer seeds than outcrossed plants (t-test P < 0.001) even when mated to the same, unrelated pollen donor, suggesting that inbreeding can produce profound maternal effects. Most importantly, our results demonstrate that progeny derived from plants with stronger expression of self-incompatibility exhibited greater levels of inbreeding depression than progeny from plants with weaker expression of self-incompatibility. Moreover, the decline in fitness (cumulative, ln-transformed) over the four inbreeding levels was steeper for the progeny of the strongly self-incompatible lineages. These empirical results suggest that inbreeding depression and mating system phenotype have the potential to coevolve.     

FITNESS CONSEQUENCES OF MATERNAL AND NONMATERNAL COMPONENTS OF INBREEDING IN THE GYNODIOECIOUS PHACELIA DUBIA

I assessed the relationship between the level of inbreeding, F, and fitness, and the effects of nonmaternal and maternal components of inbreeding on fitness in Phacelia dubia. I conducted two generations of controlled crosses and tested the performance of the F₂ progeny in field and artificial conditions covering the whole life cycle. Inbreeding significantly decreased the individual contribution of seeds to the next generation in the field, but this decrease apparently is not enough to explain the maintenance of gynodioecy. The inbred progeny contributes significantly to the population genetic structure of P. dubia. Fitness estimates and fitness components tended to decrease, usually monotonically, with F. However, nonmonotonic relationships were found in male fitness components and, in some families, in fitness estimates, seed production per fruit, and establishment. Most of the inbreeding depression takes place at the level of seed establishment in the field, but, in artificial conditions the effects of inbreeding were similar at fecundity and establishment. I studied maternal and nonmaternal components of inbreeding by testing the effects of the relatedness of maternal grandparents and parents on the performance of the progeny. Both components affected fitness. Inbreeding depression was conditioned by the level of inbreeding of the maternal plant, but this interaction varied at different fitness components. Also, the magnitude and even the direction of the relationship between fitness and F changed as a result of the combined effects of maternal and nonmaternal components of inbreeding. Such interactions can render convex or concave fitness functions, giving in the latter case the appearance of a false purging. Maternal effects of inbreeding can result from several processes: maternal investment perhaps with serial adjustments during seed development, purging of recessive deleterious genes, and nucleocytoplasmic interactions. These results illustrate the importance of maternal effects of inbreeding, and the complex effects of inbreeding on fitness. A full understanding of the fitness consequences of inbreeding and, therefore, their potential implications in the evolution of breeding systems, should take into account male and female components as well as transgenerational effects in the context of the particular environment in which fitness is evaluated.     

The cost of selfing inEncyclia cochleata (Orchidaceae)

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

Inbreeding avoidance and the evolution of gender dimorphism in Wurmbea biglandulosa (Colchicaceae)

How females establish in populations of cosexuals is central to understanding the evolution of gender dimorphism in angiosperms. Inbreeding avoidance hypotheses propose that females can establish and be maintained if cosexual fitness is reduced because they self-fertilize, and their progeny express inbreeding depression. Here we assess the role of inbreeding avoidance in maintaining sexual system variation in Wurmbea biglandulosa. We estimated costs of self-pollination, mating patterns, and inbreeding depression in gender monomorphic (cosexuals only) and dimorphic (males and females) populations. Costs of selfing, estimated from seed set of experimentally self- and cross-pollinated flowers, were severe in both males and cosexuals (inbreeding depression, sigma = 0.86). In a field experiment, intact males that could self produced fewer seeds than both emasculated males and females, whereas seed set of intact and emasculated cosexuals did not differ. Thus, pollinator-mediated selfing reduces fitness of males but not cosexuals under natural conditions. Outcrossing rates of males revealed substantial selfing (t = 0.68), whereas females and cosexuals were outcrossed (0.92 and 0.97). For males, progeny inbreeding coefficients exceeded parental coefficients (0.220 vs. 0.009), whereas for females and cosexuals these coefficients did not differ and approached zero. Differences in coefficients between males and their progeny indicate that selfed progeny express severe inbreeding depression (sigma = 0.93). Combined with inbreeding depression for seed set, cumulative sigma = 0.99, indicating that most or all selfed zygotes fail to reach reproductive maturity. We propose that present sexual system variation in W. biglandulosa is maintained by high inbreeding depression coupled with differences in selfing rates among monomorphic and dimorphic populations.     

LIMITS TO REPRODUCTIVE SUCCESS IN A PARTIALLY SELF-INCOMPATIBLE HERB: FECUNDITY DEPRESSION AT SERIAL LIFE-CYCLE STAGES

Like many angiosperms, Crinum erubescens is partially self-compatible, producing fewer seeds upon selfing than after outcrossing. In this paper we test the relative magnitude of the prefertilization and postfertilization effects of self-incompatibility, inbreeding depression, or both in a natural population of this hermaphroditic tropical herb. We characterize prefertilization effects by examining pollen tube growth, while postfertilization effects are characterized by examination of embryo abortion and seed maturation. Statistical methods are developed to test the magnitude of these effects from one life-cycle stage to the next. We find that although pollen performance in selfed flowers is lower than that in outcrossed flowers, pollen performance is low overall. Postfertilization effects attributable to inbreeding depression account for a larger proportion of the reduction in fecundity in selfed compared to outcrossed flowers. Among naturally pollinated plants, despite ample pollen deposition, the numbers of fruits and seeds set are intermediate to selfed and outcrossed treatments.     

Outcrossing rate and inbreeding depression in the perennial yellow bush lupine, Lupinus arboreus (Fabaceae)

Little is known about the breeding systems of perennial Lupinus species. We provide information about the breeding system of the perennial yellow bush lupine, Lupinus arboreus, specifically determining self-compatibility, outcrossing rate, and level of inbreeding depression. Flowers are self-compatible, but autonomous self-fertilization rarely occurs; thus selfed seed are a product of facilitated selfing. Based on four isozyme loci from 34 maternal progeny arrays of seeds we estimated an outcrossing rate of 0.78. However, when we accounted for differential maturation of selfed seeds, the outcrossing rate at fertilization was lower, ～0.64. Fitness and inbreeding depression of 11 selfed and outcrossed families were measured at four stages: seed maturation, seedling emergence, seedling survivorship, and growth at 12 wk. Cumulative inbreeding depression across all four life stages averaged 0.59, although variation existed between families for the magnitude of inbreeding depression. Inbreeding depression was not manifest uniformly across all four life stages. Outcrossed flowers produced twice as many seeds as selfed flowers, but the mean performance of selfed and outcrossed progeny was not different for emergence, seedling survivorship, and size at 12 wk. Counter to assumptions about this species, L. arboreus is both self-compatible and outcrosses ～78% of the time.     

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.     

Elevation-related variation in the population characteristics of distylous Primula nivalis affects female fitness and inbreeding depression

The population characteristics of distylous species are highly sensitive to stochastic natural selection pressure.Therefore,populations growing under different environmental conditions may vary in floral morph ratios,potentially affecting female fitness and leading to inbreeding depression.However,the variation in offspring quality among populations as a result of inbreeding depression is poorly understood in distylous species.This study investigates variations in plant density,seed mass,seed viabilityfemale fitness,and post-dispersal inbreeding depression in both sexual morphs(long-styled and shortstyled plants)of the distylous Primula nivalis that were subjected to different pollination treatments along an elevational gradient from 1657 to 2704 m a.s.l.Population characteristics(morph plant density and ratio)and fruit set were significantly affected by sexual morph and elevation.Plant density and fruitset frequencies were lower for short-styled than for long-styled plants at 2704 m a.s.l.The seeds from the cross-pollinated flowers of both morphs were higher in quality than those of self-pollinated flowers.The female fitness of seeds from cross-pollinated flowers of both morphs was higher than that of seeds from open-pollinated and self-pollinated flowers.The female fitness of seeds from long-styled flowers was higher than that of seeds from short-styled flowers at all elevations.Inbreeding depression increased with elevation among plants with short-styled flowers but not among those with long-styled flowers.Variation in the elevation-dependent mating system might influence female fitness and affect inbreeding depression in both floral morphs.In conclusion,the low quality of seeds from short-styled flowers at high elevations might decrease short-styled flower frequency,affecting population characteristics.     

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

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

THE RELATIVE IMPORTANCE OF INBREEDING AND MATERNAL SEX IN DETERMINING PROGENY FITNESS IN SIDALCEA OREGANA SSP. SPICATA,A GYNODIOECIOUS PLANT

Gynodioecious plant populations contain both hermaphrodite and female individuals. For females to be maintained they must compensate for their loss of reproductive fitness through pollen. Females may achieve compensation by producing more and/or higher quality seeds than hermaphrodites. In this study, I investigated the independent and interactive effects of maternal sexual identity and inbreeding level on fitness of the progeny of hermaphrodites and females of Sidalcea oregana ssp.spicata. Seeds produced by selling hermaphrodites and by outcrossing or sib-crossing hermaphrodites and females, were planted in the field and greenhouse. Maternal-sex effects were substantial at the juvenile stages of the life cycle; seeds of females germinated in higher proportions and produced seedlings that grew significantly faster. Inbreeding effects were manifested primarily at the adult stage of the life cycle. Outcrossed plants were significantly larger and produced more flowers per plant than sib-crossed and selfed plants growing in the greenhouse. Progeny of hermaphrodites and females appeared to respond similarly to sib-matings. The maternal-sex effects observed in Sidalcea may have been related to cytoplasmically inherited factors and could be a driving force in the maintenance of females. Inbreeding depression could play a role in determining the fitness of both sex morphs, if females experience biparental inbreeding in the field. Frequent inbreeding of hermaphrodites may not be necessary to explain the maintenance of gynodioecy in this species.     

Evidence for selective fruit abortion in Banksia spinulosa (Proteaceae)

Fruit set is consistently low in the genus Banksia (Proteaceae). One explanation for low fruit:flower ratios is that excess flowers allow plants to selectively abort inferior progeny thereby increasing the average quality of the seed crop. We examined whether self-pollinated fruits were aborted selectively in a partially self-compatible population of Banksia spinulosa. Inflorescences were divided in half and pollinated with cross pollen on both sides, self pollen on both sides or cross pollen on one side and self pollen on the other. Seed production was reduced significantly by 38% on fully self-pollinated compared to fully cross-pollinated inflorescences, indicating partial self-incompatibility or inbreeding depression. On inflorescences pollinated with both cross and self pollen, selfed seed set was reduced by 63% compared to crossed seed set. On the selfed half of these inflorescences, fewer fruits and seeds were produced, and more fruits had aborted seeds than on randomly selected halves of fully self-pollinated inflorescences. These results suggest that self-fertilized embryos were developed in the absence of outcrossed progeny but were selectively aborted when outcrossed progeny were present on inflorescences. To determine the amount of outcrossing occurring in the study population, outcrossing rates (t) of naturally pollinated plants were examined by electrophoretic assessment of progeny arrays. Values of I did not differ significantly from one, indicating complete outcrossing. High outcrossing, despite the high probability of self pollen receipt under natural conditions, is consistent with the conclusion that selective abortion occurs in B. spinulosa.     

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

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

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.     

EFFECTS OF CROSS AND SELF-FERTILIZATION ON PROGENY FITNESS IN LOBELIA CARDINALIS AND L. SIPHILITICA

Inbreeding depression, or the decreased fitness of progeny derived from self-fertilization as compared to outcrossing, is thought to be the most general factor affecting the evolution of self-fertilization in plants. Nevertheless, data on inbreeding depression in fitness characters are almost nonexistent for perennials observed in their natural environments. In this study I measured inbreeding depression in both survival and fertility in two sympatric, short-lived, perennial herbs: hummingbird-pollinated Lobelia cardinalis (two populations) and bumblebee-pollinated L. siphilitica (one population). Crosses were performed by hand in the field, and seedlings germinated in the greenhouse. Levels of inbreeding depression were determined for one year in the greenhouse and for two to three years for seedlings transplanted back to the natural environment. Fertility was measured as flower number, which is highly correlated with seed production under natural conditions in these populations. Inbreeding depression was assessed in three ways: 1) survival and fertility within the different age intervals; 2) cumulative survival from the seed stage through each age interval; and 3) net fertility, or the expected fertility of a seed at different ages. Net fertility is a comprehensive measure of fitness combining survival and flower number. In all three populations, selfing had nonsignificant effects on the number and size of seeds. Lobelia siphilitica and one population of L. cardinalis exhibited significant levels of inbreeding depression between seed maturation and germination, excluding the consideration of possible differences in dormancy or longterm viability in the soil. There was no inbreeding depression in subsequent survival in the greenhouse in any population. In the field, significant survival differences between selfed and outcrossed progeny occurred only in two years and in only one population of L. cardinalis. For both survival and fertility there was little evidence for the expected differences among families in inbreeding depression. Compared to survival, inbreeding depression in fertility (flower number) tended to be much higher. By first-year flower production, the combined effects on survival and flower number caused inbreeding depression in net fertility to reach 54%, 34% and 71% for L. siphilitica and the two populations of L. cardinalis. By the end of the second year of flowering in the field, inbreeding depression in net fertility was 53% for L. siphilitica and 54% for one population of L. cardinalis. For the other population of L. cardinalis, these values were 76% through the second year of flowering and 83% through the third year. Such high levels of inbreeding depression should strongly influence selection on those characters affecting self-fertilization rates in these two species.     

The mating system and relative performance of selfed and outcrossed progeny in Arabis fecunda (Brassicaceae)

We estimated rates of self-fertilization and inbreeding depression in the hermaphroditic perennial Arabis fecunda. Crosses were made on mesh-bagged wild plants, yielding 12 plants producing self-, outcross-, and naturally fertilized seeds that were then grown in a greenhouse. Analysis of variance indicated aboveground dry weight at 160 days differed among pollination treatments, but mean seed weight, number of seeds per fruit, and days to germination did not. For dry weight, selfed progeny have 9.4% reduction and naturally pollinated progeny a 3.7% reduction compared to outcrossed progeny, suggesting a 39% selling rate in Arabis. Significant inbreeding depression and a mixed mating system are evidence that the deleterious effects of self-fertilization are maintained through high rates of mildly deleterious mutation. Significant maternal-parent-by-pollination-treatment interactions for mean seed weight and dry weight are consistent with inbreeding depression caused by deleterious recessives and varying past maternal inbreeding.     

Inbreeding depression in an autotetraploid herb: a three cohort field study

Autotetraploids are predicted to have reduced inbreeding depression relative to diploids. However, recent theory and information on genomic changes following autopolyploidy suggest that inbreeding depression may be closer to diploids. In three consecutive years, self and outcross pollinations were conducted on autotetraploid Campanulastrum americanum, seeds were planted into native sites, and biennial offspring were followed through seed production. Inbred individuals had lower germination rates, reduced survival, were smaller, and flowered later, producing fewer fruits with fewer seeds. Inbred offspring had 6% of the cumulative fitness of outcross offspring. Although performance varied substantially among cohorts, inbreeding depression for cumulative fitness was relatively constant, with delta ranging only from 0.92 to 0.95. C. americanum, like many outcrossing species, expressed very high amounts of inbreeding depression. This supports the hypothesis that inbreeding depression of some autotetraploids may be similar to that of diploids. Furthermore, few studies have measured temporal variation in inbreeding depression. Constant inbreeding depression given a sixfold range in cohort performance suggests that inbreeding depression may be relatively robust to environmental variation experienced by natural populations.     

EFFECTS OF SERIAL INBREEDING ON FITNESS COMPONENTS IN IMPATIENS CAPENSIS

Studies of inbreeding depression in wild plants customarily compare the fitness of outcrossed progeny to progeny derived from one generation of self-pollination. We compare levels of inbreeding depression in a greenhouse in two populations of jewelweed using progeny derived from random outcrosses, one generation of self-pollination, and three generations of selling. The progeny have expected inbreeding coefficients of, respectively, 0, 0.5, and 0.875. Seedling survivorship declined linearly with the level of inbreeding in both populations. Inbreeding also increased the variability of emergence date. Maternal family membership affected early seedling performance and often interacted significantly with the level of inbreeding. In contrast, path analyses reveal that inbreeding had both negative linear and positive quadratic direct effects on seed and final plant weight, causing the highly inbred progeny to outperform progeny derived from one generation of selfing. These results suggest either the rapid purging of deleterious alleles or diminishing epistasis among the loci affecting these characters. It is not clear why the loci affecting survival responded differently.     

The Effect of Pollen Source vs. Flower Type on Progeny Performance and Seed Predation under Contrasting Light Environments in a Cleistogamous Herb

Dimorphic cleistogamy is a specialized form of mixed mating system where a single plant produces both open, potentially outcrossed chasmogamous (CH) and closed, obligately self-pollinated cleistogamous (CL) flowers. Typically, CH flowers and seeds are bigger and energetically more costly than those of CL. Although the effects of inbreeding and floral dimorphism are critical to understanding the evolution and maintenance of cleistogamy, these effects have been repeatedly confounded. In an attempt to separate these effects, we compared the performance of progeny derived from the two floral morphs while controlling for the source of pollen. That is, flower type and pollen source effects were assessed by comparing the performance of progeny derived from selfed CH vs. CL and outcrossed CH vs. selfed CH flowers, respectively. The experiment was carried out with the herb Ruellia nudiflora under two contrasting light environments. Outcrossed progeny generally performed better than selfed progeny. However, inbreeding depression ranges from low (1%) to moderate (36%), with the greatest value detected under shaded conditions when cumulative fitness was used. Although flower type generally had less of an effect on progeny performance than pollen source did, the progeny derived from selfed CH flowers largely outperformed the progeny from CL flowers, but only under shaded conditions and when cumulative fitness was taken into account. On the other hand, the source of pollen and flower type influenced seed predation, with selfed CH progeny the most heavily attacked by predators. Therefore, the effects of pollen source and flower type are environment-dependant and seed predators may increase the genetic differences between progeny derived from CH and CL flowers. Inbreeding depression alone cannot account for the maintenance of a mixed mating system in R. nudiflora and other unidentified mechanisms must thus be involved.     

INBREEDING DEPRESSION IN HYDROPHYLLUM APPENDICULATUM: ROLE OF MATERNAL EFFECTS,CROWDING, AND PARENTAL MATING HISTORY

This paper examines several aspects of the expression of inbreeding depression in an outcrossing, obligately biennial plant, Hydrophyllum appendiculatum (Hydrophyllaceae). The amount of inbreeding depression detected was small during the first year of life but increased with age and had significant effects on adult size and reproductive traits. The lack of significant inbreeding depression during early growth is likely due to the overriding influence of maternal environmental effects on seed size and seedling growth. However, as maternal effects decreased with age, the seedling's own genotype became a more important determinant of its fate. To examine whether the expression of inbreeding depression was sensitive to ecological conditions, selfed and outcrossed seedlings were grown alone or with other H. appendiculatum seedlings. No inbreeding depression was detected in the plants grown alone. In contrast, under competitive conditions, outcrossed seedlings were significantly larger than selfed seedlings by the end of the first growing season. To address whether parental mating history influences the amount of inbreeding depression expressed, I examined the consequences of two successive generations of selfing on seed set and seed weight. The amount of inbreeding depression increased following the second generation of selfing. In the first generation, seed set and seed weight differed by less than 5% between selfed and outcrossed progeny. However, both traits were 15% greater for outcrossed plants after two generations. These results indicate that the alleles responsible for the reductions in these traits were not purged and suggest the action of multiple loci with deleterious effects.     

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.