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

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

INBREEDING AND FITNESS IN THE FRESHWATER SNAIL LYMNAEA PEREGRA: AN EVALUATION OVER TWO GENERATIONS OF SELF-FERTILIZATION

Inbreeding depression was estimated from an outbreeding population of the freshwater snail Lymnaea peregra, on the basis of two successive generations of enforced selling and outcrossing, and 70 maternal lineages. Outcrossing was analyzed under two treatments, groups of two and five individuals. The fitness parameters measured included fecundity, growth, and survival. In the first generation, we contrasted three treatments (selfers vs. paired outcrossers and group outcrossers). Very similar results were obtained between the two outcrossing treatments. A strong self-fertilization depression (which includes parental fecundity and progeny fitness) was detected in the selling treatment (about 90%). In the second generation, there was again marked evidence for self-fertilization depression, with the highest contributions coming from parental fecundity and progeny hatching rate. Our results suggest that the decreased parental fecundity is a consequence of the mating system in the previous generation, although the role of partial self-incompatibility and the copulation behavior could not be ruled out. Hatching rate and early survival data are suggestive of purging of lethal mutations. Significant variation in fitness among selfing lineages was found for most fitness traits. Our experimental design also allowed to test for interactions among fitness loci. Only one trait of the nine studied behaved as expected under synergistic interactions. However we cannot rule out some purging during the experiment, which could have biased results towards linearity. Inbreeding depression was also inferred from the change of inbreeding level across generations in the same population. We obtained a value similar to the experimental estimate.     

CROSSING-DISTANCE EFFECTS IN DELPHINIUM NELSONII: OUTBREEDING AND INBREEDING DEPRESSION IN PROGENY FITNESS

Depending on its genetic causes, outbreeding depression in quantitative characters may occur first in the free-living F₁ generation produced by a wide cross. In 1981–1985, we generated F₁ progenies by hand-pollinating larkspurs (Delphinium nelsonii) with pollen from 1-m, 3-m, 10-m, or 30-m distances. From the spatial genetic structure indicated by previous electrophoretic and reciprocal transplantation studies, we estimate that these crosses range from being inbred (f ≈ 0.06) to outbred. We planted 594 seeds from 66 maternal sibships under natural conditions. As of 1992, there was strong evidence for both inbreeding depression and outbreeding depression. Progeny from intermediate crossing distances grew approximately twice as large as more inbred or outbred progeny in the first 5 yr after planting (P = 0.013, repeated measures ANOVA), and survived almost 1 yr longer on average (contrast of 3-m and 10-m treatments versus 1 m and 30 m; P = 0.028, ANOVA). Twenty maternal sibships produced flowering individuals; only four and two of these represented 1-m and 30-m crossing distances, respectively (P = 0.021, G-test). The cumulative fitness of intermediate distance sibships averaged about twice that of 1-m sibships, and five to eight times that of 30-m sibships (P = 0.017, ANOVA). Thus, even though progeny of 1-m crosses were inbred to a degree only about one-eighth that of selling, inbreeding depression approximated 50%, and outbreeding depression equaled or exceeded 50% for all fitness components.     

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.     

NATURAL SELECTION ON FLORAL TRAITS IN TWO SPECIES OF LOBELIA WITH DIFFERENT POLLINATORS

Using both multivariate and univariate regression techniques, I measured selection acting through female reproductive success in two hermaphroditic species with precise pollen placement but different pollinators: hummingbird-pollinated Lobelia cardinalis and bumblebee-pollinated L. siphilitica. Six traits were analyzed in two populations of L. cardinalis and one population of L. siphilitica: flower number, mean number of flowers open per day, inflorescence height, number of days in flower, median-flower date and nectar-stigma distance. In another study it was found that female reproductive success in one population of L. cardinalis was much less pollen limited than in the other two populations, and it was therefore expected that selection of female reproductive traits in this population would be weaker. In the univariate analyses correlations caused nearly all traits to have significant directional selection coefficients. However, in the multivariate analyses no traits in L. siphilitica experienced directional or quadratic selection. Selection acted differently in the two L. cardinalis populations. The less pollen-limited population experienced positive directional selection on flower number and median-flower date, while in the other L. cardinalis population there was positive directional selection on flower number and nectar-stigma distance and both positive directional and positive quadratic selection on height. The functional significance of floral traits in these two species and the probable effect of increased sample sizes are discussed.     

EFFECTS OF DIFFERENT LEVELS OF INBREEDING ON FITNESS COMPONENTS IN MIMULUS GUTTATUS

Recent theoretical work has shown that there can be selection favoring the maintenance of sexual reproduction and the evolution of increased recombination when deleterious mutations at different loci interact synergistically, such that the logarithm of fitness declines at a greater than linear rate with the number of harmful mutations per genome. The purpose of this experimental study was to determine whether synergism exists for genes affecting fitness components in two partially selfing populations of the monkey flower Mimulus guttatus. For each wild population, a large randomly mated base population was constructed and many independent lines, inbred to differing degrees, were extracted from this base population. Lines with expected inbreeding coefficients of 0, 0.25, 0.5, and 0.75 were raised simultaneously in the greenhouse and were scored for germination, flowering, flower production, and pollen viability. All fitness traits except germination success declined with increased inbreeding, but in spite of the substantial inbreeding depression found in this study, relatively little evidence of synergistic epistasis was found. The only trait that showed evidence of synergism was pollen viability. These results indicate that synergism is not strong for the fitness components measured in this study. The evidence for synergism from other published studies is also reviewed.     

EVOLUTIONARILY STABLE SELFING RATES OF HERMAPHRODITIC PLANTS IN COMPETING AND DELAYED SELFING MODES WITH ALLOCATION TO ATTRACTIVE STRUCTURES

I present a resource-allocation model to analyze how patterns of allocation to reproductive structures influence the evolution of selfing rates in hermaphrodites subject to competing and delayed forms of self-fertilization. The evolutionarily stable state does not depend on the mode of pollination. In contrast to previous models in which the number and the size of flowers were not considered, intermediate selfing is not evolutionarily stable with linear constraints on flower number and size. In contrast, intermediate selfing can be evolutionarily stable with nonlinear constraints on flower number and size. Optimal allocations to attractive structures increase and selfing rates decrease in the presence of inbreeding depression. In particular, stable intermediate levels of selfing may be favored when flower number is strongly constrained. Thus, nonlinear constraints on flower number and size could favor the evolution of intermediate selfing in either the delayed or the competing modes of selfing. Outcrossing is not favored in the absence of inbreeding depression, a result inconsistent with Holsinger's results in which allocation to attractive structures was not considered.     

INBREEDING AND ITS FITNESS EFFECTS IN AN INSULAR POPULATION OF SONG SPARROWS (MELOSPIZA MELODIA)

Inbreeding depression is thought to be a major factor affecting the evolution of mating systems and dispersal. While there is ample evidence for inbreeding depression in captivity, it has rarely been documented in natural populations. In this study, I examine data from a long-term demographic study of an insular population of song sparrows (Melospiza melodia) and present evidence for inbreeding depression. Forty-four percent of all matings on Mandarte Island, British Columbia, were among known relatives. Offspring of a full-sib mating (f = 0.25) experienced a reduction in annual survival rate of 17.5% on average. Over their lifetime, females with f = 0.25 produced 48% fewer young that reached independence from parental care. In contrast, male lifetime reproductive success was not affected by inbreeding. Reduced female lifetime reproductive success was mostly due to reduced hatching rates of the eggs of inbred females. Relatedness among the parents did not affect their reproductive success. Using data on survival from egg stage to breeding age, I estimated the average song sparrow egg on Mandarte Island to carry a minimum of 5.38 lethal equivalents (the number of deleterious genes whose cumulative effect is equivalent to one lethal); 2.88 of these lethal equivalents were expressed from egg stage to independence of parental care. This estimate is higher than most estimates reported for laboratory populations and lower than those reported for zoo populations. Hence, the costs of inbreeding in this population were substantial and slightly above those expected from laboratory studies. Variability in estimates of lethal equivalents among years showed that costs of inbreeding were not constant across years.     

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.     

EFFECTS OF NONRANDOM SEED ABORTION ON PROGENY PERFORMANCE IN PHASEOLUS COCCINEUS L

In this study, we sought to determine if Phaseolus coccineus normally aborts potentially viable seeds, and whether seed abortion is nonrandom with respect to progeny vigor. The ovaries of Phaseolus coccineus typically have six linearly arranged ovules. The three ovule positions at the stylar end are more likely to mature seeds than the three ovules at the basal end of the ovary. When we destroyed the developing ovules at the stylar end of the fruit after fertilization but before seed abortion, there was a significant increase in the probability that the ovules in the three basal positions would produce a mature seed. The probability of seed maturation in control fruits (no ovules destroyed) ranged from 38.3 to 42.7% over the three field seasons, whereas in the experimental fruits it ranged from 64.3 to 79.7% (similar to that of ovules at the stylar end in control fruits). We did not find any significant change in the probability of seed production in the three ovule positions in the stylar end of the ovary (the positions with high probability of seed maturation) when the basal ovules were experimentally destroyed. These results indicate that potentially viable seeds are regularly aborted in P. coccineus, with seed abortion more frequent in the basal ovule positions than at the stylar positions. In two greenhouse studies and one field study, we compared the vigor of progeny produced in the control fruits with the vigor of progeny produced in fruits where three ovules (either stylar or basal end) were destroyed. We found that the performance of the progeny from the three positions at the stylar end of the control fruits did not differ from the performance of the progeny from experimental fruits in which the three basal ovule positions were destroyed. In contrast, the progeny from the basal positions of the control fruits outperformed the progeny from the experimental fruits when the seeds were produced in the three basal ovules (stylar ovules destroyed). Our findings indicate that when the experimental treatment increased the probability of seed maturation, there was a significant decrease in the average vigor of the progeny in the remaining (basal) ovule positions.     

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.     

MAGNITUDE AND TIMING OF INBREEDING DEPRESSION IN SCOTS PINE (PINUS SYLVESTRIS L.)

Inbreeding depression is a major selective force favoring outcrossing in flowering plants. However, some self-fertilization should weaken the harmful effects of inbreeding by exposing deleterious alleles to selection. This study examines the maintenance of inbreeding depression in the predominantly outcrossing species Pinus sylvestris L. (Scots pine). Open-pollinated and self-fertilized progeny of 23 maternal trees, originating from a natural stand in southern Finland, were grown at two sites. We observed significant inbreeding depression in two of the four life stages measured. Inbreeding depression was largest for seed maturation (δ = 0.74), where seedset in open-pollinated strobili (70.9%) was about four times higher than in selfed strobili (18.3%). Inbreeding depression in postgermination survival (upto an age of 23 years) was also high (δ = 0.62–0.75). No significant differences in height (δ = 0.05) or flowering (δ = 0.14) of the trees after 23 years were observed. Cumulative inbreeding depression was high (δ = 0.90–0.94) and differed significantly among maternal families (range 0.45–1.00). The magnitude of inbreeding depression among the 23 maternal parents was not significantly correlated between early (seed maturation) and later (postgermination survival) life stages, suggesting that its genetic basis varies across the life cycle. Size differences among the progeny types diminished in time due to nonrandom size-specific mortality, causing a decrease in the inbreeding depression estimates for height over time. Our results indicate that Scots pine exhibits high levels of inbreeding depression during both early and later stages of the life cycle. It is argued that self-fertilization in Scots pine is inefficient in purging the genetic load caused by highly deleterious mutations because of the nearly complete loss of selfed individuals over time. This results in an effectively random mating outcrossing population.     

EFFECTS OF MATERNAL AND PATERNAL ENVIRONMENT AND GENOTYPE ON OFFSPRING PHENOTYPE IN SOLIDAGO ALTISSIMA L.

To predict the possible evolutionary response of a plant species to a new environment, it is necessary to separate genetic from environmental sources of phenotypic variation. In a case study of the invader Solidago altissima, the influences of several kinds of parental effects and of direct inheritance and environment on offspring phenotype were separated. Fifteen genotypes were crossed in three 5 × 5 diallels excluding selfs. Clonal replicates of the parental genotypes were grown in two environments such that each diallel could be made with maternal/paternal plants from sand/sand, sand/soil, soil/sand, and soil/soil. In a first experiment (1989) offspring were raised in the experimental garden and in a second experiment (1990) in the glasshouse. Parent plants growing in sand invested less biomass in inflorescences but produced larger seeds than parent plants growing in soil. In the garden experiment, phenotypic variation among offspring was greatly influenced by environmental heterogeneity. Direct genetic variation (within diallels) was found only for leaf characters and total leaf mass. Germination probability and early seedling mass were significantly affected by phenotypic differences among maternal plants because of genotype ( genetic maternal effects ) and soil environment ( general environmental maternal effects ). Seeds from maternal plants in sand germinated better and produced bigger seedlings than seeds from maternal plants in soil. They also grew taller with time, probably because competition accentuated the initial differences. Height growth and stem mass at harvest (an integrated account of individual growth history) of offspring varied significantly among crosses within parental combinations ( specific environmental maternal effects ). In the glasshouse experiment, the influence of environmental heterogeneity and competition could be kept low. Except for early characters, the influence of direct genetic variation was large but again leaf characters (= basic module morphology) seemed to be under stricter genetic control than did size characters. Genetic maternal effects, general environmental maternal effects, and specific environmental maternal effects dominated in early characters. The maternal effects were exerted both via seed mass and directly on characters of young offspring. Persistent effects of the general paternal environment ( general environmental paternal effects ) were found for leaf length and stem and leaf mass at harvest. They were opposite in direction to the general environmental maternal effects, that is the same genotypes produced “better mothers” in sand but “better fathers” in soil. The general environmental paternal effects must have been due to differences in pollen quality, resulting from pollen selection within the male parent or leading to pre- or postzygotic selection within the female parent. The ranking of crosses according to mean offspring phenotypes was different in the two experiments, suggesting strong interaction of the observed effects with the environment. The correlation structure among characters changed less between experiments than did the pattern of variation of single characters, but under the competitive conditions in the garden plant height seemed to be more directly related to fitness than in the glasshouse. Reduced competition could also explain why maternal effects were less persistent in the glasshouse than in the garden experiment. Evolution via selection of maternal effects would be possible in the study population because these effects are in part due to genetic differences among parents.     

MATERNAL INHERITANCE AND ITS EFFECT ON ADAPTIVE EVOLUTION: A QUANTITATIVE GENETIC ANALYSIS OF MATERNAL EFFECTS IN A NATURAL PLANT POPULATION

A mother can influence a trait in her offspring both by the genes she transmits (Mendelian inheritance) and by maternal attributes that directly affect that trait in her offspring (maternal inheritance). Maternal inheritance can alter the direction, rate, and duration of adaptive evolution from standard Mendelian models and its impact on adaptive evolution is virtually unexplored in natural populations. In a hierarchical quantitative genetic analysis to determine the magnitude and structure of maternal inheritance in the winter annual plant, Collinsia verna, I consider three potential models of inheritance. These range from a standard Mendelian model estimating only direct (i.e., Mendelian) additive and environmental variance components to a maternal inheritance model estimating six additive and environmental variance components: direct additive and environmental variances; maternal additive and environmental variances; and the direct-maternal additive () and environmental covariances. The structure of maternal inheritance differs among the 10 traits considered at four stages in the life cycle. Early in the life cycle, seed weight and embryo weight display substantial , a negative , and a positive . Subsequently, cotyledon diameter displays and of roughly the same magnitude and negative . For fall rosettes, leaf number and length are best described by a Mendelian model. In the spring, leaf length displays maternal inheritance with significant and and a negative . All maternally inherited traits show significant negative . Predicted response to selection under maternal inheritance depends on and as well as . Negative results in predicted responses in the opposite direction to selection for seed weight and embryo weight and predicted responses near zero for all subsequent maternally inherited traits. Maternal inheritance persists through the life cycle of this annual plant for a number of size-related traits and will alter the direction and rate of evolutionary response in this population.     

外源激素对风信子胚珠再生及其性细胞分化的影响

风信子（ＨｙａｃｉｎｔｈｕｓｏｒｉｅｎｔａｌｉｓＬ．）性器官分化和性细胞分化所需条件是不一样的,适合于性细胞（雌配子体）分化的外源激素水平大大低于性器官（胚珠）分化的外源激素水平。珠心细胞在低浓度的外源激素条件下可以分化性细胞;稍稍提高外源激素浓度,性细胞化化停止,代之以珠心细胞的增殖;进一步提高外源激素浓度,珠心细胞又可以分化出新的胚珠原基,进一步长大以后发育成许多新的胚珠