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1.
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.  相似文献   

2.
Theory predicts that inbreeding depression (ID) should decline via purging in self‐fertilizing populations. Yet, intraspecific comparisons between selfing and outcrossing populations are few and provide only mixed support for this key evolutionary process. We estimated ID for large‐flowered (LF), predominantly outcrossing vs. small‐flowered (SF), predominantly selfing populations of the dune endemic Camissoniopsis cheiranthifolia by comparing selfed and crossed progeny in glasshouse environments differing in soil moisture, and by comparing allozyme‐based estimates of the proportion of seeds selfed and inbreeding coefficient of mature plants. Based on lifetime measures of dry mass and flower production, ID was stronger in nine LF populations [mean δ = 1?(fitness of selfed seed/fitness of outcrossed seed) = 0.39] than 16 SF populations (mean δ = 0.03). However, predispersal ID during seed maturation was not stronger for LF populations, and ID was not more pronounced under simulated drought, a pervasive stress in sand dune habitat. Genetic estimates of δ were also higher for four LF (δ = 1.23) than five SF (δ = 0.66) populations; however, broad confidence intervals around these estimates overlapped. These results are consistent with purging, but selective interference among loci may be required to maintain strong ID in partially selfing LF populations, and trade‐offs between selfed and outcrossed fitness are likely required to maintain outcrossing in SF populations.  相似文献   

3.
We evaluated the degree of selfing and inbreeding depression at the seed and seedling stages of a threatened tropical canopy tree, Neobalanocarpus heimii, using microsatellite markers. Selection resulted in an overall decrease in the level of surviving selfed progeny from seeds to established seedlings, indicating inbreeding depression during seedling establishment. Mean seed mass of selfed progeny was lower than that of outcrossed progeny. Since the smaller seeds suffered a fitness disadvantage at germination in N. heimii, the reduced seed mass of selfed progeny would be one of the determinants of the observed inbreeding depression during seedling establishment. High selfing rates in some mother trees could be attributed to low local densities of reproductive individuals, thus maintenance of a sufficiently high density of mature N. heimii should facilitate regeneration and conservation of the species.  相似文献   

4.
Habitat fragmentation is extensive throughout the world, converting natural ecosystems into fragments of varying size, density and connectivity. The potential value of remnant trees in agricultural landscapes as seed sources and in connecting fragments has formed a fertile area of debate. This study contrasted the mating patterns of bat-pollinated Pachira quinata trees in a continuous forest to those in pasture through microsatellite-based paternity analysis of progeny. The breeding system was determined by analysis of pollen tube growth and seed production from controlled pollinations. Fitness of selfed and outcrossed seed was compared by germination and seedling growth. There was more inbreeding within pasture trees (outcrossing=0.828±0.015) compared with forest trees (0.926±0.005). Pasture trees had fewer sires contributing to mating events, but pollen dispersal distances were greater than those in the forest. Paternity analysis showed variation in outcrossing rates among pasture trees with high proportions of external and self pollen sources detected. A leaky self-incompatibility system was found, with self pollen having reduced germination on stigmas and slower growth rate through the style. Controlled pollinations also showed a varied ability to self among trees, which was reflected in the selfing rates among pasture trees shown by the paternity analysis (0–80% selfing). Self pollination resulted in lower seed set, germination and seedling growth compared with outcrossing. While remnant trees in agricultural landscapes are involved in broader mating patterns, they show increased but varied levels of inbreeding, which result in reduced fitness.  相似文献   

5.
Differential seed dispersal, in which selfed and outcrossed seeds possess different dispersal propensities, represents a potentially important individual‐level association. A variety of traits can mediate differential seed dispersal, including inflorescence and seed size variation. However, how natural selection shapes such associations is poorly known. Here, we developed theoretical models for the evolution of mating system and differential seed dispersal in metapopulations, incorporating heterogeneous pollination, dispersal cost, cost of outcrossing and environment‐dependent inbreeding depression. We considered three models. In the ‘fixed dispersal model’, only selfing rate is allowed to evolve. In the ‘fixed selfing model’, in which selfing is fixed but differential seed dispersal can evolve, we showed that natural selection favours a higher, equal or lower dispersal rate for selfed seeds to that for outcrossed seeds. However, in the ‘joint evolution model’, in which selfing and dispersal can evolve together, evolution necessarily leads to higher or equal dispersal rate for selfed seeds compared to that for outcrossed. Further comparison revealed that outcrossed seed dispersal is selected against by the evolution of mixed mating or selfing, whereas the evolution of selfed seed dispersal undergoes independent processes. We discuss the adaptive significance and constraints for mating system/dispersal association.  相似文献   

6.
We quantified inbreeding depression for fruit production, embryo vitality and seed germination in three deceptive orchids, Serapias vomeracea, S. cordigera and S. parviflora, which do not provide any reward to their pollinators, and are predicted to experience high outcrossing. Of the three species examined only S. parviflora was autonomously selfing. Both S. vomeracea and S. cordigera showed highly significant differences in fitness between selfed and outcrossed progenies, resulting in high levels of inbreeding depression, which increased in magnitude from seed set to seed germination. Inbreeding depression may promote outcrossing in Serapias by acting as a post-pollination barrier to selfing. Cumulative inbreeding depression across three stages in S. parviflora was lower that in both outcrossing species. The large difference in germination between selfed and outcrossed seeds is an important issue in conservation biology.  相似文献   

7.
The ecological and evolutionary implications of dispersal are many. Pollination type and maternal effects may affect plant fitness traits, including life-cycle traits as well as dispersal ability. This study investigated the joint influence of pollination type and maternal effects on both life-cycle traits and dispersal ability in the herb Hypochaeris radicata . We conducted experimental crosses to obtain selfed and outcrossed progeny. Individual seeds and their pappuses were measured to determine seed terminal velocity. Seed size was also used to assess maternal effects. Selfing dramatically decreased seed set, indicating that H. radicata is self-incompatible. However, the few selfed seeds produced outperformed outcrossed seeds in seed size and flowering probability, surely as a result of an effective reallocation of resources among selfed seeds. None of the life-cycle traits was affected by seed size, the estimate of maternal effects. Selfed seeds were larger and bore a smaller pappus than outcrossed seeds. As a result, dispersal ability was lower for selfed than outcrossed seeds. Several factors, such as the low proportion of plants that produced selfed seeds, the low number of selfed seeds produced per plant, and the lack of self-fertility mechanisms might act in concert to prevent the evolution of selfing in H. radicata .  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 163–170.  相似文献   

8.
Selfing or mating between related individuals in self-compatible hermaphroditic tree species may lead to inbreeding depression (ID) due to homozygosis in recessive, identical by descent alleles. In general, studies of ID in tree species have been based on comparisons of selfed individuals (produced by controlled pollination) with outcrossed individuals for quantitative traits in progeny tests. However, this approach requires a long time to quantify the extent of ID. Thus, we used an approach based on genetic markers to estimate coancestry coefficients between assigned parents from paternity analysis in two populations of the Neotropical tree Cariniana legalis. Using this method, we were able to determine which seedlings in a nursery trial originated from; (i) outcrossing between un-related trees, (ii) mating between related trees and (iii) selfing. We detected a low selfing rate (<10 %), but a substantial quantity of seedlings from mating between related parents (minimum of 35.7 %). In general, the outcrossed seedlings from unrelated parents exhibited significantly greater genetic diversity than those resulting from selfing and mating among relatives. The extent of ID varied among traits and populations. Outcrossed seedlings originating from unrelated trees generally showed greater survival than seedlings originating from selfing and related parents. Inbreeding depression was greater in the selfed seedlings than in those from mating among related parents. The results are discussed in terms of implications for genetic conservation, breeding and environmental restoration using the species.  相似文献   

9.
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.  相似文献   

10.
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.  相似文献   

11.
Many plants display limited seed dispersal, thereby creating an opportunity for sibling competition, i.e. fitness-determined interactions between related individuals. Here I investigated the consequences of intra-specific competition, by varying density and genetic composition of neighbors, on the performance of seedlings derived by selfing or outcrossing of the partially self-fertilizing plant Plantago coronopus (L.). Seedlings from eight plants, randomly selected from an area of about 50 m2 in a natural population, were used in (i) a density series with either one, four or eight siblings of each cross type per pot and (ii) a replacement series with eight plants per pot where selfed and outcrossed siblings were grown intermixed in varying frequencies. Density had a pronounced effect on plant performance. But, except for singly grown individuals, no differences were detected between selfed and outcrossed progenies in vegetative and reproductive biomass. When grown intermixed, selfed offspring were always inferior to their outcrossed relatives. The magnitude of reduction in performance was dependent on the number of outcrossed relatives a selfed seedling had to compete with, giving rise to a frequency-dependent fitness advantage to outcrossed seedlings. The major result of this study is (i) that the relative fitness of inbred progeny is strongly affected by the type of competitors (inbred or outbred) and (ii) that inbreeding depression varies according to the density and frequency of outbred plants and could be considered as a density- and frequency-dependent phenomenon. It is argued that sibling competition, due to the small genetic neighborhood of P. coronopus, might be an important selective force in natural populations of this species.  相似文献   

12.
Classical theory on mating system evolution suggests that simultaneous hermaphrodites should either outcross if they have high inbreeding depression (ID) or self‐fertilize if they have low ID. However, a mixture of selfing and outcrossing persists in many species. Previous studies with the tapeworm Schistocephalus solidus have found worms to self‐fertilize some of their eggs despite ID. The probability for selfing to spread depends on the relative fitness of selfers, as well as the genetic basis for ID and whether it can be effectively purged. We bred S. solidus through two consecutive generations of selfing and recorded several fitness correlates over the whole life cycle. After one round of selfing, ID was pronounced, particularly in early‐life traits, and the conservatively estimated lifetime fitness of selfed progeny was only 9% that of the outcrossed controls. After a second generation of selfing, ID remained high but was significantly reduced in several traits, which is consistent with the purging of deleterious recessive alleles (the estimated load of lethal equivalents dropped by 48%). Severe ID, even if it can be rapidly purged, likely prevents transitions toward pure selfing in this parasite, although we also cannot exclude the possibility that low‐level selfing has undetected benefits.  相似文献   

13.
Hermaphroditic individuals can produce both selfed and outcrossed progeny, termed mixed mating. General theory predicts that mixed-mating populations should evolve quickly toward high rates of selfing, driven by rapid purging of genetic load and loss of inbreeding depression (ID), but the substantial number of mixed-mating species observed in nature calls this prediction into question. Lower average ID reported for selfing than for outcrossing populations is consistent with purging and suggests that mixed-mating taxa in evolutionary transition will have intermediate ID. We compared the magnitude of ID from published estimates for highly selfing (r > 0.8), mixed-mating (0.2 ≤ r ≥ 0.8), and highly outcrossing (r < 0.2) plant populations across 58 species. We found that mixed-mating and outcrossing taxa have equally high average lifetime ID (δ= 0.58 and 0.54, respectively) and similar ID at each of four life-cycle stages. These results are not consistent with evolution toward selfing in most mixed-mating taxa. We suggest that prevention of purging by selective interference could explain stable mixed mating in many natural populations. We identify critical gaps in the empirical data on ID and outline key approaches to filling them.  相似文献   

14.
This study documents the comparative success of seeds and seedlings of the perennial gynodioecious-gynomonoecious weed, Silene vulgaris, in the greenhouse. The importance of experimental design is stressed by comparing two different statistical analyses of the data. Seeds were obtained from artificial pollinations in the field: self-fertilization of hermaphrodites, and cross-fertilizations of both hermaphrodites and females. One-way analysis of variance using progeny type (selfed hermaphrodites, outcrossed hermaphrodites, and outcrossed females) as the treatment effect for each seed and seedling variable showed statistically discernable differences among progeny from different cross types. The significance of this type of ANOVA resulted from a reduced error term and positively-biased F ratios. A factorial design showed no significant differences due to type of progeny in seed mass, days to germination, leaf number, area, or biomass at six weeks of age. There were, however, significant differences among seeds attributable to maternal parent for all seed and seedling variables. A higher proportion of seeds from outcrossed individuals germinated compared to that from self-fertilization. When the data were analyzed to include and partition all sources of variation, differences among offspring appeared during germination, rather than during later development. Seed mass, cross type, and sex of maternal parent all significantly affected the likelihood of germination; however, they had decreasing predictive power, respectively. Inbreeding depression in Silene vulgaris may help maintain gynodioecy; however, the pleiotropic effects of both nuclear and cytoplasmic genes for sex expression also may affect fitness and the maintenance of females.  相似文献   

15.
In rare plants that often occur in small or isolated populations the probability of selfing between close relatives is increased as a consequence of demographic stochasticity. The mode of pollination (selfing, outcrossing) may have considerable effects on seed traits and offspring performance and hence potential viability. Since current efforts aiming at the restoration of floodplain grasslands through the transfer of plant material from species-rich source stands may lead to the establishment of initially small populations consisting of founders from different populations, the present paper experimentally investigated the effects of pollen source and floral types (i.e. chasmogamous (CH) and cleistogamous (CL) flowers) on seed traits and offspring performance in three highly endangered violet species (Viola elatior, V. pumila, V. stagnina) of these grasslands. We estimated inbreeding depression and tested the performance of selfed and outcrossed offspring in two microbial environments, i.e. in soil inoculated with (i) non-sterile substrate from the same species (‘home’-conditions) and (ii) sterilised substrate.Plants produced more CL capsules than CH flowers. Pollinator exclusion had only small effects on CH seed production. CL seeds had a significantly lower mass per seed than CH seeds. This may be related to constraints in allocation or environmental conditions. Seedling growth was reduced in plants grown under ‘home’-conditions as compared to control soils. Under ‘home’-conditions, relative fitness of selfed seedlings of V. stagnina was significantly higher than that of crossed progeny. Our results suggest that high genetic differentiation among populations as a consequence of isolation may result in outbreeding depression, e.g., through biochemical or physiological incompatibilities between genes or the breaking of coadapted gene complexes. In V. stagnina, offspring fitness differed considerably between environments, but in general we found no indications for inbreeding depression in these rare species.  相似文献   

16.
Understanding the relative magnitudes of inbreeding and outbreeding depression in rare plant populations is increasingly important for effective management strategies. There may be positive and negative effects of crossing individuals in fragmented populations. Conservation strategies may include introducing new genetic material into rare plant populations, which may be beneficial or detrimental based on whether hybrid offspring are of increased or decreased quality. Thus, it is important to determine the effects of pollen source on offspring fitness in rare plants. We established pollen crosses (i.e. geitonogamous‐self, autonomous‐self, intrasite‐outcross, intersite‐outcross and open‐pollinated controls) to determine the effects of pollen source on fitness (seeds/fruit and seed mass) and early offspring traits (probability of germination, number of leaves, leaf area and seedling height) in the rare plant Polemonium vanbruntiae. Open‐pollinated, intrasite‐outcross and geitonogamous‐self treatments did not differ in fitness. However, plants receiving autonomous‐self pollen had the lowest fitness and the lowest probability of seed germination. Intersite‐outcross plants contained fewer seeds/fruit, but seeds germinated at higher frequencies and seedlings were more vigorous. We also detected heterosis at the seed germination stage. These data may imply that natural populations of P. vanbruntiae exhibit low genetic variation and little gene flow. Evidence suggests that deleterious alleles were not responsible for reduced germination; rather environmental factors, dichogamy, herkogamy and/or lack of competition among pollen grains may have caused low germinability in selfed offspring. Although self‐pollination may provide some reproductive assurance in P. vanbruntiae, the result is a reduction in germination and size‐related early traits for selfed offspring.  相似文献   

17.
Plant mating systems are known to vary within species and some immediate ecological factors have been found to be associated with the geographic distribution of selfing. The environmental condition of the maternal plant may influence the production of selfed seed relative to outcrossed seed. This study investigated the effect of late pollination on the mating system of Kalmia latifolia, a long-lived perennial shrub. A 2 × 2 experimental design was used to determine whether reproductive success declines over the course of the flowering season and whether there was an interaction between pollination time (early vs. late in the season) and pollen type (self-fertilized vs. outcrossed). An interaction of this sort would indicate context-dependent fitness of selfed seeds compared to outcrossed seeds and, thus, show an ecological influence over a plant's mating system. Relative fitness was assessed in terms of female reproductive success. Timing of pollination did not affect abortion of outcrossed seeds; however, delay in pollination increased abortion of selfed seeds by 34.7%. Thus, it appears that plants selectively aborted selfed seeds rather than outcrossed seeds and this selection was more intense at the end of the season. An ecological factor such as time of pollination may affect the mating system of K. latifolia.  相似文献   

18.
The ornithophilous species Anagyris foetida L. is a Mediterranean shrub with highly fragmented populations and a mixed mating system. In a previous study, we analyzed the first 3 years of the life cycle of two progenies (selfed and outcrossed) grown from seed obtained by hand pollination and planted in an experimental garden in 2005. In that study, we found that inbreeding depression (ID) was manifested both reproductively and vegetatively throughout the life cycle, with male reproductive function being the most affected trait. In the present study, our main aim was to check the progression of the two progenies 12 years after transplantation. For this we analyzed their survival and their vegetative and reproductive traits. According to our results, levels of ID were similar to those obtained in the previous study, with some factors decreased and thus varying with the age of the studied individuals. Vegetative parameters were found to have a greater influence than reproductive ones (δ = 0.56 vs. δ = 0.36) on overall ID. As indicated by the global ID (δ = 0.72) the populations have a mating system that is intermediate between outcrossing and a mixed system. Furthermore, the lower male reproductive capacity of the selfed individuals has been maintained over time. Our study also demonstrates the importance of studying the ID value throughout the life cycle of plants.  相似文献   

19.
How climate-change induced environmental stress may alter the effects of inbreeding in patchy populations of rare species is poorly understood. We investigated the fitness of progeny from experimental self- and cross-pollinations in eight populations of different size of Echium wildpretii, a rare endemic plant of the arid subalpine zone of the Canarian island of Tenerife. As control treatments we used open pollination and autonomous selfing. The seed set of open-pollinated flowers was 55% higher than that of autonomously selfed flowers, showing the importance of animal pollination for reproductive success. The seed set, seed mass and germination rate of seedlings of hand-selfed flowers was similar to that of hand-crossed flowers, indicating weak inbreeding depression (seed set –4.4%, seed mass –4.1%, germination –7.3%). Similarly, under normal watering there were no significant effects of inbreeding on seedling survival (–3.0%). However, under low watering of seedlings inbreeding depression was high (survival –50.2%). Seed set of open- and hand-outcrossed-pollinated flowers was higher in large than in small populations, possibly due to more frequent biparental inbreeding in the latter. However, later measures of progeny fitness were not significantly influenced by population size. We predict that increasing drought duration and frequency due to climate change and reductions of population sizes may increase inbreeding depression in this charismatic plant species and thus threaten its future survival in the longer term.  相似文献   

20.
Abstract: Inbreeding depression can decrease several fitness traits and maternal effects can strongly influence the amount of inbreeding depression. Understanding the effects of inbreeding depression on plant fitness is especially important in the context of habitat fragmentation, where plant populations become smaller and more isolated, exhibiting increasing levels of inbreeding depression. We examined the joint influence of inbreeding depression and maternal effects on life cycle traits and dispersal ability in the herb Tragopogon pratensis that grows in fragmented populations in Europe. We conducted experimental crosses to obtain selfed and outcrossed progeny in two contrasted environments. In particular, we produced a first generation of seeds and plants that were self-pollinated again to produce a second generation of seeds. Individual seeds were weighed and their pappuses measured to estimate the dispersal potential. Pollination treatment only had a significant effect on seed mass and dispersal ability. Coefficients of inbreeding depression did not differ between selfed and outcrossed plants. Seed mass had a significant effect on germination date. Environment had a significant effect on mass of the second generation of seeds and the interaction between pollination treatment and family was significant for six traits, indicating the existence of strong maternal effects in T. pratensis. Results suggest population differentiation. Overall, T. pratensis populations exhibited a good performance under selfing, in terms of life cycle traits and dispersal ability, which would allow the species to cope with problems associated with fragmentation.  相似文献   

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