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1.
Evolution to reduce inbreeding can favor disassortative (intermorph) over assortative (intramorph) mating in hermaphroditic sexually polymorphic plant species. Heterostyly enhances disassortative pollination through reciprocal placement of stigmas and anthers of morphs and appropriate pollinators. Stylar dimorphism in which there is not reciprocal anther placement may compromise disassortative mating, particularly when there is not intramorph incompatibility. Variable rates of disassortative mating along with differential female fecundity or siring success among floral morphs could lead to variation in morph ratio. We investigated mating patterns, female fecundity, and siring success of style‐length morphs in Narcissus papyraceus, a self‐incompatible but morph‐compatible species with dimorphic (long‐ and short‐styled) and monomorphic (long‐styled) populations in central and north regions of its range, respectively. We established experimental populations in both regions and exposed them to ambient pollinators. Using paternity analysis, we found similar siring success of morphs and high disassortative mating in most populations. Female fecundity of morphs was similar in all populations. Although these results could not completely explain the loss of dimorphism in the species’ northern range, they provided evidence for the evolutionary stability of stylar dimorphism in N. papyraceus in at least some populations. Our findings support the hypothesis that prevailing intermorph mating is key for the maintenance of stylar dimorphism.  相似文献   

2.
Heterodichogamy is a form of sex expression in which protandrous and protogynous individuals coexist, and is considered to be a mechanism that avoids selfing and promotes disassortative mating. We examined mating patterns in a heterodichogamous maple, Acer mono, using microsatellite markers. Parentage analysis revealed a selfing rate of only 9.8%. Disassortative mating between flowering types significantly exceeded within-type mating, but the mating patterns were better explained by flowering phenology (i.e., the temporal overlap between the female and male stages). Heterodichogamy in A. mono thus appears to promote outcrossing without requiring obligate self- or cross-incompatibility systems, although it did not guarantee disassortative mating. Multiple-regression analysis suggested that successful reproduction of pollen parents significantly increased with increased flower production and reciprocal flowering synchrony, but decreased only marginally with mating distance, although the distribution of mating distances suggested leptokurtic dispersal of pollen.  相似文献   

3.
Heterodichogamy, including protandrous (PA) and protogynous (PG) morphs, is considered a mechanism to avoid selfing and promote disassortative mating. Although morphotypes are usually present in a population at a 1:1 ratio, this ratio may be biased in a low-density population by demographic stochasticity, resulting in a deficiency of mating partners in a neighbourhood dominated by a single morph. In this study, we determined morph ratio of the heterodichogamous tree species, Juglans ailantifolia by observing flowering in a low-density population during 2?years. The morph ratio (PG: PA) of 2.56:1 deviated significantly deviated from 1:1. We genotyped 59 reproductive trees and 405 offspring derived from eight PG-mother and three PA-mother trees with 11 microsatellite markers. Paternity analysis was conducted to clarify the effects of mother morph on the proportion of intra-morph mating. Then, we applied the Bayesian mixed effect mating model (MEMM) to clarify mating system, pollen dispersal, and individual fecundity of PG- and PA-mother trees. We found that the selfing rate and the distance of pollen dispersal were not clearly different between PG- and PA-mother trees. In contrast, the proportion of intra-morph mating was higher in the majority-morph (PG) mother trees than in the minor-morph (PA) mother trees. The MEMM indicated that mean dispersal distance of PG-mother trees was larger than that of PA-mother trees with large variance. Furthermore, we observed individuals with unusually high intra-morph fecundity for majority-morph (PG) trees. These findings indicate that intra-morph mating may occur when majority-morph mothers suffer a deficiency of potential inter-morph mates.  相似文献   

4.
BACKGROUND AND AIMS: Heterodichogamy differs from normal dichogamy, in that it involves two mating types (protogyny and protandry) that occur at a 1 : 1 ratio in a population. Flowering phases of the two mating types are synchronized and reciprocal, which was considered to ensure between-type outcrossing. This study aims to quantify the flowering pattern and pollination efficacy in Juglans mandshurica, a wind-pollinated heterodichogamous tree. METHODS: The pattern of flowering phenology was monitored within individual trees and pollen traps were used to measure air-borne pollen loads during the spring in 2003 and 2004. Pollen longevity was determined by staining technique. Also a pollen supplementation experiment was performed in 2004 to assess pollen limitation of fruit production. KEY RESULTS: There was no overlap between sexual functions within individual trees. Flowering periods of the two mating types were reciprocal and synchronous in both 2003 and 2004. Air-borne pollen loads were large, and protogynous and protandrous individuals each produced a high pollination peak, consistent with the two blooming periods. Maximum pollen longevity was about 4 h for protandrous individuals, and 3 h for protogynous individuals. Pollen supplementation did not increase fruit production in either protogynous or protandrous individuals. CONCLUSIONS: Heterodichogamous flowering in Juglans mandshurica effectively avoids selfing, promotes between-type outcrossing, and leads to efficient pollination in a natural population.  相似文献   

5.
Ward M  Dick CW  Gribel R  Lowe AJ 《Heredity》2005,95(4):246-254
Despite the typically low population densities and animal-mediated pollination of tropical forest trees, outcrossing and long-distance pollen dispersal are the norm. We reviewed the genetic literature on mating systems and pollen dispersal for neotropical trees to identify the ecological and phylogenetic correlates. The 36 studies surveyed found >90% outcrossed mating for 45 hermaphroditic or monoecious species. Self-fertilization rates varied inversely with population density and showed phylogenetic and geographic trends. The few direct measures of pollen flow (N=11 studies) suggest that pollen dispersal is widespread among low-density tropical trees, ranging from a mean of 200 m to over 19 km for species pollinated by small insects or bats. Future research needs to examine (1) the effect of inbreeding depression on observed outcrossing rates, (2) pollen dispersal in a wide range of pollination syndromes and ecological classes, (3) and the range of variation of mating system expression at different hierarchical levels, including individual, seasonal, population, ecological, landscape and range wide.  相似文献   

6.
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   

7.
In sexually polymorphic species, the morphs are maintained by frequency-dependent selection through disassortative mating. In heterodichogamous populations in which disassortative mating occurs between the protandrous and protogynous morphs, a decrease in female fitness in one morph is hypothesized to drive sexual specialization in the other morph, resulting in dimorphic populations. We test these ideas in a population of the heterodichogamous species, Acer opalus . We assessed both prospective gender of individuals in terms of their allocations and actual parentage using microsatellites; we found that most matings in A. opalus occur disassortatively. We demonstrate that the protogynous morph is maintained by frequency-dependent selection, but that maintenance of males versus protandrous individuals depends on their relative siring success, which changes yearly. Seeds produced later in the reproductive season were smaller than those produced earlier; this should compromise reproduction through ovules in protandrous individuals, rendering them male biased in gender. Time-dependent gender and paternity analyses indicate that the sexual morphs are specialized in their earlier sexual functions, mediated by the seasonal decrease in seed size. Our results confirm that mating patterns are context-dependent and change seasonally, suggesting that sexual specialization can be driven by seasonal effects on fitness gained through one of the two sexual functions.  相似文献   

8.
We examined differences in pollen dispersal efficiency between 2 years in terms of both spatial dispersal range and genetic relatedness of pollen in a tropical emergent tree, Dipterocarpus tempehes. The species was pollinated by the giant honeybee (Apis dorsata) in a year of intensive community-level mass-flowering or general flowering (1996), but by several species of moths in a year of less-intensive general flowering (1998). We carried out paternity analysis based on six DNA microsatellite markers on a total of 277 mature trees forming four spatially distinct subpopulations in a 70 ha area, and 147 and 188 2-year-old seedlings originating from seeds produced in 1996 and 1998 (cohorts 96 and 98, respectively). Outcrossing rates (0.93 and 0.96 for cohorts 96 and 98, respectively) did not differ between years. Mean dispersal distances (222 and 192 m) were not significantly different between the 2 years but marginally more biased to long distance in 1996. The mean relatedness among cross-pollinated seedlings sharing the same mothers in cohort 96 was lower than that in cohort 98. This can be attributed to the two facts that the proportion of intersubpopulations pollen flow among cross-pollination events was marginally higher in cohort 96 (44%) than in cohort 98 (33%), and that mature trees within the same subpopulations are genetically more related to each other than those between different subpopulations. We conclude that D. tempehes maintained effective pollen dispersal in terms of outcrossing rate and pollen dispersal distance in spite of the large difference in foraging characteristics between two types of pollinators. In terms of pollen relatedness, however, a slight difference was suggested between years in the level of biparental inbreeding.  相似文献   

9.
Heterodichogamy is defined as the presence of two flower morphs that exhibit the male and female functions at different times among individuals within a population. Heterodichogamy is regarded as an adaptation to promote outcrossing through enhanced inter-morph mating, together with a 1:1 morph ratio. However, in highly fragmented populations, the morph ratio may be more likely to be biased by stochastic events. In such a situation, individuals of a minority morph within a population are expected to have higher reproductive success than those of a majority morph, which may suffer from pollen shortages of the minority morph. In this paper, we evaluated mating patterns and male reproductive success in a highly fragmented population of Machilus thunbergii, a putative heterodichogamous evergreen laurel tree. Results of paternity analysis indicated that the selfing rate was not clearly different between the two morphs. In contrast, the proportion of intra-morph mating was higher in the majority-morph (MM) mother trees than in the minority-morph (MF) mother trees. Bayesian estimated male reproductive success indicated that male reproductive success was higher in minority-morph (MF) than in majority-morph (MM) mother trees. These findings indicate that (1) the majority morph mothers, suffering a shortage of the opposite morph pollen, could partly compensate for the reduced reproductive success by intra-morph mating rather than by selfing, and (2) negative-frequency dependent selection may be involved in the maintenance of the two morphs.  相似文献   

10.
The evolutionary dynamics of neutral alleles under the Wright-Fisher model are well understood. Similarly, the effect of population turnover on neutral genetic diversity in a metapopulation has attracted recent attention in theoretical studies. Here we present the results of computer simulations of a simple model that considers the effects of finite population size and metapopulation dynamics on a mating-system polymorphism involving selfing and outcrossing morphs. The details of the model are based on empirical data from dimorphic populations of the annual plant Eichhornia paniculata, but the results are also of relevance to species with density-dependent selfing rates in general. In our model, the prior selfing rate is determined by two alleles segregating at a single diploid locus. After prior selfing occurs, some remaining ovules are selfed through competing self-fertilisation in finite populations as a result of random mating among gametes. Fitness differences between the mating-system morphs were determined by inbreeding depression and pollen discounting in a context-dependent manner. Simulation results showed evidence of frequency dependence in the action of pollen discounting and inbreeding depression in finite populations. In particular, as a result of selfing in outcrossers through random mating among gametes, selfers experienced a "fixation bias" through drift, even when the mating-system locus was selectively neutral. In a metapopulation, high colony turnover generally favoured the fixation of the outcrossing morph, because inbreeding depression reduced opportunities for colony establishment by selfers through seed dispersal. Our results thus demonstrate that population size and metapopulation processes can lead to evolutionary dynamics involving pollen and seed dispersal that are not predicted for large populations with stable demography.  相似文献   

11.
Gender polymorphism, plant-animal interactions, and environmental heterogeneity are the three important sources of variation in mating system and pollen dispersal patterns. We used progeny arrays and paternity analysis to assess the effects of gender type and density level on variation in mating patterns within a highly isolated population of Prunus mahaleb, a gynodioecious species. All the adult trees in the population were sampled and located. The direct estimate of long-distance insect-mediated pollination events was low (< 10%). Gender expression deeply influenced the mating system, decreasing the outcrossing rates (t(m)) and the pollen pool diversity in hermaphrodite trees. Long intermate distances (> 250 m) were significantly more frequent among female mother trees. Variation in local tree density also affected pollen pool diversity and intermate distance, with a higher effective number of fathers (k(e)) and longer intermate distances for female trees in low-density patches. A canonical correlation analysis showed significant correlations between mating variables and the maternal ecological neighbourhood. Only the first canonical variable was significant and explained 78% of variation. Outcrossing rates tended to decrease, and the relatedness among the fathers tended to increase, when mother trees grew in dense patches with high cover of other woody species and taller vegetation away from the pine forest edge. We highlight the relevance of considering maternal ecological neighbourhood effects on mating system and gene flow studies as maternal trees act simultaneously as receptors of pollen and as sources of the seeds to be dispersed.  相似文献   

12.
Hymenaea stigonocarpa is a neotropical tree that is economically important due to its high‐quality wood; however, because it has been exploited extensively, it is currently considered threatened. Microsatellite loci were used to investigate the pollen and seed dispersal, mating patterns, spatial genetic structure (SGS), genetic diversity, and inbreeding depression in H. stigonocarpa adults, juveniles, and open‐pollinated seeds, which were sampled from isolated trees in a pasture and trees within a forest fragment in the Brazilian savannah. We found that the species presented a mixed mating system, with population and individual variations in the outcrossing rate (0.53–1.0). The studied populations were not genetically isolated due to pollen and seed flow between the studied populations and between the populations and individuals located outside of the study area. Pollen and seed dispersal occurred over long distances (>8 km); however, the dispersal patterns were isolated by distance, with a high frequency of mating occurring between near‐neighbor trees and seeds dispersed near the parent trees. The correlated mating for individual seed trees was higher within than among fruits, indicating that fruits present a high proportion of full‐sibs. Genetic diversity and SGS were similar among the populations, but offspring showed evidence of inbreeding, mainly originating from mating among related trees, which suggests inbreeding depression between the seed and adult stages. Selfing resulted in a higher inbreeding depression than mating among relatives, as assessed through survival and height. As the populations are not genetically isolated, both are important targets for in situ conservation to maintain their genetic diversity; for ex situ conservation, seeds can be collected from at least 78 trees in both populations separated by at least 250 m.  相似文献   

13.
14.
  • Variation in flowering phenology is common in natural populations, and is expected to be, together with inter‐mate distance, an important driver of effective pollen dispersal. In populations composed of plants with temporally separated sexual phases (i.e. dichogamous or heterodichogamous populations), pollen‐mediated gene flow is assumed to reflect phenological overlap between complementary sexual phases. In this study, we conducted paternity analyses to test this hypothesis in the temporally dimorphic tree Acer opalus.
  • We performed spatially explicit analyses based on categorical and fractional paternity assignment, and included tree size, pair‐wise genetic relatedness and morph type as additional predictors. Because differences between morphs in flowering phenology may also influence pollination distances, we modelled separate pollen dispersal kernels for the two morphs.
  • Extended phenological overlap between male and female phases (mainly associated with inter‐morph crosses) resulted in higher siring success after accounting for the effects of genetic relatedness, morph type and tree size, while reduced phenological overlap (mainly associated with intra‐morph crosses) resulted in longer pollination distances achieved. Siring success also increased in larger trees.
  • Mating patterns could not be predicted by phenology alone. However, as heterogeneity in flowering phenology was the single morph‐specific predictor of siring success, it is expected to be key in maintaining the temporal dimorphism in A. opalus, by promoting not only a prevalent pattern of inter‐morph mating, but also long‐distance pollination resulting from intra‐morph mating events.
  相似文献   

15.
Vegetation clearing, land modification and agricultural intensification have impacted on many ecological communities around the world. Understanding how species respond to fragmentation and the scales over which functionality is retained, can be critical for managing biodiversity in agricultural landscapes. Allocasuarina verticillata (drooping sheoak, drooping she-oak) is a dioecious, wind-pollinated and -dispersed species with key conservation values across southeastern Australia. But vegetation clearing associated with agricultural expansion has reduced the abundance and spatial distribution of this species in many regions. Spatial genetic structure, relatedness among trees, pollen dispersal and mating patterns were examined in fragmented A. verticillata populations selected to represent the types of remnants that now characterise this species. Short scale spatial genetic structure (5–25 m) and relatedness among trees were observed in most populations. Unexpectedly, the two male trees closest to each female did not have a reproductive advantage accounting for only 4–15% of the seed produced in larger populations. Biparental inbreeding was also generally low (<4%) with limited evidence of seed crop domination by some male trees. More male trees contributed to seed crops in linear remnants (mean 17) compared to those from patch remnants (mean 11.3) which may reflect differences in pollen dispersal within the two remnant types. On average, pollen travels ~100 m irrespective of remnant type but was also detected to have dispersed as far as 1 km in open landscapes. Low biparental inbreeding, limited reproductive assurance for near-neighbour and probably related males and variability in the distances over which females sample pollen pools suggest that some mechanism to prevent matings between relatives exists in this species.  相似文献   

16.
DL Field  SC Barrett 《Molecular ecology》2012,21(15):3640-3643
Since Darwin's pioneering research on plant reproductive biology (e.g. Darwin 1877), understanding the mechanisms maintaining the diverse sexual strategies of plants has remained an important challenge for evolutionary biologists. In some species, populations are sexually polymorphic and contain two or more mating morphs (sex phenotypes). Differences in morphology or phenology among the morphs influence patterns of non-random mating. In these populations, negative frequency-dependent selection arising from disassortative (intermorph) mating is usually required for the evolutionary maintenance of sexual polymorphism, but few studies have demonstrated the required patterns of non-random mating. In the current issue of Molecular Ecology, Shang et al. (2012) make an important contribution to our understanding of how disassortative mating influences sex phenotype ratios in Acer pictum subsp. mono (painted maple), a heterodichogamous, deciduous tree of eastern China. They monitored sex expression in 97 adults and used paternity analysis of open-pollinated seed to examine disassortative mating among three sex phenotypes. Using a deterministic 'pollen transfer' model, Shang et al. present convincing evidence that differences in the degree of disassortative mating in progeny arrays of the sex phenotypes can explain their uneven frequencies in the adult population. This study provides a useful example of how the deployment of genetic markers, demographic monitoring and modelling can be integrated to investigate the maintenance of sexual diversity in plants.  相似文献   

17.
Mating systems have long been recognized as key factors determining genetic structure within and between populations. Outcrossing promotes genetic diversity and gene flow between populations, while inbreeding, on the other hand, decreases recombination rates, facilitating fixation of co-adapted genes. In small populations, selfing moderates pollen limitation because of low mate availability, but at the cost of increased inbreeding depression. These conflicts are of more than theoretical interest; they are critical for the management of endangered species. In order to help designing conservation strategies for the management of the gene pool of fragmented populations of Pinus cembra, a protected species in Poland, we have characterized pollen flow and mating structure using nuclear microsatellite markers. We demonstrated that P. cembra in the studied stands of the Tatra Mts. is characterized by an average outcrossing rate (t) of 0.72. Unlike with the existing approaches, using the newly developed Bayesian method, we found that population size and seasonal variation had confounding effects on outcrossing rates. In concordance with predictions, large populations showed significantly higher outcrossing rates (t?=?0.89) than smaller ones (t?=?0.51). Temporal variation revealed in the outcrossing rate might be linked with masting behavior of the species. On the other hand, we showed that outcrossing rates were not associated with a trunk diameter of a mother tree. Our study also demonstrated that biparental inbreeding is a significant component of mating system. However, we further show that pollen dispersal follows a fat-tailed distribution (with the average dispersal distance of 1,267 m) so that at least some long-distance pollen dispersal must be occurring. Overall, we conclude that the high inbreeding (both selfing and mating between relatives) found in P. cembra buffers for pollen limitation. We argue that small, isolated stands can be at risk of gene pool erosion, despite the potential for long-distance pollen and seed dispersal.  相似文献   

18.
Six microsatellite loci were used to compare the mating system and gene flow in two consecutive years of a natural, unlogged population of Symphonia globulifera in a 500 ha experimental plot in the Brazilian Amazon (Flona Tapajós). The species had a low density of reproductive trees per hectare (   d = 0.46  trees/ha). We analyzed 205 trees and 261 and 487 open-pollinated seeds from 26 and 30 mother-trees in the years 2002 and 2003, respectively. A significant spatial genetic structure was detected for the adult trees for distances up to 100 m. We observed only small interannual differences in multilocus outcrossing rate (     ,     ), biparental inbreeding (     ,     ), and paternity correlation (     ,     ). The number of pollen donors contributing to mating of each tree in both years was estimated to be low (     ). Using TwoGener analysis to calculate the density of reproductive trees and the distance of pollen dispersal for normal and exponential models, the lowest error was detected for exponential model. For this model, the estimated density of reproductive trees was lower in 2002 (     trees/ha) than 2003 (     trees/ha), resulting in a higher distance of pollen dispersal in 2002 (     m) than 2003 (     m), although these changes did not affect the outcrossing and correlated mating rates.  相似文献   

19.
While habitat fragmentation is a central issue in forest conservation studies in the face of broad-scale anthropogenic changes to the environment, its effects on contemporary mating patterns remain controversial. This is partly because of the inherent variation in mating patterns which may exist within species and the fact that few studies have replication at the landscape level. To study the effect of forest fragmentation on contemporary mating patterns, including effective pollen dispersal, we compared four native populations of the Australian forest tree, Eucalyptus globulus . We used six microsatellite markers to genotype 1289 open-pollinated offspring from paired fragmented and continuous populations on the island of Tasmania and in Victoria on mainland Australia. The mating patterns in the two continuous populations were similar, despite large differences in population density. In contrast, the two fragmented populations were variable and idiosyncratic in their mating patterns, particularly in their pollen dispersal kernels. The continuous populations showed relatively high outcrossing rates (86–89%) and low correlated paternity (0.03–0.06) compared with the fragmented populations (65–79% and 0.12–0.20 respectively). A greater proportion of trees contributed to reproduction in the fragmented ( de/d ≥ 0.5) compared with the continuous populations ( de/d  =   0.03–0.04). Despite significant inbreeding in the offspring of the fragmented populations, there was little evidence of loss of genetic diversity. It is argued that enhanced medium- and long-distance dispersal in fragmented landscapes may act to partly buffer the remnant populations from the negative effects of inbreeding and drift.  相似文献   

20.
Recent studies suggest that tropical tree species exhibit low inbreeding and high gene dispersal levels despite the typically low density of conspecifics in tropical forests. To examine this, we undertook a study of pollen gene dispersal and mating system of two Amazonian tree species. We analyzed 341 seeds from 33 trees at four microsatellite loci in a Carapa guianensis population from Brazil, and 212 seeds from 22 trees at four microsatellite loci in a Sextonia rubra population from French Guiana. Differentiation of allele frequencies among the pollen pool of individual trees was ΦFT= 0.053 (95% CI: 0.027–0.074) for C. guianensis and ΦFT= 0.064 (95% CI: 0.017–0.088) for S. rubra. The mean pollen dispersal distances were estimated at 69–355 m for C. guianensis , and 86–303 m for S. rubra , depending on the pollen dispersal model and the estimate of reproductive tree density used. The multi-locus outcrossing rate was estimated at 0.918 and 0.945, and the correlation of paternity at 0.089 and 0.096, for C. guianensis and S. rubra , respectively, while no significant levels of biparental inbreeding were detected. Comparing trees with high and low local density of conspecifics, we found no evidence for differences in inbreeding levels. The results are discussed within the framework of the emerging picture of the reproductive biology of tropical forest trees.  相似文献   

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