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1.
There has been disagreement concerning the nature of sexual polymorphisms in Narcissus, a genus of insect-pollinated geophytes native to the Mediterranean. The existence of trimorphic heterostyly in the genus has recently been confirmed, but the occurrence of distyly remains enigmatic. All sexually dimorphic species previously investigated possess two distinct style lengths but anthers of similar height. Stigma-height dimorphism does not qualify as true distyly because of the absence of a reciprocal correspondence in stigma and anther position in the floral morphs. Such reciprocal herkogamy is generally regarded as the defining feature of heterostyly. Here we report on distyly in N. albimarginatus (section Apodanthae), a rare species confined to a single mountain in northwestern Morocco. A population composed of equal numbers of long- and short-styled plants exhibited reciprocal herkogamy with lower anthers of the long-styled morph and upper anthers of the short-styled morph corresponding in height to stigmas of short- and long-styled plants, respectively. The presence of both stigma-height dimorphism and distyly in Narcissus is of general significance to theoretical models of the evolution of heterostyly.  相似文献   

2.
Here we analysed the role played by breeding systems and pollinators in the evolution of heterostyly by testing whether evolution towards heterostyly is associated with style polymorphism and changes in pollinator proficiency or breeding system variation (Darwinian hypothesis). We studied pollinators, pollen-transfer efficiency, and incompatibility systems in all seven species of Narcissus sect. Apodanthi for which we also obtained chloroplast DNA (cpDNA) sequences from three spacers to infer phylogenetic relationships. Five species are self-incompatible and within-morph cross-compatible. Heterostylous (Narcissus albimarginatus) and style-dimorphic (Narcissus cuatrecasasii) species that have a high degree of reciprocity in stigma and anther height are primarily pollinated by solitary bees. The style-monomorphic species (Narcissus watieri) and the style-dimorphic species with the least stigma-anther reciprocity (Narcissus rupicola) are both self-compatible and pollinated by butterflies, moths and hover flies. Phylogenetic reconstruction of character transitions indicates that the shift from style dimorphism to distyly is associated with a shift to bee pollination. Pollination by lepidopterans and flies is associated with stable style dimorphism and monomorphism. Evolution and maintenance of style polymorphisms in this group of species are independent of incompatibility systems. Taken together, our results strongly support the pollinator-based model for evolution of heterostyly and style length polymorphisms in general.  相似文献   

3.
We investigated the origin of stylar polymorphisms in Narcissus, which possesses a remarkable range of stylar conditions and diverse types of floral morphology and pollination biology. Reconstruction of evolutionary change was complicated by incomplete resolution of trees inferred from two rapidly evolving chloroplast regions, but we bracketed reconstructions expected on the fully resolved plastid-based tree by considering all possible resolutions of polytomies on the shortest trees. Stigma-height dimorphism likely arose on several occasions in Narcissus and persisted across multiple speciation events. As proposed in published models, this rare type of stylar polymorphism is ancestral to distyly. While there is no evidence in Narcissus that dimorphism preceded tristyly, a rapid transition between them may explain the lack of a phylogenetic footprint for this evolutionary sequence. The single instances of distyly and tristyly in Narcissus albimarginatus and N. triandrus, respectively, are clearly not homologous, an evolutionary convergence unique to Amaryllidaceae. Floral morphology was likely an important trigger for the evolution of stylar polymorphisms: Concentrated-changes tests indicate that a long, narrow floral tube may have been associated with the emergence of stigma-height dimorphism and that this type of tube, in combination with a deep corona, likely promoted, or at least was associated with, the parallel origins of heterostyly.  相似文献   

4.
There is growing appreciation that the ecological factors which impact on rates of pollen transfer can contribute significantly to reproductive trait evolution in plants. In heterostylous species, several studies support Darwin's claim that the reciprocal positions of stigmas and anthers enhance inter‐morph mating in comparison to intra‐morph mating and thus the maintenance of the polymorphism. In this study, we evaluate the relative importance of intra‐morph and inter‐morph pollen transfers in Narcissus assoanus, a species with dimorphic variation in style length but non‐reciprocity of anther positions. This stigma‐height dimorphism represents a transitional stage in theoretical models of the evolution of distyly. Seed set variation on recipient plants with donor plants of a single morph in experimental arrays in a natural population illustrate that inter‐morph cross‐pollination is more efficient that intra‐morph cross‐pollination as a result of high rates of pollen transfer from long‐styled to short‐styled plants. The observed rates of pollen transfer satisfy the theoretical conditions for the establishment of a stigma‐height dimorphism in an ancestral monomorphic long‐styled population in pollen‐limited situations. These results provide experimental evidence for the Darwinian hypothesis that enhanced inter‐morph cross‐pollination facilitates not only the maintenance of heterostyly but also the establishment of transitional forms implicated in the evolution of this polymorphism.  相似文献   

5.
异型花柱(heterostyly)是被子植物中一种特殊的花多态现象和雌雄异位形式,包括二型花柱(distyly)和三型花柱(tristyly)2种类型.据报道,在被子植物的约31个科中有异型花柱植物.该类型植物的花部特征在避免自交、促进准确的异交传粉以及通过降低雌雄功能干扰以提高亲本适合度等方面具有重要的进化意义.该文从以下3个方面总结和分析了异型花柱及其进化意义:(1)异型花柱植物的类型和花部特征、附属多态性和种群结构:(2)异型花柱植物在被子植物中的分布、起源和演化:(3)异型花柱植物的进化适应意义.结合目前作者开展的有关工作,对异型花柱植物研究中存在的一些问题进行讨论和展望,希望能为国内工作者开展该领域的研究提供一些参考.  相似文献   

6.
Heterostyly is a stylar polymorphism that has been shaped by the evolution of floral characters adapted for efficient pollen transfer. Different types of stylar polymorphism are described in which the discrete characterization of the exact polymorphic type (e.g., distyly vs. stigma-height dimorphism) requires detailed floral measurements (e.g., sex-organ reciprocity). In clonal and aquatic Nymphoides montana, although the presence of two floral morphs that contain styles of two lengths has been previously reported, no studies have quantitatively estimated the level of reciprocity and/or described the stylar condition. Morphological variations and incompatibility relationships were explored between the two morphs in three southeastern Australian populations. In this study, one population is characterized as stigma-height dimorphism (i.e., two morphs with discrete variation in stigma height but little variation in anther height), whereas the other two populations are typical distylous (i.e., two morphs for reciprocal stigma and anther height). Nymphoides montana is dimorphic in a wide range of ancillary characters, including corolla size, stigma size, shape and papillae morphology, and pollen size, number and exine sculpture. Following glasshouse pollinations, full incompatibility systems were observed in the distylous populations, whereas the stigma-height dimorphic population showed between-morph variation in the extent of incompatibility. Despite the variation in sex-organ reciprocity and incompatibility, other lines of evidence appear to assure the maintenance of the stylar polymorphism in the N. montana study populations. All populations are nearly isoplethic (i.e., both morphs in equal frequencies), which is indicative of balanced polymorphisms that appear to be maintained by legitimate pollen transfer between the morphs.  相似文献   

7.
周伟  王红 《植物学报》2009,44(6):742-751
异型花柱(heterostyly)是被子植物中一种特殊的花多态现象和雌雄异位形式, 包括二型花柱(distyly)和三型花柱(tristyly) 2种类型。据报道, 在被子植物的约31个科中有异型花柱植物。该类型植物的花部特征在避免自交、促进准确的异交传粉以及通过降低雌雄功能干扰以提高亲本适合度等方面具有重要的进化意义。该文从以下3个方面总结和分析了异型花柱及其进化意义: (1)异型花柱植物的类型和花部特征、附属多态性和种群结构; (2)异型花柱植物在被子植物中的分布、起源和演化; (3) 异型花柱植物的进化适应意义。结合目前作者开展的有关工作, 对异型花柱植物研究中存在的一些问题进行讨论和展望, 希望能为国内工作者开展该领域的研究提供一些参考。  相似文献   

8.
Narcissus, the daffodil genus, exhibits an unusual diversity of sexual systems, with populations that are monomorphic, dimorphic or trimorphic for style length. Associated with this variation are striking differences among species in floral morphology and pollination biology. This diversity provides an opportunity to investigate the evolution of mating polymorphisms, and to determine how floral morphology promotes transitions among sexual systems. Because of the absence of heteromorphic incompatibility in Narcissus, floral morphology plays a key role in governing patterns of outcrossed mating. Phylogenetic evidence indicates that stylar monomorphism is ancestral in the genus, with multiple origins of stylar polymorphism, including independent origins of stigma-height dimorphism, distyly and tristyly. Sexual polymorphisms have evolved only in lineages with narrow floral tubes that are pollinated by Lepidoptera and/or long-tongued bees. Populations of polymorphic Narcissus species are typically dominated by the long-styled morph and display imperfect reciprocity in the positions of sexual organs. These features are consequences of the unusual association between stylar polymorphism and a self-incompatibility system that permits intramorph mating.  相似文献   

9.
Darwin's early work on heterostyly and related style polymorphisms (the presence of two or three style morphs within a population) generated much interest to understand how precise interactions between ecological and genetic mechanisms influence the evolution of floral diversity. Here we tested three key hypotheses proposed to explain the evolution of heterostyly: (i) the presence of self‐incompatibility; (ii) the role of pollinators in promoting dissasortative mating; and (iii) floral architecture, which restricts pollinators’ movements and ensures more exact pollen deposition on their bodies. We combined data from experiments, field observations, and published studies to test whether evolution of style polymorphism in Narcissus is driven by the incompatibility system, pollinator guilds, or floral architecture, within a phylogenetic framework. Neither differences in pollinator environment nor the presence of genetic self‐incompatibility were correlated with presence of style polymorphism. However, our results indicate that the evolution of style polymorphism was driven by the presence of a narrow and long floral tube.  相似文献   

10.

Background and Aims

Heterostylous plants have been characterized by the presence of two or three discrete morphs that differ in their sex organ position within populations. This polymorphism is widely distributed among the angiosperms, but detailed studies are limited to few taxonomic groups. Although a small representation, evolutionary meaningful variations of the heterostylous syndrome have been reported when precise measurements of the sexual whorls were taken. A thorough exploration of groups where heterostyly has been reported should offer new opportunities to further testing the evolutionary hypotheses explaining heterostyly. Here, the traits defining heterostyly were explored in half of the species in Nivenia, the only genus of Iridiaceae where heterostyly has been reported.

Methods

Detailed morphometric analysis of the flower sexual whorls and some traits considered as ancillary are supplied to determine for each population (a) the kind of stylar polymorphism, (b) the morph ratio and (c) the degree of reciprocity between sexual whorls. Also the rates of assortative (within morph) versus disassortative (between morphs) pollen transfer were estimated by analysing pollen loads on stigmas. The association between floral phenotypic integration and the reciprocity between sexual whorls was estimated; both characteristics have been quoted as dependent on the accuracy of the fit between pollinators and flowers and therefore related to the efficiency of pollen transfer.

Key Results

Different types of polymorphism, differing in their degree of reciprocity, were found in Nivenia. Effective disassortative mating appears to be common, since (a) all dimorphic populations show equal morph-ratios (isoplethy), and (b) the pollen placed on the stigmas of each morph is likely to be coming from the other (complementary) morph. The most reciprocal populations of the heterostylous species have also the highest values of phenotypical integration.

Conclusions

Stigma height dimorphism, as opposed to distyly, is proven for the first time in Nivenia. The presence of different types of polymorphism within the genus is consistent with hypotheses of the evolution of heterostyly. The role of the pollinators as the leading force of the transition seems to be apparent, since floral integration is related to reciprocity.  相似文献   

11.
Heterostyly rarely occurs in families with strongly zygomorphic flowers. For this reason Darwin (1877) doubted whether heterostyly would occur in the Lamiaceae and recent reviews have not reported the floral polymorphism in this family. Here we describe distyly in a rare species ofSalvia restricted to bluffs and seaward canyons on Santa Rosa Island (Santa Barbara Co., California) and northwestern Baja California (Mexico).Salvia brandegeei is morphologically distylous with populations composed of equal frequencies of long-and short-styled morphs differing reciprocally in stigma and anther position. Controlled hand pollinations demonstrated no significant differences in the seed set of self, intramorph or intermorph pollinations. Unlike most heterostylous species investigated,S. brandegeei does not possess diallelic incompatibility or ancillary polymorphisms of pollen and stigmas. We propose that the evolution of distyly inS. brandegeei may have been associated with an ecological shift to a new environment in which protandry failed to prevent increased levels of geitonogamy. Heterostyly was then selected because it increased the proficiency of cross-pollination. The origin of distyly in self-compatibleS. brandegeei is consistent with Lloyd and Webb's theoretical model for the evolution of distyly.  相似文献   

12.
Distyly, a special polymorph, has evolved in many groups of angiosperms and has attracted attention since Darwin’s time. Development studies on distylous taxa have helped us to understand the evolutionary process of this polymorph, but most of these studies focus on species with narrowly tubular corolla. Here, we studied the floral development of Polygonum jucundum, a distylous species with broadly open flowers, at multiple spatial scales. Results showed that the difference in stigma height between flowers of the two morphs was caused by differences in style growth throughout the entire floral development process. The observed difference in anther heights between the two morphs was because the filaments grew faster in short-styled (SS) than in long-styled (LS) flowers in the later stages of floral development. In addition, the longer styles in LS flowers than in SS flowers was because of faster cell division in the early stages of floral development. However, SS flowers had longer filaments than LS flowers primarily because of greater cell elongation. These results indicate that floral development in P. jucundum differs from that of distylous taxa with floral tubes shown in previous studies. Further, we conclude that the presence of distyly in species with open flowers is a result of convergent evolution.  相似文献   

13.

Background and Aims

Although evolution of sexual polymorphism has been traditionally analysed using discrete characters, most of these polymorphisms are continuous. This is the case of heterostyly. Heterostyly is a floral polymorphism successfully used as a model to study the evolution of the sexual systems in plants. It involves the reciprocal positioning of anthers and stigmas in flowers of different plants within the same population. Studies of the functioning of heterostyly require the quantification of the degree of reciprocity between morphs of heterostylous species. Some reciprocity indices have been proposed previously, but they show significant limitations that need to be dealt with. This paper analyses these existing indices, and proposes a new index that aims to avoid their main problems (e.g. takes into account population variability and offers a single value per population).

Methods

The new index is based on the comparison of the position of every single sexual organ in the population with each and every organ of the opposite sex. To carry out all the calculations, a macro was programmed with MS®Visual Basic in MS® Excel. The behaviour of the index is tested using hypothetical data to simulate different situations of dimorphic populations; the index is also tested with some actual populations of different species of the genus Lithodora.

Results and Conclusions

The index of reciprocity proposed here is a sound alternative to previous indices: it compares stigma–stamen height gaps for all potential crosses in the population, it comprises stigma–stamen distance as well as dispersion, it is not skewed by the more frequent sex, and it can be meaningfully compared between populations and species. It has produced solid results for both hypothetical and natural populations.Key words: Distyly, floral polymorphism, heterostyly, reciprocity index, stylar dimorphism, Lithodora  相似文献   

14.
Knowledge of developmental pathways for achieving differences in style and anther heights, in concert with those of ancillary features accompanied with data in regard to biomass investment to male and female function, provide an excellent opportunity for examining the developmental correlations between primary and ancillary floral traits so as to understand the evolution of heterostyly. The ontogenetic relationships between bud length and anther height and between bud length and style height, and between bud length versus bud width, anther length, and number of pollen grains per anther for long-styled (LS) and short-styled (SS) morphs of P. PADIFOLIA are described. We also described the ontogenetic biomass allocation to male and female function and to corolla with elongation of buds harvested at regular intervals. We observed an early termination of stylar growth in SS buds, whereas LS styles steadily increased in size. Morph differences for relative growth rates were significant for anther height, anther length, and pollen number but not for bud width. Bud width and anther length had a negative allometric relationship with bud elongation. The relationship between bud length and number of pollen grains per anther was positive and morph differences in pollen number were detected at later stages of development. An increase in corolla mass involved a disproportionate allocation to the female function in SS flowers and male allocation was similar for the two morphs over the course of development. Our results are consistent with theoretical and empirical data for distylous species with an approach herkogamous ancestor, and with the more general hypothesis of ontogenetic lability of heterostyly, in which morph differences in style and anther heights are achieved in various ways. Variations observed in sexual investment between floral morphs suggest differences in sex expression during flower development.  相似文献   

15.
We studied a population of the distylousPalicourea padifolia (Rubiaceae) in a cloud forest remnant near Xalapa City, Veracruz, México to explore possible asymmetries between floral morphs in the attractiveness to pollinators, seed dispersers, nectar robbers, floral parasites, and herbivores. We first assessed heterostyly and reciprocal herkogamy by measuring floral attributes such as corolla length (buds and open flowers), style and anther heights, stigma and stamen lengths and the distance between the anther tip to the stigma lobe. We then estimated floral and fruit attributes such as flower size, anther height, number and size of pollen grains, fruit size, seed size, nectar production, and flower and fruit standing crops to assess differences between floral morphs in attracting and effectively using mutualistic pollinators and seed dispersers. Also, floral parasitism and nectar robbing were assessed in this study as a measure of flower attractiveness to antagonists. The system seems to conform well to classical heterostyly (e.g. reciprocal stamen/style lengths, pollen and anther dimorphism, intramorph incompatibility) yet, there were several tantalizing differences observed between pin and thrum morphs. Thrum flowers have longer corollas and larger but fewer pollen grains than pin flowers. Both morphs produced the same total number of inflorescences, developed the same number of buds, and opened the same number of flowers per inflorescence during the flowering season. Nectar production and sugar concentration were similar between floral morphs but the reward was not offered symmetrically to floral visitors throughout the day. Nectar concentration was higher in pin flowers in the afternoon. The numbers of developing, fully developed, and ripe fruits were the same between floral morphs, however, fruits and seeds were larger than those of thrums. The incidence of fly larvae was higher among thrum flowers and damage by nectar robbing was the same between floral morphs. Fruit abortion patterns of flowers manually pollinated suggest intra-morph sterility (self and intramorph incompatibility). There were no differences between morphs in fruit and seed set per flower following legitimate pollination although thrums were more leaky than the pins (intramorph compatibility).  相似文献   

16.
Background and Aims Distyly is a floral polymorphism characterized by the presence of two discrete morphs with reciprocal positioning of anthers and stigmas in flowers on different plants within the same population. Although reciprocal herkogamy and associated floral traits are generally thought to be discrete and strict polymorphisms, little is known about variation in floral traits related to the distylous syndrome within and among populations of a single species. In this study, variation in floral morphology and reciprocal positioning of the sexual organs in the distylous Primula veris (cowslip) is quantified.Methods Data were collected in ten populations occurring in two contrasting habitat types (grasslands and forests), and for each population the average level of reciprocity was assessed, the strength of the self-incompatibility system was determined, and seed production under natural conditions was quantified.Results In grassland populations, flowers showed clear distyly with low and symmetric reciprocity indices at both the lower and upper level. In forests, P. veris produced larger flowers that showed strong deviations in stigma–anther separation, especially in the L-morph. This deviation was mainly driven by variation in stigma height, resulting in high and asymmetric reciprocity indices and the occurrence of several short-styled homostylous plants. Self-incompatibility was, however, strict in both habitats, and morph ratios did not differ significantly from isoplethy. The observed shift in reciprocity in forest populations was associated with a significant reduction in seed production in the L-morph.Conclusions The results indicate that populations of P. veris show habitat-specific variation in flower morphology. Deviations from perfect reciprocal positioning of stigmas and anthers also translate into reduced seed production, suggesting that small changes in sexual organ reciprocity can have far-reaching ecological and evolutionary implications.  相似文献   

17.
A recent model by Lloyd and Webb derives conditions necessary for the evolution of distyly based on pollen-transfer probabilities between ancestral morphs and invading mutants. We used bumblebees visiting artificial flowers to measure the parameters of the model. Our findings supported the first evolutionary step proposed by the model, establishment of a stigma-height polymorphism. Conditions for the subsequent establishment of an anther-height polymorphism were not satisfied by pollen-transfer patterns alone. Because conditions for the first step are considered more onerous, however, and because the second stage depends on inbreeding depression as well as pollen-transfer patterns, we interpret our results as supporting the plausibility of the Lloyd-Webb model. Video images of bees visiting glass-sided artificial flowers demonstrate a mechanism for disassortative pollination between the ancestral and mutant morph. In general, pollen-transfer probabilities were negatively correlated with the height difference between anthers of the donor and stigma of the recipient. Style length affects bee feeding posture in such a way that disassortative pollination could feasibly occur in the absence of an anther-height polymorphism.  相似文献   

18.
  • Pollen and stigma size have the potential to influence male fitness of hermaphroditic plants, particularly in species presenting floral polymorphisms characterised by marked differences in these traits among floral morphs. In this study, we take advantage of the evolutionary transition from tristyly to distyly experienced by Oxalis alpina (Oxalidaceae), and examined whether modifications in the ancillary traits (pollen and stigma size) respond to allometric changes in other floral traits. Also, we tested whether these modifications are in accordance with what would be expected under the hypothesis that novel competitive scenarios (as in distylous‐derived reproductive system) exert morph‐ and whorl‐specific selective pressures to match the available stigmas.
  • We measure pollen and stigma size in five populations of O. alpina representing the tristyly–distyly transition.
  • A general reduction in pollen and stigma size occurred along the tristyly–distyly transition, and pollen size from the two anther levels within each morph converged to a similar size that was characterised by whorl‐specific changes (increases or decreases) in pollen size of different anthers in each floral type.
  • Overall, results from this study show that the evolution of distyly in this species is characterised not only by changes in sexual organ position and flower size, but also by morph‐specific changes in pollen and stigma size. This evidence supports the importance of selection on pollen and stigma size, which increase fitness of remaining morphs following the evolution of distyly, and raises questions to explore on the functional value of pollen size in heterostylous systems under pollen competition.
  相似文献   

19.
Heterostyly is a floral polymorphism that increase inter‐morph pollen transfer and promote disassortative mating. Breakdown of heterostyly has happened many times, either leading to dioecy or monomorphism. Mussaenda is a genus with diverse sexual systems including distyly, dioecy, floral monomorphism, and homostyly, making it an ideal system to study the evolution of floral traits and their relationship with sexual system shifts. Here, floral traits and pollen–ovule (P/O) ratios were estimated and used to test hypotheses about the relationships among P/O ratios, floral trait evolution, and sexual system shifts. Our results revealed that there is no significant difference of reciprocity in upper level sexual organs between species with different sexual systems. The reciprocity indices of the lower level sexual organs, however, are smaller in species with functional dioecy than in those with distyly. P/O ratios in dimorphic Mussaenda species were relatively lower than in monomorphic outcrossing species, but did not differ significantly between species with distyly and functional dioecy. Populational P/O ratios were negatively correlated with reciprocity index in the dimorphic species. We suggest that the loss of function in lower level organs in species with functional dioecy has resulted from less strict reciprocity. The relationship between P/O ratios and reciprocity indices strongly support that efficient disassortative pollination of distylus flowers may have promoted evolution towards low P/O ratios.  相似文献   

20.
In family Rubiaceae distyly is very common, and large variation in heterostyly characteristics has been previously documented. Analysis of these variations, even within a species, is very useful to our understanding of the evolutionary process that caused this polymorphism. For this reason, the goal of this study is to investigate the floral morphology and diallelic incompatibility system of three populations of Psychotria carthagenensis. The three studied populations of P. carthagenensis occur in forest fragments in protected areas in an urban matrix in the Municipality of Campo Grande, Mato Grosso do Sul State, Brazil. Reciprocal position of style length and stamen height was found in populations, and in general short-styled flowers were larger than long-styled flowers. All populations presented 1:1 morph ratio between short-styled and long-styled flowers. Regarding breeding system, flowers of P. carthagenensis were self-compatible and compatible within plants of the same morph, and there was no pollen limitation in the populations in any case. In only one of the populations were there differences in the extent of compatibility between morphs, with the long-styled morph being more self-compatible than the short-styled morph. The reproductive strategy of these populations can be advantageous in case of fluctuation of pollinator activity.  相似文献   

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