Stigma Height Research Articles

Population‐level variation in the expression of heterostyly in three species of Rubiaceae: does reciprocal placement of anthers and stigmas characterize heterostyly?

Heterostyly (i.e., reciprocal placement of anthers and stigmas between two or three floral morphs) is hypothesized to enhance outcrossing and reduce selfing. However, few studies have documented reciprocity among individual plants; instead, mean anther and stigma heights for floral morphs are usually reported, masking interindividual variation. We measured eight floral dimensions for individuals in five populations of three heterostylous Rubiaceae. The three methods used to quantify reciprocity yielded different conclusions regarding the degree to which populations conformed to expectations for heterostylous plants. Only Psychotria poeppigiana had stigma and, to a lesser degree, anther heights in discrete classes. Variation among plants of Bouvardia ternifolia and Psychotria chiapensis yielded a continuum of anther and stigma heights across populations. Comparison of distances between stigma and anthers indicated that only flowers of B. ternifolia had, as expected, a constant value for this distance. Finally, regression relationships between anther and stigma heights and corolla length showed that only in one population each of B. ternifolia and P. poeppigiana, and in P. chiapensis, was distance between anthers and stigmas the same across the range of corolla sizes for both floral morphs. Variation among these species in expression of heterostyly was not clearly linked to phylogenetic relationship or pollinator syndromes. Two approach herkogamous (AH) species were studied for comparison. Flowers of Psychotria brachiata were consistently AH, but flowers of P. pittieri were highly variable. Determining fitness consequences of population-level variation in sexual systems requires studies linking floral morphology to pollinator behavior and pollen transfer.

Patterns and consequences of stigma height variation in a natural population of a distylous plant, Primula sieboldii

Abstract1. In order to reveal the direction and magnitude of phenotypic selection on the stigma height of heterostylous morphs in a natural population of Primula sieboldii E. Morren (Primulaceae), morphological traits including stigma and anther heights, flowering phenology, local density of potential mating partners, size of ramet, number of pollen grains deposited from the opposite morph and seed set were measured in two flowering seasons.2. Both components of female reproductive success, pollen receipt and seed set, were significantly larger in the long‐styled than in the short‐styled morph in both years.3. Selection analyses based on bivariate correlation and path‐analysis models revealed positive relationships between stigma height and pollination or seed set in the short‐styled morph only, although the effect was statistically significant only in one year. In contrast, seed set in the long‐styled morph was correlated with ramet size.4. Morph difference in reproductive success and the natural selection revealed for higher stigmas in the short‐styled morph should be ascribed to the narrow corolla tube of the species that strictly restrict the path of the proboscis of the pollinators.

Evolution and maintenance of stigma-height dimorphism in Narcissus. II. Fitness comparisons between style morphs.

Populations of the insect-pollinated geophytes Narcissus assoanus and N. dubius (Amaryllidaceae) are commonly dimorphic for stigma height. An extensive survey of populations of the two species in SW France revealed a wide range of style-morph frequencies, particularly populations with significantly more long-styled than short-styled plants. Here we employ experimental and theoretical approaches to investigate potential selective mechanisms governing the variation in style-morph frequencies. Controlled pollination of both species demonstrated that N. assoanus is moderately self-sterile whereas N. dubius is highly self-compatible. Both intra- and intermorph crosses of N. assoanus were equally fertile, indicating that the species does not exhibit heteromorphic incompatibility. Estimates of female fertility (fruit- and seed-set) and multilocus estimates of outcrossing using allozyme markers provided no evidence of morph-specific differences in maternal components of reproductive success in natural populations of the two species. This result suggested that differences between the morphs in male fertility may be largely responsible for the observed morph-ratio variation. To investigate this hypothesis we developed a mating model that incorporates the genetics of stigma-height dimorphism and contrasting rates of assortative and disassortative mating in the style morphs. Simulation results demonstrated that stigma-height dimorphism will always be maintained when levels of disassortative mating are greater than assortative mating, and that the observed L-biased populations in Narcissus spp. probably result from greater levels of assortative mating in this morph in comparison with the S-morph.

The reproductive ecology of island populations of distylous Menyanthes trifoliata (Menyanthaceae)

Menyanthes trifoliata L. is a distylous, clonal aquatic macrophyte found in shallow bogs and river margins throughout the boreal ecosystem, including the island of Newfoundland. A combination of long-distance dispersal and colonization after deglaciation, and pollen limitation resulting from reduced pollinator diversity and abundance documented on islands, predicts the breakdown of heterostyly to favour the establishment of self-compatible homostyles on islands. To test if self-fertilizing homostyles have been selected, variation in floral characters and compatibility relationships were examined in M. trifoliata populations from the Avalon Peninsula of Newfoundland. Morph ratio and its effect on fruit and seed set were examined in nine populations. Of the seven dimorphic populations, morphs occurred in a 1:1 ratio in four populations and deviated significantly from a 1:1 ratio in three populations. The two populations monomorphic for either pin or thrum morphs set few fruits or seeds (<15%). A strictly reciprocal arrangement of stigma height and anther length was not observed between pin and thrum morphs in the majority of populations studied. Stigma-anther separation showed a bimodal distribution with few intermediate "homostylous" flowers, rather than the discreet bimodal distribution typical of distylous species. Fruit and seed set were high (>60%) in equal morph ratio populations and were not significantly correlated to stigma-anther separation, indicating that there was no selective advantage of being homostylous. All three populations tested were highly self-incompatible, confirming that there has not been a breakdown of heterostyly on the island of Newfoundland. A reduced pollinator fauna typical of island environments may have relaxed stabilizing selection for strict herkogamy between floral morphs, resulting in the observed lack of reciprocity.Key words: Menyanthes trifoliata, distyly, homostyle, reciprocal herkogamy, clonal aquatic macrophyte, island of Newfoundland.

Distyly and variation in heteromorphic Incompatibility in Gaertnera Vaginata (Rubiaceae) endemic to La Reunion Island

Documenting the floral biology of species throughout the Rubiaceae family is of particular interest since heterostyly and dioecy may have evolved more than once in this large family. Unfortunately many species in several tropical regions remain unstudied. The purpose of this paper is to describe the floral biology, the nature of self-incompatibility, morph ratios, and fecundity in natural populations of Gaertnera vaginata, a small tree endemic to the island of La Reunion in the Indian Ocean. Measurements of floral characters in populations across the entire distribution of this species showed that G. vaginata exhibits a reciprocal stigma height and anther height dimorphism characteristic of a distylous species. Pollen grain size and corolla tube length are consistently greater in short-styled plants and long-styled plants produce more pollen per flower. Controlled pollinations in a natural population showed that 25% of the short-styled plants gave at least one fruit on intramorph (illegitimate) pollination, whereas no long-styled plants set fruit on illegitimate pollination. In total, 19.4% of illegitimate pollinations produced fruit on short-styled plants. No self-pollination gave fruit on either morph and between-morph pollinations produced 92.2 and 92.8% for short and long-styled plants, respectively. Overall, short-styled plants were significantly more abundant than long-styled plants. Short-styled plants outnumbered long-styled plants in 16 of the 19 populations. In three of these populations the morph ratio was significantly different from 1:1. In two natural populations, fruit set was significantly higher on long-styled plants, although the number of seeds per fruit was not significantly different between the two morphs. The possible effect of variation in the strength of heteromorphic incompatibility on observed variation in morph abundance and the possible causes for the variation in fruit set are discussed.

DEVELOPMENT OF TRISTYLY IN PONTEDERIA CORDATA (PONTEDERIACEAE). I. MATURE FLORAL STRUCTURE AND PATTERNS OF RELATIVE GROWTH OF REPRODUCTIVE ORGANS

Pontederia cordata is a tristylous, self‐incompatible emergent aquatic. Measurements of reproductive organs in mature flowers and developing buds and analysis of relative growth rates of the styles, filaments, and floral tube were used to analyze the developmental processes that result in reciprocal positioning of reproductive parts among the style morphs. Flowers are trimerous with two series of three tepals, two series of three stamens, and a tricarpellate ovary with a single ovule. Each flower has stamens of two lengths, but the two lengths correspond to upper vs. lower position in the zygomorphic flowers, not to the two series initiated in the primordium. Morph‐dependent variation in stigma height depends on differences in style length, not ovary length. Differences among morphs in anther height result from differences in the position of filament insertion on the floral tube and differences in filament length. Styles of the three morphs develop to different lengths as a result of two distinct processes. Styles of L and M morphs develop at different rates with the L style growing more rapidly than the M style, whereas styles of the S morph have a brief period of rapid growth followed by early inhibition. Stamen growth in the S morph also differs qualitatively from that in the L and M morphs, which are distinguished from each other by quantitative differences in growth rates. Results indicate that the developmental processes that result in the complementary arrangements of organs in different morphs are morph‐specific. An attempt is made to integrate the findings of this study with the model for genetic control of tristyly.