Relationship between herkogamy,incompatibility and reciprocity with pollen–ovule ratios in Melochia (Malvaceae)

Heterostyly is a floral polymorphism that increases pollination efficiency by promoting cross-pollination and reducing pollen wastage. Efficiency in pollination has been related to plant investment in gamete production and to the pollen to ovule ratio (P/O), which has been proposed as an indication of the likelihood of enough pollen grains reaching the stigmas to result in maximum reproductive success. In heterostylous species, cross-pollination is promoted by the reciprocal position of sexual organs between morphs and a heteromorphic incompatibility system, which precludes selfing and fertilizations among plants of the same morph. Morphological features like reciprocity (between morphs) and herkogamy (within morph) together with the breeding system are thought to influence pollination quality. Therefore, a close relationship between the pollination efficiency, morphological characteristics, and incompatibility would be expected. Pollination treatments and morphological measurements were carried out to describe the breeding system, herkogamy, and reciprocity of six Melochia species. Afterward, the relation between the P/O (as a surrogate of the efficiency in pollination), and reciprocity, herkogamy and incompatibility was evaluated. Monomorphic M. nodiflora and distylous M. pyramidata are self-compatible species, whereas the rest of the species are self- and morph-incompatible. There was a positive relationship between the P/O value and the degree of herkogamy and incompatibility. However, P/O values appear to increase when higher reciprocity is found in the populations. As expected, the lower values of P/O are associated with lower levels of herkogamy and compatibility in the Melochia species studied. The relationship between the factors is discussed under different scenarios of the pollinators’ predictability.     

Trends in the reproductive biology of Venezuelan Melochia (Malvaceae) species

The breeding system, floral morphology, morph frequency, biomass allocation to floral whorls, pollination and reproductive efficiency were examined in four distylous and three monomorphic Melochia species. Monomorphic species were self-compatible and distylous species were self-incompatible. Flowers of homomorphic species were longer than those of distylous species. Herkogamy was significantly higher in pin than in thrum morphs of the two distylous species, and monomorphic species exhibited the lowest values of herkogamy. Pollen/ovule ratios were similar between monomorphic and dimorphic species, irrespective of the self-incompatibility level. Biomass allocation to flowers was biased toward non-sexual structures, attraction and support. Androecium biomass and androecium/gynoecium biomass ratio were larger in thrum than in pin flowers of distylous species indicating maleness in the thrum morph and femaleness in the pin morph. There was no clear difference between fruit set of monomorphic and dimorphic species; however, the greatest fruit set was found in the monomorphic species, M. pyramidata, which is a self-compatible species. Fruit set was significantly higher in pin than in thrum morphs in three out four distylous species and fecundity was only significantly higher in pin morphs of M. caracasana and M. parvifolia. Melochia species have generalist pollination systems. According to the taxonomic classes and number of pollinator species, M. caracasana and M. parvifolia have similar generalist pollination systems. Our comparative analyses of the characters between floral morphs of distylous species and the relationship with these characters in monomorphic species allow divergences and similarities to be established and different evolutionary trends to be postulated in the breeding systems of Melochia species. Specifically, M. parvifolia and M. caracasana are apparently biased toward monomorphy and dioecy, respectively.     

Loss of floral polymorphism in heterostylous Luculia pinceana (Rubiaceae): a molecular phylogeographic perspective

Both deterministic and stochastic forces determine the representation and frequency of floral morphs in heterostylous plant populations. Phylogeographic analysis of molecular variation can provide information on the role of historical factors, including founder events, in affecting population morph structure. Here, we investigate geographical patterns of floral morph variation in a distylous shrub Luculia pinceana (Rubiaceae) by examining the relations between floral polymorphism and molecular (cpDNA and microsatellite) variation in 25 populations sampled throughout the distribution of the species in southwest China and adjacent countries. In 19 of the 25 populations, the frequency of floral morphs was not significantly different from the expected 1:1 ratio. The remaining populations were either L‐morph biased (2) or monomorphic (4) for this form and were morphologically differentiated from the remaining populations in several floral traits, that is, corolla tube length, sex organ position and stigma‐anther separation. Phylogeographic analysis supports the hypothesis that L. pinceana was initially split into west‐central and eastern lineages in the Early Pleistocene (～1.982 Mya). A centrally located lineage composed of morph‐biased and monomorphic populations appears to have been subsequently derived from the west‐central lineage, perhaps by a founder event after the last glacial maximum. Hypotheses to explain why these populations have not returned to equilibrium morph frequencies are considered.     

Pollen and stigma size changes during the transition from tristyly to distyly in Oxalis alpina (Oxalidaceae)

• 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.

     

EXPERIMENTAL STUDIES ON THE FUNCTIONAL SIGNIFICANCE OF HETEROSTYLY

Heterostyly has been viewed as both an antiselfing device and a mechanism that increases the proficiency of pollen transfer between plants. We used experimental manipulation of the morph structure of garden populations of self-compatible, tristylous Eichhornia paniculata to investigate the function of floral polymorphism. Outcrossing rates (t), levels of intermorph mating (d), and morph-specific male and female reproductive success were compared in replicate trimorphic and monomorphic populations. In trimorphic populations, t and d averaged 0.81 (2 SE = 0.03) and 0.77 (2 SE = 0.03) respectively, with no difference in either parameter among morphs. Ninety-five percent of outcrossed seeds were therefore the result of intermorph fertilizations. Male reproductive success of the long-styled morph was low, especially in comparison with plants of the short-styled morph. Outcrossing rates for each morph were higher in trimorphic than monomorphic populations where t averaged 0.71 (2 SE = 0.01), 0.30 (2 SE = 0.04) and 0.43 (2 SE = 0.1) for the long-, mid-, and short-styled morphs, respectively. Seed set was lower in monomorphic populations, particularly those composed of the L morph, reflecting reduced pollen deposition. Floral polymorphism therefore increased both outcrossing rate and fecundity but the magnitude of the differences varied among morphs. If the ancestral condition in heterostylous groups resembled the L morph, as has been suggested, data from this study suggests that the selective basis for the establishment of floral polymorphism could have been increased pollen transfer rather than higher levels of outcrossing.     

Pollination of invasive Eichhornia crassipes (Pontederiaceae) by the introduced honeybee (Apis mellifera L.) in South China

Heterostyly functions as an outcrossing mechanism facilitating accurate pollen transfer from anthers to stigmas of particular heights as a result of the behavior of specialist pollinators. However, heterostylous plants are also visited by generalist pollinators, which may affect the plant–pollinator mutualism. Eichhornia crassipes is a tristylous invasive species, with only the mid- and long-styled morphs (M and L) found in China. We recorded flower-visiting insects in Zhuhai, Zhongshan and Nanning in South China. We hand-pollinated the two morphs to determine their compatibility. In addition, by allowing controlled insect pollination in artificial isoplethically monomorphic and bimorphic populations, we undertook a detailed analysis of pollen deposition between the floral morphs, and fruit and seed set. Ranked by relative abundance, the flower-visiting insects were: Apis mellifera, A. cerana, Lasioglossum sp. and Eristalis arvorum. Hand pollination showed that both the M and L morphs were self-compatible, but the former was probably more so than the latter. Intra-morph pollen transfer by A. mellifera within a population was significantly greater than legitimate pollen transfer between populations, suggesting that the pollen exchange between populations was limited. Seed set of the L morph was significantly greater than that of the M morph in monomorphic populations, indicating intra-morph pollen deposition in the former was higher than in the latter. The results showed that A. mellifera was the major pollinator in South China and able to pollinate E. crassipes legitimately and to promote its fruit and seed set, even though high levels of intra-morph pollination occurred.     

Skewed morph ratios lead to lower genetic diversity of the heterostylous Primula veris in fragmented grasslands

• Populations of heterostylous plant species are ideally composed of equal frequencies of two (distylous) or three (tristylous) morphologically different floral morphs. Intra-morph incompatibility helps to avoid inbreeding and to maintain genetic diversity, supporting plant fitness and long-term viability. Habitat fragmentation can lead to skewed morph ratios and thereby reduce the abundance of compatible mates. This, in turn, can result in a loss of genetic diversity. We tested whether the genetic diversity of heterostylous plants is affected by morph ratio bias using populations of the distylous grassland plant Primula veris in recently fragmented grasslands.
• We recorded morph frequencies and population sizes in 30 study populations of P. veris on two Estonian islands characterised by different degrees of habitat fragmentation. Examining variation of thousands of single nucleotide polymorphisms (SNPs) and heterostyly-specific genetic markers, we quantified overall and morph-specific genetic diversity and differentiation in these populations.
• Morph frequencies deviated more in smaller populations. Skewed morph ratios had a negative effect on the genetic diversity of P. veris in more fragmented grasslands. In the populations of better-connected grassland systems, genetic differentiation among S-morphs was higher than among L-morphs.
• Our study shows that deviations from morph balance are stronger in small populations and have a negative impact on the genetic diversity of the distylous plant P. veris. Together with the direct negative effects of habitat loss and decreased population size on the genetic diversity of plants, morph ratio bias may intensify the process of genetic erosion, thus exacerbating the local extinction of heterostylous species.

     

FLORAL VARIATION AND DISTYLY IN GUETTARDA SCABRA (RUBIACEAE)

Controversy exists as to whether the tropical shrub Guettarda scabra (Rubiaceae) is distylous. Variations in stigma and anther position and floral morphology of G. scabra were studied in a population in south Florida. Stigma and anther height have unimodal distributions, but stigma-anther separation is bimodally distributed and can be used to identify a long-styled and a short-styled morph. Stigma width varies between morphs, but anther length, pollen diameter, and stigma papillae length do not. The morphs occur in a 1:1 ratio in the two populations studied. G. scabra is self-compatible and can pollinate itself. Styles of the two morphs have similar relative growth rates in early development. Stylar growth is inhibited in the short-styled morph when buds are approximately 12 mm long. Anther height differs between morphs because of different relative growth rates and because the long-styled morph corolla tube, where the anthers are attached, stops growth before the tube of the short-styled morph. Reciprocity between morphs for average stigma and anther height falls within the range of reciprocity found in other distylous Rubiaceae. Thus G. scabra is morphologically distylous but unusual among distylous species in the variation within morphs and overlap between morphs in stigma and anther heights.     

Dimorphisms and self-incompatibility in the distylous species <Emphasis Type="Italic">Palicourea demissa</Emphasis> (Rubiaceae): possible implications for its reproductive output

Distyly has been interpreted as a mechanism that favors cross-fertilization. In this research we describe floral attributes and ancillary floral polymorphisms typically associated to heterostylous plants in Palicourea demissa (Rubiaceae), a distylous shrub of the Venezuelan Andes cloud forests. A hand-pollination experiment was done to evaluate self- and intramorph incompatibility and female reproductive output in both floral morphs. The studied population was morphologically distylous but morph differences in most ancillary floral polymorphisms and reciprocity of the sexual organ heights were found. The floral morphs were self-incompatible and did not differ in fruit set under controlled cross-pollination conditions, but at the population level they exhibited imperfect reciprocal herkogamy. Fruits and seeds of short-styled plants were larger than those of long-styled plants and fruit set was higher in short-styled plants under natural conditions, suggesting a higher reproductive potential among short-styled plants. Given the 1:1 morph ratio within the studied population, further evidence is needed to determine the influence of floral visitors and seed dispersers in the expression of heterostyly in P. demissa under natural conditions.     

茜草科二型花柱植物滇丁香传粉精确性研究

异型花柱是一种受遗传因素控制的花型多态性现象,其适应意义在于提高不同花型间传粉精确性,从而促进异交。为检验二型花柱植物滇丁香（Luculia pinceana）是如何促进异交并确定有效传粉者的类型,实验统计自然种群中长、短柱型的植株数量,测量花部特征,统计自然种群植株柱头上的型间花粉落置的数量和比例,比较型内、型间花粉的萌发率以及花粉管长度,比较不同人工授粉处理结实情况,观察传粉昆虫类型并测量虫体特征。结果表明（1）滇丁香自然居群中长、短柱型植株的数量没有显著性差异。长柱型的筒深、筒直径、开口直径、雄蕊长和柱头长均显著小于短柱型,而长柱型的叶片长和宽、雌蕊长、花药体积、花粉粒体积、柱头厚均显著大于短柱型。（2）自然种群植株柱头上型间花粉所占比例显著低于型内花粉的比例。（3）型间授粉的花粉萌发率以及花粉管长度均大于型内授粉和混合授粉。（4）型间授粉的结实率显著高于型内授粉的结实率,型内授粉具有一定的结实。（5）夜行性长喙条背天蛾（Cechenena lineosa）是滇丁香的有效传粉者,其吻长和滇丁香的花筒深相适应。本研究表明二型花柱植物滇丁香主要是通过与其花部特征相匹配的长喙天蛾实现有效的传粉,通过促进型间花粉的萌发和花粉管的伸长而有利于型间授粉结实,提高传粉精确性。     

Reproductive ecology of 21 coexisting Psychotria species (Rubiaceae): when is heterostyly lost?

Heterostyly is a genetic polymorphism in which plant populations are composed of two or more morphs that differ in stigma and anther heights. The polymorphism promotes intermorph pollen transfer, thus outcrossing. Heterostyly has been reported in 28 angiosperm families and is frequently lost in heterostylous lineages. To assess ecological factors related to shifts from heterostyly to monomorphy, we examined the reproductive ecology of heterostylous tropical shrubs of Psychotria . Among 21 species at Barro Colorado Island and the nearby Parque Nacional Soberania, Panama, 14 species were heterostylous while seven were monomorphic. A molecular phylogeny and the existence elsewhere of heterostylous populations indicated that the breakdown of heterostyly had occurred independently. Heterostylous and monomorphic species were visited by the same bee species, although visit frequencies were lower in monomorphic species. Monomorphic species had significantly lower population density and greater fruit set than did heterostylous species. Autonomous autogamy made a large contribution to fruit set in monomorphic species and was only rarely observed in heterostylous species. The results indicate monomorphic and heterostylous species produce more seeds through selfing and outcrossing, respectively. The limitation of outcrossing as a result of low population density may be related to the breakdown of heterostyly and the evolution of selfing.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 125–134.     

Is sexual organ reciprocity related to legitimate pollen deposition in distylous Pulmonaria (Boraginaceae)?

Heterostyly, i.e. the reciprocal positioning of anthers and stigmas, is a floral polymorphism that is thought to promote disassortative (i.e. between‐morph) pollination and to maintain genetic diversity within populations. Recent research, however, has shown several cases of heterostylous plant species in which the reciprocal positioning of the sexual organs varies, which may affect the likelihood of ‘legitimate’ pollination between compatible morphs, and hence morph fitness and ultimately the stability of this floral polymorphism. In this study, we investigated floral morphology and stigmatic pollen deposition patterns in nine Pulmonaria species (Boraginaceae) across Europe to test the hypothesis that sexual organ reciprocity is related to legitimate pollen deposition. We used a recently developed adaptive accuracy concept to assess the level of reciprocity and to relate inaccuracies in positioning of sexual organs to legitimate pollen deposition measured on stigmas in the field. The nine investigated Pulmonaria species showed substantial deviations from perfect reciprocity, with total inaccuracies varying between 2.9 and 20.3% relative to the squared mean of all anther and stigma heights in the population. In almost all cases, most of the inaccuracy was generated by the high‐level organs. Total pollen deposition was always higher on stigmas of long‐styled individuals, but stigmas of short‐styled individuals received significantly more legitimate pollen than stigmas of long‐styled individuals (71.23% and 38.75% respectively). We also found a weak but significant relationship between measures of inaccuracy and the level of disassortative pollination. Under these conditions of pollen flow, the distylous mating system is predicted to persist in only eleven (27%) of all sampled populations. Overall, these results indicate that deviations from perfect reciprocal herkogamy are common in Pulmonaria and have an impact on legitimate pollen deposition.     

COMPARATIVE FECUNDITY AND POPULATION COMPOSITION OF HETEROSTYLOUS AND NON-HETEROSTYLOUS SPECIES OF VILLARSIA (MENYANTHACEAE) IN WESTERN AUSTRALIA

Several species of Villarsia in Western Australia are distylous, but their floral morphology is variable and mostly differs from the expected positional relationships of stigmas and anthers. Villarsia albiflora is non-heterostylous but self-incompatible. In the distylous V. calthifolia, V. capitata, and V. lasiosperma, morph ratios were 1:1, but in some populations of V. parnassiifolia the short morph was present in excess. Ovule production per flower of the two morphs of the distylous species varied between populations, but within most populations was equal. In nine populations of the distylous species, percentage seed-set of Longs and Shorts was equal. In the five populations of the distylous species in which percentage seed-set differed between the morphs, seed-set of Longs exceeded that of Shorts. In four populations of V. parnassiifolia with Shorts in excess, percentage seed-set of the two morphs was equal; in a fifth, that of Longs exceeded that of Shorts. Percent seed-set of V. albiflora did not differ from that of the distylous species and remained constant during the flowering season despite a substantial decrease in ovule production during this period. This study demonstrated no reproductive advantage of distyly over monomorphism in the species studied.     

Floral morph composition and pollen limitation in the seed set of <Emphasis Type="Italic">Nymphoides indica</Emphasis> populations

We investigated the floral morph composition and seed set of the populations in a heterostylous aquatic plant, Nymphoides indica O. Kuntze (Menyanthaceae). The bias of floral morph ratio in the populations was negatively correlated with the seed set. In the populations with biased floral morph ratio, the pollen on the stigma of the common morphs were dominated by pollen of the same morph, resulting in reduced seed set. The addition of pollen of the opposite morph on the stigmas of the common morph resulted in an increase of fruit and seed set rates. The intermorph pollination rate and the fruit and seed set rates of Nymphoides indica were highest when two floral morphs were close in the populations. Various insect species including Lepidoptera, Diptera and Hymenoptera visited the flowers. However, geitonogamous pollination was common because of the short flying distance of the pollinators. These results indicate that the seed set of Nymphoides indica is limited by the shortage of compatible pollens and that the success of intermorph pollination is influenced by spatial distribution of the two floral morphs. As Nymphoides indica is endangered, conservation of the populations with high fecundity is proposed by us.     

Patterns of male sterility within and among populations of the distylous shrub Erythroxylum havanense (erythroxylaceae)

Distyly, a reproductive system characterized by the presence of long-styled (thrum). and short-styled (pin) individuals within a population, has been repeatedly used as a model for the study of the evolution of the reproductive systems in plants. Erythroxylum havanense is a distylous species in which most thrum plants fail to develop a fertile androecium, thus behaving as male-sterile or partially male-sterile plants. Short-styled (thrum) individuals have an increased performance as female parents, thereby compensating for their loss of male fitness. Previous studies of populations within close proximity to each other suggest that E. havanense may be involved in a process of gender specialization in which, unlike other heterostylous species, thrum plants are specializing as females and pins (long-styled) as males. In this paper we describe more general patterns of male sterility, one of the first steps in the evolution of gender specialization, among populations of the distylous shrub Erythroxylum havanense. Pollen germination differed among populations (range 0.52 ± 0.03 to 0.06 ± 0.04), and between morphs. Pollen from pin plants was almost two times (1.89) as fertile as that from thrums (0.36 ± 0.03 and 0.19 ± 0.03, pin and thrums respectively). Thrums were significantly more male sterile in four out of five populations. The population where differences between the floral morphs were not apparent showed the lowest levels of pollen fertility. Accordingly, our results indicate that populations of E. havanense show marked differences in pollen fertility and higher male sterility associated with the thrum morph. We hypothesize that differences between morphs could be explained if restorers of male sterility are linked to the distyly haplotype, while differences in genes associated with male sterility could explain the variation among populations. Overall, the prevalence of thrum-biased male sterility across populations suggests that E. havanense is subject to a process of gender specialization.     

Experimental insights on the function of ancillary pollen and stigma polymorphisms in plants with heteromorphic incompatibility

Most heterostylous plants possess a reciprocal arrangement of stigmas and anthers (reciprocal herkogamy), heteromorphic self‐incompatibility, and ancillary polymorphisms of pollen and stigmas. The topographical complementarity hypothesis proposes that ancillary polymorphisms function in the rejection of incompatible pollen thus promoting disassortative pollination. Here, we test this hypothesis by investigating patterns of pollen transfer and capture in populations of dimorphic Armeria maritima and A. pubigera and distylous Limonium vulgare (Plumbaginaceae), and by studying pollen adherence and germination patterns in A. maritima following controlled hand‐pollinations. Armeria lacks reciprocal herkogamy allowing the evaluation of the extent to which ancillary polymorphisms affect the composition of pollen loads. We compared the amounts of compatible and incompatible pollen on stigmas in natural populations and calculated the proficiencies of pollen transfer for each mating type. We detected disassortative pollination in each species, and mating types did not differ in compatible pollen capture, although cob stigmas captured more incompatible pollen. Controlled hand‐pollinations revealed the failure of incompatible pollen to adhere and germinate on stigmas. Our results provided evidence that, while structural in nature, pollen‐stigma dimorphisms are tightly associated with heteromorphic incompatibility and likely function to promote disassortative pollination, especially in the absence of reciprocal herkogamy.     

Ovarian self‐incompatibility in Narcissus papyraceus (Amaryllidaceae) is the result of a pre‐zygotic response

Ovarian self‐incompatibility, including pre‐ and post‐zygotic reactions, is a complex mechanism for which we still lack many details relating to its function and significance. The joint presence of ovarian self‐incompatibility with style polymorphism is a rare combination that is found in the genus Narcissus. Usually, style polymorphic species have heteromorphic (diallelic and linked to style length locus) incompatibility, which prevents fertilization between individuals of the same morph, thereby helping to maintain equal proportions of floral morphs in populations. However, when present, self‐incompatibility in Narcissus is not linked to style polymorphism and cross‐fertilization within each morph is possible. Hence, self‐incompatibility in Narcissus is of particular interest when attempting to unravel the nature of the rejection reaction and aiming to assess possible cryptic differences in the fertilization process in intra‐ and inter‐morph crosses, which might ultimately explain the wide variation of morph‐ratio in the field. We examined the breeding system of Narcissus papyraceus, a style‐dimorphic species that has biased morph ratios in most of its populations. We studied pollen‐tube growth in the pistil and ovule fate after experimentally controlled hand pollinations. The growth of pollen tubes in self‐ and intra‐ and inter‐morph crosses was similar up to the point of micropyle penetration in both morphs but, subsequently, a pre‐zygotic failure appeared to affect male and female gametophytes in selfed pistils. A high proportion of ovules from self‐pollinated flowers showed signs of collapse and self‐pollen tubes were blocked or behaved abnormally before entering the embryo sac. Self‐incompatibility was stronger in the long‐styled morph than in the short‐styled morph. We did not find any conclusive sign of differential functioning between intra‐ and inter‐morph cross‐pollinations in any morph. These results enable us to rule out the possible effects of pollen–pistil interactions in N. papyraceus as a cause of morph‐ratio biases and confirm the exceptional nature of the self‐incompatibility mechanism in this polymorphic species. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177 , 629–643.     

STRUCTURAL DIMORPHISM OF STIGMATIC PAPILLAE IN DISTYLOUS LINUM SPECIES

Intermorph differences in the wall structure and constituents of stigmatic papillae are described for distylous Linum pubescens and L. grandiflorum. In the long-styled morph of both species the wall portion around the apex of the papilla has a thickened cellulose-pectin layer. In the short-styled morph of L. pubescens a cap zone is interposed between the cuticle and apical portion of the papilla wall. The subcuticular cap space contains pectins and lipid particles. Similar particles are also present in deposits on the cuticle surface. Papillae of the short-styled morph in L. grandiflorum lack a cap zone and have only epicuticular lipid deposits. Other distylous Linum species in which the two morphs differ in wall structure of the papillae are L. mucronatum, L. flavum, L. perenne, L. austriacum, and L. maritimum. Studies of stigma dimorphism can help to elucidate evolutionary relations between dimorphic and monomorphic Linum species.     

Heterostyly promotes compatible pollination in buckwheats: Comparisons of intraflower,intraplant, and interplant pollen flow in distylous and homostylous Fagopyrum

Premise of the Study

Heterostyly, the reciprocal positioning of stigmas and anthers in different floral morphs, has long been thought to promote intermorph pollination. However, extensive intramorph pollination occurs commonly in heterostylous species, leading to recurrent questions about the functional and evolutionary significance of heterostyly.

Methods

To identify the sources of stigmatic pollen (autogamous [intraflower], geitonogamous [intraplant], vs. interplant), we emasculated either one flower or entire plants in experimental populations of the two closely related buckwheat species, distylous Fagopyrum esculentum and homostylous F. tataricum. Differences in pollen size allowed unambiguous identification of pollen on stigmas.

Results

Only 2.4% of F. tataricum pollen and 1.5% of F. esculentum pollen arrived successfully on compatible stigmas of other plants. In the former (homostylous) species, 71.3% of the pollen load on stigmas was autogamous, 10.8% was geitonogamous, and 17.9% was interplant. In the latter (distylous) species, 37.45% of the pollen on stigmas was autogamous, 13.8% was geitonogamous, 17.0% was intramorph, and 31.75% was intermorph. The amount of incompatible pollen arriving on stigmas was greatly decreased by both one‐flower and whole‐plant emasculations, and thus, the proportion of compatible pollen deposited increased with one‐flower emasculation and increased even more with whole‐plant emasculation.

Conclusions

Our quantification of pollen‐donor sources in these two species indicated that heterostyly in Fagopyrum esculentum provided a nearly 2‐fold fitness advantage (in terms of compatible pollination) over expected (random) pollen transfers between morphs. Because of reduced herkogamy, the homostylous F. tataricum was highly autogamous.     

Asymmetry in pollen flow promotes gender specialization in morphs of the distylous neotropical herb <Emphasis Type="Italic">Arcytophyllum lavarum</Emphasis> (Rubiaceae)

Distyly is a floral polymorphism, characterized by a reciprocal positioning between stigmas and anthers in different flowers, where two floral morphs, long-styled (pin) and short-styled (thrum) occur within the population. Distyly is suggested as one of the routes leading to the evolution of separate sexes in plants. In this evolutionary pathway, pollinators may disrupt the complementarity of pollen transfer between morphs. Consequently, the floral morphs gradually specialize as either male or female. A key process required for gender specialization in distylous plants is a deviation of the realized functional gender (i.e. the proportion of genes transmitted to the next generation via pollen donation and seed production) from the potential functional gender (i.e. the expected contribution of male and female function to reproductive success from the number of ovules or pollen grains produced by each morph). I selected the distylous herb Arcytophyllum lavarum (Rubiaceae) to determine if asymmetry in pollen flow promotes differences in seed production, pollen donation and a discrepancy between the potential and the realized functional genders in pin and thrum floral morphs. Pollen flow in A. lavarum is asymmetric and the pin morph is more efficient at performing cross-pollination than the thrum morph. Conversely, the thrum morph produced two times more seeds than the pin morph. Male and female contributions to the potential functional gender were equivalent in both morphs. However, the pin morph transmitted more genes through pollen donation and the thrum morph more through seed production than expected from their potential functional genders. These results support the hypothesis that if pollinators consistently promote asymmetric pollen flow between morphs over generations, it is possible that gender specialization may evolve to the extreme of dioecism from an original distylous condition.