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 biology of Alpinia blepharocalyx (Zingiberaceae): another example of flexistyly

The floral biology and breeding system of Alpinia blepharocalyx were studied in Yunnan province, southwest China. Our results indicate that populations of A. blepharocalyx have two floral morphs, which differ in flowering behaviour: the cataflexistylous morph in which the stigma is held erect above the dehiscent anther when anthesis begins in the morning and becomes curved under the anther at afternoon, and the anaflexistylous morph in which the receptive stigma is curved under the indehiscent anther first and moves into a reflexed superior position above the anther as it begins to shed pollen in the afternoon; the flowering period of both floral morphs is 12 hours; the stylar movements in the two floral morphs are synchronous, and they have similar traits to those found in other Alpinia species previously reported. Also, on average, cataflexistylous flowers are larger than anaflexistylous flowers, especially the labellum and corolla tube length; moreover, the P/O ratio of the two floral morphs is significantly different: the cataflexistylous morph has more pollen grains and fewer ovules than the anaflexistylous morph; the evolutionary significance of this comparison is discussed. Cataflexistylous flowers were observed to produce a lower concentration of nectar than that of anaflexistylous flowers before 11:00 a.m., but they then have higher nectar concentration; cataflexistylous flowers always have more nectar than anaflexistylous flowers. Flowers of A. blepharocalyx were visited by fewer insects at a lower frequency, including honey bees (Apis cerana cerana) and two species of carpenter bees (Xylocopa spp.). No significant differences were found in the fruit set among the hand-pollination treatments regardless of selfing (geitonogamy and autogamy) or crossing (intermorph and intramorph); but all hand-pollination treatments exhibited much higher fruit set than the controls; meanwhile, no fruit set occurred in the unpollinated bagged plants. Pollen tube growth was examined using fluorescence microscopy following self, intramorph, and intermorph hand pollinations at different times of the day; the pollen tubes of both floral morphs have the same growth rate and the capability to reach the ovary. Both the results of pollen tube growth and fruit sets under different pollination conditions show that A. blepharocalyx is self-compatible and dependent upon insects for pollination. The distinctive flexistyly mechanism of A. blepharocalyx is likely a floral mechanism that promotes effective intermorph pollen transfer.     

Enantiostyly in Chamaecrista ramosa (Fabaceae‐Caesalpinioideae): floral morphology,pollen transfer dynamics and breeding system

Enantiostyly is a form of reciprocal herkogamy, in which floral morphs present reciprocal differences in the position of sexual elements, and occurs in monomorphic and dimorphic forms. This polymorphism maximises cross‐pollination and reduces self‐pollination, being very common within the subtribe Cassiinae (Fabaceae). Nevertheless, few studies have investigated the functionality of enantiostyly, particularly in this plant group. The present study aimed to investigate enantiostyly and its functionality in Chamaecrista ramosa, a monomorphic enantiostylous shrub, in an area of coastal vegetation in northeast Brazil. Pollen deposition and capture on the body of floral visitors, the relationship of these data with floral biology and breeding system, and morph ratio were evaluated. Pollen deposition and capture occurred in specific sites of the floral visitor body, showing the functionality of enantiostyly. The floral architecture, associated with the floral visitor behaviour, resulted in indirect pollen deposition on the floral visitor body. This occurred through a loop made by the pollen upon the inner petal surface, similar that generally reported for other Cassiinae. Chamaecrista ramosa is self‐compatible, although no fruit set was observed through spontaneous self‐pollination. The occurrence and number of floral morphs was similar within clumps. Enantiostyly seems to be advantageous for this species, as it results in efficient pollen capture and deposition, reduces the chances of autogamy and maximises intermorph pollen flow.     

异型花柱植物喀什补血草的传粉者功能群与花粉转移效率

异型花柱是受遗传控制的花柱多态现象, 被达尔文认为是植物通过在传粉者体表不同部位滞落花粉以促进型间花粉准确传递的一种适应。该现象虽已受到广泛关注, 但在一些花型变异较大且不稳定的传粉系统中, 不同传粉者对各花型繁殖所产生的影响仍知之甚少。该研究以分布于新疆天山南坡的一个有同长花柱共存的异型花柱植物喀什补血草(Limonium kaschgaricum)种群为研究对象, 对其花型构成及频率、传粉者及花粉转移效率等进行了调查分析。结果表明: 1)种群中除了存在雌/雄蕊长度交互对应的长(L)/短(S)花柱型花外, 还有雌/雄蕊同长的花(H型), 且各花型花的花冠口直径、花冠筒长及花粉量等参数间无差异, 但花粉纹饰和柱头乳突细胞形态具二型性。其中, H型花的花粉和柱头形态与L型花(或S型花)的一致。2)花型内和自花授粉均不亲和; 型间授粉时, 花粉和柱头形态不同的花型间亲和, 反之不亲和。3)种群内存在长/短吻两类传粉昆虫。在以短吻传粉者为主的盛花初、中期, L和H型花柱头上的异型花粉数均显著高于S型花的, 且L和S型花高位性器官间的异型花粉传递效率高于低位性器官间的; 而在以长吻传粉者为主的盛花后期, L和S型花的柱头间异型花粉数无显著差异, 且高/低位性器官间具有相同的异型花粉转移效率; 与传粉者出现时期相对应的、在花期不同阶段开放花的结实率也明显不同。4)长/短吻昆虫具明显不同的传粉功能, 短吻昆虫只能对L和H型花进行有效传粉, 且访花频率和型间花粉转移效率较低, 为低效传粉者; 而长吻昆虫对各花型均能有效传粉, 具高的访花频率和型间花粉转移效率, 为高效传粉者。因为长吻昆虫的阶段性出现所形成的不稳定传粉系统, 使低效的短吻昆虫可能会成为种群中花型变异的驱动力, 并使S型花受到更大的选择压力。H型花克服了柱头缩入的弊端, 可能会成为不稳定传粉系统下的一个替代花型而持续存在。     

Heteromorphic incompatibility and efficiency of pollination in two distylous Pentanisia species (Rubiaceae)

BACKGROUND AND AIMS: Distyly has been hypothesized to promote cross-pollination by reducing intrafloral and geitonogamous self-pollination, and enhancing intermorph pollination. Distylous plants typically display both reciprocal herkogamy and a heteromorphic incompatibility system, which allows mating only between morphs. Distyly and its pollination consequences were examined in two Pentanisia species with long-tubed flowers which are pollinated almost exclusively by butterflies. METHODS: Anther and stigma heights were measured to quantify reciprocal herkogamy. The type of incompatibility system was determined by observing pollen tubes and seed production following controlled hand pollination. Pollen loads on pollinators and stigmas were also examined to assess the efficiency of intermorph pollen flow. KEY RESULTS: Pentanisia prunelloides and P. angustifolia exhibit reciprocal herkogamy and a host of ancillary dimorphisms, including pollen colour, exine sculpturing, stigmatic papilla shape and floral-tube pubescence. Controlled hand-pollinations revealed the presence of a strong heteromorphic incompatibility system in both species. The site of incompatibility differed between the morphs; intramorph pollen tubes were blocked in the style of the short-styled morph and on the stigmatic surface of the long-styled morph. Butterflies carried pollen from the short- and long-styled morphs primarily on their head and proboscis, respectively. Natural pollination resulted in a higher proportion of pollen transfer from long- to short-styled plants than vice versa. Nevertheless, fruit set did not differ between morphs. CONCLUSIONS: Both Pentanisia species are fully distylous. Reciprocal herkogamy results in pollen from the two morphs being carried on different locations on pollinators' bodies, which in turn promotes intermorph pollination. Intramorph pollination does not result in fertilization, because of an effective heteromorphic incompatibility system.     

Floral display size and spatial distribution of potential mates affect pollen deposition and female reproductive success in distylous Pulmonaria officinalis (Boraginaceae)

In animal‐pollinated plants, both the spatial distribution of flowering individuals and the number of flowers that an individual displays affect pollen deposition rates and female reproductive success. Heterostylous species are likely to be particularly sensitive to the contingencies of spatial distribution, as they are reproductively subdivided into distinct mating groups, which usually exhibit self‐ and intra‐morph incompatibility and differ in floral morphology. In this paper, we explore the joint effects of both spatial distribution of potential mates and floral display size on morph‐specific pollen deposition rates and seed set patterns in two natural populations of Pulmonaria officinalis, a distylous species with a weak self‐incompatibility system. Both total stigmatic pollen load and the proportion of legitimate pollen decreased with increasing spatial isolation. Legitimate (intermorph) pollen transfer was, however, asymmetric and decreased more rapidly with decreasing proximity to a compatible legitimate mating partner in the S‐morph than in the L‐morph. Total stigmatic pollen loads per flower increased with increasing floral display size, indicating that large plants are disproportionately more visited than smaller individuals. However, because legitimate pollen deposition decreased with increasing floral display size, these results also suggest that larger numbers of flowers increase the degree of geitonogamous pollination. In both the L‐ and S‐morph, seed set significantly decreased with increasing isolation from a legitimate mating partner, but in the L‐morph seed set was less dependent on the spatial distribution of the S‐morph. In addition, seed set significantly increased with floral display size in the L‐morph, but not in the S‐morph. These findings indicate that the spatial distribution of potential mates and variation in floral display size may cause morph‐specific differences in pollen deposition rates and female reproductive success.     

Impact of mate availability, population size, and spatial aggregation of morphs on sexual reproduction in a distylous, aquatic plant

In distylous, self-incompatible plants, clonal propagation, unbalanced floral morph frequencies, and reduced population size can interfere with the functioning of distyly by compromising legitimate intermorph pollinations, resulting in reduced reproductive output. Here, we examined the mating system and the impact of mate availability, population size, and spatial aggregation of morphs on reproductive output in the distylous, clonal, aquatic plant Hottonia palustris. Controlled pollinations under greenhouse conditions detected no spontaneous selfing without the action of a pollen vector (autonomous autogamy) and demonstrated very low fruit and seed development after self-pollination. Intermorph (legitimate) crossings resulted in high reproductive output in both floral morphs (long- and short-styled individuals), whereas intramorph (illegitimate) crossings decreased fruit and seed development by more than 50%, indicating that the species has partial intramorph-incompatibility. In natural populations, small population size and increasing deviation of floral morph frequencies negatively affected reproductive outcome. Individuals of the majority morph type developed significantly fewer fruit and seeds than individuals of the minority morph type. This rapid decline in fecundity was symmetrical, indicating that regardless of which morph was in the majority, the same patterns of negative frequency-dependent mating occurred. Increasing spatial isolation between compatible morphs significantly reduced fruit and seed set in both morphs similarly. This study provides clear indications of frequency- and context-dependent mating in natural populations of a distylous plant species.     

The Breeding System of Oxalis squamata,a Tristylous South American Species

Oxalis squamata is a perennial endemic herb of the Chilean-Argentine Andes previously reported as a distylous species. In this paper, analysis of floral morphology, incompatibility relations and population structure show that O. squamata possesses morphological and functional tristyly. The three floral morphs differ in calyx and corolla length, style and stamen length, pollen grain size and stylar pubescence. Results of hand pollinations showed the presence of a strong incompatibility system. Legitimate pollinations with the L and M morphs as seed parents produced approximately equal numbers of seeds per pollination. The S morph produced less than half the number of seeds per legitimate pollination compared to the other two morphs. Intra- and intermorph illegitimate pollinations produce almost no seeds. Morph representation in the population was isoplethic. Fruit and seed production did not differ significantly for the three morphs in the population. Comparison of experimental and natural production of fruits and seeds suggests that seed set in the natural population may be pollen-limited for the L and M morphs.     

Morph‐specific selection on floral traits in a polymorphic plant

Correlations between phenotypic traits are common in many organisms, but the relative importance of nonadaptive mechanisms and selection for the evolution and maintenance of such correlations are poorly understood. In polymorphic species, morphs may evolve quantitative differences in additional characters as a result of morph‐specific selection. The perennial rosette herb Primula farinosa is polymorphic for scape length. The short‐scaped morph is less damaged by grazers and seed predators but is more strongly pollen limited than the long‐scaped morph. We examined whether morph‐specific differences in biotic interactions are associated with differences in selection on two other traits affecting floral display (number of flowers and petal size) and on one trait likely to affect pollination efficiency (corolla tube width) in three P. farinosa populations. Differences in selection between morphs were detected in one population. In this population, selection for more flowers and larger petals was stronger in the short‐scaped than in the long‐scaped morph, and although there was selection for narrower corolla tubes in the short‐scaped morph, no statistically significant selection on corolla tube width could be detected in the long‐scaped morph. In the study populations, the short‐scaped morph produced more and larger flowers and wider corolla tubes. Current morph‐specific selection was thus only partly consistent with trait differences between morphs. The results provide evidence of morph‐specific selection on traits associated with floral display and pollination efficiency, respectively.     

艳丽耳草的二型花柱及异型自交不亲和系统

异型花柱是一种受遗传因素控制的花型多态性现象,包括二型花柱和三型花柱两种类型.本文以茜草科艳丽耳草(Hedyotis pulcherrima)为实验材料,通过对其野外居群的花型、花部形态及花粉特征等观察,发现艳丽耳草野外居群同时存在长花柱型花和短花柱型花,长/短花柱型花的数量比例为1∶1.两型花具有精确的交互式雌雄异位特征,并且该特征与花冠长度相关性显著.长/短花柱型花的柱头裂片长度、花粉大小及淀粉含量等具有二型性.花粉体外培养时花粉萌发率及花粉管生长速率无显著性差异.人工授粉后,艳丽耳草长/短花柱型花型间异交花粉管生长形态正常,授粉24 h后花粉管均已进入子房.而长/短花柱型花在自交及型内异交下均表现为不亲和,花粉管生长停止于柱头,花粉管顶端累积胼胝质并膨大.艳丽耳草没有无融合生殖现象,型间人工辅助异交授粉结实率为100％,显著高于自然结实率.本研究结果表明,艳丽耳草是典型的二型花柱植物,并具有异型自交不亲和系统.     

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.     

Gender specialization in <Emphasis Type="Italic">Palicourea demissa</Emphasis> (Rubiaceae), a distylous,hummingbird-pollinated treelet

Distyly has been interpreted as a mechanism that promotes cross-pollination among conspecific plants and as one of the routes leading to the evolution of dioecy. In one of the possible evolutionary pathways, pollinators may disrupt intermorph pollen flow, and, as a consequence, floral morphs may gradually specialize as either male or female (functional dioecy). Natural patterns of pollen deposition and fruit and seed production were estimated in Palicourea demissa (Rubiaceae) and used as parameters to assess functional gender differences between floral morphs. Pollen flow was asymmetrical in P. demissa. Long-styled flowers were more effective than short-styled flowers in pollen deposition towards compatible stigmas, whereas short-styled flowers were more effective in legitimate pollen receipt. Accordingly, short-styled plants produced more fruits and viable seeds than long-styled plants. The contributions of male and female function to the potential functional gender were equivalent in both morphs. However, the realized functional gender deviated significantly from the potential functional gender in both morphs, in which short-styled plants were more successful through their female function, but long-styled plants through their male function. If pollinators disrupt the complementarities of pollen transfer between the two morphs (asymmetric pollen flow), the expression of a more profitable gender is expected in each morph. Thus, our results support the hypothesis that dioecy may evolve in distylous populations through the gradual specialization of each morph as either male or female.     

云南草寇花序内不同部位的性分配

两性花植物花序内的性分配常存在差异,资源竞争、结构效应、交配环境(雌雄异熟、传粉者定向访花行为等)或授粉不均匀等几种假说可以解释这种现象。为验证上述假说,该研究以云南草寇两种表型(雄先熟型和雌先熟型)为材料,分析了其花序内不同部位(基部、中部和顶部)的每花花粉数、胚珠数、花粉/胚珠比、结实率和结籽率,花序内传粉者的定向访花行为,以及人工辅助授粉和去花处理对结实率和结籽率的影响。结果表明:两种表型花序内每花花粉数不随部位而变化,每花胚珠数、结实率和结籽率由基部到顶部依次降低,每花花粉/胚珠比由基部到顶部依次增加,表明顶部花存在偏雄的性分配。人工辅助授粉后,结实率、结籽率仍由基部到顶部依次降低,表明授粉不均匀假说不能解释云南草寇花序内不同部位结实率、结籽率的差异。去除基部和中部花后,顶部花人工辅助授粉条件下的结实率、结籽率与基部花人工辅助授粉条件下的结实率、结籽率无差异,表明云南草寇花序内不同部位结实率、结籽率的差异主要由资源竞争引起。雌先熟表型每花花粉数、花粉/胚珠比高于雄先熟表型,表明两种表型存在性分配差异。传粉者主要先访问云南草寇基部的花,然后向顶部移动。云南草寇花序内顶部偏雄的性分配可能是由资源竞争和传粉者定向访花造成的。     

STIGMATIC POLLEN LOADS IN POPULATIONS OF PONTEDERIA CORDATA FROM THE SOUTHERN U.S.

Stigmatic pollen loads were examined in four tristylous populations of Pontederia cordata from the southern U.S. to evaluate Darwin's hypothesis that floral trimorphism promotes legitimate pollination. In each population, morph frequency, pollen production, the composition of stigmatic pollen loads and seed production were estimated. Goodness-of-fit tests and a measure of the efficiency of legitimate pollination were employed to evaluate pollen load patterns. Results from the four populations demonstrate that the long-styled (L) morph usually experiences legitimate pollination, whereas the mid-styled (M) and short-styled (S) morphs often display random pollination. However, at Taylor Road (Louisiana) all three floral morphs exhibited statistically significant levels of legitimate pollination. The size of legitimate pollen loads suggests that in populations of P. cordata tristyly may have only a minor influence on the fecundity of morphs. Two hypotheses are proposed to explain the maintenance of floral trimorphism in contemporary populations of Pontederia spp. The first assumes the polymorphism is selectively neutral; the second suggests that it contributes towards the male component of fitness by increasing the distance, and hence the number of genets, that pollen can be transported to by pollinators. In clonal species, such as Pontederia cordata, where geitonogamous pollinations are promoted by large floral displays, floral mechanisms that increase pollen carryover may have selective value.     

Reproductive consequences of interactions between clonal growth and sexual reproduction in Nymphoides peltata: a distylous aquatic plant

Distyly is a sexual polymorphism in which plant populations contain two floral morphs differing in morphology and physiology. The dimorphism serves to promote animal-mediated cross-pollination between the floral morphs. Clonal propagation can interfere with the functioning of distyly by compromising intermorph pollinations, resulting in reduced fertility. Here, we investigate the relations between clonal growth and sexual reproduction in the aquatic macrophyte Nymphoides peltata (Menyanthaceae). Surveys of morph representation in 30 populations from five regions of China revealed that most populations exhibited strongly biased morph ratios and 30% contained a single floral morph. Experimental pollinations indicate that N. peltata possesses a strong dimorphic incompatibility system preventing self and intramorph fertilizations. An experiment involving the manipulation of morph ratios in an experimental population and an investigation in a natural population with strong morph substructure both provided evidence that compatible pollen dilution limits fertility. Despite constraints on the functioning of distyly in N. peltata we found no evidence for evolutionary changes to the heterostylous syndrome, as reported in Nymphoides, including populations of N. peltata in other parts of its geographical range.     

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.     

Cryptic self‐incompatibility and distyly in Hedyotis acutangula Champ. (Rubiaceae)

Distyly, floral polymorphism frequently associated with reciprocal herkogamy, self‐ and intramorph incompatibility and secondary dimorphism, constitutes an important sexual system in the Rubiaceae. Here we report an unusual kind of distyly associated with self‐ and/or intramorph compatibility in a perennial herb, Hedyotis acutangula. Floral morphology, ancillary dimorphisms and compatibility of the two morphs were studied. H. acutangula did not exhibit precise reciprocal herkogamy, but this did not affect the equality of floral morphs in the population, as usually found in distylous plants. Both pin and thrum pollen retained relatively high viability for 8 h. The pollen to ovule ratio was 72.5 in pin flowers and 54.4 in thrum flowers. Pistils of pin flowers remained receptive for longer than those of thrum flowers. No apparent difference in the germination rate of pin and thrum pollen grains was observed when cultured in vitro, although growth of thrum pollen tubes was much faster than that of pin pollen tubes. Artificial pollination revealed that pollen tube growth in legitimate intermorph crosses was faster than in either intramorph crosses or self‐pollination, suggesting the occurrence of cryptic self‐incompatibility in this species. Cryptic self‐incompatibility functioned differently in the two morphs, with pollen tube growth rates after legitimate and illegitimate pollination much more highly differentiated in pin flowers than in thrum flowers. No fruit was produced in emasculated netted flowers, suggesting the absence of apomixis. Our results indicate that H. acutangula is distylous, with a cryptic self‐incompatibility breeding system.     

Reproductive ecology of distylous Palicourea Padifolia (Rubiaceae) in a tropical montane cloud forest. I. Hummingbirds' effectiveness as pollen vectors

The adaptiveness of distyly has been typically investigated in terms of its female function, specifically pollen receipt. However, pollen loads on stigmas can only provide moderate support for Darwin's hypothesis of the promotion of legitimate crosses. To determine the effectiveness of hummingbirds as pollen vectors between floral morphs and the consequences in terms of male (pollen transfer) and female function (pollen receipt) in Palicourea padifolia (Rubiaceae), floral visitors, their foraging modes, and temporal patterns of floral visitation were observed and documented. Differences in pollen and stigma morphology, pollen flow, rates of pollen deposition, and/or stigmatic pollen loads were then evaluated for their contribution toward differences in reproductive output between floral morphs. A pollination experiment with stuffed hummingbirds that varied in bill size was done to evaluate the contribution of bill variation toward differences between floral morphs in pollen receipt and pollen transfer and female reproductive output. Anthers of long-styled flowers contained significantly more and smaller pollen grains than those of short-styled flowers, independently of corolla and anther lengths. The shape and orientation of the stigma lobes differed between morphs and were significantly longer among short-styled flowers. Hummingbird visitation rates did not differ significantly between floral morphs, and foraging movements from focal plants towards neighboring plants were independent of floral morph. Stigmatic pollen loads under field conditions and those after controlled hummingbird visitation, along with rates of pollen accumulation through the day indicated that stigmas of short-styled flowers receive proportionately more legitimate (intermorph) pollen grains than did those of long-styled flowers. However, the species of hummingbird was marginally significant in explaining variation in pollen deposition on stigmas. Lastly, intermorph pollinations of P. padifolia resulted in significant differences in fruit production between floral morphs, independent of pollination treatment and pollinator species; short-styled flowers proportionately developed almost twice the number of fruits developed by long-styled flowers.     

Flowering phenology and mating success of the heterodichogamous tree Machilus thunbergii Sieb. et Zucc (Lauraceae)

Heterodichogamy is defined as the presence of two flower morphs that exhibit male and female functions at different times among individuals within a population, and is regarded as an adaptation to promote outbreeding through enhanced intermorph pollination. In highly fragmented populations in which the morph frequency is biased, heterodichogamy may hamper population growth by reducing seed sets of the more numerous morph, and enhancing seed sets of the less numerous morph. In such situations, we hypothesize that individual plants experience greater seed sets if the opposite sexual morphs are nearby, and that individuals of a less numerous sexual morph have greater seed sets. After confirming heterodichogamy by observing flowering behavior and phenology, we tested these two hypotheses in a highly fragmented population of Machilus thunbergii, a putative heterodichogamous evergreen laurel tree. Our observations confirmed that M. thunbergii is heterodichogamous, consisting of two types of protogynous and bisexual flowers: a morning female (MF)–afternoon male morph and a morning male (MM)–afternoon female morph at the individual level. Sexual expression of the two morphs was highly synchronized and reciprocal. Investigation of seed‐set rates revealed greater rates of both morphs if the opposite morph was nearby. The less numerous sexual morph (MF) showed a greater seed‐set rate than the more numerous sexual morph (MM).     

Fertility differences among floral morphs following selfing in tristylous Eichhornia paniculata (Pontederiaceae): inbreeding depression or partial incompatibility?

Reduction in seed set following self- vs. cross-pollination in flowering plants can result from abortion of selfed offspring owing to inbreeding depression and/or partial self-incompatibility. Previous studies on tristylous Eichhornia paniculata (Pontederiaceae) indicate that reduced seed set following self-pollination generally occurs in the short- (S), but not the long-(L) or mid-styled (M) morphs. To determine whether this pattern results from morph-specific differences in inbreeding depression owing to the sheltering of deleterious alleles at the S locus and/or partial self-incompatibility, we conducted controlled hand-pollinations of the floral morphs and measured seed set and levels of seed abortion. There were no significant differences in fertilization success and seed set following self-, illegitimate, and legitimate pollinations in the L and M morphs. In contrast, in the S morph self-, intramorph and intermorph illegitimate pollinations resulted in significant reduction in seed set in comparison with legitimate pollination. This indicates that the reduced seed set observed in self-pollination is the result of partial incompatibility rather than inbreeding depression. Significantly reduced fertilization success and low levels of ovule abortion in illegitimate pollinations of S plants also supported this conclusion. Reduced fertility in the S morph may have implications for the observed loss of this morph from natural populations and the evolutionary breakdown of tristyly.