Resources limit the fecundity of three woodland herbs

Summary Effects of supplemental hand pollination on fruit set, seed number, and seed weight were examined for 3 perennial, woodland herbs, Uvularia sessilifolia, Geranium maculatum, and Maianthemum canadense. We found no evidence for pollen limitation of any measure of fecundity. Low light probably limited fecundity in Geranium, while soil nitrogen limited the number of sseds/fruit in Maianthemum. For Geranium and Maianthemum the relationship between the number of flowers and fecundity varied in differeent years. Maianthemum ramets with greater than average flower numbers had a decreased chance of reproductive failure than did ramets with few flowers.     

克隆生长对被子植物传粉过程的影响

克隆植物与其传粉者的相互作用是植物繁殖生态学的重要研究领域之一。植物克隆生长与有性繁殖通常相伴进行, 往往产生较大的花展示与复杂的克隆空间结构, 通过传粉过程对有性繁殖过程产生影响, 共同决定植物的适合度。本文回顾了克隆生长对被子植物传粉过程影响的国内外研究进展, 从植物克隆大小、花资源空间配置、克隆构型与种群遗传结构四个方面讨论了克隆生长对传粉过程的影响及其生态学与进化生物学意义。早期研究预期, 随着克隆增大, 同株异花授粉水平增加, 因而通过增大自交率或花粉阻塞效应降低植物的适合度。但是, 后来的一些模拟与野外实验研究发现, 传粉者在同一克隆内访问的花数量并不会随克隆增大而一直增加, 访花行为也主要发生在分株内; 而且分子标记的自交率组分分析也表明自交主要发生在分株内。另一方面, 人工模型模拟以及传粉者访问行为研究表明, 当花朵数量相同时, 与所有花集中生长在同一分株上相比, 将花朵分散在多个分株上的克隆生长方式不会增加, 反而降低了同株异花授粉的发生水平。如果花序内花雌雄同熟, 花朵同时提供与接收花粉, 克隆生长会使植物接收到更高比例的异交花粉, 在提高后代质量的同时不增加同株异花授粉概率。这是从传粉生物学角度对植物克隆生长习性进化的一个全新的解释。今后, 克隆植物传粉生物学研究需要针对传粉者与克隆生长之间的相互作用建立理论模型, 探究克隆大小、克隆构型、花资源空间配置模式对传粉者访问频率和行为、花粉散布、交配格局的影响。同时, 需要在自然种群中, 尤其是克隆与非克隆的近缘类群、同一物种克隆与非克隆种群开展比较研究, 利用更高效的分子标记来研究克隆生长的生态与进化意义。     

Floral distribution,clonal structure,and their effects on pollination success in a self-incompatible <Emphasis Type="Italic">Convallaria keiskei</Emphasis> population in northern Japan

In plant species, when clonal growth produces a patchy structure and flowering ramets are clustered, the amount of pollen contributing to reproductive success is often regulated by pollinator efficiency and geitonogamy. The spatial population structure may influence reproductive success. We examined the clonal structure, the spatial ramet distribution, and their combined effects on fruit set in a natural population of the insect-pollinated, self-incompatible clonal herb, Convallaria keiskei, in northern Japan. The number of shoots, flowers, and fruits in 1-m² quadrats were counted at every 5 m grid point in an established 100 × 90-m study plot. From all the quadrats where shoots existed, leaf samples were collected for allozyme analysis. Using the two spatial parameters of flowering ramet densities and genotypes, we then constructed individual-based fruit-set models. A total of 236 quadrats contained shoots, and 135 contained flowering ramets, which indicated expanded distribution of this plant throughout the study plot, while shoots, flowers and fruits all showed clustering distributions. Allozyme analysis of 282 samples revealed 94 multilocus genotypes. The largest clone extended to more than 40 m, whereas 56 genotypes were detected in only one sample. Several large clones and many small clones were distributed close to each other. Fine-scale spatial modelling revealed that the neighbouring flower numbers of different genotypes, compared with local genet or flower diversity, more influenced fruit set, in which the range of the neighbour was 14.5 m. These findings indicate that the compatible pollen dispersed by insect pollinators has a significant effect on sexual reproduction, in this C. keiskei population. Consequently, the spatial structure, which includes both genet distribution and clonal expansion by ramets, had a significant effect on pollination success.     

Consequences of clonal growth for plant mating

By affecting the number and the spatial distribution of flowering units (i.e., ramets), clonal growth can influence pollen transfer between plants and thus affect mating opportunities. In this paper I review some recent work that attests the importance of clonal growth for pollination patterns. A major aspect concerns the increase in floral display through the multiplication of flowering ramets. Although large floral displays can enhance pollinator attraction and may thus promote outcrossing, they can also increase rates of geitonogamy (i.e., pollination between flowers in the same plant). However, the latter aspect will depend on clonal architecture, a feature that greatly varies among clonal plant species. Future experimental studies and comparative analyses of rates of geitonogamy in species with clumped and intermingled distribution may allow for the evaluation of evolutionary interaction between clonal growth and floral traits that regulate mating patterns.     

Clonal growth and its effects on male and female reproductive success in <Emphasis Type="Italic">Prunus ssiori</Emphasis> (Rosaceae)

Clonal growth can increase not only floral display but also geitonogamy and may affect sexual reproduction both positively and negatively. A clonal woody species, Prunus ssiori, was partially self-incompatible according to a pollination experiment. Its main pollinators, bumble bees, were often observed to consecutively visit inflorescences within a tree. Clone identification revealed that its genets formed mutually exclusive patches. These features suggest frequent geitonogamous pollination. In a 6.24-ha plot, 212 trees belonged to 59 genets, and 42 genets consisted of a single tree, whereas the rest contained two or more clonal trees. The largest genet had 65 trees and occupied 0.4 of a hectare. Fruit set was measured in 127 inflorescences sampled from nine maternal trees at the center of the plot. Paternal genets of 107 of their 300 seeds were assigned in the plot using microsatellites. There were no selfed seeds. Male reproductive success (the probability that individual trees of each genet sired a seed) increased as tree size increased, as the distance between the trees and maternal trees decreased, when the genet did not contain the maternal trees, and when the genet consisted of a single tree. Female reproductive success (fruit set in individual inflorescences of each maternal tree) increased as the within-tree geitonogamy index, which reflected the frequency of pollination within the maternal tree, decreased. These results suggest that clonal growth reduces male reproductive success, at least, in P. ssiori, because of pollen discounting.     

Reproductive consequences of mate quantity versus mate diversity in a wind-pollinated plant

Since most pollen travels limited distances in wind-pollinated plants, both the local quantity and diversity of mates may limit female reproductive success. Yet little evidence exists on their relative contribution, despite the importance of viable seed production to population dynamics.To study how variation in female reproductive success is affected by the quantity versus the diversity of surrounding mates contributing pollen, we integrated pollination experiments, data on natural seed set and seed viability, and AFLP genetic marker data in the wind-pollinated dioecious clonal forest herb Mercurialis perennis.Pollination experiments indicated weak quantitative pollen limitation effects on seed set. Among-population crosses showed reduced seed viability, suggesting outbreeding depression due to genetic divergence. Pollination with pollen from a single source did not negatively affect reproductive success. These findings were consistent with results of the survey of natural female reproductive success. Seed set decreased with the distance to males in a female plants’ local neighborhood, suggesting a shortage of pollen in isolated female plants, and increased with the degree of local genetic diversity. Spatial isolation to other populations and population size did not affect seed set. None of these variables were related to seed viability.We conclude that pollen movement in M. perennis is likely very limited. Both male proximity and the local degree of genetic diversity influenced female reproductive success.     

Plant size,geitonogamy and seed set in Ipomopsis aggregata

Summary We used powdered fluorescent dyes to estimate receipt of self vs. outcross pollen in the self-incompatible species Ipomopsis aggregata (Polemoniaceae). Flowers on small and large plants received equal amounts of outcross pollen, whereas flowers on large plants received more self pollen, so the proportion of self pollen delivered through geitonogamy increased with plant size. In natural populations emasculation of all flowers on a plant raised average seed set per flower from 5.19 to 6.99 and also raised fruit set, though not significantly. From these results one expects a negative correlation between plant size and seeds per flower. The opposite trend was observed in a sample of plants in the field, suggesting that deleterious effects of geitonogamy on female fecundity in large plants can be overruled by other factors such as size-related fruit or seed abortion. Results are discussed in relation to the evolution of gynodioecy.     

Effects of Inbreeding,Outbreeding, and Supplemental Pollen on the Reproduction of a Hummingbird‐pollinated Clonal Amazonian Herb

Understory herbs are an essential part of tropical rain forests, but little is known about factors limiting their reproduction. Many of these herbs are clonal, patchily distributed, and produce large floral displays of nectar‐rich 1‐d flowers to attract hummingbird pollinators that may transport pollen over long distances. The aim of this study was to investigate the effects of clonality, cross‐proximity, and patchy distribution on the reproduction of the hummingbird‐pollinated Amazonian herb Heliconia metallica. We experimentally pollinated flowers within populations with self‐pollen and with pollen of different diversity, crossed flowers between populations, and added supplemental pollen to ramets growing solitarily or in conspecific patches. Only flowers pollinated early in the morning produced seeds. Selfed flowers produced seeds, but seed number and mass were strongly reduced, suggesting partial sterility and inbreeding depression after selfing. Because of pollen competition, flowers produced more seeds after crosses with several than with single donor plants. Crosses between populations mostly resulted in lower seed production than those within populations, suggesting outbreeding depression. Ramets in patches produced fewer seeds than solitary ramets and were more pollen‐limited, possibly due to geitonogamy and biparental inbreeding in patches. We conclude that high rates of geitonogamy due to clonality and pollen limitation due to the short receptivity of flowers and patchy distribution constrain the reproduction of this clonal herb. Even in unfragmented rain forests with highly mobile pollinators, outbreeding depression may be a widespread phenomenon in plant reproduction.     

Pollen limitation and distance-dependent fecundity in females of the clonal gynodioecious herb Glechoma hederacea (Lamiaceae)

Summary Pollen movement is often restricted in natural populations, and insufficient pollination is a potential constraint on sexual reproduction in outcrossing species. Seed-set should decrease with increased distance from the pollen source in outcrossing plants. This prediction was tested using females of the clonal, gynodioecious herb Glechoma hederacea in three natural populations. In controlled pollinations, both hermaphrodites and females had similar high percentages of fruit-set and seed-set. In a natural population where a female clone was isolated from the nearest hermaphroditic clone by c. 100 m, fruit-set was low (1%). In another population where hemaphroditic clones were rare and female clones had a patchy distribution, fruit-and seed-set in females were pollen-limited and decreased with increased distance from the nearest pollen source. The estimated mean pollen dispersal distance was 5.9 m when calculated on fruit-set and 5.3 m when calculated on seed-set. The most frequent pollinators were bumblebees. The mean and median distances moved by pollinators between ramets were 0.13 m and 0.05 m. In a third population where female clones were isolated from the nearest hermaphrodites by more than 200 m, fruit-set was 0%. After introduction of 16 hermaphroditic ramets in the center of the female clone, fruit-set varied between 0% and 100% in individual female ramets. Fruit-set decreased with increased distance from the pollen source. The mean and median pollen movement distances were 1.06 m and 0.54 m.     

Experimental analysis of mating patterns in a clonal plant reveals contrasting modes of self‐pollination

Hermaphrodite plants commonly practice self‐fertilization (selfing), but the mechanisms responsible vary depending on the mode of self‐pollination, pollinator behavior, and degree of clonality. Whether selfing occurs within (autogamy) or between flowers (geitonogamy) is of evolutionary significance because their fitness consequences differ. We used floral manipulations and genetic markers to determine the relative contribution of autogamy and within‐ versus between‐ramet geitonogamy to the selfing rate of the bumblebee‐pollinated, clonal herb Aconitum kusnezoffii. Data on flowering phenology and bumblebee foraging were also collected to determine opportunities for different modes of self‐pollination. Autogamy accounted for only 12% of the selfing rate with the remainder resulting from geitonogamy. Whole‐ramet emasculation of clones with multiple ramets reduced selfing by 78%, indicating that within‐ramet geitonogamy contributed significantly (68%) to total selfing. Selfing of single‐ramet plants was 44% less than multiple‐ramet plants, indicating that the contribution of between‐ramet geitonogamy was substantially less (20%) than within‐ramet geitonogamy, probably because of bumblebee foraging behavior. Our results demonstrate for the first time in a clonal plant that within‐ramet geitonogamy is substantially greater than between‐ramet geitonogamy and highlight the importance of considering the influence of clonal architecture and pollinator foraging on modes of self‐pollination.     

The impact of extensive clonal growth on fine-scale mating patterns: a full paternity analysis of a lily-of-the-valley population (Convallaria majalis)

Background and Aims

The combination of clonality and a mating system promoting outcrossing is considered advantageous because outcrossing avoids the fitness costs of selfing within clones (geitonogamy) while clonality assures local persistence and increases floral display. The spatial spread of genetically identical plants (ramets) may, however, also decrease paternal diversity (the number of sires fertilizing a given dam) and fertility, particularly towards the centre of large clumped clones. This study aimed to quantify the impact of extensive clonal growth on fine-scale paternity patterns in a population of the allogamous Convallaria majalis.

Methods

A full analysis of paternity was performed by genotyping all flowering individuals and all viable seeds produced during a single season using AFLP. Mating patterns were examined and the spatial position of ramets was related to the extent of multiple paternity, fruiting success and seed production.

Key Results

The overall outcrossing rate was high (91 %) and pollen flow into the population was considerable (27 %). Despite extensive clonal growth, multiple paternity was relatively common (the fraction of siblings sharing the same father was 0·53 within ramets). The diversity of offspring collected from reproductive ramets surrounded by genetically identical inflorescences was as high as among offspring collected from ramets surrounded by distinct genets. There was no significant relationship between the similarity of the pollen load received by two ramets and the distance between them. Neither the distance of ramets with respect to distinct genets nor the distance to the genet centre significantly affected fruiting success or seed production.

Conclusions

Random mating and considerable pollen inflow most probably implied that pollen dispersal distances were sufficiently high to mitigate local mate scarcity despite extensive clonal spread. The data provide no evidence for the intrusion of clonal growth on fine-scale plant mating patterns.     

Self‐incompatibility and post‐fertilization maternal regulation cause low fecundity in Aegle marmelos (Rutaceae)

Outbreeding confers an evolutionary advantage, and flowering plants have evolved a variety of contrivances for its maximization. However, neither fruit set nor seed set is realized to its fullest potential for a variety of reasons. The causes of low flower to fruit and seed to ovule ratios were investigated in naturally occurring bael trees (Aegle marmelos) at two sites for three seasons. The study established that the mass effect of synchronized flowering attracted a variety of insect pollinators to the generalist flowers; Apis dorsata was the most efficient pollinator. The seed to ovule ratio was low despite high natural pollination efficiency (c. 2400 pollen per stigma). Although pollination‐induced structural and histochemical changes in the style allowed many (9.5 ± 2.1) pollen tubes to grow, only cross‐pollen tubes could grow through the style. Gametophytic self‐incompatibility, manifested in the stylar zone, resulted in a significantly slower growth rate of self‐pollen tubes. The occurrence of obligate self‐incompatibility, coupled with increased self‐pollen deposition (geitonogamy), caused a significant number of flowers to abort. Fruit retention in the trees declined from 40% to 12% as a result of abortion of fruits at different stages of development. The number of mature fruits on a tree was negatively correlated (r = ?0.82) with their size. It is inferred that low natural fecundity in A. marmelos is primarily a result of obligate self‐incompatibility and strong post‐fertilization maternal regulation of allocation of resources to the developing fruits. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 572–585.     

The consequences of clone size for paternal and maternal success in domestic apple (Malus x domestica)

Clonal growth in plants can increase pollen and ovule production per genet. However, paternal and maternal reproductive success may not increase because within-clone pollination (geitonogamy) can reduce pollen export to adjacent clones (pollen discounting) and pollen import to the central ramets (pollen limitation). The relationship between clone size and mating success was investigated using clones of Malus × domestica at four orchards (blocks of 1-5 rows of trees). For each block, maternal function was measured as fruit and seed set in all rows and paternal function as siring rate estimated from isozyme profiles in the first row of the adjacent block. Expected relations between reproductive success and clone size were generated from simulations and data on pollen dispersal in this species. Siring rate per clone averaged 70% and did not increase significantly with block size, consistent with simulations of pollen dispersal under pollen discounting. Simulations also indicated that the ratio of compatible to incompatible pollen received by a tree should decline with increased block size and from the periphery to the center of blocks. However, female function was not significantly reduced among block sizes or within blocks. The results suggest that paternal function may be more sensitive to the effects of clonality than female function.     

Patterns of Solidago altissima ramet growth and mortality: the role of below-ground ramet connections

Summary For the rhizomatous perennial, Solidago altissima, I identified clonal fragments in the field, mapped ramet spatial locations, and documented patterns of ramet recruitment, growth, and mortality. Parent ramet size influenced the size and number of daughter ramets produced, and small ramets had lower survivorship and fecundity than large ramets. Similarly, small rhizomes tended to develop into small ramets, and ramets that survived to produce daughter ramets had longer parent-daughter rhizome connections than ramets that did not survive. In addition, most ramets that died during the growing season were connected to (genetically identical) ramets that persisted. There were large size inequalities among rhizomes, ramets, and clonal fragments. Inequalities in the size of ramets increased during the early part of the growing season, then decreased at the end of the season; similar patterns were observed for the growth of clonal fragments. In both instances, the decrease in size inequality could be attributed to the mortality of small individuals (ramets or clonal fragments). I found little evidence that ramet size hierarchies were structured by intraspecific competition. For example, path analyses and randomization tests indicated that size variation among S. altissima ramets was influenced little by the size of their near neighbors (but was influenced by parent size and rhizome size). In addition, within-season variation for the relative size and growth rate of individual ramets led to poor correlations between early and final ramet size; this result suggests that there was no stable hierarchy of dominant and suppressed ramets. I discuss implications of my results for contrasting interpretations of clonal plant population dynamics.     

佛坪国家级自然保护区秦岭箭竹克隆结构的SSR分析

利用SSR分子标记技术分析了佛坪国家级自然保护区秦岭箭竹(Fargesia qinlingensis)的克隆多样性和克隆结构,以探讨小尺度范围内秦岭箭竹自然居群遗传变异的分布特征,对该种开花特性、高山地区生态环境维护和大熊猫的保护提供重要依据。结果表明7对SSR引物共扩增出79个位点,其中多态性位点77个,多态位点百分率(PPB)为97.47%。秦岭箭竹的142个分株共形成107个克隆,最大克隆可达5 m。克隆多样性略高于其他克隆植物的平均值(D=0.62,G/N=0.17,E=0.68),基因型比率(G/N)、Simpson指数(D)、平均克隆大小(N/G)和Fager均匀性指数(E)分别为0.7535、0.9680、1.3271和0.5109。克隆空间结构分析表明秦岭箭竹的克隆构型为密集型,各克隆呈镶嵌性分布,同一克隆的分株排列紧密。克隆聚类分析表明各克隆之间聚类不明显,总体上来自同一样地的克隆被聚为一类。空间自相关分析显示在空间距离为36 m范围内,分株比基株有更显著的空间遗传结构,空间自相关系数r的取值范围分别为0.084—0.626和0.024—0.288,说明克隆繁殖在一定程度上限制了空间遗传结构的范围。样地内秦岭箭竹个体在空间距离小于44 m时存在显著的正相关空间结构,特别是在4 m处表现出最大的空间自相关系数(r=0.626),表明空间距离相距4 m内的个体最有可能属于同一克隆,4 m比5 m更能表现出清晰的克隆结构,X-轴截距为52.280,代表了秦岭箭竹不规则克隆的平均最小长度。秦岭箭竹的克隆多样性和克隆结构与初始苗补充、花粉散播方式和微环境差异有关。     

Characteristics of self-incompatibility in Schlumbergera truncata and S. x buckleyi (Cactaceae)

The influence of self-incompatibility (SI) on fruit set, seed set, and pollen tube growth was investigated in Schlumbergera truncata (Haworth) Moran and S.xbuckleyi (T. Moore) Tjaden. Four Schlumbergera clones were crossed in a complete diallel to verify the presence of SI. Fruit did not set when the clones were selfed or when two of the clones were crossed reciprocally, but all other outcrosses yielded fruit which contained 100–200 seeds each. Compatible outcrosses were characterized by large numbers of pollen tubes in the style and ovary cavity at 72 h after pollination. When pistils were selfed or incompatibly crossed, pollen tubes were inhibited in the upper third of the style and few pollen tubes reached the base of the style by 72 h after pollination. Schlumbergera exhibits several characteristics often associated with sporophytic SI systems (tricellular pollen and dry stigmas with elongate papillae), together with those commonly observed in gametophytic SI systems (stylar inhibition of incompatible pollen tubes and absence of reciprocal differences in outcrosses).Publication no. 3174 of the Massachusetts Agricultural Experiment Station     

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.     

克隆植物沙拐枣的母株和分株对风蚀沙埋的生理生态响应

以中国荒漠区优良的防风固沙克隆灌木沙拐枣为对象,研究了长期风蚀、沙埋环境下沙拐枣母株和克隆分株的同化枝对环境异质性的响应。结果发现:(1)风蚀母株、风蚀分株的叶绿素含量、净光合速率、气孔导度、蒸腾速率、胞间CO2浓度和水分利用效率只有沙埋分株的一半左右,导致同化枝的长度、数量、簇数也仅是沙埋分株的一半,而且风蚀母株的果实宽和果实长也都最小,但浅沙埋有利于沙拐枣的生长和繁殖,表明严峻的风蚀对母株和分株的生长与繁殖都产生了胁迫,但浅沙埋有利于沙拐枣的生长和繁殖。(2)风蚀母株倒伏后同化枝的形态特征是基部优于中部优于顶部,表明严峻风蚀下母株的死亡是从顶部-中部-底部逐渐舍弃的过程。(3)母株的全部根系以及风蚀水平根全部裸露在外但依然能够存活,间接证明沙拐枣克隆整合的方向性——不仅可在分株间进行传递,分株-母株间也可进行传递,否则遭受严峻风蚀胁迫的母株和克隆分株会直接死亡。本研究结果为沙拐枣克隆生长对风沙环境的生态适应机制提供了基础,也是对植物克隆生态学在自然异质环境中研究缺乏的有效补充。     

A ballistic pollen dispersal system influences pollination success and fruit‐set pattern in pollinator‐excluded environments for the endangered species Synaphea stenoloba (Proteaceae)

The critically endangered Synaphea stenoloba (Proteaceae) has numerous scentless flowers clustered in dense inflorescences and deploys a ballistic pollen ejection mechanism to release pollen. We examined the hypothesis that active pollen ejection and flowering patterns within an inflorescence influence the reproductive success (i.e. fruit formation) of individual flowers within or among inflorescences of S. stenoloba in a pollinator‐excluded environment. Our results showed that: (1) no pollen grains were observed deposited on the stigma of their own flower after the pollen ejection system was manually activated, indicating self‐pollination within an individual flower is improbable in S. stenoloba; (2) fruit set in the indoor open pollination treatment and the inflorescence‐closed pollination treatment indicated that S. stenoloba is self‐compatible and pollen ejection can potentially result in inter‐floral pollination success; (3) fruit set in the inflorescence‐closed pollination treatment was significantly lower than that of indoor open pollination, indicating within‐ and between‐flower pollination events in an inflorescence are most likely limited, with pollination between inflorescences providing the highest reproductive opportunity; and (4) analysis of the spatial distribution of cumulative fruit set on inflorescences showed that pollen could reach any flower within an inflorescence and there was no functional limitation on seed set among flowers located at various positions within the inflorescence. These data suggest that the pollen ejection mechanism in S. stenoloba can enhance inter‐plant pollination in pollinator‐excluded environments and may suggest adaptation to pollinator scarcity attributable to habitat disturbance or competition for pollinators in a diverse flora. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170 , 59–68.     

Pollination thoroughness and maternal yield regulation in wild radish,Raphanus raphanistrum (Brassicaceae)

Summary Under conditions where resources are limited, there are often negative correlations between components of maternal yield, or between fruit and flower production. Pollination, in turn, may vary among individuals and influence total maternal expenditure. We examined the impact of variation in pollination thoroughness upon yield components and overall plant growth in wild radish (R. raphanistrum) plants grown in the greenhouse. Plants received different pollination treatments in which 0% to 100% of all flowers produced were hand-pollinated. Fruit set was increased by hand-pollination, but rarely exceeded 30%, even when more than 50% of the flowers were pollinated. Plants receiving more thorough pollination or having greater proportion fruit set produced significantly smaller seeds. Seed number per fruit was not influenced by pollination treatment. Mean values of yield components and interactions between components often varied among plants from different maternal families. Increasing pollination thoroughness also resulted in dramatic decreases in flower production. If male fitness is related to flower number, there may be a tradeoff between maternal fecundity and successful pollen export operating at the whole-plant level in this species.