Self-pollen Deposition Research Articles

An evolutionary disruption of the buzz pollination syndrome in neotropical montane plants.

Under pollinator limitations, specialized pollination syndromes may evolve toward contrasting responses: a generalized syndrome with increased pollinator attraction, pollinator reward, and pollen transfer capacity; or the selfing syndrome with increased self-pollen deposition, but reduced pollinator attraction and pollen transfer capacity. The buzz-pollination syndrome is specialized to explore female vibrating bees as pollinators. However, vibrating bees become less-active pollinators at montane areas of the Atlantic Forest (AF) domain. This study investigated whether the specialized buzz-pollination syndrome would evolve toward an alternative floral syndrome in montane areas of the AF domain, considering a generalized and the selfing syndromes as alternative responses. We utilized a lineage within the buzz-pollinated Miconia as study system, contrasting floral traits between montane AF-endemic and non-endemic species. We measured and validated floral traits that were proxies for pollinator attraction, reward access, pollen transfer capacity, and self-pollen deposition. We inferred the evolution of floral trait via phylogenetic comparative methods. AF-endemic species have selectively evolved greater reward access and more frequently had generalist pollination. Nonetheless, AF-endemic species also have selectively evolved toward lower pollen transfer capacity and greater self pollination. These patterns indicated a complex evolutionary process that has jointly favored a generalized and the selfing syndromes. The buzz pollination syndrome can undergo an evolutionary disruption in montane areas of the AF domain. This floral syndrome is likely more labile than often assumed, allowing buzz-pollinated plants to reproduce in environments where vibrating bees are less-reliable pollinators.

Effect of robbing intensity on reproductive success of Symphytum officinale (Boraginaceae).

The intervention of nectar robbers in plant pollination systems will cause some pollinators to modify their foraging behavior to act as secondary robbers, consequently adopting a mixed foraging strategy. The influence of nectar robbing on pollinator behavior may be affected by spatio-temporal difference of robbing intensity, and consequently, may have different effects on the pollination of host plants. However, whether and how the nectar robbing might influence pollinators under different robbing intensity still needs further investigation. In this study, Symphytum officinale was used to detect the effect of nectar robbers on pollinators under different robbing intensity as well as their effects on plant reproductive success. Six robbing levels and three bumblebees with mixed foraging behaviors were used to evaluate the effect of different robbing intensity on pollinator behavior, visitation rate, flower longevity and pollen deposition. Our results indicated that the robbing rate increased gradually with the proportion of robbed flowers, but which did not affect the frequency of legitimate visits. The increase of robbing rate promoted the corolla abscission, and then enhanced the self-pollen deposition, but which had no significant effect on cross-pollen deposition. These results indicate that the overall fitness of S. officinale was improved by combined self and cross-pollination modes when visited by both pollinators and nectar robbers simultaneously. Although nectar robbing is not uncommon, its consequences for pollination in the interaction web have not been well studied. Our results emphasize the significance of indirect impacts in mediating the adaptive outcomes of species interactions.

Fine-scale genetic structure in the orchid Gymnadenia conopsea is not associated with local density of flowering plants.

Density-dependent pollinator visitation can lead to density-dependent mating patterns and within-population genetic structure. In Gymnadenia conopsea, individuals in low-density patches receive more self pollen than individuals in high-density patches, suggesting higher relatedness at low density. Ongoing fragmentation is also expected to cause more local matings, potentially leading to biparental inbreeding depression. To evaluate whether relatedness decreases with local density, we analyzed 1315 SNP loci in 113 individuals within two large populations. We quantified within-population genetic structure in one of the populations, recorded potential habitat barriers, and visualized gene flow using estimated effective migration surfaces (EEMS). We further estimated the magnitude of biparental inbreeding depression that would result from matings restricted to within 5 m. There was no significant relationship between local density and relatedness in any population. We detected significant fine-scale genetic structure consistent with isolation by distance, with positive kinship coefficients at distances below 10 m. Kinship coefficients were low, and predicted biparental inbreeding depression resulting from matings within the closest 5 m was a modest 1-3%. The EEMS suggested that rocks and bushes may act as barriers to gene flow within a population. The results suggest that increased self-pollen deposition in sparse patches does not necessarily cause higher selfing rates or that inbreeding depression results in low establishment success of inbred individuals. The modest relatedness suggests that biparental inbreeding depression is unlikely to be an immediate problem following fragmentation of large populations. The results further indicate that habitat structure may contribute to governing fine-scale genetic structure in G. conopsea.

Reduced pollination in bilateral flowers could reflect selfing avoidance

The multiple evolutionary transitions from radial to bilateral flower symmetry recorded across the angiosperms have been related to pollinator specialization and the promotion of outcross pollen transfer. In particular, the limited number of physical orientations associated with bilaterality results in a tighter morphological match between a flower and its specialized pollinators, which can enhance the transfer of incoming pollen while preventing autonomous, or even pollinator-mediated self-pollination when associated with dichogamy. In contrast, radial flowers have various planes of symmetry, which may allow pollination by a wider range of pollinators at the cost of more promiscuous pollen transfer. Here we used data from the literature and our own on pollen production, ovule production, and stigmatic pollen load compiled for 171 angiosperm species distributed worldwide to assess differences in pollen transfer in relation to flower symmetry within a phylogenetic comparative framework. We did not find evidence that bilateral and radial flowers differ in average pollen and ovule production. However, bilateral flowers tended to receive less pollen on their stigmas. This decrease in pollen receipt was most apparent among self-compatible species after accounting for the covarying effects of flower size and ovule production. Therefore, our findings indicate that the evolution of flower bilaterality, a trait we found is strongly associated with self-compatibility, comes at a quantitative pollination cost. However, they support the view that a bilateral flower morphology may provide the benefit of limiting self-pollen deposition, particularly in the absence of an alternative mechanism of selfing avoidance.

High self-paternity levels and effects of fertilised-seed number on size of strawberry fruit.

Cross-pollination can increase fruit production in both self-incompatible and self-compatible fruit crops. However, it is often unclear what proportions of the fruit crop result from cross-pollination. We quantified the proportion of cross-pollinated seeds and the proportion of fertilised seeds in two strawberry cultivars, Red Rhapsody and Sundrench, at increasing distances from a cross-pollen source. We assessed whether fully self-pollinated fruit and partly cross-pollinated fruit differed in fruit size, colour, firmness, Brix and acidity. We also assessed whether fruit size and quality were affected by the number or percentage of fertilised seeds. Almost all seeds of both cultivars resulted from self-pollination (~98%), even at only 1 m from a cross-pollen source. Distance from a cross-pollen source did not affect the proportion of partly cross-pollinated fruit or the proportion of cross-pollinated seeds per fruit. The mass and diameter of fully self-pollinated Sundrench fruit, and the redness and Brix of fully self-pollinated Red Rhapsody fruit, were higher than partly cross-pollinated fruit. Fruit mass, length and diameter increased, and acidity decreased, with increasing numbers of fertilised seeds in both cultivars. Fruit mass also increased with the percentage of fertilised seeds. Our results show that cross-pollination was not required for Red Rhapsody and Sundrench fruit production, and that cross-pollination was a rare occurrence even close to cross pollen source. Self-pollen deposition on stigmas is required to maximise the number of fertilised seeds, and consequently fruit size and quality. Our research indicates that bees improve strawberry fruit size by increasing the number of stigmas that receive pollen. Our results suggest that placing bee hives on strawberry farms during flowering and establishing nearby pollinator habitat to support wild pollinators could improve strawberry yield and fruit quality.

The influence of self-pollen deposition on female reproductive success in a self-incompatible plant, Akebia quinata.

Geitonogamy is inevitable in hermaphrodite and monecious. Even for self-incompatible species, the negative effects of self-pollen are unavoidable when geitonogamous or self-mating occurs. However, the influence of self-pollen on consecutive development of flowers (e.g., fruiting and seeding) was seldom evaluated. Here, the self-incompatible monecious species, Akebia quinata, was used to estimate the influence of self-pollen deposition. We evaluated the extent of pollen limitation and geitonogamous mating under natural conditions by count of stigmatic pollen load and pollen tracking experiment. Hand pollination with different amount and combinations of self vs. cross pollen grains was applied to detect the response of fruit and seed set. The results showed that geitonogamy and pollen limitation occurred under natural conditions in A. quinata. Carpel numbers, ratio of self- and cross-pollen, and the interactive effect of ratio of self- and cross-pollen and total mixed pollen numbers, and not total pollen grain number, determined the effect of self-pollen on female reproductive success. The effect of self-pollen depended on its intensity. In general, the transfer of self-pollen significantly affected young fruit set. However, a little self-pollen together with cross-pollen did not reduce young fruit production. Although self-incompatible plants have evolved physiological mechanisms that reduce self-fertilization, our results provide new insights into the effects of self-pollen and the adaptive significance of self-incompatible monecious species.

Why honeybees are poor pollinators of a mass-flowering plant: Experimental support for the low pollen quality hypothesis.

PremiseHoneybees dominate the flower‐visitor assemblages of many plant species, yet their efficiency in terms of the quality of pollen delivered to stigmas is largely unknown. We investigated why honeybees are poor pollinators of Aloe ferox, a self‐incompatible succulent treelet with large numbers of flowers. Honeybees are very frequent visitors to flowers of this species, yet contribute very little to seed production.MethodsWe assessed pollen loads on honeybees, studied their visitation behavior, selectively excluded birds from plants to determine direct effects of bees on pollen deposition, seed set, and ovule abortion, and used a novel “split‐pollinator” method to test whether honeybees deposit mainly low‐quality self pollen. For the latter, we captured honeybees, and with their existing pollen loads, used them to either pollinate virgin flowers on the plant on which they were caught or to pollinate virgin flowers on different plants.ResultsHoneybees cumulatively deposit as much pollen on stigmas as do birds, but our experiments showed that the pollen deposited by honeybees is mostly low‐quality self pollen that leads to substantial ovule discounting and depressed seed set.ConclusionsLack of movement among A. ferox plants during individual honeybee foraging bouts is the most likely explanation for their deposition of low‐quality self pollen on stigmas. The “split‐pollinator” method is a simple and cost‐effective technique to test the quality of pollination.

Changes in female function and autonomous selfing across floral lifespan interact to drive variation in the cost of selfing.

PremiseMorphological and developmental changes as flowers age can impact patterns of mating. At the same time, direct or indirect costs of floral longevity can alter their fitness outcomes. This influence has been less appreciated, particularly with respect to the timing of selfing. We investigated changes in stigma events, autonomous selfing, outcross seed set capacity, and autofertility—a measure representing the potential for reproductive assurance—across floral lifespan in the mixed‐mating biennial Sabatia angularis.MethodsWe examined stigma morphology and receptivity, autonomous self‐pollen deposition, and seed number and size under autonomous self‐pollination and hand outcross‐pollination for flowers of different ages, from 1 d of female phase until 14 d. We compared autonomous seed production to maximal outcross seed production at each flower age to calculate an index of autofertility.ResultsThe stigmatic lobes begin to untwist 1 d post anthesis. They progressively open, sextend, coil, and increase in receptivity, peaking or saturating at 8–11 d, depending on the measure. Autonomous seed production can occur early, but on average remains low until 6 d, when it doubles. In contrast, outcross seed number and size start out high, then decline precipitously. Consequently, autofertility increases steeply across floral lifespan.ConclusionsChanges in stigma morphology and receptivity, timing of autonomous self‐pollen deposition, and floral senescence can interact to influence the relative benefit of autonomous selfing across floral lifespan. Our work highlights the interplay between evolution of floral longevity and the mating system, with implications for the maintenance of mixed mating in S. angularis.

Pollination adaptations of group-by-group stamen movement in a meadow plant with temporal floral closure

Floral sexual organ (stamen and pistil) movements are selective adaptations that have different functions in male-female reproduction and the evolution of flowering plants. However, the significance of stamen movements in the spatial–temporal function and separation of male and female organs has not been experimentally determined in species exhibiting floral temporal closure. The current study investigated the role of slow stamen (group-by-group) movement in male-female sexual function, and the effect of stamen movement on pollen removal, male-male and male-female interference, and mating patterns of Geranium pratense, a plant with temporal floral closure. This species uses stamen group-by-group movement and therefore anther-stigma spatial–temporal separation. Spatial separation (two whorls of stamen and pistil length) was shown to be stronger than temporal separation. We found that stamen movements to the center of the flower increase pollen removal, and the most common pollinators visited more frequently and for longer durations during the male floral stage than during the female floral stage. Petal movements increased both self-pollen deposition rate and sexual interference in G. pratense. The fruit and seed set of naturally and outcrossed pollinated flowers were more prolific than those of self-pollinated flowers. Group-by-group stamen movement, dehiscence of stamens, pistil movement, and male-female spatial–temporal functional separation of G. pratense before floral temporal closure may prevent male-female and stamen-stamen interference and pollen discounting, and may increase pollen removal and cross-pollination.

The role of lateral and vertical herkogamy in the divergence of the blue- and red-flowered lineages of Lysimachia arvensis.

Herkogamy, or anther-stigma separation, is known to reduce self-pollen deposition, but little is known about the relative efficacy of different modes or conformations of herkogamy. We assessed the effectiveness of vertical versus lateral herkogamy in preventing or promoting self-pollen deposition in the annual herb Lysimachia arvensis, a plant with lineages that differ in flower colour, and in which flowers first display lateral and then vertical herkogamy. Because mating between the two lineages compromises fitness through the production of low-quality hybrid offspring, we tested the prediction that individuals sampled from sites occupied by both lineages should have flowers that promote autonomous self-pollen deposition and self-fertilization as a result of selection to reduce deleterious reproductive interference. We characterized variation in herkogamy within and among 25 pure and mixed populations of L. arvensis in its European range and assessed the effectiveness of lateral versus vertical herkogamy in avoiding self-pollen deposition. Lateral herkogamy was more effective than vertical herkogamy in limiting self-pollen deposition. In the case of vertical herkogamy, only approach herkogamy was effective. Lineages showed consistent differences in herkogamy traits. In general, angles were smaller for blue than red flowers in most populations, and blue flowers showed approach herkogamy, while red flowers showed predominantly reverse herkogamy. In sympatry, the red lineage showed a reduction of both herkogamy traits while for the blue lineage only lateral herkogamy was reduced. Our results demonstrate that pollen deposition is affected not only by the degree but also the spatial conformation of herkogamy. They also highlight reduced herkogamy as a potential mechanism for promoting reproductive assurance under pollen limitation, as well as for avoiding reproductive interference between genetically divergent lineages.

Anther Rubbing, a New Mechanism That Actively Promotes Selfing in Plants.

Self-fertilization has recurrently evolved in plants, involving different strategies and traits and often loss of attractive functions, collectively known as the selfing syndrome. However, few traits that actively promote self-fertilization have been described. Here we describe a novel mechanism promoting self-fertilization in the Brassicaceae species Erysimum incanum. This mechanism, which we called "anther rubbing," consists of autonomous, repeated, and coordinated movements of the stamens over the stigma during flower opening. We have documented anther rubbing by time-lapse videos and experimentally show that it causes self-pollen deposition on stigmas and is sufficient to achieve maximal reproductive output in E. incanum. We predict that these movements should occur in species with limited inbreeding depression, and indeed we find that inbreeding depression in seed production is negligible in this species. While many studies have documented complex floral traits that promote outcrossing, the occurrence of anther rubbing demonstrates that plants can evolve elaborate and underappreciated adaptations to promote self-fertilization.

Timing is everything: Dichogamy and pollen germinability underlie variation in autonomous selfing among populations.

The evolution of multiple floral traits often underlies the transition from outcrossing to selfing. Such traits can influence the ability to self, and the timing at which selfing occurs, which in turn affects the costs of selfing. Species that display variation in autonomous selfing provide an opportunity to dissect the phenotypic changes that contribute to variability in the mating system. In a common garden, we measured dichogamy and herkogamy in 24 populations of the protandrous mixed-mating herb Campanula americana, and related these to autonomous fruit set (autonomy). We then measured the timing of self-pollen deposition and fruit production in populations with high and low autonomy, and determined whether pollen germinability across floral development contributes to variation in autonomy. Populations that transitioned more rapidly to female phase displayed elevated autonomous selfing, but herkogamy was unassociated with autonomous selfing. Selfing occurred more rapidly in highly autonomous populations because of greater self-pollen deposition early in female phase. Pollen germinability in low-autonomy populations remained constant across floral development, but in high-autonomy populations it increased after floral anthesis and was highest near the onset of female phase. Reduced dichogamy, elevated self-pollen deposition, and higher pollen germination late in male phase contribute to both earlier selfing and greater selfing. These traits vary among populations, likely reflecting past selection on the mating system. While delayed selfing bears fewer fitness costs, the evolution of earlier selfing may be favored if self-pollen availability decreases over floral development.

Estimating pollinator performance of visitors to the self-incompatible crop-plant <i>Brassica rapa</i> by single visit deposition and pollen germination: a comparison of methods.

Estimating the pollen-deposition effectiveness of flower visitors is fundamental to understanding their performance as pollinators. While estimates of visitation rates, pollen loads, and single visit deposition (SVD) are all useful proxies for performance, and so help to reveal the relative effectiveness of different visitors, none take into account the breeding system of the plants, or the quality of pollen deposited. Here we compare the performance of visitors to the self-incompatible plant Brassica rapa (turnip) using SVD and pollen germination. We also report the first use of the staining of Brassica rapa stigma papilla cells (known to reveal a specific reaction to self-pollen) to compare self-pollen deposition between insect visitors. We found that most of the pollen grains deposited by insect visitors (and therefore counted by SVD methods) were non-germinating self-pollen. A smaller proportion of grains were outcrossed and so germinated. There was also a significant positive relationship between environmental conditions (wind speed) and pollen deposition, but not pollen germination.Both methods identified Bombus spp. as the best-performing visitors on turnip flowers, followed by Eristalis spp., whereas performance estimates for Episyrphus balteatus and ‘other hoverflies’ were no higher than controls for both methods. This study provides further insight into the methodology for estimating pollinator performance, especially in plants when only cross-pollen can germinate.

Plasticity of floral longevity and floral display in the self-compatible biennial Sabatia angularis (Gentianaceae): untangling the role of multiple components of pollination.

Plasticity of floral traits in response to pollination can enable plants to maximize opportunities for pollen import and export under poor pollination conditions, while minimizing costs under favourable ones. Both floral longevity and display are key traits influencing pollination. While pollination-induced flower wilting is widely documented, we lack an understanding of the multifactorial complexity of this response, including the influence of other pollination components, costs of extended longevity and subsequent impacts on floral display. Plasticity of floral longevity was experimentally evaluated in Sabatia angularis in response to multiple pollination factors: pollen addition, removal, and source (self, single-donor outcross, multiple-donor outcross) and timing of pollination. Effects of pollen quantity were further evaluated by exploiting variation in autonomous self-pollen deposition. Delayed pollination costs were tested comparing seed set from early versus late pollinations. Finally, I compared floral display metrics (peak floral display, time to peak flower, flowering duration, mean flowering rate) between experimentally pollinated and control plants. Floral longevity was highly plastic in response to pollen addition and its timing, and the response was dose-dependent but insensitive to pollen source. Pollen removal tended to extend floral longevity, but only insofar as it precluded pollination-induced wilting via autonomous self-pollination. Under delayed pollination, the wilting response was faster and no cost was detected. Pollination further led to reduced peak floral displays and condensed flowering periods. Floral longevity and display plasticity could optimize fitness in S. angularis, a species prone to pollen limitation and high inbreeding depression. Under pollinator scarcity, extended floral longevities offer greater opportunities for pollen receipt and export at no cost to seed set, reproductive assurance via autonomous self-pollination and larger, more attractive floral displays. Under high pollinator availability, shortened longevities lead to smaller displays that should lower the risk of geitonogamy.

Self-pollination, style length development and seed set in self-compatible Asteraceae: evidence from Senecio vulgaris L.

Background : Variation in style length has been reported in Senecio vulgaris and has been associated with outcrossing rate. Aims : To determine if (i) long styles lack germinated pollen on stigmas left to self-pollinate, (ii) successful self-pollination causes styles to stop elongating and shrink in length and (iii) seed set increases with the amount of pollen deposited on stigmas. Methods : Determined germinated self-pollen on stigmas of long and short styles after auto-self-pollination; scored style length over 48 h in self-pollinated and non-pollinated florets; recorded seed set after placing different amounts of pollen on stigmas. Results : Most long-styled florets had zero or low amounts of germinated pollen on stigmas in contrast to most short-styled florets. Styles initially elongated to the same length in self-pollinated and non-pollinated florets, then shrank in length in self-pollinated florets while continuing to elongate in non-pollinated florets. Seed set increased with number of pollen grains deposited on stigmas. Conclusions : Successful self-pollen deposition and/or germination on stigmas of S. vulgaris are indicated by presence of short styles, whereas the opposite is indicated by presence of long styles in florets left to self-pollinate. Self-pollination causes styles to shrink after initially elongating. Seed set is dependent on the amount of pollen deposited on stigmas.

The mean and variability of a floral trait have opposing effects on fitness traits.

Floral traits are essential for ensuring successful pollination and reproduction in flowering plants. In particular, style and anther positions are key for pollination accuracy and efficiency. Variation in these traits among individuals has been well studied, but less is known about variation within flowers and plants and its effect on pollination and reproductive success. Style deflexion is responsible for herkogamy and important for pollen deposition in Passiflora incarnata. The degree of deflexion may vary among stigmas within flowers as well as among flowers. We measured the variability of style deflexion at both the flower and the plant level. The fitness consequences of the mean and variation of style deflexion were then evaluated under natural pollination by determining their relationship to pollen deposition, seed production and average seed weight using structural equation modelling. In addition, the relationship between style deflexion and self-pollen deposition was estimated in a greenhouse experiment. We found greater variation in style deflexion within flowers and plants than among plants. Variation of style deflexion at the flower and plant level was positively correlated, suggesting that variability in style deflexion may be a distinct trait in P. incarnata. Lower deflexion and reduced variation in that deflexion increased pollen deposition, which in turn increased seed number. However, lower styles also increased self-pollen deposition. In contrast, higher deflexion and greater variability of that deflexion increased variation in pollen deposition, which resulted in heavier seeds. Variability of style deflexion and therefore stigma placement, independent from the mean, appears to be a property of individual P. incarnata plants. The mean and variability of style deflexion in P. incarnata affected seed number and seed weight in contrasting ways, through the quantity and potentially quality of pollen deposition. This antagonistic selection via different fitness components may maintain diverse style phenotypes.

Self-compatibility and floral traits adapted for self-pollination allow homostylous Nymphoides geminata (Menyanthaceae) to persist in marginal habitats

Many stylar polymorphic plants are equipped with reciprocal herkogamy and incompatibility systems so that each floral morph within a population can only mate with the compatible reciprocal morph. In the ecological or geographical margins of the range of stylar polymorphic taxa, homostyly may evolve in response to a lack of mate and/or poor pollinator visitation. This prediction is tested here by comparing floral traits, seed production without pollinators (i.e. autonomously), pollinator activity and ploidy level between the homostylous Nymphoides geminata that colonise geographically marginal habitats and its stylar polymorphic and self-incompatible closely related species N. montana that typically occur in natural water bodies in south-eastern Australia. Autonomous self-fertilisation is detected in glasshouse. Homostyly in N. geminata is also associated with floral adaptation to autonomous self-pollen deposition. There is a reduction in size and attractiveness of homostylous floral organs: over 80 % shorter stigma–anther separations and lower pollen:ovule ratio than the polymorphic flowers. The SEM images reveal that homostylous flowers possess similar pollen and stigma papillae morphology to those of short styles and long styles of the polymorphic flowers, respectively. Nymphoides geminata may be an example of recombinant homostyly: combining the pollen compatibility features of one morph with the stylar compatibility features of the reciprocal morph. Results of floral visitor observations and open pollinations indicate the ability of homostylous plants to reproduce successfully despite pollinator scarcity. Somatic chromosome counts indicate a possible lack of relationship between ploidy level and sexual system. Homostylous condition should provide an opportunity for reproductive assurance in environments with unreliable pollinators.

Patterns of pollen removal and deposition in Polemonium brandegeei (Polemoniaceae): the role of floral visitors, floral design and sexual interference.

The arrangement, colour, shape and size of floral parts (collectively floral design) have evolved primarily to promote mating success via animal-mediated pollen transfer. Although numerous studies have examined variation in pollinator assemblages, relatively few have examined patterns of pollen removal and deposition in the presence of fluctuating pollinators and ineffective floral visitors; therefore, net pollen removal and deposition by entire visitor assemblages are unclear. We studied the timing (diurnal or nocturnal) and effects of floral traits on pollen removal and deposition under a dynamic visitor assemblage of Polemonium brandegeei. We quantified pollen grains remaining in anthers (pollen removal) and deposited on stigmas (pollen deposition) of plants visited during either the day (07:30-20:00h) or night (20:30-07:30h) in natural populations over two flowering seasons. Pollen removal and deposition occurred both diurnally and nocturnally during our study. Increased diurnal removal and deposition coincided with peak floral visitations in 2006. This increase in pollen removal and deposition may reflect increased visits by pollen consumers, effective hawkmoth pollinators and increased self-pollen deposition due to hot, dry weather. Nonlinear effects of style length significantly affected pollen removal, with less pollen remaining in flowers with intermediate style lengths. Pollen deposition was more complex, with herkogamy and anther height affecting deposition. Further, close proximity of stigmas and anthers increased the potential for sexual interference between pollen removal and deposition. Overall, flower visitations and pollen removal and deposition varied between years and populations, but sex organ placement consistently influenced the removal and deposition of pollen.

Self-incompatibility and post-fertilization maternal regulation cause low fecundity inAegle 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.

Variation in style length and anther–stigma distance in Ixiolirion songaricum (Amaryllidaceae)

Variation in style length and anther–stigma distance (ASD) was investigated in 18 populations of Ixiolirion songaricum, an early-spring ephemeral perennial herb in northern Xinjiang. The effect of ASD on autonomous self-pollen deposition and seed set was assessed using bagging experiments. Seed production under autonomous self-pollination, cross-pollination and natural pollination was determined by manual pollination experiments. Stigmas of multiple flowers within an individual were sandwiched between two separate anther levels, and no individuals possessed approach or reverse herkogamy. Style length varied continuously among flowers, resulting in a wide range of variation in ASD. There were fewer flowers with a small ASD than with a large ASD in populations. ASD was negatively correlated with autonomous self-pollen deposition and seed set, but it was positively correlated with natural pollination seed set. Seed production under natural pollination was higher than that under autonomous self-pollination, but it was smaller than that under cross-pollination. Variation in style length and ASD among flowers was not caused by style or stamen elongation in I. songaricum, which differ from other species reported in literature that have continuous variation in style length and ASD.