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1.  Effect of Pollination on Floral Longevity and Costs of Delaying Fertilization in the Out-crossing Polygala vayredae Costa (Polygalaceae) 
Annals of Botany  2008;102(6):1043-1048.
Background and Aims
The effect of pollination on flower life span has been widely studied, but so far little attention has been paid to the reproductive consequences of delayed pollination in plants with long floral life spans. In the present study, Polygala vayredae was used to answer the following questions. (1) How does male and female success affect the floral longevity of individual flowers? (2) How does delaying fertilization affect the female fitness of this species?
Floral longevity was studied after experimental pollinations involving male and/or female accomplishment, bagging and open pollination. The reproductive costs of a delay in the moment of fertilization were evaluated through fruit set, seed–ovule ratio and seed weight, after pollination of flowers that had been bagged for 2–18 d.
Key Results
Senescence of the flowers of P. vayredae was activated by pollen reception on the stigmatic papillae, while pollen removal had no effect on floral longevity. Nonetheless, a minimum longevity of 8 d was detected, even after successful pollination and pollen dissemination. This period may be involved with the enhancement of male accrual rates, as the female accomplishment is generally achieved after the first visit. Floral life span of open-pollinated flowers was variable and negatively correlated with pollinator visitation rates. Delayed pollination had a major impact on the reproductive success of the plant, with fruit set, seed–ovule ratio and seed weight being significantly diminished with the increase of flower age at the moment of fertilization.
A strong relationship between pollination and floral longevity was observed. Flowers revealed the ability to extend or reduce their longevity, within some limits, in response to the abundance of efficient pollinators (i.e. reproductive fulfilment rates). Furthermore, with scarce or unpredictable pollinators, a long floral life span could maintain the opportunity for fertilization but would also have reproductive costs on production of offspring. Reduced female fitness late in the flower's life could shift the cost–benefit balance towards a shorter life span, partially counteracting the selection for longer floral life span potentially mediated by scarce pollination services.
PMCID: PMC2712401  PMID: 18829587
Delayed pollination; endemic species; flower longevity; life span; pollen limitation; pollination; pollinator scarcity; Polygala vayredae; Polygalaceae; reproductive consequences; secondary pollen presentation
2.  Pollination and late-acting self-incompatibility in Cyrtanthus breviflorus (Amaryllidaceae): implications for seed production 
Annals of Botany  2010;106(4):547-555.
Background and Aims
Animal pollination is typically an uncertain process that interacts with self-incompatibility status to determine reproductive success. Seed set is often pollen-limited, but species with late-acting self-incompatibility (SI) may be particularly vulnerable, if self-pollen deposition results in ovule discounting. Pollination is examined and the occurrence of late-acting SI and ovule discounting assessed in Cyrtanthus breviflorus.
The pollination system was characterized by observing floral visitors and assessing nectar production and spectral reflectance of flowers. To assess late-acting SI and ovule discounting, growth of self- and cross-pollen tubes, and seed set following open pollination or hand pollination with varying proportions of self- and cross-pollen, were examined.
Key Results
Native honeybees Apis mellifera scutellata pollinated flowers as they actively collected pollen. Most flowers (≥70 %) did not contain nectar, while the rest produced minute volumes of dilute nectar. The flowers which are yellow to humans are visually conspicuous to bees with a strong contrast between UV-reflecting tepals and UV-absorbing anthers and pollen. Plants were self-incompatible, but self-rejection was late-acting and both self- and cross-pollen tubes penetrated ovules. Seed set of open-pollinated flowers was pollen-limited, despite pollen deposition exceeding ovule number by 6-fold. Open-pollinated seed set was similar to that of the cross + self-pollen treatment, but was less than that of the cross-pollen-only treatment.
Flowers of C. breviflorus are pollinated primarily by pollen-collecting bees and possess a late-acting SI system, previously unknown in this clade of the Amaryllidaceae. Pollinators of C. breviflorus deposit mixtures of cross- and self-pollen and, because SI is late-acting, self-pollen disables ovules, reducing female fertility. This study thus contributes to growing evidence that seed production in plants with late-acting SI systems is frequently limited by pollen quality, even when pollinators are abundant.
PMCID: PMC2944973  PMID: 20647225
Amarydillaceae; Cyrtanthus breviflorus; honeybee pollination; late-acting self-incompatibility; ovule discounting; pollen limitation; pollen quantity and quality
3.  Floral traits mediate the vulnerability of aloes to pollen theft and inefficient pollination by bees 
Annals of Botany  2012;109(4):761-772.
Background and Aims
Pollen-collecting bees are among the most important pollinators globally, but are also the most common pollen thieves and can significantly reduce plant reproduction. The pollination efficiency of pollen collectors depends on the frequency of their visits to female(-phase) flowers, contact with stigmas and deposition of pollen of sufficient quantity and quality to fertilize ovules. Here we investigate the relative importance of these components, and the hypothesis that floral and inflorescence characteristics mediate the pollination role of pollen collection by bees.
For ten Aloe species that differ extensively in floral and inflorescence traits, we experimentally excluded potential bird pollinators to quantify the contributions of insect visitors to pollen removal, pollen deposition and seed production. We measured corolla width and depth to determine nectar accessibility, and the phenology of anther dehiscence and stigma receptivity to quantify herkogamy and dichogamy. Further, we compiled all published bird-exclusion studies of aloes, and compared insect pollination success with floral morphology.
Key Results
Species varied from exclusively insect pollinated, to exclusively bird pollinated but subject to extensive pollen theft by insects. Nectar inaccessibility and strong dichogamy inhibited pollination by pollen-collecting bees by discouraging visits to female-phase (i.e. pollenless) flowers. For species with large inflorescences of pollen-rich flowers, pollen collectors successfully deposited pollen, but of such low quality (probably self-pollen) that they made almost no contribution to seed set. Indeed, considering all published bird-exclusion studies (17 species in total), insect pollination efficiency varied significantly with floral shape.
Species-specific floral and inflorescence characteristics, especially nectar accessibility and dichogamy, control the efficiency of pollen-collecting bees as pollinators of aloes.
PMCID: PMC3286288  PMID: 22278414
Pollen theft; pollination efficiency; dichogamy; floral morphology; Aloe; Alooideae; Xanthorrhoeaceae; Asphodeloideae
4.  Heteromorphic Incompatibility and Efficiency of Pollination in Two Distylous Pentanisia Species (Rubiaceae) 
Annals of Botany  2004;95(3):389-399.
• 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.
PMCID: PMC4246787  PMID: 15596454
Butterfly pollination; distyly; pollen colour dimorphism; pollination; Pentanisia; reciprocal herkogamy; heteromorphic incompatibility; southern Africa; Rubiaceae
5.  A Sexually Dimorphic Corolla Appendage Affects Pollen Removal and Floral Longevity in Gynodioecious Cyananthus delavayi (Campanulaceae) 
PLoS ONE  2015;10(1):e0117149.
The floral traits of bisexual flowers may evolve in response to selection on both male and female functions, but the relative importance of selection associated with each of these two aspects is poorly resolved. Sexually dimorphic traits in plants with unisexual flowers may reflect gender-specific selection, providing opportunities for gaining an increased understanding of the evolution of specific floral traits. We examined sexually dimorphic patterns of floral traits in perfect and female flowers of the gynodioecious species Cyananthus delavayi. A special corolla appendage, the throat hair, was investigated experimentally to examine its influences on male and female function. We found that perfect flowers have larger corollas and much longer throat hairs than female flowers, while female ones have much exerted stigmas. The presence of throat hairs prolonged the duration of pollen presentation by restricting the amount of pollen removed by pollen-collecting bees during each visit. Floral longevity was negatively related to the rate of pollen removal. When pollen removal rate was limited in perfect flowers, the duration of the female phases diminished with the increased male phase duration. There was a weak negative correlation between throat hair length and seed number per fruit in female flowers, but this correlation was not significant in perfect flowers. These results suggest that throat hairs may enhance male function in terms of prolonged pollen presentation. However, throat hairs have no obvious effect on female function in terms of seed number per fruit. The marked sexual dimorphism of this corolla appendage in C. delavayi is likely to have evolved and been maintained by gender-specific selection.
PMCID: PMC4300179  PMID: 25603479
6.  Nectar robbers pollinate flowers with sexual organs hidden within corollas in distylous Primula secundiflora (Primulaceae) 
Biology Letters  2010;6(6):785-787.
Nectar robbers are thought rarely to pollinate flowers, especially those with sexual organs hidden within corollas. In this study, we examined whether robbers pollinate flowers of distylous Primula secundiflora. Distylous plants have two floral morphs. Pin flowers have long styles and short stamens, and thrum flowers have short styles and long stamens. Flowers of P. secundiflora were commonly robbed by bumble-bees, and robbing holes were always situated between high and low sexual organs for both floral morphs. We observed that pollen grains of pin flowers were removed while thrum flowers received fresh pollen grains immediately after flowers were robbed. We manipulated flowers so that only nectar robbers could visit them. This resulted in 98 per cent of thrum flowers and 6 per cent of pin flowers setting fruit, and seed number per thrum fruit was also significantly higher than per pin fruit. Our findings suggest that nectar robbers transfer pollen from pin flowers to thrum flowers effectively, and consequently increase male fitness of the pin morph and female fitness of the thrum morph. Such asymmetrical pollen flow caused by nectar robbers may act as an important selective agent in floral fitness and evolution of distyly.
PMCID: PMC3001367  PMID: 20462883
nectar robbing; distyly; reciprocity; robber-like pollinator
7.  Ecological Context of Breeding System Variation: Sex, Size and Pollination in a (Predominantly) Gynodioecious Shrub 
Annals of Botany  2007;100(7):1547-1556.
Background and Aims
Species that exhibit among-population variation in breeding system are particularly suitable to study the importance of the ecological context for the stability and evolution of gender polymorphism. Geographical variation in breeding system and sex ratio of Daphne laureola (Thymelaeaceae) was examined and their association with environmental conditions, plant and floral display sizes, and pollination environment in a broad geographic scale was analysed.
The proportion of female and hermaphrodite individuals in 38 populations within the Iberian Peninsula was scored. Average local temperature and precipitation from these sites were obtained from interpolation models based on 30 years of data. Pollination success was estimated as stigmatic pollen loads, pollen tubes per ovule and the proportion of unfertilized flowers per individual in a sub-set of hermaphroditic and gynodioecious populations.
Key Results
Daphne laureola is predominantly gynodioecious, but hermaphroditic populations were found in northeastern and southwestern regions, characterized by higher temperatures and lower annual precipitation. In the gynodioecious populations, female plants were larger and bore more flowers than hermaphrodites. However, due to their lower pollination success, females did not consistently produce more seeds than hermaphrodites, which tends to negate a seed production advantage in D. laureola females. In the northeastern hermaphroditic populations, plants were smaller and produced 9–13 times fewer flowers than in the other Iberian regions, and thus presumably had a lower level of geitonogamous self-fertilization. However, in a few southern populations hermaphroditism was not associated with small plant size and low flower production.
The findings highlight that different mechanisms, including abiotic conditions and pollinator service, may account for breeding system variation within a species' distribution range and also suggest that geitonogamy may affect plant breeding system evolution.
PMCID: PMC2759233  PMID: 17933844
Daphne laureola; environmental gradients; floral display; geographic variation; geitonogamy; gynodioecy; pollination success; sex ratio; Thymelaeaceae
8.  Effects of floral display size on male and female reproductive success in Mimulus ringens 
Annals of Botany  2011;109(3):563-570.
Background and Aims
The number of flowers blooming simultaneously on a plant may have profound consequences for reproductive success. Large floral displays often attract more pollinator visits, increasing outcross pollen receipt. However, pollinators frequently probe more flowers in sequence on large displays, potentially increasing self-pollination and reducing pollen export per flower. To better understand how floral display size influences male and female fitness, we manipulated display phenotypes and then used paternity analysis to quantify siring success and selfing rates.
To facilitate unambiguous assignment of paternity, we established four replicate (cloned) arrays of Mimulus ringens, each consisting of genets with unique combinations of homozygous marker genotypes. In each array, we trimmed displays to two, four, eight or 16 flowers. When fruits ripened, we counted the number of seeds per fruit and assigned paternity to 1935 progeny.
Key Results
Siring success per flower declined sharply with increasing display size, while female success per flower did not vary with display. The rate of self-fertilization increased for large floral displays, but siring losses due to geitonogamous pollen discounting were much greater than siring gains through increased self-fertilization. As display size increased, each additional seed sired through geitonogamous self-pollination was associated with a loss of 9·7 seeds sired through outcrossing.
Although total fitness increased with floral display size, the marginal return on each additional flower declined steadily as display size increased. Therefore, a plant could maximize fitness by producing small displays over a long flowering period, rather than large displays over a brief flowering period.
PMCID: PMC3278290  PMID: 21880660
Bumble-bee; floral display size; functional gender; geitonogamy; male selfing rate; mating system; Mimulus ringens; paternity analysis; pollen discounting; pollination; self-fertilization; siring success
9.  Adaptive Significance of Flexistyly in Alpinia blepharocalyx (Zingiberaceae): A Hand-pollination Experiment 
Annals of Botany  2007;99(4):661-666.
Background and Aims
Flexistyly is a sexual dimorphism where there are two morphs that differ in the temporal expression of sexual function and also involve reciprocal movement of the stigmatic surface through a vertical axis during the flowering period. The adaptive significance of flexistyly has been interpreted as a floral mechanism for outcrossing, but it may also function to reduce sexual interference in which styles and stigmas impede the pollen export. Here these two explanations of flexistyly were tested in Alpinia blepharocalyx through a hand-pollination experiment.
Hand-pollinations were performed in two temporal morphs and consisted of two sequential pollination treatments, namely self-pollination in the morning and inter-morph pollination in the afternoon (treatment 1) or conversely inter-morph pollination in the morning and self-pollination in the afternoon (treatment 2), and two simultaneous self- and inter-morph cross-pollination treatments either in the morning (treatment 3) or in the afternoon (treatment 4). Seed paternity was then determined to assess relative success of self- versus cross-pollen using allozyme markers.
Key Results
In the sequential pollination treatments, whether the stigmas of recipients are receptive in the morning is crucial to the success of the pollen deposited. When the cataflexistylous (protandrous) morph served as pollen recipient, early-arriving pollen in the morning can sire only a very small proportion (<15%) of seeds because the stigmas were then unreceptive. However, when the anaflexistylous (protogynous) morph served as pollen recipient, early pollen did gain a large competitive advantage over the late pollen, particularly when cross-pollen arrived first. Simultaneous self- and inter-morph cross-pollination indicated that outcross-pollen is more competitive than self-pollen on receptive stigmas.
Differential maturing of male and female organs in Alpinia blepharocalyx is sufficient for selfing avoidance, obviating the need for style movements. Instead, the upward style curvature of the cataflexistylous morph in the morning and the anaflexistylous morph in the afternoon most likely represents a means of reducing interference with pollen export.
PMCID: PMC2802932  PMID: 17237211
Alpinia; flexistyly; heterodichogamy; pollen competition; self-pollination; sexual interference
10.  Between-year Changes in Functional Gender Expression of Palicourea padifolia (Rubiaceae), a Distylous, Hummingbird-pollinated Shrub 
Annals of Botany  2004;95(2):371-378.
• Background and Aims Because distylous species have two hermaphroditic style-length floral morphs, they face two sex allocation problems: the equilibrium morph ratio and the optimal allocation to pollen and seed production in each floral morph. Gender specialization is expected among distylous species when floral morphs differ in reproductive output. However, spatio-temporal variability in female reproductive output between morphs needs to be investigated to fully understand patterns of sexual expression and gender specialization in distylous plants. Between-year variation in flower and fruit production of hummingbird-pollinated Palicourea padifolia (Rubiaceae) was examined, focusing on functional gender expression of long- and short-styled morphs and comparing their reproductive performance in five consecutive years (1998–2002).
• Methods Between-year variation in inflorescence, floral bud and fruit production was monitored and quantified. These traits were then used as parameters to determine functional gender differences between floral morphs through time.
• Key Results Inflorescence production varied among years but no significant differences were found between floral morphs. Long-styled plants initiated more floral buds per inflorescence every year than short-styled plants, suggesting higher allocation to pollinator attraction and, potentially, an increase in male fitness through pollen donation. Although fruit production was similar between morphs, their functional gender shifted across years.
• Conclusions The gender expression inconsistency across years is surprising because a number of floral characters and attributes that contribute to differently attracting and rewarding effective pollinators in P. padifolia suggest gender specialization. The evidence that morphs of distylous species might specialize in functional gender mostly comes from differences among populations in seed production and non-equilibrium morph ratios based on 1-year field population surveys. The results suggest that more sampling through time is needed to detect gender specialization among distylous species with a perennial habit.
PMCID: PMC4246829  PMID: 15546929
Cloud forest; distyly; gender expression; hummingbirds; Palicourea padifolia; Rubiaceae; sex-allocation theory
11.  Distribution of Flower Morphs, Ploidy Level and Sexual Reproduction of the Invasive Weed Oxalis pes-caprae in the Western Area of the Mediterranean Region 
Annals of Botany  2007;99(3):507-517.
Background and Aims
Oxalis pes-caprae is a widespread invasive weed in regions with a Mediterranean climate. In its native habitat (southern Africa) this species has been reported as heterostylous with trimorphic flowers and a self- and morph-incompatible reproductive system. In most of the areas invaded, only a pentaploid short-styled morphotype that reproduces mainly asexually by bulbils is reported, but this has only been confirmed empirically. This study aims to analyse the floral morph proportions in a wide distribution area, test the sexual female success, and explain the causes of low sexual reproduction of this species in the western area of the Mediterranean Basin.
Fifty-five populations of O. pes-caprae were sampled in the Iberian Peninsula and Morocco to evaluate the floral morph ratio and individual fruit set. In plants from a dimorphic population, hand-pollination experiments were performed to evaluate the effect of the pollen source on pollen tube growth through the style. The ploidy level and genome size of individuals of each floral morph were analysed using flow cytometry.
Key Results
From the populations studied 89·1 % were monomorphic, with most of them containing the short-styled (SS) floral morph, and 10·9 % were dimorphic containing long-styled (LS) and SS morphs. In some of these, isoplethy was verified but no fruit production was observed in any population. A sterile form was also recorded in several populations. Hand-pollination experiments revealed that pollen grains germinated over recipient stigmas. In intermorph crossings, pollen tubes were able to develop and fruit initiation was observed in some cases, while in intramorph pollinations, pollen tube development was sporadic and no fruit initiation was observed. All individuals within each floral form presented the same DNA ploidy level: SS plants were pentaploid and LS and the sterile form were tetraploid.
The low or null sexual reproduction success of this species in the area of invasion studied seems related with the high frequency of monomorphic populations, the unequal proportion of floral morphs in dimorphic populations and the presence of different ploidy levels between SS and LS morphs. The discovery of the occurrence of an LS floral morph and a sterile form, whose invading capacity in these areas is as yet unknown, will be valuable information for management programmes.
PMCID: PMC2802954  PMID: 17218342
Flow cytometry; genome size; heterostyly; invasive plant; Oxalis pes-caprae; ploidy level; reproductive biology; weed
12.  Recurrent modification of floral morphology in heterantherous Solanum reveals a parallel shift in reproductive strategy 
Floral morphology determines the pattern of pollen transfer within and between individuals. In hermaphroditic species, the spatial arrangement of sexual organs influences the rate of self-pollination as well as the placement of pollen in different areas of the pollinator's body. Studying the evolutionary modification of floral morphology in closely related species offers an opportunity to investigate the causes and consequences of floral variation. Here, we investigate the recurrent modification of flower morphology in three closely related pairs of taxa in Solanum section Androceras (Solanaceae), a group characterized by the presence of two morphologically distinct types of anthers in the same flower (heteranthery). We use morphometric analyses of plants grown in a common garden to characterize and compare the changes in floral morphology observed in parallel evolutionary transitions from relatively larger to smaller flowers. Our results indicate that the transition to smaller flowers is associated with a reduction in the spatial separation of anthers and stigma, changes in the allometric relationships among floral traits, shifts in pollen allocation to the two anther morphs and reduced pollen : ovule ratios. We suggest that floral modification in this group reflects parallel evolution towards increased self-fertilization and discuss potential selective scenarios that may favour this recurrent shift in floral morphology and function.
PMCID: PMC4084541  PMID: 25002701
division of labour; heteranthery; floral evolution; mating system; pollen : ovule ratio; Solanum (Solanaceae)
13.  Floral polymorphism in Chamaecrista flexuosa (Fabaceae-Caesalpinioideae): a possible case of atypical enantiostyly? 
Annals of Botany  2013;112(6):1117-1123.
Background and Aims
Reciprocal herkogamy, including enantiostyly and heterostyly, involves reciprocity in the relative positions of the sexual elements within the flower. Such systems result in morphologically and, since pollen is deposited on and captured from different parts of the pollinator, functionally distinct floral forms. Deviations from the basic pattern may modify the functionality of these mechanisms. For heterostylous species, such deviations are generally related to environmental disturbances, pollination services and/or reduced numbers of one floral morph. Deviations for enantiostylous species have not yet been reported. This study aims to investigate enantiostyly in Chamaecrista flexuosa, in particular the presence of deviations from the standard form, in an area of coastal vegetation in north-east Brazil.
Observations and investigations of floral biology, the reproductive system, pollinator behaviour, floral morphology and morphometry were performed.
Key Results
In C. flexuosa flowers, anthers of different size but similar function are grouped. The flowers were self-compatible and set fruits after every treatment, except in the spontaneous self-pollination experiment, thereby indicating their dependence on pollen vectors. The flowers were pollinated by bees, especially Xylocopa cearensis and X. grisencens. Pollen is deposited and captured from the ventral portion of the pollinator's body. Variations in the spatial arrangement of floral elements allowed for the identification of floral morphs based on both morphological and functional criteria. Using morphological criteria, morphologically right (MR) and morphologically left (ML) floral morphs were identified. Three floral morphs were identified using functional criteria: functionally right (FR), functionally central (FC) and functionally left (FL). Combinations of morphologically and functionally defined morphs did not occur in equal proportions. There was a reduced frequency of the MR–FR combination.
The results indicate the occurrence of an atypical enantiostyly in C. flexuosa. This seems to improve reproductive success by increasing the efficiency of pollen deposition and capture.
PMCID: PMC3783247  PMID: 24026440
Breeding system; Chamaecrista flexuosa; Fabaceae; Caesalpinioideae; enantiostyly; floral polymorphism; pollen flow; reciprocal herkogamy
14.  Sterile flowers increase pollinator attraction and promote female success in the Mediterranean herb Leopoldia comosa 
Annals of Botany  2012;111(1):103-111.
Background and Aims
Large floral displays have opposing consequences for animal-pollinated angiosperms: they attract more pollinators but also enable elevated among-flower self-pollination (geitonogamy). The presence of sterile flowers as pollinator signals may enhance attraction while allowing displays of fewer open fertile flowers, limiting geitonogamy. The simultaneous contributions of fertile and non-fertile display components to pollinator attraction and reproductive output remain undetermined.
The simultaneous effects of the presence of sterile flowers and fertile-flower display size in two populations of Leopoldia comosa were experimentally assessed. Pollinator behaviour, pollen removal and deposition, and fruit and seed production were compared between intact plants and plants with sterile flowers removed.
Key Results
The presence of sterile flowers almost tripled pollinator attraction, supplementing the positive effect of the number of fertile flowers on the number of bees approaching inflorescences. Although attracted bees visited more flowers on larger inflorescences, the number visited did not additionally depend on the presence of sterile flowers. The presence of sterile flowers improved all aspects of plant performance, the magnitude of plant benefit being context dependent. During weather favourable to pollinators, the presence of sterile flowers increased pollen deposition on stigmas of young flowers, but this difference was not evident in older flowers, probably because of autonomous self-pollination in poorly visited flowers. Total pollen receipt per stigma decreased with increasing fertile display size. In the population with more pollinators, the presence of sterile flowers increased fruit number but not seed set or mass, whereas in the other population sterile flowers enhanced seeds per fruit, but not fruit production. These contrasts are consistent with dissimilar cross-pollination and autonomous self-pollination, coupled with the strong predispersal inbreeding depression exhibited by L. comosa populations.
Sterile flowers enrich pollination quality by promoting pollen export and import, while limiting the mating costs of geitonogamy associated with large fertile displays.
PMCID: PMC3523656  PMID: 23131298
Anthophora; cross-pollination; geitonogamy; fertile floral display; mating cost; Leopoldia comosa; non-fertile flowers; outcrossing; pollen deposition; pollen quality; pollen removal; sterile flowers
15.  Is pollen removal or seed set favoured by flower longevity in a hummingbird-pollinated Salvia species? 
Annals of Botany  2010;106(3):413-419.
Background and Aims
The period between the beginning of anthesis and flower senescence modulates the transport of pollen by pollinators among conspecific flowers, and its length may therefore influence reproductive success. This study evaluated whether floral longevity favours pollen removal from the anthers over fecundity (seed set) in an ornithophilous species that does not undergo pollen limitation.
Field investigations were conducted on floral longevity, nectar production, pollinator behaviour, and variations in fruit set (FS), mean number of seeds per fruit (MSF) and pollen removal by hummingbirds (PR) during the anthesis of Salvia sellowiana in south-east Brazil.
Key Results
Anthesis of flowers exposed to pollinators lasted 4 d, as well as on flowers with pollen removed from the anthers or deposited on the stigma. The longevity of bagged flowers was significantly higher (approx. 9 d). FS and PR reached 87·2 and 90 %, respectively, in natural conditions. PR increased gradually over the period of anthesis; however, FS and MSF reached their maxima in the first hours of anthesis. Nectar production was continuous, but the secretion rate was reduced after pollination. The removal of nectar from non-pollinated flowers stimulated its production.
The longevity of anthesis in S. sellowiana seems to be related to the mechanism of gradual dispensing of pollen, resulting in greater male reproductive success. This is in agreement with the pollen-donation hypothesis. The small number of ovules (four) of S. sellowiana and the high frequency and the foraging mode of its pollinators may favour the selection for floral longevity driven by male fitness in this system.
PMCID: PMC2924834  PMID: 20622254
Atlantic Forest; breeding system; female reproductive success; Lamiaceae; male reproductive success; microevolution; phenotypic selection; pollen presentation; pollination biology; traplining
16.  Wind-Dragged Corolla Enhances Self-Pollination: A New Mechanism of Delayed Self-Pollination 
Annals of Botany  2007;100(6):1155-1164.
Background and Aims
Delayed self-pollination is a mechanism that allows animal-pollinated plants to outcross while ensuring seed production in the absence of pollinators. This study aims to explore a new mechanism of delayed self-pollination facilitated by wind-driven corolla abscission in Incarvillea sinensis var. sinensis.
Floral morphology and development, and the process of delayed self-pollination were surveyed. Experiments dealing with pollinator and wind exclusion, pollination manipulations, and pollinator observations were conducted in the field.
Key Results
Delayed self-pollination occurs when the abscising corolla driven by wind drags the adherent epipetalous stamens, thus leading to contact of anthers with stigma in late anthesis. There is no dichogamy and self-incompatibility in this species. The significantly higher proportion of abscised corolla under natural conditions as compared with that in wind-excluding tents indicates the importance of wind in corolla abscission. When pollinators were excluded, corolla abscission significantly increased the number of pollen grains deposited on the stigma and, as a result, the fruit and seed set. Only half of the flowers in plots were visited by pollinators, and the fruit set of emasculated flowers was significantly lower than that of untreated flowers in open pollination. This species has a sensitive stigma, and its two open stigmatic lobes closed soon after being touched by a pollinator, but always reopened if no or only little pollen was deposited.
This delayed self-pollination, which involved the movement of floral parts, the active participation of the wind and sensitive stigma, is quite different from that reported previously. This mechanism provides reproductive assurance for this species. The sensitive stigma contributes to ensuring seed production and reducing the interference of selfing with outcrossing. The pollination pattern, which combines actions by bees with indirect participation by wind, is also a new addition to ambophily.
PMCID: PMC2759251  PMID: 17881336
Ambophily; anther movement; Bignoniaceae; corolla abscission; delayed self-pollination; Incarvillea sinensis var. sinensis; reproductive assurance; stigma closure
17.  Pollen–Stigma Interference in Two Gynodioecious Species of Lamiaceae with Intermediate Individuals 
Annals of Botany  2006;100(2):423-431.
Background and Aims
Intermediate individuals (perfect flowers with very high degree of pollen abortion) in a gynodioecious plant species are very rare. A study is made of male–female relationships in each flower type and how floral characters can enhance the avoidance of ‘pollen discounting’ and ‘self-pollination’ in two gynodioecious species, Teucrium capitatum and Origanum syriacum.
The relationship between stigma receptivity and pollen viability was studied in two gynodioecious protandrous species of Lamiaceae, in addition to measuring some floral morphological characters over the life span of the flowers.
Key Results
Three plant types in each species were found: plants bearing hermaphrodite (or male fertile) flowers (MF), female (or male sterile) flowers (MS) and intermediate flowers (INT). Plant types differed in flower size, with MS types being shorter than the other two types. There was no difference in style length among plant types in T. capitatum. Stigma receptivity decayed with floral age and was negative and significantly correlated with pollen viability in the two species, and positive and significantly correlated with style length in O. syriacum but only in MS flowers of T. capitatum.
Reduction in size of floral characters is associated with male sterility, except style length in T. capitatum. MF flowers have two successive reproductive impediments: self-pollination and pollen–stigma interference. In both species, self-pollination is avoided by dichogamy (negative correlation between stigma receptivity and pollen viability), and pollen–stigma interference shows two different patterns: (1) style elongation in O. syriacum is characterized by a significant length increase, final MF dimensions are greater than those of MS dimensions, and style length is positively and significantly correlated with stigma receptivity; and (2) style movement in T. capitatum is characterized by a non-significant increase in style length, final MF floral dimensions are similar to those of MS dimensions, and there is no correlation between style length and stigma receptivity.
PMCID: PMC2735308  PMID: 16877457
Dichogamy; gynodioecy; Lamiaceae; Origanum syriacum; pollen discounting; pollen–stigma interference; pollen viability; protandry; stigma receptivity; style movement; style elongation; Teucrium capitatum
18.  Nectar Sugar Production across Floral Phases in the Gynodioecious Protandrous Plant Geranium sylvaticum 
PLoS ONE  2013;8(4):e62575.
Many zoophilous plants attract their pollinators by offering nectar as a reward. In gynodioecious plants (i.e. populations are composed of female and hermaphrodite individuals) nectar production has been repeatedly reported to be larger in hermaphrodite compared to female flowers even though nectar production across the different floral phases in dichogamous plants (i.e. plants with time separation of pollen dispersal and stigma receptivity) has rarely been examined. In this study, sugar production in nectar standing crop and secretion rate were investigated in Geranium sylvaticum, a gynodioecious plant species with protandry (i.e. with hermaphrodite flowers releasing their pollen before the stigma is receptive). We found that flowers from hermaphrodites produced more nectar than female flowers in terms of total nectar sugar content. In addition, differences in nectar production among floral phases were found in hermaphrodite flowers but not in female flowers. In hermaphrodite flowers, maximum sugar content coincided with pollen presentation and declined slightly towards the female phase, indicating nectar reabsorption, whereas in female flowers sugar content did not differ between the floral phases. These differences in floral reward are discussed in relation to visitation patterns by pollinators and seed production in this species.
PMCID: PMC3632553  PMID: 23614053
19.  Floral polymorphism and the fitness implications of attracting pollinating and florivorous insects 
Annals of Botany  2013;113(2):213-222.
Background and Aims
Floral polymorphism is frequently attributed to pollinator-mediated selection. Multiple studies, however, have revealed the importance of non-pollinating visitors in floral evolution. Using the polymorphic annual daisy Ursinia calenduliflora, this study investigated the importance of different insect visitors, and their effects on fitness, in the maintenance of floral polymorphism.
The spatial structure of a discrete floral polymorphism was characterized based on the presence/absence of anthocyanin floret spots in U. calenduliflora. A 3-year observational study was then conducted in polymorphic populations to investigate differences in visitation rates of dominant visitors to floral morphs. Experiments were performed to explore the floral preference of male and female Megapalpus capensis (the dominant insect visitor) and their effectiveness as pollinators. Next, floral damage by antagonistic florivores and the reproductive success of the two floral morphs were surveyed in multiple populations and years.
Key Results
Floral polymorphism in U. calenduliflora was structured spatially, as were insect visitation patterns. Megapalpus capensis males were the dominant visitors and exhibited strong preference for the spotted morph in natural and experimental observations. While this may indicate potential fitness benefits for the spotted morph, female fitness did not differ between floral morphs. However, as M. capensis males are very efficient at exporting U. calenduliflora pollen, their preference may likely increase the reproductive fitness of the spotted morph through male fitness components. The spotted morph, however, also suffered significantly greater costs due to ovule predation by florivores than the spotless morph.
The results suggest that pollinators and florivores may potentially exert antagonistic selection that could contribute to the maintenance of floral polymorphism across the range of U. calenduliflora. The relative strength of selection imposed by each agent is potentially determined by insect community composition and abundance at each site, highlighting the importance of community context in the evolution of floral phenotypes.
PMCID: PMC3890383  PMID: 24052554
Antagonistic selection; bee flies; community context; floral polymorphism; florivory; monkey beetles; ovule predation; pollen export; pollinator-mediated selection; Ursinia calenduliflora; Megapalpus capensis
20.  Effects of pollination limitation and seed predation on female reproductive success of a deceptive orchid 
AoB Plants  2014;6:plu031.
This research examines the contribution of plant height, number of flowers, number of stems, as well the joint impacts of mutualists and antagonists on the pollination biology and seed production of the imperiled, deceptive orchid, Cypripedium candidum. We found flowering stem height to be the only morphological feature significant in reproduction, with taller flowering stems simultaneously receiving increased pollination and decreased seed predation. Furthermore we found decreased seed mass in individuals subjected to hand-self pollination treatments. Our results may help explain the factors limiting seed production in other Cypripedium and further emphasize the importance of management in orchid conservation.
For many species of conservation significance, multiple factors limit reproduction. This research examines the contributions of plant height, number of flowers, number of stems, pollen limitation and seed predation to female reproductive success in the deceit-pollinated orchid, Cypripedium candidum. The deceptive pollination strategy employed by many orchids often results in high levels of pollen limitation. While increased floral display size may attract pollinators, C. candidum's multiple, synchronously flowering stems could promote selfing and also increase attack by weevil seed predators. To understand the joint impacts of mutualists and antagonists, we examined pollen limitation, seed predation and the effects of pollen source over two flowering seasons (2009 and 2011) in Ohio. In 2009, 36 pairs of plants size-matched by flower number, receiving either supplemental hand or open pollination, were scored for fruit maturation, mass of seeds and seed predation. Pollen supplementation increased proportion of flowers maturing into fruit, with 87 % fruit set when hand pollinated compared with 46 % for naturally pollinated flowers. Inflorescence height had a strong effect, as taller inflorescences had higher initial fruit set, while shorter stems had higher predation. Seed predation was seen in 73 % of all fruits. A parallel 2011 experiment that included a self-pollination treatment and excluded seed predators found initial and final fruit set were higher in the self and outcross pollination treatments than in the open-pollinated treatment. However, seed mass was higher in both open pollinated and outcross pollination treatments compared with hand self-pollinated. We found greater female reproductive success for taller flowering stems that simultaneously benefited from increased pollination and reduced seed predation. These studies suggest that this species is under strong reinforcing selection to increase allocation to flowering stem height. Our results may help explain the factors limiting seed production in other Cypripedium and further emphasize the importance of management in orchid conservation.
PMCID: PMC4094650  PMID: 24916060
Conservation; orchid; plant reproduction; plant–insect interactions; pollen limitation; pollination ecology; reproductive trade-offs; seed predation; supplemental pollination.
21.  Effects of pollen shortage and self-pollination on seed production of an endangered tree, Magnolia stellata 
Annals of Botany  2005;95(6):1009-1015.
• Background and Aims Pollen limitation is a significant determinant of seed production, and can result from both insufficient pollen quantity (pollen shortage) and quality (mainly relating to self-pollination). For animal-pollinated tree species with large floral displays, pollen limitation may be determined by a balance between increased pollen quantity due to increased attractiveness for pollinators, countered by increased self-pollination due to increased geitonogamy. The contributions of pollen shortage and self-pollination on seed production were quantitatively examined in the natural pollination of an insect-pollinated, dichogamous, endangered tree, Magnolia stellata, which has a large, showy floral display.
• Methods Manual self- and cross-pollinations were conducted to determine the effects of selfing on seed production. The outcrossing rate was measured using microsatellite analyses of open-pollinated seeds, and the embryo mortality rate caused by self-pollination was indirectly estimated. The frequency of ovule mortality due to pollen shortage was also inferred using the embryo mortality and ovule survival rates from natural pollination.
• Key Results The average fruit set, seed set per fruit, and ovule survival rate per tree from hand cross-pollination were 1·37, 3·15, and 3·34 times higher than those from hand self-pollination, respectively, indicating that self-pollination causes inbreeding depression for fruit and seed set. The multilocus-outcrossing rate (tm) was intermediate, 0·632, and the primary selfing rate was 0·657. This indicates that frequent geitonogamous selfing occurs. The ovule mortality rate due to pollen shortage and the embryo mortality rate due to self-pollination were estimated to be 80·8 % and 45·9 %, respectively.
• Conclusions It is concluded that seed production of M. stellata is strongly limited by both pollen shortage and self-pollination. Inefficient beetle-pollination and the automimicry system via asynchronous flowering might be responsible for the high level of pollen shortage and frequent geitonogamy. This is despite a large, showy floral display and the dichogamous system of the species.
PMCID: PMC4246754  PMID: 15753118
Magnolia stellata; pollen limitation; pollen shortage; self-pollination; fruit set; seed set; ovule survival rate; outcrossing rate; primary selfing rate; pollination; microsatellite analyses; inbreeding depression
22.  Realized tolerance to nectar robbing: compensation to floral enemies in Ipomopsis aggregata 
Annals of Botany  2009;103(9):1425-1433.
Background and Aims
Although the ecological and evolutionary consequences of foliar herbivory are well understood, how plants cope with floral damage is less well explored. Here the concept of tolerance, typically studied within the context of plant defence to foliar herbivores and pathogens, is extended to floral damage. Variation in tolerance to floral damage is examined, together with some of the mechanisms involved.
The study was conducted on Ipomopsis aggregata, which experiences floral damage and nectar removal by nectar-robbing bees. High levels of robbing can reduce seeds sired and produced by up to 50 %, an indirect effect mediated through pollinator avoidance of robbed plants. Using an experimental common garden with groups of I. aggregata, realized tolerance to robbing was measured. Realized tolerance included both genetic and environmental components of tolerance. It was hypothesized that both resource acquisition and storage traits, and traits involved in pollination would mitigate the negative effects of robbers.
Key Results
Groups of I. aggregata varied in their ability to tolerate nectar robbing. Realized tolerance was observed only through a component of male plant reproduction (pollen donation) and not through components of female plant reproduction. Some groups fully compensated for robbing while others under- or overcompensated. Evidence was found only for a pollination-related trait, flower production, associated with realized tolerance. Plants that produced more flowers and that had a higher inducibility of flower production following robbing were more able to compensate through male function.
Variation in realized tolerance to nectar robbing was found in I. aggregata, but only through an estimate of male reproduction, and traits associated with pollination may confer realized tolerance to robbing. By linking concepts and techniques from studies of plant–pollinator and plant–herbivore interactions, this work provides insight into the role of floral traits in pollinator attraction as well as plant defence.
PMCID: PMC2701760  PMID: 19304813
Compensation; herbivory; indirect effects; Ipomopsis aggregata; male reproductive success; nectar robbing; pollen donation; pollination; resistance; tolerance
23.  Variation of Pollinator Assemblages and Pollen Limitation in a Locally Specialized System: The Oil-producing Nierembergia linariifolia (Solanaceae) 
Annals of Botany  2008;102(5):723-734.
Background and Aims
Few studies have examined the dynamics of specialist plant–pollinator interactions at a geographical scale. This knowledge is crucial for a more general evolutionary and ecological understanding of specialized plant–pollinator systems. In the present study, variations in pollinator activity, assemblage composition and pollen limitation were explored in the oil-producing species Nierembergia linariifolia (Solanaceae).
Pollen limitation in fruit and seed production was analysed by supplementary hand pollination in five wild populations. Pollinator activity and identity were recorded while carrying out supplementary pollination to assess the effect of pollinators on the degree of pollen limitation. In two populations, pollen limitation was discriminated into quantitative and qualitative components by comparing supplementation and hand cross-pollination in fruit set and seed set. The effect of flower number per plant on the number of flowers pollinated per visitor per visit to a plant was examined in one of these populations as a possible cause of low-quality pollination by increasing geitonogamy.
Results and Conclusions
Although pollen limitation was evident along time and space, differences in magnitude were detected among populations and years that were greatly explained by pollinator activity, which was significantly different across populations. Floral display size had a significant effect on the visitation rate per flower. Limitation by quality clearly affected one population presumably due to a high proportion of geitonogamous pollen. The great inter-population variation in plant–pollinator interaction (both in pollinator assemblages composition and pollinator activity) and fitness consequences, suggests that this system should be viewed as a mosaic of locally selective processes and locally specialized interactions.
PMCID: PMC2712377  PMID: 18765440
Nierembergia linariifolia; Centris; Chalepogenus; pollen limitation; pollen quality; oil-producing flowers; specialized pollination; floral display; assemblage composition; geographic variation; Solanaceae; tests of equivalence
24.  How scent and nectar influence floral antagonists and mutualists 
eLife  null;4:e07641.
Many plants attract and reward pollinators with floral scents and nectar, respectively, but these traits can also incur fitness costs as they also attract herbivores. This dilemma, common to most flowering plants, could be solved by not producing nectar and/or scent, thereby cheating pollinators. Both nectar and scent are highly variable in native populations of coyote tobacco, Nicotiana attenuata, with some producing no nectar at all, uncorrelated with the tobacco's main floral attractant, benzylacetone. By silencing benzylacetone biosynthesis and nectar production in all combinations by RNAi, we experimentally uncouple these floral rewards/attractrants and measure their costs/benefits in the plant's native habitat and experimental tents. Both scent and nectar increase outcrossing rates for three, separately tested, pollinators and both traits increase oviposition by a hawkmoth herbivore, with nectar being more influential than scent. These results underscore that it makes little sense to study floral traits as if they only mediated pollination services.
eLife digest
Flowering plants have evolved a number of different approaches to reproduction. Some use their own pollen and self-fertilize, while others use pollen from other nearby plants. This fertilization by other plants is called ‘outcrossing’ and introduces new genetic variation into each generation, which is extremely important for the evolutionary process.
Some flowering plants rely on animals to help with pollination, attracting visitors with floral scents and rewarding the visitors with sugar-rich nectar. But scent and nectar also attract herbivores that damage the plants. This causes a dilemma for flowering plants, which has led some to evolve to not produce scent or to offer no nectar while masquerading as a plant that does. Previous studies into the costs and benefits of such strategies have looked at the effects of either floral scent or nectar, but no-one has uncoupled the effects of these two traits on both pollination and herbivore attack.
Kessler et al. have addressed this issue in wild tobacco plants, which can both self-fertilize and outcross, and which produce varying amounts of scent and nectar. The experiments were conducted under mesh tents and in field trials in the plant's natural habitat: the Great Basin Desert in Utah. Kessler et al. used a gene-silencing technique called ‘RNA interference’ to inhibit the production of scent or nectar, either separately or together. When grown in field trials, under conditions that prevent self-fertilization, these tobacco plants are frequently visited by a hummingbird and three species of hawkmoth. All four of these animals pollinate the tobacco plants, but one of the moths also lays eggs that hatch into caterpillars, which damage the plant. Kessler et al. monitored the effects that the loss of scent, nectar or both had on visits by each pollinator and on outcrossing.
These experiments revealed that scent is essential to attract one hawkmoth species but not for another (called Hyles lineata). Furthermore, while, the hummingbird needs nectar, the H. lineata moth does not; but this moth won't visit flowers that lack both scent and nectar. The experiments also show that, for the moth that lays its eggs on the tobacco plants, both scent and nectar increase pollination and egg laying, but nectar has a stronger effect. Thus reducing nectar, as this tobacco plant does in the wild, is one strategy that can be used to reduce herbivore attack by caterpillars. Together, these findings highlight that it is important to study both herbivores and pollinators when attempting to understand the evolution of floral traits.
PMCID: PMC4530224  PMID: 26132861
Nicotiana attenuata; Manduca sexta; Hyles lineata; Archilochus alexandri; Manduca quinquemaculata; pollination; other
25.  Resolution of male-female conflict in an hermaphroditic flower. 
The flowers of most angiosperm species are hermaphroditic. Spatial separation of male and female organs within a flower (hercogamy) is a common character traditionally interpreted as an adaptation to reduce intrafloral self-fertilization, one potential cost of hermaphroditism. Another possible cost that may lead to selection for hercogamy is physical interference between male and female floral functions. Here, I present evidence demonstrating the role of a floral character in reducing female interference with male function. The bi-lobed stigma of the bush monkeyflower closes after receiving pollen, causing increased spatial separation of the anthers and stigma ('movement' hercogamy). Experimental manipulations show that flowers with closed stigmas export more than twice as much pollen to other flowers as those in which the stigma is prevented from closing. However, stigma closure only minimally reduces the potential for intrafloral self-pollination. This study provides the first experimental evidence that selection to reduce intrafloral male female interference can be a strong selective force and can drive the evolution of floral characters usually interpreted as mechanisms to reduce self-fertilization.
PMCID: PMC1088636  PMID: 11296865

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