PROTANDRY,INCOMPATIBILITY, AND SECONDARY POLLEN PRESENTATION IN CEPHALANTHUS OCCIDENTALS (RUBIACEAE)

Cephalanthus occidentalis L. is protandrous and presents pollen secondarily on the stigma surface. Because self-pollen is present on the stigma, the degree of selling vs. outcrossing in this species will depend on 1) the phenology of pollen presentation and stigma receptivity; 2) whether the species is self-incompatible; and 3) the rates of self vs. crossed pollen tube growth. This study describes floral morphology and phenology, self-incompatibility, and pollen tube growth rates in self- and crosspollinations of C. occidentalis. Light and scanning electron microscopy were used to study stigma morphology after flower opening, while controlled pollinations tested for incompatibility. Stigmas were unreceptive initially but became receptive by the second day after flower opening. Ninety-two percent of cross-pollinated flowers set fruit, compared to 12% fruit set in self-pollinations. Pollen tubes from selfed and out-crossed pollen initially had similar growth rates. Out-crossed pollen tubes began to grow rapidly ca. 5 hr after pollination of a receptive stigma, whereas selfed pollen tubes ceased growth or grew slowly after this time. Pollen tubes from out-crossed pollen grew the length of the style within 24 hr after pollination, while selfed pollen tubes were inhibited at the stigma-style junction. Our results indicate that C. occidentalis has selfincompatibility, in addition to protandry and secondary pollen presentation. Protandry allows removal of self-pollen from the unreceptive stigma, while self-incompatibility prevents fertilization by unremoved self-pollen.     

Floral ecology of Oreocharis acaulis (Gesneriaceae): An exceptional case of “preanthetic” protogyny combined with approach herkogamy

Protogyny is supposed to represent the ancestral form of dichogamy in the angiosperms, but is rare in advanced groups such as the Asteridae, in which protandry prevails by far. Here we report on an unusual form of protogyny combined with herkogamy in a Chinese species of Gesneriaceae (Asteridae–Lamiales): Oreocharis acaulis (formerly Opithandra acaulis). This is characterized by a conspicuous protrusion of the style from the flower bud and the stigma becoming receptive before corolla opening (female-only stage; preanthetic protogyny) and both sexes staying functional during anthesis (hermaphroditic stage), with the stigma presented above the anther level (approach herkogamy). The plants studied were found to be self-compatible, but autonomous self-pollination and apomixis were not observed. Successful pollination was found to depend fully on the presence of insect pollinators (Bombus sp.). The visiting frequency was higher in the hermaphroditic stage (in which also more nectar was produced) than in the female-only stage. The out-crossed flower buds opened earlier and had a shorter flowering period than selfed flowers. Similarly, the outcross-pollen germinated earlier and the pollen tube growth was faster than in self-pollen. Anthers, pistil and corolla of O. acaulis obviously form an integrative functional unit in which the elongated style plays a key role both in pollen-dispensing and pollen-deposition. The combination of preanthetic protogyny with herkogamy has probably arisen through selection for promoting out-crossing and prolonging the exposition time of the receptive stigma in order to capture a higher amount of pollen grains. This may be understood as a strategy to cope with scarcity of pollinators in the plants’ habitat.     

The endangered Iris atropurpurea (Iridaceae) in Israel: honey-bees,night-sheltering male bees and female solitary bees as pollinators

Background and Aims

The coastal plain of Israel hosts the last few remaining populations of the endemic Iris atropurpurea (Iridaceae), a Red List species of high conservation priority. The flowers offer no nectar reward. Here the role of night-sheltering male solitary bees, honey-bees and female solitary bees as pollinators of I. atropurpurea is documented.

Methods

Breeding system, floral longevity, stigma receptivity, visitation rates, pollen loads, pollen deposition and removal and fruit- and seed-set were investigated.

Key Results

The main wild pollinators of this plant are male eucerine bees, and to a lesser extent, but with the potential to transfer pollen, female solitary bees. Honey-bees were found to be frequent diurnal visitors; they removed large quantities of pollen and were as effective as male sheltering bees at pollinating this species. The low density of pollen carried by male solitary bees was attributed to grooming activities, pollen displacement when bees aggregated together in flowers and pollen depletion by honey-bees. In the population free of honey-bee hives, male bees carried significantly more pollen grains on their bodies. Results from pollen analysis and pollen deposited on stigmas suggest that inadequate pollination may be an important factor limiting fruit-set. In the presence of honey-bees, eucerine bees were low removal–low deposition pollinators, whereas honey-bees were high removal–low deposition pollinators, because they removed large amounts into corbiculae and deposited relatively little onto receptive stigmas.

Conclusions

Even though overall, both bee taxa were equally effective pollinators, we suggest that honey-bees have the potential to reduce the amount of pollen available for plant reproduction, and to reduce the amount of resources available to solitary bee communities. The results of this study have potential implications for the conservation of this highly endangered plant species if hives are permitted inside reserves, where the bulk of Oncocyclus iris species are protected.     

The pollination biology and breeding system of Monarda fistulosa (Labiatae)

Summary Successful cross-pollination of Monarda fistulosa is the result of a complex interaction among flower opening, the pollen-bearing areas of the pollinators and/or their behavior, and the maturation of the stigmas. The flowers open continuously from 0800–2000 h providing a temporally predictable rich source of nectar and pollen. Recently opened flowers may reduce the ability of bees to discriminate between resource rich and poor patches and encourage systematic foraging within patches. The continuous opening of flowers coupled with protandry also results in some flowers of most capitula being in the staminate and others in the pistillate phase. Autogamy is highly unlikely due to strong protandry and the spatial separation of anthers and stigmas. Geitonogamy, at least that mediated by Bombus is unlikely because the pollen is spread over a relatively large area of the wings, which reduces the likelihood of a stigma contacting just deposited pollen. Because pollen is transferred from the much smaller coxal area of Anthophora and other bees that mistake the stigmas of early pistillate phase flowers for stamens some geitonogamy seems inevitable. However, the delayed receptivity of young stigmas to self-pollen decreases the likelihood of self-pollen germinating on such stigmas. Older stigmas are equally receptive to self- and cross-pollen and the number of pollen grains germinating and pollen tubes reaching the base of the style increases with flower age.     

The transmitting tissue of Nicotiana tabacum is not essential to pollen tube growth, and its ablation can reverse prezygotic interspecific barriers

The Nicotiana tabacum transmitting tissue is a highly specialized file of metabolically active cells that is the pathway for pollen tubes from the stigma to the ovules where fertilization occurs. It is thought to be essential to pollen tube growth because of the nutrients and guidance it provides to the pollen tubes. It also regulates gametophytic self-incompatibility in the style. To test the function of the transmitting tissue in pollen tube growth and to determine its role in regulating prezygotic interspecific incompatibility, genetic ablation was used to eliminate the mature transmitting tissue, producing a hollow style. Despite the absence of the mature transmitting tissue and greatly reduced transmitting-tissue-specific gene expression, self-pollen tubes had growth to the end of the style. Pollen tubes grew at a slower rate in the transmitting-tissue-ablated line during the first 24 h post-pollination. However, pollen tubes grew to a similar length 40 h post-pollination with and without a transmitting tissue. Ablation of the N. tabacum transmitting tissue significantly altered interspecific pollen tube growth. These results implicate the N. tabacum transmitting tissue in facilitating or inhibiting interspecific pollen tube growth in a species-dependent manner and in controlling prezygotic reproductive barriers.     

Floral traits mediate the vulnerability of aloes to pollen theft and inefficient pollination by bees

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.

Methods

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.

Conclusions

Species-specific floral and inflorescence characteristics, especially nectar accessibility and dichogamy, control the efficiency of pollen-collecting bees as pollinators of aloes.     

Pollination and late-acting self-incompatibility in Cyrtanthus breviflorus (Amaryllidaceae): implications for seed production

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.

Methods

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.

Conclusions

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.     

Mixed Mating System Are Regulated by Fecundity in Shorea curtisii (Dipterocarpaceae) as Revealed by Comparison under Different Pollen Limited Conditions

The maintenance of mixed mating was studied in Shorea curtisii, a dominant and widely distributed dipterocarp species in Southeast Asia. Paternity and hierarchical Bayesian analyses were used to estimate the parameters of pollen dispersal kernel, male fecundity and self-pollen affinity. We hypothesized that partial self incompatibility and/or inbreeding depression reduce the number of selfed seeds if the mother trees receive sufficient pollen, whereas reproductive assurance increases the numbers of selfed seeds under low amounts of pollen. Comparison of estimated parameters of self-pollen affinity between high density undisturbed and low density selectively logged forests indicated that self-pollen was selectively excluded from mating in the former, probably due to partial self incompatibility or inbreeding depression until seed maturation. By estimating the self-pollen affinity of each mother tree in both forests, mother trees with higher amount of self-pollen indicated significance of self-pollen affinity with negative estimated value. The exclusion of self-fertilization and/or inbreeding depression during seed maturation occurred in the mother trees with large female fecundity, whereas reproductive assurance increased self-fertilization in the mother trees with lower female fecundity.     

Sexual interference within flowers of Chamerion angustifolium

Hermaphroditism is prevalent in plants but may allow interference between male function (pollen removal and dispersal) and female function (pollen receipt and seed production) within a flower. Temporal or spatial segregation of gender within a hermaphroditic flower may evolve to reduce this interference and enhance male and female reproductive success. We tested this hypothesis using Chamerion angustifolium (Onagraceae), in which pollen removal (male) and pollen deposition (female) were measured directly on hermaphroditic and experimentally produced unisexual flowers. During a single flower visit in the field, bees deposited 159±24 (SE) pollen grains on a stigma and removed 1058±198 grains from each flower. Anther removal did not alter deposition rates. In the laboratory, bees removed 2669±273 pollen grains and deposited 209±72.3 cross-pollen and 120±28.4 facilitated self-pollen grains per visit. The presence of anthers significantly reduced cross-pollen deposition on the stigma. In contrast, pollen removal was not affected by presence of the pistil. These results suggest that within-flower interference affects female function and represents a fitness cost that can be reduced through temporal segregation of gender within the flower. Co-ordinating editor: S.-M. Chang     

The Causes of self-sterility in natural populations of the relictual angiosperm, Illicium floridanum (Illiciaceae)

BACKGROUND AND AIMS: Illicium floridanum, a species belonging to the basal extant angiosperm taxon Illiciaceae, reportedly exhibits self-incompatibility (SI). To date, the site and timing of SI within the carpel of this species remains unidentified. Thus, the objective of this research was to determine the cellular and temporal aspects of SI in I. floridanum. METHODS: Following controlled application of cross- and self-pollen in natural populations of I. floridanum, embryo sac development and temporal aspects of stigma receptivity, as well as pollen tube growth, fertilization, and embryo and endosperm development, were investigated with the aid of light and fluorescence microscopy. KEY RESULTS: Flowers of I. floridanum exhibited complete dichogamy whereby stigmas only supported cross- and self-pollen tube growth prior to anther dehiscence. In contrast to earlier reports of SI in this species, a prezygotic SI resulting in rejection of self-pollen tube growth at the stigma was absent and there were no significant differences between cross- versus self-pollen germination and pollen tube growth within the style and ovary during the first 5 d after pollination. Structural development of the four-celled embryo sac was not differentially influenced by pollen type as noted to occur in other angiosperms with late-acting ovarian SI. The ovule micropyle and embryo sac were penetrated equally by cross- and self-pollen tubes. In addition, there were no statistically significant differences in cross- versus self-fertilization. A resting zygote and multicellular endosperm at a variety of developmental stages was present by 30 d after application of cross- or self-pollen. CONCLUSIONS: In the clear absence of a prezygotic SI that was previously reported to result in differential self-pollen tube growth at the stigma, self- sterility in I. floridanum is likely due to early-acting inbreeding depression, although late-acting post-zygotic ovarian SI cannot be ruled out.     

Sites of self-pollen tube inhibition in Papaveraceae (sensu lato)

Papaver rhoeas (Papaveraceae) has a well-characterized gametophytic self-incompatibility system in which self-pollen tube growth ceases either just before, or just after, emergence from the copal aperture. Papaver flowers are unusual, however, in having flat stigmatic rays sitting directly on top of the hard ovary and no style. Immediate self-pollen arrest might be required with this floral architecture. There is much variation in floral architecture among Papaveraceae and self-incompatibility is widespread. However, there are no reports of the site of self-pollen tube inhibition in Papaveraceae other than P. rhoeas. We examined the site of self-pollen tube inhibition in four species (Argemone munita, Lamprocapnos spectabilis, Eschscholzia californica, and Platystemon californicus) representing a broad phylogenetic and morphological sample of Papaveraceae. Squash preparation was used for species with soft stigmas whereas woody tissue was sectioned with a cryostat and images were stitched into a mosaic to visualize pollen tubes on whole stigmas. For three species, self-pollen tube inhibition appeared similar to that described for P. rhoeas. Self-pollen tubes were arrested before any substantial penetration of female tissue and usually did not grow longer than 100?μm. In the fourth species, A. munita, self-pollen tubes grew up to 500?μm in length. However, self-pollen tubes appeared to grow along the stigmatic spines, and growth ceased once tubes contacted the stigma surface. Despite variation in floral architecture, rapid arrest of self-pollen tubes occurred before or just after penetration of female tissue in all species, consistent with the hypothesis that members of the family share the same incompatibility mechanism.     

Selection on pollen donors by Echium vulgare (Boraginaceae)

Fewer than 25% of the ovules of Echium vulgare in the field develop into viable seeds, even with no pollen limitation. The loss of ovules could enhance the fitness of the maternal parent, if less fit embryos are selectively aborted. Two pollination experiments were performed to examine the selectivity of maternal parents on self-pollen and different cross-pollen sources. Pollinated with one pollen genotype per flower, self-pollen was, on average, as successful in siring seeds as cross-pollen. However, the relative success of self-pollen compared to outcross-pollen differed significantly among the maternal parents. These results suggest that, under certain conditions, selfing can produce a greater number of seeds than cross-pollination. Pollen donors differed significantly in outcrossing success. The plants that were more successful in selfing were also more successful pollen donors in outcross-pollinations. No significant interaction between maternal parent and paternal genotype was found. Pollinations with a pollen mixture produced selfed and outcrossed seeds in the same ratios as the single-donor experiment. Overall, only slight differences were found between the single- and mixed-donor experiments. Pollen tube growth did not show a significant correlation with the success of the parental genotypes in the mixed-donor experiment, indicating that pollen tube growth is not the determining factor controlling the paternity of the seeds. These results are discussed with reference to possible mediating mechanisms.     

Reproductive biology of Abolboda pulchella and A. poarchon (Xyridaceae: Poales)

Background and Aims

Abolboda (Xyridaceae) belongs to the Poales, a predominantly wind-pollinated order whose phylogeny has been widely studied in recent years. The reproductive biology of Abolboda pulchella and A. poarchon was studied to determine the main pollination system of these species, providing the first experimental data on reproduction in the Xyridaceae.

Methods

A field study was conducted, including observations on the morphology and biology of the flowers, insect visits and pollinator behaviour. Experimental pollination treatments were performed to assess agamospermy, spontaneous self-pollination and self-compatibility. Pollination success was determined by pollen tube growth, and reproductive success was assessed by fruit- and seed-set.

Key Results

Abolboda pulchella and A. poarchon were pollinated by Apidae, Megachilidae and Halictidae bees. The floral resources were pollen and nectar that was produced by stylar appendages, an uncommom nectary type for monocotyledons. The species were self-compatible, and pollen tube growth from self-pollen was similar to that of cross-pollen. However, herkogamy prevented spontaneous selfing, rendering the plants dependent on the pollinator''s activity. There was no production of seeds by agamospermy.

Conclusions

Melittophily is the main pollination system of these two Abolboda species. Nectar production was first recorded here for Xyridaceae, and along with self-compatibility, herkogamy and bee pollination, is an informative characteristic that can be used in future phylogenetic analyses of the family as well as Poales.     

Pollen tube reallocation in two preanthesis cleistogamous species, Ranalisma rostratum and Sagittaria guyanensis ssp. lappula (Alismataceae)

Pollen deposition on stigmas and pollen tube growth in two apocarpous species, Ranalisma rostratum and Sagittaria guyanensis ssp. lappula (Alismataceae), were examined with fluorescence microscopy. The reallocation of pollen tubes among pistils was observed in both species. The percentage of pollinated stigmas per flower was only 22.0% in R. rostratum and 51.0% in S. guyanensis, though the seed/ovule ratios are higher than 65% in both species. The number of pollen grains on each single stigma ranged from 0 to 96 in R. rostratum, and from 0 to 125 in S. guyanensis. When more than one pollen grain deposited on a stigma, all pollen tubes grew to the ovary, but only one of them turned towards the ovule and finally entered the nucleus. The other tubes grew through the receptacle tissue into ovules of adjacent carpels whose stigmas were unpollinated or pollinated later. The intercarpellary growth of pollen tubes could be a mechanism to increase the efficiency of sexual reproduction in an apocarpous gynoecium with low pollination on the pistils.     

Floral Structure, Stigma Receptivity and Pollen Viability in Relation to Protandry and Self-incompatibility in Silky Oak (Grevillea robusta A. Cunn.)

The reproductive biology of Grevillea robusta growing underexotic conditions in Kenya and Australia is reported. The speciesshowed both protandry and a self-incompatibility mechanism.The stigma was wet and papillate with a distinct groove in themiddle. The anthers dehisced prior to anthesis, when the perianthopened. Stigmatic receptivity began 1 d after anthesis, withthe greatest pollen germination rates and longest pollen tubesobtained 2 d after anthesis. Nectar secretion commenced withpollen dehiscence and was abundant at anthesis. Most stigmaticgrooves opened widely 1–2 d after anthesis and stigmasshowed taller papillae and abundant secretion. Controlled pollinationsgave a greater fruit set from cross-pollination (5.9% in Apriland 17.5% in July) than open-pollination (0.1% in April and3.3% in July). No fruit set from self-pollination was obtainedin April, and very few fruit set for geitonogamous (two outof 1622; 0.1%) or for autogamous (one out of 2707 flowers; 0.04%)pollination treatments in July. Following self-pollination,growth of pollen tubes was poorer than in other treatments,and was generally arrested in the upper style. Cross-pollinatedflowers produced normal and straight pollen tubes, while self-pollentubes had growth abnormalities. Most of the open-pollinatedflowers were found without pollen or with only self-pollen ontheir stigmas indicating that the amount of cross-pollen reachingthe stigma under open-pollination may be a factor limiting seedproduction. Flowers shed soon after the fertilization phasewere those with ungerminated pollen or no pollen. Although avery low rate of selfing may occur, G. robusta presents a self-incompatibilitysystem and allogamy is its primary breeding behaviour.Copyright2000 Annals of Botany Company Grevillea robusta, silky oak, Proteaceae, protandry, controlled pollinations, receptivity, pollen-tube growth, self-incompatibility, pollination     

Status of self-pollen in bee pollination efficiency of white clover (Trifolium repens L.)

Flowers of white clover (Trifolium repens L.) are hermaphrodite and self-incompatible; their cross-pollination depends entirely on insect visitors, mainly bees (Apoidea). Because self-pollination of white clover occurs before flower anthesis, we determined whether selfing affected the pollination efficiency of a honeybee visit. We compared pollen deposition in emasculated and intact flowers following (1) a single honeybee visit, (2) open-pollination for a day and (3) enclosure in a cloth bag to prevent insect visits. In emasculated flowers, open-pollination resulted in more pollen deposited than after one visit (+30%) which is consistent with flowers being visited more than once by pollinators during the course of a day. On intact flowers, saturation of the stigma was achieved after the first visit of a honeybee (near 280 grains) because of self-pollination. Additional visits did not increase pollen deposits, but they improved pollen efficiency in terms of numbers of pollen tubes reaching the ovules. In such a context of easily saturated stigmas, self-pollen does not inhibit cross-pollen activity, but represents a constraint for pollination which demands multiple bee visits to each flower to achieve maximum fertilization. Received: 20 May 1997 / Accepted: 25 October 1997     

Reproductive biology ofCaesalpinia calycina andC. pluviosa (Leguminosae) of the caatinga of north-eastern Brazil

The pollination biology, breeding system and fruiting success ofCaesalpinia calycina andC. pluviosa var.sanfranciscana were studied in caatinga vegetation in Bahia, NE Brazil. The principal pollinators for both species were carpenter bees.Caesalpinia calycina is andromonoecious but inC. pluviosa all flowers are hermaphrodite. InC. calycina all selfed flowers were abscised within 72 h despite rapid self-pollen tube growth to the ovary and ovule penetration. Prevention of selfing therefore seems to be controlled by a post-zygotic mechanism. Both species had very low fruit-set and it is suggested that this is at least in part due to geitonogamous pollinations with ovule penetration by self pollen tubes.     

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

Background and Aims 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.Methods 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.Key Results 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.Conclusions 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.     

A Temporal Dimension to the Influence of Pollen Rewards on Bee Behaviour and Fecundity in Aloe tenuior

The net effect of pollen production on fecundity in plants can range from negative – when self-pollen interferes with fecundity due to incompatibility mechanisms, to positive – when pollen availability is associated with increased pollinator visitation and fecundity due to its utilization as a reward. We investigated the responses of bees to pollen and nectar rewards, and the effects of these rewards on pollen deposition and fecundity in the hermaphroditic succulent shrub Aloe tenuior. Self-pollinated plants failed to set fruit, but their ovules were regularly penetrated by self-pollen tubes, which uniformly failed to develop into seeds as expected from ovarian self-incompatibility (or strong early inbreeding depression). Bees consistently foraged for pollen during the morning and early afternoon, but switched to nectar in the late afternoon. As a consequence of this differential foraging, we were able to test the relative contribution to fecundity of pollen- versus nectar-collecting flower visitors. We exposed emasculated and intact flowers in either the morning or late afternoon to foraging bees and showed that emasculation reduced pollen deposition by insects in the morning, but had little effect in the afternoon. Despite the potential for self-pollination to result in ovule discounting due to late-acting self-sterility, fecundity was severely reduced in artificially emasculated plants. Although there were temporal fluctuations in reward preference, most bee visits were for pollen rewards. Therefore the benefit of providing pollen that is accessible to bee foragers outweighs any potential costs to fitness in terms of gender interference in this species.