Effects of soil phosphorus on pollen production,pollen size,pollen phosphorus content,and the ability to sire seeds in Cucurbita pepo (Cucurbitaceae)

To determine the effects of soil phosphorus on pollen production, pollen grain size, phosphate concentration per pollen grain, and the siring ability of pollen, two cultivars of the common zucchini (Cucurbita pepo) were grown under two soil phosphorus conditions in an experimental garden. Overall, soil phosphorus availability had a significant effect on reproductive output through the female function and on traits affecting the male function of plants (staminate flower production, pollen production per flower, and pollen grain size). In addition, pollen produced by plants in the high phosphorus soils had a higher phosphate concentration than pollen produced by plants in the low phosphorus soils. A pollen mixture experiment revealed that pollen produced by plants in the high phosphorus treatment sired significantly more seeds than pollen produced by plants in the low phosphorus treatment. This study showed that growing conditions such as soil phosphorus can influence the size of a pollen grain and its chemical composition, which, in turn, can affect its ability to sire mature seeds.     

The size and germinability of Scots pine pollen in different temperatures in vitro

The effect of genotype, the origin of genotype, and germination temperature on Scots pine pollen grain size, hydration rate, germinability, and tube growth was studied in vitro. The mean sizes of dry and germinated pollen grains varied among pollen genotypes in different ways, thus the hydration rate varied among genotypes. Pollen from Scots pine that originates in northern Finland hydrated more than pollen from a population in southern Finland. Germination temperature had no effect on the hydration rate. Germinability and tube growth rate of northern genotypes were higher at 20 °C than at 15 °C. Differences among southern genotypes were not significant. At 15 °C, the germinability and pollen tube growth rate of northern genotypes were lower than southern genotypes. At 20 °C, the differences were not significant. It appears that germination and growth of pollen from northern populations are enhanced at higher temperatures whereas pollen from southern populations is unaffected.     

The potential for selection on pollen colour dimorphisms in Nigella degenii: morph-specific differences in pollinator visitation, fertilisation success and siring ability

Two subspecies of Nigella degenii (Ranunculaceae) possess a dimorphism in pollen colour and vary extensively in frequency of the two morphs in natural populations. Here we investigate the role of selection on pollen colour during the pollination phase in the two subspecies and its potential contribution to the maintenance of this colour variation. In a combination of common garden experiments and field observations, we obtained data on pollinator visitation rates and explored the effect of pollen colour on fertilisation success and siring ability under conditions of low vs. high pollen competition. In experimental gardens, naïve pollinators responded differently to plants with different pollen colour, but the favoured morph varied between dates and locations, and colour morphs were not visited in a frequency-dependent manner. Donor plants with dark pollen had a reproductive advantage (higher seed set) in single-donor pollinations, but the realised siring ability (measured by progeny morph ratio) was highly variable between different two-donor crosses with no general bias towards the light or dark morph. Therefore, although the dark pollen type appears to have a general selective advantage in terms of fertilisation success, our data are also consistent with a scenario involving the maintenance of both colour morphs, particularly under conditions of high pollen competition, a variable genetic background and/or spatial or temporal variation in the pollinator fauna.     

Pollen tube competition as a mechanism of prezygotic reproductive isolation between Mimulus nasutus and its presumed progenitor M. guttatus

This paper considers the extent to which differences in pollen tube growth rates can provide prezygotic reproductive isolation between Mimulus nasutus and its presumed progenitor, Mimulus guttatus . Mimulus nasutus is partially cleistogamous, but its larger chasmogamous flowers offer appreciable opportunity for outcrossing. Mimulus nasutus was found to have smaller pollen grains and shorter styles than M. guttatus . No differences were observed in pollen grain germination on conspecific and heterospecific stigmas. However, pollen tube growth rates of M. nasutus were found to be much slower than those of M. guttatus in the styles of that species. Consequently, any M. nasutus pollen transferred to an M. guttatus stigma was found to be competitively disadvantaged in an M. gutattus style. By contrast, no difference in pollen tube growth rate was detected between the species when growing in M. nasutus styles, possibly because M. nasutus styles are unable to support fast pollen tube growth. We tested the prediction from the pollen tube studies that a 50:50 mix of M. guttatus and M. nasutus pollen would produce 50% hybrid seeds when M. nasutus was the maternal parent, and near to 0% hybrid seed when M. guttatus was the maternal parent. The results were found to support this prediction. We conclude that pollen–pistil interactions can effect strong reproductive isolation between these species, as M. guttatus pollen tubes have a competitive advantage over those of M. nasutus in an M. guttatus style, but not in an M. nasutus style.     

Pollen layering and male–male competition: Quantum dots demonstrate that pollen grains compete for space on pollinators

Premise

Almost nothing is known about what happens to pollen grains once they attach to pollinators, although some have postulated that pollen from different donors may form complex, two- or three-dimensional landscapes (e.g., layers or mosaics) that can facilitate male–male competition. For example, pollen that is already on pollinators may preclude the deposition of subsequent pollen grains.

Methods

Using quantum dots to mark the pollen of individual flowers, we explored the possibilities of layering and preclusion in a fly-pollinated iris, Moraea lurida.

Results and Conclusions

The proportion of labeled pollen from the last flower visited diminished in sequential pollen samples taken from the top to the bottom of the pollen load, representing the first empirical evidence for pollen layering. However, the consequences in terms of pollen preclusion were equivocal: Although the pre-existing pollen load size was not a good predictor of new pollen receipt, labeled pollen loads from the last flower visited were significantly smaller than pollen loads from the previous flower visited. Thus, pollen from the previous flower may preclude pollen placement from a subsequently visited flower, and pollen from different flowers may compete for space on pollinators.

     

Pollen competition between morphs in a pollen‐color dimorphic herb and the loss of phenotypic polymorphism within populations

Flower color polymorphism is relatively uncommon in natural flowering plants, suggesting that maintenance of different color morphs within populations is difficult. To address the selective mechanisms shaping pollen‐color dimorphism, pollinator preferences and reproductive performance were studied over three years in Epimedium pubescens in which some populations had plants with either green or yellow pollen (and anthers). Visitation rate and pollen removal and receipt by the bee pollinator (Andrena emeishanica) did not differ between the two color morphs. Compared to the green morph, siring success of the yellow morph's pollen was lower, but that of mixtures of pollen from green and yellow morphs was lowest. This difference, corresponding to in vivo and ex vivo experiments on pollen performance, indicated that pollen germination, rather than tube growth, of the green morph was higher than that of the yellow morph and was seriously constrained in both morphs if a pollen competitor was present. A rare green morph may invade a yellow‐morph population, but the coexistence of pollen color variants is complicated by the reduced siring success of mixed pollinations. Potential pollen competition between morphs may have discouraged the maintenance of multiple phenotypes within populations, a cryptic mechanism of competitive exclusion.     

POLLEN PERFORMANCE AND SEX-RATIO EVOLUTION IN A DIOECIOUS PLANT

There has been a proliferation of studies, in a variety of taxa, that have detected sex-linked or cytoplasmic genes that enhance their own transmission via sex-ratio distortion. One of the most important parameters influencing the dynamics of these elements is the magnitude of their transmission advantage. In many systems, the mechanism of sex-ratio distortion is to abort X- or Y-bearing gametes. With this mechanism, the transmission advantage associated with sex-ratio distortion is diminished when the production of male gametes limits offspring production or when competition among the gametes of different males is intense. In this study, we analyzed the outcome of pollen competition between males that produced different sex ratios in the dioecious plant, Silene alba, and estimated how the sex-ratio bias influenced the transmission properties of the sex chromosomes. We varied the intensity of pollen competition by controlling the quantity of pollen used in crosses and used a combination of single-male pollinations and pollen mixtures to evaluate the effects of multiple paternity. Paternity in pollen mixtures was estimated using allozymes. Sex-ratio bias was directly influenced by the quantity of pollen, but the magnitude of this effect was small. The relative performance of pollen from different males varied substantially, especially when there was multiple paternity. Specifically, males with biased sex ratios sired far fewer offspring of either sex in pollen mixtures. In crosses involving single males, however, these “sex-ratio” males produced the same number of offspring as other males, so the female bias caused a significant transmission advantage for X-linked genes. X-linked genes could enhance their transmission via sex-ratio distortion in Silene populations, but the magnitude of this transmission advantage will depend on the ecological circumstances that influence the opportunity for multiple paternity.     

PATERNAL EFFECTS IN INHERITANCE OF A PATHOGEN RESISTANCE TRAIT IN IPOMOEA PURPUREA

For continuously variable, polygenic characters, the response to selection depends upon the proportion of phenotypic variance that is caused by additive genetic variance, or narrow-sense heritability. Thus, a major goal of quantitative genetics is to partition phenotypic variance for a trait in a way that isolates additive genetic variance from other causes. The variance among paternal half-sib families, which is frequently used to estimate additive variance, is commonly recognized to include additive epistatic effects. However, this variance component can also include non-Mendelian paternal effects. We report here the results from a diallel crossing design used to isolate nonnuclear effects from the paternal nuclear contribution to disease resistance in the common morning glory, Ipomoea purpurea. In particular, we found that genetic variance for resistance to anthracnose, a disease caused by the fungal pathogen Colletotrichum dematium, was determined largely by a nonnuclear, additive paternal effect. We discuss potential mechanisms for this effect as well as some of their evolutionary implications.     

Environmental Effects on Components of Pollen Performance in Faramea occidentalis (L.) A. Rich. (Rubiaceae)1

Studies focusing on gametophytic competition have focused on differences in pollen performance (e.g., pollen germination rate and pollen tube growth rate) among donors in order to examine genetic sources of variation in siring success among them. Donors that produce the fastest growing pollen tubes are expected to fertilize more ovules relative to donors with slow growing pollen tubes under conditions of gametophytic competition. However, the performance of pollen in the field is known to be influenced by environmental conditions in addition to the genotype of donor plants. This field study of Faramea occidentalis was conducted to: (1) determine the effect of environmental conditions during pollination on pollen performance; (2) measure differences among pollen donors in pollen performance; (3) determine if the pollen of different donors responds differently to a variety of environmental conditions surrounding pollinated flowers; and (4) measure differences among pollen recipients in pollen performance. Single-donor crosses were made between four pollen donors and four pollen recipients under a variety of environmental conditions. Pollen performance was then quantified as the growth rate of the fastest pollen tube, the mean pollen tube growth rate, and by a pollen germination index. Pollination environment (the environment surrounding a pollinated flower) and recipient significantly affected all three measures of pollen performance. Pollen donors did not differ overall in pollen performance. However, there was significant among-donor variation in two of the five pollination environment conditions. Future studies of variation in relative siring success may benefit by considering pollination environment in addition to donor and recipient identity.     

Pollen—Tiny and ephemeral but not forgotten: New ideas on their ecology and evolution

Ecologists and evolutionary biologists have been interested in the functional biology of pollen since the discovery in the 1800s that pollen grains encompass tiny plants (male gametophytes) that develop and produce sperm cells. After the discovery of double fertilization in flowering plants, botanists in the early 1900s were quick to explore the effects of temperature and maternal nutrients on pollen performance, while evolutionary biologists began studying the nature of haploid selection and pollen competition. A series of technical and theoretic developments have subsequently, but usually separately, expanded our knowledge of the nature of pollen performance and how it evolves. Today, there is a tremendous diversity of interests that touch on pollen performance, ranging from the ecological setting on the stigma, structural and physiological aspects of pollen germination and tube growth, the form of pollen competition and its role in sexual selection in plants, virus transmission, mating system evolution, and inbreeding depression. Given the explosion of technical knowledge of pollen cell biology, computer modeling, and new methods to deal with diversity in a phylogenetic context, we are now more than ever poised for a new era of research that includes complex functional traits that limit or enhance the evolution of these deceptively simple organisms.     

Pollen competition in a natural population of Cucurbita foetidissima (Cucurbitaceae)

The pollen competition hypothesis predicts that when the number of pollen grains deposited onto stigmas exceeds the number of ovules, selection can operate in the time frame between deposition and fertilization. Moreover, because of the overlap in gene expression between the two phases of the life cycle, selection on microgametophytes may alter the resulting sporophytic generation. The extent to which pollen competition occurs in nature has been unclear, because tests of the predictions of the pollen competition hypothesis have used cultivars and/or artificial growth conditions and hand-pollination techniques. In this study we used a wild species, Cucurbita foetidissima, in its natural habitat (southern New Mexico) to determine the amount and timing of the arrival of pollen onto stigmas, the relationship between pollen deposition and seed number, and the effects of the intensity of pollen competition on progeny vigor. We found that ～900 pollen grains are necessary for full seed set and that a single visit by a pollinator results in the deposition of 653.0 ± 101.8 pollen grains. About 29% of the flowers receiving a single pollinator visit had 900 or more pollen grains on its stigma. Moreover, within 2 h of anthesis, >4000 pollen grains were deposited onto a typical stigma, indicating that multiple pollinator visits must have occurred. Fruits produced by multiple visits had greater seed numbers (206 vs. 147) than fruits produced by a single visit. Finally, the progeny produced by multiple pollinator visits were more vigorous than those produced by single visits with respect to five measures of vegetative growth (MANCOVA, Wilks' lambda = 0.96, F(6,370) = 2.54, P < 0.02. These data demonstrate that conditions for pollen competition exist in nature and support the prediction that pollen competition enhances offspring vigor.     

Pollen-tube growth rate and seed-siring success among Betula pendula clones

The aim of this study was to investigate whether genetically different pollen donors ( Betula pendula clones) differed in pollen-tube growth rate across 11 maternal plants and in vitro , and whether the differences between the donors were consistent across the recipients. To compare the seed-siring success of competing pollen donors, a two-donor hand-pollination experiment with six donors and six recipients was conducted. The experiments were performed at a plastic-house seed orchard. The donors showed significant variation in pollen-tube growth rate on all the 11 recipients. The rankings of the pollen donors were statistically consistent across different maternal plants. A significant positive correlation between pollen tube growth in vivo and in vitro was found. The seed-siring success of two competing pollen donors was unequal in 20 of 29 cases and there was a significant positive correlation between seed-siring success and pollen-tube growth rate in vivo and in vitro . The results show that fertilizations are not random and pollen competition operates in a B. pendula seed orchard population.     

Reproductive isolation and hybrid pollen disadvantage in Ipomopsis

One cause of reproductive isolation is gamete competition, in which conspecific pollen has an advantage over heterospecific pollen in siring seeds, thereby decreasing the formation of F1 hybrids. Analogous pollen interactions between hybrid pollen and conspecific pollen can contribute to post-zygotic isolation. The herbaceous plants Ipomopsis aggregata and I. tenuituba frequently hybridize in nature. Hand-pollination of I. aggregata with pollen from F1 or F2 hybrids produced as many seeds as hand-pollination with conspecific pollen, suggesting equal pollen viability. However, when mixed pollen loads with 50% conspecific pollen and 50% hybrid pollen were applied to I. aggregata stigmas, fewer than half of the seeds had hybrid sires. Such pollen mixtures are frequently received if plants of the two species and F1 and F2 hybrids are intermixed, suggesting that this advantage of conspecific over hybrid pollen reduces backcrossing and contributes to reproductive isolation.     

Incompatibility and pollen competition in Alnus glutinosa: Evidence from pollination experiments

Different types of incompatibility systems were found to operate simultaneously in alnus glutinosa in the course of numerous pollination experiments, including self-pollination and pollination with controlled pollen mixtures. Isozyme genetic markers were used to identify the pollen parent of each offspring from the mixed pollination experiments, thus allowing specification of the fertilization success of each pollen parent. In a first step, these results were compared with observations on in vitro pollen germination experiments. This comparison allows for exploration of the explanatory value of different germination media as models of germination conditions on stigmas. In most cases, the data suggest that the in vitro germination conditions resemble the fertilization conditions in vivo, at least in the sense that they favor the same pollen parents. By providing a generic and operable definition of the two basic types of incompatibility, eliminating (inability to fertilize ovules) and cryptic (resulting in lowered fertilization success of a pollen parent under competition), evidence was detected for the existence of both types of incompatibility in alnus glutinosa, where eliminating incompatibility occurred as self-incompatibility only. However, since this incompatibility seems to act primarily via pollen elimination, seed production is not likely to be negatively affected in natural populations, even for comparatively large amounts of self-pollination. This revised version was published online in July 2006 with corrections to the Cover Date.     

Male interference with pollination efficiency in a hermaphroditic orchid

Hermaphroditism can lead to both intra‐ and intersexual conflict between male and female gender functions. However, the effect that such gender conflicts have on pollination efficiency has seldom been investigated. By artificially reducing the number of available male gametes on an individual, we quantified whether male interference with pollination efficiency occurs in the self‐compatible, moth‐pollinated orchid Satyrium longicauda. We partially emasculated S. longicauda inflorescences and compared pollination success and fecundity in these plants to intact controls. Pollen in both groups of plants was colour‐labelled so that its dispersal by pollinators could be tracked directly in the field. Intact flowers on partially emasculated inflorescences exported more pollen and received more cross‐pollen and less self‐pollen than those on intact inflorescences. Proportion of fruit set per plant was similar between the two treatments; however, fruits on partially emasculated plants had proportionally more viable seeds than those on intact controls. These results provide empirical evidence that male interference with pollination efficiency can occur in a hermaphroditic plant and that such interference can compromise fecundity. The most likely mechanism for such male interference is competition for placement on the proboscis of hawkmoth pollinators. Consequently, male competition for siring success may influence the evolution of sexual systems in hermaphroditic pollinator‐dependent plants.