Pollen-pistil size correlation and pollen size-number trade-off in species of Argentinian Nyctaginaceae with different pollen reserves

Data on pollen and pistil traits from 14 Argentinean Nyctaginaceae species with starch or lipids as pollen reserves are presented. We expect differences in the traits between these two groups of species, but the same pattern within each group for (a) the relationship between pollen size and pistil length assuming that pollen tube length is predetermined by provisions in the pollen independently of the pollen reserve type, and for (b) a trade-off between size and number because available resources for male function are not unlimited. In particular we expect that (a) species with longer pistils will have larger pollen grains, (b) pollen grain size and the number of pollen grains per flower will be negatively correlated. Significant differences in the mean pollen number per flower and mean pollen size were observed between species with different pollen reserve type but not for pistil length. Then, correlation analyses were performed for species with starchy pollen or with pollen with lipids separately. Pollen size - pistil length correlation was positive and significant for species with starchy pollen but not for species with pollen with lipids. On the other hand, pollen size - number correlation was not significant for both starchy and oil-rich species. Results suggest that pollen reserve type would be a relevant factor that constraint pollen size in species of Nyctaginaceae, and that this pollen trait should be considered when studying pollen-pistil relationships, mainly between species of those families with mixed pollen reserves.     

The effects of pollen load size and donor diversity on pollen performance, selective abortion, and progeny vigor in Mirabilis jalapa

The influence of pollen competitive environment on pollen performance (pollen germination, stigmatic penetration, and pollen tube growth rate), the maturation or abortion of initiated fruit, seed size, and seedling vigor was explored by manipulating the size and diversity of stigmatic pollen loads on Mirabilis jalapa. All aspects of pollen performance significantly increased with the number of pollen grains on a stigma or pollen tubes in a style, but was not influenced by the diversity of pollen donors. Plants tended to mature single-ovulate fruits that came from flowers where pollen load size and diversity were greatest and aborted those where these were lowest. No plants from seeds resulting from pollinations with a single pollen grain survived, but other fitness measures were mostly determined by maternal plant. The data suggest that pollen performance is influenced by pollen competitive environment, and both the genetic diversity of the pollen load and number of competing pollen tubes are important determinants of seed/fruit abortion.     

Pollen–pistil relationships and pollen size-number trade-off in species of the tribe Lycieae (Solanaceae)

Pollen volume may be involved in different associations with other floral traits. Particularly, the literature indicates that pollen volume can be implicated in a functional relationship with pistil length, and that it may be affected by a trade-off with pollen number because of the subdivision of limited resources. To assess these associations, pollen volume was subjected to correlation analyses with pollen number and pistil length in 20 Argentinean and Chilean taxa of the monophyletic tribe Lycieae. Depending on the mechanisms operating on pollen size and number, the variability of these traits may be different. Therefore, their coefficients of variation were compared. Pistil length and pollen volume showed a strong positive correlation. In contrast, pollen grain size and number were significantly correlated at neither inter- nor intraspecific levels. Results suggest that pollen size and pistil length may co-evolve. The central role of this interaction is discussed. The lack of a correlated variation in pollen size and number may be related to the similar constraints experienced by the species and/or because constraints are operating at the plant level and not at the species level. Lower variance in pollen size compared with pollen number denotes that pollen size may be the main trait subjected to natural selection. Electronic Publication     

Pollen grain size variation in Caryophylloideae: a mixed strategy for pollen deposition along styles with long stigmatic areas?

The literature supports a positive correlation between pollen size and style length and that pollen size may be affected by a trade-off with pollen number. A correlation between pollen size and pollinator type has been hypothesized by some authors but not confirmed. To assess these associations, pollen grain number, pollen grain diameter and style length were measured in 53 species of Caryophylloideae that had been classified beforehand as being diurnal, nocturnal or self-pollinated. Surprisingly, many species showed two, and a few even three, distinct pollen grain size classes per individual anther. Therefore, further analyses were based on the different pollen size classes and respective pollen grain numbers. Firstly, we found no indication of a trade-off between pollen size and number. Secondly, a positive linear correlation between style length and pollen grain diameter was found using conventional analysis methods and phylogenetically independent contrasts. Thirdly, when using conventional analysis methods a strong relationship between style length and pollination mode was found, with nocturnal species having significantly longer styles and larger pollen grains than diurnal and selfing species. These results were not supported, however, when calculating comparisons of phylogenetically independent pairs. We discuss our findings in relation to intra-specific pollen polymorphism and pollen degeneration, both of which have been associated with different pollen grain sizes in Caryophylloideae. Furthermore, we discuss the possibility that pollen polymorphism might be a mixed strategy to optimise the success of pollen grains deposited at different pollen–ovule distances on long styles.     

Effects of pollen load size on seed paternity in wild radish: the roles of pollen competition and mate choice

For sexual selection to be important in plants, it must occur at pollen load sizes typical of field populations. However, studies of the impact of pollen load size on pollen competition have given mixed results, perhaps because so few of these studies directly examined the outcome of mating when pollen load size was varied. We asked whether seed paternity after mixed pollination of wild radish was affected by pollen load sizes ranging from 22 to 220 pollen grains per stigma. We examined the seed siring abilities of 12 pollen donors across 11 maternal plants. Seed paternity was statistically indistinguishable across the pollen load sizes even though, overall, the pollen donors sired different numbers of seeds. This lack of effect of pollen load size on seed paternity may have occurred because fruit abortion and early abortion or failure of fertilization of seeds increased as load size decreased. Thus, failures of fruits and seeds sired by poorer pollen donors may keep seed paternity constant across pollen load sizes.     

VARIATION IN POLLEN SIZE,FERTILIZATION ABILITY,AND POSTFERTILIZATION SIRING ABILITY IN ERYTHRONIUM GRANDIFLORUM

The mean volume of pollen grains and total pollen production varied both within and among plants of Erythronium grandiflorum. The second flowers of two-flowered plants tended to produce smaller and fewer grains than first flowers, but there was no overall relationship between mean pollen grain size and production per flower. I evaluated the effects of pollen size differences within and among plants on two components of male reproductive success: pollen tube growth and postfertilization siring ability. Pollen tubes grown in media were longer for second flowers, but were not correlated with the mean size of pollen grains, suggesting that (1) internal resource content of pollen (i.e., carbohydrates plus lipids) was not associated with the hydrated size of pollen, and that (2) pollen from second flowers contained more resources. I analyzed the growth rate and the fertilization ability of pollen growing in styles. Growth rate differed among donors and recipients, but no effects of pollen or donor characters (i.e., pollen production, grain size, and flower position) were detected. In single donor pollinations, pollen size was negatively correlated with fertilization ability across donors, and positively correlated with postfertilization siring ability of donors. A second experiment used pairs of donors; within-plant differences in pollen size and flower position had effects similar to the single donor experiment on fertilization ability, but among-plant differences were not significant. The results corroborate earlier experiments that suggest that the growth of pollen tubes in the style is probably controlled by the recipient, since donor characters had minimal effects on pollen fertilization ability. Postfertilization siring ability was not affected by within-plant differences in mean grain size and production. For among-donor differences, the number of seeds set for each donor was positively correlated with the mean grain volume, and when a donor producing large pollen fertilized ovules in an ovary, there was increased seed abortion for seeds in the same ovary sired by a second donor. In addition, the total number of seeds produced by a fruit was decreased when both donors had large pollen, apparently due to increased postfertilization abortion. Postfertilization processes appear to be influenced by paternal differences that are expressed through competition among developing seeds for maternal resources.     

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.     

Direct and indirect responses to selection on pollen size in Brassica rapa L.

Pollen size varies little within angiosperm species, but differs extensively between species, suggesting the action of strong selection. Nevertheless, the potential for genetic responses of pollen size to selection, as determined by additive genetic variance and genetic correlations with other floral traits, has received little attention. To assess this potential, we subjected Brassica rapa to artificial selection for large and small pollen during three generations. This selection caused significant divergence in pollen diameter, with additive genetic effects accounting for over 30% of the observed phenotypic variation in pollen size. Such heritable genetic variation suggests that natural selection could effect evolutionary change in this trait. Selection on pollen size also elicited correlated responses in pollen number (–), flower size (+), style length (+), and ovule number (+), suggesting that pollen size cannot evolve independently. The correlated responses of pollen number, flower size and ovule number probably reflect the genetically determined and physically constrained pattern of resource allocation in B. rapa. In contrast, the positive correlation between pollen size and style length may represent a widespread gametic‐phase disequilibrium in angiosperms that arises from nonrandom fertilization success of large pollen in pistils with long styles.     

Volume-based pollen size analysis: an advanced method to assess somatic and gametophytic ploidy in flowering plants

Pollen size is often used as a biological parameter to estimate the ploidy and viability of mature pollen grains. In general, pollen size quantification is performed one- or two-dimensionally using image-based diameter measurements. As these approaches are elaborate and time consuming, alternative approaches that enable a quick, reliable analysis of pollen size are highly relevant for plant research. In this study, we present the volume-based particle size analysis technique as an alternative method to characterize mature pollen. Based on a comparative assay using different plant species (including tomato, oilseed rape, kiwifruit, clover, among others), we found that volume-based pollen size measurements are not biased by the pollen shape or position and substantially reduce non-biological variation, allowing a more accurate determination of the actual pollen size. As such, volume-based particle size techniques have a strong discriminative power in detecting pollen size differences caused by alterations in the gametophytic ploidy level and therefore allow for a quick and reliable estimation of the somatic ploidy level. Based on observations in Arabidopsis thaliana gametophytic mutants and differentially reproducing Boechera polyantha lines, we additionally found that volume-based pollen size analysis provides quantitative and qualitative data about alterations in male sporogenesis, including aneuploid and diploid gamete formation. Volume-based pollen size analysis therefore not only provides a quick and easy methodology to determine the somatic ploidy level of flowering plants, but can also be used to determine the mode of reproduction and to quantify the level of diplogamete formation.     

Pollen-size comparisons among animal-pollinated angiosperms with different pollination characteristics

Pollen volume varies among angiosperm species over five orders of magnitude, presumably because the functional advantages of pollen size depend on each species' particular pollination and fertilization conditions. This paper reports two studies that assess whether animal-pollinated species with different pollination systems differ correspondingly in pollen size, as would be expected if pollen size affected pollen transport. Analysis of nine Pedicularis species detected significant interspecific variation in pollen size; however, the overall mean pollen volume of species pollinated primarily by nectar-collecting bees did not differ significantly from that of species pollinated by pollen-collecting bees. Similarly, comparison of bee- and bird-pollinated congeners in 16 genera (nine families) from Australia and North America found considerable variation in pollen diameter within and between genera, but this variation was not associated consistency with differences in primary pollinator type (bee>bird for three genera, bee > bird for five genera, bee相似文献   

Pollen size in Carex: The effect of different chemical treatments and mounting media

Pollen size and pollen aperture size for ten species of the genus Carex L., native to Estonia, have been measured using light microscopy. The species selected represent different sections of the genus, a range of habitats and different chromosome numbers. The effects of two basic chemical treatments, two mounting media and the effect of chemically induced dehydration with tertiary butyl alcohol (TBA) on the size of pollen grains were then recorded.

In general pollen size and pollen aperture size of the species examined is highly variable at both intraspecific and interspecific levels. Carex hirta has notably larger pollen grains than any of the other species investigated and, although correlations between size and chromosome number in the species examined are limited, it also has the highest chromosome number. Statistically significant size differences resulted from variations in chemical treatment, mounting media and tertiary butyl alcohol (TBA) induced dehydration. Acetolysed pollen grains are larger than potassium hydroxide (KOH) treated pollen grains. Pollen grains dehydrated after chemical treatment with TBA are larger than pollen grains not dehydrated. Pollen grains mounted in silicon oil are smaller than grains mounted in glycerine. But considering the great size variation of Carex pollen grains, the size changes caused by preparation procedures fall within the size variation range of the species examined.

All the samples contained a high number of deformed pollen grains and pollen grains with hardly distinguishable or no lateral apertures.     

Interspecific size and number variation in pollen grains and seeds

An interspecific correlation between pollen grain size and seed size is demonstrated by means of the phylogenetic regression, which allows for phylogenetic bias. The correlation was not explained by plant size, mass of DNA per cell, style length or breeding system, although the first three of these factors all correlated with both pollen size and seed size. Two interpretations, involving pollen competition and flower size, are discussed. There is also an interspecific correlation between pollen grain number per flower and ovule number per flower. Some consequences of these correlations for the interpretation of pollen-ovule ratios and sex allocation strategies are considered.     

Size-number trade-offs and pollen production by Papilionaceous legumes

The intra- and interspecific relations between the size and number of pollen grains were examined for 21 species of papilionaceous legumes to test for the trade-off expected from the subdivision of limited resources. We observed a tradeoff between pollen grain diameter and the number of grains produced per flower within 17 of the 21 species examined. For 12 of these species, the observed partial regression coefficient for In (pollen grain diameter) equaled -3, as expected. The remaining five species exhibited more negative partial regression coefficients than expected. Relations between pollen grain size and number that are more negative than expected may result from a decelerating relation between pollen grain size and the resource investment per grain. Flower production significantly influenced pollen production for only eight species so that pollen production seems to be determined primarily on a per flower basis. Interspecifically, pollen size also varied inversely with pollen number for the 21 species even after the effect of phylogenetic relatedness was removed. Pollen size varies relatively little compared to the number of pollen grains produced per flower and therefore may be the primary target of natural selection. The presence of a trade-off in pollen production probably constrains the evolutionary options by which flowering plants can maximize male success.     

Large pollen at high temperature: an adaptation to increased competition on the stigma?

Pollen availability is a major constraint of plant reproductive success. Because pollen size trades-off with the quantity of produced grains, the link between climate characteristics and the determination of pollen size is of fundamental importance. To minimize the rate of water loss due to desiccation, a plant should produce larger grains that also have a lower surface-to-volume ratio. We used a comparative analysis to examine the hypothesis predicting increase in pollen size as a response to desiccation intensity. To test the hypothesis, we correlated the data on pollen size with the climate characteristics, temperature and desiccation intensity of the flowering period, for 232 plant species of 11 taxonomic groups. The analysis showed a positive relationship between the pollen size and temperature, but not with the desiccation intensity. We discuss the potential mechanisms by which increased temperature is an indicator of high competition among pollen grains on the stigma, which in turn is expected to promote large pollen. Our work provides insight into the temperature dependence of pollen production in plants and reveals a link between environmental temperature and the intensity of limitation of plant reproductive success by pollen availability. The result is relevant in the context of global climate change. We also discuss why environmental temperature has to be controlled in studies dealing with pollen production, particularly in investigations of size-number trade-off.     

Pollen and stigma size changes during the transition from tristyly to distyly in Oxalis alpina (Oxalidaceae)

• Pollen and stigma size have the potential to influence male fitness of hermaphroditic plants, particularly in species presenting floral polymorphisms characterised by marked differences in these traits among floral morphs. In this study, we take advantage of the evolutionary transition from tristyly to distyly experienced by Oxalis alpina (Oxalidaceae), and examined whether modifications in the ancillary traits (pollen and stigma size) respond to allometric changes in other floral traits. Also, we tested whether these modifications are in accordance with what would be expected under the hypothesis that novel competitive scenarios (as in distylous‐derived reproductive system) exert morph‐ and whorl‐specific selective pressures to match the available stigmas.
• We measure pollen and stigma size in five populations of O. alpina representing the tristyly–distyly transition.
• A general reduction in pollen and stigma size occurred along the tristyly–distyly transition, and pollen size from the two anther levels within each morph converged to a similar size that was characterised by whorl‐specific changes (increases or decreases) in pollen size of different anthers in each floral type.
• Overall, results from this study show that the evolution of distyly in this species is characterised not only by changes in sexual organ position and flower size, but also by morph‐specific changes in pollen and stigma size. This evidence supports the importance of selection on pollen and stigma size, which increase fitness of remaining morphs following the evolution of distyly, and raises questions to explore on the functional value of pollen size in heterostylous systems under pollen competition.

     

The relationship between pollen grain size and progeny gender in dioecious Silene latifolia (Caryophyllaceae)

Summary Experiments and observations conducted during the past 90 years have provided conflicting evidence concerning the existence of a size difference between pollen grains containing an X chromosome (female-determining) and those containing a Y chromosome (maledetermining) in dioecious Silene latifolia. Were such a size difference to exist, this might explain, at least in part, the observation that X-bearing pollen tubes reach the ovary more quickly, on average, than Y-bearing pollen tubes. We tested for such a size difference by separating pollen collected from single anthers into three size classes: small, large, and random. Fruit set (number of pollinated flowers that set fruit) and seed set (number of seeds per capsule) did not differ for these three pollination treatments. Progeny sex ratios resulting from these three pollen size classes also did not differ significantly. Thus, pollen grain size is not affected by which of the two sex chromosomes is present. Our experiment is the first direct test of this relationship. Based on our results, size differences should not be invoked to explain competitive differences in male- and female-determining microgametophytes.     

Effects of soil fertility and mycorrhizal infection on pollen production and pollen grain size of Cucurbita pepo (Cucurbitaceae)

The effects of soil fertility (two levels of soil nitrogen and two levels of soil phosphorus) and mycorrhizal infection on pollen production and pollen grain size were studied in two cultivars of the common zucchini (Cucurbita pepo). Overall, soil fertility and mycorrhizal infection had significant effects on traits affecting the male function of plants (staminate flower production, pollen production per flower and pollen grain size). There were also differences between the cultivars for these male traits in all three experiments. In addition, pollen grain size decreased toward the end of the growing season. In the mycorrhiza experiment, both phosphate concentration per pollen grain and total phosphate content per anther were greater but not significantly greater in the mycorrhizal plants than in the non-mycorrhizal plants. A significant negative relationship between pollen production and pollen grain size was found in the mycorrhiza and soil phosphorus experiments, indicating that there was a trade-off between pollen production and pollen size. This study is the first to show that mycorrhizal infection has an effect on male function (pollen production and size) in addition to the well-documented effects on female function (fruit/seed production and size).     

The interaction between pollinator size and the bristle staminode of Penstemon digitalis (Scrophulariaceae)

Penstemon digitalis, a prairie species whose flowers possess a large bristle staminode, is visited by eight bee species of varying size. Two sets of field experiments involving staminode removal were performed to test pollination efficiency in relation to bee size. Our data indicate that bristle staminode presence and function are influenced by size-dependent selection on bee body size and associated pollen transporting attributes. The first experiment compared staminode presence and removal in open-pollinated flowers. Staminode removal significantly reduced total pollen deposition but had no effect on total pollen removal. The second experiment utilized single bee visits to assess the interaction between pollinator size and staminode presence on the rate of pollen deposition and removal. This experiment indicated that staminode removal resulted in fewer pollen grains deposited on stigmas and less pollen removed from anthers for both large and small bees (the contrary was true for medium bees). Although the number of pollen grains deposited was greatly reduced for large bees, staminode removal reduced deposition efficiency most notably for small bees by 41.3%, reducing female reproductive success. Staminode removal increased pollen removal efficiency most notably for medium bees by 22.7%, reducing male reproductive success. Mechanisms of staminode function are discussed.     

A model for seed size variation among plants

Summary I developed a model for seed size variation among plants assuming that the pollen captured per flower depends on both the allocation to pollen capture mechanisms per flower and the number of flowers on each plant. I showed that the optimal seed size increases with (1) the total resource allocation to reproduction, (2) decreasing outcross pollen availability, (3) decreasing probability of seedling establishment and (4) decreasing selfing rate. However, optimal seed size does not depend on the total resource allocation if the total number of pollen grains captured by a plant increases linearly with its flower number. In addition, the optimal seed size is not always positively correlated with the optimal resource allocation to pollen capture mechanisms per flower. I discussed implications of the results for seasonal decline in seed size and seed size variations among populations, such as alutitudinal variation.     

Branching habit and the allocation of reproductive resources in conifers

Background and Aims Correlated relationships between branch thickness, branch density, and twig and leaf size have been used extensively to study the evolution of plant canopy architecture, but fewer studies have explored the impact of these relationships on the allocation of reproductive resources. This study quantifies pollen cone production in conifers, which have similar basic reproductive biology but vary dramatically in branching habit, in order to test how differences in branch diameter influence pollen cone size and the density with which they are deployed in the canopy. Methods Measurements of canopy branch density, the number of cones per branch and cone size were used to estimate the amount of pollen cone tissues produced by 16 species in three major conifer clades. The number of pollen grains produced was also estimated using direct counts from individual pollen cones. Key Results The total amount of pollen cone tissues in the conifer canopy varied little among species and clades, although vegetative traits such as branch thickness, branch density and pollen cone size varied over several orders of magnitude. However, branching habit controls the way these tissues are deployed: taxa with small branches produce small pollen cones at a high density, while taxa with large branches produce large cones relatively sparsely. Conclusions Conifers appear to invest similar amounts of energy in pollen production independent of branching habit. However, similar associations between branch thickness, branch density and pollen cone size are seen across conifers, including members of living and extinct groups not directly studied here. This suggests that reproductive features relating to pollen cone size are in large part a function of the evolution of vegetative morphology and branching habit.