EVOLUTIONARILY STABLE SELFING RATES OF HERMAPHRODITIC PLANTS IN COMPETING AND DELAYED SELFING MODES WITH ALLOCATION TO ATTRACTIVE STRUCTURES

I present a resource-allocation model to analyze how patterns of allocation to reproductive structures influence the evolution of selfing rates in hermaphrodites subject to competing and delayed forms of self-fertilization. The evolutionarily stable state does not depend on the mode of pollination. In contrast to previous models in which the number and the size of flowers were not considered, intermediate selfing is not evolutionarily stable with linear constraints on flower number and size. In contrast, intermediate selfing can be evolutionarily stable with nonlinear constraints on flower number and size. Optimal allocations to attractive structures increase and selfing rates decrease in the presence of inbreeding depression. In particular, stable intermediate levels of selfing may be favored when flower number is strongly constrained. Thus, nonlinear constraints on flower number and size could favor the evolution of intermediate selfing in either the delayed or the competing modes of selfing. Outcrossing is not favored in the absence of inbreeding depression, a result inconsistent with Holsinger's results in which allocation to attractive structures was not considered.     

Trade-offs between flower number and investment to a flower in selfing and outcrossing varieties of Impatiens hypophylla (Balsaminaceae)

Floral resource allocation was compared on a whole-plant basis between two varieties of Impatiens hypophylla that differ in flower size. There were significant negative correlations between flower number and investments to a flower at both the within-population and between-variety levels. In individual flowers, var. hypophylla with larger flowers invested significantly more resources to male and pollinator-attractive functions, whereas investments to female function did not differ between the varieties. In experimental populations placed in the field, pollinators preferred the larger flowers of var. hypophylla even within the same habitat of var. microhypophylla, which has smaller flowers. There was a significant lack of observed heterozygosity only in var. microhypophylla. Thus, the outcrossing variety had more attractive but fewer flowers, while the selfing variety had less attractive but more abundant flowers.     

Production of male flowers does not decrease with plant size in insect‐pollinated Sagittaria trifolia,contrary to predictions of size‐dependent sex allocation

Abstract It has been proposed that relative allocation to female function increases with plant size in animal‐pollinated species. Previous investigations in several monoecious Sagittaria species seem to run contrary to the prediction of size‐dependent sex allocation (SDS), throwing doubt on the generalization of SDS. Plant size, phenotypic gender, and flower production were measured in experimental populations of an aquatic, insect‐pollinated herb Sagittaria trifolia (Alismataceae) under highly different densities. The comparison of ramets produced clonally can reduce confounding effects from genetic and environmental factors. In the high‐density population, 48% of ramets were male without female flowers, but in the low‐density population all ramets were monoecious. We observed allometric growth in reproductive allocation with ramet size, as evident in biomass of reproductive structures and number of flowers. However, within both populations female and male flower production were isometric with ramet size, in contrast to an allometric growth in femaleness as predicted by SDS. Phenotypic gender was not related to ramet size in either population. The results indicated that large plants may increase both female and male function even in animal‐pollinated plants, pointing towards further studies to test the hypothesis of size‐dependent sex allocation using different allocation currencies.     

Life history trade‐offs and evidence for hierarchical resource allocation in two monocarpic perennials

The evolution of floral display is thought to be constrained by trade‐offs between the size and number of flowers; however, empirical evidence for the trade‐off is inconsistent. We examined evidence for trade‐offs and hierarchical allocation of resources within and between two populations each of the monocarpic perennials, Cardiocrinum cordatum and C. giganteum. Within all populations, flower size–number trade‐offs were evident after accounting for variation in plant size. In addition, variation in flower size explained much variation in flower‐level allocation to attraction, and female and male function, a pattern consistent with hierarchical allocation. However, between population differences in flower size (C. cordatum) and number (C. giganteum) were not consistent with size–number trade‐offs or hierarchical allocation. The population‐level difference in C. cordatum likely reflects the combined influence of a time lag between initiation and maturation of flowers, and higher light levels in one population. Thus, our study highlights one mechanism that may account for the apparent independence of flower size and number in many studies. A prediction of sex allocation theory was also supported. In C. giganteum: plants from one population invested more mass in pistils and less in stamens than did plants from the other population. Detection of floral trade‐offs in Cardiocrinum may be facilitated by monocarpic reproduction, production of a single inflorescence and ease of measuring plant size.     

Invasion of unisexuals in hermaphrodite populations of animal-pollinated plants: effects of pollination ecology and floral size-number trade-offs

Abstract.— The most common sexual system in animal-pollinated plants is hermaphroditism, while some species are dioecious or gynodioecious and a very few are androdioecious. In this paper, I attempt to explain this pattern by extending previous models for the evolution of sexual systems to incorporate two main features: (1) a portion of investment in pollinator attraction contributes to only female or male function, because one sexual function of a flower is saturated with pollinator visitation earlier than the other sexual function; and (2) there are trade-offs between the size and number of flowers. The analysis was conducted to determine the conditions when females and males can increase in frequency in a hermaphroditic population, assuming either concave or convex pollinator gain curves (relation between investment to attractive structures of a flower and frequency of pollinator visits to the flower). The results suggest that both of the main factors play important roles in the evolution of plant sexual systems: uneven contribution of pollinator-attractive structures and nonlinear trade-offs between flower size and number can destabilize hermaph-roditism. When a convex pollinator gain curve was assumed, the effect of nonlinear trade-offs can produce accelerating compensation from the elimination of one sexual function, allowing males to increase for large regions of parameter space, where females could not increase. The last prediction obviously conflicts with the observed rarity of androdioecy in nature, indicating the necessity of exploring pollinator gain curves in more detail.     

Evidence for reductions in floral attractants with increased selfing rates in two heterandrous species

Although theoretical models predict low allocation to attractive structures with increased selfing in animal-pollinated plants, empirical measurement of the reproductive costs and benefits is complicated. Here, floral sex allocation was compared in two nectarless heterandrous species with different mating systems: Monochoria korsakowii (Pontederiaceae), which has moderate outcrossing rates, and Monochoria vaginalis, a predominant selfer. In both species, mirror-image flowers have one large dark-purple anther and five small yellow anthers. Experimental evidence is provided for functional differences between the two sets of anthers using data on pollinator visitation, pollen removal and deposition, and seed set after hand pollinations. Flower manipulations in bee-pollinated M. korsakowii demonstrated different functions of the two sets of anthers: the yellow (feeding) anthers function to attract pollinators, but have similar pollen performance to the purple (pollinating) anthers. Furthermore, a disproportional reduction in pollen production of the feeding anthers in the selfing species was found. This differential allocation between feeding and pollinating anthers in heterandrous species has not been recognized before. The finding of reduced allocation to attractive structures with an increase in the rate of self-fertilization supports the theory of sex allocation.     

ECOLOGICAL CONSEQUENCES AND PHENOTYPIC CORRELATES OF PETAL SIZE VARIATION IN WILD RADISH,RAPHANUS SATIVUS (BRASSICACEAE)

In this paper we examine some ecological consequences and phenotypic correlates of flower size variation in wild radish, Raphanus sativus. Mean corolla diameter varied significantly among individuals within natural populations of R. sativus in California. On the average, almost 40% of flower biomass was allocated to corolla tissue. In field experiments, pollinator visitation increased significantly with corolla size. Large flowers also accumulated more nectar when pollinators were excluded from plants. In three populations, corolla size was positively correlated with allocation to pollen per flower (either anther weight or pollen grain number), but there was usually no phenotypic relationship between corolla size and several measures of female allocation (ovule number per flower, proportion fruit set, and total seed mass per fruit). Plants growing in the field produced fewer large flowers per unit of stem, and stem biomass was negatively related to corolla size for plants grown under controlled greenhouse conditions. Male and female fitness may covary differently with allocation to attractive floral features in species such as R. sativus, where seed production is often limited by resources rather than by pollen.     

Gender plasticity in Sagittaria sagittifolia (Alismataceae), a monoecious aquatic species

 Aquatic plants are well known for their high degree of phenotypic plasticity in vegetative structures, particularly leaves. Less well understood is the extent to which their sexuality can be modified by environmental conditions. Here we investigate gender plasticity in the European clonal monoecious aquatic Sagittaria sagittifolia (Alismataceae) to determine how floral sex ratios may vary with plant size and inflorescence order. We sampled two populations from aquatic habitats in East Anglia, U.K. and measured a range of plant attributes including ramet size and the number of female and male flowers per inflorescence. The two populations exhibited similar patterns of phenotypic gender, despite contrasting patterns of total allocation to female and male flower number. Plants produced male-biased floral sex ratios but female flower number increased from the first to the second inflorescence whereas male flower number decreased. Size-dependent gender modification occurred in both populations, but the patterns of allocation to female flower production differed between the two populations. Our results are consistent with the view that monoecy is a sexual strategy that enables plants to adjust female and male allocation in response to changing environmental conditions. Received September 16, 2002; accepted October 23, 2002 Published online: March 20, 2003     

High and Dry: Drought Stress, Sex-Allocation Trade-offs, and Selection on Flower Size in the Alpine Wildflower Polemonium viscosum (Polemoniaceae)

Sex-allocation trade-offs may maintain variation in secondary sexual characteristics if such traits vary in their benefits or costs in association with different genders. In Polemonium viscosum, large flowers benefit both male and female aspects of reproduction. In this study, I explore how resource investment in flower size influences the cost of allocation to male and female function. Large flowers exact a water cost in P. viscosum under dry conditions. In an extreme drought in 1997, experimentally watered plants had higher survival and fecundity than controls. By comparing allocation patterns between plants dying from drought and survivors, I tested whether the demographic cost of large flowers increases with allocation to fecundity. Controls that died showed a positive relationship between flower size and fruit production, while survivors showed a negative relationship or trade-off. Watered plants showed no such trade-off. To test whether drought affects the relationship of corolla size to male function, I used leaf-water potential in 1998 to classify plants as stressed or unstressed. Corolla size showed positive correlations to pollen per flower regardless of drought stress. I conclude that under drought the demographic cost of producing large flowers is gender dependent, such that viability selection favors either small-flowered plants with female-biased reproduction or larger-flowered plants with male-biased reproduction.     

Floral Display in Narcissus: Variation in Flower Size and Number at the Species, Population, and Individual Levels

Floral display (the size, number, and arrangement of open flowers) influences pollinator visitation to animal-pollinated plants and should be an important determinant of reproductive success. We examined variation in the size and number of open flowers in wild daffodils (Narcissus). Our analysis of published data on 45 taxa showed that flower number varied negatively with flower diameter among Narcissus species, which supports the widespread assumption that there is a trade-off between these traits. In contrast, field measurements indicated a positive relation between flower number and diameter within two populations of Narcissus dubius, and no relation was evident after we controlled for variation in bulb size. The discrepancy between inter- and intraspecific patterns may have occurred because variable resource levels obscure trade-offs when variation in flower size is low (e.g., within species). Size-related increases in floral tube length were half as great as corresponding increases in flower diameter, a result that is consistent with stronger stabilizing selection on tube length. Staggered flowering within N. dubius inflorescences limited the mean number of open flowers to <66% of total flower number, and slow expansion by later opening flowers resulted in significant differences in flower size throughout flowering. Although pollinators preferred large flowers, experimental reductions in flower diameter did not affect seed production. Our results illustrate how the relative importance of the factors influencing floral display can vary among levels of biological organization. Interspecific variation in flower size and number appeared to be constrained by allocation trade-offs, but intraspecific variation in both traits was more greatly influenced by plant resource status. Within plants, the size and number of open flowers reflected the relative age of individual flowers and floral longevity.     

SEX ALLOCATION IN THE MONOECIOUS HERB BEGONIA SEMIOVATA

Sex-allocation models predict that the evolution of self-fertilization should result in a reduced allocation to male function and pollinator attraction in plants. The evolution of sex allocation may be constrained by both functional and genetic factors, however. We studied sex allocation and genetic variation for floral sex ratio and other reproductive traits in a Costa Rica population of the monoecious, highly selfing annual Begonia semiovata. Data on biomass of floral structures, flower sex ratios, and fruit set in the source population were used to calculate the average proportion of reproductive allocation invested in male function. Genetic variation and genetic correlations for floral sex ratio and for floral traits related to male and female function were estimated from the greenhouse-grown progeny of field-collected maternal families. The proportion of reproductive biomass invested in male function was low (0.34 at flowering, and 0.07 for total reproductive allocation). Significant among-family variation was detected in the size (mass) of individual male and female flowers, in the proportion of male flowers produced, and in the proportion of total flower mass invested in male flowers. Significant among-family variation was also found in flower number per inflorescence, petal length of male and female flowers, and petal number of female flowers. Except for female petal length, we found no difference in the mean value of these characters between selfed and outcrossed progeny, indicating that, with the possible exception of female petal length, the among-family variation detected was not the result of variation among families in the level of inbreeding. Significant positive phenotypic and broad-sense genetic correlations were detected between the mass of individual male and female flowers, between male and female petal length, and between number of male and number of female flowers per inflorescence. The ratio of stamen-to-pistil mass (0.33) was low compared to published data for autogamous species with hermaphroditic flowers, suggesting that highly efficient selfing mechanisms may evolve in monoecious species. Our results indicate that the study population harbors substantial genetic variation for reproductive characters. The positive genetic correlation between investment in male and female flowers may reflect selection for maximum pollination efficiency, because in this self-pollinating species, each female flower requires a neighboring male flower to provide pollen.     

Sexual dimorphism and gynoecium size variation in the andromonoecious shrub Caesalpinia gilliesii

The degree of sexual dimorphism in flowers and inflorescences can be evaluated early in flower development through the study of floral organ size co-variation. In the present work, the gynoecium-androecium size relationship was studied to assess the degree of sexual expression in flowers and inflorescences of the andromonoecious shrub Caesalpinia gilliesii. The co-variation pattern of floral organ sizes was compared between small and large inflorescences, under the hypothesis that inflorescence size reflected differential resource availability. Also, staminate and perfect flowers were collected from three populations and compared on the basis of gynoecium, ovule length, filament length, pollen size and number. The obtained results indicated that staminate and perfect flowers differed only in the gynoecium and ovule length, whereas filament length, pollen size, and number varied across populations. The gynoecium size was smaller and its variability was much higher in staminate than in perfect flowers, as explained by a recent hypothesis about pollinator-mediated gynoecium size selection acting upon perfect flowers. The analysis of the gynoecium-androecium size relationship during flower development, revealed a dissociation of gynoecium growth relative to other floral structures in some buds. Lower gynoecium-androecium regression slopes and smaller gynoecia length characterized smaller inflorescences, thus reflecting the fact that sexual expression was more male-biased. This trend is in agreement with a differential resource-related response at the inflorescence level, however, post-mating resource allocation and the inclusion of other modular levels may also help us to understand the variation in sexual dimorphism in this species.     

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.     

The Effects of Inflorescence Size and Flower Position on Biomass and Temporal Sex Allocation in <Emphasis Type="Italic">Lobelia sessiliflora</Emphasis>

To test the prediction of sex allocation theory that plants or flowers high in resource status emphasize the female function, we explored the variation in both biomass (the number of pollen grains and ovules) and temporal (male and female durations) sex allocation among and within plants of protandrous Lobelia sessilifolia in relation to plant size and flower position within plants. Among plants, the mean number of pollen grains and ovules per flower of a plant increased with plant size, whereas the mean P/O ratio (number of pollen grains/number of ovules ratio) decreased with plant size. The mean male duration, the mean female duration, and the mean ratio of male duration/flower longevity per flower of a plant were not correlated with plant size. Thus, large plants emphasized female function in terms of biomass sex allocation, which is consistent with the prediction of size-dependent sex allocation theory. The results for temporal sex allocation, however were inconsistent with the theory. Within plants, the mean number of pollen grains and ovules per flower at each position decreased from lower to upper flowers (early to late blooming flowers) and that of the P/O ratio increased from lower to upper flowers. The mean male duration and the mean female duration per flower decreased from lower to upper flowers, whereas the mean ratio of male duration/flower longevity increased from lower to upper flowers. The population sex ratio changed from male-biased to female-biased. Thus, later blooming flowers emphasized the male function in terms of both biomass and temporal sex allocation, consistent with the sex allocation theory, regarding the change in the population sex ratio.     

Production of male flowers does not decrease with plant size in insect-pollinated Sagittaria trifolia, contrary to predictions of size-dependent sex allocation

It has been proposed that relative allocation to female function increases with plant size in animalpollinated species.Previous investigations in several monoecious Sagittaria species seem to run contrary to the prediction of size-dependent sex allocation (SDS),throwing doubt on the generalization of SDS.Plant size,phenotypic gender,and flower production were measured in experimental populations of an aquatic,insect-pollinated herb Sagittaria trifolia (Alismataceae) under highly different densities.The comparison of ramets produced clonally can reduce confounding effects from genetic and environmental factors.In the high-density population,48％ of ramets were male without female flowers,but in the low-density population all ramets were monoecious.We observed allometric growth in reproductive allocation with ramet size,as evident in biomass of reproductive structures and number of flowers.However,within both populations female and male flower production were isometric with ramet size,in contrast to an allometric growth in femaleness as predicted by SDS.Phenotypic gender was not related to ramet size in either population.The results indicated that large plants may increase both female and male function even in animal-pollinated plants,pointing towards further studies to test the hypothesis of size-dependent sex allocation using different allocation currencies.     

Support for the predictions of the pollinator-mediated stabilizing selection hypothesis

Aims Floral traits are frequently used in traditional plant systematics because of their assumed constancy. One potential reason for the apparent constancy of flower size is that effective pollen transfer between flowers depends on the accuracy of the physical fit between the flower and pollinator. Therefore, flowers are likely to be under stronger stabilizing selection for uniform size than vegetative plant parts. Moreover, as predicted by the pollinator-mediated stabilizing selection (PMSS) hypothesis, an accurate fit between flowers and their pollinators is likely to be more important for specialized pollination systems as found in many species with bilaterally symmetric (zygomorphic) flowers than for species with radially symmetric (actinomorphic) flowers.Methods In a comparative study of 15 zygomorphic and 13 actinomorphic species in Switzerland, we tested whether variation in flower size, among and within individuals, is smaller than variation in leaf size and whether variation in flower size is smaller in zygomorphic compared to actinomorphic species.Important findings Indeed, variation in leaf length was significantly larger than variation in flower length and width. Within-individual variation in flower and leaf sizes did not differ significantly between zygomorphic and actinomorphic species. In line with the predictions of the PMSS, among-individual variation in flower length and flower width was significantly smaller for zygomorphic species than for actinomorphic species, while the two groups did not differ in leaf length variation. This suggests that plants with zygomorphic flowers have undergone stronger selection for uniform flowers than plants with actinomorphic flowers. This supports that the relative uniformity of flowers compared to vegetative structures within species, as already observed in traditional plant systematics, is, at least in part, a consequence of the requirement for effective pollination.     

Are there predictable clines in plant–pollinator interactions along altitudinal gradients? The example of Cytisus scoparius (L.) Link in the Sierra de Guadarrama (Central Spain)

Abstract. We analyse the hypothesis that predictable gradients occur in plant–pollinator interactions along altitudinal gradients due to thermal constraints on insect body size and floral traits being selected for by pollinators. Pollinators’ size should follow Bergmann's rule in mountains (larger body size at higher altitudes) and selection should, in turn, lead to larger flower size at higher altitudes. The study focuses on the flower and pollinator size variation in 11 Cytisus scoparius populations located between 680 and 1300 m a.s.l., and on the relationship between flower size and pollination success of plants in each population. Significant differences among populations were found in flower size, pollinator size and fruiting success (anova , P < 0.001 in all cases). Regression models show that (i) pollinator size increases significantly with altitude, (ii) a parallel quadratic increase in flower size and (iii) a quadratic decrease in pollination success with altitude. Moreover, there is a tendency of plants with smaller flowers in each population to have a higher percentage of flowers initiating development into fruits (ancova , P < 0.05). All these observations support the presence of the predicted gradient in the studied species and they suggest the possibility of it to be somewhat common among plants that occupy large altitudinal ranges due to the simple basis on which it could be rooted.     

A test of the effect of floral color change on pollination effectiveness using artificial inflorescences visited by bumblebees

Floral color change has been recognized as a pollination strategy, but its relative effectiveness has been evaluated insufficiently with respect to other floral traits. In this study, effects of floral color change on the visitation pattern of bumblebees were empirically assessed using artificial flowers. Four inflorescence types were postulated as strategies of flowering behavior: type 1 has no retention of old flowers, resulting in a small display size; type 2 retains old flowers without nectar production; type 3 retains old flowers with nectar; and type 4 retains color-changed old flowers without nectar. Effects of these treatments varied depending on both the total display size (single versus multiple inflorescences) and the pattern of flower-opening. In the single inflorescence experiment, a large floral display due to the retention of old flowers (types 2–4) enhanced pollinator attraction, and the number of flower visits per stay decreased with color change (type 4), suggesting a decrease in geitonogamous pollination. Type-4 plants also reduced the foraging time of bees in comparison with type-2 plants. In the multiple inflorescence experiment, the retention of old flowers did not contribute to pollinator attraction. When flowering occurred sequentially within inflorescences, type-4 plants successfully decreased the number of visits and the foraging time in comparison with type-2 plants. In contrast, floral color change did not influence the number of visits, and it extended the foraging time when flowering occurred simultaneously within inflorescences but the opening of inflorescences progressed sequentially within a plant. Therefore, the effectiveness of floral color change is highly susceptible to the display size and flowering pattern within plants, and this may limit the versatility of the color change strategy in nature.     

Why is there variation in mean seed size among plants within single populations? test of the fertilization efficiency hypothesis

We tested the fertilization efficiency hypothesis, which attempts to explain mean seed size variation among plants within single populations, by comparing the patterns of seed size variation between chasmogamous (CH) flowers and cleistogamous (CL) flowers in Impatiens noli-tangere and Viola grypoceras, respectively. The fertilization efficiency hypothesis predicts that larger plants produce larger seeds if the number of pollen grains captured by a plant increases with increased allocation of resources to its attractive structures (e.g., corolla and nectar), but with diminishing gains. Thus, seed size should depend on plant size in seeds from CH flowers because of the diminishing gains of capturing pollen in these flowers, whereas seed size should not depend on plant size in seeds from CL flowers because CL flowers need not capture outcross pollen. We found significant positive correlations between mean seed size per plant and plant size for seeds from CH flowers in both species. However, there was no significant positive correlation between these two factors for seeds from CL flowers of both species. The results of the present investigations were thus consistent with the fertilization efficiency hypothesis.     

Phylogeny determines flower size‐dependent sex allocation at flowering in a hermaphroditic family

• In animal‐pollinated hermaphroditic plants, optimal floral allocation determines relative investment into sexes, which is ultimately dependent on flower size. Larger flowers disproportionally increase maleness whereas smaller and less rewarding flowers favour female function. Although floral traits are considered strongly conserved, phylogenetic relationships in the interspecific patterns of resource allocation to floral sex remain overlooked. We investigated these patterns in Cistaceae, a hermaphroditic family.
• We reconstructed phylogenetic relationships among Cistaceae species and quantified phylogenetic signal for flower size, dry mass and nutrient allocation to floral structures in 23 Mediterranean species using Blomberg's K‐statistic. Lastly, phylogenetically‐controlled correlational and regression analyses were applied to examine flower size‐based allometry in resource allocation to floral structures.
• Sepals received the highest dry mass allocation, followed by petals, whereas sexual structures increased nutrient allocation. Flower size and resource allocation to floral structures, except for carpels, showed a strong phylogenetic signal. Larger‐flowered species allometrically allocated more resources to maleness, by increasing allocation to corollas and stamens.
• Our results suggest a major role of phylogeny in determining interspecific changes in flower size and subsequent floral sex allocation. This implies that flower size balances the male–female function over the evolutionary history of Cistaceae. While allometric resource investment in maleness is inherited across species diversification, allocation to the female function seems a labile trait that varies among closely related species that have diversified into different ecological niches.