Simulation analyses of the evolution of intra-inflorescence flowering patterns assuming selection on anthesis interval among individual flowers.

What conditions select flowering patterns within inflorescences, or variation in the anthesis interval within inflorescences among plants? Under what conditions are gradual blooming and simultaneous blooming, both traits related to floral display size, advantageous? We constructed a simulation model in which the opening times and longevities of individual flowers within inflorescences, the sizes of attractive structures of individual flowers, and the numbers of ovules and pollen grains produced by individual flowers evolve. Individual plants in the population compete for pollinators, and plants are selected by pollinators according to their floral display sizes and amounts of resources allocated to attractive structures. We found that, if the proportion of pollen on a pollinator deposited on a stigma was low, gradual blooming did not evolve even if inbreeding depression was greater than 0.5. This is because the amount of outcross-pollen on pollinators decreased at a low rate during flower visits within a single inflorescence, and the selfing rate was suppressed to a low level even if the floral display size was large. On the other hand, if the proportion of pollen deposition was high, gradual blooming evolved even if inbreeding depression was smaller than 0.5. This may be because gradual blooming can enhance pollen delivery to other plants by reducing the loss of self-pollen by geitonogamy. On the other hand, allocation ratios among floral organs (female and male organs and attractive structures) were independent of the degree of simultaneous and gradual blooming within inflorescences. We concluded that the evolution of gradual blooming is more strongly affected by the proportion of pollen on a pollinator deposited on a stigma than by inbreeding depression.

Effect of temporal changes in stamen position on reproductive success in flowers with many stamens: Manipulations of stamen position.

PREMISE: Male and female reproductive success is enhanced (increased outcrossing and seed production, respectively) by stamen movement in species that have few stamens per flower. Does such enhancement also occur in species that have many stamens per flower? METHODS: We examined the effects of stamen movement on male and female reproductive success in Anemone flaccida, which has many stamens per flower. We measured stamen movement, including temporal changes in anther-stigma and anther-anther distances. We experimentally fixed stamens in their pre- or post-movement positions. RESULTS: The anthers moved horizontally away from the stigmas with increasing flower age, thus reducing female-male interference. The dehisced anthers tended to move farther from the stigmas, while the undehisced or dehiscing anthers remained closer to them. The number of anthers touched per flower visit was higher in flowers whose stamens were fixed in the pre-movement position than in flowers whose stamens were fixed in the post-movement position or in flowers that were not manipulated. Thus, this position may promote male reproductive success. Seed production was lower for the untreated flowers than for those with stamens fixed in the post-movement position, suggesting that the post-movement stamen position is advantageous and stamen movement is suboptimal for female reproductive success. CONCLUSIONS: Stamen movement promotes male reproductive success in the early flowering stage and female reproductive success in the late flowering stage. In species having many stamens per flower, female-male interference can be reduced, but not eliminated, by stamen movement due to the conflict between female and male reproductive successes.

Pollen morphology for successful pollination dependent on pollinator taxa in a generalist plant: relationship with foraging behavior.

Pollen morphology varies at inter- and intraspecific levels. Its interaction with pollinator behavior and morphology determines the probability of successful pollination. We tested whether pollen morphology promoting successful pollination differs depending on pollinator taxa in a generalist shrub, Weigela hortensis (Caprifoliaceae). We identified flower visitors carrying pollen from anthers to stigmas and compared the spine length and diameter of the pollen grains they carried. We found that pollen on the bodies of bumble bees and hunch-back flies and the scopae of small bees (including andrenid bees) contributed to seed production. Pollen grains on the bodies of bumble bees had longer spines than those on the scopae of andrenid bees or the bodies of hunch-back flies. Pollen grains on the bodies of bumble bees and the scopae of andrenid bees had larger diameters than those on hunch-back flies. Bumble bees collected pollen grains with shorter spines and larger diameters on their corbiculae while andrenid bees collected pollen grains with shorter spines and intermediate diameters on their scopae. The differences in morphology of pollen carried by pollinators reflected the tendency of bees to collect pollen with specific morphology into corbiculae/scopae. Our findings suggest that pollen morphology has diversified to facilitate successful pollination by pollinating partners.

Evolution of mixed mating: Effects of among-parent variation in early-acting inbreeding depression coupled with overproduction of ovules.

Theoretical models were developed to propose a new mechanism enhancing mixed mating (reproduction by both outcrossing and selfing) in hermaphroditic plants; mixed mating can be maintained if there exists among-parent variation in early-acting inbreeding depression in embryos and parents can replace dead embryos by overproduction of ovules. In the two main models developed, the number of embryos produced is allowed to evolve, parents may overproduce embryos, and among-parent variation in early-acting inbreeding depression does not exist or exists. I found that mixed mating does not evolve if among-parent variation in early-acting inbreeding depression does not exist, whereas it evolves if it exists. If the degree of early-acting inbreeding depression in embryos is variable among parents, parents with the same selfing strategy suffer different effects of early-acting inbreeding depression. Specifically, overproduction of embryos may be insufficient when inbreeding depression is severe but wasteful when it is weak. Hence, it is advantageous to produce a moderate number of embryos to reduce waste of resources. Mixed mating is then advantageous to avoid great reductions in seed number caused by massive loss of selfed embryos in cases of severe inbreeding depression.

Why are deleterious mutations maintained in selfing populations? An analysis of the effects of early- and late-acting mutations by a two-locus two-allele model.

Frequencies of deleterious mutations are higher than expected in many plants. Here, by developing a two-locus two-allele model, I examine the effects of differential timing of the expression of deleterious mutations (two-stage effects) on the maintenance of mutations. I assume early- and late-acting loci to distinguish whether maintenance of mutations in populations with high selfing rates is explained better by two-stage effects of single mutations, or by separate mutations in both early- and late-acting loci. I found that, when ovules are overproduced, the stable frequency of early-acting mutations is higher if mutations also occur in a late-acting locus than if a late-acting mutation is lacking. The stable frequency of late-acting mutations is higher if mutations also occur in an early-acting locus than if an early-acting mutation is lacking. Selective interference does not account for these results because analyses in which the number of loci subject to mutations is equalized are included. Overproduction of ovules has little effect on maintenance if either early- or late-acting mutations are lacking, whereas when ovules are not overproduced, the two-stage effect does not enhance the maintenance of mutations. Hence, mutations occurring in both loci coupled with overproduction of ovules enhances the maintenance of mutations in populations with high selfing rates. The detailed mechanisms underlying the two-stage effect were also analyzed.

Optimal pollen stickiness to pollinators for maximizing paternal fitness: Increased number of recipient flowers or increased pollen deposition on recipient flowers?

A plant can sire more seeds by increasing the number of pollen recipient flowers or the amount of pollen deposited on recipient flowers. We theoretically analyzed how pollen stickiness contributes to paternal fitness through changing the pattern of pollen dispersal including both the number of recipient flowers and overall pollen deposition (the overall amount of pollen deposited on recipient flowers) in animal-pollinated plants. We developed a numerical model in which pollen stickiness to pollinators increases with production of expensive materials on pollen surfaces, and a high level of stickiness diminishes the proportions of pollen lost from a pollinator body during a flight and pollen deposited on a stigma during a visit. We found that the number of recipient flowers monotonically increased with increasing pollen stickiness allocation while overall pollen deposition was maximized at a certain amount of stickiness allocation. We demonstrated that evolutionarily stable pollen stickiness attained many recipient flowers at the expense of overall pollen deposition in most cases while it merely favored maximization of overall pollen deposition in all other cases. Sticky pollen evolved if pollinators were highly likely to drop pollen during flights and did not diffuse well. In this situation, the evolutionarily stable pattern of pollen dispersal was acquisition of many pollen recipient flowers rather than maximization of overall pollen deposition. Sticky pollen also evolved if additional sticking elements were moderately effective in increasing the force of adhesion to pollinators. Pollen stickiness has a significant effect on the pattern of pollen dispersal via the extent of pollen carryover, and our results suggest that plants maximize paternal fitness by giving pollen the optimal stickiness, which varies with pollinating partners.

Low mutation rates promote the evolution of advantageous traits by preventing interference from deleterious mutations.

We studied how mutation rates promote the evolution of advantageous traits in an asexual population. First, to examine the effects of mutation rates on the evolution of an advantageous trait (high competitive ability), we carried out simulation analyses with competition between individuals for survival. Second, to examine the mechanism underlying the promotion of advantageous trait evolution, we calculated the probabilities that new favorable effects of mutations on the phenotype were acquired and that existing favorable effects were maintained. In the simulation analyses, advantageous traits evolved in the population with a low mutation rate; however, when the mutation rate was extremely low, advantageous traits evolved slowly because few beneficial mutations occurred. Then, the numerical calculations showed that the probability of acquiring new favorable effects of mutations on the phenotype and the probability of maintaining existing favorable effects are high if the mutation rate is low. The former occurs because, if the mutation rate is high, multiple mutations may occur in a genome, and even if beneficial mutations occur, their favorable effects may be masked by simultaneously occurring deleterious mutations. However, if the mutation rate is low, it is likely that only one beneficial mutation occurs, and its favorable effect on the phenotype is direct. In conclusion, low mutation rates are advantageous because they promote favorable phenotypic effects of mutations without interference from deleterious mutations; these low rates not only prevent the occurrence of deleterious mutations but also help maintain existing beneficial mutations and promote the evolution of advantageous traits.

Maintenance of high inbreeding depression in selfing populations: Two-stage effect of early- and late-acting mutations.

High estimates of inbreeding depression have been obtained in many plant populations with high selfing rates. However, deleterious mutations might be purged from such populations as a result of selfing. I developed a simulation model assuming the presence of mutations at two sets of loci, namely, early- and late-acting loci, and the selective abortion of embryos coupled with ovule overproduction. In the model, early-acting loci are expressed during embryo initiation, and less vigorous embryos are aborted. Late-acting loci are expressed after selective abortion ends; the surviving embryos (seeds) compete, and some of them form the next generation. If mutations are allowed to occur in both early- and late-acting loci, both types increase in frequency in populations with high selfing rates. However, this phenomenon does not occur if mutations occur only in the early- or only in the late-acting loci. Consistent results are observed even if the total number of loci in which mutations are allowed to occur is the same for simulations with both early- and late-acting loci, only early-acting loci, or only late-acting loci, indicating that the presence of both types of loci is the causal factor. Thus, the two-stage effect, or occurrence of both early- and late-acting mutations, promotes the maintenance of these mutations in populations with high selfing rates.

Inherited dimorphism in cleistogamous flower production in Portulaca oleracea: a comparison of 16 populations growing under different environmental conditions.

BACKGROUND AND AIMS: Cleistogamy is considered to be an adaptive strategy resulting in plasticity in chasmogamous (CH) and cleistogamous (CL) flower production depending on environmental conditions and plant size. The aim of this study was to investigate whether CH and CL flower production in Portulaca oleracea is genetically differentiated among populations in association with climatic conditions. METHODS: First, we conducted growth experiments with P. oleracea seedlings from 16 populations under two temperature conditions. Secondly, we sowed seeds originating from the parents in the first experiment and grew the resulting plants to investigate whether flower production is heritable and whether plants in the same population show the same pattern of flower production. KEY RESULTS: Two types of plants that produced only CH or CL flowers (referred to as CH and CL plants, respectively) were mainly observed, and the growing temperature conditions did not affect flower production. The frequency of CL plants increased with a decrease in the mean temperature in the original population. The CL plants tended to begin reproduction earlier than the CH plants, and the probability that a CH plant would flower decreased under the low growing temperature condition. Thus, CL plants may have some advantages in unfavourable environments in which early reproduction is necessary due to a short growing season and/or when CH flowers cannot open due to low temperatures. The progeny originating from CH and CL plants also produced only CH and CL flowers, respectively, suggesting that there is a genetic basis for the dimorphism in flower production in P. oleracea, represented by CH and CL plants. CONCLUSIONS: In contrast to the previous hypothesis that the production of both CH and CL flowers would be plastic, the genotypes producing either CH or CL flowers occurred at different frequencies under varying climatic conditions.

Evolution towards minimum ovule size? Ovule size variations and the relative sizes of ovules to seeds.

BACKGROUND AND AIMS: Is there selection minimizing the costs of ovule production? Such selection should lead to a smaller ovule size in relation to seed size and, at the same time, smaller variation in ovule size within plants, the latter because the minimum structures and resources for functioning of ovules should be the same among ovules. Additionally, within species, ovule size should not depend on the plant's resource status. METHODS: To confirm these predictions, we examined ovule and seed production for a variety of species. KEY RESULTS: Among the 27 species studied, we found a significant negative dependence of the species mean of the coefficient of variation for plant ovule size on the ratio of the mean species seed size/mean species ovule size. Thus, the smaller the ovule size as compared with seed size, the smaller the degree of variation in ovule size. Among the 49 species studied, only two species showed significant positive dependence of mean ovule size on plant size. Although larger plants should have greater resources for ovule production, selection has not enhanced the production of large ovules in most species. CONCLUSIONS: These results suggest that there is selection minimizing the costs of ovule production.

Intraflower variation in nectar secretion: Secretion patterns and pollinator behavior in male- and female-phase flowers.

PREMISE OF THE STUDY: Plants that secrete abundant nectar attract many pollinator visits, potentially leading to a high frequency of geitonogamous pollination. However, intraflower variation in nectar volume might enhance early departure from the flower or inflorescence, reducing geitonogamy. To examine the adaptive significance of multiple nectar-producing spurs within flowers, we related intraflower variation in nectar characteristics to pollinator foraging behavior and plant reproductive success in protandrous Aquilegia buergeriana var. oxysepala. METHODS: We investigated if nectar volume and sugar mass relate to spur length and if intraflower variation in nectar characteristics differ between male- and female-phase flowers. Because male function requires more pollinator visits than female function, male-phase flowers are expected to have greater nectar volume and lower intraflower variation in nectar volume than female-phase flowers. We used intraflower variation in spur length as an indicator of variation in nectar characteristics and investigated how spur length variation affects pollinator behavior, pollen removal, and seed production. KEY RESULTS: Male-phase flowers had greater nectar volume and lower intraflower variation in nectar volume than female-phase flowers. Intraflower variation in spur length reduced the duration of bumblebee visits. Short visits reduced seed production, but increased pollen removal. CONCLUSIONS: Our results suggest that high intraflower variation in nectar production rate can reduce geitonogamy and potentially increase the genetic diversity of pollen received. Such pattern of nectar secretion within and among flowers may allow flowers to ensure effective pollen transfer or high seed production.

How Have Self-Incompatibility Haplotypes Diversified? Generation of New Haplotypes during the Evolution of Self-Incompatibility from Self-Compatibility.

I developed a gametophytic self-incompatibility (SI) model to study the conditions leading to diversification in SI haplotypes. In the model, the SI system is assumed to be incomplete, and the pollen expressing a given specificity is not fully rejected by the pistils expressing the same specificity. I also assumed that mutations can occur that enhance the rejection of pollen by pistils with the same haplotype variant and reduce rejection by pistils with other variants in the same haplotype. I found that if such mutations occur, the new haplotypes (mutant variants) can stably coexist with the ancestral haplotype in which the mutant arose. This is because pollen bearing the new haplotype is most strongly rejected by pistils bearing the same new haplotype among the pistils in the population; hence, negative frequency-dependent selection prevents their fixation. I also performed simulations and found that the nearly complete SI system evolves from completely self-compatible populations and that SI haplotypes can increase to about 40-50 within a few thousand generations. On the basis of my findings, I propose that diversification of SI haplotypes occurred during the evolution of SI from self-compatibility.

Relation between flower head traits and florivory in Asteraceae: a phylogenetically controlled approach.

UNLABELLED: • PREMISE OF THE STUDY: While much research has examined the relation between leaf traits and herbivory, very little is known about the interaction between floral traits, particularly biochemical traits, and florivory. We investigated patterns between floral traits and florivory across multiple species using phylogenetic comparative approaches to enhance our understanding of the evolution of plant-florivore interactions.• METHODS: The relation between the intensity of florivory and five biochemical traits (concentrations of carbon, nitrogen, phosphorus, water, and total phenolics) and two morphological traits (diameter and number of flower heads) were investigated in wild individuals of 18 native species of Asteraceae. The phylogenetic signals in the morphological traits and intensity of florivory were also tested.• KEY RESULTS: We found that species with higher nitrogen, water, and total phenolics and lower phosphorus concentrations in the flower heads and species with a large number and diameter of flower heads tended to be attacked by florivores. In addition, we found significant phylogenetic signals in florivory and morphological traits.• CONCLUSIONS: Our results clearly show that biochemical traits also play important roles in plant-florivore interactions, as previously shown in plant-leaf herbivore interactions. The positive relationship between florivory and total phenolics implies that phenolic compounds in flower heads may not act as a defense in the species. In addition, the observed pattern of signals in florivory might not be solely explained by the signals of the measured traits and other plant traits may also play significant roles in plant-florivore interaction in these species.

Initial invasion of gametophytic self-incompatibility alleles in the absence of tight linkage between pollen and pistil S alleles.

In homomorphic self-incompatibility (SI) systems of plants, the loci controlling the pollen and pistil types are tightly linked, and this prevents the generation of compatible combinations of alleles expressing pollen and pistil types, which would result in self-fertilization. We modeled the initial invasion of the first pollen and pistil alleles in gametophytic SI to determine whether these alleles can stably coexist in a population without tight linkage. We assume pollen and pistil loci each carry an incompatibility allele S and an allele without an incompatibility function N. We assume that pollen with an S allele are incompatible with pistils carrying S alleles, whereas other crosses are compatible. Ovules in pistils carrying an S allele suffer viability costs because recognition consumes resources. We found that the cost of carrying a pistil S allele allows pollen and pistil S alleles to coexist in a stable equilibrium if linkage is partial. This occurs because parents that carry pistil S alleles but are homozygous for pollen N alleles cannot avoid self-fertilization; however, they suffer viability costs. Hence, pollen N alleles are selected again. When pollen and pistil S alleles can coexist in a polymorphic equilibrium, selection will favor tighter linkage.

Difference in defense strategy in flower heads and leaves of Asteraceae: multiple-species approach.

Although a vast number of studies have investigated defenses against herbivores in leaves, relatively little is known about defenses in flowers. Using wild individuals of 34 species of Asteraceae, we investigated differences in five traits that are thought to affect the intensity of herbivory (C, N, P, water, and total phenolic contents). Combinations of these traits between flower heads and leaves were studied as well. We also evaluated phylogenetic patterns of flower head and leaf traits. Flower heads had higher P and lower total phenolics than leaves. Water and C contents were negatively correlated both in the flower heads and leaves. N, P, and water contents were positively correlated in the flower heads, whereas this pattern was not found in the leaves. Thus, the traits we measured were more tightly inter-correlated in flower heads than in leaves. Because the flower heads had a lower total phenolic content, the relative allocation of defensive compounds could not be explained solely by fitness values of the organs. Perhaps plants employ an escape strategy rather than a defense strategy to cope with floral herbivores and higher allocation in P may enhance their escape from herbivores by improving the growth rate of flower heads, though our result might be affected in part by the plasticity of plants growing at different sites. Moreover, we found weak phylogenetic signals in the defensive traits. Because we found significant differences in the flower head traits, these weak signals may imply that the traits we measured evolved frequently.

Effects of floral display size, local open raceme density, patch size, and distance between patches on pollinator behaviour in Salvia nipponica.

Flowers cluster at various spatial scales, so pollinators use information from multiple scales when foraging in natural plant populations. Little is known about the effects of interactions between scales or their relative strength. We examined bumblebee foraging behaviour in a natural population of Salvia nipponica in 10 and 7 patches in 2019 and 2020, respectively. We recorded within-patch factors (display size of racemes and local open raceme densities) and patch-level factors (patch size and distance from the nearest patch) and analysed their relationships with pollinator behaviour. The numbers of visits per raceme and flower were mainly affected by the interaction of patch size and raceme density; they were higher in locations with lower raceme density in larger patches. The ratio of flowers visited to all open flowers in a raceme during a raceme visit, which relates to a bumblebee's choice to leave a raceme, was mainly affected by the interaction of display size and local open raceme density; in 2019 it was higher in racemes with smaller display sizes, while in 2020 the strength and direction of the relationship depended on the open raceme density. These results suggest that pollinators relied on the sizes of flower clusters at different spatial scales when visiting and leaving racemes and adjusted their responses to the sizes of flower clusters depending on the distances between clusters. Therefore, it is important to evaluate factors at various spatial scales and their interactions to fully understand pollinator behaviour in natural plant populations.

With whom is the gene in conflict in offspring production? Synthesis of the theories of intragenomic and parent-offspring conflict.

In offspring production, with whom are the maternally derived (madumnal), paternally derived (padumnal), and maternal genes in conflict? I developed a model, in which those genes independently regulate resource absorption of developing offspring, and offspring with a high realized resource absorption rate may become large, but may suffer abortion due to overgrowth. I analyzed two cases: maternal control is weak (maternal genes cannot completely inhibit the resource demand by the madumnal or the padumnal genes) and is strong (maternal genes can completely inhibit it). I found that, under weak maternal control, the maternal genes inhibit resource absorption, but the madumnal and padumnal genes enhance it if the abortion cost of overgrowth is low. The maternal and madumnal genes inhibit resource absorption, but the padumnal genes enhance it if the cost is high. Under strong maternal control, the maternal genes inhibit resource absorption, but the madumnal and padumnal genes enhance it irrespective of the degree of abortion cost. I also found that the effects of offspring abortion on an ESS size and number of offspring when independent are large under weak maternal control, but are moderated under strong maternal control.

Optimal defense strategy against herbivory in plants: conditions selecting for induced defense, constitutive defense, and no-defense.

To examine the conditions selecting for induced defense, constitutive defense, and no-defense, we developed a model of plant defense strategy against herbivory. In the model, a plant consists of two modules between which signal inducing defense compounds can be translocated. We assume three strategies: plants produce defense compounds responding to herbivory (induced defense), they have the compounds beforehand (constitutive defense), and they never produce the compounds (no-defense). We found that no-defense is optimal if the amount of biomass lost due to herbivory is small because of the growth cost of having defense compounds. The constitutive defense is optimal if the amount of biomass lost is not so small and the probability of herbivory is high. If the biomass loss is not so small but the probability of herbivory is low, the induced defense or no-defense is optimal. When the induced defense is optimal, the probability of herbivory necessarily increases in plants once herbivory has occurred. If the probability stays the same, no-defense is optimal. Thus, the behavior of herbivores, i.e., whether they remain around a plant and attack it repeatedly, affects the evolution of the induced defense.

Overproduction and selective abortion of ovules based on the order of fertilization revisited.

Given that seeds fertilized by slowly growing pollen are of low quality genetically, we theoretically reanalyzed the hypothesis that plants selectively abort ovules fertilized later to enhance the mean quality of resulting seeds. We assumed that both superior and inferior pollen exist, the superior pollen growing faster to fertilize ovules, resulting in seeds of higher quality than those of ovules fertilized by inferior pollen. We developed two models to determine the conditions under which selective abortion is favored. In the first model, ovules in one flower are fertilized by pollen grains that arrive at different times, with each visit bringing both fast- and slow-growing pollen. In the second model, ovules in two flowers are fertilized by all pollen grains that arrive at the same time. In the first model, we found that selective abortion based on the order of fertilization is never advantageous irrespective of the duration of the time lag between the two visits. Rather, random abortion is possibly favored. In the second model, although selective abortion based on the order of fertilization can be advantageous, the parameter region favoring it is rather restricted. This is because overproduction can be advantageous only if the quantity of the superior pollen is not limited in one flower but is limited in the other flower. In addition, the degree of overproduction was very low, implying that the merit of overproduction (increase in the number of superior seeds) is low compared to the cost of overproducing ovules. These results suggest that selective abortion of ovules based on the order of fertilization is not as advantageous as previously considered.

Ovule and seed production patterns in relation to flower size variations in actinomorphic and zygomorphic flower species.

Zygomorphic flower species tend to show lower flower size variation than actinomorphic flower species. Have these differences also brought an association in ovule and seed production that has arisen due to natural selection in these species? Flowers were collected from 29 actinomorphic and 20 zygomorphic flower species, and fruits were collected from 21 actinomorphic and 14 zygomorphic flower species in Miyagi and Aomori prefectures, in Japan. The coefficient of variations (CVs) of flower sizes, mean ovule sizes of flowers, ovule numbers of flowers and mean seed sizes of fruits were calculated. The CV of flower sizes was marginally different between the floral symmetry types; tending to be lower in the zygomorphic flower species than in the actinomorphic flower species. The CVs of mean ovule sizes and ovule numbers of flowers increased with increase in the CV of flower sizes in the actinomorphic flower species but not in the zygomorphic flower species. Mean ovule number of flowers tends to increase with increase in mean flower size in the actinomorphic flower species but not in the zygomorphic flower species. The degrees in variations in ovule size and number of flowers were influenced by the interaction of floral symmetry type and flower size variation, suggesting that floral symmetry also has brought an evolutionary association in ovule production by flowers.