Differential siring success of Pgi genotypes in Clarkia Unguiculata (Onagraceae)

Competition among pollen grains for the fertilization of ovules can play an important role in determining the male and female reproductive success of flowering plants. To examine the influence of pollen-donor genotype on male reproductive success, hand-pollinations were conducted on Clarkia unguiculata and the siring success of pollen-donor plants was compared between donors homozygous for different allelomorphs of the allozyme PGI (phosphoglucoisomerase). Donors homozygous for the B allele sired more seeds than C-allele donors. Single-donor crosses indicated that C-donor-sired seeds are aborted more often than are B-donor-sired seeds, suggesting that the B-allele donor's advantage in mixed pollinations was a result of differential abortion. A negative relationship between pollen load and the siring success of B-allele donors implies that pollen from B-allele donors has reduced performance relative to C-allele donors when pollen loads are high. These data demonstrate consistent differences in siring success between individuals homozygous for different alleles at a single locus and suggest that variation at the Pgi locus may be maintained by a post-pollination trade-off.     

Movement herkogamy in Kosteletzkya virginica: Effect on reproductive success and contribution to pollen receipt and reproductive assurance

Herkogamy may act as a mechanism to reduce interference between the reproductive functions of female and male organs, but too great a distance between stigmas and anthers may also decrease the outcross pollen transfer and male and female fitness. Our goal was to quantify the effect of movement herkogamy in Kosteletzkya virginica on reproductive success and its contribution to pollen receipt and reproductive assurance. Field investigations and hand-pollination experiments were performed to test: (1) the relationship between herkogamy before curvature of un-pollinated lobes and reproductive success under pollinator exclusion; (2) the contribution of movement herkogamy to pollen receipt; and (3) the relationship between pollen load and seed set. We noted that variation in herkogamy before curvature of un-pollinated lobes did not affect the outcome of autogamous self-pollination measured by fruit set and mean number of seeds per fruit. Under natural conditions, 39% of pollen grains deposited per flower were from autogamous self-pollination via movement herkogamy resulting in contact between anthers and stigmas. The relationship between the amount of pollen grains and the number of seeds per capsule was highly significant. Flowers producing four or five seeds received more pollen grains than those producing only one seed. Our data provides experimental evidence that movement herkogamy, which decreases the separation between anthers and stigmas and results in delayed self-pollination, contributes to pollen receipt and further enhances seed set when pollinators are scarce or absent.     

Movement herkogamy in Kosteletzkya virginica: Effect on reproductive success and contribution to pollen receipt and reproductive assurance

Herkogamy may act as a mechanism to reduce interference between the reproductive functions of female and male organs, but too great a distance between stigmas and anthers may also decrease the outcross pollen transfer and male and female fitness. Our goal was to quantify the effect of movement herkogamy in Kosteletzkya virginica on reproductive success and its contribution to pollen receipt and reproductive assurance. Field investigations and hand-pollination experiments were performed to test: (1) the relationship between herkogamy before curvature of un-pollinated lobes and reproductive success under pollinator exclusion; (2) the contribution of movement herkogamy to pollen receipt; and (3) the relationship between pollen load and seed set. We noted that variation in herkogamy before curvature of un-pollinated lobes did not affect the outcome of autogamous self-pollination measured by fruit set and mean number of seeds per fruit. Under natural conditions, 39% of pollen grains deposited per flower were from autogamous self-pollination via movement herkogamy resulting in contact between anthers and stigmas. The relationship between the amount of pollen grains and the number of seeds per capsule was highly significant. Flowers producing four or five seeds received more pollen grains than those producing only one seed. Our data provides experimental evidence that movement herkogamy, which decreases the separation between anthers and stigmas and results in delayed self-pollination, contributes to pollen receipt and further enhances seed set when pollinators are scarce or absent.     

Why do stigmas move in a flexistylous plant?

Flexistyly is a recently documented stylar polymorphism involving both spatial and temporal segregation of sex roles within hermaphroditic flowers. Using the experimental manipulation of stigma movement in self-compatible Alpinia mutica, we tested the hypothesis that selection for reducing interference between male and female function drives the evolution and/or maintenance of stigma movement. In experimental arrays, anaflexistylous (protogynous) flowers served as pollen donors competing for mating opportunities on cataflexistylous (protandrous) flowers. The pollen donors were either manipulated so their stigmas could not move or were left intact, and their success was determined using allozymes to assess the paternity of recipient seeds. We found that manipulated flowers sired a significantly smaller proportion of seeds, showing that stigma movement in unmanipulated plants increased male fitness. This result was strongest under conditions in which pollen competition was expected to be highest, specifically when pollinators visited multiple donor plants before visiting recipient flowers.     

文冠果花粉供体的多样性对果实产出的影响

很多植物的花,其柱头上接受的花粉供体多样性数量通常与果实成熟和种子产出有关。关于提高花粉多样性能否增加植物的生殖产出问题,文献中时常有相互矛盾的结论。本研究通过对自交不亲和性植物文冠果(Xanthoceras sorbifolium)进行1、2、3、4种花粉供体传粉实验,调查了不同花粉供体数量对文冠果雌性生殖的影响。结果表明,增加亲和花粉供体的多样性,一定程度上提高了花序上果实的坐果数量,同时也增加了果实中种子的数量及单粒种子重量。对文冠果授以4种亲和性花粉供体后,花序上成熟果实的平均数最多;3种亲和性花粉供体传粉后,每个果实中的种子数最多,单粒种子也最重。如果混合花粉中含有自交花粉,那么果实将败育。除花粉多样性影响雌性生殖产出外,母本基因型也具有重要影响,即增加文冠果种植品系的多样性,将有助于提高果实和种子产量。本研究还讨论了花粉遗传多样性效应、母本交配选择及果实选择性败育假说。     

Sequential variation in the components of reproductive success in the distylousJasminum fruticans (Oleaceae)

A 2-year study of three natural populations of the distylousJasminum fruticans showed that mean fruit and seed production were significantly greater in shortstyled plants (thrums) than in long-styled plants (pins). In this study, we investigated the role of four sequential factors which may differentially influence fruit and seed set in the two floral morphs: (1) differences in flowering phenology, (2) a limitation of pollen transfer towards pins, (3) a differential capacity of the two morphs to act through famale and male function and (4) differential fruit abortion in the two morphs. Fruit set was significantly influenced by differences in flowering phenology although there were no differences in flowering time between the two morphs. supplementary pollinations in a natural population significantly increased fruit set and reduced the difference in fruit set between the two morphs in relation to controls, indicating a limitation on pollen transfer which was most severe towards pin stigmas. In reciprocal crosses, seed set was significantly dependent on the paternal and maternal identity of the pin parent. There was no significant variation among thrums for their performance as male or female parent. Furthermore, individual pin plants with relatively high percentage seed set as female parents gave poor seed set as male parents and vice-versa. Whereas fruit removal had no effect on seed number in thrum plants, a greater proportion of viable seeds were produced on pin plants which were left to naturally mature their fruits than on pins which had fruits artificially removed, suggesting the occurrence of selective fruit abortion in pins but not in thrums. The initially greater maternal fitness of thrums due to their greater success as pollen recipients may thus be opposed by increased viable seed set in the pins due to factors acting after the pollination stage. The relative reproductive success of floral morphs in the distylousJ. fruticans is thus differentially influenced by ecological factors occurring at different stages of the reproductive process.     

Analysis of a chalcone synthase mutant in Ipomoea purpurea reveals a novel function for flavonoids: amelioration of heat stress

Flavonoids are thought to function in the plant stress response and male fertility in some, but not all, species. We examined the effects of a self-fertile chalcone synthase null allele, a, for the effects of heat and light stress on fertilization success and flower production in Ipomoea purpurea. Pollen recipients and pollen donors of both homozygous genotypes exhibit reduced fertilization success at high temperatures, indicating that high temperature acts as a stress-lowering fertilization success. Homozygous aa individuals exhibit reduced male and female fertilization success, compared to AA individuals, at high temperatures but not at low temperatures. In addition, aa individuals produce fewer flowers than AA individuals at low temperatures, but not at high temperatures. These results suggest that flavonoids alleviate heat stress on fertilization success. They also suggest that pleiotropic effects at the A locus may explain the low frequency of the a allele in natural populations.     

Effects of forest fragmentation on male and female reproductive success in<Emphasis Type="Italic"> Cestrum parqui</Emphasis> (Solanaceae)

In this paper we evaluate the effects of forest fragmentation on male (pollen removal, pollen load, and pollen tubes) and female reproductive success (fruit- and seed-set) of Cestrum parqui, a self-incompatible, pollination-specialist plant species. We also measure focal individual conspecific density to account for possible density-related effects that could influence the response variables. We calculate an index which incorporates male and female fitness and gives an integrated assessment of overall reproductive success. Forest fragmentation strongly affected the amount of pollen grains on stigmas and number of pollen tubes as well as seed-set, decreasing from continuous forest to small forest fragments, whereas focal individual conspecific density failed to explain any of the variability for the studied variables. Declines in overall reproductive success (i.e. male and female) in small forest fragments are ascribed to decreases in both the quality and quantity of pollination. Self-incompatibility coupled with a specialist pollination system may be particularly important traits determining the negative fragmentation effects observed in C. parqui. Logarithmic regression models described the behaviour of the variables along the fragmentation size gradient, allowing us to detect a threshold below which the effects of fragmentation begin to negatively affect reproductive success in C. parqui. Our results emphasize the importance of evaluating both components of the total plant fitness, as well as including simultaneously several aspects of pollination and reproduction processes when assessing the effects of forest fragmentation on plant reproductive success.     

Plant Parenthood in Milkweeds: A Direct Test of the Pollen Donation Hypothesis

Abstract For hermaphroditic plant species whose fruit production is limited by maternal resources, the "pollen donation hypothesis" views large floral displays as an adaptation to enhance the probability of fathering seeds on other plants. This hypothesis has frequently been used to describe the evolution of large floral displays in milkweeds ( Asclepias ). Most tests of the pollen donation hypothesis, however, have used indirect measures, such as flower production or pollen removal, to estimate male reproductive success. To test the pollen donation hypothesis directly, we performed a paternity analysis and determined the number of seeds sired by individual genotypes in a natural population of poke milkweed, A. exaltata , in southwestern Virginia. Seeds sired (male success) and seeds produced (female success) were significantly correlated with flower number per plant (for male success: r = 0.32, P < 0.05; for female success: r = 0.66, P < 0.0001). Functional gender of plants that reproduced both as males and females (N = 17) was not correlated with flower number per plant ( r = 0.35, P>0.05), indicating that plants with large floral displays did not reproduce primarily as males. Percent fruit-set and seed number per fruit were higher in 1986, when levels of pollinarium removal also were higher. Furthermore, several umbels that experienced high pollinator activity selectively matured fruits that contained many seeds. We argue that the evolution of large floral displays in milkweeds is the result of selection to increase overall reproductive success rather than male reproductive success alone.     

Pollen and Resource Limitation in Veratrum nigrum L. （Liliaceae）, an Andromonoecious Herb

Pollen limitation and resource limitation were invoked to account for the pattern that flowering plants produce more flowers and ovules than fruits and seeds. This study aimed to determine their relative importance in Veratrum nigrum, a self-compatible, perennial, andromonoecious herb. In order to determine whether female production was limited by pollen grains on stigmas or by available resources, we performed supplemental hand pollination in three populations, male-flower-bud removal in three other populations, and emasculation of hermaphroditic flowers in still another population, resulting in a total of seven populations experimentally manipulated. Across the three populations, supplemental hand pollination did not significantly increase fruit set, seed number per fruit, and total seed production per individual, nor did emasculation of hermaphroditic flowers. Taken together, our results suggest that pollen grains deposited on stigmas were abundant enough to fertilize all the ovules. Male-flower-bud removal significantly increased the mean size of hermaphroditic flowers in all three populations. Female reproductive success was increased in one population, but not in the other two populations possibly due to heavy flower/seed predation. We concluded that the female reproductive success of V. nigrum was not limited by pollen grains but by available resources, which is consistent with Bateman＇s principle. Furthermore, the female reproduction increase of male-flower-bud removal individuals might suggest a trade-off between male and female sexual functions.     

Differential male reproductive success in Douglas fir

Summary Differential male reproductive success was studied in clones at two seed orchards of Douglas fir, Pseudotsuga menziesii (Mirb.) Franco. The performance of tester pollen parents was compared in controlled pollinations with two-parent pollen mixes. Marker pollen homozygous for a rare IDH allele was the genetic marker in each pollen mix. The resulting seeds were analyzed electrophoretically. At both seed orchards, the proportion of seeds sired by tester pollen significantly varied among the tester pollen parents. Tester pollen parents did not perform the same across all seed parents. The significant interaction effect was evidence of male-female complementarity. These results suggest a genetic basis to differential male reproductive success in Douglas fir.     

Effects of floral display size on male and female reproductive success in Mimulus ringens

Background and Aims

The number of flowers blooming simultaneously on a plant may have profound consequences for reproductive success. Large floral displays often attract more pollinator visits, increasing outcross pollen receipt. However, pollinators frequently probe more flowers in sequence on large displays, potentially increasing self-pollination and reducing pollen export per flower. To better understand how floral display size influences male and female fitness, we manipulated display phenotypes and then used paternity analysis to quantify siring success and selfing rates.

Methods

To facilitate unambiguous assignment of paternity, we established four replicate (cloned) arrays of Mimulus ringens, each consisting of genets with unique combinations of homozygous marker genotypes. In each array, we trimmed displays to two, four, eight or 16 flowers. When fruits ripened, we counted the number of seeds per fruit and assigned paternity to 1935 progeny.

Key Results

Siring success per flower declined sharply with increasing display size, while female success per flower did not vary with display. The rate of self-fertilization increased for large floral displays, but siring losses due to geitonogamous pollen discounting were much greater than siring gains through increased self-fertilization. As display size increased, each additional seed sired through geitonogamous self-pollination was associated with a loss of 9·7 seeds sired through outcrossing.

Conclusions

Although total fitness increased with floral display size, the marginal return on each additional flower declined steadily as display size increased. Therefore, a plant could maximize fitness by producing small displays over a long flowering period, rather than large displays over a brief flowering period.     

Effects of foliar herbivory by insects on the fitness of Raphanus raphanistrum: damage can increase male fitness

Generally, effects of herbivory on plant fitness have been measured in terms of female reproductive success (seed production). However, male plant fitness, defined as the number of seeds sired by pollen, contributes half of the genes to the next generation and is therefore crucial to the evolution of natural plant populations. This is the first study to examine effects of insect herbivory on both male and female plant reproductive success. Through controlled field and greenhouse experiments and genetic paternity analysis, we found that foliar damage by insects caused a range of responses by plants. In one environment, damaged plants had greater success as male parents than undamaged plants. Neither effects on pollen competitive ability nor pollinator visitation patterns could explain the greater siring success of these damaged plants. Success of damaged plants as male parents appeared to be due primarily to changes in allocation to flowers versus seeds after damage. Damaged plants produced more flowers early in the season, but not more seeds, than undamaged plants. Based on total seed production, male fitness measures from the first third of the season, and flower production, we estimated that damaged and undamaged plants had equal total reproductive success at the end of the season in this environment. In a second, richer environment, damaged and undamaged plants had equal male and female plant fitness, and no traits differed significantly between the treatments. Equal total reproductive success may not be ecologically or evolutionarily equivalent if it is achieved differentially through male versus female fitness. Genes from damaged plants dispersed through pollen may escape attack from herbivores, if such attack is correlated spatially from year to year.     

PATERNITY ANALYSIS IN A NATURAL POPULATION OF ASCLEPIAS EXALTATA: MULTIPLE PATERNITY,FUNCTIONAL GENDER,AND THE “POLLEN-DONATION HYPOTHESIS”

Recently, some evolutionary biologists have argued that selection on the male component of fitness shapes the evolution of reproductive characters in angiosperms. Floral features, such as inflorescence size, that lead to increased insect visitation without a concomitant increase in seed production are viewed as adaptations to enhance the probability of fathering seeds on other plants. In tests of this “pollen donation hypothesis,” male reproductive success has usually been measured indirectly by flower production, pollinator visitation, or pollen removal. We tested the pollen donation hypothesis directly by quantifying the number of seeds sired by individual genotypes in a natural population of poke milkweed, Asclepias exaltata, in southwestern Virginia. Multiple paternity was low within fruits, a fact which allowed us to use genotypes of progeny arrays to identify a unique pollen parent for 85% of the fruits produced in the population. Seeds sired (male success) and seeds produced (female success) were significantly correlated with flower number per plant (for male success, r = 0.32, P > 0.05; for female success, r = 0.66, P > 0.001). While the number of pollinaria removed, the usual estimator of male success in milkweeds, was highly correlated with numbers of seeds sired (r = 0.47; P > 0.001), it was even more highly correlated with numbers of seeds produced (r = 0.71, P > 0.001). Analysis of functional gender indicated that plants with many flowers did not behave primarily as males. In fact, individuals with the highest total reproductive success contributed equally as males and females. Furthermore, estimates of gender based on numbers of flowers produced or pollinaria removed overestimated the number of functional males in the population. In pollen-limited species, such as many milkweeds, proportional increases in both male and female reproductive success indicate the potential for selection to shape the evolution of large floral displays through both male and female functions.     

Microevolution of S-allele frequencies in wild cherry populations: respective impacts of negative frequency dependent selection and genetic drift

Negative frequency dependent selection (NFDS) is supposed to be the main force controlling allele evolution at the gametophytic self-incompatibility locus (S-locus) in strictly outcrossing species. Genetic drift also influences S-allele evolution. In perennial sessile organisms, evolution of allelic frequencies over two generations is mainly shaped by individual fecundities and spatial processes. Using wild cherry populations between two successive generations, we tested whether S-alleles evolved following NFDS qualitative and quantitative predictions. We showed that allelic variation was negatively correlated with parental allelic frequency as expected under NFDS. However, NFDS predictions in finite population failed to predict more than half S-allele quantitative evolution. We developed a spatially explicit mating model that included the S-locus. We studied the effects of self-incompatibility and local drift within populations due to pollen dispersal in spatially distributed individuals, and variation in male fecundity on male mating success and allelic frequency evolution. Male mating success was negatively related to male allelic frequency as expected under NFDS. Spatial genetic structure combined with self-incompatibility resulted in higher effective pollen dispersal. Limited pollen dispersal in structured distributions of individuals and genotypes and unequal pollen production significantly contributed to S-allele frequency evolution by creating local drift effects strong enough to counteract the NFDS effect on some alleles.     

Seed paternity in flowering plants: an evolutionary perspective

The ultimate importance of paternal contributions to fitness and of post-pollination selection in flowering plants have remained elusive, largely because of the technical difficulty of assigning paternity. I review empirical studies that use heritable markers to investigate per-fruit seed paternity in natural populations and after experimental multiple-donor pollination. Thirty-one studies covering 23 species from 16 plant families document that in natural populations seeds from a single fruit are often fathered by multiple pollen donors (5 species from 5 families), that donors can differ significantly in seed-siring success (8 species from 6 families), that variation in pollen tube growth rates can be heritable (n = 1 out of 4 studies), that donor and recipient genotypes can simultaneously affect paternity (n = 2), and that temporal order of pollen deposition (n = 1) and environmental effects(n = 2) affect the outcome of pollen competition. These studies also investigate the role of post-pollination selection in the avoidance of inbreeding and for species boundaries. Most studies of male reproductive success in plants to date base on isozyme electrophoresis. The availability in the last decade of highly polymorphic molecular markers such as microsatellite DNA has been expected to open new possibilities to investigate competition and selection during the gametophytic phase. Yet, to date, there is still need for greater data wealth on seed paternity to test theories of sex allocation and to gain deeper understanding of floral trait evolution and of the evolutionary consequences of post-pollination selection in flowering plants.     

An Examination of the Function of Male Flowers in an Andromonoecious Shrub Capparis spinosa

The pollen donor and pollinator attractor hypotheses are explanations for the functions of the male flowers of andromonoecious plants. We tested these two hypotheses in the andromonoecious shrub Capparis spinosa L. (Capparaceae) and confirmed that pollen production and cumulative volume and sugar concentration of nectar do not differ between male and perfect flowers. However, male flowers produced larger anthers, larger pollen grains and smaller ovaries than perfect flowers. Observations on pollinators indicated that two major pollinators (Xylocopa valga Gerst and Proxylocopa sinensis Wu) did not discriminate between flower morphs and that they transferred pollen grains a similar distance. However, there were more seeds per fruit following hand pollination with pollen from male flowers than from perfect flowers. Individuals of C. spinosa with a larger floral display (i.e. bearing more flowers) received more pollen grains on the stigma of perfect flowers. Female reproductive success probably is not limited by pollen. These results indicate that male flowers of C. spinosa save resources for female function and that they primarily serve to attract pollinators as pollen donors.     

Selection on floral design in Polemonium brandegeei (Polemoniaceae): female and male fitness under hawkmoth pollination

Plant-pollinator interactions promote the evolution of floral traits that attract pollinators and facilitate efficient pollen transfer. The spatial separation of sex organs, herkogamy, is believed to limit sexual interference in hermaphrodite flowers. Reverse herkogamy (stigma recessed below anthers) and long, narrow corolla tubes are expected to promote efficiency in male function under hawkmoth pollination. We tested this prediction by measuring selection in six experimental arrays of Polemonium brandegeei, a species that displays continuous variation in herkogamy, resulting in a range of recessed to exserted stigmas. Under glasshouse conditions, we measured pollen removal and deposition, and estimated selection gradients (β) through female fitness (seeds set) and male fitness (siring success based on six polymorphic microsatellite loci). Siring success was higher in plants with more nectar sugar and narrow corolla tubes. However, selection through female function for reverse herkogamy was considerably stronger than was selection through male function. Hawkmoths were initially attracted to larger flowers, but overall preferred plants with reverse herkogamy. Greater pollen deposition and seed set also occurred in reverse herkogamous plants. Thus, reverse herkogamy may be maintained by hawkmoths through female rather than male function. Further, our results suggest that pollinator attraction may play a considerable role in enhancing female function.     

Quantitative and qualitative consequences of reduced pollen loads in a mixed‐mating plant

Greater pollination intensity can enhance maternal plant fitness by increasing seed set and seed quality as a result of more intense pollen competition or enhanced genetic sampling. We tested experimentally these effects by varying the pollen load from a single pollen donor on stigmas of female flowers of Dalechampia scandens (Euphorbiaceae) and measuring the effects on seed number and seed mass. Seed set increased rapidly with pollen number at low to moderate pollen loads, and a maximum set of three seeds occurred with a mean pollen load of 19 pollen grains. We did not detect a trade‐off between the number of seeds and seed mass within a fruit. Seed mass increased with increasing pollen load, supporting the hypothesis of enhanced seed quality via increased pollen‐competition intensity or genetic sampling. These results suggest that maternal fitness increases with larger pollen loads, even when the fertilization success is already high. Our results further highlight the importance of high rates of pollen arrival onto stigmas, as mediated by reliable pollinators. Comparing the pollen‐to‐seed response curve obtained in this experiment with those observed in natural populations suggests that pollen limitation may be more severe in natural populations than predicted from greenhouse studies. These results also indicate that declines in pollinator abundance may decrease plant fitness through lowered seed quality before an effect on seed set is detected.     

Absence of conspecific pollen advantage in the dynamics of an Ipomopsis (Polemoniaceae) hybrid zone

The frequency of hybrid formation in angiosperms depends on how often heterospecific pollen is transferred to the stigma and on the success of that heterospecific pollen at fertilizing ovules. Even if heterospecific pollen is capable of effecting fertilization it may perform poorly when conspecific pollen is also available on the stigma. We applied pollen mixtures to stigmas to determine how pollen interactions affect siring success and the frequency of hybrid formation between two species of Ipomopsis (Polemoniaceae) in Colorado. Plants of both parental species and natural hybrids were pollinated with I. aggregata and I. tenuituba pollen in ratios of 100:0, 80:20, 50:50, 20:80, and 0:100 by mass. Plants were homozygous for different alleles at an isozyme marker, allowing us to distinguish the type of pollen parent for 2166 viable seeds from 273 fruits. In contrast to studies of many other hybridizing taxa, there was no evidence of an advantage to conspecific pollen, nor did composition of the stigmatic pollen load affect seed set. Instead, the frequency of seeds sired by a given species was proportional to its representation in the pollen load. In this hybrid zone, both the frequency of first-generation hybrid formation and the relative male fitness of the two parental species should be predictable from the rates of pollen transfer to stigmas.