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1.
Among all ovules produced by a plant population in an area of arbitrary size, the expectation and the variance is derived for the fraction of pollinated ovules. Particular emphasis is put on the use of parameters which are accessible to direct measurement. The results are applied to the problem of finding the sex ratio in a dioecious plant population which guarantees maximum population fitness, i.e. the maximum number of pollinated ovules per plant. The optimal sex ratio always shows a female excess. The excess increases as either the average number of pollen grains per male plant or the population density increases.  相似文献   

2.
Theory predicts that plant defensive traits are costly due to trade-offs between allocation to defense and growth and reproduction. Most previous studies of costs of plant defense focused on female fitness costs of constitutively expressed defenses. Consideration of alternative plant strategies, such as induced defenses and tolerance to herbivory, and multiple types of costs, including allocation to male reproductive function, may increase our ability to detect costs of plant defense against herbivores. In this study we measured male and female reproductive costs associated with induced responses and tolerance to herbivory in annual wild radish plants (Raphanus raphanistrum). We induced resistance in the plants by subjecting them to herbivory by Pieris rapae caterpillars. We also induced resistance in plants without leaf tissue removal using a natural chemical elicitor, jasmonic acid; in addition, we removed leaf tissue without inducing plant responses using manual clipping. Induced responses included increased concentrations of indole glucosinolates, which are putative defense compounds. Induced responses, in the absence of leaf tissue removal, reduced plant fitness when five fitness components were considered together; costs of induction were individually detected for time to first flower and number of pollen grains produced per flower. In this system, induced responses appear to impose a cost, although this cost may not have been detected had we only quantified the traditionally measured fitness components, growth and seed production. In the absence of induced responses, 50% leaf tissue removal, reduced plant fitness in three out of the five fitness components measured. Induced responses to herbivory and leaf tissue removal had additive effects on plant fitness. Although plant sibships varied greatly (49–136%) in their level of tolerance to herbivory, costs of tolerance were not detected, as we did not find a negative association between the ability to compensate for damage and plant fitness in the absence of damage. We suggest that consideration of alternative plant defense strategies and multiple costs will result in a broader understanding of the evolutionary ecology of plant defense.  相似文献   

3.
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.  相似文献   

4.
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.  相似文献   

5.
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.  相似文献   

6.
Based on the hypothesis that both plant size and local conspecific density influence allocation to female/male functions, we explored the relationship between plant height, local conspecific density, sexual expression, and fruit production in the andromonoecious shrub Caesalpinia gilliesii. We quantified the total number of perfect and staminate flowers, the pollen received and fruits produced per plant in two populations, and estimated phenotypic gender and fruit set. Local density failed to explain phenotypic gender, nevertheless, plant height and fruit set increased with local density in one population where, in addition, the slopes for the size-dependent sex allocation curve were steeper. As observed for other plant species, this suggests that between population differences in resource availability is the main underlying factor for the observed population differences in the size-dependent allocation pattern to flowers and fruits. On the other hand, the number of staminate and perfect flowers per plant increased with plant height and the fastest increase of staminate flowers resulted in a male-biased size-dependent sex allocation strategy in both populations. Since pollination intensity was not correlated with plant height in any population, the observed allocation strategy cannot be attributed to differences in pollen availability between different sized individuals, but to differences in plant size. Finally, because fruit set and total fruit number increased with plant height in one population, the obtained results provide further evidence that animal-pollinated, andromonoecious species may exhibit a male-biased size-dependent sex allocation strategy, which may favor female fecundity.  相似文献   

7.
Aims We explore the possible role of leaf size/number trade-offs for the interpretation of leaf size dimorphism in dioecious plant species.Methods Total above-ground biomass (both male and female) for three herbaceous dioecious species and individual shoots (from both male and female plants) for three woody dioecious species were sampled to record individual leaf dry mass, number of leaves, dry mass of residual above-ground tissue (all remaining non-leaf biomass), number of flowers/inflorescences (for herbaceous species) and number of branches.Important findings For two out of three woody species and two out of three herbaceous species examined, male plants produced smaller leaves but with higher leafing intensity—i.e. more leaves per unit of supporting (residual) shoot tissue or plant body mass—compared with females. Male and female plants, however, did not differ in shoot or plant body mass or branching intensity. We interpret these results as possible evidence for a dimorphic leaf deployment strategy that promotes both male and female function, respectively. In male plants, capacity as a pollen donor may be favored by selection for a broadly spaced floral display, hence favoring relatively high leafing intensity because this provides more numerous axillary meristems that can be deployed for flowering, thus requiring a relatively small leaf as a trade-off. In one herbaceous species, higher leafing intensity in males was associated with greater flower production than in females. In contrast, in female plants, selection favors a relatively large leaf, we propose, because this promotes greater capacity for localized photosynthate production, thus supporting the locally high energetic cost of axillary fruit and seed development, which in turn requires a relatively low leafing intensity as a trade-off.  相似文献   

8.
I investigated the effects of display size and flowering phenology on fruit set in Aucuba japonica, an understory dioecious shrub pollinated by opportunistic insects. Natural variations in display size, flowering phenology, and fruit set were monitored in 1997. A hand-pollination experiment was also conducted to check whether pollen limitation was a factor in fruit set in the field. Increases in floral display size did not affect fruit set; the proportion of flowers that set fruit was almost constant irrespective of the total number of flowers per inflorescence, the total number of inflorescences per plant, and the total number of flowers per plant. The hand-pollination experiment showed that fruit set was not pollen limited despite the low mating probability that resulted from the combination of dioecism and the species' dependence on opportunistic pollinators. This was due, in part, to the fact that female flowers did not have a predetermined period of receptivity, but instead remained receptive until they received pollen. In contrast, flowering phenology did affect fruit set. Fruit set was most abundant when male and female flowering was most abundant. This suggests there was some degree of pollen limitation during the part of the flowering season when male flowers were scarce.  相似文献   

9.
Bateman’s principle states that male fitness is usually limited by the number of matings achieved, while female fitness is usually limited by the resources available for reproduction. When applied to flowering plants this principle leads to the expectation that pollen limitation of fruit and seed set will be uncommon. However, if male searching for mates (including pollen dissemination via external agents) is not sufficiently successful, then the reproductive success of both sexes (or both sex functions in hermaphroditic plants) will be limited by number of matings rather than by resources, and Bateman’s principle cannot be expected to apply. Limitation of female success due to inadequate pollen receipt appears to be a common phenomenon in plants. Using published data on 258 species in which fecundity was reported for natural pollination and hand pollination with outcross pollen, I found significant pollen limitation at some times or in some sites in 159 of the 258 species (62%). When experiments were performed multiple times within a growing season, or in multiple sites or years, the statistical significance of pollen limitation commonly varied among times, sites or years, indicating that the pollination environment is not constant. There is some indication that, across species, supplemental pollen leads to increased fruit set more often than increased seed set within fruits, pointing to the importance of gamete packaging strategies in plant reproduction. Species that are highly self-incompatible obtain a greater benefit relative to natural pollination from artificial application of excess outcross pollen than do self-compatible species. This suggests that inadequate pollen receipt is a primary cause of low fecundity rates in perennial plants, which are often self-incompatible. Because flowering plants often allocate considerable resources to pollinator attraction, both export and receipt of pollen could be limited primarily by resource investment in floral advertisement and rewards. But whatever investment is made is attraction, pollinator behavioral stochasticity usually produces wide variation among flowers in reproductive success through both male and female functions. In such circumstances the optimal deployment of resources among megaspores, microspores, and pollinator attraction may often require more flowers or more ovules per flower than will usually be fertilized, in order to benefit from chance fluctuations that bring in large number of pollen grains. Maximizing seed set for the entire plant in a stochastic pollination environment might thus entail a packaging strategy for flower number or ovule number per flower that makes pollen limitation of fruit or seed set likely. Pollen availability may limit female success in individual flowers, entire plants (in a season or over a lifetime), or populations. The appropriate level must be distinguished depending on the nature of the question being addressed.  相似文献   

10.
Floral traits that increase attractiveness to pollinators are predicted to evolve through selection on male function rather than on female function. To determine the importance of male-biased selection in dioecious Wurmbea dioica, we examined sexual dimorphism in flower size and number and the effects of these traits on pollinator visitation and reproductive success of male and female plants. Males produced more and larger flowers than did females. Bees and butterflies responded to this dimorphism and visited males more frequently than females, although flies did not differentiate between the sexes. Within sexes, insect pollinators made more visits to and visited more flowers on plants with many flowers. However, visits per flower did not vary with flower number, indicating that visitation was proportional to the number of flowers per plant. When flower number was experimentally held constant, visitation increased with flower size under sunny but not overcast conditions. Flower size but not number affected pollen removal per flower in males and deposition in females. In males, pollen removal increased with flower size 3 days after flowers opened, but not after 6 days when 98% of pollen was removed. Males with larger flowers therefore, may have higher fitness not because pollen removal is more complete, but because pollen is removed more rapidly providing opportunities to pre-empt ovules. In females, pollen deposition increased with flower size 3 days but not 6 days after flowers opened. At both times, deposition exceeded ovule production by four-fold or more, and for 2 years seed production was not limited by pollen. Flower size had no effect on seed production per plant and was negatively related to percent seed set, implying a tradeoff between allocation to attraction and reproductive success. This indicates that larger flower size in females is unlikely to increase fitness. In both sexes, gamete production was positively correlated with flower size. In males, greater pollen production would increase the advantage of large flowers, but in females more ovules may represent a resource cost. Selection to increase flower size and number in W. dioica has probably occurred through male rather than female function. Received: 15 June 1997 / Accepted: 12 February 1998  相似文献   

11.
Male and hermaphrodite flowers from the andromonoecious perennial Solanum carolinense were compared. Hermaphrodite flowers were 11–16% heavier than males, and this difference was consistent for two clones studied. Hermaphrodite flowers contained a greater amount of nitrogen than did male flowers, but males had higher percent nitrogen. Male and hermaphrodite flowers had equal investment in male structures: stamen mass, number of pollen grains, mass of pollen grains ejected by vibration, and pollen size were all similar for the two flower types. In contrast, male flowers had reduced masses of disfunctional female structures: ovaries of males were ⅓ as massive, and styles 1/7; as massive, as those of hermaphrodites. These results are consistent with the hypothesis that male flowers (which occur where fruit set is unlikely) increase male fitness through pollen donation, while saving resources that would otherwise be invested in hermaphrodite flowers that abscise.  相似文献   

12.
The effect of nectar robbing on plant fitness is poorly understood and restricted to a few plant species. Furthermore, the available studies generally evaluate the effects of nectar robbing on female fitness, disregarding the male component. Here we measured the effects of the nectar-robbing bumblebees on male (measured as pollen analogue flow distance) and female (measured as seed production) reproductive success in the insect-dependent Polygala vayredae, a narrow endemic species from the pre-Pyrenees (Spain). Intense nectar robbing by bumblebees significantly reduced the nectar available to legitimate pollinators in the studied population, and this reduction affected both male and female fitness. Significant differences were observed in fluorescent dye dispersion between robbed and non-robbed flowers within the population. Fluorescent dyes from non-robbed flowers were dispersed to larger distances and over a larger number of flowers when compared with robbed ones. Moreover, significant differences were observed in both fruit set and seed ovule ratios between the two groups, with non-robbed flowers presenting higher reproductive outcomes. However, no effect on seed weight was detected among treatments. The data obtained suggest that in this species, nectar robbing has important indirect and negative effects on plant fecundity, through both male and female functions, due to a modification in the foraging behaviour of legitimate visitors.  相似文献   

13.
Does the mode of self-pollination affect the evolutionarily stable allocation to male vs. female function? We distinguish the following scenarios. (1) An ‘autogamous’ species, in which selfing occurs within the flower prior to opening. The pollen used in selfing is a constant fraction of all pollen grains produced. (2) A species with ‘abiotic pollination’, in which selfing occurs when pollen dispersed in one flower lands on the stigma of a nearby flower on the same plant (geitonogamy). The selfing rate increases with male allocation but a higher selfing rate does not mean a reduced export of pollen. (3) An ‘animal-pollinated’ species with geitonogamous selfing. Here the selfing rate also increases with male allocation, but pollen export to other plants in the population is a decelerating function of the number of simultaneously open flowers. In all three models selfing selects for increased female allocation. For model 3 this contradicts the general opinion that geitonogamous selfing does not affect evolutionarily stable allocations. In all models, the parent benefits more from a female-biased allocation than any other individual in the population. In addition, in models 2 and 3, greater male allocation results in more local mate competition. In model 3 and in model 2 with low levels of inbreeding depression, hermaphroditism is evolutionarily stable. In model 2 with high inbreeding depression, the population converges to a fitness minimum for the relative allocation to male function. In this case the fitness set is bowed inwards, corresponding with accelerating fitness gain curves. If the selfing rate increases with plant size, this is a sufficient condition for size-dependent sex allocation (more allocation towards seeds in large plants) to evolve. We discuss our results in relation to size-dependent sex allocation in plants and in relation to the evolution of dioecy.  相似文献   

14.
A detailed examination was made of the numbers of pollen grains and microgametophytes occurring on stigmas of a dioecious, wind-pollinated species, Staberoha banksii. The number of pollen grains per pistil ranged from 0 to over 100; and the mean pollen load size per pistil per plant varied from 4.4 to 28. Distance to the fourth nearest male plant was used as a measure of local pollen availability and accounted for 63.5% of the between-plant variation in pollen load size. In contrast, the number of microgametophytes did not show a leptokurtic decline with increasing distance from a pollen source. On average there were 5.19 microgametophytes per ovule, which is higher than all comparable data for insect-pollinated taxa, and 66% of the stigmas had two or more microgametophytes. Given that there is a single ovule per flower, there is considerable potential for gametophytic competition. Seedset was very high, averaging 93%, and showed no significant decline in isolated female plants. We suggest that the quantity of pollen is not limiting fertilization in this population and that wind may therefore be highly effective as a pollen vector.  相似文献   

15.
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.  相似文献   

16.
Teak is a timber tree that is widely distributed in the tropics. Several studies on pollination and reproductive biology have been conducted, but generally information on flowering phenology and annual variation in total pollen production per tree is lacking. The reproductive phenology as well as flower‐ and pollen grain production of individuals in a population is important to theoreticians, field biologists and plant breeders, as they determine the distribution of genotypes within populations and influences the degree of differentiation among populations. This study reports flowering phenology and variation in total flower, fruit‐ and pollen production per tree in teak in a 25‐year‐old plantation across three consecutive years (2006–2008). The results show that the date of onset and end of flowering was highly variable across years. The longest flowering period of 93 days was observed in 2006. There was an asynchrony in the number of open flowers due to differences in time of anthesis among individuals (± 2 days) and inflorescences within individuals (± 6 h). The production of pollen grains per tree in 2007 was 33%, i.e. 16% more compared to 2006 and 2008. The fruit production per tree was 42% and 27% higher in 2007 compared to 2006 and 2008. Concentration of pollen grains (both on jelly‐coated microscopic slides and stigmas) were highest between noon and 2 pm. At this time, the stigmatic pollen load ranged between 4–8 pollen grains per stigma, which is sufficient for fruit development. The study concludes that the asynchronization of the flower opening might give rise to a high amount of self‐pollination in the stand, ultimately leading to poor fruit setting. Also, the large production of flowers and pollen per tree induced geitonogamy and decreased female fitness, as T. grandis is preferentially an out‐crossing species.  相似文献   

17.
Herbivory is an important selection pressure in the life history of plants. Most studies use seed or fruit production as an indication of plant fitness, but the impact of herbivory on male reproductive success is usually ignored. It is possible that plants compensate for resources lost to herbivory by shifting the allocation from seed production to pollen production and export, or vice versa. This study examined the impact of herbivory by Helix aspersa on both male and female reproductive traits of a monoecious plant, Cucumis sativus. The effects of herbivory on the relative allocation to male and female flowers were assessed through measurements of the number and size of flowers of both sexes, and the amount of pollinator visitation. We performed two glasshouse experiments; the first looked at the impact of three levels of pre-flowering herbivory, and the second looked at four levels of herbivory after the plants had started to flower. We found that herbivory during the flowering phase led to a significant increase in the number of plants without male flowers. As a consequence there was significantly less pollen export from this population, as estimated by movement of a pollen analog. The size of female flowers was reduced by severe herbivory, but there was no affect on pollen receipt by the female flowers of damaged plants. The decrease in allocation to male function after severe herbivory may be adaptive when male reproductive success is very unpredictable.  相似文献   

18.
慈姑花的开放式样及其花粉流   总被引:3,自引:0,他引:3  
对一野生慈姑(SagitariatrifoliaL.)居群的性表达状态及其花粉流散布规律进行了观测。慈姑雌雄同株,单株雄花数是雌花数的3.6倍。雌雄花单花期均为1d,但二者开放式样不同。尽管雌花的数目较少,但在1d内开放较多的花,其花的开放速率比雄花快。在居群水平上表现出雌雄花比例随时间大幅度波动。在传粉者进行的2612次访花活动中,传粉者表现出偏“雌性”访问,对每朵雌花的访问是雄花的1.4倍。用番红染色法标记慈姑的花粉运动表明,花的不同开放式样影响花粉流的散布。当雄花数占高比例时,花粉主要在邻近的个体间散布;而雄花的比例较低、雌花相对较多时,花粉远距离散布的强度加大。  相似文献   

19.
Summary Flower and fruit characters were measured in ten female, five male and five fruiting male selections of A. deliciosa var deliciosa (A. Chev) Liang and Ferguson. Flowers from female vines had functional pistils, which contained many ovules. Stamens appeared to be fully developed but produced only empty pollen grains. Flowers from male vines had functional stamens that produced high percentages of pollen grains with stainable cytoplasmic contents. Pistils did not contain ovules and were generally small with vestigial styles. Fruiting male vines had both staminate and bisexual flowers. Staminate flowers were similar to those found on strictly male vines. Bisexual flowers produced ovules and stainable pollen. Pistils were smaller than in pistillate flowers. Although the three flower sexes differed in style length, ovary dimensions and ovules per carpel, staminate and bisexual flowers were similar in number of flowers per inflorescence, stamen filament length, pollen stainability, inflorescence rachis length and carpel number, and differed from pistillate flowers in these characters. The three flower sexes had similar sepal and petal numbers. The fruit of fruiting males were considerably smaller than those of females. Low ovule number appears to be the major factor limiting fruit size in the fruiting males studied. Prospects for developing hermaphroditic kiwifruit cultivars through breeding are discussed.  相似文献   

20.
The influence of pollen competitive environment on pollen performance (pollen germination, stigmatic penetration, and pollen tube growth rate), the maturation or abortion of initiated fruit, seed size, and seedling vigor was explored by manipulating the size and diversity of stigmatic pollen loads on Mirabilis jalapa. All aspects of pollen performance significantly increased with the number of pollen grains on a stigma or pollen tubes in a style, but was not influenced by the diversity of pollen donors. Plants tended to mature single-ovulate fruits that came from flowers where pollen load size and diversity were greatest and aborted those where these were lowest. No plants from seeds resulting from pollinations with a single pollen grain survived, but other fitness measures were mostly determined by maternal plant. The data suggest that pollen performance is influenced by pollen competitive environment, and both the genetic diversity of the pollen load and number of competing pollen tubes are important determinants of seed/fruit abortion.  相似文献   

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