Local plant density,pollination and trait–fitness relationships in a perennial herb

Both differences in local plant density and phenotypic traits may affect pollination and plant reproduction, but little is known about how density affects trait–fitness relationships via changes in pollinator activity. In this study we examined how plant density and traits interact to determine pollinator behaviour and female reproductive success in the self‐incompatible, perennial herb Phyteuma spicatum. Specifically, we hypothesised that limited pollination service in more isolated plants would lead to increased selection for traits that attract pollinators. We conducted pollinator observations and assessed trait–fitness relationships in a natural population, whose individuals were surrounded by a variable number of inflorescences. Both local plant density and plant phenotypic traits affected pollinator foraging behaviour. At low densities, pollinator visitation rates were low, but increased with increasing inflorescence size, while this relationship disappeared at high densities, where visitation rates were higher. Plant fitness, in terms of seed production per plant and per capsule, was related to both floral display size and flowering time. Seed production increased with increasing inflorescence size and was highest at peak flowering. However, trait–fitness relationships were not density‐dependent, and differences in seed production did not appear to be related to differences in pollination. The reasons for this remain unclear, and additional studies are needed to fully understand and explain the observed patterns.     

Allocation to reproduction following experimental defoliation in Platanthera bifolia (Orchidaceae)

The roles of herbivory and pollination success in plant reproduction have frequently been examined, but interactions between these two factors have gained much less attention. In three field experiments, we examined whether artificial defoliation affects allocation to attractiveness to pollinators, pollen production, female reproductive success and subsequent growth in Platanthera bifolia L. (Rich.). We also recorded the effects of inflorescence size on these variables. We studied the effects of defoliation on reproductive success of individual flowers in three sections of inflorescence. Defoliation and inflorescence size did not have any negative effects on the proportion of opened flowers, spur length, nectar production or the weight of pollinia. However, we found that hand-pollination increased relative seed production and defoliation decreased seed set in most cases. Interactions between hand-pollination and defoliation were non-significant indicating that defoliation did not affect female reproductive success indirectly via decreased pollinator attraction. Plants with a large inflorescence produced relatively more seeds than plants with a small inflorescence only after hand-pollination. The negative effect of defoliation on relative capsule production was most clearly seen in the upper sections of the inflorescence. In addition to within season effects of leaf removal, defoliated P. bifolia plants may also have decreased lifetime fitness as a result of lower seed set within a season and because of a lower number of reproductive events due to decreased plant size (leaf area) following defoliation. Our study thus shows that defoliation by herbivores may crucially affect reproductive success of P. bifolia.     

REPRODUCTIVE SUCCESS AND INFLORESCENCE SIZE OF CALOPOGON TUBEROSUS (ORCHIDACEAE)

Reproductive success of Calopogon tuberosus, which produces no nectar, was investigated in relation to inflorescence size and dispersion pattern. Mean inflorescence size was 2.56 (range 1–10). A bagging experiment showed that insects are required for pollen transfer and that fruits are produced from self-, geitonogamous, and cross-pollinations; fruit set was not 100%. Fruit set of nonmanipulated plants was limited by the number of pollinator visits. Reproductive success increased with increasing inflorescence size, although not above theoretical predictions. However, the probability of producing no fruit or contributing no pollinia decreased with increasing inflorescence size since sequential flowering increased the probability of a pollinator visit to the inflorescence over the blooming period. Large inflorescences did not provide a greater pollinator attraction than small ones, because inflorescences only presented a few open flowers at a time. In addition, flowers on plants growing in clumps of 2–8 plants had a higher probability of setting fruit, apparently because of increased pollinator attraction. Although there are obvious selective advantages for large inflorescences, the sequential flowering habit, and low resource availability may reduce the advantages of large inflorescence size at our study site.     

Changes in Florets’ Vertical Direction within Inflorescence Affects Pollinator Behavior,and Fitness in <i>Trifolium repens</i>

Ecological interactions between flowers and pollinators greatly affect the reproductive success. To facilitate these interactions, many flowers are known to display their attractive qualities, such as scent emission, flower rewards and floral vertical direction, in a rhythmic fashion. However, less is known about how plants regulate the relationship between these flower traits to adapt to pollinator visiting behavior and increase reproduction success. Here we investigated the adaptive significance of the flower bending from erect to downward in Trifolium repens. We observed the flowering dynamic characteristics (changes of vertical direction of florets, flowering number, pollen grain numbers, pollen viability and stigma receptivity over time after blossom) and the factors affecting the rate of flower bending in T. repens. Then we altered the vertical direction of florets in inflorescence of different types (upright and downward), and compared the pollinator behaviors and female reproductive success. Our results showed that florets opened sequentially in inflorescence, and then bend downwards slowly after flowering. The bending speed of florets was mainly influenced by pollination, and bending angle increased with the prolongation of flowering time, while the pollen germination rate, stigma receptivity and nectar secretion has a rhythm of “low-high-low” during the whole period with the time going. The visiting frequency of all the four species of pollinators on upward flowers was significantly higher than that of downward flowers, and they especially prefer to visit flowers with a bending angle of 30°–60°, when the flowers was exactly of the highest flower rewards (nectar secretion and number of pollen grains), stigma receptivity and pollen germination rate. The seed set ratio and fruit set ratio of upward flowers were significantly higher than downward flowers, but significantly lower than unmanipulated flowers. Our results indicated that the T. repens could increase female and male fitness by accurate pollination. The most suitable flower angle saves pollinators’ visiting energy and enables them to obtain the highest nectar rewards. This coordination between plants and pollinators maximizes the interests of them, which is a crucial factor in initiating specialized plant-pollinator relationships.     

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

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

Spatial and temporal dispersion patterns of pollinators and their relationship to the flowering strategy of Yucca whipplei (Agavaceae)

Summary A field investigation of the mutualistic interaction between a monocarpic perennial plant, Yucca whipplei, and its host-specific pollinator and seed predator, Tegeticula maculata (Lepidoptera: Prodoxidae), was conducted to determine how the resource utilization pattern and population dynamics of the pollinator have influenced the evolution of the flowering and fruiting pattern of the plant. Although the temporal pattern of emergence of pollinators results in a relatively close tracking of flower abundance within a season, the ratio of pollinators to open flowers does vary significantly within a season, as well as between seasons. At any point in time during the flowering season, the population of adult yucca moths is distributed evenly among the available flowers, so that the number of pollinators on an inflorescence is directly proportional to the number of open flowers available. The relative isolation of individual flowering plants appears to have little effect on the distribution of pollinators among inflorescences. The number of fruits initiated on a plant is directly proportional to the number of flowers produced, and is also partially determined by the time of flowering. Yucca whipplei always produces many more flowers than fruits. Most flowers are not fertilized, and the plants also generally abort and abscise immature fruits after flowering. Fruit production of at least some plants, however, appeared limited by pollination. It is also expected that in some years the relative abundance of pollinators will be low enough that most plants will be pollinator-limited. It is suggested that the pattern of flowering and fruiting of this species has evolved in response to the unpredictability of pollinator availability, both within and between seasons. Resource uncertainty and selection acting on the male component of fitness may also be involved.     

Up and down asymmetrical world for flower blooming.

Form of plant body shows vertical polarity. Photosynthetic organs deploy upwards to seek light. Root system extends downward for water. In addition to this major polarity, flowers has similar features, because they bloom in the world where up and down asymmetry dominates. Many flowering plants co-evolved with pollinator animals. Success of their reproduction is linked to the shape of flower organs in many ways. Orientation of inflorescence and each individual flowers, arrangement and shape of flower organs exhibit various up and down asymmetry. Some flowers mimic female of its pollinator animal. In such case, posture of the target animal is copied to the flower. Since animal posture and shape have vertical polarity, flower happens to equip same kind of polarity. Tropic response of pistils is another up and down feature that improves fitness of flowering plants. Certain lily flowers show phototropism to bend pistil upward. Azalea flower depends on gravity as the major environmental cue, and light as the secondary signal for up. Molecular machinery for those tropic responses seems to be shared with other tropism expressed in shoot and root. However, certain differences are found in distribution of sediment amyloplast, or spatial allocation of photo-sensing and bending site. Tropic responses result in adaptation of those flowers against behavior of pollinator animals and terrestrial environment, where gravity affects living organisms and interaction among them.     

THE FLORAL ECOLOGY OF PLATANTHERA BLEPHARIGLOTTIS

The floral ecology of a central Maine population of Platanthera blephariglottis was studied during the 1979–1981 flowering seasons. Although this species possesses characteristics typical of moth-pollinated plants, only diurnal pollinators were documented, primarily Hesperiidae, Pieridae, and Apidae. Pollinators were scarce throughout the study, yet bagging experiments indicated vectors were necessary for successful capsule set. The average capsule set per plant was lowest (32.2%) in 1979 when 362 plants bloomed and highest in 1980 (80.2%) when 202 plants flowered. Capsule set was found to be limited by pollinator activity. The adaptations of P. blephariglottis, including sequential flowering, length of receptivity, nectar production and inflorescence size, enable consistently successful reproduction regardless of microhabitat. This species is well-adapted to reproduce in environments experiencing periods of unfavorable weather and low pollinator abundance.     

The effect of pollination on resource allocation among sexual reproduction,clonal reproduction,and vegetative growth in <Emphasis Type="Italic">Sagittaria potamogetifolia</Emphasis> (Alismataceae)

In order to evaluate the effects of pollination on resource allocation in the marsh herb Sagittaria potamogetifolia, experimental manipulation of pollination efficiency on the pattern of resource allocation was accessed by the proportion of dry weight measurements of sexual, vegetative, and clonal organs. In trials where half of the flowers were pollinated, a significant increase of resource allocated to sexual production and decrease to vegetative production resulted compared to plants that received no pollination. In trials where pollination was 100%, these two reproductive components showed the same trend, but less dramatically. This may support the idea that the trade-offs would be more pronounced when the resource was scarce. Besides, a higher inflorescence production with a lower fruit reproduction occurred as a consequence of decreased pollination level. This increased inflorescence production may be a mechanism to promote outcrossing by enhancing floral attraction or by synchronizing reproductive activity with insect pollinators. Examination for possible trade-offs in resource allocation revealed that there was also a trade-off caused by pollination between fruits plus flowers and bulbils production, which might have detrimental effects on the survival of individuals and populations, but promotes outcrossing and genetic variability.     

Refuse attracts? Effect of refuse dumps of leaf‐cutting ants on floral traits

The nutrient‐rich organic waste generated by ants may affect plant reproductive success directly by enhancing fruit production but also indirectly, by affecting floral traits related with pollinator attraction. Understanding how these soil‐nutrient hot spots influence floral phenotype is relevant to plant–pollination interactions. We experimentally evaluated whether the addition of organic waste from refuse dumps of the leaf‐cutting ant Acromyrmex lobicornis (Hymenoptera: Formicidae: Attini) alters floral traits associated with pollinator attraction in Eschscholzia californica (Ranunculales: Papaveraceae), an entomophilous herb. We analysed flower shape and size using geometric morphometric techniques in plants with and without the addition of refuse‐dumps soil, under greenhouse conditions. We also measured the duration of flowering season, days with new flowers, flower production and floral display size. Plants growing in refuse‐dumps soil showed higher flower shape diversity than those in control soil. Moreover, plants in refuse‐dumps soil showed bigger flower and floral display size, longer flowering season, higher number of flowering days and flower production. As all these variables may potentially increase pollinator visits, plants in refuse‐dumps soil might increase their fitness through enhanced attraction. Our work describes how organic waste from ant nests may enhance floral traits involved in floral attraction, illustrating a novel way of how ants may indirectly benefit plants.     

Fruit to flower ratios and trade-offs in size and number

Summary Hermaphroditic flowering plants commonly produce many more flowers than fruits. To interpret this observation, I suggest an explanation based on trade-offs occurring between the size and the number of flowers and fruits that mature when resources are limiting. I make this suggestion concrete with the aid of a phenotypic model analysed using the ESS approach. The model includes resource allocations to male and female reproductive structures at the time of flowering and to fruit maturation. The formulation allows for non-linear relations between opportunities for fertility gain on the one hand and flower and fruit size and number on the other. Results of the model indicate that fewer fruits will be matured than flowers produced, even in the absence of previously suggested factors such as pollinator limitation and bet-hedging in the face of environmental variability. Model results emphasize the distinction between flower versus inflorescence contributions to male and female reproductive success. Analysis also shows that the fruit to flower ratio in hermaphroditic plants will be lower than that in female plants, in accord with broad taxonomic surveys. Finally, results obtained lead to empirically testable predictions about the relationship between flower size, fruit size and the fruit to flower ratio. Methods to test these predictions are discussed.     

Carbon source for reproduction in a spring ephemeral herb, Corydalis ambigua (Papaveraceae)

1.  The carbon source for reproduction in plants may differ between flowering and fruiting stages. To clarify how spring ephemerals use current photosynthetic products for reproduction, the allocation patterns of photosynthate at flowering and fruiting and the effects of resource limitation on reproductive performance in Corydalis ambigua were assessed.
2.  A ¹³C tracing experiment revealed that about 20% of the current photosynthetic carbon was used for reproduction at both flowering and fruiting. The proportion of ¹³C allocated to fruits was constant irrespective of the light level. In contrast, ¹³C translocation to tubers increased at fruiting, and this trend was accelerated when plants were shaded.
3.  Defoliation treatment significantly reduced nectar production and tuber mass, while seed production was not affected. Therefore, when carbon assimilation was limited, carbon was preferentially allocated to current reproduction (seeds) rather than to pollinator attraction (nectar) or storage (tuber).
4.  If seed production is partly supported by carbohydrate reserved in the old tissue of tubers, nectar and seed production may not compete strongly for carbon sources. In contrast to the ability of high seed production, the susceptibility of nectar production to current photosynthesis indicates that seed production of this species is basically limited by pollen capture.
5.  Therefore, temporal separation of resource pool for reproduction may mitigate the joint limitation of seed production between pollinator attraction and resource availability. Temporal variation of the sink–source balance of storage organ is crucial to understand the cost of reproduction in perennial plants.     

Pollen limitation and natural selection on floral characters in the yellow monkeyflower, Mimulus guttatus

In flowering plants, pollen limitation has been proposed to intensify selection on floral characters important in pollinator attraction, but may also select for traits that increase seed set through autonomous selfing. Here, a factorial design (+/- pollen addition, +/- pollinator removal) was used to investigate how the pollination environment affects selection on floral morphology via female fitness in a mixed-mating population of the yellow monkeyflower, Mimulus guttatus (Phrymaceae). Female fitness was strongly pollen-limited, with supplementally pollinated plants setting 37% more seeds than open-pollinated individuals. Strong positive selection was found on flower length, weak positive selection on flower width : length ratio and no selection on stigma-anther distance in both open-pollinated and supplementally pollinated treatments. By contrast, flowers with relatively narrow corollas and low stigma-anther distances were favored in the pollinator exclusion treatment. These results provide mixed support for the idea that pollen limitation intensifies selection on floral characters. Despite strong phenotypic selection, natural pollen limitation did not mediate selection on characters associated with either pollinator attraction or self-fertilization. However, the novel pattern of selection on severely pollen-limited plants suggests that reproductive assurance against pollinator loss may have been directly involved in the floral evolution of closely related selfing taxa.     

Effects of pollen reward removal on fecundity in a self-incompatible hermaphrodite plant

The pollen of hermaphrodite plants is often utilised by flower-visiting animals. While pollen production has obvious benefits for plant male fitness, its consequences for plant female fitness, especially in self-incompatible hermaphrodite species, are less certain. Pollen production could either enhance seed production though increased pollinator attraction, or reduce it if ovules are discounted by deposition of self pollen, as can occur in species with late-acting self-incompatibility. To test the effects of pollen reward provision on female fitness, we artificially emasculated flowers in two populations of the succulent Aloe maculata (Asphodelaceae), which has a late-acting self-incompatibility system, over the course of its flowering period. Flowers of this species are visited by sunbirds (for nectar) and native bees (for pollen and nectar). We measured floral visitation rates, floral rejection rates, pollen deposition on stigmas and fruit and seed set in both emasculated and non-emasculated plants. We found that flowers of emasculated plants suffered reduced visitation and increased rejection (arrival without visitation) by bees, but not by sunbirds; had fewer pollen grains deposited on stigmas and showed an overall decrease in fruit set and seed set. Rates of seed abortion were, however, greatly reduced in emasculated flowers. This study shows that pollen rewards can be important for seed set, even in self-incompatible plants, which have been assumed to rely on nectar rewards for pollinator attraction. Seed abortion was, however, increased by pollen production, a result that highlights the complexity of selection on pollen production in hermaphrodite flowers.     

开花时间与伴生种对鹤首马先蒿传粉和生殖成功的影响

开花物候是物种间相互作用的重要生活史特征和适合度因子,在全球气候变化的背景下而备受关注.为探讨开花时间如何存种内和种间水平上影响植物的传粉和生殖成功,我们连续3年(2003-2005)对不同花期和伴生种存在情况下的鹤首马先蒿(Pedicularis gruina)的传粉者访花忠实度、受粉率、坐果率、单果种子产量和果实被啃食频率进行了比较研究.结果表明鹤首马先蒿的坐果率主要受其传粉环境的影响:在没有伴生种时,不同时期鹤首马先蒿的坐果率没有显著差异,34-38％的花可以坐果;在有伴生种存在时既可以显著提高其坐果率,也可显著降低其坐果率,这取决于传粉者类型以及伴生种密穗马先蒿(P.densispica)花期的差异.密穗马先蒿具有花蜜和花粉双重报酬,在群落中可以作为主体物种吸引传粉者并间接促进与其伴生的鹤首马先蒿的传粉和生殖成功.同样无蜜的管花马先蒿(P.siphonantha)和鹤首马先蒿伴生,则是通过提高群落水平对传粉者的吸引力进而直接促进鹤首马先蒿的传粉和生殖成功.此外,研究结果也表明开花时间对坐果率没有显著影响,但是显著影响单果种子产量和果实被啃食的频率.在相同的传粉条件下早花期植株单果种子产量显著高于晚花期的种子产量,同时早花期的果实被啃食的频率显著增加.     

克隆生长对被子植物传粉过程的影响

克隆植物与其传粉者的相互作用是植物繁殖生态学的重要研究领域之一。植物克隆生长与有性繁殖通常相伴进行, 往往产生较大的花展示与复杂的克隆空间结构, 通过传粉过程对有性繁殖过程产生影响, 共同决定植物的适合度。本文回顾了克隆生长对被子植物传粉过程影响的国内外研究进展, 从植物克隆大小、花资源空间配置、克隆构型与种群遗传结构四个方面讨论了克隆生长对传粉过程的影响及其生态学与进化生物学意义。早期研究预期, 随着克隆增大, 同株异花授粉水平增加, 因而通过增大自交率或花粉阻塞效应降低植物的适合度。但是, 后来的一些模拟与野外实验研究发现, 传粉者在同一克隆内访问的花数量并不会随克隆增大而一直增加, 访花行为也主要发生在分株内; 而且分子标记的自交率组分分析也表明自交主要发生在分株内。另一方面, 人工模型模拟以及传粉者访问行为研究表明, 当花朵数量相同时, 与所有花集中生长在同一分株上相比, 将花朵分散在多个分株上的克隆生长方式不会增加, 反而降低了同株异花授粉的发生水平。如果花序内花雌雄同熟, 花朵同时提供与接收花粉, 克隆生长会使植物接收到更高比例的异交花粉, 在提高后代质量的同时不增加同株异花授粉概率。这是从传粉生物学角度对植物克隆生长习性进化的一个全新的解释。今后, 克隆植物传粉生物学研究需要针对传粉者与克隆生长之间的相互作用建立理论模型, 探究克隆大小、克隆构型、花资源空间配置模式对传粉者访问频率和行为、花粉散布、交配格局的影响。同时, 需要在自然种群中, 尤其是克隆与非克隆的近缘类群、同一物种克隆与非克隆种群开展比较研究, 利用更高效的分子标记来研究克隆生长的生态与进化意义。     

INFLORESCENCE ARCHITECTURE: HOW FLOWER NUMBER,ARRANGEMENT, AND PHENOLOGY AFFECT POLLINATION AND FRUIT-SET

The adaptive significance of different types of inflorescences in flowering plants has been largely ignored. The few published studies investigating adaptive aspects of floral displays suggest that numbers of flowers and their arrangement in space and time determine levels of pollination and fruit-set in natural populations. The frequently conflicting demands placed on inflorescence architecture have led to an evolutionary compromise that maximizes the genetic contribution of an individual plant to the next generation. These conflicting demands include pollinator attraction vs. ovary competition, fruit dispersal vs. fruit predation, and reproductive vs. vegetative resource allocation. In most cases, the inflorescence size most successful in fruit production is also the most frequent in natural populations. In addition to quantity of offspring, inflorescence architecture affects, and in turn is affected by, the quality of offspring that result from selfing vs. outcrossing.     

Determinants of seed production in Geranium maculatum

Summary We experimentally examined factors limiting seed production in two populations of the perennial woodland herb Geranium maculatum in central Illinois, USA. To test the pollinator-limitation hypothesis, we compared the seed production of plants whose flowers were supplementarily pollinated with outcross pollen to that of control plants receiving natural pollination only. To test if fruit production by early flowers suppresses fruit and seed formation by late flowers, a third group of plants was prevented from producing seed from the first 50% of the flowers to open (stigmas were excised at flower opening). Finally, to test if seed maturation and flower initiation are correlated with photosynthetic capacity, we performed a defoliation experiment in which either the stem leaves within the inflorescence, the stem leaves below the inflorescence, or the rosette leaves were removed during late flowering. Plants that reccived supplemental pollination produced 1.5–1.6 times more seeds than control plants. We found no difference between hand-pollinated plants and controls in mortality, flowering frequency or number of flowers produced in the year following the experiment. In both control and hand-pollinated plants, the fruit set and total seed production of early flowers were more than twice as high as those of late flowers. In one of the two populations, plants whose early flowers were prevented from setting seed produced significantly more seeds from their late flowers than did control plants. Seed predation was low and did not differ between early and late flowers. Leaf removal did not affect seed number or size in the year of defoliation, nor did it reduce survival or flower production in the subsequent year. This suggests that the plants were able to compensate for a partial defoliation by using stored resources or by increasing photosynthetic rates in the remaining leaves. Taken together, the results demonstrate that both pollinator activity and resource levels influence patterns of seed production in G. maculatum. While seed production was pollinatorlimited in both populations, a seasonal decline in resource availability was apparently responsible for the low seed production by late flowers.     

The reproductive biology of a riparian Mediterranean shrub, Nerium oleander L. (Apocynaceae)

This study presents data on the reproductive characteristics of Nerium oleander, an evergreen, sclerophyllous shrub inhabiting the banks of temporary streams in all countries around the Mediterranean seaboard. Two southern Spanish populations (El Garrobo and Grazalema) were selected to investigate flowering, dispersal and seed germination phenologies, together with morphological and functional traits of flowers, fruits and seeds. Observations on pollinator visitation, fruit set, and seedling survival were also carried out. Plants produce massive flower displays of showy, tubular flowers void of nectar that rely on insect cheating for pollination. Hand-pollinations and seed germination tests demonstrated full self-compatibility. Automatic selfing is prevented by spatial separation of stigma and anthers, and pollinators are thus necessary for reproduction. Low pollinator visitation rates are the rule, as indicated by the fact that observations spread over 2 years yielded only eight insect records. Percentage fruit set of open-pollinated flowers was found to be extremely low (0.1–4.9%), while hand-pollination increased fruiting levels to 34–50%. Direct and indirect evidence point to consistently pollen-limited reproduction in this species. If a large number of flowers within any inflorescence are pollinated artificially, however, resource limitations operate and pods from ovaries pollinated late in the season are aborted. On average, the maximum of ovaries developing into fruit within any inflorescence is 4. It is suggested that, in this nectarless species, about 80% of total are ‘excess’ flowers which contribute to increase pollinator attraction. Nerium oleander pollen is aggregated into a sticky substance, which allows the fertilization of many ovules after a single, probably rare, pollination event. Fruits are many seeded follicles (181 seeds per fruit on average), so that individual plants often liberate thousands of seeds in spite of the low percent fruit set. The hairy, water-dispersed seeds are released during the rainiest season in the year, and germination follows rapidly. All seedlings tracked died before completing 1 year of life, mostly because of desiccation during the first summer drought. Pollination by deceit in this Mediterranean species is discussed in terms of phylogenetic constraints within the mainly tropical family Apocynaceae.     

Bateman’s principle and plant reproduction: The role of pollen limitation in fruit and seed set

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.