The economy of pollen dispersal in flowering plants.

Mating success of flowering plants depends strongly on the efficiencies of pollen removal from flowers and its subsequent dispersal to conspecific stigmas. We characterized the economy of pollen dispersal in flowering plants by analysing pollen fates and their correlates for 228 species. The mean percentage of pollen removed from flowers (removal efficiency) varied almost twofold according to the type of pollen-dispersal unit, from less than 45% for orchids and milkweeds with solid pollinia, to greater than 80% for species with granular monads or sectile (segmented) pollinia. The mean percentage of removed pollen reaching stigmas (pollen transfer efficiency, PTE) varied from 2.4% for species with separate monads to 27.0% for orchids with solid pollinia. These values tended to be higher in plants with single pollinator species and in those with non-grooming pollinators. Nectar production increased removal efficiency, but did not influence PTE. Among types of pollen-dispersal units, the net percentage of produced pollen that was dispersed to stigmas varied negatively with removal efficiency and positively with PTE, indicating the relative importance of the latter for overall pollen economy. These findings confirm the key importance of floral traits, particularly pollen packaging, for pollen dispersal outcomes and highlight the under-appreciated pollination efficiency of non-grooming pollinators.

Pollinators differ in their contribution to the male fitness of a self-incompatible composite.

PREMISE: Reproductive fitness in plants is often determined by the quantity and quality of pollen transferred by pollinators. However, many fitness studies measure only female fitness or rely on proxies for male fitness. Here we assessed how five bee taxon groups affect male fitness in a prairie plant by quantifying pollen removal, visitation, and siring success using paternity assignments and a unique pollinator visitation experiment. METHODS: In Echinacea angustifolia, we measured per-visit pollen removal for each pollinator taxon and estimated the number of pollen grains needed for successful ovule fertilization. Additionally, we directly measured pollinator influence on siring by allowing only one bee taxon to visit each pollen-donor plant, while open-pollinated plants acted as unrestricted pollen recipients. We genotyped the resulting offspring, assigned paternity, and used aster statistical models to quantify siring success. RESULTS: Siring success of pollen-donor plants differed among the five pollinator groups. Nongrooming male bees were associated with increased siring success. Bees from all taxa removed most of the flowering head's pollen in one visit. However, coneflower-specialist bee Andrena helianthiformis removed the most pollen per visit. Female fitness and proxy measures of male fitness, such as pollinator visitation and pollen removal, did not align with our direct quantifications of male fitness. CONCLUSIONS: Our results illustrate the need for more studies to directly quantify male fitness, and we caution against using male fitness proxy measures. In addition, conservation efforts that preserve a diverse pollinator community can benefit plants in fragmented landscapes.

Flower orientation influences floral temperature, pollinator visits and plant fitness.

Effective insect pollination requires appropriate responses to internal and external environmental cues in both the plant and the pollinator. Helianthus annuus, a highly outcrossing species, is marked for its uniform eastward orientation of mature pseudanthia, or capitula. Here we investigate how this orientation affects floral microclimate and the consequent effects on plant and pollinator interactions and reproductive fitness. We artificially manipulated sunflower capitulum orientation and temperature in both field and controlled conditions and assessed flower physiology, pollinator visits, seed traits and siring success. East-facing capitula were found to have earlier style elongation, pollen presentation and pollinator visits compared with capitula manipulated to face west. East-facing capitula also sired more offspring than west-facing capitula and under some conditions produced heavier and better-filled seeds. Local ambient temperature change on the capitulum was found to be a key factor regulating the timing of style elongation, pollen emergence and pollinator visits. These results indicate that eastward capitulum orientation helps to control daily rhythms in floral temperature, with direct consequences on the timing of style elongation and pollen emergence, pollinator visitation, and plant fitness.

Pollination intensity and paternity in flowering plants.

BACKGROUND: Siring success plays a key role in plant evolution and reproductive ecology, and variation among individuals creates an opportunity for selection to act. Differences in male reproductive success can be caused by processes that occur during two stages, the pollination and post-pollination phases of reproduction. In the pollination phase, heritable variation in floral traits and floral display affect pollinator visitation patterns, which in turn affect variation among plants in the amount of pollen exported and deposited on recipient stigmas. In the post-pollination phase, differences among individuals in pollen grain germination success and pollen tube growth may cause realized paternity to differ from patterns of pollen receipt. The maternal plant can also preferentially provision some developing seeds or fruits to further alter variation in siring success. SCOPE: In this review, we describe studies that advance our understanding of the dynamics of the pollination and post-pollination phases, focusing on how variation in male fitness changes in response to pollen limitation. We then explore the interplay between pollination and post-pollination success, and how these processes respond to ecological factors such as pollination intensity. We also identify pressing questions at the intersection of pollination and paternity and describe novel experimental approaches to elucidate the relative importance of pollination and post-pollination factors in determining male reproductive success. CONCLUSIONS: The relative contribution of pollination and post-pollination processes to variation in male reproductive success may not be constant, but rather may vary with pollination intensity. Studies that quantify the effects of pollination and post-pollination phases in concert will be especially valuable as they will enable researchers to more fully understand the ecological conditions influencing male reproductive success.

Effects of pollination and postpollination processes on selfing rate in Mimulus ringens.

PREMISE OF THE STUDY: Selfing rates vary widely within and among populations of self-compatible flowering plants. This variation is often attributed to differences in the amount and timing of self and outcross pollen deposition on stigmas, as well as to the influence of postpollination mechanisms that control fertilization success. This study explores the relative importance of pollination and postpollination processes in determining selfing rates in monkeyflower, Mimulus ringens. METHODS: We hand-pollinated flowers on 17 unrelated mothers with pollen from one of three experimental treatments intended to replicate field conditions: (1) simultaneous deposition of 50% self pollen and 50% outcross pollen from 5 unrelated donors; (2) self pollen followed 15 min later by application of an equal amount of outcross pollen from five unrelated donors; and (3) outcross pollen from 5 unrelated donors followed 15 min later by application of an equal amount of self pollen. We genotyped 757 progeny at 8 polymorphic microsatellite loci and used paternity exclusion to determine whether each seedling was selfed or outcrossed. KEY RESULTS: When self and outcross pollen arrived simultaneously, and when self pollen arrived 15 min prior to outcross pollen, the observed proportions of self and outcross progeny did not deviate from the expected 1:1 ratio. However, when outcross pollen was applied 15 min prior to self pollen, there was a significant excess of outcross progeny. CONCLUSIONS: Selfing rate in Mimulus ringens is influenced by small differences in the timing of pollen arrival, but not by nonrandom postpollination sorting.

Pollen competition in style: Effects of pollen size on siring success in the hermaphroditic common morning glory, Ipomoea purpurea.

PREMISE OF THE STUDY: Pollen size varies greatly among flowering plant species and has been shown to influence the delivery of sperm cells to the eggs. Relatively little is known, however, about the functional significance of within-species genetic variation in pollen size. This study tests whether pollen size influences the relative siring success of a pollen donor during in vivo pollen competition experiments. METHODS: We used two groups of Ipomoea purpurea plants genetically divergent in their pollen sizes and applied equal number of pollen grains from one large-pollen and one small-pollen donor onto the same stigma. Using microsatellite genetic markers, we identified the pollen parent of each of the resulting progeny to determine the relative siring success of the competing donors. Competitions between donors of equal-sized pollen served as a control. KEY RESULTS: Differences in pollen size significantly affected the relative siring success of a pollen donor; larger-grained individuals outcompeted smaller-grained competitors but not equal-sized competitors. Relative siring success, however, sometimes varied across different pollen recipients. CONCLUSIONS: Pollen size can influence the relative siring success of different individuals competing on the same stigma during postpollination processes. However, other factors, such as pollen-pistil interaction and environmental conditions, are likely to influence these competitions as well.