The time required for a bumble bee to visit a flower is affected by the length of the bee's glossa and its body weight, and by the depth of the flower and the volume of nectar it contains. Probing time is comprised of two components: access time and ingestion time. Access time increases linearly with flower depth, but ingestion time varies with flower depth only in flowers deeper than the length of the bee's glossa, due to a decline in the rate of ingestion of nectar. Probing time therefore incr
eases gradually with increasing depth for flowers shallower than the bee's glossa, but beyond that depth it increases much more rapidly. The relation of probing time to flower depth influences the foraging efficiency and choice of flowers by bumble bees.
In a riparian population of Erythronium americanum (Liliaceae) in central New Jersey, experimentally self-pollinated plant produced markedly fewer fruit and fewer seeds per fruit than hand-outcrossed and open pollinated plants, even though differences were not evident between pollen tubes that penetrated stigmas from self or foreign pollen. This weak self-compatibility and a positive relation between the percentage of seeds set by outcrossed plants and the distance between pollen donor and recip
ient plants indicate that this population could be susceptible to inbreeding depression.
Limited resources for seed development apparently constrained maximal seed production, based on low seed set (40.6%) by hand-pollinated plants and positive correlations for these plants between plant size and the number and size of seeds set. In contrast, naturally-pollinated plants set a smaller proportion of their ovules, suggesting that limited pollinator service reduced the quantity of seeds produced in this population. Free-foraging bees usually removed more than half of the available pollen in a single visit, so that individual plants probably have few opportunities to disseminate their pollen.
Even though sexually reproductive ramets produce only a single flower per year, less than a third of variation in floral morphology is associated with variation in plant size. Within the flower, the sizes of some closely associated structures, such as the style and ovary, and the anthers and filaments, vary essentially independently of one another. Production of nectar and pollen, the ultimate attractors of pollinating insects, was positively correlated with flower size.
The diversification of many flowering plant families has been attributed to adaptive radiation of pollination and mating systems accompanying changes in ecology and life history. Reproductive traits in seed plants therefore provide a potentially rich source of diversity for comparative and phylogenetic studies. Here we address three topics in reproductive biology: floral allocation strategies, mating systems and life history, and the origin of complex reproductive syndromes using various compara
tive methods. Results from these studies generally complement and inform those obtained from previous microevolutionary work at the population level. Historical inferences concerning reproductive character evolution in some taxa can be hampered by topological uncertainties in tree reconstruction and a lack of resolution in molecular phylogenies. Future insights into the ecology and evolution of plant reproductive adaptations using comparative approaches will require well resolved phylogenies, particularly at the species level.
The diversification of many flowering plant families has been attributed to adaptive radiation of pollination and mating systems accompanying changes in ecology and life history. Reproductive traits in seed plants therefore provide a potentially rich source of diversity for comparative and phylogenetic studies. Here we address three topics in reproductive biology: floral allocation strategies, mating systems and life history, and the origin of complex reproductive syndromes using various compara
tive methods. Results from these studies generally complement and inform those obtained from previous micro- evolutionary work at the population level. Historical inferences concerning reproductive character evolution in some taxa can be hampered by topological uncertainties in tree reconstruction and a lack of resolution in molecular phylogenics. Future insights into the ecology and evolution of plant reproductive adaptations using comparative approaches will require well resolved phylogenics, particularly at the species level.
В книге: Floral Biology. Studies on Floral Evolution in Animal-Pollinated Plants / Editors: Lloyd D.G., Barrett S.C.H. USA: Chapman & Hall, 1996. 410 pp. AID: 3323343442. Book P. 140–190. AID: 9009225588. Часть книги
Inflorescence architecture and floral morphology vary extensively within the Poaceae, but the functional significance of this variation remains largely unknown. As grasses are wind-pollinated, their inflorescence diversity probably reflects alternate solutions to manipulating airstreams to enhance pollen export and import. We tested this hypothesis with two field experiments that contrasted pollen removal and receipt by compact and diffuse inflorescences.
In the ‘aggregation’ experiment, we tied
together panicle branches of two species with diffuse inflorescences, creating more compact inflorescences. Aggregation reduced pollen removal from both species, probably by increasing boundary-layer thickness. The effects of inflorescence aggregation differed between the two species in a manner that is consistent with pollen-size differences, which could affect the ability of pollen grains to pass through the thickened boundary layer around stigmas.
The ‘staking’ experiment constrained inflorescence motion and revealed that culm characteristics contribute to the interaction between grass inflorescences and airstreams. In particular, inflorescence oscillation principally serves pollen removal for species with compact inflorescences, but is of primary importance in pollen receipt for species with diffuse architectures.
These results suggest that inflorescence architecture interacts with wind in a complex manner to facilitate pollination and supports the hypothesis that the extensive diversity of inflorescence architecture within the Poaceae represents evolutionary solutions to the physical constraints of wind pollination.
The aerodynamics of wind pollination selects for an intimate relation between form and function in anemophilous plants. Inflorescence architecture and floral morphology vary extensively within the Poaceae, but the functional implication of this variation remains largely unknown. Here we quantify associations between floret, culm, and inflorescence characteristics for 25 grass species in Kananaskis, Alberta, Canada, and consider whether different architectures may implement unique mechanisms to a
id pollination. The species cluster into four categories defined by all combinations of floret size (small vs. large) and inflorescence architecture (compact vs. diffuse). Species differed significantly for all 18 traits that we measured, with 12 traits differing only between floret‐size classes, three differing only between inflorescence types, and three differing among both (independently or by an interaction). Based on these morphological associations, we discuss the aerodynamic and functional consequences of each category for wind pollination. The independence of inflorescence and floral traits has probably allowed exploration of all possible combinations of inflorescence architecture and floret size during the evolution of the Poaceae.
Although not 'a professed botanist', Charles Darwin made seminal contributions to understanding of floral and inflorescence function while seeking evidence of adaptation by natural selection. This review considers the legacy of Darwin's ideas from three perspectives. First, we examine the process of floral and inflorescence adaptation by surveying studies of phenotypic selection, heritability and selection responses. Despite widespread phenotypic and genetic capacity for natural selection, only
one-third of estimates indicate phenotypic selection. Second, we evaluate experimental studies of floral and inflorescence function and find that they usually demonstrate that reproductive traits represent adaptations. Finally, we consider the role of adaptation in floral diversification. Despite different diversification modes (coevolution, divergent use of the same pollen vector, pollinator shifts), evidence of pollination ecotypes and phylogenetic patterns suggests that adaptation commonly contributes to floral diversity. Thus, this review reveals a contrast between the inconsistent occurrence of phenotypic selection and convincing experimental and comparative evidence that floral traits are adaptations. Rather than rejecting Darwin's hypotheses about floral evolution, this contrast suggests that the tempo of creative selection varies, with strong, consistent selection during episodes of diversification, but relatively weak and inconsistent selection during longer, 'normal' periods of relative phenotypic stasis.
Pollen dispersal by animals varies extensively because of differences in pollinator visitation rates among plants, dissimilar pollination by the various pollinators that visit individual plants, and stochastic variation in deposition as an individual pollinator disperses a plant’s pollen to subsequently visited recipient flowers. Such variation reduces expected female and male success if seed production decelerates with increasing pollen receipt, because less than average receipt diminishes mean
seed production more than copious pollination increases it (Jensen’s inequality). We report empirical studies of the nature and magnitude of pollen dispersal variance, which provide the basis for a numerical model of the consequences of dispersal for expected seed production. Model fitting revealed that dispersal of Brassica napus pollen by bumblebees and especially butterflies exhibited much more variation than is expected of a binomial process and was best modeled as a beta‐binomial process with a constant mean. Overdispersion arose primarily during pollen dispersal by individual insects, since differences between individuals of the same pollinator type were limited. Our model revealed variance limitation as a previously unrecognized, substantial, and ubiquitous component of pollen limitation of seed production. Variance limitation should select for floral traits that increase pollinator visitation, reduce dispersal variance, or reduce the postpollination nonlinearities that cause Jensen’s inequality.
Background and AimsPollen-collecting bees are among the most important pollinators globally, but are also the most common pollen thieves and can significantly reduce plant reproduction. The pollination efficiency of pollen collectors depends on the frequency of their visits to female(-phase) flowers, contact with stigmas and deposition of pollen of sufficient quantity and quality to fertilize ovules. Here we investigate the relative importance of these components, and the hypothesis that floral
and inflorescence characteristics mediate the pollination role of pollen collection by bees.MethodsFor ten Aloe species that differ extensively in floral and inflorescence traits, we experimentally excluded potential bird pollinators to quantify the contributions of insect visitors to pollen removal, pollen deposition and seed production. We measured corolla width and depth to determine nectar accessibility, and the phenology of anther dehiscence and stigma receptivity to quantify herkogamy and dichogamy. Further, we compiled all published bird-exclusion studies of aloes, and compared insect pollination success with floral morphology.Key ResultsSpecies varied from exclusively insect pollinated, to exclusively bird pollinated but subject to extensive pollen theft by insects. Nectar inaccessibility and strong dichogamy inhibited pollination by pollen-collecting bees by discouraging visits to female-phase (i.e. pollenless) flowers. For species with large inflorescences of pollen-rich flowers, pollen collectors successfully deposited pollen, but of such low quality (probably self-pollen) that they made almost no contribution to seed set. Indeed, considering all published bird-exclusion studies (17 species in total), insect pollination efficiency varied significantly with floral shape.ConclusionsSpecies-specific floral and inflorescence characteristics, especially nectar accessibility and dichogamy, control the efficiency of pollen-collecting bees as pollinators of aloes.
Background. The production of flowers, fruits and seeds demands considerable energy and nutrients, which can limit the allocation of these resources to other plant functions and, thereby, influence survival and future reproduction. The magnitude of the physiological costs of reproduction depends on both the factors limiting seed production (pollen, ovules or resources) and the capacity of plants to compensate for high resource demand.
Methods. To assess the magnitude and consequences of reproduc
tive costs, we used shading and defoliation to reduce photosynthate production by fully pollinated plants of a perennial legume, Oxytropis sericea (Fabaceae), and examined the resulting impact on photosynthate allocation, and nectar, fruit and seed production.
Key Results. Although these leaf manipulations reduced photosynthesis and nectar production, they did not alter photosynthate allocation, as revealed by 13C tracing, or fruit or seed production. That photosynthate allocation to reproductive organs increased >190 % and taproot mass declined by 29 % between flowering and fruiting indicates that reproduction was physiologically costly.
Conclusions. The insensitivity of fruit and seed production to leaf manipulation is consistent with either compensatory mobilization of stored resources or ovule limitation. Seed production differed considerably between the two years of the study in association with contrasting precipitation prior to flowering, perhaps reflecting contrasting limits on reproductive performance.
BackgroundMost angiosperms present flowers in inflorescences, which play roles in reproduction, primarily related to pollination, beyond those served by individual flowers alone. An inflorescence's overall reproductive contribution depends primarily on the three-dimensional arrangement of the floral canopy and its dynamics during its flowering period. These features depend in turn on characteristics of the underlying branching structure (scaffold) that supports and supplies water and nutrients t
o the floral canopy. This scaffold is produced by developmental algorithms that are genetically specified and hormonally mediated. Thus, the extensive inflorescence diversity evident among angiosperms evolves through changes in the developmental programmes that specify scaffold characteristics, which in turn modify canopy features that promote reproductive performance in a particular pollination and mating environment. Nevertheless, developmental and ecological aspects of inflorescences have typically been studied independently, limiting comprehensive understanding of the relations between inflorescence form, reproductive function, and evolution.ScopeThis review fosters an integrated perspective on inflorescences by summarizing aspects of their development and pollination function that enable and guide inflorescence evolution and diversification.ConclusionsThe architecture of flowering inflorescences comprises three related components: topology (branching patterns, flower number), geometry (phyllotaxis, internode and pedicel lengths, three-dimensional flower arrangement) and phenology (flower opening rate and longevity, dichogamy). Genetic and developmental evidence reveals that these components are largely subject to quantitative control. Consequently, inflorescence evolution proceeds along a multidimensional continuum. Nevertheless, some combinations of topology, geometry and phenology are represented more commonly than others, because they serve reproductive function particularly effectively. For wind-pollinated species, these combinations often represent compromise solutions to the conflicting physical influences on pollen removal, transport and deposition. For animal-pollinated species, dominant selective influences include the conflicting benefits of large displays for attracting pollinators and of small displays for limiting among-flower self-pollination. The variety of architectural components that comprise inflorescences enable diverse resolutions of these conflicts.
Background and AimsThe Orchidaceae have a history of recurring convergent evolution in floral function as nectar production has evolved repeatedly from an ancestral nectarless state. However, orchids exhibit considerable diversity in nectary type, position and morphology, indicating that this convergence arose from alternative adaptive solutions. Using the genus Disa, this study asks whether repeated evolution of floral nectaries involved recapitulation of the same nectary type or diversifying i
nnovation. Epidermis morphology of closely related nectar-producing and nectarless species is also compared in order to identify histological changes that accompanied the gain or loss of nectar production.MethodsThe micromorphology of nectaries and positionally equivalent tissues in nectarless species was examined with light and scanning electron microscopy. This information was subjected to phylogenetic analyses to reconstruct nectary evolution and compare characteristics of nectar-producing and nectarless species.Key ResultsTwo nectary types evolved in Disa. Nectar exudation by modified stomata in floral spurs evolved twice, whereas exudation by a secretory epidermis evolved six times in different perianth segments. The spur epidermis of nectarless species exhibited considerable micromorphological variation, including strongly textured surfaces and non-secreting stomata in some species. Epidermis morphology of nectar-producing species did not differ consistently from that of rewardless species at the magnifications used in this study, suggesting that transitions from rewardlessness to nectar production are not necessarily accompanied by visible morphological changes but only require sub-cellular modification.ConclusionsIndependent nectary evolution in Disa involved both repeated recapitulation of secretory epidermis, which is present in the sister genus Brownleea, and innovation of stomatal nectaries. These contrasting nectary types and positional diversity within types imply weak genetic, developmental or physiological constraints in ancestral, nectarless Disa. Such functional convergence generated by morphologically diverse solutions probably also underlies the extensive diversity of nectary types and positions in the Orchidaceae.
Background and AimsLarge floral displays have opposing consequences for animal-pollinated angiosperms: they attract more pollinators but also enable elevated among-flower self-pollination (geitonogamy). The presence of sterile flowers as pollinator signals may enhance attraction while allowing displays of fewer open fertile flowers, limiting geitonogamy. The simultaneous contributions of fertile and non-fertile display components to pollinator attraction and reproductive output remain undetermin
ed.MethodsThe simultaneous effects of the presence of sterile flowers and fertile-flower display size in two populations of Leopoldia comosa were experimentally assessed. Pollinator behaviour, pollen removal and deposition, and fruit and seed production were compared between intact plants and plants with sterile flowers removed.Key ResultsThe presence of sterile flowers almost tripled pollinator attraction, supplementing the positive effect of the number of fertile flowers on the number of bees approaching inflorescences. Although attracted bees visited more flowers on larger inflorescences, the number visited did not additionally depend on the presence of sterile flowers. The presence of sterile flowers improved all aspects of plant performance, the magnitude of plant benefit being context dependent. During weather favourable to pollinators, the presence of sterile flowers increased pollen deposition on stigmas of young flowers, but this difference was not evident in older flowers, probably because of autonomous self-pollination in poorly visited flowers. Total pollen receipt per stigma decreased with increasing fertile display size. In the population with more pollinators, the presence of sterile flowers increased fruit number but not seed set or mass, whereas in the other population sterile flowers enhanced seeds per fruit, but not fruit production. These contrasts are consistent with dissimilar cross-pollination and autonomous self-pollination, coupled with the strong predispersal inbreeding depression exhibited by L. comosa populations.ConclusionsSterile flowers enrich pollination quality by promoting pollen export and import, while limiting the mating costs of geitonogamy associated with large fertile displays.
Background and Aims Plant species differ greatly in the three-dimensional arrangements of their flowers (inflorescence architecture). However, the nature of selection responsible for this diversity is poorly understood. Studies that examine among-species variation suggest that inflorescence architecture affects pollinator behaviour, and so should influence plant mating. However, few studies consider the consequences of within-population architectural variation for pollinator behaviour.Methods We
manipulated inflorescence architecture of Delphinium glaucum to contrast bumblebee responses to normal and one-sided (secund) inflorescences.Key Results The ‘dimensionality’ of manipulated inflorescences did not affect the number of flowers that bees visited; however, bees moved upward proportionally more on secund inflorescences.Conclusions This study shows that realistic within-population variation in inflorescence architecture can manipulate pollinator behaviour. These results bear important consequences for plant mating success and the coordinated evolution of inflorescence architecture and floral specialization within inflorescences. These results also question why secund inflorescences are rare, for which we propose four testable explanations.