Pollination Accuracy Research Articles

Nectar and floral morphology differ in evolutionary potential in novel pollination environments.

Plants can evolve rapidly after pollinator changes, but the response of different floral traits to novel selection can vary. Floral morphology is often expected to show high integration to maintain pollination accuracy, while nectar traits can be more environmentally sensitive. The relative role of genetic correlations and phenotypic plasticity (PP) in floral evolution remains unclear, particularly for nectar traits, and can be studied in the context of recent pollinator changes. Digitalis purpurea shows rapid recent evolution of corolla morphology but not nectar traits following a range expansion with hummingbirds added as pollinators. We use this species to compare PP, heritability, evolvability and integration of floral morphology and nectar in a common garden. Morphological traits showed higher heritability than nectar traits, and the proximal section of the corolla, which regulates access to nectar and underwent rapid change in introduced populations, presented lower integration than the rest of the floral phenotype. Nectar was more plastic than morphology, driven by highly plastic sugar concentration. Nectar production rate showed high potential to respond to selection. These results explain the differential rapid evolution of floral traits previously observed in this species and show how intrafloral modularity determines variable evolutionary potential in morphological and nectar traits.

Staminal hairs increase pollinator attraction and pollination accuracy in Tradescantia fluminensis (Commelinaceae).

Staminal hairs are the particular appendages of stamens, which may affect pollinator foraging behaviour and pollen transfer. However, experimental evidence of the functions of staminal hairs in pollination remains scarce. Here, we conducted staminal hair manipulation experiments in Tradescantia fluminensis (Commelinaceae) to investigate their effects on visitation and pollen transfer by bees. Our observations revealed that both visitation rates and visit duration of honeybees (Apis cerana) to control flowers were significantly higher than that of hairless flowers. Moreover, removing the staminal hairs significantly decreased pollen deposition by honeybees (A. cerana), but did not affect pollen removal. The staminal hair was similar in length to the stamen and the pistil of T. fluminensis. The staminal hairs provide more footholds for honeybees, and they lay prone on the staminal hairs to collect pollen, which increased the accuracy of pollination through the consistent pollen placement and pick-up on the ventral surface of honeybees. These results showed that the staminal hairs in T. fluminensis may represent an adaptation to attract pollinators and enhance pollination accuracy.

Flower angle favors pollen export efficiency in the snowdrop Galanthus nivalis (Linnaeus, 1753) but not in the lesser celandine Ficaria verna (Huds, 1762)

ABSTRACT Flower angle is crucially important for accurate pollination and flower protection against abiotic factors. Evolutionary factors shaping floral traits are particularly strong for bilaterally symmetric flowers because these flowers require more pollination accuracy than radially symmetrical flowers. We experimentally investigated the flower angle in the snowdrop’s (Galanthus nivalis) radially symmetrical, early-blooming downward flowers. Bumblebees were able to gather significantly more pollen grains from downward flowers than from upward flowers, but female traits (fertility in the field) seem unaffected by flower angle. Similar experiments with radially symmetrical, later flowering Lesser celandine (Ficaria verna) upward flowers showed no differences in bees’ abilities to gather pollen in upward vs downward-facing flowers. The downward angle of snowdrop flowers is an adaptation that increases the ability of insects to collect more pollen grains under unfavorable early spring weather conditions when pollinators are scarce.

Floral recovery from accidents

Pollination Plants in the field layer may be susceptible to damage, with their stems bent or twisted by wind, falling objects, or trampling. For flowers specialized to attract insect pollinators, such accidents can cause changes in orientation that could decrease the chances of successful pollination. In experiments with a range of herbaceous species, Armbruster and Muchhala compared the postinjury responses of bilaterally symmetrical flowers with those of radially symmetrical flowers. Bilateral flowers mostly showed the ability to correct their position though reorientation of the floral stem or the floral sexual organs themselves, resulting in restoration of pollination efficiency. Few radially symmetrical flowers showed the same ability because their orientation had little effect on pollination accuracy. New Phytol. 10.1111/nph.16482 (2020).

Floral reorientation: the restoration of pollination accuracy after accidents.

Plants sometimes suffer mechanical injury. The nonlethal collapse of a flowering stalk, for example, can greatly reduce plant fitness if it leads to 'incorrect' floral orientation and thus reduced visitation or poor pollination. When floral orientation is important for accurate pollination, as has been suggested for bilaterally symmetrical flowers, we predict that such flowers should have developmental and/or behavioural mechanisms for restoring 'correct' orientation after accidents. We made observations and conducted experiments on 23 native and cultivated flowering plant species in Australia, South America, North America and Europe. We found that flowers with bilateral symmetry usually have the capacity to reorient after accidents, and that this is manifested through rapid bending and/or rotation of pedicels or sexual organs or slower peduncle bending. Floral reorientation restores pollination accuracy and fit with pollinators. However, experimental floral misorientation in eight species with radially symmetrical flowers showed that, with one exception, they had little capacity to reorient their flowers, in line with expectations that the orientation of radially symmetrical flowers does not substantially affect pollination accuracy. Our results suggest that quick corrective reorientation of bilaterally symmetrical flowers is adaptive, highlighting a little-studied aspect of plant-pollinator interactions and plant evolution.

Accurate position exchange of stamen and stigma by movement in opposite direction resolves the herkogamy dilemma in a protandrous plant, Ajuga decumbens (Labiatae).

Herkogamy is an effective way to reduce sexual interference. However, the separation of stigma and anther potentially leads to a conflict because the pollen may be placed in a location on the pollinator different from the point of stigma contact, which can reduce pollination accuracy. Floral mechanisms aiming to resolve this conflict have seldom been explored. The floral biology of protandrous Ajuga decumbens was studied to uncover how the herkogamy dilemma can be resolved. Flower anthesis was divided into male, middle, female and wilting phases. The positions of stigma and stamen were dissimilar in different flower development stages. We measured the distance of the stamen and stigma to the lower corolla lip at different floral phases, which was the pollinators’ approaching way. The pollen viability, stigma receptivity, pollen removal and pollen deposition on stigma were investigated at different phases. During the male phase, the dehisced anthers were lower than the stigma, located at the pollinators’ approaching way, and dispersed most pollen with high viability. As the flower developed, the anthers moved upwards, making way for pollen deposition during the female phase. Meanwhile, the stigma becomes receptive by moving into the way and consequently was deposited with sufficient pollen. The position exchange of the stamen and stigma created a dynamic herkogamy at the floral phase with different sexual functions. This floral mechanism effectively avoided sexual interference and maintained pollination accuracy. In Ajuga, the movement herkogamy might be of adaptive significance in response to the changes in the pollination environment.

Multiple Effects of Drought on Pollination and Mating-System Traits inDalechampia scandens

Premise of research. Selfing rates in mixed-mating plant species are often found to fluctuate greatly across time and space. Environmentally induced changes in floral traits may mediate changes in selfing rates through several mechanisms, including direct effects via changes in traits influencing autofertility rates and indirect effects via changes in traits affecting the rate of pollinator visitation and/or the efficiency of cross-pollination. In this study, we tested how experimentally induced drought affected traits related to these three components of plant mating systems.Methodology. We subjected two populations from each of two species in the Dalechampia scandens species complex to a series of experimental drought events in the greenhouse. We measured drought effects on advertisement (signaling) traits, reward traits, herkogamy, dichogamy, autofertility, and pollination accuracy and compared these across populations and species.Pivotal results. Blossom size (advertisement and reward traits) and dich...

The mean and variability of a floral trait have opposing effects on fitness traits.

Floral traits are essential for ensuring successful pollination and reproduction in flowering plants. In particular, style and anther positions are key for pollination accuracy and efficiency. Variation in these traits among individuals has been well studied, but less is known about variation within flowers and plants and its effect on pollination and reproductive success. Style deflexion is responsible for herkogamy and important for pollen deposition in Passiflora incarnata. The degree of deflexion may vary among stigmas within flowers as well as among flowers. We measured the variability of style deflexion at both the flower and the plant level. The fitness consequences of the mean and variation of style deflexion were then evaluated under natural pollination by determining their relationship to pollen deposition, seed production and average seed weight using structural equation modelling. In addition, the relationship between style deflexion and self-pollen deposition was estimated in a greenhouse experiment. We found greater variation in style deflexion within flowers and plants than among plants. Variation of style deflexion at the flower and plant level was positively correlated, suggesting that variability in style deflexion may be a distinct trait in P. incarnata. Lower deflexion and reduced variation in that deflexion increased pollen deposition, which in turn increased seed number. However, lower styles also increased self-pollen deposition. In contrast, higher deflexion and greater variability of that deflexion increased variation in pollen deposition, which resulted in heavier seeds. Variability of style deflexion and therefore stigma placement, independent from the mean, appears to be a property of individual P. incarnata plants. The mean and variability of style deflexion in P. incarnata affected seed number and seed weight in contrasting ways, through the quantity and potentially quality of pollen deposition. This antagonistic selection via different fitness components may maintain diverse style phenotypes.

Floral specialization and angiosperm diversity: phenotypic divergence, fitness trade-offs and realized pollination accuracy.

Plant reproduction by means of flowers has long been thought to promote the success and diversification of angiosperms. It remains unclear, however, how this success has come about. Do flowers, and their capacity to have specialized functions, increase speciation rates or decrease extinction rates? Is floral specialization fundamental or incidental to the diversification? Some studies suggest that the conclusions we draw about the role of flowers in the diversification and increased phenotypic disparity (phenotypic diversity) of angiosperms depends on the system. For orchids, for example, specialized pollination may have increased speciation rates, in part because in most orchids pollen is packed in discrete units so that pollination is precise enough to contribute to reproductive isolation. In most plants, however, granular pollen results in low realized pollination precision, and thus key innovations involving flowers more likely reflect reduced extinction rates combined with opportunities for evolution of greater phenotypic disparity (phenotypic diversity) and occupation of new niches. Understanding the causes and consequences of the evolution of specialized flowers requires knowledge of both the selective regimes and the potential fitness trade-offs in using more than onepollinator functional group. The study of floral function and flowering-plant diversification remains a vibrant evolutionary field. Application of new methods, from measuring natural selection to estimating speciation rates, holds much promise for improving our understanding of the relationship between floral specialization and evolutionary success.

In the right place at the right time: Parnassia resolves the herkogamy dilemma by accurate repositioning of stamens and stigmas

Spatial (herkogamy) and temporal (dichogamy) separation of pollen presentation and stigma receptivity have been interpreted as reducing interference between male and female functions in hermaphroditic flowers. However, spatial separation leads to a potential conflict: reduced pollination accuracy, where pollen may be placed in a location on the pollinator different from the point of stigma contact. To understand better how herkogamous flowers resolve this conflict, a study was made of a subalpine herb, Parnassia epunctulata, the nectariferous flowers of which exhibit sequential anther dehiscence (staggered pollen presentation) and stamen movements; usually one newly dehisced anther is positioned each day over the central gynoecium, while the older stamens bend away from the central position. The open flowers were visited by a variety of pollinators, most of which were flies. Seed set was pollinator-dependent (bagged flowers set almost no seeds) and pollen-limited (manual pollination increased seed set over open pollination). Analyses of adaptive accuracy showed that coordinated stamen movements and style elongation (movement herkogamy) dramatically increased pollination accuracy. Specifically, dehiscing anthers and receptive stigmas were positioned accurately in the vertical and horizontal planes in relation to the opposite sexual structure and pollinator position. By contrast, the spatial correspondence between anthers and stigma was dramatically lower before the anthers dehisced and after stamens bent outwards, as well as before and after the period of stigmatic receptivity. It is shown for the first time that a combination of movement herkogamy and dichogamy can maintain high pollination accuracy in flowers with generalized pollination. Staggered pollen and stigma presentation with spatial correspondence can both reduce sexual interference and improve pollination accuracy.

Do specialized flowers promote reproductive isolation? Realized pollination accuracy of three sympatric Pedicularis species

Interest in pollinator-mediated evolutionary divergence of flower phenotype and speciation in plants has been at the core of plant evolutionary studies since Darwin. Specialized pollination is predicted to lead to reproductive isolation and promote speciation among sympatric species by promoting partitioning of (1) the species of pollinators used, (2) when pollinators are used, or (3) the sites of pollen placement. Here this last mechanism is investigated by observing the pollination accuracy of sympatric Pedicularis species (Orobanchacae). Pollinator behaviour was observed on three species of Pedicularis (P. densispica, P. tricolor and P. dichotoma) in the Hengduan Mountains, south-west China. Using fluorescent powder and dyed pollen, the accuracy was assessed of stigma contact with, and pollen deposition on, pollinating bumble-bees, respectively. All three species of Pedicularis were pollinated by bumble-bees. It was found that the adaptive accuracy of female function was much higher than that of male function in all three flower species. Although peak pollen deposition corresponded to the optimal location on the pollinator (i.e. the site of stigma contact) for each species, substantial amounts of pollen were scattered over much of the bees' bodies. The Pedicularis species studied in the eastern Himalayan region did not conform with Grant's 'Pedicularis Model' of mechanical reproductive isolation. The specialized flowers of this diverse group of plants seem unlikely to have increased the potential for reproductive isolation or influenced rates of speciation. It is suggested instead that the extreme species richness of the Pedicularis clade was generated in other ways and that specialized flowers and substantial pollination accuracy evolved as a response to selection generated by the diversity of co-occurring congeners.

Macroevolutionary patterns of pollination accuracy: a comparison of three genera

We hypothesize that pollination efficiency selects for equal distances between the pollinator reward and the anthers, and the stigmas, creating an adaptive ridge. We predict that this fitness surface governs the divergence of many plant species. We use the theory of adaptive accuracy, precision and mean optimality to assess how close populations lie to the hypothesized adaptive ridge and which factors contribute to departure from the optimum. Patterns of accuracy of pollen placement and receipt were compared across species in three study systems, Dalechampia (Euphorbiaceae), Collinsieae (Plantaginaceae) and Stylidium (Stylidiaceae), in order to assess the roles of stamen/stigma imprecision and population mean departure from the optimum in the generation of floral inaccuracy. We found that population mean departure from the optimum was the most important factor in Dalechampia, female imprecision and departure from the optimum were about equally important factors in Collinsieae, and stamen and stigma imprecision were equally important in Stylidium, with virtually no departure from the optimum. Possible reasons for imprecision and departure from the optimum were assessed using phylogenetically informed methods, indicating important roles of limited floral integration in the generation of imprecision, and conflicting selective pressures, associated with outcrossing, in the generation of departure from the optimum.

Floral isolation betweenAquilegia formosaandAquilegia pubescens

The acquisition of floral nectar spurs is correlated with increased species diversity across multiple clades. We tested whether variation in nectar spurs influences reproductive isolation and, thus, can potentially promote species diversity using two species of Aquilegia, Aquilegia formosa and Aquilegia pubescens, which form narrow hybrid zones. Floral visitors strongly discriminated between the two species both in natural populations and at mixed-species arrays of individual flowers. Bees and hummingbirds visited flowers of A. formosa at a much greater rate than flowers of A. pubescens. Hawkmoths, however, nearly exclusively visited flowers of A. pubescens. We found that altering the orientation of A. pubescens flowers from upright to pendent, like the flowers of A. formosa, reduced hawkmoth visitation by an order of magnitude. In contrast, shortening the length of the nectar spurs of A. pubescens flowers to a length similar to A. formosa flowers did not affect hawkmoth visitation. However, pollen removal was significantly reduced in flowers with shortened nectar spurs. These data indicate that floral traits promote floral isolation between these species and that specific floral traits affect floral isolation via ethological isolation while others affect floral isolation via mechanical isolation.

Zygomorphy and Its Causes. I

Previous articleNext article FreeZygomorphy and Its Causes. ICharles RobertsonCharles Robertson Search for more articles by this author PDFPDF PLUS Add to favoritesDownload CitationTrack CitationsPermissionsReprints Share onFacebookTwitterLinkedInRedditEmail SectionsMoreDetailsFiguresReferencesCited by Volume 13, Number 6Jun., 1888 Article DOIhttps://doi.org/10.1086/326285 Views: 28Total views on this site Citations: 9Citations are reported from Crossref Journal History This article was published in the Botanical Gazette (1876-1991), which is continued by International Journal of Plant Sciences (1992-present). PDF download Crossref reports the following articles citing this article:W. Scott Armbruster, Nathan Muchhala Floral reorientation: the restoration of pollination accuracy after accidents, New Phytologist 227, no.11 (Apr 2020): 232–243.https://doi.org/10.1111/nph.16482Somayeh Naghiloo, Dirk U. Bellstedt, Regine Claßen‐Bockhoff Pollination biology in Roepera (Zygophyllaceae): How flower structure and shape influence foraging activity, Plant Species Biology 35, no.11 (Dec 2019): 72–80.https://doi.org/10.1111/1442-1984.12262Yang Dong, Jing Liu, Peng-Wei Li, Chao-Qun Li, Tian-Feng Lü, Xia Yang, Yin-Zheng Wang, Michael Purugganan Evolution of Darwin’s Peloric Gloxinia (Sinningia speciosa) Is Caused by a Null Mutation in a Pleiotropic TCP Gene, Molecular Biology and Evolution 35, no.88 (Apr 2018): 1901–1915.https://doi.org/10.1093/molbev/msy090J. L. Blanco-Pastor, C. Ornosa, D. Romero, I. M. Liberal, J. M. Gómez, P. Vargas Bees explain floral variation in a recent radiation of Linaria, Journal of Evolutionary Biology 28, no.44 (Mar 2015): 851–863.https://doi.org/10.1111/jeb.12609Charles B. Fenster, W. Scott Armbruster, Michele R. Dudash Specialization of flowers: is floral orientation an overlooked first step?, New Phytologist 183, no.33 (Jul 2009): 502–506.https://doi.org/10.1111/j.1469-8137.2009.02852.xFavio González, Dennis Wm. Stevenson Perianth development and systematics of Aristolochia, Flora 195, no.44 (Dec 2000): 370–391.https://doi.org/10.1016/S0367-2530(17)30995-7 Peter K. Endress Symmetry in Flowers: Diversity and Evolution Endress, International Journal of Plant Sciences 160, no.S6S6 (Jul 2015): S3–S23.https://doi.org/10.1086/314211Heather C. Proctor, Robert L. Baker, Darryl T. Gwynne Mating behaviour and spermatophore morphology: a comparative test of the female-choice hypothesis, Canadian Journal of Zoology 73, no.1111 (Nov 1995): 2010–2020.https://doi.org/10.1139/z95-237David G. Lloyd Progress in understanding the natural history of New Zealand plants, New Zealand Journal of Botany 23, no.44 (Dec 1985): 707–722.https://doi.org/10.1080/0028825X.1985.10434239