Patterns of pollen removal and deposition in Polemonium brandegeei (Polemoniaceae): the role of floral visitors,floral design and sexual interference

The arrangement, colour, shape and size of floral parts (collectively floral design) have evolved primarily to promote mating success via animal‐mediated pollen transfer. Although numerous studies have examined variation in pollinator assemblages, relatively few have examined patterns of pollen removal and deposition in the presence of fluctuating pollinators and ineffective floral visitors; therefore, net pollen removal and deposition by entire visitor assemblages are unclear. We studied the timing (diurnal or nocturnal) and effects of floral traits on pollen removal and deposition under a dynamic visitor assemblage of Polemonium brandegeei. We quantified pollen grains remaining in anthers (pollen removal) and deposited on stigmas (pollen deposition) of plants visited during either the day (07:30–20:00 h) or night (20:30–07:30 h) in natural populations over two flowering seasons. Pollen removal and deposition occurred both diurnally and nocturnally during our study. Increased diurnal removal and deposition coincided with peak floral visitations in 2006. This increase in pollen removal and deposition may reflect increased visits by pollen consumers, effective hawkmoth pollinators and increased self‐pollen deposition due to hot, dry weather. Nonlinear effects of style length significantly affected pollen removal, with less pollen remaining in flowers with intermediate style lengths. Pollen deposition was more complex, with herkogamy and anther height affecting deposition. Further, close proximity of stigmas and anthers increased the potential for sexual interference between pollen removal and deposition. Overall, flower visitations and pollen removal and deposition varied between years and populations, but sex organ placement consistently influenced the removal and deposition of pollen.     

Mechanisms of pollen deposition by insect pollinators

Summary Studies of pollen dispersal in insect-pollinated plants have often documented highly leptokurtic patterns of pollen deposition that can increase the likelihood of long-distance mating. To examine potential causes of highly leptokurtic deposition, we introduce four functions that arise when (1) the duration of pollinator visits to pollen sources is limited, (2) the rate of pollen deposition varies randomly among pollinators and/or among visits, (3) the rate of pollen deposition changes monotonically over time or (4) pollen is carried in layers or compartments on the pollinator's body that differ in deposition rate. Maximum likelihood techniques were used to fit deposition functions to data obtained from honey bees (Apis mellifera L.) visiting mustard plants (Brassica campestris L.) that contained a marker gene. Each of the alternative leptokurtic functions fit the experimental data better than a simple exponential function and the best-fit function predicted a mean pollen dispersal distance more than three times greater than the exponential. We argue that studies of pollen deposition need to test a broader range of deposition models to assess outcrossing distance in plant populations accurately.     

Correlation between aerobiological and phytogeographical investigations in the Florence area

Summary The aim of our research work was to evaluate the results from our pollen count (p.c.) carried out at the Allergological Centre of Nuovo Osp. San Giovanni di Dio in Florence, in the light of orogeographical, phytogeographical and meteorological data. The highest p.c. was yielded by Graminaceae which appear in small quantities early in the year but increase considereably starting from April. Oleaceae too showed high pollen concentration but over a shorter lenght of time. The p.c. for Compositae and Urticaceae was lower. It should be noted that Cupressaceae yielded constantly high p.c. in February–March. Investigations on the allergenic flora of the surroundings were carried out concentrically in order to find out the closest and the farthest sources of pollen production. The species investigated come under 7 types of vegetation: xerophilous woods, mesophilous woods, conifers woods, parks and gardens, rural areas, meadows, border areas-weeds. A phytosociological study of the types of vegetation has made it possible to evaluate the single species quantitatively. The different sources of pollen production can be spotted by mapping these types of vegetation. The seasonal characterization of pollen production is better defined by means of periodical phenological checking of the existing allergenic flora along carefully planned routes through the areas next to the sampling station.      

ANGIOSPERM WOOD FROM THE UPPER CRETACEOUS OF CENTRAL CALIFORNIA: PART I

This paper is the first in a series about a collection of 40 Upper Cretaceous woods from central California. The wood is in the form of fragments of stems and roots embedded in nodules of collophane rich in microorganisms, pollen, and spores. Two new species are described: Ulminium pattersonensis and Ulminium mulleri. Evidence presented indicates that these two specimens are similar to woods of the Lauraceae and fall well within the range of variation of woods of that family. Because the taxonomy of the Lauraceae is in need of revision, meaningful comparisons of fossil plant parts with modern taxa were found to be unfeasible.     

Nonrandom mating in Clarkia gracilis (Onagraceae): a case of cryptic self-incompatibility

In plants capable of both self-fertilization and outcrossing, the selfing rate depends on the proportion of self pollen in pollen loads and on the relative postpollination success of self pollen in siring offspring. While the composition of pollen loads is subject to unpredictable variation, paternity success of self vs. outcross pollen following pollen deposition may be controlled by maternal plants. This study examined postpollination paternity success in Clarkia gracilis ssp. sonomensis, in which deposition of self pollen is common. Pure loads of self and outcross pollen produced similar numbers of mature seeds, but equal mixtures of self and outcross pollen yielded more than three times as many outcrossed offspring as selfed offspring. The finding that the paternity success of self pollen depends on whether it is in competition with outcross pollen helps to explain an earlier finding that the selfing rate in experimental populations was highest when pollinator activity was lowest. Cryptic self-incompatibility allows paternity by self pollen when outcross pollen is unavailable.     

Ragweed and mugwort pollen in Szczecin, Poland

Ragweed (genus Ambrosia) and mugwort (Artemisia vulgaris) pollen grains are known to be very potent aeroallergens, often noted to enter into cross reactions. The aim of the study was to analyse ragweed and mugwort pollen release in Szczecin (western Poland) during the period 2000–2003. Measurements were performed by the volumetric and gravimetric method. Pollen seasons were defined as the periods of 90% of the total catch. Of the 4 years studied, the lowest concentration of ragweed pollen was observed in 2000. In 2000, the annual ragweed pollen count was very high, threefold higher than in 2001. There was a high Ambrosia pollen count in 2003, with the highest daily value of 84 grains/m³. The mugwort pollen season started in the third 10-day period of July and lasted to the end of August in all of the years studied. Analysis of pollen deposition from different Szczecin city’s districts showed that the highest exposure to ragweed pollen allergens occurred in the Majowe district, which is related to the presence of numerous plants of Ambrosia in that district. The mugwort pollen deposition was more abundant in the Żelechowa district, which is an area with villas and gardens. Statistically significant correlations were found between the ragweed pollen count in the air and the maximum wind speed, air temperature and relative humidity and between the mugwort pollen count in the air and air temperature and relative humidity.     

Patterns and sources of variation in pollen deposition and pollen tube formation in flowers of the endemic monoecious shrub Cnidoscolus souzae (Euphorbiaceae)

Pollen deposition and pollen tube formation are key components of angiosperm reproduction but intraspecific variation in these has rarely been quantified. Documenting and partitioning (populations, plants and flowers) natural variation in these two aspects of plant reproduction can help uncover spatial mosaics of reproductive success and underlying causes. In this study, we assess variation in pollen deposition and pollen tube formation for the endemic monoecious shrub Cnidoscolus souzae throughout its distribution range in Mexico, and determine how this variation is structured among populations, plants and flowers. We also infer the relative importance of pollen quantity and quality in determining pollination success in this species. While we found no evidence suggesting that pollen receipt limits C. souzae reproduction across 19 populations, we did find extensive variation in pollen load size and pollen tube number per flower. Total variation in pollen receipt and pollen tube number was mostly explained by intra‐individual and among‐population variance. Furthermore, pollen load size had a stronger effect on the number of pollen tubes at the base of the style than pollen germination rate, suggesting that pollen quantity may be more important than quality for pollen tube success in C. souzae. Our results suggest that both small within‐plant flower differences and broad‐scale differences in community attributes can play an important role in determining pollination success. We emphasise the need to evaluate patterns and sources of variation in pollen deposition and pollen tube formation as a first step in understanding the causes of variation in pollination success over broad spatial scales.     

AERODYNAMICS OF WIND POLLINATION IN SIMMONDSIA CHINENSIS (LINK) SCHNEIDER

Wind tunnel analyses of Simmondsia chinensis (Link) Schneider or “jojoba” were conducted to quantify the behavior of airborne pollen grains around individual branches and leaves and near individual carpellate flowers. Field data (wind velocity) were used to ensure a correspondence between wind tunnel and natural conditions. Based upon the visualization of individual pollen grain trajectories, it is concluded that pollen deposition on stigmatic surfaces is influenced by large-scale aerodynamic patterns, generated by foliage leaves, and small-scale airflow patterns, formed around and by floral parts and stigmas. Leaves are seen to deflect airborne pollen grains into trajectories that can intersect ambient airflow at 90° angles, showering decumbent carpellate flowers with pollen. Similarly, flowers can deflect pollen upward and downwind, toward other flowers. The extent of floral bract and sepal recurvature is shown to influence the extent of pollen deposition by determining the characteristic airflow pattern around stigmas. Available evidence concerning the relatively recent evolutionary transition to anemophily in Simmondsia is interpreted within the context of morphological adaptations and exaptations favoring wind pollination.     

Pollinator preference and pollen viability mediated by flower color synergistically determine seed set in an Alpine annual herb

Gentiana leucomelaena manifests dramatic flower color polymorphism, with both blue‐ and white‐flowered individuals (pollinated by flies and bees) both within a population and on an individual plant. Previous studies of this species have shown that pollinator preference and flower temperature change as a function of flower color throughout the flowering season. However, few if any studies have explored the effects of flower color on both pollen viability (mediated by anther temperature) and pollinator preference on reproductive success (seed set) in a population or on individual plants over the course of the entire flowering season. Based on prior observations, we hypothesized that flower color affects both pollen viability (as a function of anther temperature) and pollen deposition (as a function of pollinator preference) to synergistically determine reproductive success during the peak of the flowering season. This hypothesis was tested by field observations and hand pollination experiments in a Tibetan alpine meadow. Generalized linear model and path analyses showed that pollen viability was determined by flower color, flowering season, and anther temperature. Anther temperature correlated positively with pollen viability during the peak of the early flowering season, but negatively affected pollen viability during the peak of the mid‐ to late flowering season. Pollen deposition was determined by flower color, flowering season (early, or mid‐ to late season), and pollen viability. Pollen viability and pollen deposition were affected by flower color that in turn affected seed set across the peak of the flowering season (i.e., when the greatest number of flowers were being pollinated). Hand pollination experiments showed that pollen viability and pollen deposition directly influenced seed set. These data collectively indicate that the preference of pollinators for flower color and pollen viability changed during the flowering season in a manner that optimizes successful reproduction in G. leucomelaena. This study is one of a few that have simultaneously considered the effects of both pollen viability and pollen deposition on reproductive success in the same population and on individual plants.     

Natural levels of pollination intensity and effects of pollen loads on offspring quality in females of Thalictrum pubescens (Ranunculaceae)

Females of Thalictrum pubescens produce stamens containing sterile pollen. Earlier studies have shown the presence of stamens does not increase seed set through increased visitation by insects. Insects may, however, increase pollen deposition on stigmas and increase pollen competition. This paper examines: 1) pollen deposition levels in natural populations and 2) whether larger pollen loads lead to higher quality offspring. The majority of stigmas in two populations received less than 15 grains, but a small proportion had large loads. The latter may represent the occasional insect visit. These visits may provide the opportunity for pollen tube competition. In the greenhouse, flowers receiving heavy-pollen loads had higher seed set than flowers receiving light-pollen loads. Seeds from the two treatments were the same mass, had similar germination times, and seedlings had the same mass. In conclusion, it does not appear that pollen tube competition can account for the maintenance of stamens in females of T. pubescens.     

Effect of pollination success on floral longevity in the orchid Calypso bulbosa (Orchidaceae)

The lifespan of an individual flower is often affected by pollination success. Species differ regarding whether male function (pollen removal), female function (pollen deposition), or both trigger floral senescence. We studied senescence in the singleflowered, deceptive orchid Calypso bulbosa by manipulating the degree of male and female reproductive success. We found that deposition of any amount of pollen resulted in dramatic changes in shape and color within 4 d, whereas unmanipulated flowers and those that had had pollinia removed remained unchanged for 8-11 d after treatment. Selection may favor the reproductive function that is less easily satisfied as the trigger for senescence, because a flower that senesces after accomplishment of this function is likely to have already succeeded at the more easily satisfied one. Deceptive (i.e., rewardless) flowers are more likely to satisfy male than female function since the latter requires that a pollinator be fooled twice, first to pick up pollen and second to deposit it. A survey of naturally pollinated Calypso showed that male function, pollinium removal, was more likely to occur than female function, deposition (95% vs. 66% of visited flowers); thus floral senescence in Calypso is triggered by achievement of the function less likely to succeed. Studies of senescence triggers in species in which female function is more likely to be achieved than male are necessary to further test this hypothesis.     

14. Lake Sedmo Rilsko (Bulgaria): Lateglacial vegetation history

The length of the pollen deposition period covered by an individual moss sample is a matter of discussion. Here we evaluate (1) how many years of pollen deposition are contained in individual moss samples, (2) which factors (i.e. moss type, geographical location and thickness of sample) affect the pollen content, and (3) whether the time span of the pollen deposited in the moss is related to the pollen type. The pollen deposition period covered by 20 individual moss samples adjacent to pollen traps was estimated using the accumulation method. The investigated moss samples usually contain pollen from a period of two to five years. The length of the time period covered by the moss depends on the thickness of the moss cushion and, therefore, we recommend estimating the time period over which the moss has been growing by the increment method. Compared to pollen traps, mosses tend to accumulate more Poaceae and bisaccate grains (especially Picea) but less Cyperaceae pollen. Betula and pollen tetrads tend to be equally represented in mosses and traps. This study demonstrates that moss samples may contain sufficient pollen to be a reliable modern analogue for Quaternary peat and sediment samples but cannot be used to calculate pollen accumulation rates.     

Mating system and the hybridization between self-compatible Phlox cuspidata and self-incompatible Phlox drummondii

Mating system can impact the frequency of hybridization and therefore the maintenance of species diversity. I evaluate the effects of weak self-incompatibility (SI) in Phlox cuspidata and SI in Phlox drummondii on mating success within species and on hybridization dynamics between species under controlled conditions. The effects of SI on hybridization frequency were assessed by manipulating the relatedness of conspecific pollen and the relative timing of pollen deposition in mixed-donor interspecific pollinations. Selfing as opposed to outcrossing increased hybridization by 16% in P. cuspidata maternal plants and by 48% in P. drummondii maternal plants because self pollen did not compete as well against heterospecific pollen. The relative timing of conspecific versus heterospecific pollen deposition also impacted hybridization. In self-compatible P. cuspidata, the deposition of self pollen 5 h earlier than heterospecific pollen decreased hybridization by 28%. In self-incompatible P. drummondii, a 5 h delay in the deposition of compatible conspecific pollen increased hybridization by 32%. In this hybrid system, early self-pollination can decrease hybridization (but increase inbreeding) by P. cuspidata maternal plants, and SI may increase hybridization by P. drummondii maternal plants.     

Evaluation of pollinator effectiveness based on pollen deposition and seed production in a gynodieocious alpine plant,Cyananthus delavayi

Examining variations in pollinator effectiveness can enhance our understanding of how pollinators and plants interact. Pollen deposition and seed production after a single visit by a pollinator are often used to estimate pollinator effectiveness. However, seed production is not always directly related to pollen deposition because not all pollen grains that are deposited on a stigma are compatible or conspecific. In the field, we tested pollinator effectiveness based on pollen deposition and the resulting seed production after single visits by different pollinator groups in a gynodieocious alpine plant Cyananthus delavayi (Campanulaceae). Our results showed that mean pollen deposition was generally inconsistent with mean seed production when comparisons were performed among different pollinator groups and sexes. In general, the correlations were not significant between pollen deposition and seed production in both perfect and female flowers after single visits by halictid bees, bumble bees, and hoverflies. We suggest seed set of virgin flowers after single visits is a more reliable indicator of pollinator effectiveness than pollen deposition and would be a better indicator of pollinator effectiveness for future studies.     

Palaeoecological studies at the Mesolithic site at Bedburg-Königshov near Cologne, Germany

The Late Glacial and early Holocene palaeovegetation is reconstructed by analysis of pollen and macroscopic plant remains from sediments of the lower river Erft valley, near Cologne, exposed by the Garzweiler opencast lignite mine. The study was carried out in parallel with the archaeological excavation of the Mesolithic site of Bedburg-Königshoven, located within a former meander of the Erft. During the Younger Dryas period, the study region was thinly wooded. Relatively open pine woods existed during the Preboreal, becoming more closed during the Boreal. Later in the Boreal,Corylus and taxa of the mixed oak woodland (Ulmus, Quercus, Tilia andFraxinus) appeared. The Atlantic period was characterised by mixed oak woods with dominantUlmus. At the beginning of the Subboreal,Ulmus declined andTilia became the predominant element of the woods. The Mesolithic settlement existed during the early and middle part of the Preboreal period. At that time, the woodland of the Erft valley was mainly composed ofPinits, Belula and somePopulus. The pollen results do not show any evidence of the activity of Mesolithic people. Local vegetational changes of the filling process of the Erft meander were described and interpreted. The vegetational development of the study region is compared with that of neighbouring regions.     

Analysis of sticky generative cell mutants reveals that suppression of callose deposition in the generative cell is necessary for generative cell internalization and differentiation in Arabidopsis

In flowering plants, double fertilization between male and female gametophytes, which are separated by distance, largely depends on the unique pattern of the male gametophyte (pollen): two non-motile sperm cells suspended within a tube-producing vegetative cell. A morphological screen to elucidate the genetic control governing the strategic patterning of pollen has led to the isolation of a sticky generative cell (sgc) mutant that dehisces abnormal pollen with the generative cell immobilized at the pollen wall. Analyses revealed that the sgc mutation is specifically detrimental to pollen development, causing ectopic callose deposition that impedes the timely internalization and differentiation of the generative cell. We found that the SGC gene encodes the highly conserved domain of unknown function 707 (DUF707) gene that is broadly expressed but is germline specific during pollen development. Additionally, transgenic plants co-expressing fluorescently fused SGC protein and known organelle markers showed that SGC localizes in the endoplasmic reticulum, Golgi apparatus and vacuoles in pollen. A yeast two-hybrid screen with an SGC bait identified a thaumatin-like protein that we named GCTLP1, some homologs of which bind and/or digest β-1,3-glucans, the main constituent of callose. GCTLP1 is expressed in a germline-specific manner and colocalizes with SGC during pollen development, indicating that GCTLP1 is a putative SGC interactor. Collectively, our results show that SGC suppresses callose deposition in the nascent generative cell, thereby allowing the generative cell to fully internalize into the vegetative cell and correctly differentiate as the germline progenitor, with the potential involvement of the GCTLP1 protein, during pollen development in Arabidopsis.     

Does natural deposition of pine pollen affect the ovipositional behavior of Frankliniella occidentalis and Frankliniella fusca?

Tree pollen, especially Pinus spp. (Pinaceae), is shed in large quantities every spring in North America. Pine pollen deposition onto leaves was found to significantly influence the ovipositional behaviors of certain thrips species (Thysanoptera: Thripidae) in peanut and tomato leaf choice and no‐choice tests. Pine pollen (Pinus elliottii Engelm.) increased the oviposition rate 2.9‐fold for Frankliniella occidentalis (Pergande) (western flower thrips) and 1.6‐fold for Frankliniella fusca (Hinds) (tobacco thrips) in choice tests averaged over both plant species. These results support the idea that pollen has a greater impact on F. occidentalis behavior than on F. fusca behavior. The most dramatic increase was in peanut, where F. occidentalis only oviposited on leaves dusted with pollen, suggesting that the addition of pollen stimulated this flower thrips to lay eggs on a poor host‐plant part. The impact of pollen on the rate of oviposition by thrips is important because it is the early‐instar nymphs that acquire tomato spotted wilt virus (TSWV), which these two thrips species vector. In a laboratory bioassay, the addition of pine pollen to TSWV‐infected peanut foliage increased the percentage of infected F. fusca after one generation.     

Pine pollen dehiscence relative to thrips population dynamics

A two‐part study related tree pollen dehiscence and thrips population dynamics in the field, and directly evaluated effects of pine pollen deposition on Frankliniella spp. (Thysanoptera: Thripidae) reproduction on tobacco, Nicotiana tabacum L. (Solanaceae), under laboratory conditions. Ambient pollen and thrips were monitored in Tift County, GA, USA, from early spring to late summer/early fall from 2005 to 2008. Correlation analyses were conducted using weekly means of thrips collected on sticky traps compared to weekly pollen counts from a Burkard air sampler and pollen deposition sheets. There were significant positive correlations between pollen and thrips counts on later dates, suggesting that if a relationship exists between pine pollen dehiscence and thrips population dynamics, it will be delayed. Over all years combined, the first spring time peak in Frankliniella fusca (Hinds) counts on sticky traps occurred at 227 accumulated degree days (dd) after the first peak in pine pollen, approximately the dd required to complete a single F. fusca generation. Two subsequent thrips peaks occurred at approximately one and two F. fusca generations following the first peak. After 10 weeks following the final peak in pollen, there appears to be no more effect of pollen on thrips population dynamics. A tobacco leaf, lightly dusted with Slash pine [Pinus elliottii Engelmann (Pinaceae)] pollen to mimic natural deposition, showed a significant increase in the number of offspring produced per Frankliniella occidentalis (Pergande) and F. fusca female. The total offspring produced increased five‐fold in F. fusca and 22‐fold in F. occidentalis in 15 days on the pollen‐treated leaves over the untreated leaves.     

Does the morphological fit between flowers and pollinators affect pollen deposition? An experimental test in a buzz‐pollinated species with anther dimorphism

Some pollination systems, such as buzz‐pollination, are associated with floral morphologies that require a close physical interaction between floral sexual organs and insect visitors. In these systems, a pollinator's size relative to the flower may be an important feature determining whether the visitor touches both male and female sexual organs and thus transfers pollen between plants efficiently. To date, few studies have addressed whether in fact the “fit” between flower and pollinator influences pollen transfer, particularly among buzz‐pollinated species. Here we use Solanum rostratum, a buzz‐pollinated plant with dimorphic anthers and mirror‐image flowers, to investigate whether the morphological fit between the pollinator's body and floral morphology influences pollen deposition. We hypothesized that when the size of the pollinator matches the separation between the sexual organs in a flower, more pollen should be transferred to the stigma than when the visitor is either too small or too big relative to the flower. To test this hypothesis, we exposed flowers of S. rostratum with varying levels of separation between sexual organs, to bumblebees (Bombus terrestris) of different sizes. We recorded the number of visits received, pollen deposition, and fruit and seed production. We found higher pollen deposition when bees were the same size or bigger than the separation between anther and stigma within a flower. We found a similar, but not statistically significant pattern for fruit set. In contrast, seed set was more likely to occur when the size of the flower exceeded the size of the bee, suggesting that other postpollination processes may be important in translating pollen receipt to seed set. Our results suggest that the fit between flower and pollinator significantly influences pollen deposition in this buzz‐pollinated species. We speculate that in buzz‐pollinated species where floral morphology and pollinators interact closely, variation in the visitor's size may determine whether it acts mainly as a pollinator or as a pollen thief (i.e., removing pollen rewards but contributing little to pollen deposition and fertilization).     

The first approach towards recognising allergy-provoking flora in Belluno and its relationship with allergic phenomena

Summary The town of Belluno has a high number of wild and cultivated plants whose pollen has various degrees of importance from an allergological point of view.In this preliminary study we have taken into consideration the distribution and the flowering period of these species, correlating them with the data on the quantity and type of pollen collected by a sampler placed on the roof of the City Hospital, and also with the results of the allergometric tests (Prick Tests) carried out in the Broncho-Pneumological Department during the first 9 months of 1990.The wild specis of spring-flowering trees which can be found in the woods surrounding the city (Fagaceae, Betulaceae, Corylaceae) and along the gravel banks of the Piave river (Salicaceae) as well as the Graminaceae which are widespread both in the urban area and in the fields outside the city and some cultivated species (Robinia, Tilia, Acer) which flower a little later, are the species which produce most pollen and which are held responsible for allergies. However, only the pollen of the Graminaceae gave rise to cases of mono-sensitivity (16%) (apart from one case of Betulaceae and Corylaceae and one case ofArtemisia as well asDermatophagoides) (9.7%). For the other species, the positive reactions were associated with the positive reactions to the pollen of other species belonging to different families and/or to other factors.