Reproductive ecology and pollen representation among neotropical trees

Three years of pollen trapping data from Barro Colorado Island, Panama, are compared with local vegetation inventories. Two hypotheses relating pollen representation to ‘messy’ pollination and flower form are tested. Dioecious taxa were found to be over‐represented in pollen spectra compared with their occurrence in local forests. Similarly, anemophilous and ‘messy’ pollination types were found to be over‐represented. While anemophilous taxa were the best dispersed pollen types, zoophilous taxa were also well‐represented in dispersal classes of 20–40 m and > 40 m. Thus pollen arriving to lake sediments is most likely to be from anemophilous species or those zoophilous species exhibiting ‘messy’ pollination syndromes. Pollination mechanisms will predictably bias the fossil record against certain flower morphologies. These data are of significance to those using the fossil pollen record to reconstruct the timing and sequence of angiosperm evolution. These data help prioritize plants to be included in modern pollen reference collections and to focus the search for ‘unknown’ types on most‐likely candidate families.     

花粉形态与风媒植物母体地理分布间的关系

本文对部分风媒植物花粉形态特征,及其母体植物地理分布情况进行了对照研究。在此基础上发现,同一分类系统内,小个体花粉者,其母体植物地理分布趋向于较优越环境。对于具气囊花粉而言,具较大气囊者,或者花粉本体较小者,其母体植物趋向于分布在低纬度或较低海拔环境。并且,产大个体或球状花粉者,往往具有较强的单种成林倾向。本文首次指出了花粉形态与母体植物地理分布间存在着一定的统计学关系,并首次将动态概念引入植物生态学领域,认为花粉形态特征作为一项重要的参变量,通过种间生存竞争过程对母体植物适应倾向施加影响,进而影响物种的生活习性。那些在花粉的传播及授粉过程中稍占优势的物种,其自然分布域趋向于比较优越的环境。本文仅作为一种尝试,试图为动态植物生态学研究,乃至植物生态演化学研究提供一个新的研究思路。     

OPTIMUM POLLEN AND FEMALE RECEPTOR SIZE FOR ANEMOPHILY

A set of mathematical models is developed to describe the relationship between the sizes of pollen grains and female receptors which result in maximum anemophilous pollen transfer. The models predict anemophilous plants should have Stokes numbers in the range of 1.0 to 2.72. The Stokes number is a function of both pollen size and effective receptive surface size. The models were tested with data gathered for 28 assumed anemophilous species and 20 known zoophilous species. The models identify anemophily with an accuracy of approximately 75%. If pollen size alone is used to determine anemophily, then accuracy drops to only 50%. The Stokes number criterion reported here could help in the determination of anemophily of extant and fossil plant taxa.     

The influence of climate changes in Platanus spp. pollination in Spain and Italy

There is evidence of a significant increase in air temperature in the northern hemisphere over recent decades, with consequent changes for anemophilous pollen. In this work we present the effects of climatic change on Platanus spp. pollination in different areas of Italy and Spain, characterized by different climates. In particular, the historical series of pollen monitoring and meteorological data of two Italian stations, Perugia (1982 – 2003) and Torino (1985 – 2003), and two Spanish stations, Santiago de Compostela (1992 – 2003) and Vigo (1994 – 2003), were analysed. The changes recorded in all stations included the timing and behaviour of pollen release. However, no or minimal influence on the total pollen emission was found. Research has linked the changes in phenological events to an increase in temperature, moreover in this study temperature changes are believed to be mainly responsible for the variations recorded in the pollen season of Platanus. A previous start of pollination (?0.66; ?1.21; days/year) is reported in both Italian stations where the temperatures have significantly increased and a delay of 0.2 – 0.8 days/year in Spanish stations where a different trend of temperature is recorded. Other important data is given regarding the type of discharge of pollen grains during the pollen season. Pollination curves are examined by two statistical shape parameters (kurtosis and skewness) which show that pollen release is more gradual with higher temperatures or faster under colder conditions. A regression analysis links the atmospheric pollen presence to mean air temperature.     

Airborne behaviour of <Emphasis Type="Italic">Echium</Emphasis> pollen

Species of Echium are clearly entomophilous, but they release great amounts of pollen into the atmosphere with its consequent anemophilous transport, because their high pollen production, the smallness of the grains, and the exserted position of their anthers. Using three volumetric airborne sporetraps in Extremadura (SW of Spain) between 1994 and 1998, we found that Echium pollen reached pollen concentrations similar to or greater than other anemophilous plants. The main pollination period appeared from April to June. The maximum peak daily concentration reached 35.9 grains/m³ and the annual recorded totals showed interannual variations between 64.2 and 614.4. Correlations were calculated between the daily pollen concentrations and the meteorological parameters rain, temperature, wind direction and velocity and relative humidity. Wind direction seems to be significant, warm dry air seems to facilitate the release of pollen into the atmosphere and increase its concentrations. Hourly pollen concentration reached a maximum between 11:00 and 12:00 and a minimum at 07:00, and the patterns were very similar in the three localities studied. This would indicate that the presence of Echium pollen in the atmosphere is related to the processes of anthesis of the populations near the traps, and would not correspond to a model of transport from distant zones.     

Abiotic pollen and pollination: Ecological, functional, and evolutionary perspectives

The transport and capture of pollen in ~20% of all angiosperm families occurs in air and water. In other words, pollination is abiotic and occurs via the fluid media, not an animal vector. Whereas some early concepts considered abiotic pollination to be largely a stochastic phenomenon, there is sufficient evidence to indicate that wind pollination (i.e. anemophily) and water pollination (i.e. hydrophily) have deterministic features and are sophisticated fluid dynamic solutions to the problem of pollen release, dispersal, and capture.An abiotic pollination syndrome is defined in which there is spatial or temporal separation of carpellate and staminate flowers, which are drab, a reduction in perianth parts, stigmas and anthers are exposed to the fluid, and typically unclumped pollen may be produced in large amounts. Separate pollination syndromes are defined for anemophilous (i.e. wind-pollinated), ephydrophilous (i.e. surface-pollinated), and hydrophilous (i.e. submarine-pollinated) plants. Distinctions are based on habitat and physical conditions for pollination, pollen size, shape, and ultrastructure, morphology and ultrastructure of stigmas, and outcrossing rates. For example, anemophilous pollen are spherical and small, ephydrophilous pollen are spherical or reniform and large, while hydrophilous pollen are filiform (i.e. filamentous) or functionally filiform. The pollination mechanisms and mechanics associated with these syndromes reveals a strong evolutionary relationship between plant morphology and fluid dynamics.     

STARCH IN ANGIOSPERM POLLEN GRAINS AND ITS EVOLUTIONARY SIGNIFICANCE

Pollen grains of 990 species were examined and measured to test four predictions: (1) “Primitive” angiosperms will have starch-containing pollen; more advanced families will have starchless pollen. (2) Where Hymenoptera and Diptera use pollen nutritionally, there will be selection of starchless (oilcontaining) pollen, particularly where pollen is the only reward for visitors. (3) Conversely, in autogamous species, anemophilous species, and those pollinated by Lepidoptera or birds, the energetically more economical accumulation of starch will be seen. (4) Small pollen grains will be more likely to be starchless (oil-rich); starch-containing grains will tend to be larger. Prediction 1 is only partially supported by the results; contemporary representatives of primitive families often show starchless grains, and so do most “advanced” families. Strong intra-familial resemblances are found (and some between groups of families), but other families show mixtures of “starchy” and “starchless” species. The latter are more likely to show a fit between food reserve and pollination system. Prediction 2 is well supported. Prediction 3 is supported in that autogamous and anemophilous species show relatively high proportions of starchy grains, but the picture is complicated for them by apparent selection for starchlessness (oil-richness) in small pollen grains (prediction 4), and for the lepidopteran- and bird-pollinated species by selection for large starchless (or very large starchy grains) in species whose pollen tubes must traverse long styles.     

Live pine pollen in rainwater: reconstructing its long-range transport

Raindrops brim with pollen even when there is no ambient local pollen. How does this nonlocal pollen get inside rain? The likely answer is long-range transport beneath or inside clouds. To test this hypothesis, we captured rain-delivered pollen on Ocracoke Island, NC, USA over a 12-day interval before local pine pollen release then reconstructed its trajectory and its atmospheric exposure conditions. Findings were as follows: four rain episodes yielded a total of 632 pollen grains of which 6.7% germinated. To find pollen sources, we first identified pollen-releasing forested areas using a predictive heat sum equation for each rain episode. Next, we constructed the backward trajectory for air parcels carrying rain-delivered pollen from those forests using the MLDP atmospheric transport and dispersion model. Nonlocal sources were located at distances up to 300 km from Ocracoke Island and distances lessened with each successive episode. Below-cloud transport time was 8 and 17 h for Episodes A and B, respectively. Pollen grains were exposed to harsh atmospheric conditions during below-cloud transport, yet some grains still germinated. Atmospheric turbulence patterns changed for each episode, so distance from pollen source was poorly correlated with pollen transport time. Pollen germination was not closely correlated with either distances or transport time. In-cloud transport was more likely for pollen sampled during Episodes C and D. Pine pollen, although rarely allergenic, brings fresh insights into how atmospheric events can trigger human respiratory distress.

     

The development of composite dispersal functions for estimating absolute pollen productivity in the Swiss Alps

Considering the complexity of real-world pollen dispersal, a single set of parameters may be inadequate to model pollen dispersal, especially as dispersal occurs on both local and regional scales. Here we combine more than one dispersal function into a composite dispersal function (CDF). The function incorporates multiple parameters and different modes of pollen transportation, and thus has the potential to better simulate the relationship between deposited pollen and the surrounding vegetation than would otherwise be possible. CDFs based on different dispersal functions and combinations of dispersal functions were evaluated using a pollen-trap dataset from the Swiss Alps. Absolute pollen productivity (APP) was estimated at 7,700 ± 2,000 grains cm⁻² year⁻¹ for Larix decidua, 13,500 ± 1,900 grains cm⁻² year⁻¹ for Picea abies and 95,600 ± 17,700 grains cm⁻² year⁻¹ for Pinus cembra (with 95% confidence level). The results are consistent with previous APP estimates made from the same dataset using different methods.     

Poaceae pollen grain size as a tool to distinguish past grasslands in South America: a new methodological approach

Despite the dominance of grasslands during the last glacial period, especially in South America, the highly uniform morphology of Poaceae pollen grains has so far allowed only very few palynological studies based on Poaceae pollen. In our study we compare two methods of distinguishing between South American grassland ecosystems based on quantitative morphology of Poaceae pollen grains. We investigated data sets from Páramo in southern Ecuador, Campos de Altitude and Campos in south-eastern and southern Brazil as well as data sets from the Pampa in Argentina by measuring the pollen grain length, grain width, pore diameter and annulus width. Firstly we investigated the potential influence of chemical treatment of pollen grains on pollen grain size as well as the measurement setting for defining the boundary conditions for using Poaceae pollen grains in a palaeoecological investigation. Finally the measured pollen grain parameters were analyzed by comparison of average grain length using statistical tests. This approach reveals highly significant differences in average grain size between all grassland ecosystems. Assuming that a certain grain size range can be assigned to a certain Poaceae taxon, conclusions about differences and similarities in taxa composition can be derived. We used two methods of multivariate data analysis. One uses the pollen grain parameters directly for a Principle Component Analysis (PCA). The other is an already established method in grassland ecology which defines parameter based pollen grain types to investigate similarities between grassland ecosystems. Both approaches confirm the results of the grain length analysis. In this work we demonstrate that the method we developed has the potential to provide acquisition of so far inaccessible information on spatial and temporal patterns and dynamics of South American grasslands.     

Some New Sporomorphs from the Upper Tertiary of Nigeria

A simple model for the immission of pollen grains from large area sources into the atmospheric convective boundary layer is presented.

The terrain is assumed to be homogeneous (like a large plain) and the sources are assumed to be uniform over horizontal distances of the order of 10 km.

The model requires the production rate of pollen to be specified, as well as the temperature as a function of time or analogous atmospheric parameters.

The results compare well with data measured in the Po Valley, Italy.     

Pollen dispersal by <Emphasis Type="Italic">Artemisia tridentata</Emphasis> (Asteraceae)

While the biophysics of anemophilous pollen dispersal is understood in principle, empirical studies for testing such principles are rare, particularly in native ecosystems. This paper describes mechanisms underlying the dispersal of Artemisia pollen in a Wyoming sagebrush steppe. The relationships between meteorological variables and pollen flux were defined during the 1999 Artemisia flowering season, and detailed processes at the individual plant level were experimentally tested in the field in 2000. Results indicated that Artemisia pollen presentation is continuous but with early morning maxima. Atmospheric pollen concentrations and potential dispersal rates are controlled at diurnal time scales by individual flower development together with characteristic changes in temperature/humidity and wind speeds, at multi-day scales by frontal weather patterns, and at week-long scales by flowering phenology.     

Atmospheric conditions during high ragweed pollen concentrations in Zagreb,Croatia

We examined the atmospheric conditions favourable to the occurrence of maximum concentrations of ragweed pollen with an extremely high risk of producing allergy. Over the 2002–2009 period, daily pollen data collected in Zagreb were used to identify two periods of high pollen concentration (> 600 grains/m³) for our analysis: period A (3–4 September 2002) and period B (6–7 September 2003). Synoptic conditions in both periods were very similar: Croatia was under the influence of a lower sector high pressure system moving slowly eastward over Eastern Europe. During the 2002–2009 period, this type of weather pattern (on ~ 70% of days), in conjunction with almost non-gradient surface pressure conditions in the area (on ~ 30% of days) characterised days when the daily pollen concentrations were higher than 400 grains/m³. Numerical experiments using a mesoscale model at fine resolution showed successful multi-day simulations reproducing the local topographic influence on wind flow and in reasonable agreement with available observations. According to the model, the relatively weak synoptic flow (predominantly from the eastern direction) allowed local thermal circulations to develop over Zagreb during both high pollen episodes. Two-hour pollen concentrations and 48-h back-trajectories indicated that regional-range transport of pollen grains from the central Pannonian Plain was the cause of the high pollen concentrations during period A. During period B, the north-westward regional-range transport in Zagreb was supplemented significantly by pronounced horizontal recirculation of pollen grains. This recirculation happened within the diurnal local circulation over the city, causing a late-evening increase in pollen concentration.     

Characteristics of Poaceae pollen grains as a tool to assess palaeoecological grassland dynamics in South America

Despite the dominance of grasslands during the last glacial period, especially in South America, the highly uniform morphology of Poaceae pollen grains has so far allowed only very few palynological studies based on Poaceae pollen. In our study we compare two methods of distinguishing between South American grassland ecosystems based on quantitative morphology of Poaceae pollen grains. We investigated data sets from Páramo in southern Ecuador, Campos de Altitude and Campos in south-eastern and southern Brazil as well as data sets from the Pampa in Argentina by measuring the pollen grain length, grain width, pore diameter and annulus width. Firstly we investigated the potential influence of chemical treatment of pollen grains on pollen grain size as well as the measurement setting for defining the boundary conditions for using Poaceae pollen grains in a palaeoecological investigation. Finally the measured pollen grain parameters were analyzed by comparison of average grain length using statistical tests. This approach reveals highly significant differences in average grain size between all grassland ecosystems. Assuming that a certain grain size range can be assigned to a certain Poaceae taxon, conclusions about differences and similarities in taxa composition can be derived. We used two methods of multivariate data analysis. One uses the pollen grain parameters directly for a Principle Component Analysis (PCA). The other is an already established method in grassland ecology which defines parameter based pollen grain types to investigate similarities between grassland ecosystems. Both approaches confirm the results of the grain length analysis. In this work we demonstrate that the method we developed has the potential to provide acquisition of so far inaccessible information on spatial and temporal patterns and dynamics of South American grasslands.     

On the causes of variability in amounts of airborne grass pollen in Melbourne,Australia

In Melbourne, Australia, airborne grass pollen is the predominant cause of hay fever (seasonal rhinitis) during late spring and early summer, with levels of airborne grass pollen also influencing hospital admissions for asthma. In order to improve predictions of conditions that are potentially hazardous to susceptible individuals, we have sought to better understand the causes of diurnal, intra-seasonal and inter-seasonal variability of atmospheric grass pollen concentrations (APC) by analysing grass pollen count data for Melbourne for 16 grass pollen seasons from 1991 to 2008 (except 1994 and 1995). Some of notable features identified in this analysis were that on days when either extreme (>100 pollen grains m⁻³) or high (50–100 pollen grains m⁻³) levels of grass pollen were recorded the winds were of continental origin. In contrast, on days with a low (<20 pollen grains m^-3) concentration of grass pollen, winds were of maritime origin. On extreme and high grass pollen days, a peak in APC occurred on average around 1730 hours, probably due to a reduction in surface boundary layer turbulence. The sum of daily APC for each grass pollen season was highly correlated (r = 0.79) with spring rainfall in Melbourne for that year, with about 60% of a declining linear trend across the study period being attributable to a reduction of meat cattle and sheep (and hence grazing land) in rural areas around Melbourne. Finally, all of the ten extreme pollen events (3 days or more with APC > 100 pollen grains m⁻³) during the study period were characterised by an average downward vertical wind anomaly in the surface boundary layer over Melbourne. Together these findings form a basis for a fine resolution atmospheric general circulation model for grass pollen in Melbourne’s air that can be used to predict daily (and hourly) APC. This information will be useful to those sectors of Melbourne’s population that suffer from allergic problems.     

Entwicklungsgeschichte und Ultrastruktur von Pollenkitt und Exine bei nahe verwandten entomophilen und anemophilen Sippen derOleaceae,Scrophulariaceae, Plantaginaceae undAsteraceae

Different members of the Angiosperm familiesOleaceae, Scrophulariaceae, Plantaginaceae andAsteraceae with variable modes of pollination were studied in relation to ultrastructure of pollenkitt and exine. In entomophilous representatives the mostly electron-dense and homogeneous pollenkitt is located on the outside layer of the exine, making the pollen sticky. In related anemophilous taxa the pollenkitt mostly is electrontransparent and inhomogeneous; it becomes inactive by remaining in the loculus or sinking to the bottom of the exine caves; this makes the pollen powdery. According to such criteria,Fraxinus ornus andRhinanthus serotinus are entomophilous, whileFraxinus excelsior is anemophilous. The pollen ofLathraea squamaria eventually becomes powdery and suited for wind dispersal. The broad transition field between entomo- and anemophily is demonstrated byPlantago andArtemisia. Plantago lanceolata is anemophilous, whilePl. serpentina andPl. media are amphiphilous.Artemisia nitida andA. mutellina are entomophilous,A. absinthium, A. pedemontana andA. gabriellae are amphiphilous, but tend towards entomo- (A. absinthium) or anemophily (A. gabriellae), whileA. chamaemelifolia and especiallyA. vulgaris are anemophilous.

     

Correlation between pollen morphology and pollination mechanisms in the Hydrocharitaceae

The pollen morphology of 11 genera and 11 species of the Hydrocharitaceae and one species of the Najadaceae was studied using scanning and transmission electron microscopies, and the exine structures and sculptures are discussed in relation to pollination mechanisms and the molecular phylogeny. The pollen grains of the Hydrocharitaceae are spherical, inaperturate, and form monads or tetrads, while those of the Najadaceae are elliptical, inaperturate, and form monads. The entomophilous genera Egeria, Blyxa, Ottelia, Stratiotes, and Hydrocharis share pollen grains that have projections like spines or bacula. The anemophilous genus Limnobium has reticulate pollen grains. The hypohydrophilous genera Thalassia and Najas are characterized by pollen grains with reduced exine structures. The pollen-epihydrophilous genera Elodea and Hydrilla have tightly arranged small spinous pollen grains, and the male flower-epihydrophilous genera Enhalus and Vallisneria have reduced reticulate or gemmate exines. Character state reconstruction of the exine structures and sculptures using a molecular phylogenetic tree suggests that variation in the exine is generally correlated with the pollination mechanism; the selective pressures acting on the pollination mechanisms have reduced the exine structure in hypohydrophilous plants and resulted in various exine sculptures that are adapted to the different pollination mechanisms in entomophilous, anemophilous, and pollen-epihydrophilous plants.     

Types of pollen dispersal units in orchids,and their consequences for germination and fertilization

The various pollen dispersal units (PDU) found in orchids are discussed together with possible evolutionary trends and the consequences for germination and fertilization. Orchids with monad and tetrad pollen form more complex dispersal units by means of pollenkitt, elastoviscin, a callosic wall, common walls or a combination of these. Evolutionary trends include (1) from pollenkitt to elastoviscin; (2) from monad to tetrads and multiples of tetrads; (3) from partially dehydrated (<30 %) to partially hydrated (>30 %) pollen; and (4) from monad pollen to PDUs with many pollen grains. The biological consequences concern both male and female reproductive systems. Some features of the male side are present in all orchids irrespective of the pollen dispersal unit, whereas other characters are found only in orchids with pollinia; the same applies for the female counterpart. Pollen grains of orchids with pollinia germinate at least 24 h after pollination because the pollen grains/tetrads must swell and make space for the growth of pollen tubes.     

Seed hairs of poplar trees as natural airborne pollen trap for allergenic pollen grains

The present article deals with the efficacy of seed hairs of poplar trees (Populus spp.) as a potent natural airborne pollen trap. Different species of Populus are commonly found planted along the streets in the cities of North China. The seed hairs and pericarp of poplar trees were collected from the trees and on the ground in Beijing Botanical Garden of Chinese Academy of Sciences and around Miyun Reservoir during May 2005 for pollen analysis. Different pollen spectra are recorded from different samples and are characterised by dominant occurrence of pollen grains of arboreal and anemophilous plants. In addition, pollen grains of non‐arboreal plants including grasses are also found trapped. Among the 46 trapped pollen grains, 26 are known to be allergenic. This study suggests that poplar seed hairs possibly make people feel uncomfortable due to the presence of allergenic pollen trapped in the hairs.