A long-term study of winter and early spring tree pollen in the Tulsa, Oklahoma atmosphere

Since 1986 the atmosphere in Tulsa, Oklahoma has been monitored for airborne pollen and spores with a Burkard 7-day spore trap situated on the roof of a building at The University of Tulsa. The present study specifically examined the early spring tree pollen season for several local taxa and the occurrence of pre-season pollen during December and January. Knowledge of the local pollen season will help identify the presence of out-of-season pollen and possible long distance transport (LDT) events. Average daily concentrations of airborne pollen for species ofBetula, Quercus, Ulmus, and Cupressaceae were determined for each year from 1987 to 1996. The data showed that during the early spring the precise pollination periods for these allergenic tree species are highly variable. There were considerable variations in start date, season length, peak concentration, date of peak, and cumulative season total. The start dates forUlmus, Betula, andQuercus varied by 30 days or more, while the early spring Cupressaceae pollen showed the least variation in start date (only 23 days). More research is needed to understand the mechanisms which govern the onset and magnitude of pollen release. Although several reports have documented episodes of long distance transport (LDT) of pollen, the actual contribution of out-of-season or out-of-region pollen to local air spora is poorly known. The current study also re-examined the LDT ofJuniperus ashei pollen in Oklahoma.Juniperus pollen appeared in the Tulsa atmosphere on 40% of the days in December and January with concentrations as high as 2400 pollen grains/m³ of air; however, no local populations ofJuniperus pollinate at this time of the year. High concentrations occurred on days with southerly winds suggesting thatJuniperus ashei populations in southern Oklahoma and Texas were the pollen source. Since no local pollen is present in the Tulsa atmosphere in December and January, this example of LDT has been easy to document.     

土壤与大气Cu处理下迎春的耐性和富集特征研究

作为北京市常见园林灌木树种之一，迎春(Jasminum nudiflorum)因其在早春独特的观赏性而深受市民喜爱。Cu污染是北京市较为严重的重金属污染类型之一。为探讨迎春对城市Cu污染的修复作用，该文通过模拟北京市土壤和大气Cu污染条件，采用盆栽试验，设置9种不同浓度的土壤和大气Cu处理，以验证迎春Cu富集能力及生理生长特性。结果表明：(1)土壤和大气沉降处理均能显著增加迎春根、茎、叶中的Cu含量，其中土壤贡献率为63.48%～96.99%。各处理中Cu含量均表现为根>茎>叶。(2)大气处理下光化学转化效率(F_v/F_m)和相对叶绿素含量(SPAD值)提高，初始荧光(F₀)降低，光合能力增强，而土壤处理及土壤和大气双重处理则对迎春的光合作用产生抑制影响。(3)与大气处理相比，土壤处理及土壤和大气双重处理导致活性氧(ROS)积累增多，膜脂过氧化作用加剧，丙二醛(MDA)含量大幅升高，抗氧化酶活性与脯氨酸(PRO)含量逐渐下降，造成生物膜系统损伤。(4)低浓度Cu处理对迎春生长有促进作用，而高浓度Cu处理(SHA...     

Levels of Ambrosia pollen in the atmospheric spectra of Catalan aerobiological stations

Ambrosia pollen is known as an importantallergen in North America, and more recently in someEuropean countries. From 1989 to 1995, the Ambrosia pollen levels detected at the stationsmonitored by the Aerobiological Network of Catalonia(Xarxa Aerobiològica de Catalunya, XAC) wereinsignificant. In 1996, a considerable althoughtemporary increase in the concentration of this pollenwas detected in the atmosphere over Girona, Barcelona,Bellaterra, Manresa, and Tarragona. Most of the Ambrosia pollen collected in 1996 was concentrated ina single day. Its appearance on that day wasattributed to long range transportation, caused byunusual conditions of atmospheric circulation, fromthe Lyon region in France where the species isabundant. This is the only day where concentrations ofAmbrosia pollen that may be dangerous to humanhealth have been reached. Ambrosia coronopifolia is the most abundantspecies of the genus in Catalonia, and although rare,its expansion is favoured by the fact that it growsthrough rhizomes and sprouts easily. It is, therefore,important to monitor the growth of its population andthe release of its pollen in order to predict theappearance of pollen levels that may precipitateallergic symptoms.     

Interspecific pollen transfer as a mechanism of competition: Consequences of foreign pollen contamination for seed set in the alpine wildflower,Polemonium viscosum

Summary Earlier studies showed that under natural conditions foreign pollen receipt is correlated with loss of seed set in populations of P. viscosum. Here, we report on experimental pollinations using foreign pollen from two co-flowering species that share pollinators with P. viscosum. Experiments were designed to ascertain whether improper pollen transfer causes reduced seed set under controlled conditions and, if so, to determine the stage at which interference occurs. In flowers pre-treated by pollination with either Mertensia or Castilleja pollen, adherence of subsequently deposited conspecific pollen grains was unaffected. However, conspecific grains had significantly lower germination and ovule fertilization success in flowers receiving foreign pollen than in controls. These results suggest that pollinator infidelity in high alpine plant communities can reduce female reproductive success of P. viscosum by inhibiting the performance of conspecific pollen grains. Negative effects of Mertensia and Castilleja pollen were comparable despite large interspecific differences in pollen grain morphology. Accordingly, susceptibility to foreign pollen interference appears to be an intrinsic property of P. viscosum that is not dependent on the unique structural properties of pollen produced by competitive species.     

Examining <Emphasis Type="Italic">Ambrosia</Emphasis> pollen episodes at Poznań (Poland) using back-trajectory analysis

The pollen grains of Ambrosia spp. are considered to be important aeroallergens in parts of southern and central Europe. Back-trajectories have been analysed with the aim of finding the likely sources of Ambrosia pollen grains that arrived at Poznań (Poland). Temporal variations in Ambrosia pollen at Poznań from 1995–2005 were examined in order to identify Ambrosia pollen episodes suitable for further investigation using back-trajectory analysis. The trajectories were calculated using the transport model within the Lagrangian air pollution model, ACDEP (Atmospheric Chemistry and Deposition). Analysis identified two separate populations in Ambrosia pollen episodes, those that peaked in the early morning between 4 a.m. and 8 a.m., and those that peaked in the afternoon between 2 p.m. and 6 p.m.. Six Ambrosia pollen episodes between 2001 and 2005 were examined using back-trajectory analysis. The results showed that Ambrosia pollen episodes that peaked in the early morning usually arrived at Poznań from a southerly direction after passing over southern Poland, the Czech Republic, Slovakia and Hungary, whereas air masses that brought Ambrosia pollen to Poznań during the afternoon arrived from a more easterly direction and predominantly stayed within the borders of Poland. Back-trajectory analysis has shown that there is a possibility that long-range transport brings Ambrosia pollen to Poznań from southern Poland, the Czech Republic, Slovakia and Hungary. There is also a likelihood that Ambrosia is present in Poland, as shown by the arrival of pollen during the afternoon that originated primarily from within the country.     

Water Relations Only Partly Explain the Distributions of Three Perennial Plant Species in a Semi-arid Environment

The water relations and stomatal conductances of three perennial plant species, Stipa tenacissima L., Anthyllis cytisoides L., and Retama sphaerocarpa (L.) Boiss., dominant on the upper slopes, mid-slopes and floor of a valley, respectively, in semi-arid south-east Spain, were investigated to test the hypothesis that differences in plant-soil water relations could account for the different distributions of each species in the catena. Diurnal measurements of water potential (Ψ_w), relative water content (RWC) and stomatal conductance (g_s) of leaves were made over one year. Leaf temperature, air humidity, wind-speed and incident quantum flux density were measured concurrently. Soil water content was determined gravimetrically at 0 – 5 cm and 15 – 20 cm depths. Measurements of Ψ_w, RWC and g_s were analysed according to meteorological conditions, based on the maxima for daily air temperature and atmospheric saturation water vapour deficit and on soil moisture content. The hypothesis that plant-soil water relations can explain the distribution of the three species along the catena from valley side to valley floor was rejected for Anthyllis and Stipa but confirmed for Retama. This revised version was published online in July 2006 with corrections to the Cover Date.     

Association of allergic rhinitis or asthma with pollen and chemical pollutants in Szeged,Hungary, 1999–2007

The effect of biological (pollen) and chemical air pollutants on respiratory hospital admissions for the Szeged region in Southern Hungary is analysed. A 9-year (1999–2007) database includes—besides daily number of respiratory hospital admissions—daily mean concentrations of CO, PM₁₀, NO, NO₂, O₃ and SO₂. Two pollen variables (Ambrosia and total pollen excluding Ambrosia) are also included. The analysis was performed for patients with chronic respiratory complaints (allergic rhinitis or asthma bronchiale) for two age categories (adults and the elderly) of males and females. Factor analysis was performed to clarify the relative importance of the pollutant variables affecting respiratory complaints. Using selected low and high quantiles corresponding to probability distributions of respiratory hospital admissions, averages of two data sets of each air pollutant variable were evaluated. Elements of these data sets were chosen according to whether actual daily patient numbers were below or above their quantile value. A nonparametric regression technique was applied to discriminate between extreme and non-extreme numbers of respiratory admissions using pollen and chemical pollutants as explanatory variables. The strongest correlations between extreme patient numbers and pollutants can be observed during the pollen season of Ambrosia, while the pollen-free period exhibits the weakest relationships. The elderly group with asthma bronchiale is characterised by lower correlations between extreme patient numbers and pollutants compared to adults and allergic rhinitis, respectively. The ratio of the number of correct decisions on the exceedance of a quantile resulted in similar conclusions as those obtained by using multiple correlations.     

Ragweed pollen: The aeroallergen is spreading in Italy

Presently in Europe, ragweed pollen as an aeroallergen is not as important as Poaceae,Parietaria or Betulaceae, even if in some countries the plant is beginning to influence the local composition of the airborne pollen spectra. In northern Italy, the presence of ragweed airborne pollen has only been reported since the beginning of the 1980’s and it is increasingly spreading from year to year. Given this situation, the allergologists have begun to regard the potential risk of sensitisation to ragweed pollen with much attention. Up to now, such pollen has not been included in the routine allergological tests. In 1995 in some sites of northern Italy (Turin, Milan, Trieste), the concentration values of ragweed pollen were remarkable (∼ 20–30 p/m³) and on the increase with respect to the previous years. This investigation aims at focusing the atmospheric concentration trend on this new aeroallergen (Ambrosia sp.) in Italy from 1991 throughout 1995.     

Airborne ragweed (Ambrosia artemisiifolia L.) pollen concentrations in Croatia, 2002–2004

The aim of this study was to determine the onset, length and end of the ragweed pollen season, taking into account diurnal, day-to-day, monthly and annual pollen variations, the effect of some meteorological parameters on atmospheric pollen concentrations and possible differences in the airborne pollen season and concentration due to sampling site. Airborne pollen was collected at three sites in central Croatia (Zagreb, Samobor and Ivanić Grad) during three pollen seasons (2002–2004). Seven-day Hirst-type volumetric pollen traps were used for pollen sampling. Ragweed pollen was the third most abundant pollen type to occur in the atmosphere of central Croatia. Total Ambrosia pollen concentration was the highest in the 2003 pollen season and the lowest in 2004 at all sampling sites. Maximum emissions were restricted to August and September. Intradiurnal periodicity showed a peak from 1000 to 1200 hours. The concentration of ragweed pollen during the pollen season was greatly influenced by temperature and precipitation: on rainy days accompanied by temperature decline, the air pollen concentration decreased abruptly. The results of this study are aimed at helping to alleviate the symptoms of allergic reactions in individuals with ragweed pollen hypersensitivity, thus improving their quality of life.     

Multiple ice-age refugia in a southern beech of South America as evidenced by chloroplast DNA markers

Chloroplast DNA polymorphisms were analysed to infer the post-glacial history of Nothofagus nervosa (Phil.) Dim. et Mil., a species endemic to South American Temperate Forests. Two hypotheses were postulated to explain the current distribution of the species in relation to the refugia proposed after palynological studies and the time elapsed since Last Glacial Maximum. If refugia were located only in the Coastal Mountains, as pollen records suggests, long-distance dispersal events should be invoked to explain the current occurrence of the species in southern latitudes. The alternative hypothesis is the existence of cryptic refugia in southern latitudes. A total of 26 populations covering the entire geographical range of the species in Chile and Argentina were analysed through PCR–RFLP. Five haplotypes were identified, and a very low intrapopulation variation was observed together with a high gene differentiation (G _ST=0.93). The haplotypes showed a highly structured geographic distribution, separating populations located in the Pacific Coastal Mountains from those coming from the easterly located Andes Mountains. Moreover, among the populations of the Andes Mountains a north–south variation in the distribution of haplotypes was found. The results strongly suggest the persistence of the species in several ice-age refugia. The possible location of such refugia is discussed in combination with the available palynological data.     

Concordant clines and significant correlation between floral and pollen characters in Asarum heterotropoides var. heterotropoides (Aristolochiaceae)

To examine geographic differentiation in Asarum heterotropoides var. heterotropoides in Hokkaido Is. in the northern Japan, in which two putative cryptic species have been suspected to exist, extensive and detailed morphological research on 794 individuals from 44 populations throughout Hokkaido Is. was performed. Among the characters examined, the angle between and tip shape of the calyx lobes and the supratecta of the pollen grains were significantly correlated and were found in similar geographic clines. Among them, the pollen showed two discrete states in almost distinct distribution. Multidimensional scaling analysis showed that individuals within each of the two pollen types had different trends in flower characters. Consequently, we assumed a cline from south to north on Hokkaido Is. For the causes of the cline, the two hypotheses were proposed, primary geographic differentiation or extensive introgressive hybridization between two distinct geographical species existing in the past.     

In vitro pollen germination and transient transformation of<Emphasis Type="Italic">Zea mays</Emphasis> and other plant species

To study pollen-specific gene expression, fast and convenient methods involving in vitro pollen germination and bombardment with promoter deletion constructs are needed. Unfortunately, because of variation of pollen germability and tube growth, conducting these experiments is often unsatisfying for many plant species, including maize, especially when pollen is collected at different times of the day or season. We have overcome these problems by defining a novel medium (PGM) that guarantees germination efficiencies of more than 90% for maize pollen from at least 7 genotypes (A188, AC 3572 C, B73, H99, Hi-II, Q2, Tx232). This medium is also suitable to germinate pollen of other monocot species, such asPennisetum americanum andTradescantia species, and dicot species, such asArabidopsis thaliana, Arachis hypogaea, Columnea oesterdiana, Nicotiana tabacum, Phaseolus vulgaris, Pisum sativum, Solanum lycopersicum, Solanum tuberosum, andVicia faba. On average, reproducible germination rates ranging from 50–100% were observed with all plant species tested. In addition, we report a transient transformation assay using the luciferase (Luc) reporter gene. Biolistic parameters were defined to obtain reproducibleLuc activity measurements after bombarding thick-walled pollen, such as maize pollen. For comparison, samples of germinated maize and tobacco pollen were bombarded with the reporter gene under control of the constitutive ubiquitin-and pollen-specificZmMADS2 maize promoters. The important parameters necessary to apply both in vitro pollen germination and transient transformation for a large range of plant species are discussed. An erratum to this article is available at .     

Nitrogen and Carbon Concentrations, and Stable Isotope Ratios in Mediterranean Shrubs Growing in the Proximity of a CO2 spring

Seasonal changes in foliage nitrogen (N) and carbon (C) concentrations and δ¹⁵N and δ¹³C ratios were monitored during a year in Erica arborea, Myrtus communis and Juniperus communis co-occurring at a natural CO₂ spring (elevated [CO₂], about 700 μmol mol⁻¹) and at a nearby control site (ambient [CO₂], 360 μmol mol⁻¹) in a Mediterranean environment. Leaf N concentration was lower in elevated [CO₂] than in ambient [CO₂] for M. communis, higher for J. communis, and dependent on the season for E. arborea. Leaf C concentration was negatively affected by atmospheric CO₂ enrichment, regardless of the species. C/N ratio varied concomitantly to N. Leaves in elevated [CO₂] showed lower δ¹³C, and therefore likely lower water use efficiencies than leaves at the control site, regardless of the species, suggesting substantial photosynthetic acclimation under long-term CO₂-enriched atmosphere. Leaves of E. arborea showed lower values of δ¹⁵N under elevated [CO₂], but this was not the case of M. communis and J. communis foliage. The use of the resources and leaf chemical composition are affected by elevated [CO₂], but such an effect varies during the year, and is species-dependent. The seasonal dependency and species specificity suggest that plants are able to exploit different available water and N resources within Mediterranean sites. This revised version was published online in July 2006 with corrections to the Cover Date.     

<Emphasis Type="Italic">Cyperaceae</Emphasis> and <Emphasis Type="Italic">Juncaceae</Emphasis> pollination measured in the air at two sites in SW Spain

The presence of airborne Cyperaceae and Juncaceae pollen was quantified using volumetric aerobiological traps over a 10-year period at two sites in SW Spain separated by 60 km (Badajoz 10 years, Mérida 3 years). The Pearson correlation coefficient was calculated between the daily and hourly concentrations. The values of the principal meteorological parameters—temperature, rainfall, relative humidity, and speed and direction of the wind—were calculated during the study period, and with the accumulated values corresponding to the period prior to pollination. The beginning and ending dates and the duration of the main pollen season (MPS) were also analyzed. Even though both families are anemophilous, the presence of their pollens in the air was low. The annual accumulated daily concentrations were in the range 1.8–15.8 for Juncaceae and 111.8–473.9 for Cyperaceae—values far lower than any other anemophilous pollen type. The Cyperaceae pollen concentration peaked between 09:00 and 12:00. The meteorological factor most closely related to its daily variations was found to be the wind direction, showing that location of the sources is of great importance. The results lend support to the hypothesis of a limitation of allogamous reproduction in favour of vegetative multiplication in both families. Nevertheless, the principal source of the airborne Cyperaceae pollen was found to be Scirpus holoschoenus, whose pollen is distinguishable from the rest. Hence, because of its large production of pollen, this species can be characterized as anemophilous and allogamous. Rainfall in the preceding autumn seemed to be responsible for the amount of Cyperaceae pollen in the air, since a lack of rain was found to be associated with lower densities in the traps. For the Juncaceae, it seems that the temperatures of the preceding December constituted the most limiting meteorological parameter.     

Daily variations of Urticaceae pollen count and influence of meteoclimatic parameters in East Perugia during 1989

Summary Urticaceae pollen monitoring was carried out, in the atmosphere of East Perugia during the summer of 1989. In summer the Urticaceae pollen shows a fluctuating diurnal trend, which was analysed daily over six 4-hour periods. The high peaks of thistaxon were recorded in the middle of the day in July and August, and in the morning in June. The percentage pattern of the daily mean pollen concentration of Urticaceae was compared with mean temperature, wind speed, relative humidity and rain, both in the non-rainy days and rainy ones. The best correlations came out comparing pollen concentration and meteoclimatic parameters during non-rainy days. The interpretation of the influence of rainfalls is very difficult owing to their peculiar features in summer.     

Spatial Segregation of the Sexes and Nutrients affect Reproductive Success in a Dioecious Wind-pollinated Grass

In many dioecious plant species in which spatial distributions of males and females have been examined, the sexes are spatially segregated – usually along an environmental gradient. Unless pollen is uniformly distributed in a population, spatial segregation of the sexes should reduce the average mating success of individuals. In three Californian populations of Distichlis spicata – a wind-pollinated grass species that exhibits spatial segregation of the sexes – I examined patterns of pollen movement and the effects of pollen load and nutrient availability on seed set to determine whether spatial segregation of the sexes actually reduces mating success for both males and females. In two of the populations, pollen dispersal was restricted, and pollen augmentation consistently, significantly increased seed set. However, in the third population – which had the lowest seed set – I found that although there were some indications of pollen limitation, pollen dispersal was not restricted, and seed production was limited primarily by nutrient availability. These results imply that in some populations of D. spicata nutrient limitation on the production of seeds by females may be sufficiently strong that spatial segregation of the sexes causes a fairly low cost to reproductive success compared with a more random distribution of the sexes. However, in other populations, pollen does limit mating success, and the spatial segregation of males and females in these populations is reducing the fecundity of both males and females.     

Novel polymorphic microsatellite loci and patterns of pollen-mediated gene flow in an ex situ population of <Emphasis Type="Italic">Eurycorymbus cavaleriei</Emphasis> (Sapindaceae) as revealed by categorical paternity analysis

Understanding the patterns of contemporary, pollen-mediated gene flow is of great importance for designing appropriate conservation strategies. In this study, ten novel polymorphic microsatellite loci were isolated for the rare dioecious tree, Eurycorymbus cavaleriei, and the patterns of pollen dispersal were investigated in an ex situ conserved population. A combination of microsatellite markers with high-collective exclusion power (0.932) was used to assign paternity to 240 seeds collected from eight maternal trees. The average effective pollen dispersal distance (δ) was 292.6 m and the frequency distribution of pollen movement suggested extensive pollen movement in the population. The effective pollen donors per maternal tree (N _ep) ranged from 5 to 10, and the most isolated maternal trees were observed with the largest number of N _ep = 10. Although a trend of near-neighbor mating was found in seven of eight maternal trees, no significant correlations were detected between the average effective pollen dispersal distance (δ) and the geographic distances (d1 and d2) between maternal and male trees. The increased average effective distance of pollen dispersal and number of N _ep for isolated maternal trees might be a compound consequence of low density and long-distance flight of pollinators of this species. The conservation implications of these results are discussed.     

Ambrosia pollen in the air of Lublin, Poland

Studies on Ambrosia pollen concentrations were carried out in Lublin in the period 1995–2004. The effects of a number of meteorological factors were analysed. In the first period of the study, the gravimetric method was used (1995–1999), while in the second period, the volumetric method was applied. The results show an increasing trend in the amount of airborne pollen. The Ambrosia pollen season in Lublin lasts from August to October. Over a period of 5 years, the highest number of pollen grains was recorded in September (53%), followed by August (44%) and October (3%). There were wide variations in annual totals. The annual total pollen counts was 167–1180 grains, with the peak value in 2002. Maximum daily pollen concentrations (56–312 pollen grains m⁻³) were recorded in the first half of August and in the first half of September. On the days when high Ambrosia pollen concentrations occurred, the temperature was above 21°C and the winds were mainly from the southeast, south and east. Maximum intradiurnal concentrations of pollen grains occurred in the afternoon hours. These results indicate, to some degree, that Ambrosia pollen is transported for long distances before descent.     

Discovery of a life history shift: precocious flowering in an introduced population of Prosopis

The genus Prosopis contains many valuable, long-lived, multi-purpose legume trees, some of which are also invasive species. The time of first flowering is important for increasing production of sweet, protein-rich pods in plantations, especially under short rotations, and affects the rate of spread as a weed. Trees generally begin flowering at 3–5 years of age. However, seedlings from a small seedlot collected from an introduced population in southern Mauritania were observed to begin flowering at 3–4 months after germination. This is unknown in the native range of any Prosopis species, and appears to represent a major evolutionary event triggered after naturalization. This paper reports a detailed investigation with seed collected from Aleg, Brakna region, Mauritania, in 1998 (Prosopis sp. ‘Aleg’). Two experiments were established, in glasshouses at Cirad, Montpellier, France, and Coventry University, UK. Flowering began at the two sites 104 and 169 days after sowing, respectively, and 97.5% of plants had flowered at Coventry after 306 days. Flowers produced abundant pollen with 50–60% viability indicated by FDA staining and 40–50% germination on an artificial medium. No pods were formed. All plants analysed were diploid. Morphologically, these juvenile plants were similar to American species of section Algarobia, and exhibited leaf characters typical of P. pallida. However, due to the large variation in morphology within this section of the genus, further studies are required to confirm the actual species. This is the first record of precocious flowering in Prosopis, and has been found only in a single, introduced population. Implications for genetic improvement of tropical Prosopis, and particularly the spread of these species as invasive weeds, are discussed. Such a life history shift in the increased reproductive ability of a species following introduction, with potentially significant environmental effects, may not be restricted to Prosopis and merits further detailed investigation.     

Mating System in a Low-density Natural Population of the Dioecious Wind-pollinated Patagonian Cypress

The mating system of a species is an essential factor that determines the genotypic frequencies of its populations in each generation. Thus, knowledge in this regard is important in the application of practical measures in conservation and forest tree breeding. Austrocedrus chilensis (D.DON) PIC. SER. ET BIZZARRI is a member of the Cupressaceae native to the Andean–Patagonian Forest of Argentina and Chile with regional economic importance. It is a dioecious tree with pollen and seeds distributed by wind. Isozyme markers were utilized to study the mating system of this species through the survey of individual effective pollen clouds in a natural low-density population. If random mating occurs, effective pollen clouds that pollinate single seed trees are expected to be genetically homogeneous. The classical HardyWeinberg test of panmixia was also performed. Although it failed to reject random mating, the analysis of the pollen clouds showed that this mating pattern does not occur. Evidence of locally distributed heterogeneous pollen clouds is presented. The two methods are critically compared.