POLLEN-POLLEN AND POLLEN-STYLE INTERACTIONS DURING POLLEN TUBE GROWTH IN ERYTHRONIUM GRANDIFLORUM (LILIACEAE)

Using pairs of pollen donors, I analyzed the growth of pollen tubes growing in different channels of the same style in Erythronium grandiflorum. After 24 hr the length of pollen tubes of randomly selected pollen donors was affected by the presence of other donors. The mean pollen tube lengths of donors did not differ when taken across all of the donor pairings, but in individual pairs, pollen from one donor was often significantly longer than pollen from the other donor when tested across several recipients. Pollen tube lengths were also consistently longer for pollen paired with pollen from the same donor than when paired with pollen from other donors, apparently because of mutual stimulation between the pollen populations from the same plant. In a second experiment, the amount of pollen tube attrition after five days of growth of pollen from a donor growing near (within 3 m) the recipient depended upon the source of other pollen growing in the same style. Local pollen experienced more attrition paired with self pollen than when paired with outcross pollen. Pollen from different outcross distances also modified the attrition of local pollen, but local pollen usually outcompeted pollen from greater distances. Since the growth of local pollen was modified depending upon the source of other pollen growing in the same style, it is probable that recipient styles are selectively inhibiting pollen tubes to produce the patterns of pollen tube interaction observed. The results from these two experiments indicate that the amount of attrition for pollen can be dependent on the composition of the pollen pool. Both direct pollen tube interactions and mediation by the stylar tissue appear to affect the growth rate and attrition of pollen in Erythronium.     

Pollen-pistil size correlation and pollen size-number trade-off in species of Argentinian Nyctaginaceae with different pollen reserves

Data on pollen and pistil traits from 14 Argentinean Nyctaginaceae species with starch or lipids as pollen reserves are presented. We expect differences in the traits between these two groups of species, but the same pattern within each group for (a) the relationship between pollen size and pistil length assuming that pollen tube length is predetermined by provisions in the pollen independently of the pollen reserve type, and for (b) a trade-off between size and number because available resources for male function are not unlimited. In particular we expect that (a) species with longer pistils will have larger pollen grains, (b) pollen grain size and the number of pollen grains per flower will be negatively correlated. Significant differences in the mean pollen number per flower and mean pollen size were observed between species with different pollen reserve type but not for pistil length. Then, correlation analyses were performed for species with starchy pollen or with pollen with lipids separately. Pollen size - pistil length correlation was positive and significant for species with starchy pollen but not for species with pollen with lipids. On the other hand, pollen size - number correlation was not significant for both starchy and oil-rich species. Results suggest that pollen reserve type would be a relevant factor that constraint pollen size in species of Nyctaginaceae, and that this pollen trait should be considered when studying pollen-pistil relationships, mainly between species of those families with mixed pollen reserves.     

How does secondary pollen presentation affect the fitness of Polygala vayredae (Polygalaceae)?

Secondary pollen presentation is the relocation and presentation of pollen in floral structures (termed pollen presenters) other than the anthers. These pollen presenters are often found close to the stigma and have been hypothesized to increase the accuracy of pollen transfer, although no experimental studies have been done. We examined the function of the pollen presenter and its efficiency in pollen dispersal, female fitness, and the degree of interference created by self-pollen in the shrublet Polygala vayredae, an insect-pollinated species with secondary pollen presentation. Herkogamy, a mechanism generally involved in the reduction of self-interference, was also evaluated. Significant pollen was lost (49% of total pollen) during the secondary relocation in the pollen presenter. However, pollen was exported from the pollen presenter, and subsequent pollen losses were similar to those in species with primary pollen presentation. Despite the presence of a self-incompatibility system, the numbers of developed pollen tubes as well as fruit and seed production were significantly reduced by the self-pollen interference created at the stigmatic papillae level. The extent of herkogamy correlated positively with female fitness. The secondary pollen presentation mechanism may in fact be an accurate system for pollen transport, but it may also have its costs. Further comparative studies involving species with primary and secondary pollen presentation are needed to fully understand the advantages and disadvantages of secondary pollen presentation.     

Reproductive isolation and hybrid pollen disadvantage in Ipomopsis

One cause of reproductive isolation is gamete competition, in which conspecific pollen has an advantage over heterospecific pollen in siring seeds, thereby decreasing the formation of F1 hybrids. Analogous pollen interactions between hybrid pollen and conspecific pollen can contribute to post-zygotic isolation. The herbaceous plants Ipomopsis aggregata and I. tenuituba frequently hybridize in nature. Hand-pollination of I. aggregata with pollen from F1 or F2 hybrids produced as many seeds as hand-pollination with conspecific pollen, suggesting equal pollen viability. However, when mixed pollen loads with 50% conspecific pollen and 50% hybrid pollen were applied to I. aggregata stigmas, fewer than half of the seeds had hybrid sires. Such pollen mixtures are frequently received if plants of the two species and F1 and F2 hybrids are intermixed, suggesting that this advantage of conspecific over hybrid pollen reduces backcrossing and contributes to reproductive isolation.     

阴山山脉东段花粉通量及其与表土花粉比较研究

阴山山脉东段蛮汉山和大青山山顶草甸、桦木林、虎榛子灌丛、人工油松林及杂草草原5个植被类型花粉通量与表土花粉对比研究发现,不同植被类型及同一植被类型不同样点花粉通量和花粉浓度差异明显,捕捉器样品花粉源区面积小于表土样品,能较好地反映样品点周围植被组成,但受虫媒植物和局地植物花粉影响,与表土样品花粉组合差异明显;表土样品中松属、桦属、蒿属及藜科花粉含量多高于捕捉器样品,表明这些花粉不仅易于传播,而且在表土中具有较强的保存能力;禾本科花粉具低代表性,花粉保存能力较低.地层花粉分析中应引起注意.     

Pollen–pollen interactions in Betula pendula in vitro

Direct pollen interactions, as well as interactions mediated by a recipient, can have a remarkable influence on pollen fertilization ability. Under conditions of pollen competition it could be advantageous if pollen grains interfered with the germination of other pollen. The aim of this study was to find out if there are direct negative or positive pollen–pollen interactions between pollen grains from genetically slightly different donors. The in vitro germinability of the pollen from several Betula pendula Roth clones was investigated. The pollen interactions between the clones were examined pairwise by using equal pollen mixtures. In three of the eight cases the germination percentage of the pollen mixture was significantly higher than the average germination percentage of the separate clones that formed the mixture, which indicates some type of interaction between the pollen populations. We found only positive interactions between the pollen of clones. This study also documented density-dependent germination of pollen grains in vitro (=pollen population effect). Adding an aqueous pollen extract to the incubation medium increased the germination percentages of poorly germinating pollen and small pollen populations. Germination-stimulating effects were found to exist both with fresh and dead pollen. Such direct pollen–pollen interactions could be explained by specific water-soluble substances diffusing from pollen grains.     

燕山南部花粉散布特征

燕山南部花９粉用布特性研究表明,空气中的花粉反映的是植物花期的季节性变化,表 粉是植物多年花粉散布的混合,冲积物花粉与空气中和表土花粉显著不同,是取样点上游表土花粉的混合,与取样点周围植被关系较小。因此,根据冲积物花粉恢复古植被,反映的是整个流域的植被面貌,而不是某一点的植被特征。     

Sunny-side up: flower heliotropism as a source of parental environmental effects on pollen quality and performance in the snow buttercup, Ranunculus adoneus (Ranunculaceae)

Floral traits affect mating success via their influence on the microenvironment in which sexual reproduction occurs as well as their impact on pollinator attraction. Here we investigate the importance of flower heliotropism as a source of parental environmental effects on pollen quality and performance. Flowers of the snow buttercup, Ranunculus adoneus, closely track the sun's rays. We experimentally restrained flowers to test for effects of heliotropism on pollen quality and performance after pollination. When equivalent amounts of pollen were transferred to recipient pistils, pollen from solar-tracking donor flowers exhibited a 32% advantage in germination compared to pollen from stationary (tethered) donor flowers. By the end of anthesis, pistils of tracking flowers contained 40% more germinating pollen grains and 44% more pollen tubes midway down the style than pistils of stationary ones. Solar tracking had no direct effect on pollen tube growth. The greater amount of germinating pollen in tracking flowers accounted for the treatment effect on pollen tube density. A survey of pollen receipt and pollen germination in naturally tracking flowers indicated that solar tracking primarily affects pollen tube density by promoting pollen germination rather than pollen deposition. We conclude that flower heliotropism, by enhancing the paternal environment for pollen development and the maternal environment for pollen germination, represents a source of positive parental environmental effects on pollen performance in snow buttercups.     

Microgametophyte population sizes and plant reproductive output in the insect-pollinated Prunella grandiflora (Lamiaceae)

Intraspecific variation in pollen deposition and number of pollen tubes per style is rarely quantified, but is essential for assessing the occurrence of pollen limitation and pollen competition and their evolutionary implications. Moreover, pollen deposition, pollen tube growth, and the fate of fertilized ovules are rarely distinguished in field studies. Here we present such a study in eight natural populations of Prunella grandiflora. We quantified microgametophyte population sizes and inferred pollen limitation when the number of fertilizable ovules exceeded pollen tubes, and assessed seed set and fate after open pollination. Two and three populations had on average significantly fewer pollen grains and pollen tubes per flower, respectively, than the fixed number of fertilizable ovules per fruit, while one population experienced significant pollen competition. Style length was positively correlated with the number of pollen tubes. While pollen availability was very variable, seed abortion was significantly less frequent in denser populations, and in one population the proportion of well-developed seeds was significantly, positively correlated with the number of pollen tubes in the style. Less pollen deposition, lower numbers of pollen tubes reaching the base of the style, lower pollen quality and therefore increased abortion of fertilized ovules can all reduce seed set in natural P. grandiflora stands. Substantial intraspecific variability implies that microgametophyte competition also occurs in this species. Finally, style morphology may affect pollen receipt.     

Feeding friend and foe: ample pollen mitigates the effects of pollen theft for a gynodioecious plant,Polemonium foliosissimum (Polemoniaceae)

Background and AimsMost angiosperms rely on pollinators to transport pollen and effect fertilization. While some floral visitors are effective pollinators, others act as thieves, consuming pollen but effecting little pollination in return. The importance of pollen theft in male and female reproductive success has received little attention. Here, we examined if pollen consumption by flies altered pollen receipt and exacerbated pollen limitation for a bumblebee-pollinated plant, Polemonium foliosissimum (Polemoniaceae).MethodsTo examine the effect of pollen-thieving flies, we took a three-pronged approach. First, we used single-visit observations to quantify pollen removal and pollen deposition by flies and bumblebees. Second, we manipulated pollen in the neighbourhood around focal plants in two years to test whether pollen reduction reduced pollen receipt. Third, we combined pollen reduction with hand-pollination to test whether pollen thieving exacerbated pollen limitation. Polemonium foliosissimum is gynodioecious in most populations in the Elk Mountains of central Colorado, USA. Thus, we also tested whether pollen theft affected hermaphrodites and females differently.ResultsFlies removed significantly more pollen and deposited less pollen per visit than did bumblebees. Reduction of pollen in the neighbourhood around focal plants reduced pollen receipt in both years but only nearly significantly so in 2015. In 2016, plants were significantly pollen-limited; hand-pollination significantly increased seeds per fruit for both hermaphrodites and females. However, the reduction of pollen around focal plants did not exacerbate pollen limitation for either hermaphrodites or females.ConclusionsOur results suggest that plants tolerate significant consumption of pollen by thieves and pollinators by producing ample pollen to feed both and fertilize available ovules. Our results demonstrate that pollen limitation in P. foliosissimum is driven by lack of effective pollinators rather than lack of pollen. Teasing out these effects elucidates the relative importance of drivers of reproductive success and thus the expected response to selection by different floral visitors.     

The effects of pollen load size and donor diversity on pollen performance, selective abortion, and progeny vigor in Mirabilis jalapa

The influence of pollen competitive environment on pollen performance (pollen germination, stigmatic penetration, and pollen tube growth rate), the maturation or abortion of initiated fruit, seed size, and seedling vigor was explored by manipulating the size and diversity of stigmatic pollen loads on Mirabilis jalapa. All aspects of pollen performance significantly increased with the number of pollen grains on a stigma or pollen tubes in a style, but was not influenced by the diversity of pollen donors. Plants tended to mature single-ovulate fruits that came from flowers where pollen load size and diversity were greatest and aborted those where these were lowest. No plants from seeds resulting from pollinations with a single pollen grain survived, but other fitness measures were mostly determined by maternal plant. The data suggest that pollen performance is influenced by pollen competitive environment, and both the genetic diversity of the pollen load and number of competing pollen tubes are important determinants of seed/fruit abortion.     

Pollen nutritional content and digestibility for animals

This paper reviews the literature concerning digestion and nutrient content of pollen. Four topics are addressed in detail: 1) The mechanism of pollen digestion by animals; 2) The efficiency of mechanical and digestive removal of pollen content by various animals; 3) Range and taxonomic distribution of pollen nutrients, and 4) Adaptive hypotheses proposed to associate pollen chemistry with pollinator reward. Studies on the mechanism(s) of pollen digestion remain inconclusive, but suggest that differences in digestibility among pollen types may reflect differences in pollen wall porosity, thickness, and composition. Although hummingbirds reportedly digest pollen very poorly, most animals studied, including those that do not regularly consume pollen, can digest 50–100% of ingested grains. Overlooked and recent research of pollen protein content shows that pollen grains may contain over 60% protein, double the amount cited in some studies of pollen-feeding animals. Adaptive hypotheses that associate pollen starch and pollen caloric content with pollinator reward remain unsubstantiated when critically viewed through the lens of phylogeny.     

Environmental Effects on Components of Pollen Performance in Faramea occidentalis (L.) A. Rich. (Rubiaceae)1

Studies focusing on gametophytic competition have focused on differences in pollen performance (e.g., pollen germination rate and pollen tube growth rate) among donors in order to examine genetic sources of variation in siring success among them. Donors that produce the fastest growing pollen tubes are expected to fertilize more ovules relative to donors with slow growing pollen tubes under conditions of gametophytic competition. However, the performance of pollen in the field is known to be influenced by environmental conditions in addition to the genotype of donor plants. This field study of Faramea occidentalis was conducted to: (1) determine the effect of environmental conditions during pollination on pollen performance; (2) measure differences among pollen donors in pollen performance; (3) determine if the pollen of different donors responds differently to a variety of environmental conditions surrounding pollinated flowers; and (4) measure differences among pollen recipients in pollen performance. Single-donor crosses were made between four pollen donors and four pollen recipients under a variety of environmental conditions. Pollen performance was then quantified as the growth rate of the fastest pollen tube, the mean pollen tube growth rate, and by a pollen germination index. Pollination environment (the environment surrounding a pollinated flower) and recipient significantly affected all three measures of pollen performance. Pollen donors did not differ overall in pollen performance. However, there was significant among-donor variation in two of the five pollination environment conditions. Future studies of variation in relative siring success may benefit by considering pollination environment in addition to donor and recipient identity.     

Pollen performance, cell number, and physiological state in the early-divergent angiosperm Annona cherimola Mill. (Annonaceae) are related to environmental conditions during the final stages of pollen development

Pollen performance is an important determinant for fertilization success, but high variability in pollen behavior both between and within species occurs in different years and under varying environmental conditions. Annona cherimola, an early-divergent angiosperm, is a species that releases a variable ratio of bicellular and tricellular hydrated pollen at anther dehiscence depending on temperature. The presence of both bi- and tricellular types of pollen is an uncommon characteristic in angiosperms and makes Annona cherimola an interesting model to study the effect of varying environmental conditions on subsequent pollen performance during the final stages of pollen development. In this work, we study the influence of changes in temperature and humidity during the final stages of pollen development on subsequent pollen performance, evaluating pollen germination, presence of carbohydrates, number of nuclei, and water content. At 25?°C, which is the average field temperature during the flowering period of this species, pollen had a viability of 60-70?%, starch hydrolyzed just prior to shedding, and pollen mitosis II was taking place, resulting in a mixture of bi- and tricellular pollen. This activity may be related to the pollen retaining 70?% water content at shedding. Temperatures above 30?°C resulted in a decrease in pollen germination, whereas lower temperatures did not have a clear influence on pollen germination, although they did have a clear effect on starch hydrolysis. On the other hand, slightly higher dehydration accelerated mitosis II, whereas strong dehydration arrested starch hydrolysis and reduced pollen germination. These results show a significant influence of environmental conditions on myriad pollen characteristics during the final stages of pollen development modifying subsequent pollen behavior and contributing to our understanding of the variability observed in pollen tube performance.     

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.     

Variations in mugwort (Artemisia spp.) airborne pollen concentrations at three sites in central Croatia, in period from 2002 to 2003

In spite of the low atmospheric pollen levels, Artemisia sensitisation and allergy has been reported widely. The aim of the study was to determine the length of pollen season, intradiurnal, daily and monthly pollen variation, and the effect of some meteorological parameters on atmospheric pollen concentrations in Central Croatia. Seven-day Hirst volumetric pollen and spore traps were used for pollen sampling. The Artemisia pollen season lasted from the end of July until the end of September with the highest concentrations in August. The percentage of the total pollen count ranged from 0.52% to 0.92%. The intradiurnal peak occurred between 10 a.m. and 12 a.m. Statistical analysis showed a significant correlations between higher air temperature and high pollen concentration as well as high precipitation and low pollen concentration. Results of this study are expected to help in preventing the symptoms of allergic reaction in individuals with Artemisia pollen hypersensitivity.     

干旱区石羊河流域河水孢粉组合特征

石羊河流域初春和仲夏两季不同地点 ,洪水期、平水期 30个河水样孢粉分析显示 ,没有人类影响的情况下 ,河水样孢粉组合是由河流上游径流区的植物孢粉和采样点附近植被孢粉组成的 ,前者在孢粉组合中至少占30 %～ 6 0 %。河水搬运孢粉的能力非常强 ,可长距离、大量地搬运孢粉。孢粉组合中河水贡献孢粉的含量较高 ,如 :云杉属 (PiceaDietr.)花粉河水的贡献率在中下游可达 16 .5 %和 7.7%。采样时间、地点影响河水的孢粉组合和浓度 ,洪水期是孢粉搬运的主要时期。河水经过水库后 ,其中孢粉绝大多数沉积在水库中。     

Do pollen carryover and pollinator constancy mitigate effects of competition for pollination?

Pollinator constancy and pollen carryover are both thought to mitigate competitive effects that result when shared pollinators cause loss of pollen to heterospecific flowers. I present analytical and simulation models to investigate how pollinator constancy and pollen carryover interact with each other and with the relationship between pollen receipt and seed set to determine pollination success in competitive environments. With inconstant pollinators, increased pollen carryover reduces variance in pollen receipt without affecting average pollen receipt. Consequently, for flowers requiring at least a threshold quantity of pollen for success, rare flowers with inconstant pollinators benefit from reduced carryover, especially for high pollen receipt thresholds, whereas common flowers benefit from increased carryover, especially for low receipt thresholds. Pollinator constancy is predicted to increase pollen receipt, especially if pollen carryover rates are low. As a result, increased pollinator constancy reduces the range of pollen receipt thresholds for which carryover is beneficial. Similarly, for flowers whose pollination success is a convex function of pollen receipt, carryover is expected to increase fecundity if pollinators are inconstant, but with even a low degree of pollinator constancy, carryover reduces fecundity. These results predict that rare plants with many ovules per flower benefit from dispersing aggregations of pollen, especially if their pollinators exhibit constancy, whereas plants with inconstant pollinators and low thresholds of pollen receipt benefit from pollen grains dispersing individually to increase the number of flowers reached by the pollen.     

Variability within the 10-Year Pollen Rain of a Seasonal Neotropical Forest and Its Implications for Paleoenvironmental and Phenological Research

Tropical paleoecologists use a combination of mud-water interface and modern pollen rain samples (local samples of airborne pollen) to interpret compositional changes within fossil pollen records. Taxonomic similarities between the composition of modern assemblages and fossil samples are the basis of reconstructing paleoclimates and paleoenvironments. Surface sediment samples reflect a time-averaged accumulation of pollen spanning several years or more. Due to experimental constraints, modern pollen rain samples are generally collected over shorter timeframes (1–3 years) and are therefore less likely to capture the full range of natural variability in pollen rain composition and abundance. This potentially biases paleoenvironmental interpretations based on modern pollen rain transfer functions. To determine the degree to which short-term environmental change affects the composition of the aerial pollen flux of Neotropical forests, we sampled ten years of the seasonal pollen rain from Barro Colorado Island, Panama and compared it to climatic and environmental data over the same ten-year span. We establish that the pollen rain effectively captured the strong seasonality and stratification of pollen flow within the forest canopy and that individual taxa had variable sensitivity to seasonal and annual changes in environmental conditions, manifested as changes in pollen productivity. We conclude that modern pollen rain samples capture the reproductive response of moist tropical plants to short-term environmental change, but that consequently, pollen rain-based calibrations need to include longer sampling periods (≥7 years) to reflect the full range of natural variability in the pollen output of a forest and simulate the time-averaging present in sediment samples. Our results also demonstrate that over the long-term, pollen traps placed in the forest understory are representative samples of the pollen output of both canopy and understory vegetation. Aerial pollen traps, therefore, also represent an underutilized means of monitoring the pollen productivity and reproductive behavior of moist tropical forests.     

Effects of pollen competitive environment on pollen performance in Mirabilis jalapa (Nyctaginaceae)

 We examined the influence of pollen competitive environment on pollen performance in Mirabilis jalapa. We used the number of pollen grains and the number of pollen tubes per pistil as measures of pollen competition. Pollen germination, pollen tube penetration into the style, and pollen tube growth rates were used as measures of pollen performance. All three measures of pollen performance were affected by the competitive environment. Pollen germination was greatest at intermediate pollen load sizes. The percentage of germinated pollen grains that penetrated the stigma and grew into the style decreased with pollen load size. Pollen tube growth rate in the style was greater and more variable with larger numbers of pollen tubes in the style. Controlling for the degree of selection at the stigma indicated that pollen-pollen or pollen-style interactions were the likely causes of increased growth rates. Received: 28 October 1996 / Revision accepted: 24 January 1997