EVOLUTIONARY AND ECOLOGICAL SIGNIFICANCE OF STARCH STORAGE IN POLLEN OF THE ARACEAE

Starch content was qualitatively assessed for pollen of 79 of the 111 currently recognized genera of the family Araceae—one of three monocot families known to exhibit both starchy and starchless pollen. Although 73% of the genera investigated had exclusively starchy pollen, character correlation suggests that starchless pollen is the primitive type for the family Araceae, as well as for monocots in general. Pollen starch content is a highly conservative character at the generic level in Araceae; only a single genus (Schismatoglottis) clearly exhibits both character states. The distribution of starchy pollen among aroid genera is consistent with what have here been termed Bakers' Starch Laws. Aroid pollen below a certain critical diameter—17-25 μm—is almost invariably starchless. Larger pollen is nearly always starchy, except where insect pollinators may use pollen nutritionally. There is strong evidence that the trend from starchless to starchy pollen in Araceae is reversible, according to the constraints imposed by the aforementioned factors.     

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.     

Pollen starch reserves in tomato relatives: Ecophysiological implications

The presence or absence of starch in microspore development and in pollen grains was recorded in eleven wild tomato species (Solanum sect. Lycopersicon) and two close relatives (S. lycopersicoides and S. sitiens). In all the species starch started to accumulate in the early microspore bicellular stage and continued until the cytoplasm was filled. At flower anthesis, pollen grains were mostly starchless in the wild tomatoes, except in S. pennellii, which had starchy pollen. Starchy pollen is also present in the two related species. The latter two species had larger pollen grains and grow in drier environments than the other species. The heterogeneity of pollen starch content among all these species, supposed to have the same pollination mechanism, is a new finding supporting the idea that starch content and pollination mechanism do not necessarily influence each other. The presence of starchy pollen in the self‐incompatible species, which grow in the driest environments, raises questions regarding the relationship between carbohydrates content and pollen survival.     

叶下珠科花粉组织化学、花粉数和胚珠数及其与传粉者关系的研究

植物花粉中营养物质的储存形式以及单花的花粉数与胚珠数被认为与其传粉系统有一定的联系。本文研究了叶下珠科部分种类的花粉组织化学、花粉数和胚珠数, 以及它们与传粉者之间的关系。结果显示: 在叶下珠科内, 植物花粉所含营养成分和传粉者之间存在相关性: 蛾类传粉的类群主要属“淀粉型”花粉, 蝇类和蜂类传粉的类群主要属“非淀粉型”花粉。蝇类传粉和蛾类传粉的植物花粉数没有一定规律。蝇类传粉的类群比蛾类传粉的类群胚珠数少, 这可能是由于蝇类携带花粉能力及传粉精确性均较小, 导致植物以减少胚珠数来适应的结果。对同一属内不同生活型植物的花粉数比较, 发现乔木的单花花粉数高于灌木, 灌木的单花花粉数明显高于草本。这可能是由于不同生活型的植株, 其花朵大小不同, 导致花粉数出现明显差别。另外, 通过扫描电子显微镜对花粉形态的观察, 发现蝇类传粉的类群和蛾类传粉的类群间的花粉表面纹饰存在显著差异。     

Systematic significance of pollen arrangement in MICROSPORANGIA OF POACEAE AND Cyperaceaei REVIEW AND OBSERVATIONS ON REPRESENTATIVE TAX A

An anatomical survey of anthers of the Poaceae and Cyperaceae and two related families, the Restionaceae and Flagellariaceae, was conducted to determine the taxonomic distribution and possible phylogenetic significance of pollen arrangement. An unusual pollen arrangement was known from a small number of taxa in the Poaceae and Cyperaceae, in which a single, uniseriate cylinder of pollen grains is arranged in the anther locule such that each grain is in contact with the tapetum (termed here “peripheral” pollen). This contrasts with the prevailing arrangement in other angiosperms in which the locule contains a relatively large number of pollen grains in no special configuration, with many interior grains that never touch the tapetum (termed here “central” pollen). A total of 48 species in these four families was examined in this study, and observations on pollen arrangement for numerous additional species in these families were gleaned from the literature. We confirm that the peripheral arrangement is predominant in Poaceae and Cyperaceae, although both families also include species with central pollen, whereas only central pollen is found in Restionaceae and Flagellariaceae. In the peripheral arrangement, the pore of the pollen grain or pseudomonad, when observed, is in contact with the tapetum, but it has not been definitively demonstrated that this is always the case. Peripheral pollen in the Poaceae and Cyperaceae is nonhomologous because of the presence of pseudomonads in the latter family. It remains unexplained why peripherally arranged pollen or pseudomonads, with the attendant reduction in the number of pollen grains, should be associated with anemophily in these two families.     

POLLEN-OVULE RATIOS AND BREEDING SYSTEM EVOLUTION IN SOLANUM SECTION BASARTHRUM (SOLANACEAE)

Pollen-ovule ratio, pollen grains per flower, pollen grain volume, pollen biomass, ovules per ovary, seeds per fruit, seed volume, seed biomass, and corolla size were studied to reveal patterns of variation within a monophyletic group. Solanum sect. Basarthrum includes 22 neotropical species of two clades, one having small, few-seeded red fruits, and the other having larger, many-seeded green fruits. The former includes self-incompatible species and a dioecious species, while the latter includes self-incompatible species, self-compatible (and autogamous) species, and a domesticate (the “pepino”). Although the pollen quantity and ovule quantity of the self-incompatible species are significantly higher in the green-fruited subgroup than in the red-fruited subgroup, the pollen-ovule ratios of the self-incompatible species are not significantly different between the two subgroups, suggesting parallel evolution of the pollen-ovule ratio. Furthermore, the pollen-ovule ratio tracks the breeding system: self-incompatible species have significantly higher pollen-ovule ratios than self-compatible species, resulting both from fewer pollen grains and more ovules of the latter. The pollen-ovule ratio of the dioecious species is among the highest of all, resulting only from fewer ovules. The pepino is self-compatible but has a pollen-ovule ratio like its wild self-incompatible progenitors and shows wide variation in seed production and pollen quality, but not in pollen quantity. Among all species, pollen size and quantity are inversely proportional, as are seed size and quantity.     

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。     

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.     

Pollen,ovules, and pollination in pea: Success,failure, and resilience in heat

Field pea (Pisum sativum), a major grain legume crop, is autogamous and adapted to temperate climates. The objectives of this study were to investigate effects of high temperature stress on stamen chemical composition, anther dehiscence, pollen viability, pollen interactions with pistil and ovules, and ovule growth and viability. Two cultivars (“CDC Golden” and “CDC Sage”) were exposed to 24/18°C (day/night) continually or to 35/18°C for 4 or 7 days. Heat stress altered stamen chemical composition, with lipid composition of “CDC Sage” being more stable compared with “CDC Golden.” Heat stress reduced pollen viability and the proportion of ovules that received a pollen tube. After 4 days at 35°C, pollen viability in flower buds decreased in “CDC Golden,” but not in “CDC Sage.” After 7 days, partial to full failure of anthers to dehisce resulted in subnormal pollen loads on stigmas. Although growth (ovule size) of fertilized ovules was stimulated by 35°C, heat stress tended to decrease ovule viability. Pollen appears susceptible to stress, but not many grains are needed for successful fertilization. Ovule fertilization and embryos are less susceptible to heat, but further research is warranted to link the exact degree of resilience to stress intensity.     

Contrasting variation within and covariation between gender-related traits in autogamous versus outcrossing species: Alternative evolutionary predictions

We present several predictions concerning the expression of genetic variation in, and covariation among, gender-related traits in perfect-flowered plant taxa with different breeding systems. We start with the inference that the pollen:ovule (P/O) ratio in obligately autogamous species (in which the ovules in a flower are fertilized only by the pollen it produces) should be under much stronger stabilizing selection than in outcrossing taxa. Consequently, we predict that obligately autogamous taxa should exhibit lower genetic coefficients of variation in the P/O ratio. Nevertheless, genetic variation in both pollen and ovule production per flower might persist within autogamous as well as outcrossing populations. In autogamous taxa, genotypes with relatively few pollen grains and ovules per flower (but producing relatively high numbers of flowers) and genotypes with comparatively high numbers of gametes per flower (but producing relatively few flowers) could co-exist if lifetime flower production is selectively neutral. In contrast, in outcrossers, the maintenance of genetic variation in ovule and pollen production per flower might result predominantly from their ability to maintain variation in phenotypic and functional gender. Given genetic variation in primary sexual traits, we predict that the genetic correlation between investment in male and female gametes per flower should qualitatively differ between selfers and outcrossers. We predict a positive genetic correlation between pollen and ovule production per flower in obligately autogamous taxa, primarily because strong stabilizing selection on the P/O ratio should select against the gender specialists that would be necessary to effect a negative genetic correlation between mean pollen and ovule production per flower. Moreover, the fact that autogamous individuals are 50% female and 50% male means that gender-biased phenotypes cannot be functionally gender-biased, preventing selection from favouring phenotypic extremes. In contrast, in outcrossing taxa, in which functionally male- and female-biased genotypes may co-exist, the maintenance of contrasting genders could contribute to the expression of negative genetic correlations between pollen and ovule production per flower. We discuss these and a number of corollary predictions, and we provide a preliminary empirical test of the first prediction. This revised version was published online in July 2006 with corrections to the Cover Date.     

Aperture variation in the pollen of Nelumbo (Nelumbonaceae)

The evolutionary and developmental origin of tricolpate pollen is of great interest because pollen of this kind defines a major clade of angiosperms (eudicots), a clade that is also well supported by molecular data. We examined evidence that tricolpate and monosulcate pollen types are produced alongside each other in the anthers of Nelumbo flowers, as has previously been reported. Observations of pollen in situ within individual anthers revealed mainly tricolpate pollen produced in tetrahedral tetrads, but also a small percentage of clearly aberrant pollen grains that have a great variety of aperture configurations. Previously published evidence for tetragonal tetrads is not supported, and previously reported monosulcate grains are part of a continuum of variation among the aberrant grains in aperture number, position and form. Other eudicots show similar variability in their pollen apertures. The variation in the pollen of Nelumbo is not exceptional, and may not be more significant than variation seen in the other taxa with regard to the origin of the tricolpate and tricolpate‐derived pollen characteristic of eudicots. Nevertheless further studies of aberrant pollen in Nelumbo and other eudicots, together with comparisons of pollen development in “normal” eudicots and closely related species that show radical, and developmentally fixed, reorganization of apertures and pollen polarity, may be helpful in understanding the processes that controlled the transition from the monosulcate to the tricolpate condition.     

Anemophilous plants select pollen from their own species from the air

In wind-tunnel experiments, Niklas (1985) has demonstrated the ability of anemophilous plants to select pollen from their own species from the airstream. However, there have been no field experiments to establish whether this operates in nature. We surveyed the pollents on the stigmas of four different, coextensive, dioecious anemophilous species surrounded by a Pinus radiata plantation. The alien pine pollen was overrepresented relative to background levels on only one of the four species. For all three indigenous species with both male and female plants in the area, the highest proportion — in all cases more than 40% — of the pollen found on their stigmas came from their own species. One indigenous species lacked male plants in the area; consequently, the results from this species are difficult to interpret. However, for all four species, there was at least 15% pine pollen, and some pollen of other indigenous species. These results suggest that there is some pollen selection, but that the mechanisms are may be not as effective as Niklas has suggested.     

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.     

Pollen grain viability in Parietaria judaica L. during the long blooming period and correlation with meteorological conditions and allergic diseases

Abstract

Pollen grain viability and starch presence in pollen were followed during the long blooming period (May-November) of Parietaria judaica, the most widespread pellitory in Italy, responsible for many cases of allergic diseases. Observations were carried out near Siena (Tuscany), in the years 1978 and 1979.

Viability is high in late spring and early autumn, and pollen grains are starchy. The presence of starchless grains is always related to low viability: a production of pollen with a low viability occurs at the beginning and end of blooming, and also in summer, during drought periods. Pollen grain viability varies widely during the blooming period, and its variations are correlated to meteorological conditions, mainly rainfall and temperature.

Allergic diseases due to pellitory, however, are mainly reported between April and July, and on the other hand a small amount of patients states that they suffer all the year round. The discrepancies between the periods of viable pollen production and of declared symptoms by patients, are discussed.     

POLLEN-OVULE RATIOS IN THE CRUCIFERAE

Several recent studies have revealed sources of variation in pollen-ovule ratios that might limit their usefulness as breeding system indicators. In this survey, pollen-ovule ratios were determined for 66 crucifer taxa (Cruciferae) with known breeding systems to establish whether a clear distinction could be made between the pollen-ovule ratios of allogamous and autogamous crucifers and whether this was sufficient to allow a standard to be set for the Cruciferae. Both the mean number of pollen grains per flower and the pollen-ovule ratio were significantly larger in the allogamous taxa than in the autogamous taxa, whereas the mean number of ovules per flower was not significantly different. However, the pollen-ovule ratio was found to be a much better indicator of the breeding system than the mean number of pollen grains by itself, based on the degree of overlap between the values for the allogamous and autogamous taxa. Variation in the amount of pollen produced by different sets of stamens and a correlation between the pollen-ovule ratio and seed size were identified as potential sources of error in calculating and interpreting the pollen-ovule ratios. This survey suggests that the pollen-ovule ratios of allogamous crucifers are typically greater than 3,500, that those of autogamous crucifers are typically less than 1,000, and that these standards can be used for indicating the breeding systems of other species in the Cruciferae.     

Pollen production in anemophilous species of the Poaceae family in Tetouan (NW Morocco)

Total pollen production per inflorescence and per square meter were studied in anemophilous species of the Poaceae family, to determine the relative contribution of each species to the total load of pollen released and to identify the species with the highest potential pollen emission. This was done by calculating the number of pollen grains per flower and per inflorescence and by estimating the density of inflorescences in an area of one square meter. Pollen production per inflorescence varied between 12,000 and 15 million grains, often being higher in the perennial species. Mountainous, grassland, riverside, and littoral areas contributed similarly to the amount of pollen grain production per square meter. Species such as Gaudinia fragilis, Hordeum murinum, Lolium rigidum, and Trisetaria panicea, with high pollen records, were found in all the studied areas. Aerobiological analysis will be undertaken to determine the main pollen season of Poaceae species. The results obtained are of great importance for respiratory health management in this region.     

FLORAL PARAMETERS,BREEDING SYSTEM AND POLLINATOR TYPE IN TRICHOSTEMA (LABIATAE)

Diversity in flower size and degree of exsertion of anthers and stigma from the corolla in the California species of Trichostema appear to be correlated with breeding system and pollinator type (bee vs. bird). Autogamous (self-pollinating) species unlike xenogamous (cross-pollinating) species lack spatially separate anthers and stigma and nototribic pollination. The outcrossing species have significantly larger flowers, significantly larger nectar volumes and significantly higher pollen-ovule ratios than do selfing species. Because autogamous species are less reliant on flower visitors to facilitate pollination, there may be relaxed selection for large nectar producing flowers. Pollen-ovule ratios are correlated with breeding system and reflect the efficiency of pollen transport. Data on floral parameters suggest xenogamous species expend more energy per floral unit and are less efficient seed producers than closely related autogamous species.