Pollen histochemistry and pollen : ovule ratios in Zingiberaceae

BACKGROUND AND AIMS: Pollen grains of 37 species from 11 genera in the family Zingiberaceae were examined to assess qualitatively starch or lipid contents; the pollen grain and ovule numbers per flower and pollen : ovule ratios were also counted and calculated. Pollen : ovule ratios were studied to reveal patterns of variation in the Zingiberaceae. METHODS: Freshly open flowers with dehiscing anthers were collected at random from plants growing in natural habitats or in botanical gardens. Presence of lipids or starch in pollen grains was tested by Sudan solution and IKI solution, respectively, and examined under a microscope. To estimate the pollen and ovule numbers per flower, one anther from each bud was carefully dissected and all pollen grains were counted; ovaries were carefully dissected out of each flower and counted. Whenever possible, at least 10-15 buds were used in the determination. KEY RESULTS: Thirty-three of all the 37 species examined had starchy pollen. Starch was not found in only four species and lipid was not found in only one species; among the four tribes in subfamily Zingiberoideae, all species of Zingibereae and Globbeae had pollen with no starch, Alpineae and Hedychieae had pollen with and without starch, whereas, all species of subfamily Costoideae had starchy pollen with abundant lipids. The mean pollen : ovule ratios in the members of the Zingiberaceae investigated range from 3.25 +/- 1.56 to 616.52 +/- 117.83. CONCLUSIONS: The pollen nutrition types seemed not related to mating systems. The pollen : ovule ratios in members of the Zingiberaceae with the same breeding system are noticeably lower than those recorded by previous authors. The lower pollen : ovule ratios in this family are presumed to be related to the highly efficient pollination systems, mediated by pollen which can be quite glutinous and the relatively large stigma area. In most of the Alpinia species the anaflexistylous flowers have much larger numbers of pollen grains and higher pollen : ovule ratios than the cataflexistylous flowers. There are significant differences in mean pollen grain numbers and pollen : ovule ratios between different life forms but ovule numbers are approximately the same.     

东北杜鹃花属植物花粉形态的研究

利用光学显微镜和扫描电子显微镜对东北杜鹃花属(Rhododendron Linn.)6种, 1变种, 1变型的花粉形态进行了观察和比较研究。该属花粉粒均为四合花粉。四合花粉呈正四面体排列。花粉粒近球形。每粒花粉均具3沟,在四合花粉粒上,相邻花粉粒上的沟相接,有些种可见沟内的萌发孔。光镜下可见花粉外壁为内外两层,厚度约相等。不同的种在花粉粒大小上有区别。在扫描电镜下观察可见不同的种花粉表面具有不同的纹饰。即该属在花粉形态上具有种的特性。     

东北杜鹃花科4属植物花粉形态的研究

利用光学显微镜和扫描电子显微镜对东北杜鹃花科甸杜属(Chamaedaphne Moench.)、杜香属(Ledum Linn.)、松毛翠属(Phyllodoce Salisb.)、越橘属(Vaccinium Linn.)4属6种, 1变种的花粉形态进行了观察和比较研究。以上4属花粉粒均为四合花粉。四合花粉呈正四面体排列。花粉粒近球形。每粒花粉均具3沟,在四合花粉粒上,相邻花粉粒上的沟相接,有些种可见沟内的萌发孔。光镜下可见花粉外壁为内外两层,厚度约相等。不同属种的花粉粒在大小上有区别,花粉粒平均直径为34.0~43.7μm。在扫描电镜下观察可见不同的种花粉表面具有不同的纹饰。为分类学提供了孢粉学方面的证据。     

Pollen of Ceratostema (Ericaceae, Vaccinieae): tetrads without septa

The pollen morphology of two species of the Neotropical genus Ceratostema (Ericaceae) was examined by light, scanning and transmission electron microscopy. The Ceratostema species examined have 3-colporate pollen grains united in permanent tetrahedral tetrads that show a common condition encountered in the Ericaceae. But the septal exine was absent between two neighboring grains in each pollen tetrad of Ceratostema. The pollen tetrads without septa are the first report for the Ericaceae as well as other angiosperm families.     

Beyond pollen:ovule ratios: Evolutionary consequences of pollinator dependence and pollination efficiency for pollen and ovule production in angiosperms

Premise

The relative per-flower production of ovules and pollen varies broadly with angiosperm mating systems, with outcrossing types commonly producing more pollen grains per ovule than selfing types. The evolutionary causes of this variation are contentious, especially the relevance of pollination risk. Resolution of this debate may have been hampered by its focus on pollen:ovule (P:O) ratios rather than on the evolution of pollen and ovule numbers per se.

Methods

Using published mean ovule and pollen counts, we analyzed associations with the proportion of removed pollen that reaches stigmas (pollen-transfer efficiency) and differences between pollinator-dependent and autogamous forms within and among species. Analyses involved Bayesian methods that simultaneously considered variation in pollen and ovule numbers and accounted for phylogenetic relatedness. We also assessed the utility of P:O ratios as mating-system proxies and their association with female outcrossing rates.

Results

Median pollen number declined consistently with pollen-transfer efficiency among species, whereas median ovule number did not. Similarly, in both intraspecific and interspecific analyses, pollinator-dependent plants produced more pollen than autogamous plants, whereas ovule production did not differ statistically. Distributions of P:O ratios overlapped extensively for self-incompatible and self-compatible species and for different mating-system classes, and P:O ratios correlated weakly with outcrossing rate.

Conclusions

Our findings demonstrate that pollinator dependence and pollination efficiency commonly influence the evolution of pollen number per flower but have more limited effects on ovule number. P:O ratios provide ambiguous, possibly misleading, information about mating systems, especially when compared among clades.

     

Breeding systems in Balsaminaceae in relation to pollen/ovule ratio,pollination syndromes,life history and climate zone

• Pollen/ovule (P/O) ratios are often used as proxy for breeding systems. Here, we investigate the relations between breeding systems and P/O ratios, pollination syndromes, life history and climate zone in Balsaminaceae.
• We conducted controlled breeding system experiments (autonomous and active self‐pollination and outcrossing tests) for 65 Balsaminaceae species, analysed pollen grain and ovule numbers and evaluated the results in combination with data on pollination syndrome, life history and climate zone on a phylogenetic basis.
• Based on fruit set, we assigned three breeding systems: autogamy, self‐compatibility and self‐incompatibility. Self‐pollination led to lower fruit set than outcrossing. We neither found significant P/O differences between breeding systems nor between pollination syndromes. However, the numbers of pollen grains and ovules per flower were significantly lower in autogamous species, but pollen grain and ovule numbers did not differ between most pollination syndromes. Finally, we found no relation between breeding system and climate zone, but a relation between climate zone and life history.
• In Balsaminaceae reproductive traits can change under resource or pollinator limitation, leading to the evolution of autogamy, but are evolutionary rather constant and not under strong selection pressure by pollinator guild and geographic range changes. Colonisation of temperate regions, however, is correlated with transitions towards annual life history. Pollen/ovule‐ratios, commonly accepted as good indicators of breeding system, have a low predictive value in Balsaminaceae. In the absence of experimental data on breeding system, additional floral traits (overall pollen grain and ovule number, traits of floral morphology) may be used as proxies.

     

Study on the Pollen Morphology of the Family Berberidaceae

The pollen grains of 33 species representing 11 genera of the family Berberidaceae, mostly from China, were examined with the light microscope and scanning electron microscope. Their characteritic details can be used for generic diagnosis. A pollen key to the genera based on these observations is presented. Based on the morphology, the pollen grains can be grouped into the following three types: 1. The tetrad pollen type found only in the genus Sinopodophyllum. 2. The spiraperturate pollen type found in the genera Berberis and Mahonia. 3. The tricolpate pollen type found in the genera Diphylleia, Jeffersonia, Nandina, Dysosma, Caulophyllum, Leontice and Epimedium. A diagnostic key to the pollen grains of genera in the Berberidaceae. 1. Pollen grains single 2. Pollen grains spiraperturate .................... Berberis L., Mahonia L. 2. Pollen grains tricolpate 3. Exine with spinose sculpture ........................ Diphylleia Michx. 3. Exine with non-spinose sculpture 4. Exine with striate or striate-reticulate sculpture ...... Jeffersonia Barton 4. Exine with reticulate sculpture 5. Exine around colpus with distinct thickening ........ Nandina Thunb. 5. Exine around colpus without thickening 6. Pollen grains larger (45—50)×(32.5—37.5)μ 7. Colpus with membrane ................ Dysosma R. E. Woodson 7. Colpus without membrane ................ Caulophyllum Michx. 6. Pollen grains smaller (25—550)×(20—527.5) μ 7. Pollen grains prolate-perprolate .................... Leontice L. 7. Pollen grains spheroidal-prolate...Epimedium L., Podophyllum L. 1. Pollen grains tetrad ............................ Sinopodophyllum Ying     

Relationships of pollen size,pistil length and pollen tube growth rates in Rhododendron and their influence on hybridization

Summary Pollen size and pistil length data have been collected for 93 species of Rhododendron (Ericaceae) belonging to a number of different subgeneric taxa. For a sample of eight species in section Vireya, pollen tube growth in the style after selfor interspecific pollination has been quantified. Pollen volume and the time taken for pollen tubes to reach the ovary were both related to pistil length. Pollen-tube growth rates were generally greater for species with longer pistils and larger pollen. Increasing temperature increased the rate of pollen-tube growth. There was no detectable effect of pollen tube density on tube growth rate in the style. After interspecific pollinations tube growth rates in foreign styles could be faster or slower than in self styles. A semisterile individual with two viable pollen grains per tetrad and a plant grafted as scion to a longer-styled stock both showed more rapid pollen-tube growth than expected on the basis of pistil size. Data collected for 26 species in section Vireya showed that where extreme disparity of pollen/pistil size causes failure of interspecific crosses, one or more bridging species with intermediate pollen/pistil size can generally be selected.     

Pollen morphology in delimiting subgenera and species of the genus <Emphasis Type="Italic">Cybianthus</Emphasis> s.l. (Myrsinoideae–Primulaceae)

Pollen grains of species of the subgenera of Cybianthus s.l. were analyzed in order to determine if pollen morphology is consistent with the currently accepted infrageneric classification, and if it can provide characters useful for distinguishing species or groups of species. Acetolyzed pollen grains were analyzed, measured, described and photomicrographed, while non-acetolyzed pollen grains were analyzed using scanning electron microscopy. The pollen grains of all species investigated were isopolar monads, small- to medium-sized, subprolate or prolate, 3-colporate, and rugulate or rugulate-perforate. The different analytical methods (statistical analyses, exploratory analysis of the data and cluster analysis) used demonstrated that pollen morphology is informative in delimiting this genus. However, the pollen characters are generally quite uniform among the subgenres of Cybianthus s.l. Thus, the use of the pollen attributes in the separation of the subordinate species to different subgenera is not effective. Pollen morphology did not give support to current subgeneric systematics, but it did support the genus Cybianthus s.l.     

Pollen tube reallocation in two preanthesis cleistogamous species, Ranalisma rostratum and Sagittaria guyanensis ssp. lappula (Alismataceae)

Pollen deposition on stigmas and pollen tube growth in two apocarpous species, Ranalisma rostratum and Sagittaria guyanensis ssp. lappula (Alismataceae), were examined with fluorescence microscopy. The reallocation of pollen tubes among pistils was observed in both species. The percentage of pollinated stigmas per flower was only 22.0% in R. rostratum and 51.0% in S. guyanensis, though the seed/ovule ratios are higher than 65% in both species. The number of pollen grains on each single stigma ranged from 0 to 96 in R. rostratum, and from 0 to 125 in S. guyanensis. When more than one pollen grain deposited on a stigma, all pollen tubes grew to the ovary, but only one of them turned towards the ovule and finally entered the nucleus. The other tubes grew through the receptacle tissue into ovules of adjacent carpels whose stigmas were unpollinated or pollinated later. The intercarpellary growth of pollen tubes could be a mechanism to increase the efficiency of sexual reproduction in an apocarpous gynoecium with low pollination on the pistils.     

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。     

Pollen heteromorphism is pervasive in <Emphasis Type="Italic">Thalictrum</Emphasis> (Ranunculaceae)

Among the angiosperms, features of pollen morphology such as grain size, aperture number and surface ornamentation display striking variation. It is less well appreciated that pollen morphology may vary within and among populations of the same species as well as within individual plants. In some species, individual plants produce multiple types of fertile pollen grains (called pollen heteromorphism). Aspects of pollen morphology, such as aperture number, are likely to affect fertilization success with different morphologies favored in different local competitive and ecological environments. This study surveys variation in pollen grain morphology among species throughout the genus Thalictrum. Pollen from individuals of 36 species was rehydrated from herbarium specimens, and light microscopy was used to quantify pollen grain aperture number and size. I find that pollen aperture-number heteromorphism is present within all Thalictrum species studied, and distributions of aperture-number morphs vary both within and among species. This study provides an example of significant pollen heteromorphism within a genus that also varies widely for pollination mode and sexual system.     

Pistil anatomy and pollen tube development in Polygala vayredae Costa (Polygalaceae)

Low seed ovule ratios have been observed in natural populations of Polygala vayredae Costa, a narrowly endemic species from the oriental pre-Pyrenees. To evaluate physical and nutritional constraints and pollen tube attrition in this endemic species, stigma and style anatomy, as well as pollen tube development along the pistil were investigated using light and fluorescence microscopy. The structural morphology of the stigmatic region was also examined with scanning electron microscopy. Pollen grains that reached the stigmatic papillae came into contact with a lipid-rich exudate and germinated easily. Although a large number of pollen grains reach the stigmatic papillae, few pollen tubes were able to grow into the style towards the ovary. The style was hollow, with the stylar channel beginning a few cells below the stigmatic papillae. Initially, the stylar channel area was small compared to other levels of the style, and was surrounded by lipid-rich, highly metabolic active cells. Furthermore, lipid-rich mucilage was detected inside the stylar channel. At subsequent style levels towards the ovary, no major reserves were detected histochemically. The reduced intercellular spaces below the stigmatic papillae and the reduced area of the stylar channel at its commencement are suggested to physically constrain the number of pollen tubes that can develop. In subsequent levels of the style, the stylar channel could physically support a larger number of pollen tubes, but the lack of nutritional reserves cannot be disregarded as a cause of pollen tube attrition. Finally, the number of pollen tubes entering the ovary was greater than the number of ovules, suggesting that interactions occurring at this level play a major role in the final reproductive outcome in this species.     

Reproductive ecology in the endemic Primula apennina Widmer (Primulaceae)

Abstract

Disassortative mating in distylous self-incompatible species should result in the equilibrium of morph types in natural populations. Deviation from isoplethy may affect pollen transfer, and in isolated populations it could lead to Allee effect and genetic drift. Pollen limitation has been found to occur in several distylous species, for which mating opportunities are actually reduced to half population. In this study, we investigated the reproductive features and pollination ecology of the narrow endemic Primula apennina. We recorded equilibrium of morph frequencies in the studied population, reflecting the comparable fecundity found in the two morphs. Long-styled flowers produce more pollen grains of smaller size than short-styled ones: we hypothesize that in thrum flowers, pollen is more easily removed by the insect pollinator Macroglossum stellatarum, resulting in equal pollen amounts carried to both short styles and long styles. This lower pollen transfer efficiency from long-styled to short-styled flowers is also reflected in legitimate pollen–ovule ratio values. Despite results show no evidence of imminent threats to population persistence at study site, the strict dependence on one or very few pollinator species, and ecological traits, may increase extinction risks in the long-term period.     

Pollen morphology of the genus Euphrasia L. (Orobanchaceae) from Pakistan and Kashmir and its taxonomic significance

Pollen morphology of 25 species of the genus Euphrasia L., belonging to the family Orobanchaceae from Pakistan has been examined by light and scanning electron microscope. It is a stenopalynous genus. Pollen grains are usually radially symmetrical, isopolar, tricolpate, sub-prolate to prolate-spheroidal rarely oblate-spheroidal. Sexine slightly thinner than nexine or as thick as nexine. Tectum coarse to fine retipilate or rugulate–retipilate. On the basis of exine ornamentation three distinct pollen types viz., Euphrasia foliosa, Euphrasia multiflora, and Euphrasia incisa are recognized. Pollen morphology is significantly helpful at specific level within the pollen type.     

12种十字花科植物花粉形态研究

采用扫描电子显微镜对十字花科8属12种植物的花粉形态特征进行了观察分析.其中8种为首次报道.观察结果表明:4族8属12种十字花科植物花粉形态的一般特征为:花粉粒长球形至超长球形;极面观三裂圆形,少数四裂圆形;外壁纹饰为网状或细网状纹饰;三沟,偶四沟.12种植物的花粉可划分为5种类型:(1)花粉粒长球形,网状纹饰,三沟;(2)花粉粒超长球形,网状纹饰,三沟;(3)花粉粒长球形,细网状纹饰,三沟;(4)花粉粒超长球形,细网状纹饰,三沟;(5)花粉粒长球形,三或四沟,网状纹饰.十字花科花粉形态特征在不同族间、不同属间以及同属不同种间均没有明显规律,在十字花科分类过程中,不宜单独以花粉形态特征作为依据.     

Pollen morphology of the genus Nothofagus and its taxonomic significance

The present paper describes the pollen morphology of 26 species and 2 varieties in Nothofagus from Australia, New Zealand, New Caledonia, New Guinea and South America. Pollen grains were all examined with light microscope (LM), scanning electron microscope ( SEM ) and transmission electron microscope(TEM). A comparative analysis of pollen exine ultrastructure was made for some species. The results are summarised as follows: Pollen grains are oblate to peroblate, 5～8-short-colpate, rarely 4- or 9-colpate; colpi generally thickened at margins; pollen surface spinulose. The exine ultrastructure of Nothofagus differs considerably from that of the other genera in the Fagaceae. The pollen grains of the species examined here show great differences in shape, size, colpal number and characteristics of colpi at margins and could be divided into three distinct types, i.e. N. brassii type; N. menziesii type and N. fusca type.     

Pollen morphology of some selected plant species from Horton Plains,Sri Lanka

The pollen morphology of four species: Gaultheria leschenaultii (Ericaceae), Psychotria zeylanica (Rubiaceae), Rubus leucocarpus (Rosaceae) and Symplocos bractealis (Symplocaceae) from Horton Plains, Sri Lanka was investigated using LM and SEM. Psychotria zeylanica is endemic to Sri Lanka and the other three taxa are common in Horton Plains. Gaultheria leschenaultii has tetrads with 3-colporate, coarsely granular to granulo-rugulate pollen grains; in the polar region and at the colpus margin the exine is smooth with micro-rugulate structure. Psychotria zeylanica has 3- (or 4-) colporate, reticulate-spinulose to granular pollen grains. Rubus leucocarpus has 3-colporate, striate-perforate pollen grains and Symplocos bractealis has 3- (or 4-) colporate pollen grains with a verrucate-perforate exine. Pollen grains of these four species are very indicative of human impact and palaeoclimate.     

Embryological studies of Oxalis debilis Kunth

The study of pollen and female gametophyte development of Oxalis debilis was carried out to elucidate the reasons for the absence of seed production in this species. The formation of an incomplete callosic wall separating the microspores of the tetrad was observed; therefore, cytoplasm connections were present between the microspores. Microspores of different sizes and with different grades of vacuolation were observed in the mature anther. Only few microspores divide mitotically and form the generative and vegetative cell. The cytoplasm of the vegetative cell may accumulate different substances in reserve. Pollen grains are 8–12?colpate, and the morphology and electron density of the exine varies. Because flowers in full anthesis have all ovules at the megasporocyte stage with the nucellar epidermis intact, fecundation could not occur. The production of only a few viable pollen grains and the absence of successful megagametophyte development prevent fertilization and account for the absence of fruits and seeds formation.     

Mating systems in two species ofPotentilla from Alaska

Aspects of reproductive ecology have been studied in two species ofPotentilla (Rosaceae) from interior Alaska. Data were collected to determine mating system traits such as seed: ovule and pollen: ovule ratios, and experiments were carried out in the field in order to study the effect of caging, emasculation, and manual self-pollination on seed set and seed quality. Mating system strategies differ between the two species in the populations examined. High pollen: ovule ratio, high seed: ovule ratio, good autodeposition ability, and unpredictable pollination in concert with self-incompatibility indicate thatP. hookeriana in the population studied has adopted a more or less pseudogamous mating system. In contrast, the data suggest thatP. uniflora in the studied population is approaching a sexual, facultatively outbreeding species. Because both species exhibit ploidy variation, a trait common among agamospermous plants, it is concluded that both species are likely to be facultatively agamospermous but that sexual reproduction is much more important in the population ofP. uniflora than it is inP. hookeriana.