A numerical approach to pollen sculpture terminology

We present a key synthesizing pollen-sculpture terminology for grains having perforations or lumina of any size and spatial distribution. The key uses quantitative criteria to distinguish five non-overlapping qualitative terms (reticulate, microreticulate, foveolate, scrobiculate, and punctate). In addition, the range of quantitative variation encompassed by these qualitative terms is highlighted, including ways to express and compare that variation using computerized image analysis of SEM micrographs. Finally, a mathematical analysis combined with image-analysis measurements is used to explore the relationships between the terms outlined above and two closely related terms, tectate and semitectate.     

The classification and evolution of pollen types ofPlanchonioideae (Lecythidaceae)

The pollen type ofPlanchonioideae is unique among the angiosperms in its syntricolpatism associated with the marginal ridge, marginal groove, and polar cushion; it has been studied byJ. Muller, who classified it into three types, mainly with a light microscope. In the present study, pollen grains of the six constitutent genera were studied with a scanning electron microscope, five additional pollen types were recognized. Of the eight pollen types the one with sealed polar ectoapertures and weakly developed marginal ridges, without marginal grooves and polar cushions is regarded as the most primitive. Evolutionary trends proceeded with the opening of the polar ectoapertures, the complication of the polar cushions and marginal grooves. In the most advanced form, the marginal ridges and marginal grooves coalesced to form two concentric rings, whereas the polar cushions were secondarily reduced.     

Sporoderm ultrastructure ofLophosoria andCyatheacidites (Filicopsida): Systematic and evolutionary implications

The distinctive spores produced byLophosoria, an extant monotypic fern, are examined ultrastructurally and correlated with the sporoderm of fossilCyatheacidites. The morphological and ultrastructural similarity of the two taxa provide additional information that can be used to trace the evolutionary history of this spore type.     

Intraspecific polymorphism of pollen sculpture inParis tetraphylla A. Gray (Liliaceae)

Paris tetraphylla has a greater diversity of pollen exine sculpture than was previously thought. On the basis of 357 plants from 52 localities of Japan, we distinguished seven sculpture types that are tentatively referred to here as G₁, G₂, G₃, R₁, R₂, R₃ and S. G₁, G₂ and G₃ have gemmate sculpture in which the gemmae are characteristically large, medium and small in size, respectively; likewise, R₁, R₂ and R₃, have coarsely to finely ruglate sculpture; and S, scrobiculate sculpture. In general, any one of the seven types always or predominantly occurs in each plant, thus allowing us to make comparisons among plants and further among local populations. We thus found that G₂ and G₃, respectively, are the most and the second most widespread and occur in plants of 41 and 22 respectively, of the localities examined, although their frequencies differ with locality. Generally R₃ occurs in plants distributed at higher latitudes, while R₂ is rather widespread and common to plants growing on high mountains. G₁ and R₁ are restricted to plants occurring south of Ibaraki Pref. and north of Gunma Pref., respectively. Such localized distributional patterns of some types may reflect the reproductive nature and the history of geographical isolation inParis tetraphylla over the Japanese Islands.     

A functional interpretation ofLactuceae (Compositae) pollen

Pollen morphology and ultrastructure inLactuceae pollen is considered in relation to the accomodation of volume changes, pollination biology and exine-held substances. Echinate pollen grains, such as those ofCatananche, are shown to accomodate volume changes by folding along the colpi and possibly by volume changes in the cavea. The different patterns of echinolophate pollen respond in different ways. Folding along the colpi is important inScorzonera andTragopogon and to a limited extent inCichorium andEpilasia whilst inScolymus the colpi are almost immobilized. Movements of the lacunar floors take over the harmomegathic function to compensate for lack of colpus mobility. Bulging of the intine at the apertures and changes in the size of the cavea may account for part of the volume change accomodated in any pollen type. Echinolophate pollen is interpreted as being a superior means of regulating volume changes with the most economical and mechanically efficient use of wall material which has evolved independently in several tribes ofCompositae.     

Invasive tapetum and tricelled pollen inAmbrosia trifida (Asteraceae,tribeHeliantheae)

Staminate flowers of giant ragweed,Ambrosia trifida L. (Asteraceae, tribeHeliantheae, subtribeAmbrosiinae) were processed into resin and sectioned 1–2 µm thick. The invasive (amoeboid) anther tapetum remains parietal until microspores are released from tetrads, then it swells and invades the locule, merging gradually into a single protoplast that flows among the microspores. After the tapetal membrane ruptures at late microspore stage, tapetal debris fills the locule, then disappears as pollen matures. Pollen becomes tricelled before anthesis. The two sperm cell nuclei are slender and wormlike. The present report supports the two generalizations that invasive tapetum and tricelled pollen are attributes of theAsteraceae.     

Significance of palynology for phylogeny ofAnnonaceae: Experiments with removal of pollen characters

Phylogenetic analyses based on morphology placeAnaxagorea and other taxa with granular monosulcate pollen, as in otherMagnoliales, at the base ofAnnonaceae. Taxa with columellar tetrads, granular tetrads, and inaperturate monads form a derived clade. To test the systematic importance of palynology, we analyzed the data set with pollen characters removed. The result was lower resolution and a different rooting of the family, betweenUvariopsis and other groups with columellar tetrads.Anaxagorea and other monosulcates are higher in the tree, implying that granular monosulcate pollen, laminar stamens, and irregular endosperm ruminations are reversals. This rooting is highly unparsimonious when pollen characters are included, and only weakly supported over theAnaxagorea rooting when pollen is excluded. Together with preliminary molecular analyses, these experiments confirm the special value of palynology in systematics ofAnnonaceae. This paper is dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer on the occasion of his 70th birthday     

A comparison of the pollen wall ultrastructure of aureoid and non-aureoidSenecio species (Asteraceae) in North America

The structure of the pollen wall as revealed by transmission electron microscopy is presented for 29 speciesSenecio including 19 aureoid and ten non-aureoid species. The previous report of a helianthoid wall structure (exine with internal foramina) is confirmed for all the aureoid species examined while all of the non-aureoid taxa showed a senecioid pattern. The data provide support for the treatment of the aureoid complex as a distinct taxon.     

Exine ultrastructure of in situ peltasperm pollen from the Rhaetian of Germany and its implications

Morphology and exine ultrastructure of pollen grains of Triassic peltasperms have been studied for the first time. Pollen grains of Antevsia zeilleri from the Rhaetian of Germany are of the Cycadopites-type and monosulcate; the sculpturing is the same in the apertural and non-apertural areas. The proximal exine includes a row of lacunae covered by a solid, thick tectum and underlined by a foot layer. Pillars are hanging from the tectum between the lacunae. The exine is thinning to a homogeneous layer in the apertural region. The latter is bordered by thicker alveolate areas of the exine, in places resembling a saccus-like ultrastructure. The endexine includes white-line-centred lamellae. The exine ultrastructure is compared with that of pollen of Permian peltasperms. Although pollen types ascribed to Permian peltasperms are completely different in their general morphology, a transformation can be hypothesized by ultrastructural data from Permian Vesicaspora into Triassic Cycadopites extracted from pollen sacs of Antevsia. Comparison with Cycadopites of non-peltaspermalean (Ginkgoalean, Cycadophyte) and unknown affinities has been accomplished. The exine ultrastructure is distinctive enough to differentiate among peltaspermalean, cycadalean and bennettitalean Cycadopites; some ultrastructural features are shared with pollen of modern Ginkgo biloba. More ultrastructural data are needed as well as numerous sections of pollen grains are necessary to reveal original unchanged ultrastructure.     

凤仙花属棒凤仙亚属(凤仙花科)植物花粉形态的分类学意义

棒凤仙亚属代表着凤仙花属最先分化出的一支,以多年生草本,多于5朵花的总状花序,侧生萼片4枚,3沟花粉,心皮四室且每室具1枚种子,果实棒锤状,种子椭圆形以及三沟花粉其极面观为三角形而区别于其它种类。该亚属主要分布于中国南部,少数种类辐射分布至华中和中南半岛地区。已有报道证实花粉的形态特征对凤仙花属植物的属下分类具有重要意义,但对棒凤仙亚属的种类少有涉及。为了进一步探讨该亚属花粉的形态特征及其分类学价值,该研究应用扫描电镜技术对国产棒凤仙亚属植物的14种以及凤仙花亚属的窄萼凤仙花的花粉进行了观察。结果表明：棒凤仙亚属的花粉以单粒存在,具3条萌发沟,辐射对称,极面观为三角形或三角状圆形,花粉粒外壁具均匀的网状纹饰,网脊平滑或具波状边缘,网眼中具不同密度的颗粒状突起。棒凤仙亚属的花粉形态与水角属的花粉形态相似,表现出了其原始性;其花粉形态特征与花部形态性状等相关性差,对该亚属的组级分类意义较小,但对于该亚属的种间界定具有重要意义;而其花粉的极面观形状、赤道面观形状以及网状纹饰等变异相对稳定,对于界定该亚属具有重要的分类学价值。     

New views of tapetum ultrastructure and pollen exine development in Arabidopsis thaliana

Background and Aims

The Arabidopsis thaliana pollen cell wall is a complex structure consisting of an outer sporopollenin framework and lipid-rich coat, as well as an inner cellulosic wall. Although mutant analysis has been a useful tool to study pollen cell walls, the ultrastructure of the arabidopsis anther has proved to be challenging to preserve for electron microscopy.

Methods

In this work, high-pressure freezing/freeze substitution and transmission electron microscopy were used to examine the sequence of developmental events in the anther that lead to sporopollenin deposition to form the exine and the dramatic differentiation and death of the tapetum, which produces the pollen coat.

Key Results

Cryo-fixation revealed a new view of the interplay between sporophytic anther tissues and gametophytic microspores over the course of pollen development, especially with respect to the intact microspore/pollen wall and the continuous tapetum epithelium. These data reveal the ultrastructure of tapetosomes and elaioplasts, highly specialized tapetum organelles that accumulate pollen coat components. The tapetum and middle layer of the anther also remain intact into the tricellular pollen and late uninucleate microspore stages, respectively.

Conclusions

This high-quality structural information, interpreted in the context of recent functional studies, provides the groundwork for future mutant studies where tapetum and microspore ultrastructure is assessed.     

Molecular phylogeny ofIva (Asteraceae,Heliantheae) based on chloroplast DNA restriction site variation

Iva s.str. (comprising ten species) was examined by cpDNA restriction site variation to determine phyletic relationships within the group. The results were compared with relationships proposed from other data. A total of 86 restriction site mutations was detected, 47 of which proved phylogenetically informative. A single most parsimonious tree was obtained using both Wagner and Dollo parsimony. The tree revealed three main lineages that are congruent with the three chromosome lineages (base numbers of x = 16, 17, 18). The monophyly of the x = 16 and 18 groups was supported strongly by molecular data, while the monophyly of x = 17 lineage was only supported moderately. Relationships among the three lineages indicate that the sect.Iva is paraphyletic because sect.Linearbractea is nested within it. Both morphological data and the secondary chemical data are in agreement with the proposed cpDNA phylogeny. Because of this agreement, sect.Iva is revised such that,I. axillaris was excluded and positioned within the newly proposed sect.Rhizoma. Patterns and rates of cpDNA evolution were also examined. The results indicated an uneven evolution in the chloroplast genome with different rates of cpDNA evolution in at least a few species ofIva. However, the evolutionary clock hypothesis can not be rejected within most of the lineages inIva.     

Studies on the ultrastructure and taxonomy of the genusTetraselmis (Prasinophyceae)

Comparative ultrastructural investigations on many isolates ofTetraselmis from Japan and the Pacific coast of North America, and on cultures from the Culture Centre of Algae and Protozoa, Cambridge, England, have revealed that the species have characteristic fine structural features of the pyrenoid. Using the pyrenoid structure as a basic character it is proposed that the genus be subdivided into four subgenera,Tetraselmis, Prasinocladia, Tetrathele andParviselmis. In the present paper, species of the subgenusTetraselmis, includingT. cordiformis, T. ascus, T. convolutae andT. astigmatica sp. nov., are described in detail.     

New details on the morphology of fossil onagraceous pollen grains

Fossil onagraceous pollen grains from two Upper Miocene localities in E. Austria were investigated by LM and EM. Exine structure and sculpture as well as viscin threads suggest affinities with the extant genusCircea.     

Studies on the ultrastructure and taxonomy of the genusTetraselmis (Prasinophyceae)

Ultrastructural investigations on many isolates ofTetraselmis have revealed that the species have characteristic fine structural features of the pyrenoid and it was proposed (Horiet al., 1982) that the genus be subdivided into four subgenera. In the present study of this series, species of the subgenusPrasinocladia, includingTetraselmis marina, T. verrucosa andT. verrucosa f.rubens, are described in detail.     

Comparative pollen morphology and ultrastructure of the Callitrichaceae

The Callitrichaceae are an aquatic family of dicots that include the single, geographically cosmopolitan genus Callitriche. Callitriche contains 40-50 terrestrial, amphibious, and obligately submersed species, and it is the only known genus in the plant kingdom with co-occurring aerial and hydrophilous pollination syndromes. Pollen morphology and ultrastructure were described for 13 Callitriche species using scanning electron and transmission electron microscopy. Representative taxa of each growth form were examined; these included three terrestrial species (C. deflexa, C. peploides, and C. nuttallii), nine amphibious species (C. brutia, C. cophocarpa, C. cophocarpa-stagnalis hybrid, C. cribrosa, C. hamulata, C. heterophylla var. heterophylla, C. lusitanica, C. marginata, and C. trochlearis), and one obligately submersed species (C. truncata). Of the amphibious taxa, C. heterophylla var. heterophylla and C. trochlearis had internal geitonogamy, a type of internal self-fertilization. Pollen from all taxa was spheroidal, small, intectate, and lacked well-defined apertures. Taxa primarily differed with respect to exine thickness, surface ornamentation, and the presence or absence of aperture-like regions. The pollen of terrestrial species, as well as that of C. marginata, had well-developed exines with thick sculptured and basal layers. In general, amphibious taxa produced pollen with distinct, but thinner, exines than that of terrestrial taxa. Pollen of the amphibious taxa with internal geitonogamy had a thicker basal layer than species without internal geitonogamy, whereas the overall exine was reduced in C. hamulata and absent in C. brutia and C. lusitanica. Pollen of the obligately submersed C. truncata also lacked an exine. These palynological data were correlated with growth habits and related pollination biologies, as well as with phylogenetic interpretations of Callitrichaceae. Exine reduction or loss has evolved at least twice in the family, and it is associated with aneuploid reduction in chromosome number.     

Pollen grain morphology ofCoris (Primulaceae)

Pollen grain morphology, sculpturing, and wall ultrastructure are investigated in two species ofCoris (Primulaceae),C. monspeliensis L. andC. hispanica Lange. The study includes both acetolysed and unacetolysed pollen. No evidence of any major palynological difference is recorded between these two species, apart from a somewhat larger pollen inC. monspeliensis. However,Coris can be distinguished from the remaining members of thePrimulaceae by the conjunction of relatively large pollen grains, prominent margo, and particular tectal pattern causing a reticulate surface with minute luminal perforations decreasing towards the colpi. From both these distinctive features, and others typically primulaceous, some evolutionary considerations are inferred. Finally, the higher proportion of irregular grains inC. hispanica is interpreted in light of environmental stress.     

The remarkable genus<Emphasis Type="Italic"> Coptosapelta</Emphasis> (Rubiaceae): pollen and orbicule morphology and systematic implications

The pollen morphology and distribution of orbicules were investigated in ten species of the genus Coptosapelta (Rubiaceae) using light, scanning electron and transmission electron microscopy. In general, Coptosapelta pollen is three- to five-pororate and suboblate to oblate-spheroidal. The sexine ornamentation varies from psilate to reticulate, and the pollen-wall stratification lacks columellae. Droplets are observed on the inner nexine surface. Distinct orbicules are found in five species. Orbicules are often lobed and appear to be aggregated and embedded in the tapetal remnants of the locule wall. Ultrastructurally, a single, flattened core is frequently observed. Coptosapelta has a unique pollen type within Rubiaceae and does not resemble possibly related genera. Pollen characteristics provide additional evidence supporting earlier findings that Coptosapelta represents an isolated lineage in Rubiaceae.     

A study on pollen grains ofCrocus cartwrightianus (Iridaceae)

Crocus cartwrightianus andC. cartwrightianus cv.albus pollen have been studied from structural and ultrastructural points of view and the germination assayed in vivo and in vitro.C. cartwrightianus pollen is regularly shaped and sized, has a low percentage of anomalous grains and has a high germination rate in vitro, whileC. cartwrightianus cv.albus is less regularly shaped with some variation in size and has a high percentage of anomalous grains and a low germination percentage in vitro. Ultrastructural observations have revealed, in the pollen of both the taxa, the presence of a thin elongated vegetative nucleus and a generative cell surrounded by a thin membrane. However,C. cartwrightianus pollen shows a thicker intine, andC. cartwrightianus cv.albus shows numerous pollen germination anomalies which are in common withC. sativus.     

Experimental results about Platanus pollen deposition

A comparison between the aerobiological and floristic data carried out for Platanus are reported. The source was constituted by 60 Platanus trees. When they were pollinating we calculated the total pollen prodution of the source, controlling the number of male inflorescences for each plant, the number of anthers for each inflorescence and the number of pollen for each anther. Aerobiological data were obtained with 16 Durham gravimetric samplers positioned over an area of approximately 25 km² at variable distances and in different directions to capture pollen transported by winds coming from any direction. In the first year of analysis, we carried out two kind of aerobiological monitoring positioning a Hirst volumetric trap near the closest gravimetric sampler. The two methods showed a similar trend in Platanus pollen profiles. In the following years, only Durham gravimetric samplers were used to study pollen dispersion. The data showed that there were high concentrations of pollen on the soil near the source while at distances higher than 800 m pollen concentration decreased dramatically. About 1/4 of all the pollen produced fell in an area within 400 m from the source and at a distance of 2750 m only 9 pollen grains/cm² fell throughout the entire season. This revised version was published online in June 2006 with corrections to the Cover Date.