Apertural chambers inGeranium: Development and ultrastructure

Pollen grains ofGeranium robertianum andG. pratense are tricolpate. At the time of the vacuolated microspore stage intine protrusions are formed at each aperture. Each aperture becomes separated from the vegetative cytoplasm by a thick ectintine layer. Starch grains are enclosed in the protrusions and do not participate in pollen tube growth.     

Pollen morphology and functional dioecy inSolanum (Solanaceae)

Dioecy has evolved independently several times in the large, mostly tropical genusSolanum. In all cases of dioecy inSolanum functionally male flowers have normal anthers, normal pollen and reduced stigmas while functionally female flowers have stigmas and anthers that appear normal but contain non-functional, usually inaperturate pollen. The inaperturate pollen has living cytoplasm, but apparently never germinates and it has been hypothesised that the pollen in these functionally female flowers is retained as a pollinator reward. Pollen morphology is compared in twelve of the thirteen known dioecious species ofSolanum, and some stages in the the development of inaperturate pollen in the anthers of functionally female flowers ofSolanum confertiseriatum of Western Ecuador are examined. Observations on the development and morphology of inaperturate pollen in functionally female flowers ofSolanum are related to hypotheses about the evolution of dioecy in the genus.     

Morphological and histochemical differentiation of the pollen wall ofBetula pendula Roth,during dormancy up to anthesis

Summary Anthers ofBetula pendula were collected at regular intervals during the dormancy period until anthesis. Ultrathin sections of maturing pollen grains were especially stained for polysaccharides and proteins and examined with TEM to determine whether structural or/and chemical changes in the pollen wall occur during the dormancy period of the plant life cycle. At the beginning of the dormancy period, the microspore wall consists of a well developed tectum, columellae and a foot layer. Spinules and supratectal elements are prominent. Microchannels are present in the tectum but not obvious in the foot layer. Some of the columellae are not clearly connected with the foot layer, but some connections are evident. Pores are filled with a thick fibrillar network flocculent material. The cytoplasm is packed with starch grains and lipid globules. The stainability for acidic and neutral polysaccharides and protein was tested, and variations in the pollen wall are illustrated. As temperature increased towards the end of dormancy and before anthesis there is obvious differentiation in the morphology of the pollen wall. The granular fibrillar layer beneath the pore and the Zwischenkörper are the most variable part of the wall. Different histochemical reactions observed in different layers at the aperture sites indicate different functions of these layers.     

凤仙花花药发育中多糖和脂滴组织化学研究

以不同发育时期的凤仙花花药为实验材料,采用组织化学方法,对花药发育中的结构变化及多糖和脂滴物质分布进行观察。结果表明:(1)凤仙花的花药壁由6层细胞组成,包括1层表皮细胞,2层药室内壁细胞,2层中层细胞和1层绒毡层细胞。其中绒毡层细胞的形态不明显,很难与造孢细胞区分,且在小孢子母细胞时期退化。(2)在小孢子母细胞中出现了一些淀粉粒,但减数分裂后,早期小孢子中的淀粉粒消失,又出现了一些小的脂滴;随着花粉的发育,小孢子形成大液泡,晚期小孢子中的脂滴也消失;小孢子分裂形成二胞花粉后,营养细胞中的大液泡降解、消失,二胞花粉中又开始积累淀粉;接近开花时,成熟花粉中充满细胞质,其中包含了较多的淀粉粒和脂滴。(3)在凤仙花的花药发育中,绒毡层细胞很早退化,为小孢子母细胞和四分体小孢子提供了营养物质;其后的中层细胞退化则为后期花粉发育提供了营养物质。     

Ultrastructural studies on plastids of generative and vegetative cells inLiliaceae 3. Plastid distribution during the pollen development inGasteria verrucosa (Mill.) Duval

Summary This paper describes the development of pollen grains ofGasteria verrucosa from the late microspore to the mature two-cellular pollen grain. Ultrastructural changes and the distribution of plastids as a result of the first pollen mitosis have been investigated using light and electron microscopy. The microspores as well as the generative and the vegetative cell contain mitochondria and other cytoplasmic organelles during all of the observed developmental stages. In contrast, the generative cell and the vegetative cell show a different plastid content. Plastids are randomly distributed within the microspores before pollen mitosis. During the prophase of the first pollen mitosis the plastids become clustered at the proximal pole of the microspore. The dividing nucleus of the microspore is located at the distal pole of the microspore. Therefore, the plastids are not equally distributed into both the generative and the vegetative cell. The possible reasons for the polarization of plastids within the microspore are briefly discussed. The lack of plastids in the generative cell causes a maternal inheritance of plastids inGasteria verrucosa.     

Exploding pollen in Montrichardia arborescens (Araceae)

Pollen grains of Montrichardia are inaperturate with psilate ornamentation. The pollen wall is formed by a thin ectexine and an extraordinarily thick intine. In living as well as in dead pollen grains contact with water leads to a rapid swelling of the intine followed by an explosive opening of the exine. Within a few seconds a thick tube is formed, which is not the pollen tube. The pollen protoplast is situated at the tip of the tube. These intine tubes are interpreted as pollen connecting tools to keep pollen grains together and adhere them to the cuticle of the hairless pollinators.     

Why does some pollen lack apertures? A review of inaperturate pollen in eudicots

Apertures are key characters of pollen grains with systematic importance in angiosperms. They function as sites for pollen tube exit, water uptake, transfer of recognition substances and accommodation of volume changes. Not all pollen has apertures; inaperturate pollen (lacking obvious apertures) characterizes many angiosperm groups, especially in early divergent angiosperms and monocots, but also eudicots. In order to expand our knowledge of the systematic distribution, possible functional significance and development of inaperturate pollen in angiosperms, this review focuses on inaperturate and cryptoaperturate (with hidden apertures) pollen in the large eudicot clade, which comprises about 75% of present‐day angiosperm species. It includes new TEM observations of inaperturate pollen from four exemplar taxa selected from different parts of the eudicot phylogeny. Two categories of inaperturate (including cryptoaperturate) pollen occur in eudicots. (1) Sterile attractant or feeding pollen associated with functional dioecy has evolved iteratively at least six times in conjunction with complex breeding systems in the core eudicots. (2) Fertile pollen has evolved numerous times independently throughout eudicots, though generally in a relatively small number of individual taxa. Notable exceptions are the petaliferous crotonoid Euphorbiaceae s.s., in which fertile inaperturate pollen occurs in c. 1500 species, and two subfamilies of Apocynaceae s.l. (Secamonoideae and Asclepiadoideae) with c. 2500 species with fertile inaperturate pollen in pollinia. Fertile inaperturate pollen is sometimes (but not always) associated with an aquatic habit, parasitism, insectivory, heterostyly, anemophily or pollinia. Most fertile inaperturate pollen has a thin exine, or the exine is largely restricted to isolated components (muri, protuberances, subunits) separated by thinner areas which probably function as apertures. In cryptoaperturate pollen, the aperture is covered by continuous exine which probably has a protective function, similar to an operculum. Developmentally, inaperturate pollen is not associated with any particular tetrad type or meiotic spindle orientation (unlike some apertures) due to the absence of a colpal shield of endoplasmic reticulum or other organelles and hence is independent of microsporogenesis type. The lack of a colpal shield during the tetrad stage of development permits complete deposition of first primexine and then exine around each microspore, possibly mediated by the action of the DEX1 protein. © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 155 , 29–48.     

Viral particles in developing pollen grains of Olea europaea

Double-walled tubules containing rows of isodiametric virus particles were observed in developing pollen grains of Olea europaea L. cultivar Correggiolo. Sometimes the tubules are contained in another double-walled tubular structure or in a tubular endoplasmic reticulum cistern. The viruses are present in the cytoplasm from the microspore mother cell stage up to the microspore stage but just before the first haploid mitosis they are to be found only in the pores, inside the evaginations formed by the plasmalemma. During the last phase of pollen grain development, after the germinative pores are completed, the viruses disappear.Abbreviations ER endoplasmic reticulum     

Pollen evolution and systematics inAnnonaceae with special reference to the disulcate Australian endemic genera

All genera ofAnnonaceae endemic in Australia (Ancana, Fitzalania, Haplostichanthus) show almost exactly the same type of disulcate (disulculate) pollen with intact exine extending over the sulci. Tetrad stages inHaplostichanthus andAncana reveal a latudinal subequatiorial orientation of the two sulci at the proximal hemisphere. Sometimes they fuse into a ±zonosulcate aperture.Fissistigma pollen grains are ±globose and have a flattened pole with a central elevation and a concentric groove, covered by a somewhat reduced exine. This palynological characters give further support for separating the generaAncana andFissistigma. Germination was observed inHaplostichanthus where the pollen tube emerges at one of the two sulci and inFissistigma where the flattened part breaks up during germination. The aperture types described here are obviously transitional stages between aperturate and inaperturate pollen grains and are discussed in regard to pollen evolution.     

Pollen morphology in Asiatic species ofTrillium

Pollen grains of five species and two interspecific hybrids ofTrillium from eastern Asia have been examined with scanning electron and transmission electron microscopy. All species from eastern Asia, excludingT. govanianum, are characterized by inaperturate pollen and an unstratified and granulated exine structure. Pollen of the Himalayan species,T. govanianum, is, however, monocolpate and intectate. The palynological evidence suggests that the Asiatic species, excludingT. govanianum, are closely related to each other and toT. erectum in North America, andT. govanianum is independent of the other species in the genus.     

Microsporogenesis and exine structure in Trochodendron aralioides Siebold and Zuccarini (Trochodendraceae)

This study aimed to elucidate the anther wall development, pollen wall development, and exine structure of Trochodendron aralioides Siebold and Zuccarini, a tree with primitive vessels but long considered to lack vessel elements in its wood. The anther wall is the basic type: epidermis, endothecium layer, three middle layers, and tapetum. The anther tapetum is glandular and cells are uniseriate. Microspore mother cells undergo meiosis with simultaneous cytokinesis to produce tetrahedral tetrads enclosed within a callose wall. Before development of the protectum, primexine is inserted against the callose, and the plasma membrane is invaginated. Then, the probacula are elongated under the protectum and arise basally from the plasma membrane. The foot layer formation is concomitant with callose wall dissolution. The foot layer is thick, and the endexine is thin. The foot layer and the endexine are both continuous. The intine is initially formed in the vacuolated microspore stage. Hollow Ubisch bodies are observed on the inner surface of the tapetum in free microspore stage. Pollen grains are tricolporate and 2-celled at the time of shedding. The numerous anthers of a single flower are at different development stages in both protandrous and protogynous individuals.     

Pollen morphology and pollen-wall proteins (localization and enzymatic activity) inSesamothamnus lugardii (Pedaliaceae)

The pollen grains ofSesamothamnus lugardii Stapf (Pedaliaceae of subdesert regions of SE tropical Africa) are associated in acalymmate tetrads (cross wall cohesion), with a tectate and perforate exine and 8–12 colpi. The pollen wall consists of an ectexine with a complete, perforate and ample tectum, columellated infratectum and clearly interrupted and fragmented foot layer. The endexine is built of scanty lamellae and granules. The intine is bistratificate, with a homogeneous, fibrillate layer (endintine or intine-2) and a heterogeneous, more lax and channeled layer (exintine or intine-1). Test for glycoprotein is particularly positive in the homogeneous internal intine and channels of external intine. On the other hand acid phosphatase has been localized in the exine and channeled external intine layers. These observations confirm the general interpretation of the distribution of wall compounds.     

The ultrastructure and reproductive significance of Lasiostrobus microspores

The normally inaperturate microspores of the Pennsylvanian cone Lasiostrobus are found to rarely possess trilete sutures, confirming the subequatorial position of the sacci. Sporoderm ultrastructure compares favourably with saccate pollen of some extant members of the Coniferales. The shedding of grains in the microspore stage suggests that microgametophyte evolution had passed the stage of prothallial cell production, and that Lasiostrobus is an advanced member of the Paleozoic gymnosperm complex.     

Development and cytochemistry of pollen and tapetum in Mitriostigma axillare (Rubiaceae)

The development of pollen grains and tapetum in Mitriostigma axillare (Rubiaceae) was studied from anther primordium to dehiscence. Anthers were freeze-cracked and studied with SEM. Embedded anthers were sectioned and studied with LM and TEM. Cytochemistry was performed in order to distinguish the different layers of the sporoderm and to determine its chemical nature at different development stages. The pollen grains remained as tetrads by partial fusion of the exine, probably because of reduced callose septa during the stage of microspore tetrads within callose envelopes. Characteristic features of the sporoderm were an irregular foot layer, an endexine composed of amalgamated granules, a transient granular-fibrous layer beneath the endexine, and a thin intine. During maturation of the exine, the endexine became chemically different from the ectexine. All layers of the sporoderm were reduced in thickness due to stretching during the engorgement of the pollen grains prior to dehiscence. The pollen grains were colpoidorate with a reticulate to microreticulate tectum covered with a scanty surface coating. The mature pollen grains were binucleate and contained a lot of starch grains. Thick intineous onci protruded through the apertures and formed papillae. About 50% of the microspores were aborted. The tapetum was of secretory type, probably with cycles of hyperactivity and protrusions of the cells into the locular cavity. No syncytium was formed and there were neither orbicules nor tapetal membrane.     

Anther development, microsporogenesis and microgametogenesis in Heliconia (Heliconiaceae, Zingiberales)

Anther development, microsporogenesis and microgametogenesis in several species of Heliconia were investigated as part of a complementary embryological study of the Heliconiaceae. All studied Heliconia species present bithecate and tetrasporangiate anthers with fertile pollen grains; only H. rivularis, a natural hybrid, presented sterile pollen grains of variable size and no content. The anther wall has an uniseriate epidermis and endothecium, the latter with helicoidal thickenings, although some cells of the middle layers also showed thickenings; the biseriate tapetum is of amoeboid non-syncytial type, since the tapetum cells did not fuse together forming a true plasmodium. The microsporogenesis is successive leading to isobilateral tetrads. The inaperturate pollen grains had a very reduced exine consisting of a thin, more or less continuous layer with small spines upon; the pollen grain shape is variable among the species, all of them presenting heteropolar pollen, except H. angusta with isopolar ones. Most of these characteristics were shared with other studied Zingiberales, although more studies need to be done.     

Sporoderm development inNymphaea mexicana (Nymphaeaceae)

Mature pollen grains ofNymphaea mexicana have a verrucate proximal surface, a psilate distal surface and an anazonasulculus (encircling-sulcate aperture). The developmental events of microspores and tapetal cells were observed with TEM and SEM. Radially oriented substructural elements are seen in the microspore surface coating ofNymphaea mexicana from the early tetrad stage through the whole exine development. These elements, being the structural units of the microspore surface matrix (glycocalyx), are associated with sporopollenin precursor accumulation. In young free microspores, radially oriented elements are observed at both proximal and distal poles as a palisade between the endexine and plasmalemma.—Several points are discussed: (1) the initial and mature forms of exine substructure elements; (2) the significance of exine substructure for realisation of morphogenetic processes; (3) the ways by which verrucate and psilate sculpture patterns are developed.     

Dioecism of the genusCallicarpa (Verbenaceae) in the Bonin (Ogasawara) Islands

The morphology and function of the flowers of three species of the genusCallicarpa, C. glabra, C. nisnimurae andC. subpubescens, endemic to the Bonin Islands, revealed that all three species were dioecious. Male plants had short-styled flowers with sterile ovaries, while female plants had long-styled flowers with non-germinating inaperturate pollen grains. Such dioecism has never been reported from the family Verbenaceae. This curious sexual system with non-germinating pollen grains may have been maintained as a reward to insect pollinators.     

水鳖科9属15种植物花粉形态的研究

应用光学显微镜、扫描电镜和透射电镜对水鳖科Hydrocharitaceae 9属15种植物的花粉形态进行 了观察。水鳖科植物花粉为圆球形至近椭球形,无萌发孔或偶为单沟萌发孔,外壁纹饰通常为小刺状纹 饰,刺密集或稀疏,花粉表面具瘤状、疣状、颗粒状、皱波状突起或光滑。外壁由覆盖层、柱状层和基层组 成。覆盖层厚或较薄,柱状层小柱发育不明显,基层薄。水鳖科植物在花粉大小、纹饰类型、刺的长短、 密度、形态、萌发孔的有无以及花粉壁的结构等方面表现出了较为明显的差异,这些特征对探讨类群间 关系具有较重要意义。由于黑藻属Hydrilla和Stratiotes属花粉较为特殊,支持将它们各自作为一个独立 的族处理。水鳖科植物花粉外壁纹饰和结构特点表明该科与水雍科Aponogetonaceae、泽泻科Alismataceae 和花蔺科Butomaceae等近缘,而该科植物花粉大多无萌发孔等则反应了该科与茨藻目Najadales植物有密切联系。     

The pollinium ofLoroglossum hircinum (Orchidaceae) between pollination and pollen tube emission

The structure of the massulae composing the pollinium ofLoroglossum hircinum was studied before pollination and 12 and 24 hours afterwards. The grains are grouped in tetrads closely packed in massulae. The exine is only present on the outside of the massulae. The intine consists of two layers: a compact layer surrounding the pollen grain and a looser layer surrounding the pollen grain and a looser layer surrounding the tetrad. Twelve hours after pollination, pollen volume and the space between the tetrads increase due to vacuolization. Twenty-four hours after pollination, pollen volume and tetrad spacing are higher due to vacuolization and some grains have emitted pollen tubes. Pollen growth due to vacuole formation, and the absence of common walls between adjacent tetrads lead to crumbling of the massulae. The mature pollen grain does not have apertures: the site of pollen tube emission is determined after pollination. The first grains to germinate are those in the centre of the massula. The vegetative cell nucleus is the first to enter the pollen tube; the generative cell elongates and undergoes the second haploid mitosis shortly after entering the pollen tube.     

笔竹大小孢子发生及雌雄配子体发育研究

为探讨笔竹(Pseudosasa viridula)结实率低的原因,该文通过采用石蜡切片的方法结合显微技术对笔竹大小孢子发生及雌雄配子体的发育过程进行研究.结果表明:(1)笔竹的雄蕊多为3枚,极少有6枚,每枚花药具有4个花粉囊.(2)花药壁发育为基本型,由4层细胞构成,由外向内依次为表皮细胞、药室内壁细胞、中层细胞和绒...