The tapetum inSchizaea pectinata (Schizaeaceae) and a comparison with the tapetum inPsilotum nudum (Psilotaceae)

A combination tapetum consisting of a cellular, parietal component and a plasmodial component occurs inSchizaea pectinata. A single, tapetal initial layer divides to form an outer parietal layer which maintains its cellular integrity until late in spore wall development. The inner tapetal layer differentiates into a plasmodium which disappears after the outer exospore has developed. In the final stages of spore wall development, granular material occurs in large masses and is dispersed as small granules throughout the sporangial loculus. No tapetal membrane develops. Comparisons are drawn with the combination tapetum found inPsilotum nudum.     

A comparison of tapetal structure and function in pteridophytes and angiosperms

This article is based on a survey of literature with particular reference to the structure and functioning of the tapetum during sporogenesis in angiosperms and pteridophytes. Some original observations onSchizaea pectinata andSelaginella japonica are also included. In the past, superficial similarities in the structure and function of pollen grains and spores have dominated the research approach, but patterns of development and the functioning of the tapetum during sporogenesis in angiosperms and pteridophytes are shown to differ quite significantly and raises questions relating to homologies. It is suggested that direct comparisons may no longer be appropriate.     

蕨类植物紫萁绒毡层细胞凋亡的细胞学特征

绒毡层凋亡过程是小孢子发生中的重要事件,以往的研究主要集中在被子植物,蕨类植物尚未见此方面的报道。该研究首次采用透射电镜和免疫荧光技术对蕨类植物紫萁(Osmunda japonica Thunb.)绒毡层细胞凋亡的细胞学过程进行了观察,以明确紫萁绒毡层细胞的发育类型和凋亡特征,为蕨类植物绒毡层细胞凋亡的深入研究以及孢子发育研究提供依据。结果显示：(1)紫萁的绒毡层属于复合型,即外层绒毡层为分泌型,该层细胞发育过程中液泡化,营养物质被吸收;内层绒毡层为原生质团型,经历了细胞凋亡的过程。(2)绒毡层内层细胞在凋亡过程中细胞壁和细胞膜降解,细胞质浓缩且空泡化;细胞核内陷、变形,染色质浓缩凝聚,形成多数小核仁,DAPI荧光由强变弱;线粒体、质体、内质网、高尔基体等细胞器逐渐退化,液泡中多包含纤维状物、絮状物、黑色嗜锇颗粒和小囊泡等;出现多泡体、多膜体和细胞质凋亡小体,上述特征与种子植物绒毡层凋亡特征基本一致。(3)与种子植物相比,紫萁绒毡层的细胞凋亡开始得早,在整个凋亡过程中没有核凋亡小体的产生;除了产生孢粉素外,绒毡层细胞内产生了大量的丝状物质、絮状物质和电子染色暗的颗粒物,这些物质可能用于...     

Embryology ofMalaxis saprophyta,with comments on the systematic position ofMalaxis (Orchidaceae)

InMalaxis saprophyta, anther wall development corresponds to the Monocotyledonous type. The uninucleate tapetum is of secretory type and the endothecium develops U- and V-shaped thickenings on the inner tangential and radial walls. Cytokinesis is simultaneous; tetrahedral, isobilateral and T-shaped tetrads are formed which are compactly aggregated in pollinia. At anthesis the microspore tetrads are 2-celled. The ovule is anatropous, bitegmic and both integuments are dermal in origin. A single hypodermal cell develops directly into a megaspore mother cell. Embryo sac development is predominantly monosporic and less often bisporic. Irrespective of the type of development, the mature embryo sac is 6-nucleate. Although double fertilization occurs, the primary endosperm nucleus degenerates. Embryogeny is of the Onagrad type. The mature embryo lacks differentiation into cotyledon, plumule and radicle. The reticulate seed coat is formed entirely by the outer layer of outer integument. There are three sterile and three fertile valves in the ovary. Although initially parenchymatous, the entire three sterile valves in the ovary and the upper half of the three fertile valves become sclerified after fertilization. The embryological characters support the disputed systematic position ofMalaxis within subtribeMalaxidinae ofEpidendreae.     

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.     

车桑子小孢子发生与雄配子体发育

为了解干热河谷区车桑子(Dodonaea viscosa)胚胎学特征及其结籽率低的原因,采用常规石蜡切片法和电镜扫描技术对车桑子小孢子发生、雄配子体发育和花粉的形态特征进行了观察。结果表明,车桑子花药具有4个花粉囊。完整的花药壁从外到内依次为表皮、药室内壁、2～3层中层细胞和绒毡层;绒毡层类型是腺质绒毡层。花药成熟期,中层、绒毡层均退化消失。小孢子母细胞进行同时型胞质分裂;四分体为四面体型结构。成熟的花粉为二细胞型。花粉近球形,外壁密布颗粒状纹饰,具有3条不构成合沟的萌发沟。雄性生殖发育过程出现的异常现象可能是干热河谷地区车桑子结籽率低的原因之一。     

开口箭大小孢子发生及雌雄配子体发育

利用石蜡切片技术,对百合科植物开口箭(Tupistra chinensis Baker)大小孢子发生及雌雄配子体发育进程进行胚胎学观察分析,以明确开口箭胚胎发育的特征,为百合科植物的研究提供生殖生物学依据。结果表明:(1)开口箭花药具有4个药室,花药壁的发育方式为基本型,由表皮、药室内壁、中层及绒毡层组成;绒毡层发育类型为分泌型,到四分体花药阶段绒毡层细胞开始解体退化,花药成熟时完全消失。(2)花粉母细胞减数分裂为连续型,依次形成二分体、四分体,四分体为左右对称形;成熟花粉为2-细胞花粉,具单萌发沟。(3)子房3室,倒生型胚珠6枚,双珠被,薄珠心;在花部的分化早期,由珠心顶端表皮下方分化出雌性孢原细胞,孢原细胞经过一次平周分裂形成周缘细胞和造孢细胞,造孢细胞发育为大孢子母细胞;大孢子母细胞第一次减数分裂后形成二分体,珠孔端的二分体孢子退化,合点端的二分体孢子继续第二次分裂,形成两个子细胞依次发育为二核胚囊、四核胚囊和八核胚囊;开口箭的胚囊发育类型为葱型。     

Studies in the embryology ofHeliantheae (Compositae)

The embryology ofLagascea mollis, Zinnia angustifolia andGalinsoga parviflora has been studied. The anther archesporium is hypodermal and consists of a single row of 6–8 cells, there are two layers below the epidermis of the anther and a periplasmodial tapetum. Ripe pollen grains are tricolpate and 3-celled. The ovary contains a single ovule, but in a few cases ofGalinsoga parviflora two ovules have been found. The female archesporium is unicellular, but sometimes more than one archesporial cell occurs inLagascea mollis andZinnia angustifolia. The embryo sac development is of the Polygonum type, the synergids are hooked, antipodal cells show great variation. The endosperm development is cellular inGalinsoga parviflora and peripheral layer persists in the mature seed. The embryo development conforms to the Senecio variation of the Asterad type. The pericarp structure and cmbryological features support the disputed systematic position ofLagascea withinHeliantheae. Part of a Ph.D. thesis, accepted at Andhra University.     

Entwicklungsgeschichte und Ultrastruktur des Pollenkitts beiTilia (Tiliaceae)

The pollenkitt ofTilia platyphyllos andT. tomentosa is produced during the tetrade-stage by the tapetum plastides. After tapetum degeneration pollenkitt and accompanying substances are deposited predominantly in the intrabacular spaces of the exine, and only in small quantities on the surface of the exine itself.

     

The tapetum: Its form,function, and possible phylogeny inEmbryophyta

It appears that the tapetum is universally present in land plants, even though it is sometimes difficult to recognize, because it serves mostly as a tissue for meiocyte/spore nutrition. In addition to this main function, the tapetum has other functions, namely the production of the locular fluid, the production and release of callase, the conveying of P.A.S. positive material towards the loculus, the formation of exine precursors, viscin threads and orbicules (= Ubisch bodies), the production of sporophytic proteins and enzymes, and of pollenkitt/tryphine. Not all these functions are present in all land plants:Embryophyta. Two main tapetal types are usually distinguished in theSpermatophyta: the secretory or parietal type and the amoeboid or periplasmodial type; in lower groups, however, other types may be recognized, with greater or lesser differences. A hypothetical phylogenesis of the tapetum is proposed on the basis of its morphological appearance and of the nutritional relations with meiocytes/spores. The evolutionary trends of the tapeta tend towards a more and more intimate and increasingly greater contact with the spores/pollen grains. Three evolutionary trends can be recognized: 1) an intrusion of the tapetal cells between the spores, 2) a loss of tapetal cell walls, and 3) increasing nutrition through direct contact in narrow anthers.     

Embryology ofKyllinga monocephala (Cyperaceae) and its systematic position

Aspects of the life history ofKyllinga monocephala are described. Anther wall development corresponds to the Monocot type. The endothecium shows spiral thickenings. The tapetum is glandular and has uninucleate cells. Ubisch granules are present. Mature pollen grains (pseudomonads) are 3-celled at maturity. Ovules are bitegmic, crassinucellate and develop a funicular obturator. The embryo development conforms to theJuncus-variation of the Onagrad type. Endosperm, seed coat and pericarp are described.     

Flower development inSpirodela polyrrhiza (Lemnaceae)

Flowering under experimental conditions inSpirodela polyrrhiza, and the development of the spatha, stamina and pistil are described and illustrated by microphotographs. During the development of the microsporangia the existence of the middle layer between endothecium and tapetum has been documented for the first time inLemnaceae.     

Genetic control of male fertility in Arabidopsis thaliana: structural analysis of premeiotic developmental mutants

We have taken a mutational approach to identify genes important for male fertility in Arabidopsis thaliana and have isolated a number of nuclear male/ sterile mutants in which vegetative growth and female fertility are not altered. Here we describe detailed developmental analyses of four mutants, each of which defines a complementation group and has a distinct developmental end point. All four mutants represent premeiotic developmental lesions. In ms3, tapetum and middle layer hypertrophy result in the degeneration of microsporocytes. In ms4, microspore dyads persist for most of anther development as a result of impaired meiotic division. In ms5, degeneration occurs in all anther cells at an early stage of development. In ms15, both the tapetum and microsporocytes degenerate early in anther development. Each of these mutants had shorter filaments and a greater number of inflorescences than congenic male-fertile plants. The differences in the developmental phenotypes of these mutants, together with the non-allelic nature of the mutations indicate that four different genes important for pollen development, have been identified.     

鹅毛竹大小孢子及雌雄配子体发育

利用扫描电镜、透射电镜、石蜡切片,对鹅毛竹的花芽分化、大、小孢子及雌、雄配子体的发育进行了详细观察.结果发现:鹅毛竹花药具4个药室,花药壁由表皮、药室内壁、中层、绒毡层4层结构组成,花药壁发育为单子叶型,绒毡层为腺质型,小孢子母细胞减数分裂中的胞质分裂为连续型,产生左右对称型小孢子.鹅毛竹成熟花粉大多2细胞型,都具1个萌发孔.鹅毛竹子房为单子房,子房1室,侧膜胎座,一个倒生胚珠,双珠被,薄珠心.大孢子母细胞由一个雌性孢原细胞直接发育而成,大孢子四分体呈线型,合点端一个大孢子分化为功能大孢子,由功能大孢子经过3次有丝分裂形成8核胚囊,发育类型为蓼型,位于核点端的3个细胞核进行多次分裂形成多个反足细胞.至此,成熟胚囊形成.并就鹅毛竹不结实的原因进行了探讨.     

Orbicules and the ektexine are homologous sporopollenin concretions inSpermatophyta

During microsporogenesis sporopollenin becomes accumulated independently by the only two sporopollenin-producing cell types in the higher plants—the anther tapetum and the microspores—not only within the exine, but also within the orbicules. In Spermatophytes usually only a single orbicule type is found, but sometimes, e.g., inEuphorbia palustris, two different types exist within a single species. Very interestingly, most orbicules are strikingly similar in their structure, sculpture, shape and dimension to the respective ektexine: e.g., smooth sporopollenin globules are found near the totally smooth exines ofEupomatia laurina andPentaphragma sinense respectively, while inDelonix elata andEuphorbia palustris the orbicules often look like some incomplete tectum pieces. All these parallelisms can be attributed to the homologous and therefore highly similar genetic information to form sporopollenin in the sporogeneous tissue and the anther tapetum. The orbicules should be seen neither as sporopollenin storage, as sporopollenin transfer vehicle, nor as sporopollenin surplus.     

Embryology ofEriocaulon setaceum (Eriocaulaceae)

Eriocaulon setaceum can be characterized by: young microsporangium wall with epidermis, endothecium (with fibrous thickenings), and glandular tapetum (uninucleate cells); pollen grains 3-celled, spiraperturate; embryo sac development according to the Polygonum type and with antipodal cyst; endosperm nuclear; embryo small, with incipient differentiation into cotyledonary and epicotylary loci; seed coat mainly from the inner layers of the integuments; pericarp 2-layered and membranous. Embryologically, theEriocaulaceae are nearer to theXyridaceae than to otherFarinosae. Their elevation to the rank of an order,Eriocaulales, therefore appears justified.     

Ontogenesis of monad pollen inPterostylis plumosa (Orchidaceae,Neottioideae)

Pterostylis plumosa (Neottioideae) is an orchid with monosulcate monad pollen. Tetrads may be isobilateral, decussate, tetrahedral, rhomboid or T-shaped, but all pollen grains have a similar shape. Those belonging to the same tetrads are contiguous from microspore release to opening of the anther, with the furrow oriented inwards. Sporophytic proteins are present outside the furrow. The tapetum is of the parietal type without orbicles. The increase in pollen grain size between meiosis and maturity is only three-fold. The generative cell is spherical when the pollen is mature. These features are discussed in relation to the primitive nature of the species.     

Embryology ofCrocus thomasii (Iridaceae)

The embryology ofCrocus thomasii is described. Male meiosis is of simultaneous type, and gives rise to starchy microspores which develop into lipoid pollen grains; these are two-celled and show a spinulate acolpate, abaculate exine lacking apertures. The tapetum is glandular and its cells become bi- or sometimes multinucleate. The ovule is anatropous and bitegmic; the inner integument forms the micropyle. Megasporogenesis is heteropolar with starch accumulation in the functional chalazal megaspore. Embryo sac development conforms to thePolygonum type. The endosperm development is nuclear. The embryo develops according to the Caryophyllad type. In the ripe seed it is differentiated and enveloped by a starchy cellular endosperm. The embryological characters observed strongly favour a close relation betweenC. thomasii andC. sativus.     

中国特有种苦绳(广义夹竹桃科)的大小孢子发生和雌雄配子体发育及其分类学意义

近年来的分子系统学把狭义萝藦科和狭义夹竹桃科合并为广义夹竹桃科,包括5个亚科和25个族,但亚科和族间的亲缘关系较为复杂,亟待多学科证据澄清。本文利用常规石蜡切片技术观察了马利筋亚科南山藤属中的中国特有植物苦绳(Dregea sinensis var. sinensis)的孢子发生和配子体发育,结合已有资料比较了5个亚科的胚胎学特征。结果表明:(1)苦绳的花药由一对侧生并列药室组成,各有一个花粉团。(2)花药壁有6层,由外至内分别为表皮、2层药室内壁、中层和2层绒毡层,花药壁发育模式属于多层型。(3)绒毡层细胞单核,排成2列,为腺质型; 在小孢子四分体形成时期,药室内壁发生明显纤维状加厚; 花药成熟时,位于药室远轴最外侧处的花药壁发生断裂,准备散粉。(4)小孢子母细胞减数分裂中,胞质分裂方式为连续型,小孢子四分体排列方式为左右对称; 成熟花粉粒为3-细胞型,排列紧密,形成花粉团。(5)雌蕊含有两枚离生心皮,具边缘胎座,胚珠倒生,单珠被,薄珠心,蓼型胚囊。本文观察到的这些胚胎学特征为牛奶菜族提供了新资料。同时,胚胎学特征在5个亚科间的区别和联系,支持广义夹竹桃科的成立。     

葱兰大小孢子发生和雌雄配子体发育及其系统学意义

采用常规石蜡切片技术,对石蒜科葱兰的花药壁发育、大小孢子的发生和雌雄配子体的发生过程进行了研究,并对葱兰属、石蒜科、百合科以及葱科的胚胎学特征进行比较讨论。结果表明:(1)葱兰花药四室,药壁由表皮、药室内壁、中层和绒毡层组成;药壁发育类型为单子叶型,绒毡层的类型为分泌型;花粉成熟时药室内壁径向加长并纤维状加厚,表皮宿存;小孢子母细胞在减数分裂过程中胞质分裂为连续型,小孢子四分体排列方式主要为四面体型,还有少数一些为左右对称型,成熟花粉为2-细胞型。(2)葱兰的雌蕊3心皮合生,子房下位,中轴胎座,3室,每室具2列倒生胚珠;胚珠双珠被,厚珠心,具蓼型胚囊。(3)葱兰属的胚胎学特征与石蒜科的其他种类存在较大的差异,如葱兰属花药壁发育为单子叶型,而石蒜科花药壁发育主要为双子叶型,但葱兰属的这些胚胎学特征却和百合科较为相似。