花粉粒外壁成层现象、外壁外层的结构和被子植物的进化

显花植物的花粉粒外壁被认为由外壁外层所覆盖着的外壁内层组成。这是在形态学和个体发育上所规定的解释。外层内层沉积在可能由质膜起源的切向膜上。外壁外层沉积在没有居间膜(可是,这膜有时存在,但不是切向膜)的原外壁中,在成熟状态中,外壁内层可能或不可能保存具片层的,可能或不可能被一薄的清晰区与外壁外层分隔开,可能或不可能在染色性能和溶解度上与外壁外层有所区别。在这一方面,例如番荔枝科(Annonaceae)和肉豆蔻教(Myristicaceae)花粉粒具片层的内部层次被认为是外壁内层的,跟裸子植物和其他被子植物的外壁内层同源。一如蕨类植物孢子外壁[译者注:原文为花粉外壁(exine),应为孢子外壁(exospore)]的内部基本上具片层的区同源,只有在个体发育上能够证明没有切向膜支持着内部的外壁区时才可以认为花粉是缺乏外壁内层。     

琼榄小孢子发生、雄配子体发育及花粉粒形态

利用光学显微技术和电镜扫描技术研究了琼榄的小孢子发生、雄配子体发育和花粉粒形态以增加广义心翼果科的胚胎学和孢粉学资料。主要结果如下：（1）花药四孢囊;（2）花药壁四层,从外到内分别为表皮、具纤维性加厚的药室内壁、退化早的中层和细胞具2~4核的分泌型绒毡层;（3）小孢子母细胞胞质分裂同时型,形成四面体型排列的小孢子四分体;（4）成熟花粉粒为二细胞型;（5）花粉粒具3个隐形萌发孔,外壁为网状纹饰。琼榄与心翼果属的小孢子发生和雄配子体发育特征非常相似,稍有不同。琼榄的花粉粒形态特征与同属其它种基本相同。     

中国特有属牛筋条属的花粉形态与其系统位置

从发育的角度研究了中国特有单种属DichotomanthesKurz及与其系统学研究有关的外类群Prinsepiautilis的花粉形态 ,扫描电镜观察显示Dichotomanthes花粉粒自脱离四分体胼胝质膜开始至成熟二核花粉粒不同发育时期 ,花粉形态和外壁纹饰未见变化 ,仅花粉体积随成熟度增加而有所增大。而Prinsepiautilis ,其花粉粒刚脱离四分体时形状和成熟花粉明显不同 ,成熟花粉极面观为三裂圆形 ,赤道面观为圆形 ,外壁具清晰的平行条纹 ,但幼嫩花粉粒的形状很特别 ,极面观为深三裂圆形 ,赤道面观亦见花粉在两条沟之间下陷而沟部外突 ,明显为角萌发孔花粉 ,且花粉体积较成熟者小 ,而外壁纹饰同成熟者相比无根本性差异。前述两种植物花粉在不同成熟期体积有明显差异 ,而外壁纹饰在不同成熟期不存在质的变化并相对稳定 ,说明花粉外壁纹饰这一性状在蔷薇科中具有较为重要的分类学意义。DichotomanthesKurz具典型Rosaceae花粉的三孔沟结构 ,外壁具条纹 -穴状纹饰。将其孢粉学特征同Rosaceae 4个亚科有关类群的同类资料相比较 ,并结合其它形态解剖与细胞学等研究结果 ,支持将Dichotoman thes置入Maloideae下而不赞同将其另立亚科或置于Prunoideae之下。此外 ,由于Prinsepiautilis的花粉在其发育初期具角萌发孔花粉 ,与Cunoniacea     

重楼属花粉形态的研究

本文对重楼属(paris L.)18个种和变种或变型的花粉形态进行了光学显微镜和电镜的观察和初步比较研究。就所观察过的种类看,花粉粒为扁球形,极面观椭圆形。左右对称。具一远极沟。外壁两层,外层厚于或等于内层。外壁纹饰有穴状、网状和皱-网状之分。不同的种类,其花粉大小和外壁纹饰均或多或少有所不同,其中有些种类的外壁纹饰比较特殊,可以作为区分种类甚至作为分组的依据或参考。     

沿阶草亚科(百合科)的花粉形态及其属间亲缘关系的探讨

本文对山麦冬属Liriope 3种、沿阶草属Ophiopogon 24种和球子草属Peliosanthes 2种植物的花粉,用扫描电镜和透射电镜观察了它们的花粉的形态以及外壁的超微结构。发现它们的花粉外壁纹饰和结构明显的可分为两类：1．皱波状类型：具穿孔类型,山麦冬属和沿阶草属大体属这一类,其外壁外层具覆盖层、柱状层和基层、内层不明显,内壁明显。 2．瘤状突起类型：球子草属大体属这一类,其瘤状突起大小不均,外壁外层无覆盖层, 柱状层为大小不均的小柱,内层不明显,内壁明显。 这表明山麦冬属和沿阶草属的关系密切,但它们与球子草属的关系则较远。本文还论述了这三属的花粉形态与外部形态之间的相关性,并讨论了这三属之间的进化关系。 山麦冬属、沿阶草属和球子草属的外壁纹饰和结构等特征均支持将这三属分成沿阶草族和球子草族的观点。     

华中铁角蕨孢子纹饰形成的超微结构观察

利用透射电子显微镜对铁角蕨科(Aspleniaceae)华中铁角蕨(Asplenium sarelii Hook.)孢子及其纹饰的形成过程进行观察。结果表明:①华中铁角蕨孢子囊发育为薄囊蕨型;②孢子外壁表面光滑,远极面的外壁厚约0.8~1.1μm,近极面的外壁厚约1.4~1.8μm;③孢子周壁厚度约4~5μm,染色较外壁深,分为内层和外层;内层紧帖外壁表面,其上具柱状、瘤状或疣状突起;外层向外隆起形成脊状纹饰的轮廓,脊的下方具空腔,脊的顶端具翅;④铁角蕨型与鳞毛蕨型孢子外壁和周壁纹饰的形成过程具有相似性;⑤孢子的成熟度对于孢子形态的研究是至关重要的,只有完全成熟的孢子的表面纹饰才是稳定的。     

长梗苦草花粉粒的电镜观察

通过扫描电镜和透射电镜首次对沉水植物梗苦草（Vallisneria longipedunculata)的花粉粒进行了观察,其花粉粒的细胞壁非常薄,无萌发孔,但有2－3个具有较质丝的凹穴。花粉粒细胞壁的覆盖层十分不明显。具散生的颗粒;外壁内层较为厚实,但柱状结构分化不明显,整体呈海绵状;花粉内壁较厚。在花粉粒内部,有大量的单粒和复粒的淀粉粒,但未见到半复粒。     

芝麻核雄性不育系ms86-1微粉发生的细胞学观察

芝麻(Sesamum indicum)核雄性不育系ms86-1姊妹交后代表现为可育、部分不育(即微粉)及完全不育(简称不育)3种类型。不同育性类型的花药及花粉粒形态差异明显。Alexander染色实验显示微粉植株花粉粒外壁为蓝绿色, 内部为不均一洋红色, 与可育株及不育株花粉粒的染色特征均不相同。为探明芝麻微粉发生机理, 在电子显微镜下比较观察了可育、微粉、不育类型的小孢子发育过程。结果表明, 可育株小孢子母细胞减数分裂时期代谢旺盛, 胞质中出现大量脂质小球; 四分体时期绒毡层细胞开始降解, 单核小孢子时期开始出现乌氏体, 成熟花粉时期花粉囊腔内及花粉粒周围分布着大量乌氏体, 花粉粒外壁有11–13个棱状凸起, 表面存在大量基粒棒, 形成紧密的覆盖层。不育株小孢子发育异常显现于减数分裂时期, 此时胞质中无脂质小球出现, 细胞壁开始积累胼胝质; 四分体时期绒毡层细胞未见降解; 单核小孢子时期无乌氏体出现; 成熟花粉时期花粉囊腔中未发现正常的乌氏体, 存在大量空瘪的败育小孢子, 外壁积累胼胝质, 缺乏基粒棒。微粉株小孢子在减数分裂时期可见胞质内有大量脂质小球, 四分体时期部分绒毡层发生变形, 单核小孢子时期有部分绒毡层开始降解; 绒毡层细胞降解滞后为少量发育进程迟缓的小孢子提供了营养物质, 部分小孢子发育为正常花粉粒; 这些花粉粒比较饱满, 表面有少量颗粒状突起, 但未能形成覆盖层, 花粉囊腔中及小孢子周围存在少量的乌氏体。小孢子形成的育性类型与绒毡层降解是否正常有关。     

中国红豆杉科花粉形态的研究

红豆杉科(Taxaceae),花粉近球形,有时稍扁或稍长。直径为20．8-45．8μm。具远极 薄壁区,或不典型的乳头状突起。外壁两层,内外层厚度相等,有时层次不明显。在光学显微 镜下,外壁表面粗糙,或具微弱的颗粒状纹饰。在穗花杉(Amentotaxus argotaenia)这个种里,有些花粉粒具残存气囊。在扫描电镜下,外壁表面具粗瘤和细瘤两种纹饰类型。透射电镜本科各属代表种观察表明,本科花粉外壁内层具片状结构,外层由单层瘤状纹饰分子构成,有 些种细瘤连接形成覆盖层。根据花粉形态资料,可将本科植物分为2个族： 白豆杉族(仅包括白豆杉一个属)和红豆杉族(包括红豆杉属和榧树属)。 鉴于穗花杉属花粉的特殊性,建议独立上升为穗花杉科Amentotaxaceae。     

辣椒花药培养胚状体发生的组织学和细胞学研究

采用荧光显微镜、扫描电镜和透射电镜技术．系统研究了辣椒花药培养胚状体发生的组织学和细胞学变化特征。辣椒单个花药中花粉发育具有强烈的不同步性。随着培养时期的变化．不同时期花粉的百分率也发生变化。处于单核靠边期的小孢子培养以后按两种发育途径之一进行发育。在多数情况下,孢子体不对称分裂,产生典型双核花粉。胚性花粉粒是由营养核的重复分裂形成的。当小孢子从四分体中释放出来．特殊类型的外壁已经形成。在随后的花粉发育过程中．小孢子体积增大,外壁继续加厚。培养24h后,小孢子体积增大。胚性发生的小孢子表现出两种不同的形态变化。当胚状体发育到心形胚时．胚状体的表皮细胞排列规则。用光学和电子显微镜分析了小孢子胚状体形态形成过程．及胚状体诱导后细胞组织发生的一系列结构变化的时序性特征,这些变化主要影响质体、液泡室、细胞壁和细胞核,进一步分化的程序模拟合子胚的发育。     

Aerodynamics of saccate pollen and its implications for wind pollination

Pollen grains of many wind-pollinated plants contain 1-3 air-filled bladders, or sacci. Sacci are thought to help orient the pollen grain in the pollination droplet. Sacci also increase surface area of the pollen grain, yet add minimal mass, thereby increasing dispersal distance; however, this aerodynamic hypothesis has not been tested in a published study. Using scanning electron and transmission electron microscopy, mathematical modeling, and the saccate pollen of three extant conifers with structurally different pollen grains (Pinus, Falcatifolium, Dacrydium), we developed a computational model to investigate pollen flight. The model calculates terminal settling velocity based on structural characters of the pollen grain, including lengths, widths, and depths of the main body and sacci; angle of saccus rotation; and thicknesses of the saccus wall, endoreticulations, intine, and exine. The settling speeds predicted by the model were empirically validated by stroboscopic photography. This study is the first to quantitatively demonstrate the adaptive significance of sacci for the aerodynamics of wind pollination. Modeling pollen both with and without sacci indicated that sacci can reduce pollen settling speeds, thereby increasing dispersal distance, with the exception of pollen grains having robust endoreticulations and those with thick saccus walls. Furthermore, because the mathematical model is based on structural characters and error propagation methods show that the model yields valid results when sample sizes are small, the flight dynamics of fossil pollen can be investigated. Several fossils were studied, including bisaccate (Pinus, Pteruchus, Caytonanthus), monosaccate (Gothania), and nonsaccate (Monoletes) pollen types.     

美乐多（Melodorum fruticosum Lour．）花粉形态观察

利用扫描电子显微镜（SEM）和透射电子显微镜（TEM）观察了美乐多（Melodorum fruticosum）的花粉形态特征。美乐多花粉为球形或扁圆形的单粒花粉,外壁纹饰为微褶皱状,有点状凹陷,无任何萌发孔或萌发沟。花粉外壁由外壁外层包括覆盖层（连续）、覆盖下层、基足层（1～3层薄片层结构,偶断裂或扭曲至6～10层）和外壁内层（连续）组成。其中,覆盖下层,其厚度为整个花粉外壁厚度的1／2,为混合型结构,即小柱状和颗粒状同时存在,但以颗粒状为主。花粉内壁分为内壁外层和内壁内层,其厚度逐渐变薄。美乐多的花粉特征（单粒、无萌发孔或沟、覆盖层连续、基足层为薄片层结构、花粉外壁内层薄等）与紫玉盘族其他类群一致。     

Effects of ovular secretions on pollen in Pseudotsuga menziesii (Pinaceae)

Effects of ovular secretions on pollen grains were examined in Pseudotsuga menziesii. The exine is cast off in the micropylar canal. A membranelike structure covers parts of pollen grains and appears to protect them. The outer intine consists of fibrous materials, but it also shows a thicker filamentous appearance in some ovules during pollen elongation. The inner intine is electron-dense. Its fibrous nature is occasionally visible. Dissolution of the outer intine varies in amount and manner in ovules from different trees. The plasma membrane near the pollen wall alternatively appears normal and distorted. These different morphologies of the outer intine and of the plasma membrane are considered to result from secretions from the ovule. The outer intine may contain electron-dense globules that are formed in the tube cell and traverse the inner intine. Pollen tube formation appears to be triggered by a secretion from the ovule. Cross-pollinated grains are less distorted compared with self-pollinated grains.     

The Structure of Pollen Wall and Its Relation to Po41en Aggregation in Cymbidium goeringii (Rchb. F.) Rchb. F.

The pollen wall of tetrads located in different positions of a mature pollinium of Cymbidium goeringii was examined with the electron microscope, and the compositions of wall materials were also tested with different histochemical methods. In all tetrads of a pollinium, the pollen wall can be distingished into an exine and an intine, but the exine may be varied greatly according to the tetrad position in a pollenium. The part of the pollen wall (the outer wall) of the external tetrads, lying close, to the tapetum, is composed of two layers, i.e. the exine, and the intine. Theexine consists of tectum, granulate ectexine and endexine, without foot layer. The intine is cellulose in nature. In the outer wall between different groups of: tetrads and in the inner wall within an individual tetrad, the structure of ectexine becomes simple and the deposition of sporopollenin is roduced The degree of reduction of ectexine nicreases from the outer to inner tetrads in several external layers of a pollinium, and even the internal tetrads have a reduced ectexine or lack of it. The present study also demonstrates that the mechanism of pollen aggregation into a pollinium is built on a combined effect of the following features: (1) connected bridges formed' by intine between two pollens within a tetrad, (2) formation of cytoplasmic channels between two pollens within a tetrad, (3) incomplete cell wall formation within a tetrad, (4) little size of tetrads and compact arrangement of mature tetrads and (5) a sticky viscin material surrounded on the outside of a pollinium.     

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。     

Pectin distribution pattern in the apertural intine of Euphorbia peplus L. (Euphorbiaceae) pollen

The apertural inner layer (intine) of Euphorbia L. pollen grains has a characteristic but original structure that has paired thickenings, one on either side of the colpus. To determine the nature and role of this intine layer, pollen grains of Euphorbia peplus L. were germinated in vivo and in vitro. The germination process involves wall changes that facilitate formation of the pollen tube and its subsequent growth. In the thickenings of the intine of E. peplus, the unesterified pectin epitopes are more densely localised in the inner part of the middle intine. No such epitopes are located in the intine portion adjacent to the plasma membrane (cellulosic endintine). Unesterified pectin epitopes are also localised in the outer part of the intine but are restricted to the centre of the aperture, around and in the pore. The de-esterification of pectins is very advanced at the time of dehiscence and pollen germination. The stratification of the aperture intine may take the following pathway at the time of germination: the thin outer zone of the intine in the pore region becomes disorganised and undergoes dissolution with liberation of unesterified and esterified pectins; the middle intine thickenings undergo an important elastic modification, but without liberation of unesterified pectins; the cellulosic inner intine is the progenitor of the pollen tube wall. This special intine of E. peplus is an adaptation to the hydration process preceding germination, increasing intine and pollen grain wall elasticity.     

三种黄花石蒜植物的花粉形态比较

利用光学显微镜和扫描电镜对三种开黄花的石蒜属(Lycoris Herb.)植物的花粉形态进行观察和比较.结果表明:三种植物花粉均为两侧对称,赤道面观呈舟形或肾形,极面观为椭圆形或长椭圆形,萌发孔类型为远极单槽,槽长几达两端;三种植物花粉外壁具网状纹饰,且在网眼底壁及网脊多附有小乳突.通过花粉特征及其他性状的比较研究,对三种黄花石蒜植物的分类鉴定和系统演化关系进行了初步探讨.     

Pollen wall development in Vigna vexillata I. Characterization of wall layers

Light and electron microscope observations characterized the layers that comprise Vigna vexillata L. pollen walls, and identified the timing of their development. Exine sculpturings form an unusually coarse ektexinous reticulum. The structure of the ektexine is granular; this differs from the columellate/tectate type of structure typical of most angiosperm pollen. The ektexine overlies a homogeneous-to-lamellar, electron-dense endexine, which in turn surrounds a thick, microfibrillar intine. Pollen grains are triporate and operculate, with Zwischenkörper and thickened intine underlying the apertures. The ektexine forms during the tetrad period of microspore development, the endexine and Zwischenkörper during the free microspore stage, and the intine during the bicelled (pollen) stage. Coarsely reticulate exine sculpturings and the granular structure of the patterned exine wall of the pollen grains are features that make this species suitable for detailed studies of pollen wall pattern formation.     

Pollen Morphology of Tundra Shrubs and Submarginal Plants from Barrow, Alaska

Investigation of plant morphological features, pollen, and habitat have been made for two shrub species from Barrow, Alaska, namely Dryas integrifoila M. Vahl and Salix rotundifolia Trautv., both of which are endemic to the Arctic floristic area. The former species has small lanceolate or plate leaves, whereas the latter has rounded leaves with distinct veins, rich in vitamin C. Both have dwarf and sprawling habits. Pollen studies showed that the pollen grains of the two species are spheroidal to sub-spheroidal or prolate. The type of aperture was tricolporate; pollen size 26.3-31.3 μm; ornamentation finely reticulate under a light microscope （LM） and strlate-reticulate under a scanning electron microscope （SEM） for D. integrafoila and finely reticulate under the LM and SEM for S. rotundifolia. Comparisons were made between the pollen from the same species from Arctic collections with those from China and Japan. Investigation of pollen morphology of tundra plants can provide significant data for comparative studies of fossil pollen and for the reconstruction of paleovegetation and paleoclimate in the Barrow area.     

Cupressus arizonica pollen wall zonation and in vitro hydration

The structure of Cupressus arizonica pollen at different degrees of hydration was examined by using cytochemical staining and light (LM) and scanning electron (SEM) microscopy. Most pollen grains are inaperturate and a minority are provided with an operculate pore enveloped by a concave annulus. Intine consists of: 1) a thin polysaccharidic outer layer, 2) a large polysaccharidic middle layer that is spongy and bordered by a mesh of large and branched fibrils, and 3) an inner cellulosic thick layer with callose concentrated on the inner side, which forms a shell around the protoplast. The protoplast is egg-shaped with PAS positive cytoplasm and prominent nucleus. Exine splits during hydration and is cast off according to three major steps: 1) the split opens like a mouth and the underlying intine is expelled by swelling like a balloon, 2) the protoplast enveloped by the inner intine is sucked in the outgrowing side, and 3) the backside of the intine gets rid of the exine shell. In water containing salts, exine is rapidly released and the middle intine may expand up to break the outer layer, with disgregation of the spongy material and release of the intine shell including the protoplast. In water lacking salts, the sporoderm hydration and breaking are negatively influenced by the population effect. Pollen when air dried after the exine release become completely flat owing to disappearance of the middle intine layer which may be restored by dipping pollen in water. The results are discussed in relation to the functional potentialities of the sporoderm.