The Pollen Morphology and Exine Ultrastructure of Paeonia L in China

The pollen morphology of 9 species of Paeonia L. has been investigated with both light microscope and scanning electron microscope. In addition, the exine structure of pollen grains of Paeonia suffruticosa and P. lactiflora was examined by transmission electron microscope. Tricolporoidate aperture is an important character of the pollen grains of the Paeonia. The surface of the exine is characterized by reticulate, foveolate and irregularly tuberculate-foveolate sculpture under the SEM. Thin sections of the pollen of this genus shows that the layers of exine are complete i.e. a perforate rectum to semitectum, columellae and foot layers. The endexine is continuous, considerably thickened in the aperture areas and relatively thin or indistinct in the mesocolpia. Paeonia has been placed in Ranunculaceae. But since the beginning of this century many authors have suggested to separating Paeonia from Ranunculaceae. Pollen marphology supports such separation. In Ranunculaceae most pollen grains are tricolpate or have other types of aperture, and exine with spinules and perforations between them. In electron microscopy, the ektexine contains a foot layer, columellae, and perforate rectum, the columellar layer with two types of columellae; the endexine is generally thin. However, the columellar layer of Paeonia has only monomorphic columellae. Some authors considered that there is a close relationship between Paeonia and the Dilleniaceae, but these also differ in the characters of the pollen grains. In Paeonia the constriction of the colpus in equator is in some degree similar to that of Theaceae (Camellia sasanqua Thunb.), Guttiferae (Hypericum L.), Actinidiaceae and Rosaceae. But in the other respects they are quite different. In sum, the pollen morphology of Paeonia is unique. So the palynological information supports Takhtajan's view that Paeonia should be elevated to a family (Paeoniaceae) or order (Paeoniales).     

五列木科和肋果茶科花粉外壁超微结构及其与山茶科的系统学关系

借助光学显微镜、扫描电镜和透射电镜对五列木科Ｐｅｎｔａｐｈｙｌａｃａｃｅａｅ和肋果茶科Ｓｌａｄｅｎｉａｃｅａｅ花粉进行了观察,并与山茶科Ｔｈｅａｃｅａｅ若干属的花粉进行了详细的比较,同时参考了猕猴桃科Ａｃｔｉｎｉｄｉａｃｅａｅ的花粉特征。研究表明,肋果茶科的花粉与山茶科的花粉具有较多的相似性,如其形状和大小与山茶科中厚皮香亚科的几乎一致;而花粉的表面纹饰（粗糙至模糊的皱波状纹饰）则与山茶属,Ｆｒｅｚｉｅｒａ属和厚皮香亚科部分属的花粉纹饰相类似。从花粉外壁的结构看,肋果茶的花粉与石笔木属和山茶属的更接近,表现为：花粉外壁层次分化明显,复盖层和柱状层均较厚,外壁内层薄。而五列木科花粉除了形状和萌发孔类型与山茶科的相似外,其纹饰特征和外壁结构均与山茶科的差异较大,如五列木科花粉表面近光滑,外壁覆盖层较簿,柱状层很不发达,外壁内层相对较厚等。孢粉学上认为,肋果茶科与山茶科具有更为密切的关系,五列木科则与山茶科较疏远。支持把肋果茶科作为一个属置于山茶科内或作为山茶科的一个亚科     

Pollen morphology in thePortulacaceae (Centrospermae)

Pollen of 110 species from 18 genera in thePortulacaceae has been examined by light and scanning electron microscopy, and a representative number by transmission electron microscopy. Three basic pollen types were found: 3-colpate with thick tectum and foot layer with prominent unbranched columellae and an extremely thin endexine; pantoporate with thick tectum and foot layer with branched columellae enclosing pores and an endexine that is one to two layers thick; pantocolpate with thin tectum and foot layer with broad, short unbranched columellae and an inconspicuous endexine. All pollen types, however, have a spinulose and tubuliferous/punctate ektexine. Also, all the genera except three,Calandrinia H.B.K.,Montia L. andTalinum Adanson are stenopalynous. There is, however, no absolute correlation between pollen morphology and geographical distribution, although both the major centre of palynological diversity and the majority of all species with tricolpate grains occur in South America.     

国产蔷薇科李亚科的花粉形态

本文报道了国产李亚科（Prunoideae）10属11种植物花粉形态。花粉近球形至长球形,极轴 20.00－44.12um,赤道轴门17.85-36.95um。极面观三裂圆形,赤道面观椭圆形至圆形。具三孔沟,内孔常为长方形。沟较长,两端较窄。但在 Sinoplagiospermum uniflora中除了3孔沟外,还有周孔沟。花粉外壁明显分化为覆盖层和柱状层,在光学显微镜下表面常模糊,扫描电镜下外壁纹饰均为条纹状。抱粉学结合形态学和细胞学证据说明本亚科为一单系发生的类群。Prinsepia utilis 和Sinoplagiospermum uniflora（Prinsepia uniflora）这两个种在花粉特征和外壁纹饰上差异很大,从而支持将 Prinsepia和 Sinopmplagiospermum分别处理为两个属但不支持将广义的Princepia（含Princepia和Sinoplagiospermum）独立为亚科。此外,由于Exochorda的外壁纹饰同Prunoideae中较原始的类群（如 Laurocerasus）相近,结合细胞学和形态学证据,支持将 Exochord移置至Prunoidea之下。     

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。     

A Study on Pollen Morphology and Ultrastructure of Subfamily Castaneoideae (Fagaceae) in China

The present paper describes the pollen morphology of 45 species of the subfamily Castaneoifeae (including genera Castanea, Castanopsis and Lithocarpus) from China. The pollen grains were all examined with light microscope, scanning electron microscope and transmission electron microscope. Pollen grains of the subfamily are prolate, subprolate or perprolate, (14.7-23.1)× (8.4 -18.9) μm in size, 3-colporate, the exine in 2-layered, 0.9-1.9μm thick, indistinctly ornate, striate-rugulose or crass-striate, sexine and nexine almost equal in thickness, the sexine consists of tectum, bacules and endonexine under TEM. On the basis of very similar pollen shape, pollen size, type of aperture and exine structure and also other characteristics of plant morphology of the genera Castanea, Castanopsis and Lithocarpus, the present authors tend to support the opinion that they all fall into the samesubfamily, Castaneoideae.     

Pollen morphology of the tribe Cynoglosseae of Boraginoideae (Boraginaceae) in China

Pollen morphology of 16 Chinese species representing 7 genera in the tribe Cynoglosseae of Boraginoideae(Boraginaceae) was examined under LM and SEM, and 5 species under TEM. Pollen grains are cocoon-shaped, rarely subprolate, prolate or ovoid, very small, 7～ 15.7μm× 3.5 ~ 13.9 μm in size, P/E = 1.6～2.02; 3-colporate apertures alternate with 3-pseudocolpi, with equatorial endocingulus except those in Bothriospermum; exine surface is usually smooth, with or without perforations in two poles, rarely with tuberculate ornamentation; exine is rather thin and includes ectexine and endexine, while ectexine consists of imperforate tectum, columellae and foot-layer. The tribe Cynoglosseae has many common characters of pollen grains, but there are some differences among genera. A key to the genera is given based on pollen morphology. Bothriospermum, without endocingulus, may be a primitive genus in this tribe, and its pollen morphology is more similar to that of Eritrichieae than to Cynoglosseae, thus it seems more reasonable to put it into Eritrichieae. Solenanthus may be the most advanced genus in the tribe Cynoglosseae for its pollen grains of sub-isopolar and ovoid shape.     

Palynological diversity and major evolutionary trends in Cyperaceae

Pollen and orbicule morphology of 84 species, representing 52 genera from all tribes and subfamilies are investigated, in order to assess the systematic value of palynological data and to determine palynological evolutionary trends in Cyperaceae. A total of 90% of the species are examined for the first time with scanning electron microscopy. Pollen grains of Cyperaceae are oblate spheroidal to perprolate in shape, inaperturate to polyporate with opercula or pontopercula on pori or colpi. We distinguished seven different sexine ornamentation patterns. Orbicules occur in all species investigated. Pollen morphological variation within Cyperaceae is considerable and includes dispersal unit; number, location and degree of differentiation of apertural zones; and sexine ornamentation patterns. In subfamily Mapanioideae both tribes can be characterized by palynological synapomorphies. However, in subfamily Cyperoideae, the observed pattern of variation does not fit the most recent molecular phylogeny due to high levels of homoplasy and polymorphism in major pollen characters.     

Pollen morphology of the subfamily Aurantioideae (Rutaceae)

The Aurantioideae is one of seven subfamilies of the Rutaceae consisting of two tribes, the Clauseneae, containing five genera, and the Citreae, with 28 genera. Each tribe contains three subtribes. The pollen morphology of the subfamily Aurantioideae is described and illustrated for the first time based on light and scanning electron microscopy. Five pollen types have been recognised in the subfamily, based mainly on aperture number and exine ornamentation. The pollen grains show a high degree of intergeneric variation. Pollen grains of Clauseneae are 3-colporate, microstriate or microstriato-reticulate, whereas pollen grains of Citreae are almost always 4/5 colporate with exines varying from microperforate to coarsely reticulate. Congruence between pollen types and the currently accepted classification is discussed, as well as the systematic implications of pollen morphology for the subfamily.     

Studies on Pollen Morphology of Pulsatilla Mill.

Pollen grains of 18 species of Pulsatilla in Ranunculaceae distributed in Asia and Europe were examined by LM and SEM, and exine ultrastructure of tricolpate pollen grains of P. chinensis and of pantoporate pollen grains of P. campanella was examined by TEM. Pulsatilla pollen is divided into four major types based on the aperture character, i.e. tricolpate, di- and tricolpate, pantocolpate and pantoporate. The revolutionary trend of pollen types is as follows: tricolpate→pantocolpate→pantoporate. Surface spinulate and perforate. According to density and size of sptnulae and distribution of perforation, the pollen grains of the genus can be divided into two groups. Thin sections of P. chinensis and P. campanella show endexine thickened at colpi and ora. Ektexine consists of a foot layer, a collumellae layer and a continuous, perforate tectum. The columallae layer is thicker than foot layer and tectum. Pollen morphology of Pulsatilla is similar to that of Anemone, but different in the distribution of spinules and perforation. Pollen information supports Wang’s view about systematic arrangment of species of Pulsatilla in China.     

螺序草属(茜草科)的花粉形态

首次对螺序草属(Spiradiclis B1.)的11种和1变型共16份植物的花粉样品在光镜和电镜下进行了观察。结果表明,本属植物花粉为单粒,小型和中型花粉,近球形至椭球形,辐射对称,常具3孔沟,偶为4孔沟,外壁表面纹饰为孔状。花粉外壁由不连续的具穿孔的覆盖层、柱状结构和基层组成,内外壁明显。此外,本属植物的花粉具有花粉2型和“芽状”结构的现象。螺序草属植物的花粉形态在属下分类上有一定的局限性,但是在个体的系统发育上却可以提供极有价值的信息,如具有花粉单型的种类比花粉二型的种类进化。     

A Note on Remarkable Exines in Acalyphoideae (Euphorbiaceae)

As part of an extensive study of pollen of Euphorbiaceae that combines transmission electron microscopy with scanning electron microscopy, distinctive exines are reported and documented for certain Acalyphoideae. Cheilosa and Neoscortechinia, which comprise the tribe Cheiloseae, are the only Acalyphoideae with an echinate tectum, but their apertures and exine structure do not support a relationship to Oldfieldioideae. In Ditaxis, one of the three mesocolpia is much smaller than the other two and the pollen can be easily distinguished from all other Euphorbiaceae. In SEM, the tectum of Pycnocoma appears almost complete, but in TEM the exine consists of irregular, mostly discrete tectal elements that narrow to points (=columellae) at the interface with the threadlike footlayer. The operculate grains of Alchornea and Boquillonia have exines with a poorly differentiated double layer of columellae in the mesocolpium, but nearer the endoaperture the lower tier becomes greatly elongated and appears to rest directly on the endexine. Plukenetia polyadenia has a complete tectum and a thick exine with a predominant infratectum of large, occasionally branched columellae that nearer the tectum are combined with densely spaced granules. Plukenetia penninervia has a reticulate tectum of crenate muri and short, sparse columellae. The pollen diversity in Acalyphoideae suggests that the subfamily, many tribes and even genera may not be monophyletic.     

Notes on the morphology and fine structure of the exine of some pollen types in dipterocarpaceae

The pollen morphology of nine species belonging to six genera of Dipterocarpaceae is described in detail with light, scanning electron and transmission electron microscopy, while additional data on twenty-eight species are briefly discussed. In Dipterocarpoideae, grains are tricolpate and the exine consists of a thin, often laminated basal layer and an outer, sculptured layer in which columellae and tectum may either be recognizable or fused into a tilioid structure. The surface of the tectal ridges often show a highly characteristic grooved or crenelated microsculpture. In Monotoideae the grains are tricolporate and four layers can be recognized, the inner two being probably equivalent to endexine s.s. and footlayer, the outer two forming a fairly coarse tilioid reticulate sculpture.The evolutionary relations between the pollen types and their relations to taxonomy, wood anatomy and fruit morphology are discussed. A fair amount of correspondence is obvious, although discrepancies, especially with anatomical evidence, exist also.     

Pollen morphology and evolution in the genus Orobanche L. s.l. and its allied genera (Orobancheae/Orobanchaceae) in Turkey

The pollen morphology of 120 samples of the tribe Orobancheae, representing four genera and 40 species of Cistanche, Diphelypaea, Orobanche and Phelipanche native to Turkey, has been studied by light and scanning electron microscopy. Pollen of the tribe Orobancheae is typically isopolar, radially asymmetrical, oblate spheroidal or prolate and belongs to one of the following basic pollen types: inaperturate, tricolpate and syncolpate. Also, pollen heteromorphism is widespread among pest species of Orobanche and Phelipanche. Pollen characters display considerable variation among genera and species, but some characters are significant at the levels of genera and sections. The results of the SEM study show that there are various exine ornamentation types in these genera. The obtained results support the division of the traditionally circumscribed genus Orobanche sensu lato into two genera, Phelipanche and Orobanche. The pollen morphological features are discussed on the basis of molecular phylogeny of the taxa.     

中国紫草科厚壳树亚科的花粉形态及其系统学意义

为探讨厚壳树亚科在紫草科中的系统位置,用扫描电子显微镜研究了国产厚壳树亚科Ehretioideae的厚壳树属Ehretia、轮冠木属Rotula和基及树属 Carmona 3属12种植物的花粉形态,其扫描电镜资料属首次报道。从本文并结合文献报道的双柱紫草属 Coldenia的花粉资料,可得知该亚科花粉为长球形 、近球形,稀球形,赤道部位具有不同程度的膨大;萌发孔类型为三孔沟和三拟假沟二者交替排列,稀具三孔;外壁表面具微弱的小穿孔,稀光滑或皱波状纹饰。厚壳树亚科Ehretioideae与破布木亚科     

木通科、大血藤科花粉壁的超微结构研究

应用透射电子显微镜(TEM)观察了木通科Decaisnea,Sinofr-anchetia,Holboellia,Stauntonia属以及大血藤科Sargentodoxa属共18种植物花粉壁的超微结构。所观察的木通科和大血藤科植物具较发达的覆盖层和柱状层;外壁内层以及内壁均在萌发沟处明显增厚;基层通常不甚发达。与扫描特征相对应的覆盖层结构特征,显示出类群的特异性。在Stauntonia属,覆盖层富于形态变化,反映出该属在木通科中较进化的地位;大血藤(Sarg-entodoxa cuneata)花粉壁结构隶属木通型花粉结构,表明大血藤科与木通科的密切关系。     

Pollen ultrastructure of Panax(the ginseng genus, Araliaceae),an eastern Asian and eastern NorthAmerican disjunct genus

Pollen of ten species of Panax and six species of Aralia was examined in light microscopy and scanning and transmission electron microscopy. Grains of both genera have similar complex apertures, short columellae, and overlapping tectal sculptures, suggesting a close relationship. Most species of Panax have pollen characterized by striato-reticulate tecta, short columellae, thick foot layers, costa ectocolpi, and lalongate endoapertures. The eastern North American P. trifolius, commonly known as the dwarf ginseng, has a distinctive pollen morphology and exine structure, supporting the hypothesis of its phylogenetically isolated position. Pollen of the eastern Asian P. ginseng (ginseng) can be distinguished from the eastern North American P. quinquefolius (American ginseng) by differences in ultrastructure. The monophyly of the three medicinally important species, P. ginseng, P. notoginseng, and P. quinquefolius, suggested by triterpenoid data, is not supported by pollen data. The results of the pollen study are generally congruent with those from the sequences of nuclear ribosomal DNA.     

水雍科植物的花粉形态研究

应用光学显微镜、扫描电镜和透射电镜对世界水雍科6种植物的花粉形态进行了观察。水雍科植物花粉为舟形或船形,具远极单沟萌发孔类型,外壁纹饰通常为浅网状至网状,稀为小刺状纹饰,外壁外层由覆盖层、柱状层和基层组成,覆盖层厚。水雍科植物花粉外壁纹饰表现了从小刺状向网状的过渡。该科花粉为远极单沟,覆盖层厚,具小刺等特征反映了其与水鳖科和泽泻目花蔺科的密切联系,而该科花粉外壁纹饰多为网状则与茨藻目植物接近。这一结果支持将水雍科从茨藻目中分出作为一个独立的目处理的观点。此外,水雍科植物的花粉大小、纹饰类型、网眼大小与深浅等方面的差异对种级分类有一定意义。     

Pollen morphology and ultrastructure of Marathrum schiedeanum (Podostemaceae)

The Podostemaceae, or river-weeds, comprise 46 genera and 270 species of dicots and are the largest family of strictly aquatic angiosperms. Despite the large size, specialized habitats, and enigmatic morphology of the family, relatively little is known about the palynology of Podostemaceae. In the current paper, pollen morphology and ultrastructure of Marathrum schiedeanum are described. Pollen grains are relatively small, spheroidal, and tricolpate to spiraperturate. The exine has a microechinate ornament, a tectate-granular sexine and a relatively thick nexine in non-apertural regions, and a semitectate sexine and thinner nexine in apertural regions. Although aperture variation occurs in the family, this is the first report of the spiral aperture type in Podostemaceae. The spiraperturate condition appears to be derived in river-weeds, as does the granular pollen wall, which represents a reduction of the typical columellae found in eudicots.