五味子科植物花形态及其系统学意义

用光镜和扫描电镜观察了五味子科Ｓｃｈｉｓａｎｄｒａｃｅａｅ２３种、６变种和１变型的９１份花粉的形态。其中５ 、４变种和１变型的花粉形态为首次报道。该科花粉为单花粉,异极,辐射对称,扁球形至超扁球形,其萌发沟的排列格局在被子植物中是独一无二的。若按萌发沟的数目划分,该科花粉可分为３沟型和６沟型两个类型。外壁为网状纹饰,网眼的大小和网脊的宽度与花的形态有一定相关性。研究结果不支持Ｐｒａｇｌｏｗｓｋｉ对     

Pollen morphology of the genus Nothofagus and its taxonomic significance

The present paper describes the pollen morphology of 26 species and 2 varieties in Nothofagus from Australia, New Zealand, New Caledonia, New Guinea and South America. Pollen grains were all examined with light microscope (LM), scanning electron microscope ( SEM ) and transmission electron microscope(TEM). A comparative analysis of pollen exine ultrastructure was made for some species. The results are summarised as follows: Pollen grains are oblate to peroblate, 5～8-short-colpate, rarely 4- or 9-colpate; colpi generally thickened at margins; pollen surface spinulose. The exine ultrastructure of Nothofagus differs considerably from that of the other genera in the Fagaceae. The pollen grains of the species examined here show great differences in shape, size, colpal number and characteristics of colpi at margins and could be divided into three distinct types, i.e. N. brassii type; N. menziesii type and N. fusca type.     

中国南五味子属植物花粉形态的研究

本文用光学显微镜和扫描电子显微镜研究了国产南五味子属2个组7种花粉,记述了它们的形态,讨论了该属花粉形态的特征。该属花粉异极六沟,是属的鉴别上的一个重要特征,其中三条沟常在一极汇合成三合沟,本文报道亦有四合沟的现象,认为具合沟的一极是远极。南五味子属Kadsura Kaempf.ex Juss.集中分布于东亚至东南亚,约有24种,我国种类最为丰富,已知有13种,大都为药用经济植物。本属植物的花粉形态在国内尚未见有报道,本文试图通过对本属广布种和我国一些特有种的花粉形态的观察,探讨这个属花粉形态的特征,并与其他有关类群的花粉比较,给植物分类提供依据,为解决某些争议性问题提供资料,同时提出作者的观点。     

A study on pollen morphology of the genus Vitis L.

The pollen grains of 32 species, 1 subspecies and 4 varieties of Vitis and 1 species of Muscadinia were scanned with Hitachi S-800 scanning electron microscope and described in the present paper. In Vitis, the pollen morphology is found to be valuable for the identification of species, though it is of little significance in supraspecific classification and systematic consideration. The ornamentation of Vitis could be divided into four types: ( 1 ) The pollen grains are oblate or rarely spheroidal (P/E =0.97 ～ 1.08) in shape, with colpi wide enough to show dense granulae inside. The pollen grains of V. coignetiae Pull. Ex Planch. belong to this type. (2) The pollen grains are prolate to subprolate (P/E = 1.25 ～ 1.76 ), with sparse granulae within the colpi. This type includes l0 species, 1 subspecies and 3 varieties, such as V. hui Cheng etc. (3) The pollen grains of V. romanetii Roman du Caill. ex Planch. and V. davidii (Roman du Caill. ) Foex. var. ferruginea Merr. et Chun are prolate (P/E= 1.87 ～ 2.1), with colpi narrow and linear-shaped, but rhombic in the equational region. (4) The pollen grains are prolate or perprolate (P/E= 1.30 ～ 2. 0), with narrow and linear-shaped colpi. The species with this type of pollen grains are Muscadiniarotundifolia (Michx.) Small and 20 species of Vitis.     

Pollen Morphology of Loxostemon (Cruciferae) in China

The present paper deals with the pollen morphology of 10 species and 1 variety of Loxostemon in China. The pollen grains were all examined under light microscope. The pollen grains of Loxostemon are subspheroidal, spheroidal or prolate, 18--33×11.8-28 μ in size, 3-colpate, colpi 15-21 μ long and 1-2 μ wide. The exine is 1.5-3 μ thick with two indistinct or distinct layers. All the pollen grains are generally reticulate under light microscope. They are distinctly or obscurely and finely reticulate. L. axillus and L. repens are generally similar in gross morphology, but the pollen grains of these two species are different. The pollen grains of L. axillus are regularly polygonally reticulate, colpi are acute-ended and the exine is about 3 μ thick, whereas those of L.repens are irregularly polygonally reticulate, colpi are enlarged at both ends and the exine is about 2.8 μ thick. L. incanus and L. stenolobus appear to have similar gross morphology, but the pollen grains of the former have exines with two distinct layers and a densely and finely reticulate ornamentation and those of the latter have exines with two indistinct layers and a flexuosely reticulate ornamentation.     

Pollen Morphology of the Genus Malus and Its Taxonomic and Evolutionary Significance

The pollen morphology of 26 species and 5 hybrids of the genus Malus was investigated with aid of SEM. It is found that the pollen morphology of Malus is rather similar in shape, size, position and number of aperture and exine sculpture. The characters of polle morphology of sections and series are as follows: Sect. Malus Ser. Baccatae: Striae regular, parallel to colpi, and conjunct at pole; Ser. Pumilae: the same as in Ser. Baccatae, but more or less curved near pole. Sect. Docyniopes: Striae regular and parallel to colpi, but bent near pole. Sect. Chloromeles: Striae irregular, dense and interlock. Sect. Sorbormalus: Striae irregular, sparse and not interlock. Ser. Sieboldiance: pollen grains prolate, with perforation among striae, colpi narrow; Ser. Kansuenses: Pollen grains spheroidal, colpi wide in the middle but narrow at both ends, striae relatively dense and regular, mostly dichotomous, perforation present; Ser. Yunnanenses; Pollen grain spheroidae, colpi wide in the middle but narrow at both ends, striae obviously irregular, less dichotomous, perforation absent. The major evolutionary trend of exine sculpture of pollen may be from densely thin-striate to sparsely striate with perforation. Characters of the exine sculpture of hybrids can be used to recognize the rela-tionship between the parental species.     

中国壳斗科栎亚科花粉形态研究

本文应用光学显微镜与扫描电子显微镜观察了壳斗科所属栎亚科2属26种2变种的花粉形态,并以本科其它属一些代表种作对比,试图从孢粉学方面为栎亚科的系统分类提供依据。 花粉外壁纹饰在扫描电镜下可分三种类型,即水青冈型(含水青冈属),栎型(含栎属和三棱栎属),栗型(含栗属、锥属和柯属),花粉的类型与植物形态分类的三个亚科一致。 栎亚科的花粉外壁纹饰在扫描电镜下为颗粒聚集成的种种形状。栎属可区分为四种类型,即颗粒状、颗粒——蠕虫状、聚颗粒及芽孢状,其中类型一、二、三为常绿种类,类型四为落叶种类。花粉纹饰在常绿与落叶种类之间有较明显的差异,而青冈亚属和栎亚属之间却没有界线。因此,栎属仍以包含两个亚属为宜。三棱栎属花粉纹饰则介于栎属的类型二与三之间。     

Taxonomy of the Genus Oreocharis (Gesneriaceae)

The genus Oreocharis as circumscribed here consists of 27 species including 5 varieties, of which 5 species and 4 varieties are described as new in the present paper. In the work analysed were the external morphology and geographic distribution and examined under SEM were pollen exine of 22 species and seed coat of 16 species. As a result, three types of the corolla, two types of the anther, three types of the pollen exine and three types of the seed coat are distinguished here in the paper. It is discovered that the corolla in the genus is relatively stable, though diverse, and highly correlated with the characters of pollen grains and seeds. The corolla clearly bilabiate but constricted at the throat, occurring in O. auricula, O. cordatula, O. aurantiaca, etc., for an example, is correlated with smooth, reticulate pollen exine and partial tectum and the reticulate and smooth seed coat. For this reason the subdivision of the genus in the paper is mainly based on the characters of the corolla, but combined with those of the anther, pollen and seed coat. The genus is divided into four sections in the present classification. Dasydesmus Craib, based on a single species. O. bodinieri, is reduced here, and the reasons are given. The genus is distributed mainly in the subtropics, and less frequently in the tropics, of China south of 32.5°N and east of 98.5°E, with only two species beyond the border, O. hirsuta in Thailand (only a single locality in Chiengmai) and O. aurea also found in north Vietnam (see Fig. 1, Table 3). Sect. 1. Stomactin (Clarke) Fritsch. Corolla urceolate-tubular, constricted at the throat, with limb distinctly bilabiate; anthers broad-oblong; seed coat reticulate, smooth, rarely minutely tuberculate; pollen exine fine-reticulate, tectum partial and smooth, luminae slightly unequal in size. Sect. 2. Orthanthera K. Y. Pan Corolla campanulate or campanulate-tubular; anthers broad-oblong; seed coat reticulate, muri smooth, rarely spiny-processed; pollen exine fine-reticulate, with partial and smooth tectum and luminae slightly unequal in size, rarely exine insular and fine-tuberculate, tectum perforate. Setc. 3. Oreocharis Corolla thin-tubular; anthers broad-oblong; seed coat densely spinyprocessed, rarely fine-tuberculate; pollen exine insular, densely spiny-processed, rarely finereticulate and smooth, luminae unequal in size. Sect. 4. Platyanthera K. Y. Pan Corolla campanulate; anthers hippocrepiform; seed coat densely spiny-processed; pollen exine fine-reticulate, tectum perforate, luminae small, nearly equal in size. In the section Stomactin, although the constriction of corolla at its throat is a specialized character, the characters of seed coat, pollen grains and anthers are apparently primitive. Therefore it may be said at least that more primitive characters are preserved in the section. In the section Oreocharis, on the contrary, the characters of corolla, seed coat and pollen exine are all advanced. And in the section Platyanthera, the seed coat, pollen (with perforate tectum) and anthers have developed rather specialized characters.     

囊萼紫草属与滇紫草属花粉形态比较研究

本文借助光学显微镜和扫描电镜研究了囊萼紫草属3种和滇紫草属12种植物的花粉形态。囊萼紫草属的花粉为哑铃形或茧形,中等大小,P／E比为1．6一1．67,三孔沟,内孔横长;具小刺状纹饰。滇紫草属的花粉为近长球形或近卵球形,P／E为l—1．23;三孔沟或三合沟孔,内孔一般纵长,具皱波状纹饰,在皱波上具密集的小瘤或微颗粒。从花粉形态的角度,本文支持把囊萼紫草属从滇紫草属(广义)中分离出来的观点。值得注意的是,在滇紫草属的花粉中首次观察到了一种比较少见且特化的花粉即单极三合沟孔的花粉。     

Pollen Morphology of the Family Zingiberaceae in China—Pollen Types and Their Significance in the Taxonomy

Pollen morphology of 89 species and 3 varieties belonging to 18 genera (out of 150 spcies in 20 genera) of Zingiberaceae in China was studied under both light microscope and scanning electron microscope. Pollen grains of Zingiberaceae are spherical, subspherical, ovoid and prolate, 36-225 μm in size, nonaperturate or aperturate (spiraperturate, porate). Pollen grains are almost not resistant to acetolysis. The wall is composed of a very thin exine and a thick intine. The exine is psilate, spinate, cerebelloid-areolate, striate, verrucate and foveolate. According to the presence or the absence of aperture and differential ornamentations, two types and six subtypes are recognized: I. The type Nonaperturate: (85 species and 3 varieties in 18 genera). Four subtypes can be recognized within the type based on the characteristics of the exine sculpture. These are: (1) The subtype Psilate, in which, the exine is nearly smooth (including: Hedychium, Curcuma, Kaempferia, Caulokaempferia coenobilis, Boesenbergia rotunda, Stahlianthus, Amomum compactum, Etingera, Hornstedtis, Rhynchanthus). (2) The subtype Spinate, which comprises two groups: (A) The group Short-spinate, pollen grains with smaller spines (Globba), (B) The group Long-spinate, pollen grains with longer spines (Alpinia, Amomum, Plagiostachys, Roscoea, Cautleya, Boesenbergia fallax, Caulokaempferia yunnanensis). (3) The subtype Cereblloid-areolate, pollen grains of which are spherical or subspherical, with cerebelloid sculpture (Zingiber Sect. Zingiber). (4) The subtype Striate, pollen grains of which are prolate or oliveshaped, and striate (Zingiber Setc. Cryptanthium). II. The type Aperturate, in which pollen grains are acetilysis-resistant and possess distinct apertures (mixed colpate-porate or forate), including two subtypes: (1)The subtype Mixed colpate and Porate. Pollen grains are both 3-colpate and 1-3-porate, and usually with one long spiral, two short (straight or slightly curved) colpi and 1-3-poris. The exine is verrucate or not, nearly sinuolate (Costus speciosus, C. tonkinensis, C. lacerus). (2) The subtype porate, whose grains are 6-8-porate and exine is foveolate (Costus megalobractea). The taxonomic significance of the pollen types in the family Zingiberaceae is also discussed.     

中国芸香科植物的花粉形态

本文用光学显微镜对我国21属52种2变种芸香科(Rutaceae)植物的花粉形态进行了详细地观察研究,其中对6属6种进行了扫描电镜观察。本科花粉具沟孔,是花粉单类型的科之一。根据萌发孔的数目和外壁纹饰特点,本文将芸香科的花粉分为具3孔沟和4-6孔沟两个花粉类型,并根据各属花粉形态特征,作出了花粉形态分属检索表。此外,本文还对花椒(Zanthoxylum L.)和Fagara两属的合并问题及花椒与吴茱萸(Evodia Scop.)两属之间的亲缘关系等问题进行了讨论。     

Pollen Morphology of Asarum in China

The present paper describes the pollen morphology of 27 species, 4 varieties and 1 form of Asarum from China. The pollen grains were all examined under light microscope and scanning electron microscope. The apertures of pollen grains of Asarum are variable and so distinct as to allow the identification of individual subgenera and species. The exine ornamentation is compound and may be divided into two types: reticulate or cerebelloid under verrucae. The pollen grains of Subgen. Asarum and Subgen. Heterotropa are distinguishable, which supports the taxonomical subdivision based on the gross morphology. The differences in the pollen morphology between A. caulescens Maxim. and A. sieboldii Miq. in China and in Japan are discussed and some comparisons in the pollenmorphology between several species similar in the gross morphology are also made.     

Pollen morphology in the genus Coffea (Rubiaceae) and its taxonomic significance

The pollen morphology of 38 representative species and varieties within the genus Coffea L. is described. Eight pollen types, placed in two major groups, have been identified on the basis of the number of colpi, colpus characteristics, exine morphology and pollen size. Differences in pollen structures of the species studied do not correspond to present taxonomic groupings, but support a current proposal to place some of the species in the genus Paracoffea Leroy. Among the four sections in the genus, the section Eucoffea contains a high degree of pollen polymorphism.     

A palynotaxonomic study of the genus Filipendula (Rosaceae)

Pollen grains from 15 species (18 taxa) of the genus Filipendula were examined with light and scan-ning electron microscopy. It was revealed that the pollen grains are isopolar, tricolporate, with scabrate or scabrate-microechinate surface. The pollen morphology was compared with the conventional classification sys-tems of the genus by different authors, and supported Shimizu's system (1961), in which the genus was divided into three subgenera. The monotypic subgen. Hypogyna is characterized by pollen lacking fastigium and thickened costae colpi. The other monotypic subgen. Filipendula differs from others by pollen having larger grain, larger pore size, longitudinally elliptic fastigium and thickened costae colpi. The largest subgen. Ulmaria is distinguished by pollen having rounded or latitudinally elliptic fastigium and thickened costae colpi. Sectional classification was not supported by the pollen morphology due to insufficient variability.     

栝楼属花粉形态研究及其在分类学上的意义

本文收集了国内外栝楼属Trichosanthes 31种1变种,在光学显微镜和扫描电镜下,进行了花粉粒形态的比较观察。该属花粉粒为3孔沟型,外壁表面纹饰可分为四个类型,即：疣状或皱波状(小苞组),粗网状(大苞组),细网状或光滑(叶苞组),近光滑或皱波状(王瓜组)。这四个类型的划分与植物形态分类基本一致,可作为分组及分种的依据之一。花粉特征支持将叶苞组分为叶苞亚组和柔毛亚组,如叶苞亚组有明显的沟,而柔毛亚组没有。     

Micromorphological features of the seed surface of Schisandraceae and their systematic significance

Micromorphological features of the seed surface of the Schisandraceae are reported for the first time. One hundred and seventeen seed samples from 92 populations, representing 24 species, seven varieties and one form of the Schisandraceae, were examined and photographed under scanning electron microscope (SEM). Micromorphological features of seed surfaces of the Schisandraceae are little affected by the habitats in which plants grow, and are quite constant within species, therefore they can be used as reliable diagnostic characters to distinguish species. The following taxonomic treatments are not supported by micromorphological features observed: ( 1 ) To reduce Schisandra wilsoniana A. C. Smith to S. henryi Clarke; (2) To reduce S. neglecta A. C. Smith, S. arisanensis Hayata, S. viridis A. C. Smith, S. sphenanthena Rehd. & Wils., S. gracilis A. C. Smith, S. micrantha A. C. Smith and S. lancifolia var. polycarpa Z. He to S. elongata ( Bl. ) Baill.; (3) To reduce S. henryi var. longipes (Merr. & Chun) A. C. Smith, S. tomentella A. C. Smith and S. pubescens var. pubinervis (Rehd. & Wils.)A. C. Smith to S. pubescens Hemsl. & Wils.; (4) To reduce S. rubriflora(Franch.)Rehd. & Wils., S. flaccidiramosa C. R. Sun, S. incarnata Stapf, S. sphaerandra Stapf, S. sphaerandra f. pallida A. C. Smith and S. glaucescens Diels to S. grandiflora ( Wall. ) Hook. f. & Thoms.; (5) To reduce Schisandra wilsoniana to S. bicolor Cheng; (6) To reduce S. lancifolia var. polycarpa to S. neglecta; (7) To raise S. henryi var. longipes to S. longipes(Merr. & Chun)R. M. K. Saunders; (8) To reduce Kadsura polysperma Yang to K. heteroclita(Roxb. )Craib. The relationship between Schisan-dra Michx. and Kadsura Kaempf. ex Juss. revealed by the micromorphological features of seed surface is very similar to that revealed by pollen morphology, namely these two genera might have originated from a common ancestor and then evolved along two different routes. However the probability of a more complicated relationship between the two genera than we have known hitherto can not be excluded. The general evolutionary level of Kadsura seems to be higher than that of Schisandra. Therefore, the viewpoint that Kadsura is more primitive than Schisandra is not supported.     

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.     

南五味子属花的形态及其系统学意义

借助扫描电镜观察了南五味子属(Kadsura)中黑老虎(K.coccinea)、南五味子(K.japonica)和狭叶南五味子(K.angustifolia)雄花和雌花的形态发生过程, 三种植物花发育早期相似而发育后期出现分异。根据花形态发生早期的相似性, 支持南五味子属为单系起源;根据花形态发生后期的分异状态, 支持将南五味子属分为离蕊南五味子亚属(Subgenus Cosbaea)和南五味子亚属(Subgenus Kadsura)。借助扫描电镜观察了短梗南五味子(K.borneensis)和披针叶南五味子(K.lanceolata)花的形态, 将南五味子和狭叶南五味子的花与短梗南五味子和披针叶南五味子的花比较, 赞成在南五味子亚属下设南五味子组(Section Kadsura)和南洋南五味子组(Section Sarcocarpon)。根据在狭叶南五味子中观察到的两性花痕迹和在五味子属(Schisandra)东亚五味子(S.elongata)花中观察到的两性花, 本文认为现存五味子科(Schisandraceae )植物的单性花可能由具两性花结构的祖先演化而来。根据五味子科植物柱头与八角科(Illiciaceae)植物柱头的相似性, 不支持将五味子科从八角目(Illiciales)中分出而成立五味子目(Schisandrales)的观点。     

毛茛叶报春组一存疑种花粉形态的多样性研究

对分布于浙江西北部的毛茛叶报春组(sect. Ranunculoides)一存疑种5个居群共50个植株的花粉形态进行了研究观察。结果表明该存疑种的花粉形态特征在种下具高度多样化,根据萌发孔类型、数量及组成的不同,其花粉可划分为多沟型、沟多孔少型、孔多沟少型、散孔型和环沟型5种类型;而且居群间花粉的大小及花粉类型的组成也存在较大的差异;此外,同一个植株或同一个花粉囊的花粉也可能具有多种类型的花粉。本文进一步证明了花粉形态特征在种内可能存在多样化的现象。花粉形态特征支持该存疑种为一个独立的新物种,它很可能是由具多沟型和散孔型花粉的亲本杂交进化而来。     

Microsporogenesis, microgametogenesis and pollen morphology of Passiflora spp. (Passifloraceae)

Microsporogenesis, microgametogenesis and pollen morphology of six species of genus Passiflora L. belonging to three subgenera ( Passiflora , Dysosmia , Decaloba ) were studied with light and scanning microscopy; P. caerulea was also examined with transmission microscopy. The tapetum is secretory, microspore tetrads are tetrahedral and pollen grains are two-celled when shed. Small Ubisch bodies are attached to a peritapetal membrane; they are a product of tapetal activity and the rough endoplasmic reticulum (ERr) appears to be involved in their origin. The pollen grains of all the species are subspheroidal, zonocolpate, geminicolpate. Each pair of colpi anastomoses at the poles. The exine is semitectate, reticulate, heterobrochate. The muri are simplibaculate, wavy. The lumina have clavate bacula of varying height. The colpus structure is similar to that of the lumina but generally with fewer and smaller bacula. Lumina size and amount of bacula inside the lumina vary between subgenera. The grains from subgenera Passiflora and Dysosmia differ from those of Decaloba in their size and number of colpi. The pollen and microsporangium morphology of the species of subgenera Passiflora and Dysosmia are more similar than those of subgenus Decaloba . The results are discussed in relation to the current taxonomic classification.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society of London , 2002, 139 , 383–394.