Pollen Characters of Old World Gesneriaceae (Cyrtandroideae)

Pollen of 108 species out of 18 genera (from all tribes of the Old World Gesneriaceae [subf. Cyrtandroideae]) was examined using light, scanning and transmission electron microscopy. The pollen grains are small sized, isopolar and 3-colpate or 3-colporate. In equatorial view they are mostly spheroidal (rarely suboblate or oblate), in polar view circular or subangular. The most variable character is the exine structure and -sculpture. The tectum is perforate, microreticulate, coarsely reticulate or rugulate. Further important characters of the tectum are (a) the presence or absence of conical supratectate sculptural elements, and (b) the width of lumina being either equal or exhibiting different at the apo- and mesocolpium. Ten exine types are distinguished. Some genera and species can be well referred to a special exine type, e.g., Aeschynanthus, Epithema, Stauranthera grandiflora; in other genera several exine types occur, e.g., in the large and heterogeneous genus Didymocarpus. The pollen morphology of the two large subfamilies Cyrtandroideae and Gesnerioideae is compared.     

国产爵床科芦莉花族植物的花粉形态

报道了国产爵床科Acanthaceae芦莉花族Ruellieae芦莉花亚族Ruelliinae 2属7种、假杜鹃亚族Barlerinae 1属3种和马蓝亚族Strobilanthinae 16属34种植物扫描电镜下的花粉形态.芦莉花亚族的地皮消属Pararuellia和喜花草属Eranthemum的花粉均为圆球形,具3孔或3孔沟,外壁为不同的网状结构; 假杜鹃亚族的假杜鹃属Barleria的花粉为长球形,具3孔沟,外壁亦为网状结构;马蓝亚族植物(包含广义的马蓝属Strobilanthes s.l.)花粉形态多样,结构复杂.依据花粉萌发孔和外壁纹饰特征,可将马蓝亚族16属植物和上述两亚族3属植物的花粉形态归纳成3大类型: 1. 具3孔类型.其中又有(1)外壁具网状纹饰者,见于地皮消属; (2)外壁具芽胞状纹饰者,见于黄猄草属Championella; (3)外壁具刺状(棒状)纹饰者,见于南一笼鸡属Paragutzlaffia、叉花草属Diflugossa和假蓝属Pteroptychia.2. 具3孔沟及具3孔沟与假沟类型(肋条带型).其中又有(1)具3孔沟和网状纹饰者,见于喜花草属和假杜鹃属; (2)具刺状(棒状)纹饰者,见于南一笼鸡属、叉花草属和假蓝属; (3)具3孔沟与假沟,外壁纹饰具节隔、肋条带状或网状,网眼纵向排列成行,网眼内有细网纹者,见于耳叶马蓝属Perilepta、马蓝属Pteracanthus(大部分)、金足草属Goldfussia、紫云菜属Strobilanthes(部分)和合页草属Sympagis; (4)具3孔沟与假沟类型,肋条带状,但不具节隔,外壁纹饰网状,网眼不成行或不明显纵向排列,网内无细网纹者,见于尖蕊花属Aechmanthera、板蓝属Baphicacanthus、马蓝属(部分)和糯米香属Semnostachya; (5)具双脊及细网状纹饰者,见于环毛紫云菜Strobilanthes cycla.3. 具(4-)5孔沟及假沟类型(肋条带型),外壁具网状或拟网状纹饰,见于腺背蓝属Adenacanthus.另外兰嵌马蓝属Parachampionella、山一笼鸡属Gutzlaffia和肖笼鸡属Tarphochlamys的花粉有无萌发孔尚不清楚,有待进一步研究.综上所述,芦莉花族植物的花粉形态具有较高的多样性,是重要的分类性状.利用花粉形态特征能较好地区分高级分类群如亚科、族以及亚族,有时也有助于阐明类群之间的相互关系,甚至也能用于区分属、种和阐明其关系.     

刺五加、无梗五加花粉的亚显微形态学研究

用扫描电子显微镜观察了刺五加和无梗五加花粉的亚显微形态特征。花粉粒的赤道面观呈近圆形,极面观呈三裂圆形,具三孔沟。无梗五加、刺五加雄株的花粉表面纹饰为网状;刺五加两性株的花粉的表面纹饰亦呈网状,但有些花粉网脊不连续。无梗五加的花粉粒大小为26.5μm×27.4μm;刺五加雄株的花粉粒大小为21.1μm×22.1μm;刺五加两性株的花粉粒大小约为21.6μm×23.9μm。研究结果表明,无梗五加和刺五加雄株花粉的亚显微形态特征支持传统的属级和种级水平的分类;刺五加两性株某些花粉粒亚显微形态的独特表现可能与其发育不正常有关。     

Pollen Morphology of Guttiferae in China

The present paper deals with the pollen morphology of 23 species belonging to 6 genera of Guttiferae occurred in China. All pollen grains have been observed under light microscope, and those of 11 species in 5 genera have been examined with SEM as well. The pollen grains of Guttiferae are subspheroidal, spheroidal or prolate, (17–43) × (15–26)μ in size. Mostly 3-eolporate, less frequently 4-colporate or 5-colporate. The exine is 2-layered, but the occassionally demarcation of these layers is generally indistinct. The exine is 0.8–2.6 μ in thickness. The oramentations of all the pollen grains is generlly finely reticulate under the light microscope. Pollen morphology of the 11 species of 5 genera observed under the SEM is stressed. Their exine oramentations may be classified into four major groups, ie. reticulate, finely reticulate, baculate and perforate. Calophyllum: pollen grains reticulate and rugulo-reticulate. Cratoxylon: pollen grains reticulate, with finely and densely granulate over entire muri. Gareinia: pollen grains baculate, reticulate, and exine with perforate oramentation; in reticulate grains, muri with spinulation. Hypericum: pollen grains reticulate, perforate and finely reticulate. Mesua: pollen grains finely reticulate. From the comparison of pollen morphology, Hypericaceae has much connection with Guttiferae (sensu stricto). Because their pollen grains are all 3-eolporate and the exine is reticulate or oerforate under light mieroseope, but it is different between the pollen grains of Hypericum and those of Guttiferae.     

Pollen morphology of the tribe Ruellieae (Acanthaceae) from

Pollen morphology of seven species in two genera in the Ruelliinae, three species in one genus in the Barlerinae and 34 species in 16 genera in the Strobilanthinae (Ruellieae, Acanthaceae), were observed under SEM. Pollen grains in Ruellieae, particularly in Strobilanthinae (including Strobilanthes s.l.) exhibit great diversity and are most eurypalynous in the family. In light of the aperture types and exine ornamentation patterns, pollen grains of the genera examined fall into three major types: 1. 3-porate pollen, which includes, (1) pollen with reticulate exine ornamentation (Pararuellia); (2) pollen with gemmate exine ornamentation (Championella); (3) pollen with echinate exine ornamentation (Paragutzlaffia, Diflugossa and Pteroptychia). 2. 3-colporate pollen or 3-colporate pollen with pseudocolpi, which includes, (1) 3-colporate pollen with reticulate exine ornamentation (Eranthemum and Barleria); (2) 3-colporate pollen with pseudocolpi and ribbed or banded, and septate exine ornamentation, the lumina rounded, finely reticulate inside and arranged in longitudinal rows (Perilepta, Pteracanthus, Goldfussia, Sympagis and some species of Strobilanthes); (3) 3-colporate pollen with ribbed or banded yet not septate exine ornamentation, the lumina neither distinctly rounded and finely reticulate inside, nor conspicuously arranged in longitudinal rows (Aechmanthera, Baphicacanthus, Semnostachya and some species of Pteracanthus); and (4) 3-colporate pollen with ribbed and finely reticulate exine ornamentation but with broader bands, each band with two ridges (Strobilanthes cycla). 3. (4-)5-colporate pollen with pseudocolpi and reticulate or ornate exine ornamentation (Adenacanthus). No distinct apertures were observed in three genera, Parachampionella, Gutzlaffia and Tarphochlamys. Pollen morphological characters in the Acanthaceae can be used not only to distinguish taxa of higher ranks (subfamily, tribe and subtribe) and elucidate their relationships, but sometimes can also be used to distinguish genera and species.     

Pollen morphology of the genus Gagea (Liliaceae) in Iran

Pollen grains of 26 species of the genus Gagea distributed in Iran were examined by light and scanning electron microscopy. Detailed pollen morphological characteristics are given for these species. Among the studied species, the newly described G. iranica together with G. olgae and G. graminifolia possess the smallest pollen grains, and the widely distributed G. lutea the largest ones. Our studies show that the sculpturing of exine provides valuable characters for separating the species, sometimes even for closely related ones, and delimitation of natural groups within the genus. The exine of the genus Gagea is in most cases perforated upon tectum or rarely tectate-columellate. The muri are solid or structured, compound, simpli-, dupli- or pluricolumellate. It seems that the structure of muri is very important in recognizing natural groups within the genus. Tectal perforations vary from <0.2 to 2.0 μm in diameter among the studied species. Regarding sculpturing of the exine in proximal face, four basic types of pollen grains can be distinguished: reticulate, microreticulate, foveolate and perforate. Within the reticulate type there is sufficient variation in exine structure at distal face to describe three subtypes: reticulate, microreticulate and perforate. A diagnostic key is given for all studied taxa based on palynomorphological characters. For a limited number of populations of selected Iranian species of Gagea, further aspects of pollen biology were studied. It seems that populations with ploidy levels other than diploidy, show a low percentage of pollen fertility. Moreover, the rate of pollen fertility is correlated with the manner of the reproduction in certain species.     

国产爵床科鳞花草族1属、穿心莲族2属和爵床族8属植物的花粉形态

报道了中国爵床科Acanthaceae爵床亚科Ruellioideae鳞花草族Lepidagathideae 1属、穿心莲族Andrographideae 2属和爵床族Justicieae 8属共14种植物的花粉形态。根据花粉萌发孔类型和外壁纹饰特征,可将这些植物的花粉归为4个类型: (1)具3孔沟,外壁具网状纹饰。该类型见于色萼花属Chroesthes (采自广西样品)和穿心莲属Andrographis植物。(2)具3孔沟及假沟(肋条带型)类型,外壁具网状纹饰,网眼纵向排列成行,网眼内有细网纹。该类型见于色萼花属(采自云南样品)和鳔冠花属Cystacanthus。(3)具3孔沟,沟两侧各有1条假沟,外壁具网状纹饰。该类型见于瑞丽山壳骨Pseuderanthemum shweliense、狗肝菜属Dicliptera和驳骨草属Gendarussa植物。(4)具2孔沟,沟两侧具斑块。该类型见于山壳骨属Pseuderanthemum 1种、纤穗爵床属Leptostachya、鳄嘴花属Clinacanthus、鸭嘴花属Adhatoda、杜根藤属Calophanoides和野靛棵属Mananthes植物。     

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.     

Pollen morphology of Chinese Glochidion (Phyllanthaceae) and its taxonomic implications

The pollen morphology of twenty‐five Chinese species belonging to two sections of Glochidion and one species from Phyllanthus subgen. Phyllanthodendron was examined using scanning electron microscopy (SEM). The results showed that pollen grains of Glochidion are isopolar, spheroidal to rarely subprolate in shape, (3)4–5(6)‐colporate in aperture pattern, and tectate‐reticulate to tectate‐rugulate in exine ornamentation. Fusion of colpus ends was observed in some pollen grains of G. pseudo‐obscurum and G. wrightii. The pollen grains of Phyllanthus dunnianus are similar to those of Glochidion species in their spheroidal shape and four‐colporate aperture pattern, but are slightly larger in size, and show tectate‐perforate exine ornamentation. In Glochidion, pollen morphology appears to be useful for distinguishing several macromorphologically similar. Additionally, palynological characters provide evidence for the taxonomy of the supergenus Phyllanthus s.l.     

Taxonomic implications of pollen exine morphology in infrageneric classification of Scutellaria (Lamiaceae)

Scutellaria L. in Lamiaceae subfamily Scutellarioideae is a subcosmopolitan genus with about 360 species. According to the latest subgeneric classification, two subgenera are recognized: S. subgen. Scutellaria and S. subgen. Apeltanthus. We studied pollen grains from 29 species of Scutellaria belonging to both subgenera and from both the Old and the New World to evaluate their taxonomic importance for the infrageneric classification of the genus. Pollen grains were acetolysed and studied by scanning electron microscope (SEM) and light microscope (LM). All examined pollen grains are isopolar, tricolpate, suboblate, oblate spheroidal, prolate spheroidal to subprolate. The exine is mainly bireticulate perforate, but with variations in lumina shape and size as well as perforation size and uniformity. Based on the exine sculpturing, two subtypes are recognized within the bireticulate pollen type. In subtype I the primary reticulum of the exine consists of angular muri; the lumen has an irregular polygonal shape and the perforations of the secondary reticulum are more or less uniform and of the same size. This pollen subtype was observed in species of subgenus Scutellaria, sectt. Scutellaria p.p., Anaspis, Perilomia and Salazaria. In subtype II the primary reticulum of the exine consists of a more or less curved murus; the lumen is rounded or obtusely angular and the secondary reticulum consists of one or more large central perforations surrounded by some smaller ones. This pollen subtype was observed in species of subgenus Scutellaria sectt. Scutellaria p.p., Salviifoliae and in all members of subgenus Apeltanthus. The results of this study reveal that the exine ornamentation is a diagnostic character useful for the infrageneric classification of Scutellaria.     

Pollen morphology supports the transfer of Wendlandia (Rubiaceae) out of Rondeletieae

Pollen morphology of 27 species, eight subspecies and one variety of Wendlandia was studied using scanning electron microscopy (SEM). Wendlandia pollen are monads, radiosymmetric, small in size, tricolporate (rarely tetracolporate or bicolporate) and spheroidal (rarely subprolate or suboblate) in equatorial view. The compound aperture consists of ectocolpus, mesoporus and endocolpus. In addition, reticulate sexine and granular nexine were observed. The pollen wall ultrastructure of two Wendlandia spp. was examined by transmission electron microscopy (TEM). The exine consists of the tectum, columellae, foot layer and endexine. The endexine is thickened into a costa around the aperture. The intine forms a protruding oncus at the aperture. The palynological characters show a remarkable uniformity among the Wendlandia spp. Differences with Rondeletia, the main genus of tribe Rondeletieae, exist in the exine pattern, the endoaperture and the pollen wall structure. Our observations indicated that the endoaperture type and the structure of the pollen wall of Wendlandia were similar to those of the Gardenieae–Pavetteae–Coffeeae–Octotropideae clade, which provided palynological evidence for a closer relationship of Wendlandia to subfamily Ixoroideae and the transfer of Wendlandia out of Rondeletieae. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 128–141.     

Pollen morphology of some selected plant species from Horton Plains,Sri Lanka

The pollen morphology of four species: Gaultheria leschenaultii (Ericaceae), Psychotria zeylanica (Rubiaceae), Rubus leucocarpus (Rosaceae) and Symplocos bractealis (Symplocaceae) from Horton Plains, Sri Lanka was investigated using LM and SEM. Psychotria zeylanica is endemic to Sri Lanka and the other three taxa are common in Horton Plains. Gaultheria leschenaultii has tetrads with 3-colporate, coarsely granular to granulo-rugulate pollen grains; in the polar region and at the colpus margin the exine is smooth with micro-rugulate structure. Psychotria zeylanica has 3- (or 4-) colporate, reticulate-spinulose to granular pollen grains. Rubus leucocarpus has 3-colporate, striate-perforate pollen grains and Symplocos bractealis has 3- (or 4-) colporate pollen grains with a verrucate-perforate exine. Pollen grains of these four species are very indicative of human impact and palaeoclimate.     

Studies on the Pollen Morphology of the Genus Brassica (Cruciferae) in China

The present paper reports the pollen morphology of 36 species and varieties of Brassica L. in China mostly for the first time. They were examined both under LM and SEM. Pollen grains of Brassica L. in China are long-spheroidal, subspheroidal, spheriodal, oblate, rarely reniformis or subsquare. According to the aperture characters pollen grains may be divided into four types: 2,3,4-colpate, 3-colpate, 3,4-colpate, 3,4-colporate. The exine sculpture may also be divided into three types: 1, reticulate, 2, cerebroid-reticulate, 3, foveolatereticulate. Detailed characters of pollen grains for each taxon are presented in Table 1. The 2,3,4-colpate type of pollen grains may be considered as primitive one and the 3,4-colpate or 3,4-colporate type of pollen grains is the most advanced in Brassica. The tricolpate aperture and reticulate exine have been reported by the previous authors, whereas, 3,4-colporate and cerebroid-reticulate or foveolate-reticulate pollen grains are newly reported.     

Pollen Morphology of Parnassia L. （Parnassiaceae） and Its Systematic Implications

The pollen morphology of 28 species of Parnassia L. was investigated with light microscopy and scanning electron microscopy (SEM). The shape of pollen grains in this genus varies from subspheroidal to prolate in equatorial view and is three-lobed circular in the polar view. Pollen grains are usually radially symmetrical, isopolar, tricolporate or syntricolporate, with reticulate sculpture. The pollen characteristics among species are fairly similar to each other. Morphological information regarding the pollen grains shows that Parnassia is a natural genus. Based on exine ornamentation observed under SEM, three types of pollen grains were recognized: (i) type I, with foveolate-reticulate sculpture; (ii) type II, with a finely reticulate sculpture; and (iii) type III, with a coarsely reticulate sculpture. Most sections of this genus have one type of sculpture of pollen morphology, but Sect. Nectarotrilobos has three types of sculpture and Sect.Saxifragastrum has two types of sculpture. All three types of sculpture can be found in Southwest China,with species with the longest (Parnassia delavayi Franch.) and shortest (Parnassiafaberi Oliv.) colpi,implying that Southwest China is the center of diversification of the genus.     

Pollen Morphology of Plumbaginaceae in China

The present paper describes the pollen morphology of 30 species belonging to 7 genera of Plumbaginaceae from China. The pollen grains were all examined under light microscope, and those of some species under scanning and transmission electron microscope. The pollen grains of the family are subspheroidal, prolate or oblate, (37.5-74.5)× (40.4-81.9)μ in size, 3-colpate, rarely 4-6-colpate, only pancolpate in Ceratostigma willmottianum. The exine 2-layered, 2.0-7.4μ thick, sexine thicker than nexine, verrucate, reticulate or coarsely reticulate. On the basis of the morphology, two types of pollen grains are distinguished in the family: (1) The pollen grains are 3-(rarely 4-6 ) or pancolpate, the exine verrucate. They are found in the tribe Plumbagineae (inculuding the genera Ceratostigma, Plumbago and Plumbagella). (2) The pollen grains are all 3-colpate, the exine reticulate or coarsely reticulate. They are found in the tribe Staticeae (including the genera Acantholimon, Ikonnikovia, Goniolimon and Limonium ).     

中国东北牻牛儿苗科（Geraniaceae）花粉形态与系统分类学研究

本文对中国东北牛儿苗科Ｇｅｒａｎｉａｃｅａｅ老鹳草属Ｇｅｒａｎｉｕｍ和牛儿苗属Ｅｒｎｄｉｕｍ植物计２属,１４种,２变种及１变型的花粉形态进行了较系统的研究。并对本科花粉形态进行了分类处理,编制了花粉形态科种检索表。结果表明,本科花粉如按萌发孔类型、外壁雕纹式样可以分为二大类：即三孔沟,外壁雕纹瘤状（Ｇｅｒａｎｉｕｍ）三沟,外壁雕纹网状（Ｅｒｏｄｉｕｍ）。在老鹳草属种及种下等级的分类处理中,花粉粒大小、萌发沟长度、外壁覆盖层网眼均匀性、直径大小、网脊上纹饰及外壁雕纹瘤纹突起均匀性、大小及瘤上纹饰都具有较大的分类学意义。花粉形态分类与经典分类对本科分类群的处理意见相吻合。大大丰富了本科分类微形态的证据。     

Exine Development in Illicium Religiosum Sieb. et Zucc. (Illiciaceae)

The exine development in Illicium was investigated using transmission electron and field emission scanning electron microscopy. The protectum and procolumellae appear on protruding sites of the microspore cytoplasm in the early tetrad stage. The protectum takes the form of a reticulate pattern with perforations within the callosic wall. After dissolution of the callosic wall, the central part of muri rises to form tectal ridges. The developing tectum, shows an echinate appearance in sectional view and has perforations at both sides around each lumen. There are two kinds of columellae; those forming continuous rings around each lumen and others which are individual rods standing beneath the tectum. The present developmental study in Illicium showed that the initial simple reticulate pattern formed within the callosic wall develops into the complex reticulate exine pattern of the differentiating tectum during the free microspore stage. The tectum has an angular shape with perforations and is supported by the two kinds of columellae.     

A Study on Pollen Morphology of Tribe Rhamneae (Rhamnaceae)in China

Pollen morphology of 25 Chinese species, representing 6 five genera in the tribe Rhamneae of Rhamnaceae, was examined under LM and SEM. Pollen grains are subspheroidal or suboblate, obtuse-triangular in polar view, 3-colporate, with the polar axis (P)10.4-28.7μm long and equatorial axis (E) 11.3-30.5μm long, P/E = 0.81-1.11. Colpi generally narrow and long, ora lalongate, with two ends connected with the thinned part of exine, forming a H-shape. Four thickenings were observed where colpi and ora cross, but sometimes the thickenings were indistinct or formed a ring. The stratification indistinct. The ornamentation of exine obscure or indistinctly reticulate under LM, but rugulose, short-striate, rugulose-foveolate or reticulate under SEM. The exine ultrastructure of Rhamnus ussuriensis was examined under TEM. It consists of imperforate tectum, granular layer, foot layer and endexine. The tectum is rather thick with uneven surface, while the granular layer is rather thin. A key to the genera based on pollen characteristics is provided and the general pollenmorphology of five genera, i.e. Sageretia, Rhamnus, Hovenia, Colubrina and Alphitonia, is described respectively. Based on pollen size, the four thickenings and the ornamentation, the five genera under study may be distinguished from each other. For example, pollen grains are smaller in Sageretia; the four thickenings are larger and distinct in Hovenia and Alphitonia. However, the ornamentation is short-striate under SEM in Hovenia and Alphitonia, while reticulate in the Colubrina and Rhamnus ( Subgen. Rhamnus) .Two different concepts of Rhamnus exist among taxonomists. Heppeler and Suessenguth divided Rhamnus into two subgenera (Subgen. Frangula and Subgen. Rhamnus), whereas Grubov separated Subgen. Frangula from Rhamnus as an independent genus. According to the pollen morphology, the separationis reasonable.     

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。