Study on the Pollen Morphology of the Family Berberidaceae

The pollen grains of 33 species representing 11 genera of the family Berberidaceae, mostly from China, were examined with the light microscope and scanning electron microscope. Their characteritic details can be used for generic diagnosis. A pollen key to the genera based on these observations is presented. Based on the morphology, the pollen grains can be grouped into the following three types: 1. The tetrad pollen type found only in the genus Sinopodophyllum. 2. The spiraperturate pollen type found in the genera Berberis and Mahonia. 3. The tricolpate pollen type found in the genera Diphylleia, Jeffersonia, Nandina, Dysosma, Caulophyllum, Leontice and Epimedium. A diagnostic key to the pollen grains of genera in the Berberidaceae. 1. Pollen grains single 2. Pollen grains spiraperturate .................... Berberis L., Mahonia L. 2. Pollen grains tricolpate 3. Exine with spinose sculpture ........................ Diphylleia Michx. 3. Exine with non-spinose sculpture 4. Exine with striate or striate-reticulate sculpture ...... Jeffersonia Barton 4. Exine with reticulate sculpture 5. Exine around colpus with distinct thickening ........ Nandina Thunb. 5. Exine around colpus without thickening 6. Pollen grains larger (45—50)×(32.5—37.5)μ 7. Colpus with membrane ................ Dysosma R. E. Woodson 7. Colpus without membrane ................ Caulophyllum Michx. 6. Pollen grains smaller (25—550)×(20—527.5) μ 7. Pollen grains prolate-perprolate .................... Leontice L. 7. Pollen grains spheroidal-prolate...Epimedium L., Podophyllum L. 1. Pollen grains tetrad ............................ Sinopodophyllum Ying     

蜘蛛抱蛋亚族的花粉形态

本文研究并报道了百合科蜘蛛抱蛋亚族的花粉形态,主要结论如下：1．开口箭属含两种花粉类型即：(1)单槽、椭圆形;(2)无萌发孔、球形。具单槽、椭圆形花粉的种类含单型性核型,花形较小,柱头小而花柱短,雄蕊高于柱头;无萌发孔、球形花粉的种类则为单型性核型,花形较大,柱头常高度膨大,雄蕊常着生在花被筒基部。2．万年青属花粉为椭圆形,具单槽;蜘蛛抱蛋属花粉则为球形,无萌发孔;开口箭属花粉一部分与万年青属相似,另一部分则与蜘蛛抱蛋属雷同。3．蜘蛛抱蛋亚族中最原始的花粉类型为单槽椭圆形,具孔状或网状外壁纹饰。球形,无萌发孔,外壁具皱波状、瘤状或芽孢状纹饰的花粉为派生类型。4．蜘蛛抱蛋亚族种下花粉形状、萌发孔类型和外壁纹饰基本稳定,可作为分种特征。     

Pollen Morphology in the Subtribe Aspidistrinae (Liliaceae s. l.)

Pollen morphology in the subtribe Aspidistrinae is reported and the main viewpoints are summarized as follows: (1) Two major types of pollen grains, i.e., monosulcate,ellipsoidal pollen and inaperturate, spheroidal pollen, are identified in the genus Tupistra. The species with monosulcate, ellipsoidal pollen usually have monomodal karyotypes, brightcolored flower, smaller stigma, and stamens inserted at upper or middle part of perianth tube, while the inaperturate and spheroidal pollen is always found in the species with bimodal karyotypes, dingy-colored flower, swollen stigma and stamens inserted at the base of perianth tube. (2) Pollen grains in the genus Rhodea are monosulcate and ellipsoidal, while those in the genus Aspidistra are inaperturate and spheroidal, correspondingly similar to the second type of pollen grains in the genus Tupistra. (3) The most primitive pollen in the subtribe Aspidistinae is regarded as monosulcate and ellipsoidal, having perforate or reticulate exine sculpture. The inaperturate, spheroidal pollen with verrucate, gemmate or rugulate exine sculpture is considered derived; (4) Unlike those reported in other groups of the family Liliaceae, the infraspecific pollen shape, aperture type and exine sculpture in Aspidistrinae are basically stable and may serve as a taxonomic character.     

A New species of the Genus Tupistra (Convallariaceae), with Reference to Its Karyotype and Pollen Morphology

A species of the genus Tupistra,T．pingbianensis J．L．Huang et X．Z．Liu,is described as new,and its karyotype and pollen morphology are reported． The bimodal karyotype of this species is formulated as 2n＝38＝14m+10sm+8st+6t,and pollen grains are spheroidal,inaperturate,with rugulate exine sculpture．Based on its floral structure,karyotype and pollen morphology,T．pingbianensis J．L．Huang et X．Z．Liu is considered phylogenetically closely related to T．fungilliformis Wang et Liang．Whether the repense rhizome,solitary leaves of T．pingbianensis J．L．Huang et X．Z．Liu indicate a close relationship between the genus Tupistra and the genus Aspidistra needs further studies．     

开口箭属一新种及其核型和花粉形态

文章报道了新种屏边开口箭T．pingbianensis J．L．Huang et X．Z．Liu及其核型和花粉形态。屏边开口箭的核型公式为2n=38=14m+lOsm+8st+6t,具明显的二型性,属3C型。花粉粒为球形,无萌发孔沟,外壁具皱块状纹饰。从花结构、核型和花粉形态上看,屏边开口箭和伞柱开口箭T．fungilliformis Wang et Liang亲缘关系近,而且伞柱开口箭比屏边开口箭在一系列特征上都较特化。     

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.     

槭树科花粉形态及其系统学意义

报道了槭树科(Aceraceae)槭属21组33种和金钱槭属2种植物的花粉形态。本科花粉近球形至长球形,极面观为三裂圆形。从花粉萌发孔类型看,金钱槭属Dipteronia具三孔沟,槭属（Acer)除4组具三孔沟外,其余均为三沟。从花粉外壁纹饰看,金钱槭属2个种和槭属的大多数种为条纹状,罕为细条纹－拟网状和皱波状。通过花粉形态分析,并结合其它方向的证据,我们认为：（1）Sect.Palmata,Sect.Spicata和Sect.Microcarpa可能是槭属中与金钱槭属关系最密切的类群;（2）A.distalum和A.nipponicum代表了近缘的两个单种组;（3）A.pseudoplatanus不同于Sect.Acer的其它4个种而与A.saccharum可能存在更为密切的关系;（4）Sect.Carpinifolia(细条纹－拟网状外壁纹饰）和Sect.Negundo(皱波状外壁纹饰）则可能代表了槭属中最特化的类群;（5）按槭树科花粉三孔沟到三沟的演化规律,Dipteronia较Acer原始。     

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     

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 (Parnassia faberi Oliv.) colpi,implying that Southwest China is the center of diversification of the genus.     

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.     

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 of Astragalus section Hymenostegis (Fabaceae) and evaluation of its systematic implications

Astragalus is with nearly 3000 described species the largest genus of flowering plants. So far analyses of pollen characters have only been conducted for a few species of the groups within the genus. Here we analyse pollen grains of 22 species representative for Astragalus section Hymenostegis using scanning electron microscopy. We found the basic shape of the pollen grains to be oblate-spheroidal and apertures to be tricolpate as for other eudicots. The sculpturing pattern of the exine is micro-reticulate. Pollen grains show low morphological variation among different species of this section, but differences occur between sections of the genus. We conclude that the vast morphological differentiation that occurred during the rapid radiation of section Hymenostegis was not accompanied by comparable differentiation in pollen morphology.     

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.     

中国北部绣线菊属的花粉形态及分类学意义

本文用光学显微镜和扫描电镜观察了中国北部绣线菊属ＳｐｉｒａｅａＬ．２１种及６个变种的花粉形态,描述了外壁纹饰的形态特征,指出了外壁纹饰特征在分类上的意义。绣线菊属的花粉为球形至长球形,外壁纹饰为条网状至条纹状。外壁纹饰类型不具有属内分组意义,但对一些种类具有一定的分类价值。花粉形态支持将曲枣绣线菊Ｓ．ｆｌｅｘｕｏｓａＦｉｓｃｈ。并入石蚕叶绣线菊Ｓ．ｃｈａｍｏｅｄｒｙｆｏｌｉａＬ．的观点［Ｍａｘｉｍｏｗｉｃｚ,Ｃ．Ｊ．（１８７９）、Ｒｅｈｄｅｒ,Ａｉ．（１９４０）］,不同意将蒙古绣线菊毛枝变种Ｓ．ｍｏｎｇｏｌｉｃａＭａｘｉｍ．ｖａｒ．ｔｏｍｅｎｔｕｌｏｓａＹｕ提升为贺兰山绣线菊Ｓ．ｔｏｍｅｎｔｕｌｏｓａ（Ｙｕ）Ｙ．Ｚ．Ｚｈａｏ［赵一之（１９８７）］。     

Phylogeny of holoparasitic Orobanche (Orobanchaceae) inferred from nuclear ITS sequences

Orobanche is the largest genus among the holoparasitic members of Orobanchaceae. We present the first molecular phylogenetic analysis (using nuclear ITS sequences) that includes members of all sections of Orobanche, Gymnocaulis, Myzorrhiza, Trionychon, and Orobanche. Orobanche is not monophyletic, but falls into two lineages: (1) the Orobanche group comprises Orobanche sect. Orobanche and the small Near Asian genus Diphelypaea and is characterized by a chromosome base number of x=19 and (2) the Phelipanche group contains Orobanche sects. Gymnocaulis, Myzorrhiza, and Trionychon and possesses a chromosome base number of x=12. The relationships between these two groups and to other genera such as Boschniakia or Cistanche remain unresolved. Within the Orobanche group, Orobanche macrolepis and Orobanche anatolica (including Orobanche colorata) constitute two phylogenetically distinct lineages. Intrasectional structurings proposed by some authors for O. sect. Orobanche are not confirmed by the molecular data. In most cases, intraspecific sequence divergence between accessions, if present, is negligible and not correlated with morphological or ecological traits. In a few cases, however, there is evidence for the presence of cryptic taxa.     

延龄草科四属花粉形态的研究

对延龄草科重楼属Paris L.延龄草属Trillium L.Medeola L.属和Scoliopus Torr.属以及百合科百合属Ｌilium L.和萱草属Ｈemerocallis L.花粉进行了光镜和电镜的观察和比较。结果表明,在延龄草科中,重楼属,Medeola属和Scoliopus属花粉具有较多的相似性：具单一远极沟,极面观椭圆形,外壁为网状纹饰或皱波－网状纹饰。而延龄草属花粉与它们的差异则较大,为圆形,无萌发孔,外壁非网状纹饰,而是颗粒状,疣状,芽孢状,星状和微刺状等纹饰。百合属中的Lilium michauxii和萱草属中的金针菜Hemerocallis citrina花粉均为单一远极沟,外壁表面为网状纹饰。但其网脊特别粗,具不规则突起并形成棱角（H.citina)或由许多排列成念珠状的、形状不一的分子组成（Lilium michauxii),结构特别复杂。就其萌发孔类型和纹饰特征看,它们与延龄草科中的重楼属,Medeola属和Scoliopus属花粉具有较多的性。花粉外壁的层次结构在延龄草科各属之间（延龄草属除外）,甚至在延龄草科与百合科之间均没有多大的差异。     

12种十字花科植物花粉形态研究

采用扫描电子显微镜对十字花科8属12种植物的花粉形态特征进行了观察分析.其中8种为首次报道.观察结果表明:4族8属12种十字花科植物花粉形态的一般特征为:花粉粒长球形至超长球形;极面观三裂圆形,少数四裂圆形;外壁纹饰为网状或细网状纹饰;三沟,偶四沟.12种植物的花粉可划分为5种类型:(1)花粉粒长球形,网状纹饰,三沟;(2)花粉粒超长球形,网状纹饰,三沟;(3)花粉粒长球形,细网状纹饰,三沟;(4)花粉粒超长球形,细网状纹饰,三沟;(5)花粉粒长球形,三或四沟,网状纹饰.十字花科花粉形态特征在不同族间、不同属间以及同属不同种间均没有明显规律,在十字花科分类过程中,不宜单独以花粉形态特征作为依据.     

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     

中国独行菜属9种植物花粉形态扫描电镜观察

对中国产独行菜属9种植物的花粉微形态作了扫描电镜下的观察和研究。结果表明,花粉粒为超长球形或长球形,具三沟,外壁纹饰均为网状纹饰。根据外壁纹饰可分为3种类型：（1）网眼为不规则形状,网眼边缘常有刺突;（2）网眼为圆形或近圆形,网眼边缘较光滑;（3）网眼为多边形或不规则多边形,网眼边缘较光滑,偶有刺突。