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.     

国产蔷薇科绣线菊亚科的花粉形态

本文报道了国产蔷薇科绣线菊亚科(Spiraeoideae)8属9种植物花粉形态。该亚科花粉近球形,极面观三裂圆形。花粉通常较小,最小花粉见于Spiraeapurpurea,为1487(1275～1658)μm×1517(1275～1734)μm,其中Physocarpusamurensis的花粉比较大,为2797(1989～3825)μm×3091(2677～3366)μm。三孔沟,内孔为长方形,部分种内孔明显外突。沟较长,两端较窄,在Physocarpusamurensis观察到合沟现象。合沟出现时通常三沟在一极汇合,在另一极常不汇合。花粉外壁具穴状、条纹-穴状、条纹-网状、条纹状等数种纹饰。迄今为止,在Rosaceae观察到的所有花粉外壁纹饰中,除较为特化的刺状纹外,其余纹饰在该亚科均存在。在Rosaceae所有的花粉外壁纹饰中,穴状纹饰可能是较为原始的类型,条纹状-网状和条纹状依次是较为进化的,而刺状是最进化的纹饰。在Spiraea中,外壁纹饰有相对进化的条纹-网状和条纹状,然而未观察到穴状这一最原始类型,这表明该属及其属下种的分化程度可能较高。但因Spiraea所含种数较大,这一推论有待进一步深入论证。此外,花粉形态特征支持Takhtajan(1997)关于Spiraeoideae亚科下族与属的排序。     

国产蔷薇科蔷薇亚科的花粉形态

对国产蔷薇科（Ｒｏｓａｃｅａｅ） 蔷薇亚科（Ｒｏｓｏｉｄｅａｅ） １０ 属１２ 种植物的花粉进行了光镜和扫描电镜观察。本亚科花粉呈单粒存在, 花粉近球形至长球形, 极面观常呈三裂圆形, 赤道面观椭圆形至圆形。花粉大小为（１６－０６μｍ ～３１－８８μｍ） ×（１５－３０μｍ ～３５－７０μｍ） 。三孔沟, 内孔为长方形, 沟较长, 两端较窄。其中, 矮地榆Ｓａｎｇｕｉｓｏｒｂａ ｆｉｌｉｆｏｒｍｉｓ 的花粉形态较为特殊, 每条沟的中部特别宽, 向两端则逐渐变窄, 形成近乎菱形图案, 并且在沟的中央覆盖着近椭圆形的沟膜, 将沟部分隔开来。从而在光镜下, 其极面观为不等的六裂圆形。但该种花粉依然属于３ 孔沟类型。花粉外壁在扫描电镜下见有穴状、皱波或疣状－ 穴状、条纹－ 穴状、条纹－ 网状、条纹状、刺状、条纹－ 刺状等纹饰变化。与外部形态特征和染色体基数复杂化相适应, 本亚科的外壁纹饰与其余亚科相比变化也最为多样。本文在对Ｒｏｓｏｉｄｅａｅ 的部分国产属种进行花粉形态报道的同时, 也对一些有关系统学问题进行了讨论。     

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１２μｍ,赤道轴１７８５～３６９５μｍ。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Ｓｉｎｏｐｌａｇｉｏｓｐｅｒｍｕｍｕｎｉｆｌｏｒａ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。孢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。Ｐｒｉｎｓｅｐｉａｕｔｉｌｉｓ和Ｓｉｎｏｐｌａｇｉｏｓｐｅｒｍｕｍｕｎｉｆｌｏｒａ（Ｐｒｉｎｓｅｐｉａｕｎｉｆｌｏｒａ）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Ｓｉｎｏｐｌａｇｉｏｓｐｅｒｍｕｍ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Ｅｘｏｃｈｏｒｄａ移置至Ｐｒｕｎｏｉｄｅａｅ之下     

国产蔷薇科苹果亚科的花粉形态

对国产蔷薇科（Ｒｏｓａｃｅａｅ）苹果亚科（Ｍａｌｏｉｄｅａｅ）１６属１８种及１变型进行了植物花粉形态的光学显微镜和扫描电镜观察。本亚科花粉呈单粒存在,花粉近球形至长球形,极面观常呈三裂圆形,赤道面观椭圆形至圆形。本亚科的花粉在全科中是最大的,极轴为１８．６２～４５．６５μｍ,赤道轴为１８．３６～４０．８μｍ。最大花粉见于Ｃｒａｔａｅｇｕｓ ｐｉｎｎａｔｉｆｉｄａ,为（２８．８５～４５．６５）μｍ     

Pollen Morphology of Hydrocotyloideae and Saniculoideae (Apiaceae) in China

Pollen grains of 30 species and 2 varieties from China, belonging to 5 genera (out of 36 species in 5 genera) of Hydrocotyloideae and Saniculoideae, were examined with light microscope and scanning electron microscope. 1.Six types of the pollen shape are recognized in Hydrocotyloideae and Saniculoideae from China, and their evolutionary trend is from rhomboidal→subspheroidal→ellipsoidal→subrectangular→superrectangular to equatorially-constricted. 2.Pollen grains of Hydrocotyloideae are mostly ellipsoidal with P / E ratio 1.07-1.6, the polar axis 22.5-46μm long, the equatorial axis 13.75-27.5μm long, and with the size index 21.1-31.8.They are angulapeturate in the great majority, and the exine sculpture is recticulate under SEM. Those of Saniculoideae are mostly superrectangular or rectangular, with P/E ratio 1.3-2.1, the polar axis 35-65μm long, the equatorial axis 17.5-42.5μm long, and with the size index 28.2-49.7. They are planaperturate in the great majority and the exine sculpture is striate-reticulate or cerebro-reticulate under SEM. Therefore, the latter might be more advanced than the former. 3.Pollen grains of Dickinsia Franch., endemic to China, are ellipsoidal,with P/E ratio 1.5,the polar axis 31.25-37.5μm long, the equatorial axis 20-25μm, and with the size index 27.81. They are angulaperturate, and the exine sculpture is cerebroid-reticulate under SEM. The genus is greatly similar to the members of Hydrocotyloideae in pollen morphology, which supports the treatment of the genus asa member of Hydrocotyloideae by Handel-Mazzetti (1933).     

羊蹄甲属中国特有种的花粉形态学

对羊蹄甲属植物11种中国特有种的花粉形态进行了研究。结果表明,花粉萌发孔类型均为三孔沟,少数还有合沟;花粉均具半覆盖层,表面纹饰有皱波状、皱波状-穿孔、疣状、孔穴-穿孔、穿孔-网状。依据Larsen(1975)的划分,这11种植物的花粉隶属于Integrifolia型、Glauca型和Curtisii型,我们根据花粉纹饰的差异在Integrifolia型下进一步划分了Auma亚型和Paucinervata亚型。     

国产芋属花粉形态

通过光镜和扫描电子显微镜观察了芋属(Colocasia Schott)6种、1变种、1品种的花粉形态,其中贡山芋C．gaoligongensis,龚氏芋C．gongii,李氏香芋C．lihengiae,花叶芋C．bic-olor4个种为首次报道。本属花粉为圆球形,无萌发孔。外壁纹饰分为两类：大野芋C．giga-ntea外壁光滑无刺,其余各种均具刺。花粉大小为20．07—32．76μm。     

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 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 ).     

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 of Dioscorea in China

This paper deals with pollen morphology of 33 species belonging to 5 sections of Dioscorea in China. They were examined under both LM and SEM. The pollen grains of Sect. Stenophora are monocolpate with the size of 17-25.5×26.8-39.1μm, the exine is reticulate, striate, parforate-reticulate or cerebro-reticulate, while those of the other sections are bicolpate, reticulate or cerebro- reticulate with the size of 19.2-26.7×13.9-23.1μm. Based on palynological data Sect. Stenophora may be considered primitive and Sect. Enantiophyllum the most advanced in Dioscorea.     

Pollen Morphology of Embothrieae (Proteaceae)

This is the first of two papers detailing pollen morphology and evolution with the tribe Embothrieae comprising eight genera and ca. 56 spp. The present paper examines pollen of subtribes Buckinghaminae (Buckinghamia; 2 spp., Opislhiolepis; 1 sp.), Stenocarpinae (Strangea; 3 spp.), Stenocarpus (ca. 27 spp.) and Lomatiinae (Lomatia; ca. 12 spp.) in the light microscope and scanning and transmission electron microscopes. Pollen is medium-sized, oblate, foveolate to microreticulate to reticulate, and predominantly columellate with a complex modified postvestibulate aperture morphology. Pollen data indicate ties between Lomatia and Stenocarpus on the one hand and Stenocarpus and Strangea on the other. Though Buckinghamia and Opislhiolepis have been placed in the same subtribe, the unique combination of pollen features in each suggests only a remote relationship to each other as well as to remaining Embothrieae. Comparisons to the remaining genera of Embothriae (Embothrium, Oreocallis, Telopea) and overall analysis of pollen evolution within the tribe are detailed in the subsequent paper.     

中国紫葳科花粉形态的研究

对紫葳科18属27种花粉进行了光镜和扫描电镜的观察,其中有些种是第一次观察或报道。紫葳科花粉明显地属多型花粉,萌发孔类型和纹饰特征多种多样。萌发孔基本上可分为3（－4）沟,3（－4）孔沟,3孔,多沟及散沟等类型;纹饰则可分为皱波状饰,穴状纹饰,穴－网状纹饰,网状纹饰,刺状纹饰以及表面光滑等。值得注意的是在菜豆树属Rademoachera Zoll.et Moritzi中,菜豆树R．sinica和滇菜豆树R.yunnanensis虽为同一个属,但其萌发孔类型和纹饰特征截然不同,菜豆树花粉为3孔,具网状纹饰;而滇菜豆树花粉为3孔沟,具穴状纹饰。就花粉形态学而言,紫葳科也许是多元发生的。     

中国木犀属花粉形态研究及其系统学意义

观察了中国木犀属4组21种植物的花粉形态。花粉为圆球形或近长球形,外壁表面均具网状纹饰。大多数种类都具有三孔沟的萌发孔,极少四孔沟,或六萌发孔。内孔明显或不明显,有9种植物同时存在具三孔沟和具三沟二种类型的花粉及中间过渡类型(显脉木犀同时存在具三拟孔沟和具四拟孔沟)。很多特点表明,木犀属花粉的萌发孔在木犀科中处于由简单萌发孔向复合萌发孔过渡的阶段。根据网眼从沟间区中央到沟边是否变细,可将本属花粉分为两大类型,与分组有一定的对应关系。综合一些花粉特征,可发现本属花粉形态差异较明显,与一般形态差异有一定的相关性。花粉形态在木犀属具有一定的系统学意义。     

大戟科叶下珠属花粉形态的研究

对叶下珠属Phyllanthus Linn.13种植物花粉进行了光学显微镜和扫描电镜的观察。研究表明,本属花粉虽然在大小、形状和外壁纹饰等方面有不同程度的差异,但差异最大的还是在于萌发孔类型上。可以把本属花粉分为3孔沟,4－5孔沟,多孔沟,散沟（沟短而纺锤形或近圆形）和散孔等类型。这些萌发孔类型虽然与李秉滔在中国植物志第44卷中所划分的组不很吻合,但或多或少有一定的相关性,并且对于属下的分类具有一定的意义。在外壁纹饰方面,均为网状纹饰,可分为具粗网眼,细网眼和中等网眼等几种类型。但纹饰类型在组间或种间的差异不大明显。     

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.     

Pollen Morphology in Tribe Lactuceae (Compositae)

The Lactuceae contain two basic pollen types, echinolophate and echinate. Most taxa have echinolophate, tricolporate pollen. Internally, most ektexines are composed of a perforate spiny tectum, several levels of columellae, a cavus, and a foot layer. In lacunae, the columellae are reduced to a single level and the cavus is often absent. Highly modified echinolophate pollen grains are found in Scolymus, Scorzonera and Tragopogon. Scolymus, Catananche, Scorzonera, and Tolpis have distinctive exine stratification patterns. Exines of Catananche and, to a lesser extent, those of Tragopogon contain internal foramina like those found in the Heliantheae. Echinate pollen is found in all subtribes and is probably ancestral. However, some echinate grains are probably derived.     

Chromosome Numbers of Brassiceae (Cruciferae) in China

Reported in this paper are chromosome numbers of 41 species and varieties of Brassiceae in China, belonging to 5 genera. The chromosome numbers of 15 Species and varieties are first reported and some problems concerning chromosome numbers of Brassiceae are also discussed.     

中国鸡矢藤属花粉形态

用扫描电子显微镜对中国鸡矢藤属(Paederia L.)6种1变种植物的花粉进行观察。结果表明:鸡矢藤属植物的花粉均为单粒,辐射对称,小型或中型,极面观3-裂圆形至钝三角形,赤道面观呈长圆球形或近长球形,具3个萌发沟,无内萌发孔。外壁纹饰网状、细网状或穿孔状,孔边缘具小刺状突起或无。臭鸡矢藤(P.foetida)和白毛鸡矢藤(P.pertomentosa)具有花粉二型现象,其中白毛鸡矢藤(P.pertomentosa)是首次报道。花粉二型现象与花柱二型现象可能没有直接的关联性,与前人的观点一致。推测鸡矢藤属外壁纹饰的可能演化趋势为:穿孔、网状、细网状→粗网状;网眼内无棒状突起→网眼内有棒状突起。鸡矢藤属花粉的外壁纹饰变化较大,且无内萌发孔,是茜草科花粉过渡类型的特征。