The Pollen Morphology in Relation to the Taxonomy and Phylogeny of Fagaceae

The present paper deals with the pollen morphology of 103 species belonging to sixgenera-Castanea, Castanopsis, Lithocarpus, Quercus, Fagus and Trigonobalanus in threesubfamilies-Castaneoideae, Quercoideae and Fagoideae. All pollen grains were examinedunder light microscope and scanning electron microscope, and those of some species were ex-amined under transmission electron microscope. The results may be summarised as follows:1. Pollen morphology of Fagaceae, seems to support division of the family into threesubfamilies. Fagoideae, Castaneoideae and Quercoideae.2. Four types of pollen grains are recognized in Fagaceae:1) Fagus-type (representative genus: Fagus): pollen granis are oblate-sphaeroidal,(31.5-39.9) x (35.7 46.2) μm in size, 3(-4)-colporate, peritreme or goniotreme, granulate-ornate under LM., granulate or verrucate under SEM.2) Trigonobalanus-type (T. doichangensis): pollen grains are suboblate-sphaeroidal,(23.1-29.4) ×(25.2-29.4) μm in size, 3-colporate, goniotreme, obscurely granulate-ornateunder LM, densely granulate or verrucate under SEM.3) Quercus-type (Quercus): pollen grains are subspheroidal-subprolate, (21-44.3)× (16.8-39.9) μm in size. 3-colporoidate (-3-colpate), peritreme, crassgranulate or finely-gra-nulate under LM, tuberculate verrucate or spinate under SEM.4) Castanea-type (including Castanea, Castanopsis, Lithocarpus): Pollen grains areprolate-supraprolate, (14.7-23.1)×(10.5-16.8)μm in size; 3-colporate, peritreme, obscu-rely ornate or subpsilate, under LM, rugulose, striate-rugulate or crass-striate under SEM.3. Pollen grains of Cyclobalanopsis age very similar to those of Quercus, and there-fore we support the treatment of Cyclobalanopsis as a subgenus of Quercus.4. On the basis of shape, type of aperture and exine structure, pollen of Trigonobalanusis distinguishable from those of the other genera in Fagaceae and it may be a new type of Fagaceae;5. On the basis of pollen morphology, morphological characters and geological stratification a scheme of phylogeny of Fagaceae is here presented.     

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。     

Pollen Morphology of Quercus L. in China

The present paper describes the pollen morphology of 31 species of the genus Quercus from China. The pollen grains were all examined with light microscope and scanning electron microscope, and those of some species under transmission electron microscope. Pollen grains of the genus are spheroidal or subspheroidal, 16.8-50.4μm in diameter, 3colporoidate or 3-colpate. The exine is 2-layered, 0.4-1.8μm thick, sexine thicker than nexine, granulated tuberculate or verrucate.     

东北杜鹃花属植物花粉形态的研究

利用光学显微镜和扫描电子显微镜对东北杜鹃花属(Rhododendron Linn.)6种, 1变种, 1变型的花粉形态进行了观察和比较研究。该属花粉粒均为四合花粉。四合花粉呈正四面体排列。花粉粒近球形。每粒花粉均具3沟,在四合花粉粒上,相邻花粉粒上的沟相接,有些种可见沟内的萌发孔。光镜下可见花粉外壁为内外两层,厚度约相等。不同的种在花粉粒大小上有区别。在扫描电镜下观察可见不同的种花粉表面具有不同的纹饰。即该属在花粉形态上具有种的特性。     

东北杜鹃花科4属植物花粉形态的研究

利用光学显微镜和扫描电子显微镜对东北杜鹃花科甸杜属(Chamaedaphne Moench.)、杜香属(Ledum Linn.)、松毛翠属(Phyllodoce Salisb.)、越橘属(Vaccinium Linn.)4属6种, 1变种的花粉形态进行了观察和比较研究。以上4属花粉粒均为四合花粉。四合花粉呈正四面体排列。花粉粒近球形。每粒花粉均具3沟,在四合花粉粒上,相邻花粉粒上的沟相接,有些种可见沟内的萌发孔。光镜下可见花粉外壁为内外两层,厚度约相等。不同属种的花粉粒在大小上有区别,花粉粒平均直径为34.0~43.7μm。在扫描电镜下观察可见不同的种花粉表面具有不同的纹饰。为分类学提供了孢粉学方面的证据。     

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 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).     

山茶属的花粉形态及其分类学意义

本文对山茶属(Camellia L.)18个组27个代表种的花粉进行了光学显微镜、扫描电镜和大部分进行了透射电镜的观察,并对花粉大小、形状、外壁纹饰和超微结构等作了综合的比较。结果表明,其形状和大小差异不很明显。花粉壁的层次、厚度和不同层次的比例,如复盖层与柱状层以及外壁外层与外壁内层的比例等也没有显著的差异。但花粉壁的纹饰特征在扫描电镜下显示出较大的差异,具有较重要的分类学价值。根据其纹饰特征,可以把山茶属花粉大致区分为六个类型:皱疣状纹饰;粗瘤状纹饰;颗粒状至皱-颗粒状纹饰;穴-网状纹饰;皱沟状纹饰及皱波-念珠状纹饰。孢粉学上区分的类型不完全与各个学者从植物形态上划分的组相一致。其中管蕊茶 Camellia lanceolata的纹饰非常特殊(粗瘤状),其外壁表面很象牛心果(Annona reticulata)的外形特征,为其重新分类提供孢粉学依据。     

我国睡莲科花粉形态的研究

本文对我国睡莲科5属11种2变种的花粉形态进行了系统的研究。每种花粉都通过光学显微镜和扫描电镜进行观察和照相。同时对莲和白睡莲二种花粉作了超薄切片,对它们的外型结构进行了透射电镜的观察和照相。并结合有关分类学和孢粉学资料,讨论了本科的原始性、异质性及演化上的问题。     

杉科植物花粉形态的研究

杉科(Taxodiaceac)是裸子植物的重要科之一。全科共有10属16种。本文用光学显微镜和扫描电镜观察了本科7属10种植物花粉,并用透射电镜研究了其中5个属代表种的外壁内部的超微结构。     

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.     

紫草亚科附地菜族的花粉形态研究

本文用光学显微镜和扫描电子显微镜系统观察了附地菜族5个属26种植物花粉形态。五属是附地菜属、车前紫草属、滨紫草属、山茄子属和皿果草属。观察证明属间花粉形态有明显的差异。花粉资料为族内属的划分和皿果草属的建立提出了花粉学证据。     

中国石蒜属植物花粉形态的研究

利用光学显微镜和扫描电镜对我国石蒜属ＬｙｃｏｒｉｓＨｅｒｂ．１１种植物花粉形态进行系统地研究,并对其中６种具有代表性植物花粉进行了透射电镜观察。该属植物花粉舟形或肾形;萌发孔为远极单槽;在光学显微镜和扫描电镜下观察,外壁具网状纹饰;在透射电镜下观察,外壁包括外壁－１和外壁－２两层,外壁－１是由半覆盖层、柱状层和基层组成,外壁－２很薄。根据花粉形态及其他器官特征,对本属植物分类地位也进行了一些讨论。     

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.     

旌节花科及其相关类群花粉形态的研究

对旌节花科及其相关类群五桠果科、猕猴桃科、水东哥科、金莲木科、省沽油科和山茶科等7科19属37种花粉进行了光镜（LM）和扫描电镜（SEM）的观察和比较。其中有些种类是第一次观察和报道。研究表明,上述各科花粉多为近球形至球形,少数近扁球形或近长球形。极面观三裂圆形或近三裂圆形。直径20－30（－35μm）,但厚皮香亚科和省沽油科银鹊树属Tapiscia的花科特别小,仅12.5-20μm。3孔沟,罕有3沟或4孔沟。外萌发孔（沟）相对地较长而宽,内孔多数较发达。花粉外壁纹饰变化大,从模糊而粗糙的细突起到皱波状纹饰,细孔状纹饰,蜂巢状－穴状纹饰,穴－网状纹饰,网状纹饰和颗粒状纹饰等。从形状、大小、萌发孔类型和外壁纹饰的比较看,虽然上述各科花粉均在不同程度上具有一定的相似性,但其中以旌节花科、省沽油科和山茶科中厚皮香科的花粉具有更多的相似性,尤其表现在外壁纹饰特征上。就孢粉学而言,也许旌节花科与省沽油科和山茶科中的厚皮香亚科的关系相对地较为密切。不过,其孢粉学特征所表现出的亲缘关系似乎不如在分子生物学中所表现出的明显。     

蓼科花粉外壁超微结构的研究

用扫描电镜和透射电镜对蓼科6属15种的花粉壁构造的特点进行了观察和研究。结果表明：(1)外壁纹饰有如下几种：颗粒-穿孔,微刺-穿孔,微刺-穿孔-光滑,颗粒-穿孔-光滑,细网状,粗网状,皱块状,鼓锤状;(2)外壁结构分化成两个明显的层次,即外壁外层以及外壁内层。其外壁外层由覆盖层、柱状层和基层组成。然而,由于每一部分发育状况的不同而导致外壁结构有各种变化,如无覆盖层或无外壁内层。     

须芒草族植物花粉形态的观察

在光学显微镜下和扫描电镜下对禾本科须芒草族（Ａｎｄｒｏｐｏｇｏｎｅａｅ）中分隶于８个亚族３４个属的３６种植物的花粉进行形态面容和比较研究。结果显示,本族植物花粉形态较为一致,花粉近形或扁球形,单萌发孔,孔,周围加厚,具盖,外壁表面散布有颗粒。这表明其是一个自然类群。总体来讲,芬烨大的演化分异,只是表面纹饰的和芬烨大小有一;定的差异。纹饰可分为三种类型粗糙型,不明显疠状突起才明显疠状突起型。花粉开矿     

泽泻科的花粉形态研究

本文对泽泻科11属27种代表植物的花粉进行了光学显微镜、扫描电镜和透射电镜观察。在系统描述了该科及各属植物花粉形态的基础上,将泽泻科植物的花粉划分为3种类型,即少果泽苔草型、慈菇型和泽泻型。根据花粉形态特征的比较,并依据泽泻科植物祖先类群的花粉具有船形、具单沟萌发孔、花粉外壁具明显的刺状纹饰、覆盖层完整无通道等特征,作者认为泽泻科植物花粉形态的如下演化趋势是明显的：由船形演化为卵球形、球形和多面体球形;由单沟萌发孔经过一无孔的中间类型演化为散孔类型;孔膜由光滑演化为具颗粒和小刺;萌发孔不内陷进化到内陷;花粉粒外壁的刺状纹饰逐渐过渡为颗粒状纹饰或者消失,以及覆盖层由无通道到具细通道和通道。     

国产爵床科山牵牛属6种、叉柱花属和老鼠簕属各1种植物的花粉形态

报道了国产爵床科Acanthaceae山牵牛属Thunbergia(山牵牛亚科Thunbergioideae)6种、叉柱花属Staurogyne(瘤子草亚科Nelsonioideae)和老鼠簕属Acanthus(老鼠簕亚科Acanthoideae)各1种植物在扫描电镜下的花粉形态。山牵牛属植物的花粉粒为圆球形,均具螺旋状萌发孔,外壁纹饰以光滑或颗粒状为主,偶具棒状突起。具螺旋状萌发孔被认为是该科独特而较原始的花粉特征。叉柱花属的花粉粒为圆球形,具3孔沟,外壁平滑。老鼠簕属的花粉粒为长球形,具3沟,外壁具细网状纹饰或具小穿孔。花粉形态特征支持传统上将上述3属置于3个不同亚科的处理。     

A Study on Pollen Morphology of Eucommia ulmoides Oliver

Eucommia ulmoides Oliver is endemic to China. The pollen morphology and exine ultrastructure were examined under LM, SEM and TEM. Pollen grains are prolate, polar axis 30.5-54.8 μm long, with the average 32.7 μm, equatorial axis 27.8-31.3 μm long, with the average 29.2 μm, tricolporoidate, but sometimes the outline of ora could be observed and elliptic in shape. Colpi are narrow, uequal in length, often two long and one short or two short and one long, sometimes rather irregularly arranged, with indistinct and thin colpus membrance. Exine psilate under LM, granulate under SEM, and shortly baculate under TEM. Tectum is thin with dense and small granules, columellae layer consists of short bacules, and foot layer very thick. Some taxonomists (Cronquist, 1968) consider that Eucommiaceae is related to Hamamelidales, but others (Takhtajan, 1969) to Urticales. The Urticales is of the porate type of pollen grains, while Eucommiaceae of tricolporoidate type, and thus the former is more advanced than the latter. Pollen grains of some members of Hamamelidales, tricolporoidate, are similar to those of Eucommiaceae. We therefore consider that Eucommiaceae is related to Hama-melidales.