茨藻科、角果藻科植物花粉形态的研究

在水生植物茨藻目中,茨藻科Najadaceae和角果藻科Zannichelliaceae植物由于其结构、尤其是花部结构的极度简化和趋同适应,系统关系仍不清楚。本文应用光学显微镜、扫描电镜和透射电镜对上述2科4属的10种代表植物的花粉形态进行了观察,系统描述了各属、种的花粉形态。结果表明角果藻科植物花粉均无萌发孔,外壁纹饰通常为拟网状至浅网状。花粉形态的研究结果支持将Vleisia作为角果藻属Zannichellia的近缘属处理的观点。茨藻科植物花粉外壁光滑、稍皱波状或为疣状,具颗粒状纹饰,萌发孔的有无在种间存在变异。花粉形态的研究结果表明角果藻科与眼子菜科Potamogetonaceae近缘,茨藻科可能与水蕹科Aponogetonaceae或水鳖科Hydrocharitaceae中具单沟花粉的类群有一定联系,而与角果藻科和眼子菜科则相去甚远。     

水鳖科9属15种植物花粉形态的研究

应用光学显微镜、扫描电镜和透射电镜对水鳖科Hydrocharitaceae 9属15种植物的花粉形态进行 了观察。水鳖科植物花粉为圆球形至近椭球形,无萌发孔或偶为单沟萌发孔,外壁纹饰通常为小刺状纹 饰,刺密集或稀疏,花粉表面具瘤状、疣状、颗粒状、皱波状突起或光滑。外壁由覆盖层、柱状层和基层组 成。覆盖层厚或较薄,柱状层小柱发育不明显,基层薄。水鳖科植物在花粉大小、纹饰类型、刺的长短、 密度、形态、萌发孔的有无以及花粉壁的结构等方面表现出了较为明显的差异,这些特征对探讨类群间 关系具有较重要意义。由于黑藻属Hydrilla和Stratiotes属花粉较为特殊,支持将它们各自作为一个独立 的族处理。水鳖科植物花粉外壁纹饰和结构特点表明该科与水雍科Aponogetonaceae、泽泻科Alismataceae 和花蔺科Butomaceae等近缘,而该科植物花粉大多无萌发孔等则反应了该科与茨藻目Najadales植物有密切联系。     

水雍科植物的花粉形态研究

应用光学显微镜、扫描电镜和透射电镜对世界水雍科6种植物的花粉形态进行了观察。水雍科植物花粉为舟形或船形,具远极单沟萌发孔类型,外壁纹饰通常为浅网状至网状,稀为小刺状纹饰,外壁外层由覆盖层、柱状层和基层组成,覆盖层厚。水雍科植物花粉外壁纹饰表现了从小刺状向网状的过渡。该科花粉为远极单沟,覆盖层厚,具小刺等特征反映了其与水鳖科和泽泻目花蔺科的密切联系,而该科花粉外壁纹饰多为网状则与茨藻目植物接近。这一结果支持将水雍科从茨藻目中分出作为一个独立的目处理的观点。此外,水雍科植物的花粉大小、纹饰类型、网眼大小与深浅等方面的差异对种级分类有一定意义。     

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。     

中国茨藻科植物种皮微形态特征及其系统学意义

本文应用扫描电镜对国产茨藻科植物的外种皮和内种皮微形态特征进行了研究,结果表明茨藻科植物在种子表面细胞形态、表面纹饰和内种皮内层细胞形态及小瘤状突起的大小和密度等方面存在差异,可作为种级水平分类的依据,并首次发现外种皮微形态特征也与内种皮内层小瘤状突起一样具有重要的系统学意义。根据种皮微形态特征的研究结果,我们支持Miki(1937)和Shaffer-Fehre(1991b)等关于茨藻科与水鳖科近缘,而不应放在眼子菜目的观点。对弯果茨藻Najas ancistrocarpa A．Br．ex Magnus和草茨藻N.graminea Del.外种皮的研究结果表明其外种皮细胞表面具瘤状颗粒而非网状加厚。     

中国紫草科破布木属花粉形态和外壁超微结构

为了深入探讨紫草科（Boraginaceae）的分类问题,用光学显微镜和扫描电子显微镜观察了该科破布木属（CordiaL.) 10种植物的花粉形态和外壁超微结构。发现该属花粉具三孔、三孔沟、三拟孔沟和三合沟4种萌发孔类型。外壁表面具微刺状纹饰、刺状纹饰、网状纹饰和不规则的条纹网状纹状。破布木属的花粉特征表明,该属花粉在紫草科中既是独特的分类群,又是比较原始的属种。     

中国无患子科的花粉形态及其系统学意义

利用扫描电镜对国产无患子科(狭义,Sapindaceae)23属30种1变种植物的花粉形态进行了观察。结果显示,该科花粉粒多为扁球形,部分为长球形,少数为球形或近球形,极面观多为三角形;从花粉萌发孔类型看,大多数种类具三沟孔,有的形成合沟,少数仅具三孔而无沟;从花粉外壁纹饰看,多数花粉外壁具网状或条纹状雕纹,少数花粉的外壁具刺状或颗粒状纹饰。花粉形态特征支持文冠果亚科(Xanthoceroideae)以及广义鳞花木属(Lepisanthes)概念,并支持仍将茶条木属(Delavaya)置于车桑子亚科(Dodonaeoideae)。观察发现黄梨木(Boniodendron minus)与栾树属(Koelreuteria),龙眼属(Dimocarpus)、荔枝属(Litchi)与韶子属(Nephelium)从花粉形态上表现出较近的亲缘关系。基于花粉形态特征编制了国产无患子科分属检索表。     

中国锦葵科15属26个分类群植物花粉形态及其系统学意义

该研究对中国锦葵科15属22种、2变种、2变型及外类群椴树科华椴的花粉形态特征进行了光学显微镜和扫描电镜观察与比较分析,其中7属9种、2变种、2变型植物的花粉为首次光学显微镜观察报道,8属10种、1变种、2变型植物的花粉为首次扫描电镜观察报道,为该科植物的科下分类阶元(族、亚族、属)的划分及其演化关系的探讨提供花粉形态学依据。结果显示：（1）中国锦葵科15属植物的花粉粒均较大,圆球形,直径约54.4~211.5 μm,表面具刺状纹饰,具散孔或孔沟;其演化趋势为从中等大小、3孔沟、具基部膨大的短尖刺的花粉向体积增大、多散孔、具基部平的长刺（有时具二型刺）的花粉演化。（2）锦葵科花粉的形态特征在科、族、亚族和属的划分上有较大的分类学意义,从花粉形态特征角度支持将锦葵科植物划分为4个族。翅果麻属和木槿族成员曾被放入木棉科中,但本研究表明翅果麻属和木槿族的花粉形态符合锦葵科的特征而与木棉科存在明显差异,因此认为宜将这2类群归于锦葵科,并将翅果麻属放到木槿族中。（3）花粉学证据支持棉族和锦葵族具较紧密的亲缘关系,锦葵族中的苘麻亚族可能是该科最原始的类群,梵天花族可能比木槿族更进化。     

中国桑寄生科的孢粉学研究

系统报道了中国桑寄生科Loranthaceae33种5变种植物的花粉形态,并与澳大利亚 2属6种植物的花粉形态做了比较。通过光学显微镜和扫描电镜观察,国产桑寄生科花粉外壁 纹饰可明显分为两个类型:一种类型为刺状或条状纹饰,另一种为颗粒状纹饰,这与该科的鞘 花族和桑寄生族两个族相吻合。在鞘花族类型中,3合沟、钝刺状或条状纹饰的花粉是基本类 型,合半沟或孔沟形,刺状纹饰的花粉是较进化的类型;在桑寄生族类型中,等极、3合沟、 颗粒状纹饰的花粉是基本类型,异极、副合半沟-合半沟、3沟形和沟形-短沟形或沟孔形、粗 糙或模糊颗粒状纹饰的花粉是较进化类型。根据萌发孔和纹饰可将桑寄生族类型花粉分为3个 类群:类群I包括五蕊寄生属Dendrophtho、梨果寄生属Scurrula、钝果寄生属Taxillus和大苞 寄生属Tolypanthus;类群II仅包括离瓣寄生属Helixanthera;类群III也仅1属,桑寄生属Lor anthus。在这3个类群中,类群I属于基本的类型,属间花粉差别较小,其中梨果寄生属和钝 果寄生属花粉差别最小,显示出较近的亲缘关系;类群II和类群III皆是较进化类型。     

中国天南星科花粉形态的研究

利用扫描电镜对天南星科Araceae22属28种(除Arum maculatum产自德国外,其余均产自中国) 及菖蒲科Acoraceae 1属2种植物的花粉形态进行了观察。结果显示天南星科花粉形态在科内变异很 大。花粉粒形状从球形、近球形、椭球形到扁球形和橄榄形;萌发孔类型有散孔型、具薄壁区型、环沟型 或无萌发孔;外壁纹饰为小穴状、网状、肋条状、条纹状、疣状、具刺或光滑。主要依据花粉形态方面的证 据探讨了崖角藤属Rhaphidophora、麒麟叶属Epipremnam 、龟背竹属Monstera 3属的属间关系以及犁头尖属Typhonium属下分类中存在的一些问题。     

Correlation between pollen morphology and pollination mechanisms in the Hydrocharitaceae

The pollen morphology of 11 genera and 11 species of the Hydrocharitaceae and one species of the Najadaceae was studied using scanning and transmission electron microscopies, and the exine structures and sculptures are discussed in relation to pollination mechanisms and the molecular phylogeny. The pollen grains of the Hydrocharitaceae are spherical, inaperturate, and form monads or tetrads, while those of the Najadaceae are elliptical, inaperturate, and form monads. The entomophilous genera Egeria, Blyxa, Ottelia, Stratiotes, and Hydrocharis share pollen grains that have projections like spines or bacula. The anemophilous genus Limnobium has reticulate pollen grains. The hypohydrophilous genera Thalassia and Najas are characterized by pollen grains with reduced exine structures. The pollen-epihydrophilous genera Elodea and Hydrilla have tightly arranged small spinous pollen grains, and the male flower-epihydrophilous genera Enhalus and Vallisneria have reduced reticulate or gemmate exines. Character state reconstruction of the exine structures and sculptures using a molecular phylogenetic tree suggests that variation in the exine is generally correlated with the pollination mechanism; the selective pressures acting on the pollination mechanisms have reduced the exine structure in hypohydrophilous plants and resulted in various exine sculptures that are adapted to the different pollination mechanisms in entomophilous, anemophilous, and pollen-epihydrophilous plants.     

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.     

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 Microula Benth. and Allied Genera (Boraginaceae)

Pollen grains of 16 species of Microula Benth. and six species of three related genera were examined under LM and SEM, and four of them also under TEM. Pollen grains of Microula and three related genera are dumb-bell-shaped, 3-colporate apertures alternate with three pseudocolpi. Pollen grains are very small,ranging from 12.18 x 7.13 μm to 6.36 x 3.66μm. In general, colpi with os are wider and shorter, rhomboid, but sometimes they are equal to pseudocolpi in length. Colpus margins are regularly or irregularly tooth-like. The surface of colpi is psilate or processed. Ora are circular or lalongated in outline, protruded or not; surface of os membrane is smooth or scabrid. The exine is usually indistinctly layered under LM. The exine surface is psilate, and more or less perforate. TEM examination shows that the pollen wall is differentiated into exine and intine: the exine includes ectexine and endexine, while the ectexine consists of tectum, columellae and foot-layer. However, there are differences in constriction of equatorial area, apertural characters, ornamentation and exinous ultrastructure between these genera. Pollen morphology indicates that the genus Microula Benth. is primitive, directly related to the genus Actinocarya Benth .; the genus Asperugo L. Is more advanced. The genus Eritrichium Schrad. which has two ora or one os and is anisopolar, represents the most advanced group among them. Noteworthily, the diorate phenomenon is found for the first time not onlyin the genus but in the family Boraginaceae.     

Phylogenetic Studies in the Monocot Subclass Alismatidae: Evidence for a Reappraisal of the Aquatic Order Najadales

Within the angiosperm subclass Alismatidae (= superorder Alismatiflorae), contemporary taxonomists have often assigned the families Hydrocharitaceae and Najadaceae to different orders. The Najadaceae are presumably allied to a variety of aquatic families in the order Najadales, whereas the Hydrocharitaceae have been segregated as the order Hydrocharitales or placed within the order Alismatales. Analyses of DNA sequence data from the chloroplast gene rbcL, however, indicate that Najadaceae have a much closer phylogenetic relationship to Hydrocharitaceae than to families of the "Najadales" (Cymodoceaceae, Potamogetonaceae, Ruppiaceae, Scheuchzeriaceae, Zannichelliaceae, Zosteraceae). This association supports previous studies based upon examination of floral structure and seed coat anatomy. The rbcL sequence data also indicate that the Zosteraceae and Potamogetonaceae are closely related families. The rbcL sequence of Zostera is actually more similar to that of Potamogeton richardsonii than is the sequence of the latter to a congener, Potamogeton amplifolius. The marine, dioecious, hydrophilous genus Zostera has acquired a number of distinctive adaptations, but probably diverged relatively rapidly from freshwater Potamogetonaceae. Molecular data place Ruppiaceae as a sister group to the marine Cymodoceaceae and do not support the commonly accepted merger of Ruppiaceae and Potamogetonaceae.     

Phylogeny of the family hydrocharitaceae inferred fromrbcL andmatK gene sequence data

The family Hydrocharitaceae, with 15 genera and ca. 80 species, shows a remarkable morphological diversity which presumably developed as an adaptation to their aquatic habitat. This is particularly true in the case of the many different kinds of pollination mechanisms. To gather more basic information regarding the adaptive evolution of Hydrocharitaceae, we have carried out a phylogenetic analysis based on the sequences of therbcL andmatK. Our resulting neighbor-joining distance tree provides the following insights: (1) none of the previous classification systems were supported by molecular phylogenetic tree; (2) Najas (Najadaceae), which has never been included in Hydrocharitaceae except in Shaffer-Fehre's (1991) system based on seed coat structures, is an ingroup of Hydrocharitaceae; (3) Limnocharitaceae and Alismataceae are sister groups of Hydrocharitaceae; (4) the three marine genera,Halophila, Enhalus andThalassia, are monophyletic; and (5) a peculiar pollination mechanism specific to Hydrocharitaceae (Hydrocharitaceae-epihydrophily), underwent a parallel evolution.     

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.