Characters of Leaf Epidermis and Their Systematic Significance in Menispermaceae

The epidermal characters of mature leaves of 29 genera, 50 species and 3 varieties (totally 56 samples) representing all the 5 tribes in the family Menispermaceae were examined under the light microscope. The main conclusions are as follows: (1) The shape of the epidermal cells is polygonal or irregular, and the anticlinal walls are straight or waved in the family. In some genera a special arrangement of epidermal cells is named as “rosette-cell arrangement" for the first time. The lower epidermal cells were found to have a papilla in Stephania, Diploclisia and Legnephora . (2) In some genera, the anticlinal walls are oblique, rather than perpendicular, to the surface. (3) The stomatal apparatuses, generally restricted to the lower surface of the leaves, were assigned to anomocytic, staurocytic, cyclocytic, anisocytic and actinocytic types, and their distribution on the epidermis may be of diffuse pattern or island congregating pattern. (4) The cells of both upper and lower epidermis in the tribe Pachygoneae and Fibraureae are generally polygonal with straight or arched anticlinal walls, and the stomatal apparatuses are usually staurocytic and actinocytic. The cyclocytic stomatal apparatus was found only in two genera of the tribe Pachygoneae. By contrast, the epidermal cells of the tribes Anomospermeae, Tinosporeae and Menispermeae are generally irregular with waved anticlinal walls, and the stomatal apparatuses are predominantly anomocytic. These correlated characters are of much significance in delimiting tribes within the Menispermaceae, and also provide evidence for studies on systematic relationships of several genera.     

甘草属植物叶表皮特征及其系统学意义

应用叶片离析法和石蜡切片法,在光学显微镜下观察了国产甘草属植物10种1变种共16个代表植物的叶表皮特征。统计并测量叶表皮气孔大小、气孔密度及气孔指数。结果表明,该属植物叶表皮细胞形态(表面观)为多边形和不规则形,垂周壁式样为平直、浅波缘、深波缘;气孔器有6种类型,以无规则型、不等型、短平列型为主,一种植物往往具有2~3种气孔器;毛状体为单毛和鳞片毛两种类型。以上特征在属以下、种以上的分类等级具有一定规律性,有的也表现出种间差异,对甘草属属以下等级的划分有重要价值,同时也为本属一些分类群间的系统关系的探讨提供佐证。     

防己科植物的叶表皮特征及其系统学意义

利用光学显微镜观察了防己科29尾50种3变种的叶表皮形态。结果表明：（1）该科植物叶表皮细胞形状为多边形或不规则形,垂周壁式样为平直和波状缘;首次发现一些属的表皮细胞排列成玫瑰花环式样;在Stephania等属发现其下表皮细胞中央有乳突。（2）气孔器为无规则型、环列型、十字型、辐射型和不等细胞型,几科均出现在下表皮,在表皮上的分布有散生和岛状聚集两种式样。（3）在五族中,Pachygoneae族和天仙藤族的上、下表皮细胞多为多边形,垂周壁平直,气孔器多为十字型和辐射型;而另外三族的上、下表皮细胞多为无规则形,垂周壁波状,气孔器则以无规则型为主,这些相关性状对于科下族的划分具有重要价值,同时也为一些属的系统学关系提供了佐证。     

鹿药属植物叶表皮特征及其系统学意义

采用光学显微镜和扫描电镜对鹿药属12种植物的叶表皮进行了观察,首次报道了12种鹿药属(Sm ilaci-na)植物叶表皮的微形态特征。结果表明:气孔器普遍存在于叶的下表皮,少数种的上表皮也有分布,均为不规则形。叶表皮细胞形状为多边形或不规则形,垂周壁式样可区分为近平直、浅波状和波状。在扫描电镜下,叶表皮气孔器外拱盖内缘为近平滑、浅波状或波状;角质膜条纹状,有的条纹隆起,有的条纹上附有颗粒和晶簇。气孔器的分布、气孔器外拱盖内缘形态以及角质膜等特征对该属部分种的区分具有一定的参考价值。     

A Study on Foliar Epidermis in Betulaceae

Light and scanning electron microscopes were used to examine foliar epidermal features such as the shape of epidermal cells, the type of stomatal apparatus and outer stomatal rim in 38 species of 6 genera of the Betulaceae. Four types of stomatal apparatus on mature leaves are recognized in this survey: anomocytic, cyclocytic, non-typical actinocytic and brachyparacytic. The foliar epidermal characters and character states are taxonomically useful for the identification of genera and the division of tribes. The generalized evolutionary scheme for foliar epidermal features is fairly closely correlated with the generic relationships proposed in other studies (Abbe, 1935, 1974; Brunner & Fairbrothers, 1979; Hall, 1952; Kikuzava, 1982; Kuprianova, 1963)and it is clear that foliar epidermal morphology can be an important taxonomic character added to others for determining evolutionary relationships. The explanations of generic relationships are based on our hypothesis that anomocytic stomatal apparatus, uni-layered outer stomatal rim, having not T-pieces at the polar region of guard cells and straight anticlinal wall of lower epidermal cell are from the ancestor (Fig. 1). Sinuolate or sinuate anticlinal wall of lower epidermal cells and non-typical actinocytic stomatal apparatus, present in Corylus, Ostryopsis, Ostrya and Carpinus of the Betulaceae, correlates with the separation of Coryleae from Betuleae. Betuleae including Alnus and Betula is considered primitive with more primitive characters. Of Coryleae, Carpinus might be the most advanced, with brachyparacytic stomatal apparatus and double-layered outer stomatal rim.     

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。     

骨碎补科植物叶表皮特征及其系统学意义

基于骨碎补科(Davalliaceae)植物属的界定和属下等级的划分一直存在较大争议,本研究首次对骨碎补科6属39种(秦仁昌系统)植物的叶表皮进行了扫描电镜观察。结果显示,骨碎补科有9种类型的角质层,其中,阴石蕨属(Humata)的角质层有密集的孔状凹陷结构;Wibelia条纹突起较厚,排列整齐且细密紧致;广义钻毛蕨属(Davallodes)内存在多种角质层类型,需要在属下进一步细分。本研究还根据角质层特征讨论了骨碎补科与一些近缘种类的关系。角质层特征是骨碎补科内种属分类的重要依据,而保卫细胞形状和气孔密度,均不能用来界定骨碎补科的属和种。本研究按角质层分类的结果与Kato和Tsutsumi的分子系统学分类观点一致。     

中国柴胡属植物叶表皮特征及系统学意义

利用光学显微镜和扫描电镜对我国柴胡属（Bupleurum L．）13种（含1变种）植物的叶表皮进行了观察,首次报道了它们的微形态特征。结果表明：除了大叶柴胡（B．longiradiatum Turczaninow）气孔器仅存在于下表皮,其余12种柴胡的上、下表皮都存在气孔器,气孔器类型包括不规则型和不等型两种。叶表皮细胞形状为多边形或不规则型,垂周壁式样可区分为平直一弓形、浅波状。保卫细胞壁加厚明显,极端联合形成极层结构。在扫描电镜下,气孔器内陷于表皮细胞间。角质膜条纹状,有的条纹隆起,有的条纹上附有蜡质胶状分泌物和鳞片。光镜和电镜下叶表皮柴胡属微形态表现出相当高的多态性;而在特定的分类群中,又表现为高度的一致性,为种间分类提供了新的证据。     

The Structural Features of Leaf Epidermis in Oryza and Their Systematic Significance

The rice genus (Oryza L. ) belongs to the grass family(Poaceae) and contains 24 annual or perennial species, including two cultivated rice species, i.e., the Asian rice ( O. sativa L. ) and African rice (O. glaberrima Steud. ), and 22 wild species distributed throughout the tropics of the world. Species in this genus have been extensively studied by scientists with different approaches, including morphological characterization and cytological and molecular investigations. The leaf epidermis is an important morphological character which has been studied for taxonomic identification and studies on systematic relationships of species, particularly in grasses. In this study, morphological features of the leaf epidermis of 23 rice species were observed through light microscopy. The results showed that some characters of the rice leaf epidermis had significant diversity between species and these characters were valuable for the identifying Oryza species, and for assessing systematic relationships in the genus. For example, O.schlechteri, O.ridleyi, O.longiglumis, O.granulata, and O. rneyeriana had elliptic stomatal complexes, whereas the other species had rhombic stomatal complexes. In most cases, papillae on the surface of the epidermis were variable in size and distribution between species. The size of papillae varied from small ( 1.5～4.4µm in diameter), medium-sized (9～18µm), to large (21～30µm) , and the pattern of papillary size and distribution were very useful for identification of rice species. In addition, the number and location of the small papillae in stomatal complexes were particularly different between species. Based on the following combinations of leaf-epidermic characters, i.e., the size and distribution of papillae on the abaxial surface of the epidermis, the number and location of the small papillae in stomatal complexes, and the shape of stomatal complexes, the 23 studied Oryza species could be divided into three major groups. The first group comprises O.longiglumis, O.ridleyi, O.meyeriana, and O.granulata. In these species, neither large nor medium-sized papillae, in some cases extremely rare small papillae, were found on the surfaces of epidermis, and there were no small papillae found in stomatal complexes. All species in the first group had elliptic stomatal complexes. The second group consists of O.brachyantha, diploid and tetraploid O.officinalis, O.minuta, O.eichingeri, O. punctata, O.latifolia, O.alta, O.grandiglumis, O.rhizomatis, and O.australiensis. In these species usually no large papillae were observed, but medium-sized and densely populated small papillae were found to cover the surface of epidermis, and at least four small papillae were found in stomatal complexes (in guard cells) of most species. The third group contains O.sativa, O.nivara, O.rufipogon, O.longistaminata, O. glumaepatula, O.meridionalis, O.barthii, O.glaberrima and O. schlechteri. The abaxial leaf epidermis of these species was usually covered with large papillae, medium-sized, and small papillae. In addition, more than 4 (usually 6～8 ) small papillae were found in guard cells or/and subsidiary cells of the stomatal complexes. Most species in the second and third groups had rhombic stomatal complexes. These results agree mostly with previous re-ports on the biosystematic studies of rice species by applying other methodologies.     

山茶属的叶表皮形态及其分类学意义

在光学显微镜下观察了山茶属（Camellia)19个组36个代表种的叶表皮形态,发现上述植物的叶上下表皮细胞表面观形状均为不规则形,气孔器仅在下表皮存在,且一致为环列型;对上述植物中18个组30个种的叶下表皮进行扫描电镜观察,其特片为：气孔形状为梭形或长卵形,气孔外拱盖内缘光滑或呈浅波状,波状,气孔器附近的角质膜平坦或皱褶,表面光滑或有多种纹饰,这些特征可作为区分种或变种的依据,但与植物的外部形态等特征没有相关性。     

石竹属植物叶表皮特征及其系统学意义

用光学显微镜和扫描电镜,对石竹属12个种的植物叶表皮特征进行观察,统计并测量叶表皮气孔大小、气孔密度及气孔指数。结果表明,石竹属植物叶表皮细胞形态(表面观)为长方形和不规则多边形。乳突只存在于针叶石竹、高石竹和细茎石竹中。按气孔形状可将其分为了3个类型:椭圆形、卵圆形和长方形。研究结果对石竹属的系统分类和种间亲缘关系研究具有一定的参考价值。     

中国无患子科的叶表皮特征及其系统学意义

在光学显微镜和扫描电镜下,对狭义无患子科25属33种植物以及七叶树科2种植物和槭树科2属4种植物的叶表皮特征进行了观察,发现这三科植物叶表皮细胞形状为多边形或无规则形,垂周壁一般为平直、弓形或浅波状.气孔器主要有6种类型,通常仅分布在下表皮(倒地铃Cardiospermum halicacabum和伞花木Eurycorymbus cavaleriei偶尔可在上表皮观察到),除无规则型(如波叶异木患Allophylus caudatu、掌叶木Handeliodendron bodinieri、茶条木Ddavaya toxocarpa等16种)最为常见外,辐射型(细子龙Amesiodendronchinense、七叶树Aesculus chinensis、金钱槭Dipteronia sinensis等11种)和环列型(如龙眼Dimocarpus longan、荔枝Litchi chinensis、韶子Nephelium chryseum等6种)也较为普遍,短平列型(赛木患 Aphania oligophylla、绒毛番龙眼Pometia tometttosa、干果木Xerospemuan bonii、假山萝Harpullia cupaniodes)和十字型(广西檀栗Pavieasia kwangsiensis)仅在少数几属中出现,不等细胞型仅出现在倒地铃Cardioslx,mmm halicacabum中.在扫描电镜下,角质层多具有条纹、鳞片或颗粒等特征;三科植物的气孔外拱盖内缘大多为浅波状.本文试图通过对三科的叶表皮特征,尤其是气孔器类型等形态特征的观察,寻求有意义的叶表皮性状,为进一步探讨无患子科尤其是掌叶木属、茶条木属与七叶树科乃至与槭树科亲缘关系提供资料.研究结果从叶表皮特征的角度说明了这三个科在系统上的密切联系.     

茄科天仙子族的叶表皮特征及其系统学意义

在光学显微镜和扫描电镜下,观察了茄科天仙子族７属２１种及其邻近类群２属３种共２４种植物的叶表皮特征。结果表明：气孔器在各种植物的上、下表皮均有分布,多为无规则型,也有不等细胞型;叶表皮细胞形状有不同规则形、十字形、近长圆形和不规则多边形,垂周壁较平直、浅波状、波状或深波状。在扫描电镜下可见：叶表皮气孔器外拱盖内缘近平滑、浅波状或波状;角质膜多为具条纹,也有同时具条纹和颗粒或鳞片,偶尔仅具颗粒和鳞片     

Structural Characters of Leaf Epidermis in Hyoscyameae (Solanaceae) and Their Systematic Significance

nifH-gfp 表达载体的构建及其在Enterobacter gergoviae 57-7中的表达     

地黄属（Rehmannia）和崖白菜属（Triaenophora）植物的叶表皮形态特征

利用扫描电子显微镜对车前科(P lantaginaceae)地黄属(Rehmannia)和崖白菜属(Triaenophora)8种植物成熟叶片的背面进行了观察。地黄属和崖白菜属的气孔都突出于叶表面,气孔类型均为无规则型;毛被有2种类型,即单列多细胞疏柔毛 单列多细胞腺毛和单列多细胞棉毛 单列多细胞腺毛。气孔和毛被在种间无明显的差异,不能作为分种依据,但毛被在属间具有明显的差异,具有一定的属间分类意义。     

Cytological Studies of the Genus Nomocharis and Its Related Genera

The somatic cells of 12 taxa of the genera Nomocharis, Lilium, Fritillaria and Cardiocrinum from north-western Yunnan were karyomorphologically studied. The 12 taxa are similar to each other in chromosomal morphology at the resting nucleus stage, which belongs to the complex chromocenter type. They are also similar to each other in morphology of mitotic prophase chromosomes, which belongs to the interstitial type. Their chromosome numbers are all 2n = 24, and centromeric terminalization values range from 79.9- 82.2%. The above results show that the four genera have close relationships. These 12 taxa, however, exhibit some similarities and differences in the position of secondary constrictions seemingly showing the karyological characters of each genus in the area. Firstly, on m(sm)-chromosomes, secondary constrictions close to centromeres appear stable in one of pairs in both Nomocharis( 6 taxa) and Lilium (4 taxa); In Cardiocrinum giganteum, they were found distant from centromeres on short arms in one of pairs; None was observed in Fritillaria cirrhosa. Secondly, on st (t)- chromosomes, remarkable secondary constrictions were generally found on long arms in Nomocharis, C. giganteum, F. cirrhosa and the high mountain habitants of Lilium, such as L. souliei, while they were always observed on short arms in L. nepalense, L. taliense and L. brownii, which grow in the lower altitude region. So the Nomocharis and high mountain members of Lilium have similary distribution of secondary constrictions, and theyare considered to be most closely related.     

Character of Leaf Epidermis and Their Systematic Significance of Daphne and Wikstroemia (Thymelaeaceae)

Fifty six species (varieties) of Daphne, forty five species (varieties) of Wikstroemia and twelve species of outgroups from Thymelaeaceae, with ten phenotypic characters were selected, to study the leaf epidermal microfeatures by using light microscopy, Results reveal that Daphne and Wikstromia are consistent in most of leaf features. The epidermal cell are polygonal and irregular in shape. The patterns of anticlinal wall are straight, curved and U shape. The cell sizes range in 30-40μm × 10-20μm. Some species have singly hair on surface. Stomatal distribution is random, size range in 20-36μm × 17-25μm, most are broadly elliptical and narrowly elliptical, stomatal frequency are 100-250S/mm2, and the anomocytic stomatal type is dominate. It is apparently that many stomatal and epidermal features in Daphne and Wikstromia are crossed, and no distinguishable features are suitable for separating this two genera. Results from epidermal characters analysis don’t support monophyletic status of Daphne and Wikstroemia. Eriosolena should not be include in the genus Daphne as the special papillae only found in Eriosolena composita rather than in other genera of Thymeleaceae. In compare with other genera, Rhamnoneuron, Eriosolena, Stelleropsis, Daphnopsis and Edgeworthia in Thymelaeoideae, Daphne and Wikstroemia might be derived and originated latter based on the stomatal features.     

变豆菜属植物叶表皮微形态及分类学意义

对变豆菜属(Sanicula L.)6种13个居群植物的叶表皮特征进行光学显微观察,并测定了相应的数量指标。结果表明,变豆菜属植物叶表皮形态较为一致:上表皮几乎全为多边形,垂周壁平直或弓形弯曲;下表皮多为不规则形,垂周壁浅波状、波状至深波状;气孔散生,均为不等型。但表皮细胞垂周壁式样、气孔长短轴之比等细微特征在种间存在差异,可以区分种类。因此,叶表皮特征对于变豆菜属种类的划分和亲缘关系的确定具有一定的分类学意义。     

Kingdonia,Embryology and Its Systematic Significance

The embryological studies show that Kingdonia is similar to Circaeaster while different from other members of the Ranunculales (sensu Takhtajan, 1980) in development of endosperm and embryo. We consider that there is close systematic relationship between Kingdonia and Circaeaster.