基于不同方法的毛茛族(毛茛科)导管穿孔板比较研究

导管分子的演化与植物的进化相关联。毛茛科是联系基部被子植物和核心真双子叶植物的关键类群,对其导管穿孔板的研究具有重要的系统学意义。为研究毛茛科毛茛族不同属植物导管穿孔板类型及其与环境的相关性,该文利用新的扫描电子显微镜和半薄切片技术,对该族7属8种植物次生木质部导管状分子进行了比较观察。结果表明:木质部管状分子以单穿孔板为主,除了茴茴蒜、鸦跖花和角果毛茛外,其他类群均具有梯状穿孔板;侧金盏花属的蜀侧金盏花导管分子穿孔板的类型多样,有单穿孔板、梯形穿孔板、买麻藤式穿孔板、网状穿孔板和梯-网混合型穿孔板;碱毛茛属的圆叶碱毛茛具单穿孔板、梯形穿孔板和网状穿孔板;在水毛茛属水毛茛的根和茎中未离析出导管,半薄切片显示其具原始的环纹导管。对不同类型导管纹孔膜进行了观察,发现有对列纹孔、互列纹孔,纹孔膜完整或有残余。毛茛族植物导管长度的多样性较高。该文还对不同导管类型的适应性意义和毛茛族的系统位置进行了探讨,揭示了导管分子微形态结构特征与其所处的环境存在一定的适应性。太白美花草、蜀侧金盏花、鸦跖花的导管类型与高寒环境相关,角果毛茛、水毛茛的导管类型分别与其旱生、水生环境相关。     

黄三七(毛茛科)茎次生木质部导管穿孔板的研究

利用扫描电子显微镜对东亚特有植物黄三七（Souliea vaginata（Maxim.） Franch.）茎的次生木质部离析材料进行了观察,结果表明,黄三七茎次生木质部中的导管分子端壁上具网状穿孔板（麻黄式穿孔板）、梯状穿孔板、网状-梯状混合穿孔板、网状-梯状-单穿孔混合型穿孔板、梯状-单穿孔混合型穿孔板及单穿孔板,同时也观察到了端壁多穿孔板和侧壁穿孔板,并对不同类型穿孔板中纹孔膜的残留也进行了观察。其中,网状穿孔板、各种过渡类型的穿孔板均为毛茛科植物中首次报道。根据观察结果,对导管分子穿孔板的演化及黄三七属植物的系统位置进行了分析。     

黄三七( 毛茛科) 茎次生木质部导管穿孔板的研究*

利用扫描电子显微镜对东亚特有植物黄三七( Souliea vaginata (Maxim. ) Franch. ) 茎的次生木质部
离析材料进行了观察, 结果表明, 黄三七茎次生木质部中的导管分子端壁上具网状穿孔板( 麻黄式穿孔
板) 、梯状穿孔板、网状- 梯状混合穿孔板、网状- 梯状- 单穿孔混合型穿孔板、梯状- 单穿孔混合型穿
孔板及单穿孔板, 同时也观察到了端壁多穿孔板和侧壁穿孔板, 并对不同类型穿孔板中纹孔膜的残留也进
行了观察。其中, 网状穿孔板、各种过渡类型的穿孔板均为毛茛科植物中首次报道。根据观察结果, 对导
管分子穿孔板的演化及黄三七属植物的系统位置进行了分析。     

领春木茎次生木质部中导管穿孔板的变异

领春木Euptelea pleiosperma Hook. f. &; Thoms.隶属领春木科Eupteleaceae。该科为东亚特有的单型科,其系统位置一直颇有争议。本文对中国产领春木茎次生木质部中导管穿孔板的变异进行了观察,以期为它的系统位置提供进一步的解剖学证据。结果表明,领春木茎次生木质部中包括无明显穿孔板的管胞状导管和典型的导管两种类型。在无明显穿孔板的导管中,穿孔中的纹孔膜全部或部分消失,但穿孔无规则排列或聚集,不形成具典型的形态特征的穿孔板;在典型的导管中,穿孔板形态变异较大,包括几个类型:网状穿孔板(含麻黄式穿孔板)、网状和梯状混合型穿孔板、梯状穿孔板、梯状穿孔板向单穿孔板的过渡。在上述导管穿孔板类型中,只有梯状穿孔板的穿孔中可以观察到纹孔膜的残余。在领春木次生木质部中也观察到了端壁多穿孔板及侧壁穿孔板。根据观察结果,我们认为领春木次生木质部导管穿孔板的许多特征说明该科可能处于毛茛目中比较原始的系统位置。     

花楸(Sorbus pohuashanensis)导管分子穿孔板的类型及演化

对花楸茎次生木质部进行离析研究,发现其导管穿孔板有两种类型,即网状穿孔板和单穿孔板,并且还有5种过渡类型,具不同类型穿孔板的导管分子有3种类型,即两端均为网状穿孔板的导管分子;一端为网状穿孔板,另一端为单穿孔板的导管分子;两端的均为单穿孔板的导管分子,在花楸个体发育过程中穿孔板类型的演化重演了系统发育过程中导管网状穿孔板演化成单穿孔板的过程。     

余甘子次生木质部导管分子观察研究

运用细胞图象分析系统及显微照相的方法,对余甘子次生木质部导管分子进行观察研究.结果发现,余甘子次生木质部导管分子中存在着许多不同的样式,导管分子大多数具尾;其穿孔板存在着两种类型:(1)两端均为1个单穿孔板;(2)一端为1个单穿孔板另一端为2个单穿孔板;(3)极少数的导管分子具有特殊的内含物;(4)管间纹孔式为互列纹孔式;(5)导管射线间纹孔式为混合型纹孔与横列刻痕状纹孔以及梯状穿孔.     

两种麻黄根与茎次生木质部的解剖与进化和干旱环境的关系

用光镜及扫描电镜对两种麻黄根、茎次生木质部进行了解剖研究,结果表明：轴向系统主要由导管和管胞组成。横向系统由细胞壁木质化了的射线薄壁组织细胞组成。导管直径甚小,多孔式穿孔板,并存在导管与管胞之间的管状分子类型,推断麻黄属是裸子植物中最早出现导管的类群;管胞中有一些两头尖、胞腔小、具缘纹孔含纹孔塞的长分子,可视作纤维状管胞,使管胞的输导作用被削弱,而支持功能被加强;射线异型多列,已不具备裸子植物具较窄射线的特点。导管与管胞并存,恰好起到了一般沙生被子植物具宽窄两种类型的导管、复孔率高等典型的对干旱环境的适应特征的作用,茎中导管分子的长度和宽度均小于根,这与茎部需要较强的机械支持力相一致。     

国产球盖蕨科植物管状分子的比较研究

利用扫描电镜观察了国产球盖蕨科10种植物,鳞毛蕨科6种植物的管状分子,结果显示:它们的管状分子端壁和侧壁的形态及结构分别相同,且侧壁具有穿孔板。它们具有4种类型的管状分子:(1)梯状穿孔板,无穿孔板的二型性现象;(2)梯状穿孔板,具有二型性现象;(3)梯状-网状混合穿孔板;(4)大孔状穿孔板。穿孔板仅存在于端壁的管状分子为导管分子,而端壁和侧壁形态、结构相似,有或无穿孔板的管状分子为管胞,蕨类植物中的管状分子主要为管胞,这与传统观点不同。管状分子的形态特征表明:球盖蕨科是鳞毛蕨群的成员,但不是原始成员,可能属于其中较为进化的类群,与鳞毛蕨科有许多共同特征,但仍存在较大差异,所以将其作为独立的科是合理的,推测球盖蕨科中的鱼鳞蕨属是比较进化的属,柄盖蕨属相对原始,红腺蕨属的系统位置应介于二者之间。     

早期维管植物两种特殊的后生木质部管胞

研究现代植物维管系统中木质部的管状分子(tracheary element),包括管胞(tracheid)和导管分子(vessel member),特别是研究它们的次生壁加厚类型是植物解剖学的一个重要内容。     

国产对囊蕨亚科（蹄盖蕨科）植物的管状分子

利用扫描电镜观察了国产蹄盖蕨科（Athyriaceae）对囊蕨亚科（Deparioideae）10种植物及双盖蕨属（Diplazium Sw．）3种植物根状茎的管状分子。结果显示,这些管状分子端壁和侧壁的形态及结构分别相同且侧壁具有穿孔板（多穿孔板）。根据穿孔板的形态特征,将该亚科的管状分子分为5种类型：（1）梯状穿孔板,无穿孔的二型性现象：（2）梯状穿孔板,有穿孔的二型性现象：（3）网状穿孔板：（4）梯状-网状混合的穿孔板：（5）大孔状穿孔板。按照纹孔膜残留的程度又可分为3种：部分区域有完整的纹孔膜、残留呈网状或线状以及很少或无纹孔膜残留。结合前人的研究资料,发现蕨类植物的管状分子与被子植物的导管分子在形态和输导机理上存在明显差异,管胞和导管分子不能仅仅根据纹孔膜的存在与否来确定,而应根据穿孔板存在于端壁还是侧壁进行判断,即穿孔板仅存在于端壁的管状分子为导管分子：端壁和侧壁形态及结构分别相同,有或无穿孔板的管状分子为管胞。由此可以推测蕨类植物和裸子植物中输导水分和矿物质的管状分子主要为管胞。单叶双盖蕨属（Triblemma（J．Sm．）Ching）与双盖蕨属管状分子的特征并不相似,显示了将单叶双盖蕨属从双盖蕨属独立出来归人对囊蕨亚科的合理性。根据管状分子的特征,推测假蹄盖蕨属（Athyriopsis Ching）和蛾眉蕨属（Lunathyrium Koidz．）可能是比较进化的属,而介蕨属（Dryoathyrium Ching）相对比较原始,单叶双盖蕨属的系统位置应介于假蹄盖蕨属与介蕨属之间。     

国产对囊蕨亚科(蹄盖蕨科)植物的管状分子

利用扫描电镜观察了国产蹄盖蕨科(Athyriaceae)对囊蕨亚科(Deparioideae)10种植物及双盖蕨属(Diplazium Sw.)3种植物根状茎的管状分子。结果显示, 这些管状分子端壁和侧壁的形态及结构分别相同且侧壁具有穿孔板(多穿孔板)。根据穿孔板的形态特征, 将该亚科的管状分子分为5种类型: (1)梯状穿孔板, 无穿孔的二型性现象; (2)梯状穿孔板, 有穿孔的二型性现象; (3)网状穿孔板; (4)梯状-网状混合的穿孔板; (5)大孔状穿孔板。按照纹孔膜残留的程度又可分为3种: 部分区域有完整的纹孔膜、残留呈网状或线状以及很少或无纹孔膜残留。结合前人的研究资料, 发现蕨类植物的管状分子与被子植物的导管分子在形态和输导机理上存在明显差异, 管胞和导管分子不能仅仅根据纹孔膜的存在与否来确定, 而应根据穿孔板存在于端壁还是侧壁进行判断, 即穿孔板仅存在于端壁的管状分子为导管分子; 端壁和侧壁形态及结构分别相同, 有或无穿孔板的管状分子为管胞。由此可以推测蕨类植物和裸子植物中输导水分和矿物质的管状分子主要为管胞。单叶双盖蕨属(Triblemma(J. Sm.) Ching)与双盖蕨属管状分子的特征并不相似, 显示了将单叶双盖蕨属从双盖蕨属独立出来归入对囊蕨亚科的合理性。根据管状分子的特征, 推测假蹄盖蕨属(Athyriopsis Ching)和蛾眉蕨属(Lunathyrium Koidz.)可能是比较进化的属, 而介蕨属 (Dryoathyrium Ching)相对比较原始, 单叶双盖蕨属的系统位置应介于假蹄盖蕨属与介蕨属之间。     

慈姑不定根中导管分子的观察

慈姑导管仅在根中出现。根的后生木质部中央导管由顶端平截、单穿孔的网纹导管分子连接组成;周围较小的导管分子和管胞有从梯纹管胞向导管分子演化的各种过渡类型;有一至多个梯形穿孔或单穿孔发生在导管分子的端壁或侧壁,并有分枝型导管分子存在,特别在根与主茎连接处尤为明显。管胞亦有分枝与不分枝的类型。     

Vessels in ferns: structural, ecological, and evolutionary significance

We have studied macerated xylem of ferns, supplemented by sections, by means of scanning electron microscopy (SEM) in a series of 20 papers, the results of which are summarized and interpreted here. Studies were based mostly on macerations, but also on some sections; these methods should be supplemented by other methods to confirm or modify the findings presented. Guidelines are cited for our interpretations of features of pit membranes. Fern xylem offers many distinctive features: (1) presence of numerous vessels and various numbers of tracheids in most species; (2) presence of vessels in both roots and rhizomes in virtually all species; (3) presence of specialized end walls in vessels of only a few species; (4) multiple end-wall perforation plates in numerous species; (5) lateral-wall perforation plates in numerous species; (6) porose pit membranes associated with perforation plates in all species; and (7) pit dimorphism, yielding wide membrane-free perforations alternating with extremely narrow pits. Multiple end wall perforation plates and lateral wall perforation plates are associated with the packing of tracheary elements in fascicles in ferns: facets of tips of elements contact numerous facets of adjacent elements; all such contacts are potential sites for conduction by means of perforations. This packing differs from that in primary xylem of dicotyledons and monocotyledons. Porosities in pit membranes represent a way of interconnecting vessel elements within a rhizome or root. In addition, these porosities can interconnect rhizome vessel elements with those of roots, a feature of importance because roots are adventitious in ferns as opposed to those of vascular plants with taproots. Fully-formed or incipient (small-to-medium sized porosities in pit membranes) perforation plates are widespread in ferns. These are believed to represent (1) ease of lysis of pit membranes via pectinase and cellulase; (2) numerous potential sites for perforation plate formation because of fasciculate packing of tracheary elements; (3) evolution of ferns over a long period of time, so that lysis pathways have had time to form; (4) lack of disadvantage in perforation plate presence, regardless of whether habitat moisture fluctuates markedly or little, because ferns likely have maintaining integrity of water columns that override the embolism-confining advantage of tracheids. Although all ferns share some common features, the diversity in xylem anatomy discovered thus far in ferns suggests that much remains to be learned.     

Vessel-bearing stems of ASOVINEA TIANIIgen. et sp. nov. (Gigantopteridales) from theUpper Permian of Guizhou Province, China

Permineralized gigantopterid stems of Vasovinea tianii Li et Taylor gen. et sp. nov. were collected from the Upper Permian of Guizhou Province, China. They are slender and bear prickles, trichomes, and compound hooks. Internally, the stems have a sparganum cortex, eustele, and secondary xylem. The mesarch protoxylem tracheids have annular to helical thickenings, and metaxylem tracheary elements have scalariform and/or transversely elongated, bordered pits, while those of the secondary xylem have scalariform to circular bordered pits. Importantly, the inner part of the secondary xylem has large vessel elements with foraminate-like perforation plates. The hooks and other morphological and anatomical characteristics are similar to those found in gigantopterids, suggesting that Vasovinea is a member of the Gigantopteridales. The vegetative plant is reconstructed from permineralized stems and Gigantopteris-type leaves based on the anatomical similarities and intimate association. The eustele, secondary xylem, and other features support the placement of the order among the seed plants. Ecologically, Vasovinea is suggested to have been a vine or liana that used compound hooks to climb among the trees in a Permian tropical rain forest. The occurrence of vessels could have been an efficient adaptation to allow the slender stems to conduct sufficient water to the large Gigantopteris-type leaves.     

Discovery of vessels in Tetracentron (Trochodendraceae) and its systematic significance

Through SEM observation, we found that there are vessels as well as tracheids in the secondary wood of Tetracentron sinense Oliv. The vessel elements are as narrow and as long as or slightly shorter than the tracheids and generally have 1 to 3 very long, or sometimes relatively short and oblique end-wall perforation plates; such perforation plates are also present on the lateralwalls. The perforation plates of the vessels include scalariform and reticulate-scalariform types, with various degrees of membrane remnants present in the end walls.     

木通科4属植物导管穿孔板的比较研究

运用扫描电子显微镜法(SEM)对木通科(Lardizabalaceae)4属植物茎的次生木质部导管分子进行观察,结果表明:(1)端壁均具有单穿孔板;(2)串果藤属的导管分子具有丰富的穿孔板类型,包括网状、梯状、单穿孔及过渡类型,穿孔具有网状、丝状、片状的纹孔膜残余;大血藤属和八月瓜属的导管分子具有相似的特征,端壁具有梯状、单穿孔及梯—单混合穿孔板;野木瓜属只具有单穿孔板;(3)侧壁上具有穿孔板,多为梯状或梯—网混合类型(除了野木瓜属);(4)野木瓜属的导管侧壁具有独特的螺旋状加厚。各属导管的不同特征为木通科的系统演化提供比较可靠的依据。