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。     

蹄盖蕨科5种植物管状分子的研究

采用扫描电镜观察了国产蹄盖蕨科3属5种植物的管状分子,发现安蕨属、拟鳞毛蕨属和蹄盖蕨属管状分子结构类似,具体可分为3种类型:(1)梯状穿孔板,无穿孔板二型性现象;(2)梯状穿孔板,有穿孔板二型性现象;(3)梯状-网状混合穿孔板.除长江蹄盖蕨不具有梯状穿孔板,有穿孔板二型性现象外,其余4种均具有上述3种类型.该结果支持安蕨属、拟鳞毛蕨属和蹄盖蕨属三者之间有亲密关系的观点,并在前人基础上提出了新的管胞定义:管胞是指维管植物木质部中存在的一类狭长中空,端部圆凸或尖削,不具有明显端壁,侧壁具有多个侧壁穿孔板,纹孔膜从缺失到不同程度存在的死细胞.     

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。     

蹄盖蕨科三属植物管状分子的研究

扫描电镜观察假冷蕨属4种、冷蕨属3种、蹄盖蕨属3种植物根状茎中的管状分子,结果显示:这些管状分子端壁和侧壁的形态、结构相同且侧壁具有穿孔板(多穿孔板)。根据穿孔板的类型和穿孔的纹孔膜残留程度,我们发现假冷蕨属与蹄盖蕨属亲缘关系较近,进化地位较高,冷蕨属的进化地位相对原始。     

黑龙江省蹄盖蕨科（Athyriaceae）植物的RAPD分析及其系统学意义

在形态分类研究的基础上,利用RAPD技术,对黑龙江省蹄盖蕨科植物6属9种共15个种群进行了遗传多样性分析,分析结果表明：（1）假冷蕨属应包括在蹄盖蕨属中（2）角蕨属与蹄盖蕨属,羽节蕨属与冷蕨属亲源关系较近（3）短肠蕨属为独立一属,与其它属亲源关系较远。根据黑龙江省蹄盖蕨科植物的DNA水平的研究结果,再结合其形态学特征,建议将黑龙江省蹄盖蕨科划分为3个亚科：冷蕨亚科（Cystopterioideae）包括2个属：冷蕨属（Cystopteris）和羽节蕨属（Cemnocarpium）。蹄盖蕨亚科（Athyrioideae）包括3个属：蹄盖蕨属（Athyrium）、假冷蕨属（Pseudocystopteris）和角蕨属（Cornopteris）。双盖蕨亚科（Diplazioideae）包括短肠蕨属（Alhntodia）。     

蹄盖蕨科的亚科划分的修订

蹄盖蕨科Athyriaceae是蕨类植物中复杂的大科,分子系统学的研究证据表明它是一个自然类群。前人根据染色体的基数,将蹄盖蕨科划分为3个亚科,但没有得到分子证据的支持;本文依据分子系统学的研究结果,再结合形态特征,将该科重新划分为5个亚科：冷蕨亚科、蹄盖蕨亚科、对囊蕨亚科、双盖蕨亚科和轴果蕨亚科。     

类叶升麻(毛茛科)次生木质部管状分子的研究

利用扫描电子显微镜对毛良科类叶升麻（Artaea asiaticaHara）根和根状茎次生木质部中的管状分子进行观察．发观其管状分子类型丰富,主要有：管胞、管胞状导管、纤维导管和典型的导管分子,其中管胞、管胞状导管和纤维导管为在该类群中首次报道;在导管分子中．存在着梯状穿孔板、网状穿孔板、混合型穿孔板和单穿孔板．其中网状穿孔板和混合型穿孔板为在陔类群中的首次报道;对其导管分子上的侧壁穿孔板、多穿孔板和纹孔膜残余也进行了描述。根据类叶升麻次生木质部中多变的管状分子类型,认为以往积累的有关毛茛科植物管状分子类型及导管穿孔板类型是小个面的,因此以该性状为参考作出的有关某一个类群的原始性和进化性的推论也是不可靠的。同时探讨了不同类型管状分子作类叶升麻不同器官的分布与其生理功能和生态环境的关系,同时将该植物作为毛莨科的代表类群．与其它基邴类群植物导管分子进行了比较。     

黑龙江省蹄盖蕨科(Athyriaceae)植物的RAPD分析及其系统学意义

在形态分类研究的基础上,利用RAPD技术,对黑龙江省蹄盖蕨科植物6属9种共15个种群进行了遗传多样性分析,分析结果表明：（1）假冷蕨属应包括在蹄盖蕨属中（2）角蕨属与蹄盖蕨属,羽节蕨属与冷蕨属亲源关系较近（3）短肠蕨属为独立一属,与其它属亲源关系较远。根据黑龙江省蹄盖蕨科植物的DNA水平的研究结果,再结合其形态学特征,建议将黑龙江省蹄盖蕨科划分为3个亚科：冷蕨亚科(Cystopterioideae)包括2个属：冷蕨属(Cystopteris)和羽节蕨属(Gymnocarpium)。蹄盖蕨亚科(Athyrioideae)包括3个属：蹄盖蕨属(Athyrium)、假冷蕨属(Pseudocystopteris)和角蕨属(Cornopteris)。双盖蕨亚科(Diplazioideae)包括短肠蕨属(Allantodia)。     

蹄盖蕨科的亚科划分

蹄盖蕨科的建立对蕨类植物的分类起了很大的推动作用,然而,即使按秦仁昌的狭义概念,它仍然是一个极其复杂的大科。自秦仁昌把该科划分为21个属以后,属级水平上的分类问题比较清楚了,但亚科、亚属和组的划分至今仍不成功。本文主要根据该科三个染色体基数X=42、41和40,结合孢子囊群着生的位置及其它形态特征,将其划分为3个亚科：冷蕨亚科、蹄盖蕨亚科和双盖蕨亚科。     

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。     

SEM studies on vessels in ferns. 2. Pteridium

Xylem from roots and rhizomes of two infraspecific taxa of Pteridium aquilinum was studied by means of scanning electron microscopy (SEM). All tracheary elements proved to be vessels. End wall perforation plates were all scalariform, lacked pit membrane remnants in at least the central part of the perforation plate, and varied with respect to width of bars, from wide to tenuous, and with respect to presence of pit membrane remnants. In addition, porose pit membranes on walls that are likely all lateral vessel-to-vessel walls must be considered to be perforations also, although different from those on end walls. Lateral wall perforation plates, hypothesized by one worker on the basis of tylosis presence but denied by another on the basis of light microscopy, were confirmed by demonstration of pores with SEM. In addition, lateral walls of Pteridium vessels bear some grooves interconnecting pit apertures; this feature is newly figured by SEM for ferns. Lateral wall pitting that is not porose may either have striate thickenings of the primary wall or be smooth. Vessel presence and degree of specialization in Pteridium vessels may bear a relationship to the wide ecological tolerances of the genus.     

PRESENCE OF VESSELS IN WOOD OF SARCANDRA (CHLORANTHACEAE); COMMENTS ON VESSEL ORIGINS IN ANGIOSPERMS

Sarcandra is the only genus of Chloranthaceae hitherto thought to be vesselless. Study of liquid-preserved material of S. glabra revealed that in root secondary xylem some tracheary elements are wider in diameter and have markedly scalariform end walls combined with circular pits on lateral walls. Examination of these wider tracheary elements with scanning electron microscope (SEM) demonstrated various degrees of pit membrane absence in the end walls. Commonly a few threadlike fibrils traverse the pits (perforations); these as well as intact nature of pit membranes in pits at ends of some perforation plates are evidence that lack of pit membranes does not result from damage during processing. Some perforations lack any remnants of pit membranes. Although perforation plates and therefore vessels are present in Sarcandra roots, no perforations were observed in tracheary elements of stems or lignotubers. Further, stem tracheids do not have the prominently scalariform end walls that the vessel elements in roots do. Presence of vessels in Sarcandra removes at least one (probably several) hypothetical events of vessel origin that must be postulated to account for known patterns of vessel distribution in angiosperms, assuming that they are primitively vesselless. Seven (perhaps fewer) vessel origin events in angiosperms could account for these patterns; two of those events (Nelumbo and monocotyledons) are different from the others in nature. Widely accepted data on trends of vessel specialization in woody dicotyledons yield an unappreciated implication: vessel specialization has happened in a highly polyphyletic manner in dicotyledons, and therefore multiple vessel origins represent a logical extension backward in time. If a group of vesselless dictyoledons ancestral to other angiosperms existed, they can be hypothesized to have had a relatively homogeneous floral plan now that Sarcandra-like plants no longer need be imagined within that group. Sarcandra and other Chloranthaceae show that the borderline between vessel absence and presence is less sharp than generally appreciated.     

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.     

Sem studies on vessels in ferns. 12. Marattiaceae, with comments on vessel patterns in eusporangiate ferns

Scanning electron microscopy (SEM) of tracheary elements of roots of five species from four genera of Marattiaceae and of the rhizome of one species revealed vessel elements present in all. The secondary wall framework of perforation plates is the same as that of lateral wall pitting for vessel elements in all species. Thus, no specialization is present in perforation plates of Marattiaceae compared to the simplified morphology of perforation plates of some leptosporangiate ferns (e.g., Dryopteridaceae, Polypodiaceae, and Pteridaceae). The difference between lateral wall pitting and perforation plates in tracheary elements of Marattiaceae cannot be seen by light microscopy (in which pit membranes are transparent), but is evident with SEM. Diversity in structure of perforation plates (especially the alternation of wide and narrow perforations within a plate) and presence of web-like pit membrane remnants are evident. Vessels are widespread in both leptosporangiate and eusporangiate ferns, although specialization in perforation plates (e.g., bars few and more widely spaced in lateral wall pitting of a given vessel element) is to be expected only in ferns of habitats with marked fluctuation in water availability. Vessels of Marattiaceae lack such specializations and are thus are correlated with the mesic habitats characteristic for the family.     

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.     

Origins and nature of vessels in Monocotyledons. 14. Vessellessness in Orontioideae (Araceae): adaptation or relictualism?

SEM studies of tracheary elements of subfamily Orontioideae (Lysichiton, Orontium, Symplocarpus) of Araceae show unexpected features. The plants are entirely vesselless. There are small pores in pit membranes of end walls of tracheids in roots and stems, but pit membranes remain intact. End wall pit membranes of stems have a coarse fibrillar texture, somewhat reminiscent of (but different from) those of Nymphaeaceae and Cabombaceae. Acoraceae, which are also vesselless, represent the first branch of the monocot tree, according to phylogenies, and the orontioids form the next branch. Vessellessness is therefore a potentially plesiomorphic feature in monocots, but it may also be related to the highly mesic habitats of Acoraceae and the orontioids. Various other non‐submersed monocots have vesselless or near‐vesselless xylem. Sectioned xylem of Orontioideae is also very suggestive of stages in the development of the pit membranes of both end walls and lateral walls of tracheids: open networks of cellulosic fibrils apparently precede the addition of denser fibrillar meshes, key information in assessing to what extent perforations in scalariform perforation plates of vascular plants may stop formation at the open network stage, and to what extent a thicker pit membrane experiences lysis and disintegration as the vessel element matures.