Origin and nature of vessels in monocotyledons. 5. Araceae subfamily Colocasioideae

Tracheary elements from macerations of roots and stems of one species each of five genera of Araceae subfamily Colocasioideae were studied by means of SEM (scanning electron microscopy). All of the genera have vessel elements not merely in roots, as previously reported for the family as a whole, but also in stems. The vessel elements of stems in all genera other than Syngonium are less specialized than those of roots; stem vessel elements are tracheid-like and have porose pit membrane remnants in perforations. The perforations with pit membrane remnants demonstrate probable early stages in evolution of vessels from tracheids in primary xylem of monocotyledons. The vessel elements with such incipient perforation plates lack differentiation in secondary wall thickenings between perforation plate and lateral wall, and such vessel elements cannot be identified with any reliability by means of light microscopy. The discrepancy in specialization between root and stem vessel elements in genera other than Syngonium is ascribed to probable high conductive rates in roots where soil moisture fluctuates markedly, in contrast with the storage nature of stems, in which selective value for rapid conduction is less. Syngonium stem vessels are considered adapted for rapid conduction because the stems in that genus are scandent. Correlation between vessel element morphology and ecology and habit are supported. Although large porosities in vessel elements facilitate conduction, smaller porosities may merely represent rudimentary pit membrane lysis.     

The anatomy of lotus fibers found in petioles of Nelumbo nucifera

Lotus fibers are the isolated helical secondary cell wall thickenings from tracheary elements of lotus (Nelumbo nucifera Gaertn) petioles. In this study the anatomical characteristics of lotus petioles and microstructures of tracheary elements were studied using light microscopy (LM) and scanning electron microscopy (SEM). The results show that vascular bundles of lotus petioles are scattered throughout ground tissue. Their tracheary elements are of various sizes and there are several patterns of secondary wall thickening present. However, only secondary thickening in a ribbon-like helical pattern can be drawn out from the petiole to form lotus fibers for subsequent utilization. Study of the microstructure of the tracheary elements reveals that there are two pit structures present in the end walls in addition to pits with intact pit membranes: those with porose or web-like remnants pit membrane and those that lack pit membranes. This is an indication of the transitional stage between tracheids and vessel elements. This study provides supportive evidence that lotus fibers are found in both helically thickened tracheids and helically thickened primitive vessels.     

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.     

The tracheid-vessel element transition in angiosperms involves multiple independent features: cladistic consequences

Current definitions of tracheids and vessel elements are overly simple. These definitions are based on light microscope studies and have not incorporated information gained with scanning electron microscopy (SEM) or transmission electron microscopy (TEM). Current definitions are based primarily on angiosperms, especially eudicots, and were devised before many basal angiosperms were carefully studied. When all sources of information are taken into account, one can recognize changes in six characters in the evolution of tracheids into vessel elements in angiosperms (or vice versa) as well as in other groups of vascular plants. There is an appreciable number of taxa in which all criteria for vessel origin are not met, and thus incipient vessels are present. At the very least, vessel presence or absence should not be treated as a single binary character state change in construction of cladistic matrices. Increase in conductive area of an end wall by means of lysis of progressively greater areas of pit membrane and increase in pit area on the end wall (as compared to pit area on equivalent portions of lateral walls) are considered the most important usable criteria for recognizing intermediacy between tracheids and vessel elements. Primitive character states in vessel elements are briefly discussed to differentiate them from changes in character states that can be regarded as intermediate between tracheids and vessel elements.     

Origins and nature of vessels in monocotyledons. 2. Juncaginaceae and Scheuchzeriaceae

SEM studies of roots and rhizomes of Triglochin (one species) and Maundia (monotypic) of Juncaginaceae and the sole species of Scheuchzeriaceae, Scheuchzeria palustris, reveals that vessels are present not only in roots, as previously reported, but also in rhizomes. The perforations contain pit membranes with pores of various sizes. Striate pit membranes, like those previously seen in Acorus, occur on pit remnants in peforations and on pit membranes of lateral walls in all genera studied. Grooves interconnecting pit apertures are illustrated for root tracheary elements of Triglochin; this is believed to be a first report of this feature for monocotyledons. The tracheary elements of Juncaginaceae and Scheuchzeriaceae are similar in their thick walls and narrow slitlike pits, lending support to the close relationship between the two families often claimed.     

PIT MEMBRANE REMNANTS IN PERFORATION PLATES OF PRIMITIVE DICOTYLEDONS AND THEIR SIGNIFICANCE

Perforations of vessel elements characteristically retain remnants of pit membranes (primary walls) in woods of species of more than 30 families of dicotyledons. Scanning electron microscopy is necessary to demonstrate presence and type of membrane remnant. Species with these remnants in perforations given in earlier literature as well as those newly reported here are listed. Perforation membrane remnants may take the form of flakes, strands, or webs, and particular types may characterize particular families (e.g., strands or bands in Illiciaceae). Some families have abundant perforation membrane remnants (e.g., Chloranthaceae, Illiciaceae). Where membranes are nearly intact, they are porose and closely resemble the porose pit membranes on end walls of Tetracentron tracheids. In Tetracentron, however, tracheary elements are monomorphic, so vessel origin cannot yet be said to have occurred. Membrane remnants in perforations are regarded as a relictual primitive feature that should be added to the list of primitive character states claimed for vessel elements in angiosperms; alternative hypotheses are considered and discussed, and evidence from DNA phylogenies is needed. In vessel-bearing dicotyledons with membrane remnants in perforations, many perforations are relatively clear, but an appreciable proportion of perforation plates do have membrane remnants.     

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.     

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。