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.     

LECLERCQIA COMPLEXA (LYCOPSIDA,MIDDLE DEVONIAN): ITS ANATOMY,AND THE INTERPRETATION OF PYRITE PETRIFACTIONS

The development and/or modification of special acid etching and macerating techniques permits pyritized fossils of Leclercqia complexa to be separated into their carbonaceous wall remains and pyrite lumen—pit casts. These carbonized and pyritic portions can then be studied separately by both light and SEM microscopy and information from both modes compared with that obtainable from carbonaceous compression fossils of L. complexa. This combination of techniques, preservation modes and methods of analysis, allows a synthesis of information obtainable from each, as well as providing a check on errors of interpretation due to preservation mode and method of observation. The developmental sequence of the secondary wall thickenings of the protoxylem elements is shown to be annular to connected annular and spiral, with both exarch and mesarch maturation. The metaxylem shows a transition through reticulate to multiseriate round, oval, and elongate bordered-pitted tracheids. Different wall facets of separated pyrite tracheid casts can be examined by the SEM and variations in wall structure and pitting characterized. Cells of the outer cortex are 6 to 10 times longer than wide, have thickened vertical walls and end walls that vary from nearly horizontal to angles of overlap of 50 degrees.     

PITTING IN PSILOPHYTON DAWSONII,AN EARLY DEVONIAN TRIMEROPHYTE

Tracheids of Psilophyton dawsonii Banks, Leclercq, and Hueber 1975 from calcareous pebbles of late Early Devonian age on the Gaspé Peninsula, eastern Canada, are shown to have scalariform bordered and circular bordered pits. Macerated tracheids embedded in Spurr epoxy resin and sectioned at 2 μm; peels, ground sections, SEM and petrographic observations all indicate that interconnections between the scalariform bars are composed of secondary wall material and that the patterns produced are not artifacts of preservation. The interconnections produce a wide range of patterns from simple vertical strands, to reticula, to more extensive interconnections that outline circular openings. The circular openings result in scalariform bordered pits that are considered to be multiaperturate. P. dawsonii is believed to illustrate the most complex pitting yet demonstrated conclusively in Early Devonian time. Among trimerophytes P. forbesii, tracheids of P. charientos, and Hostinella from Röragen, Norway lack only the circular bordered pits.     

Examination of morphological changes in the first formed protoxylem in Arabidopsis seedlings

We examined morphological changes in the first-formed protoxylem vessels in Arabidopsis seedlings. Between 2.5 and 8 days after imbibition, mean hypocotyl and root length increased 1.52 and 23.3 times, respectively. In the 2.5-day-old seedlings, two continuous protoxylem vessels were present in the hypocotyl-root axis. In the 8-day-old upper hypocotyls, six protoxylem vessels were observed, and in the lower hypocotyls, four protoxylem vessels and one or two metaxylem vessels were observed. In the 8-day-old roots, there were two protoxylem vessels and two or three metaxylem vessels. Two protoxylem vessels in the hypocotyls connected to two metaxylem vessels in the roots of 8-day-old seedlings. At the 0.3-mm part below the hypocotyl-root boundary, the mean intervals of neighboring annular secondary wall thickenings in protoxylem vessels in 8-day-old roots were 12.9% larger than those in 2.5-day-old roots. In more apical parts of 8-day-old roots, the mean intervals fluctuated between 1.71 and 2.29 μm. In 8-day-old seedlings, metaxylem vessels were formed between 0.4 mm above the hypocotyl-root boundary and 17 mm below the boundary. The intervals in these regions were not extended so much as protoxylem vessels were collapsed. The first-formed protoxylem vessels presumably retain their water-conductive function after metaxylem formation. Received: April 10, 2001 / Accepted: November 7, 2001     

The Orientation and Localization of Cortical Microtubules in Differentiating Conifer Tracheids during Cell Expansion

Arrays of cortical microtubules (MTs) on radial walls in differentiatingtracheids of Taxus cuspidata were randomly oriented when primarywalls formed. The orientation of MTs changed progressively fromlongitudinal to transverse as cells expanded. During formationof primary walls, MTs in differentiating tracheids disappearedlocally at sites of future intertracheal bordered pits. In furtherdifferentiated tracheids, circular bands of MTs were observedaround the edges of developing bordered pits. (Received July 17, 1996; Accepted November 11, 1996)     

STUDIES OF PALEOZOIC FERNS: TUBICAULIS STEWARTII SP. NOV. AND EVOLUTIONARY TRENDS IN THE GENUS

Eggert , Donald A. (Yale U., New Haven, Conn.) Studies of Paleozoic ferns: Tubicaulis stewartii sp. nov. and evolutionary trends in the genus. Amer. Jour. Bot. 46(8): 594–602. Illus. 1959.—Tubicaulis stewartii, a new species of the order Coenopteridales is described. The specimen was derived from the Upper Pennsylvanian of Berryville, Illinois, and is characterized by having a lacunar middle cortex, a well-developed integumentary system bearing uniseriate hairs, and xylem parenchyma organized into vertically anastomosing strands. In addition, multiseriate (somewhat transitional to reticulate) bordered pitting is present in the petiolar metaxylem elements, while those of the stem stele are multiseriate scalariform. The habit is intermediate between that of a form such as Osmunda and a tree fern, having an upright tapering stem which gives off prominently decurrent petioles in a 2/5 divergence. A reinvestigation of the type specimen of the most closely allied species, T. multiscalariformis, of Upper-Middle Pennsylvanian age, has shown that it has similar features in the cortex, metaxylem, and integumentary layers. Tubicaulis multiscalariformis and T. stewartii form a distinct group in the 6 species now known, whose evolution has most likely involved the retention of a more primitive form of pitting (multiseriate scalariform) with parenchymatization of the xylem. The remaining species of the genus have not developed xylem parenchyma but have developed circular bordered pitting. The relationships of the genus to other genera in the Coenopteridales remain obscure.     

PROCAMBIUM VS. CAMBIUM AND PROTOXYLEM VS. METAXYLEM IN POPULUS DELTOIDES SEEDLINGS

The concept of a procambium-cambium continuum was examined in Populus deltoides by following its development in serially sectioned bud and stem tissues. As in other species, the term cambium is used to refer to that part of the continuum associated with the formation of secondary vascular tissues; i.e., with secondary growth. However, that part of the continuum associated with the formation of primary vascular tissues is subdivided to facilitate interpretation of the consecutive stages of primary xylem differentiation. Thus, the procambium as envisioned by other authors is subdivided into procambium, initiating layer, and metacambium, all of which develop acropetally and in complete continuity. The procambium is derived from the residual meristem in the form of acropetally developing strands and traces. The initiating layer is represented by the first, tangentially separated, periclinal divisions that delineate the position of the prospective cambium. The metacambium is a later stage during which additional periclinally dividing cells unite the initiating layer into a tangentially continuous meristem within a trace bundle. After establishment of the initiating layer, the procambial trace is completely phloem dominated. Protoxylem differentiation begins in an originating center at the base of the leaf primordium and it progresses basipetally to form the protoxylem pole. Cells of the initiating layer do not contribute to the formation of either protoxylem or protophloem. However, those cells of the initiating layer directly opposite the protoxylem pole divide precociously and later differentiate to metaxylem, thus forming a radial file of protoxylem-metaxylem elements. Protoxylem elements of lateral traces are longitudinally continuous with the protoxylem of their parent traces, whereas those of a central trace are longitudinally continuous with the metaxylem of its parent trace. Metaxylem is formed later than protoxylem and it is derived from the metacambium. Metaxylem does not form a continuous system with protoxylem of the same trace because of the different temporal and spatial origins of the two kinds of xylem. Rather, metaxylem is longitudinally continuous with secondary xylem of older traces below. An attempt was made to determine the functional significance of the pattern of protoxylem and metaxylem differentiation in relation to primary and secondary plant development.     

Arrangement of cortical microtubules during formation of bordered pit in the tracheids ofTaxus

Summary The arrangement of cortical microtubules during the development of the secondary wall and bordered pits in the tracheids ofTaxus was examined by immunofluorescence and electron microscopy. The cambial region of radial longitudinal sections of developing young shoots (2–3 years old) contains cells at various stages of differentiation from cambial cells to tracheids. At the early stage of formation of bordered pits, circular bands of microtubules were seen to be associated with the inner edge of the border of the developing pit. In other regions than the pit secondary wall of uniform thickness was laid down, and obliquely oriented cortical microtubules ran parallel to one another. These cortical microtubules also covered the surface of the border of the developing pit on the side facing the center of the cell. As the border of the pit developed, a circular band of MTs remained associated with the inner edge of border, suggesting that the MTs were involved in the formation of the rim of the bordered pit, extending the initial border thickening, which consisted of concentrically oriented cellulose microfibrils. After completion of the formation of the bordered pit, helical thickenings became apparent. The obliquely oriented microtubules were organized in bands parallel to one another, being superimposed on the helical thickenings. The involvement of MTs in the formation of bordered pits and helical thickening is discussed.     

SEM studies on vessels in ferns. 17. Psilotaceae

Perforation plates are reported in aerial and subaerial axes of Psilotum nudum and in aerial axes of Tmesipteris obliqua. In Psilotum, both perforations lacking pit membranes and perforations with pit membrane remnants were observed. Perforation plates in Psilotum may consist wholly of one type or the other. In Tmespteris, perforations have threadlike pit membranes or consist of porose pit membranes. Wide perforations alternating with narrow pits, a conformation observed in various ferns, were observed in Psilotum (subaerial axes). In Psilotum, perforations are more common in metaxylem than in protoxylem; perforations in protoxylem consist of primary wall areas containing small circular porosities or relatively large circular to oval perforations. There are no modifications in the secondary wall framework of protoxylem or metaxylem in Psilotum or Tmesipteris that would permit one to distinguish presence of perforations or perforation plates with light microscopy, and scanning electron microscopy (SEM) is required for demonstration of porose walls or perforations. The tracheary elements of the Psilotaceae studied have no features not also observed in other ferns with SEM.     

A new Bennettitalean genus from the Middle Jurassic of the Mikhailovskii Rudnik locality (Kursk Region,Russia)

Bennettitalean leaves with unusual morphology were found in the Middle Jurassic (Bathonian) deposits of Mikhailovskii Rudnik locality (Kursk Region, European Russia). This morphotype derived on the basis of Pterophyllum-like pinnate leaves by coherent coalescence of segments, through which superficial similarity of leaves of the new genus to Nilssoniopteris Nath., possessing entire leaf blade, is achieved. Each of coalescent segments was completely isolated from neighboring segments by massive cuticular wall. This character distinguishes leaves of the new genus from coherent leaves, known in other gymnosperms. The unique preservation type characteristic of plant remains of the Mikhailovskii Rudnik locality, allowed us to study some details of anatomical structure of leaf segments in the new genus. In particular, it was established, that palisade mesophyll in segments was poorly developed, and metaxylem in vascular bundles comprises tracheids with scalariform to circular bordered pits.     

Examination of morphological changes in the first formed protoxylem in Arabidopsis seedlings

We examined morphological changes in the first-formed protoxylem vessels in Arabidopsis seedlings. Between 2.5 and 8 days after imbibition, mean hypocotyl and root length increased 1.52 and 23.3 times, respectively. In the 2.5-day-old seedlings, two continuous protoxylem vessels were present in the hypocotyl-root axis. In the 8-day-old upper hypocotyls, six protoxylem vessels were observed, and in the lower hypocotyls, four protoxylem vessels and one or two metaxylem vessels were observed. In the 8-day-old roots, there were two protoxylem vessels and two or three metaxylem vessels. Two protoxylem vessels in the hypocotyls connected to two metaxylem vessels in the roots of 8-day-old seedlings. At the 0.3-mm part below the hypocotyl-root boundary, the mean intervals of neighboring annular secondary wall thickenings in protoxylem vessels in 8-day-old roots were 12.9% larger than those in 2.5-day-old roots. In more apical parts of 8-day-old roots, the mean intervals fluctuated between 1.71 and 2.29 microm. In 8-day-old seedlings, metaxylem vessels were formed between 0.4 mm above the hypocotyl-root boundary and 17 mm below the boundary. The intervals in these regions were not extended so much as protoxylem vessels were collapsed. The first-formed protoxylem vessels presumably retain their water-conductive function after metaxylem formation.     

梣属11种植物次生木质部附物纹孔的电镜观测

应用扫描电镜对梣属11种植物次生木质部导管附物纹孔的分布和形态进行了详细的观察,应用Carnoy2.0软件和扫描电子显微镜采集的照片,测定了附物纹孔丰富度指标和纹孔数量特征指标。电镜观察表明,梣属11种研究植物次生木质部导管分子侧壁附物纹孔的分布和形态变化较大,附物纹孔丰富度指标的统计描述进一步证实附物纹孔的分布变化大;3个附物纹孔丰富度指标分别与管间具缘纹孔数量特征指标的逐步回归分析表明,导管侧壁附物纹孔丰富度2个指标,即导管外纹孔口附物频率与导管纹孔腔附物频率随纹孔口面积百分比的增大而增大,推测梣属植物附物纹孔丰富度与纹孔几何构造及数量特征有关。研究认为,附物纹孔在梣属植物稳定存在,是可界定梣属植物的木材解剖性状。     

扁圆封印木（相似种）茎干的解剖特征

贵州省水城矿区晚二叠世煤核中扁圆封印木（相似种Ｓｉｇｉｌｌａｒｉａ ｃｆ．ｂｒａｒｄｉｉＢｒｏｎｇｎ．）茎干的主要解剖特征如下：管状中柱,具多边形薄壁细胞组成的髓。初生木质部成环带状,外缘呈规则的齿槽状,向心式发育。次生木质部显束状特征,横切面管胞为方圆至长方形,纵切面为梯状壁增厚,并具流苏纹。射线１—２列细胞宽,数个至十余个细胞高。叶迹起源于初生木质部外缘的槽中,中始式,但以向心发育为主。     

Myosin,microtubules, and microfilaments: co-operation between cytoskeletal components during cambial cell division and secondary vascular differentiation in trees

The immunolocalisation of unconventional myosin VIII ('myosin') in the cells of the secondary vascular tissues of angiosperm (Populus tremula L. x P. tremuloides Michx. and Aesculus hippocastanum L.) and gymnosperm (Pinus pinea L.) trees is described for the first time and related to other cytoskeletal elements, as well as to callose. Both myosin and callose are located at the cell plate in dividing cambial cells, whereas actin microfilaments are found alongside the cell plate; actin and tubulin are both associated with the phragmoplast. Myosin and callose also localise to the plasmodesmata-rich pit fields in the walls of living cells, which are particularly abundant within the common walls between ray cells and between ray cells and axial parenchyma cells in the phloem and xylem. In those xylem ray cells that contact developing vessel elements and tracheids, myosin, tubulin, actin and callose are localised at the periphery of developing contact and cross-field pits; the respective antibodies also highlight the bordered pits between vessels and between tracheids. The aperture of the bordered pits, whose diameter diminishes as the over-arching border of these pits develops, also houses myosin, actin and tubulin. Myosin, actin and callose are also found together around the sieve pores of sieve elements and sieve cells. We suggest that an acto-myosin contractile system (a 'plant muscle') is present at the cell plate, the sieve pores, the plasmodesmata within the walls of long-lived parenchyma cells, and at the apertures of bordered pits during their development.     

In vitro induction of secondary xylem-like tracheary elements in calli of hybrid poplar (Populus sieboldii × P. grandidentata)

The formation of tracheary elements was induced in calli derived from petioles of hybrid poplar (Populus sieboldii × P. grandidentata) after 10 days of culture on medium that lacked auxin but contained 1 μM brassinolide. Some differentiated cells formed broad regions of cell walls and bordered pits, which are typical features of tracheary elements of secondary xylem. Other differentiated cells resembled tracheary elements of primary xylem, with spiral or reticulate thickening of cell walls. The tracheary elements that developed in calli were formed within cell clusters. This induction system provides a new model for studies of the mechanism of differentiation of secondary xylem cells in vitro.     

WOOD ANATOMY OF BUBBIA (WINTERACEAE), WITH COMMENTS ON ORIGIN OF VESSELS IN DICOTYLEDONS

Quantitative and qualitative features of wood anatomy are reported for ten collections of seven species of Bubbia. Variations on the basic plan for Winteraceae can be interpreted in terms of taxonomic and ecological distinctions. Tracheid length is correlated with plant size and habit: tracheids are shortest in shrubs. Tracheid wall thickness and ray cell wall thickness distinguish species. Ray cell procumbency and multiseriate ray width increase with age. Growth rings occur only in a species from stream margins. SEM studies reveal absence of a warty layer within tracheids. Helical thickenings are absent. Presence of these two features in Pseudowintera may be correlated with the cool temperate habitats of that genus. Overlap areas of tracheids in Bubbia show various degrees of scalariform pitting, ranging from none (B. semecarpoides) to abundant presence (B. balansae). Perforation-like pits in tracheids of the latter prove, with SEM studies, to have pit membranes containing porosities less than 1 μm in diameter. Scalariform pitting on overlap areas is absent in earlier secondary xylem and increases during later secondary xylem. Scalariform lateral wall pitting can occur in abnormally wide tracheids formed after pauses in cambial activity. These facts show that primitive dicotyledon woods like those of Bubbia can activate genetic information for scalariform end wall patterns and lateral wall pitting such as primitive vessels show without the intervention of paedomorphosis. Paedomorphosis in dicotyledon woods is held still to apply only to special herbaceous and herblike growth forms, not to primarily woody plants. Progenesis (in xylem, loss of secondary xylem) is not held to be necessary to account for the scalariform patterns seen in tracheary elements of primitive dicotyledons. Reasons are given for rejection of the hypothesis that Winteraceae and other woody dicotyledons (Amborella, Sarcandra, Tetracentron, Trochodendron) are secondarily vesselless.     

Cell Wall Thickening of the Xylem Tracheary Elements in Different Zones of the Adventitious Root of Allium cepa L.

Cell wall thickness of the xylem tracheary elements was measuredin the proto- and metaxylem of the Allium cepa L. adventitiousroot. Measurements were taken in root fragments of known age(1, 3, 5, 7 and 9 d) located in either the basal or medio-apicalzone. Tracheary elements in the protoxylem matured within ashorter period of time than those in the metaxylem. Final cellwall thickness was greater in metaxylem than in protoxylem components.The cell wall thickening in the tracheary elements in both proto-and metaxylem was more rapid in the basal zone of the root thanin the medio-apical zone. Additionally, cell walls of the maturetracheary elements were thicker in the basal zone than in areasfurther from the bulb. Allium cepa, onion, root, cell wall, xylem maturation     

Structural analysis of K<Superscript>+</Superscript> dependence in <Emphasis Type="SmallCaps">l</Emphasis>-asparaginases from <Emphasis Type="Italic">Lotus japonicus</Emphasis>

Wood is composed of various types of cells and each type of cell has different structural and functional properties. However, the temporal and spatial diversities of cell wall components in the cell wall between different cell types are rarely understood. To extend our understanding of distributional diversities of cell wall components among cells, we investigated the immunolabeling of mannans (O-acetyl-galactoglucomannans, GGMs) and xylans (arabino-4-O-methylglucuronoxylans, AGXs) in ray cells and pits. The labeling of GGMs and AGXs was temporally different in ray cells. GGM labeling began to be detected in ray cells at early stages of S₁ formation in tracheids, whereas AGX labeling began to be detected in ray cells at the S₂ formation stage in tracheids. The occurrence of GGM and AGX labeling in ray cells was also temporally different from that of tracheids. AGX labeling began to be detected much later in ray cells than in tracheids. GGM labeling also began to be detected in ray cells either slightly earlier or later than in tracheids. In pits, GGM labeling was detected in bordered and cross-field pit membranes at early stages of pit formation, but not observed in mature pits, indicating that enzymes capable of GGM degradation may be involved in pit membrane formation. In contrast to GGMs, AGXs were not detected in pit membranes during the entire developmental process of bordered and cross-field pits. AGXs showed structural and depositional variations in pit borders depending on the developmental stage of bordered and cross-field pits.