Distinctive tracheid microstructure in stems of Victoria and Euryale (Nymphaeaceae)

Scanning electron microscopy (SEM) photographs of thick sections from liquid‐preserved stems of Victoria cruziana and Euryale ferox show accretions of coarse fibrils on pit membranes of tracheids. The first‐deposited fibrils are randomly orientated; on top of them (facing the tracheid lumina) are axially orientated coarse fibrils. The two systems are interconnected. Axially orientated fibrils were more extensively observed in Euryale than in Victoria and tips of fibrils in Euryale extend over the pit apertures onto secondary wall surfaces. Tracheid–parenchyma interfaces bear rudimentary coarse fibrils on the tracheid side. End walls of Victoria tracheids have highly porose pit membranes, thinner and less complex than those of the lateral intertracheid walls. The structures reported in Victoria and Euryale are consistent with those concurrently reported for stems of other Nymphaeaceae. Although also present in Cabombaceae, the coarse fibrils are otherwise not reported for stems of angiosperms and are not yet reported in roots of any species. Pit membrane remnants in perforation plates of various woody dicotyledons represent a nonhomologous phenomenon. The accretions of coarse fibrils in stem tracheids of Nymphaeaceae do not appear to enhance conduction, although they do contain porosities interconnecting tracheids. Removal of pit membrane remnants from perforation plates of primitive dicotyledon woods by hydrolysis does, on the contrary, suggest conduction enhancement. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 52–57.     

Wood anatomy of daphniphylla ceae: Ecological and phylogenetic considerations,review of pittosporalean families

Wood anatomy of 16 collections representing three species containing eight subspecies of the single genusDaphniphyllum is analyzed quantitatively and qualitatively.Daphniphyllum has vessels angular to roundish in transection, scalariform perforation plates, scalariform to opposite lateral wall vessel pitting, tracheids with fully bordered pits, heterocellular multiseriate and uniseriate rays, diffuse axial parenchyma and, in one taxon, chambered crystals in axial parenchyma cells. Growth rings, narrower vessels, and more numerous, vessels per square mm characterize taxa from cooler habitats. All of the taxa have highly mesomorphic woods. Comparisons are made between Daphniphyllaceae and the other families of Thorne's Pittosporales (Balanopaceae, Bruniaceae, Buxaceae, Byblidaceae, Geissolomataceae. Grubbiaceae, Myrothamnaceae, Pittosporaceae, Roridulaceae, and Tremandraceae). These families are most comparable to hamamelidoid or rosoid families; other similarities or relationships for these families may exist, but are less conspicuous or less close. The families cited may form a plexus, characterized by primitive xylary and other features, comparable to Annonales (Magnoliales) as products of an early radiation of dicotyledons.     

The earliest wood and its hydraulic properties documented in c. 407‐million‐year‐old fossils using synchrotron microtomography

We document xylem structure and hydraulic properties in the earliest woody plant A rmoricaphyton chateaupannense gen. nov. & sp. nov. based on c. 407‐million‐year‐old fossils from the Armorican Massif, western France. The plant was small, and the woody axes were narrow and permineralized in pyrite (FeS₂). We used standard palaeobotanical methods and employed propagation phase contrast X‐ray synchrotron microtomography (PPC‐SRμCT) to create three‐dimensional images of the wood and to evaluate its properties. The xylem comprised tracheids and rays, which developed from a cambium. Tracheids possessed an early extinct type of scalariform bordered pitting known as P‐type. Our observations indicate that wood evolved initially in plants of small stature that were members of Euphyllophytina, a clade that includes living seed plants, horsetails and ferns. Hydraulic properties were calculated from measurements taken from the PPC‐SRμCT images. The specific hydraulic conductivity of the xylem area was calculated as 8.7 kg m^?1 s^?1 and the mean cell thickness‐to‐span ratio (t/b)² of tracheids was 0.0372. The results show that the wood was suited to high conductive performance with low mechanical resistance to hydraulic tension. We argue that axis rigidity in the earliest woody plants initially evolved through the development of low‐density woods. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 423–437.     

Observations on the vegetative anatomy of Austrobaileya: habital, organographic and phylogenetic conclusions

For the single species of Austmbaileya (Austrobaileyaceae), quantitative and qualitative data are offered on the basis of a mature stem and a root of moderate diameter. Data available hitherto have been based on stems of small to moderate diameter, and roots have not previously been studied. Scanning electron microscope (SEM) photographs are utilized for roots, and show compound starch grains. Roots lack sclerenchyma but have relatively narrow vessels and abundant ray tissue. Recent phylogenies group Austrobaileyaceae with the woody families Illiciaceae, Schisandraceae, and Trimeniaceae (these four may be considered Illiciales), and somewhat less closely with the vesselless families Amborellaceae and Winteraceae and the aquatic families Cambombaceae and Nymphaeaceae. The vessel-bearing woody families above share vessels with scalariform perforation plates; bordered bars on plates; pit membrane remnants present in perforations; lateral wall pitting of vessels mostly alternate and opposite; tracheids and/or septate fibre-tracheids present; axial parenchyma vasicentric (sometimes abaxial); rays Heterogeneous Type I; ethereal oil cells present; stomata paracytic or variants of paracytic. Although comparisons between vessel-bearing and vesselless families must depend on fewer features, Amborellaceae and Winteraceae have no features incompatible with their inclusion in an expanded Illiciales.     

Wood anatomy of theSarraceniaceae; ecological and evolutionary implications

The wood of theSarraceniaceae has a considerable number of primitive features including scalariform perforation plates, long and oblique end walls, scalariform lateral wall pitting, solitary vessels, tracheids with scalariform pitting, and diffuse axial parenchyma. Vessel elements in the genusHeliamphora have the greatest number of primitive features, while vessel elements inDarlingtonia andSarracenia appear to have modifications relating to temperate climates. The wood anatomy suggests thatHeliamphora is growing in a habitat more similar to the original habitat for the family thanDarlingtonia andSarracenia. The wood of theSarraceniaceae is similar to the wood of theTheales.     

STRUCTURE OF TRACHEIDS IN THREE SPECIES OF LYCOPODIUM

The structure of tracheids in Lycopodium lucidulum, L. clavatum, and L. tristachyum was studied with the light microscope. Protoxylem development is at least sometimes and possibly always mesarch in indeterminate axes of all three species. Centrifugally formed protoxylem elements are reticulate and discontinuities in the secondary walls of these elements are sometimes conspicuously bordered. Wall thickenings of first formed protoxylem elements consist mainly of indirectly connected rings. Late centripetally formed protoxylem elements and transitional elements have a reticulate secondary wall pattern. The narrowest metaxylem elements have circular bordered pits while in wider metaxylem elements pits are bordered and may vary from circular to scalariform. Pitting is uniseriate to triseriate in tracheids of all three species, and intermittent tetraseriate pitting was occasionally observed in L. lucidulum. Crassulae occur in tracheids of the three species, and in L. clavatum an additional framework, probably representing thickened compound middle lamella, is also present. Pits often appear helically arranged, and in all three species pits are connected by thin areas in the secondary wall. Macrofibrils approximately 0.5 μ wide were observed in tracheids of the three species. In L. clavatum the arrangement of macro-fibrils was predominantly bidirectional.     

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。     

THE VASCULAR CAMBIUM AND XYLEM DEVELOPMENT IN HIBISCUS LASIOCARPUS

Cumbie, B. G. (U. Missouri, Columbia.) The vascular cambium and xylem development in Hibiscus lasiocarpus. Amer. Jour. Bot. 50(9): 944–951. Illus. 1963.—Circumferential growth of the vascular cambium, as determined primarily by an analysis of the secondary xylem, in Hibiscus lasiocarpus, an herbaceous dicotyledon, occurred through both radial and oblique anticlinal divisions. Divisions to produce segments were less frequent. Although the fusiform initials usually elongated somewhat between successive divisions, this accounted for very little increase in circumference of the cambium. A fusiform initial underwent a specific pattern of anticlinal divisions, determined primarily by its length, at the beginning of cambial activity. There was no loss of fusiform initials, except by ray formation. Most new rays originated only after considerable secondary xylem had been formed. The findings are discussed in relation to circumferential growth of the vascular cambium in woody dicotyledons.     

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.     

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.     

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.     

FLORA OF THE LOWER CRETACEOUS CEDAR MOUNTAIN FORMATION OF UTAH AND COLORADO. PART III: ICACINOXYLON PITTIENSE N. SP

Icacinoxylon pittiense, a new species of angiospermous wood from the Lower Cretaceous Cedar Mountain Formation of Utah is described and compared with similar fossil and modem woods. It is distinguished from other species of Icacinoxylon by its thick-walled fiber-tracheids with their walls making up at least 50% of the total diameter of the cells, conspicuous bordered pits with obliquely crossing extended apertures on both the tangential and radial walls of its fiber-tracheids, scalariform perforation plates with as few as four or greater than 30 bars, transitional opposite to scalariform pitting on its vessel walls, thick-walled ray cells, and distinct sheath or border cells in its rays. Icacinoxylon pittiense is the first species of this genus to be reported from Cretaceous sediments. This wood is of special interest because very few angiosperm woods have been reported from lower Cretaceous strata.     

RELATION OF DWARFMISTLETOE (ARCEUTHOBIUM) TO THE XYLEM TISSUE OF CONIFERS. II. EFFECT OF THE PARASITE ON THE XYLEM ANATOMY OF THE HOST12

Srivastava , L. M., and K. Esau , (U. California, Davis.) Relation of dwarfmistletoe (Arceuthobium) to the xylem tissue of conifers. II. Effect of the parasite on the xylem anatomy of the host. Amer. Jour. Bot. (48(3): 209–215. Illus. 1961.—The changes in the xylem anatomy induced by dwarfmistletoe infection were studied in 7 coniferous species. The most pronounced abnormalities are observed in the shape and size of the infected rays. Because of the presence of parasite tissue, the rays assume a hypertrophied appearance; moreover, they fuse to form large composite rays. The union of rays involves intrusive growth of ray cells and displacement of fusiform initials. Some division of fusiform initials also occurs. Rays may increase in number and they may contain more host cells than normal rays. Axial tracheids in infected host woods differ more or less strongly from those of noninfected woods. They may be shorter, wider, and more irregular in shape than the axial tracheids in healthy wood. The samples of xylem from infected pines had a larger number of resin canals than those from healthy trees. Resin canals were also found in infected Tsuga, which normally lacks these structures.     

Anatomical Characteristics of the Stem of Sigillaria cf. brardii from Coal Balls in Guizhou Province,China

The stem specimens of Sigillaria cf. brardii were collected from the coal balls of Upper Permian in Shuicheng Coal Mines in Guizhou Province. The main anatomical characteristics of Sigillaria cf. brardii are described as follows: The stem is siphonostelic, with pith composed entirely of polygonal parenchyma cells, there are secondary walls in some pith cell cavities these secondary walls show the characters of cell division. Surrounding the pith is the continuous cylindrical primary xylem which consists entirely of tracheids. The outermost, and part are the protoxylem elements show spiral secondary thickenings. In cross section, the outer edge of exarch primary xylem appears regularly sinuous, with trace of mesarch leaf originating from the furrows. The centripetal metaxylem is characterized by scalariform wall thickenings on the tracheids, and delicated strands of secondary wall materials extending between abjacent bars, these structures are called fimbris, or williamson striations, and are characteristic in lepidodendrids. The secondary xylem consists of tracheids and vascular rays. The tracheids, too, have scalariform wall thickenings and fimbris. The rays are one-to twocell width and several to more than ten cells in height.     

壳斗科次生木质部水分输导分子间连接通道结构的研究

壳斗科次生木质部水分运输聚合体及其连接通道结构,根据水分运输方向可分纵向和横向两大系统。水分纵向运输通道有导管分子间的穿孔,导管分子侧壁间的管间纹孔,导管分子与无穿孔管状分子间的侧壁纹孔,无穿孔管状分子与无穿孔管状分子间的侧壁纹孔。     

Study on the Convergent Passages of Water Conducting Elements in Secondary Xylem of Fagaceae

Structural variations in the convergent passages of water conducting elements were described in secondary xylem of 70 species of Fagaceae. Two types of water conducting systems were recognized on the ground of water-flow directions. The convergent passages of the longitudinal system mainly exhibited in the forms as perforations between the ends of Vessel elements, lateral wall pitting between the lateral walls of vessel elements, pitting be-tween vessel elements and imperforate tracheary elements, and pitting between imperforate tracheary elements. The convergent passages in the transverse system consisted of pitting between vessel elements and ray cells, pitting between imperforated tracheary elements and ray cells and simple pitting between ray cells. In addition, the roles of vasicentric tracheids and broad rays played in terms of conducting effectiveness and efficiency as proposed by Zimmermann were discussed.     

Single‐vessel flow measurements indicate scalariform perforation plates confer higher flow resistance than previously estimated

During vessel evolution in angiosperms, scalariform perforation plates with many slit‐like openings transformed into simple plates with a single circular opening. The transition is hypothesized to have resulted from selection for decreased hydraulic resistance. Previously, additional resistivity of scalariform plates was estimated to be small – generally 10% or less above lumen resistivity – based on numerical and physical models. Here, using the single‐vessel technique, we directly measured the hydraulic resistance of individual xylem vessels. The resistivity of simple‐plated lumens was not significantly different from the Hagen–Poiseuille (HP) prediction (+6 ± 3.3% mean deviation). In the 13 scalariform‐plated species measured, plate resistivity averaged 99 ± 13.7% higher than HP lumen resistivity. Scalariform species also showed higher resistivity than simple species at the whole vessel (+340%) and sapwood (+580%) levels. The strongest predictor of scalariform plate resistance was vessel diameter (r² = 0.84), followed by plate angle (r² = 0.60). An equation based on laminar flow through periodic slits predicted single‐vessel measurements reasonably well (r² = 0.79) and indicated that Baileyan trends in scalariform plate evolution maintain an approximate balance between lumen and plate resistances. In summary, we found scalariform plates of diverse morphology essentially double lumen flow resistance, impeding xylem flow much more than previously estimated.     

Fossil wood of Eristophyton sp. from the Carboniferous deposits of northern Russia (Arkhangelsk region)

Permineralised wood of Eristophyton sp. is first described from the Carboniferous deposits of the Arkhangelsk region, northern Russia. The specimens used in the study show scalariform thickening of the metaxylem tracheids both on radial and tangential walls. Eristophyton sp. indicates well preserved elements of secondary xylem: uni-, rarely biseriate xylem rays up to 15–16 cells high; uni-, multiseriate tracheid pitting only on radial walls; 1–8 contiguous cross-field pits and their inclined narrow apertures. A brief review and comparison with known anatomically preserved plants from the Lower Carboniferous of different localities of Scotland, France, USA and Poland is discussed.