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IAWA Bulletin Ns, Vol. 4 (2-3),1983 79 INTERVASCULAR PIT

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IAWA Bulletin Ns, Vol. 4 (2-3),1983 79 INTERVASCULAR PIT IAWA Bulletin n.s., Vol. 4 (2-3),1983 79 INTERVASCULAR PIT MEMBRANES IN ULMUS AND CELTIS NATIVE TO THE UNITED STATES· by E. A. Wheeler Department of Wood and Paper Science, North Carolina State University, Raleigh, North Carolina 27650, U.S.A. Summary Intervascular pit membranes in Ulmus ameri­ Studies to date have resulted in the generali­ cana L. resemble those of other dicotyledons as sation that the membranes in bordered pit pairs they are ofunifonn thickness, lack visible open­ of dicotyledons exhibit primary wall texture, ings, and have randomly arranged microfibrils. that is, they have randomly arranged micro­ In contrast, some of the intervascular pit mem­ fibrils, lack visible openings, and are of unifonn branes in U. alata Michx. , U. thomasii Sarg., thickness (Panshin & DeZeeuw, 1980; Schmid, Celtis laevigata Willd., C occidentalis L., and 1965). This generalisation is based on examina­ C reticulata Torr. have a central thickened re­ tion with the transmission electron microscope gion, a torus. Evidence from ultrathin sections of fewer than a dozen species of angiospenns. suggests such thickenings are fonned primarily In contrast, the intertracheary pit membranes in vascular tracheids. Microfibrils are clearly of conifers generally are differentiated into a visible in the membranes in the outer sapwood central thickened region, the torus, and periph­ of U. americana, U. alata, and C laevigata , while erally, a margo with radiating strands of micro­ in the inner sapwood, encrustives are present fibrils and often openings readily seen with the on some membranes. There is variation in how electron microscope. There exists variation in infilled and thickened the heartwood mem­ how well defined the torus is and how large the branes are. The variability in pit membrane ap­ gaps, if any, in the margo are, but Bauch et al. pearance in these U. S. species of Ulmaceae fur­ (1972) found that of the 47 conifer genera ther indicates that the structure of intervascular they examined only Thuja and Thujopsis con­ pit membranes is more diverse than previously tained pit membranes without obvious central recognised. thickening. They referred to such pits as being of the 'hardwood type'. The pit membranes of Introduction the gymnospennous groupsCycadales, Gnetales In wood, liquid primarily moves from vessel and Ginkgoales also lack torus-margo differen­ to vessel or from tracheid to tracheid through tiation (Liese, 1965). bordered pit pairs. Consequently, pit structure, There have been three recent reports of pit particularly pit membrane structure, affects membranes with a central thickened area in water transport in the living tree, the drying of angiospenns. Parameswaran and Liese (1981) lumber and its treatment with preservatives, found torus-like thickenings in the pit mem­ and has been studied by both botanists and branes of interfibre pits in Prunus and Pyrus wood technologists (e.g. Bailey, 1913; Cote, (Rosaceae). Ohtani and Ishida (1978) described 1958; Schmid, 1965; Bauch et aI., 1972; Bon­ the occurrence of intervessel pit membranes ner & Thomas, 1972; Meylan & Butterfield, with a torus in three species of Daphne and 1982). The accepted definition of a pit is 'a four species of Osman thus (Oleaceae). Parame­ recess in the secondary wall of a cell, together swaran and Gomes (1981) described a torus-like with its external closing membrane' (IAWA, region in intervessel pits of Ligustrum, another 1964). Generally, two complementary pits are member of the Oleaceae. fonned in adjacent cells resulting in a pit pair. During the course of a study whose purpose Bordered pits are characterised by the second­ is to characterise the fine structure of sapwood ary wall overarching the pit membrane. The and heartwood of some dicotyledonous trees primary walls of the two adjacent cells and the of the southeastern United States (Thomas, middle lamella are considered to fonn the basic 1976; Wheeler & Thomas, 1981; Wheeler, 1981, structure of the pit membrane. 1982), pit membranes with a central thickened • Paper No. 8634 of the JourfialSeries of the Agricultural Research Service, Raleigh, North Caro­ lina, U.S.A. 80 IAWA Bulletin n.s., Vol. 4 (2-3),1983 area were observed in Ulmus alata Michx. and with platinum at an angle of approximately 20° Celtis laevigata WiIld. (Ulmaceae). This paper in a high vacuum evaporator prior to examina­ describes the intervascular pit membranes from tion with a Siemens Elmiskop electron micro­ the sapwood and heartwood of U. americana, scope. U. alata, and C laevigata and further documents To ascertain if torus-like thickenings were of greater variability in dicotyledonous pit mem­ more general occurrence within the Ulmaceae branes than previously recognised. Intervascular direct carbon replicas (DCR's) were prepared pit membranes from the sapwood of Ulmus from sapwood samples ofU. S. Celtis and Ulmus thomasii, Celtis occidentalis, and C reticulata available in the David A. Kribs Collection at are also described. N.C. State University. All samples are herba­ There are six tree species of Ulmus native to rium vouchered (US, A) and were collected as the United States, all naturally restricted to the part of Project I of the State University of New eastern United States. Of the six species of Cel­ York, Syracuse (xylarium abbreviation: BWCw). tis naturally occurring in the United States, on­ The goal of Project I was to collect samples of ly C laevigata and C occidentalis of the eastern all tree species native to the U. S. and each sam­ U. S. frequently become large enough to pro­ ple is accompanied by data on the site, tree size duce marketable timber (Elias, 1980). Woods and vigour. Specimens with sapwood are Ulmus of the Ulmaceae native to the U. S. are described americana: BWCw 8014, U. thomasii: BWCw as distinctive as they are ring-porous with the 8024, Celtis laevigata var. texana: BWCw 8690, late wood pores arranged in wavy tangential C occidentalis: BWCw 8159, C reticulata (na­ (ulmiform) bands. Vascular tracheids are asso­ tive to the western U. S.): BWCw 8580, 8667. ciated with the vessel elements. This cell type is As examination of ultrathin sections indi­ described as 'an imperforate cell resembling cated the torus was formed primarily in vascu­ in form and position a small vessel member' lar tracheids, samples of all species listed above (lAW A, 1964).' CarIquist (1961) refers to vas­ were macerated to see if there was a correlation cular tracheids as 'degenerate vessel elements'. between frequency of vascular tracheids and Details of the wood anatomy of tree species of presence of a torus. Samples were macerated Ulmus and Celtis native to the U.S. are discus­ by placing them in a solution of equal parts of sed in a separate paper (Wheeler, ms. in prep.). hydrogen peroxide and glacial acetic acid and warming them for 2-3 days in a 50°C oven. Materials and Methods Wood samples were obtained from the dia­ Observations meter breast height region of one mature tree Ulmus americana L. (American Elm) - All each of Ulmus americana L., U. alata Michx., intervascular pit membranes in the outer sap­ and Celtis laevigata WiIld. felled specifically for wood had randomly arranged micro fibrils, no this study. Direct carbon replicas (Cote et aI., visible openings (Fig. I) and were of uniform 1964) of tangential and radial surfaces of (I) thickness (Fig. 2). In air-dried preparations the the outermost two growth rings-the area in microfibrils appeared somewhat coarser than in which most water conduction in the standing the solvent-dried preparations. At times, the tree would occur; (2) inner sapwood-the area membrane had aspirated and the imprint of the with living parenchyma cells (as determined by aperture was noticeable. staining for starch with 12 KI), but with little or In the inner sapwood (ring 6) most mem­ no water conduction; (3) heartwood just to the branes looked as they did in the outer sapwood. inside of the sapwood-heartwood boundary; and However, encrusting materials had begun to ac­ (4) heartwood approximately one inch from cumulate in some; microfibrils were completely the pith. In U. alata the heartwood zone was obscured or deposits were confined to a central not large, so no distinction was made between area beneath the aperture (Fig. 3). inner and outer heartwood. Prior to replication, In the outer heartwood, a few un encrusted the wood was either air-dried or solvent-dried pit membranes with microfibrils as distinct as by being placed for one week in methanol fol­ in the sapwood were observed. Others were lowed by one week in acetone in a soxhlet ex­ completely coated by deposits and there often tracting apparatus prior to drying. Solvent dry­ appeared to be a concentration of material at ing prevents displacement of the pit membrane the position corresponding to the pit aperture to one side of the pit chamber as would occur (Fig. 4). In the inner heartwood, all membranes during air-drying (Thomas & Nicholas, 1966). had encrustives to some degree. The outline of Ultrathin sections were prepared from never­ the aperture was apparent in both solvent-dried dried, air-dried, and solvent-dried wood embed­ and air-dried samples. The appearance of the ded in a mixture of n-butyl and methylmetha­ encfustives on some membranes (Fig. 4, and crylate (75: 25). These sections were shadowed see also Figs. 9 & 17) indicates that extractives .
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