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Cedrus, Keteleeria, Pseudolarix, and Pseudo- Key Words: Abies, Larix

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Cedrus, Keteleeria, Pseudolarix, and Pseudo- Key Words: Abies, Larix IAWA Journal, Vol. 15 (4), 1994: 399-406 FUSIFORM PARENCHYMA CELLS IN THE YOUNG WOOD OF PINACEAE, AND THEIR DISTINCTION FROM MARGINAL PARENCHYMA by Shuichi Noshiro and Tomoyuki Fujii Wood Anatomy Laboratory, Forestry and Forest Products Research Institute, Tsukuba Norin, P. O. Box 16, Ibaraki 305, Japan Summary Fusiform parenchyma cells found in sev­ ray tracheids, and the proportion of ray tra­ eral genera of Pinaceae are described and cheid pit border types. Recently, Anagnost compared with marginal parenchyma. Fusi­ et al. (1994) restudied all these features using form parenchyma cells are mostly fusiform world-wide sam pIes of Picea and Larix, and in shape, with occasional smooth horizontal confirmed the results of Bartholin (1979) that walls. They form discontinuous tangential ray tracheid pit border types were the most bands in complete or incomplete circ1es in the reliable characteristic. However, because of innermost growth rings of Larix, Abies, and restricted growth in branches, it was difficult Tsuga. Fusiform parenchyma always con­ to use these stemwood characteristics in iden­ tains resinous material, and is more conspic­ tifying branchwoods. uous in branchwoods than in stern woods. Modern branchwood materials gathered Marginal parenchyma cells were observed in for comparative purposes, however, revealed Cedrus, Keteleeria, Pseudolarix, and Pseudo­ the conspicuous occurrence of resinous paren­ tsuga as weIl as in Larix, Abies, and Tsuga, chyma cells in the inner growth rings of Larix and very rarely in Picea. Marginal paren­ and their absence in Picea. Occurrence of chyma cells are scattered along growth ring resinous parenchyma ceHs was also observed boundaries. They are always in strands with in the Quaternary branchwoods, and was used nodular horizontal walls with conspicuous to distinguish Larix branchwoods from Picea simple pits. Cell wall structure of these two branchwoods. Resinous parenchyma cells in types of parenchyma differs in the intensity Larix were fusiform, and differed from strand of the birefringence of the secondary walls. parenchyma occurring along growth ring Fusiform parenchyma cells are distinct from boundaries of several Pinaceous genera. We marginal parenchyma with which they were observed fusiform parenchyma in the young previously confused, and should be regarded stemwood of Larixand branchwood of Abies as a new component of coniferous wood. and Tsuga (also Pinaceae); fusiform parenchy­ Key words: Abies, Larix, Picea, Tsuga, Pina- ma was most conspicuous in Larix branch­ ceae, fusiform parenchyma cells, young wood. Budkevich (1956, 1961) did a com­ wood, marginal parenchyma. prehensive study of the wood anatomy of Pinaceae, and described the presence ofaxial Introduction parenchyma in the latewood of Abies, Cedrus, While identifying Quaternary fossil branch­ Keteleeria, Larix, Pseudolarix, Pseudotsuga, woods, the first author encountered difficulty Tsuga and Ducampopinus (most taxonomists in distinguishing between branchwood of inc1ude this genus in Pinus, e.g. Farjon 1984; Larix and Picea. Stemwoods of these two Mabberley 1987), and absence ofaxial paren­ genera have been distinguished by transition chyma from Cathaya, Picea and Pinus. He from earlywood to latewood, pit seriation of distinguished axial parenchyma with nodular longitudinal tracheids, number of epithelial end walls from axial parenchyma with smooth cells surrounding horizontal resin canals, end walls, and described parenchyma dis­ presence or absence of spiral thickenings in tribution in Abies, Pseudotsuga, Larix, and Downloaded from Brill.com09/25/2021 12:26:19PM via free access 400 IAWA Journal, Vol. 15 (4), 1994 Table 1. Occurrence of fusiform and marginal parenchyma in the secondary xylem of Pinaceae. Species Specimen fusifo11ll parenchyma marginal nurnber parenchyrna growth ring Abies firma Sieb. et Zucc. branch 1WTw-12974 + 2-3 ± do branch 1WTw-12975 + 2-6 do branch 1WTw-12976 ± 2 do stern 1WTw-433 do stern TWfw-14714 ++ 2-3 Abies sachalinensis (Fr. Schrn.) Masters branch 1WTw-5694 ++ 1-6 ± do branch 1WTw-5695 ++ 1-7 ± do branch 1WTw-5695 ++ 1-7 ± Cedrus deodara (Roxb.) G. Don branch 1WTw-12963 ++ do branch 1WTw-12964 + do branch 1WTw-12965 ++ Keteleeria davidiana (Bertr.) Beissn. branch 1WTw-12962 ++ do banch 1WTw-12969 ++ do branch 1WTw-12970 ++ Larix gmelinii (Rupr.) Rupr. var. japonica (Regel) Pilger branch 1WTw-2906 ++ 1-3 ++ do branch 1WTw-2913 ++ 1-3 ++ do branch 1WTw-2914 ++ 2-3 ++ Larix kaempferi (Lamb.) Carr. branch 1WTw-2905 ++ 1-3 ++ do branch 1WTw-2911 ++ 1-4 ++ do branch 1WTw-2912 ++ 1-6 ++ do stern K-l60 + 1-2 ++ do stern K-l84 + 2-3 + do stern Sn-l27 + 2-3 + do stern Sn-37 + 2 ++ do stern Sn-48 + 4-6 + Picea glehnii (Fr. Schrn.) Masters branch 1WTw-3903 do branch 1WTw-2907 do branch 1WTw-2908 Piceajezoensis (Sieb. et Zucc.) Carr. branch 1WTw-2904 do branch 1WTw-2909 do branch 1WTw-2910 Picea maximowiczii Regel ex Masters stern 1WTw-835 ± do stern Kan-9135 Pinus densiflora Sieb. et Zucc. branch 1WTw-12966 do branch 1WTw-12967 do branch 1WTw-12968 Pseudolarix amabilis (Nelson) Rehder branch 1WTw-12977 ++ Pseudotsuga japonica (Shirasawa) Beissn. branch 1WTw-12971 ++ do branch 1WTw-12972 ++ do branch 1WTw-12973 ++ Tsuga canadensis (L.) Carr. branch 1WTw-6790 ++ 3-4 ++ do branch 1WTw-6791 + 3-4 ++ Tsuga diversifolia (Maxim.) Masters branch 1WTw-6795 ++ 1-4 ++ do branch 1WTw-6796 ++ da branch 1WTw-6797 ++ do branch 1WTw-6798 + 3-4 ++ do stern 1WTw-593 + 8 + -: not observed; ±: less than ten per specirnen; +: incornplete circlc; ++: cornplete circle. Downloaded from Brill.com09/25/2021 12:26:19PM via free access Noshiro & Fujii - Fusiform parenchyma in Pinaceae 401 Cedrus. He also noted a more marked occur­ ture of parenchyma cells was observed both rence ofaxial parenchyma in branchwoods between cross nicols of a polarised light mi­ than in stemwoods. However, he did not croscope and with a phase contrast micro­ mention fusiform parenchyma cells. Our ob­ scope on thin transverse and tangential sec­ servations indicate a distinction between fusi­ tions (0.5 microns) of branchwoods and form parenchyma with rare smooth horizon­ stemwoods of Larix that were prepared using tal walls and marginal stranded parenchyma epoxy-embedding. with nodular end walls. Fusiform parenchy­ ma cells in secondary xylem are so far known to occur in Ephedra (Esau 1964) and several Results dicotyledons (panshin & De Zeeuw 1980; Fusiform parenchyma cells occur in Abies IAW A Committee 1989), but not in conifers firma, Abies sachalinensis, Larix kaempferi, (Phillips 1948; Greguss 1955; Panshin & De Larix glehnii, Tsuga canadensis, and Tsuga Zeeuw 1980). diversifolia, and were not observed in the In this paper we describe the morphology other six genera studied (Table 1). They are and distribution of fusiform parenchyma in mostly found in the latter half of the earlywood Pinaceae, and compare it to marginal stranded or in the latewood, and are occasionally scat­ parenchyma. tered throughout the innermost growth rings (Figs. 1, 6, 9). They usually form irregular, Materials and Methods discontinuous tangential bands in complete or Branchwoods and stemwoods of 14 spe­ partial circles within growth rings, mostly cies of nine genera of Pinaceae (47 sampies) occurring as single cells. These cells always were studied (Table 1). Sampies were collect­ contain resinous material, and can be easily ed from plantations and natural habitats in recognised in transverse sections (e.g., Figs. Japan. We were unable to collect materials of 2, 3). The diameters of these parenchyma Cathaya and Nothotsuga. Nomenclature fol­ cells are equal to those of the surrounding lows that of Farjon (1984, 1990). Specimens tracheids, or slightly sm aller (Fig. 2), and were sectioned on a sliding microtome for their length exceeds 1 mm (Fig. 7). They have light microscopy, including pith and inner­ sparse simple pits on radial walls, and lack most rings, and were double-stained with pits on tangential walls (Figs. 4, 8). They are Safranin and Gentian Violet. Cell wall struc- basically fusiform in shape without subdivi- -) Figs. 1-5. Fusiform and marginal parenchyma in the branchwood of Larix kaempferi. - 1: TS x 20; fusiform parenchyma making complete circles in the innermost rings. - 2: RLS x 200; mar­ ginal parenchyma cells in astrand (m), and a fusiform one with rcsinous content (f). - 3: TS x 400; fusiform parenchyma cens with resinous content, and a marginal one with simple pits on a hori­ zontal wall (arrow). - 4: RLS x 400; sparse simple pits on a radial wall of a fusiform parenchy­ ma cell. - 5: TLS x 400; nodular horizontal walls (arrows) and simple pits on tangential walls of a marginal parenchyma cello - Scale bar = 200 11m in Fig. 1, and 50 11m in Figs. 2-4. -) -) Figs. 6-8. Fusiform parenchyma in the branchwood of Abies sachalinensis. - 6: TS x 40; fusi­ form parenchyma in the latewood nearly parallel to growth ring boundaries. - 7: RLS x 100; fusiform parenchyma cells (f) more than 1 mm long. - 8: TLS x 400; a smooth horizontal wall in fusiform parenchyma and absence of pits on tangential walls. - Fig. 9. Fusiform parenchyma in the branchwood of Tsuga diversifolia. TS x 40. - Figs. 10 & 11. Marginal parenchyma in the branchwood of Pseudolarix amabilis. - 10: TS x 200; marginal parenchyma cells (arrows) on growth ring boundaries.- 11: TLS x 200; nodular horizontal walls and simple pits on tangential and radial walls. - Sc ale bar = 100 11m in Figs. 6, 7 & 9, and 50 11m in Figs. 8, 10 & 11. Downloaded from Brill.com09/25/2021 12:26:19PM via free access 402 IAWA Journal, Vol. 15 (4), 1994 Downloaded from Brill.com09/25/2021 12:26:19PM via free access Noshiro & Fujii - Fusiform parenchyma in Pinaceae 403 Downloaded from Brill.com09/25/2021 12:26:19PM via free access 404 IAWA Journal, Vol. 15 (4), 1994 Figs. 12 & 13. Cell wall structure of fusifollTI and marginal parenchyma in branchwood of Larix kaempferi. - 12: Phase-contrast micrograph. - 13: Polarised light micrograph showing fusifollTI (f) and marginal (m) parenchyma. si on (Fig.
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