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Development of Vascular Cambium and Compression Wood Formation in the Shoot of Young Spruce (Picea Jezoensis V Ar

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Development of Vascular Cambium and Compression Wood Formation in the Shoot of Young Spruce (Picea Jezoensis V Ar IAWA Bulletin n.s., Vol. 7 (1),1986 21 DEVELOPMENT OF VASCULAR CAMBIUM AND COMPRESSION WOOD FORMATION IN THE SHOOT OF YOUNG SPRUCE (PICEA JEZOENSIS V AR. HONDOENSIS) by Nobuo Yoshizawa, Yujiro Tanaka and Toshinaga Idei Faculty of Forestry, Utsunomiya University, Utsunomiya, Japan Summary In the course of the righting movement in However, little is known on the development young spruce trees (Pieea jezoensis Carr. var. of the cambium cylinder associated with the hondoensis Rehd.) inclined at 45°, the occur­ righting movement of the current shoot. Saiki rence of compression wood associated with the and Tachida (1973) reported in their observa­ development of vascular cambium in the shoot tions of the current shoots of Pinus thunbergii was observed. In shoots, the recovery first took which exhibits rapid righting movements that place at the mid point, a few days after inclina­ no compression wood cells occur within a spe­ tion. The observations of serial cross sections cific distance of the shoot apex. A correlation taken from the apex downward revealed no ap­ between the development of vascular cambium preciable difference in the development of the and that of compression wood within the in­ procambium-cambium continuum between clined current shoots is of great interest. This the upper- and underside of the shoot. The for­ paper is focused on the response of xylem tis­ mation and structure of primary tracheary ele­ sues to the stimulus of inclination in associa­ ments were similar, irrespective of the site of tion with the early development of the cambial the procambium in the shoot. No compression system. wood cells occurred before the vascular cam­ bium cylinder was complete. The stimulus of Materials and Methods compression wood formation is received only Materials in this experiment were taken from by the differentiating secondary xylem tissues the current shoots of 5 years old spruce trees derived from the cambium cylinder. (Pieea jezoensis Carr. var. hondoensis Rehd.), Key words: Apex, cell division, righting move­ which were inclined at 45° in late May, and the ment, secondary xylem, tracheary elements. righting movement in the current shoots fol­ lowing inclination was monitored using photo­ Introduction graphic records taken periodically. The process of cell differentiation begins The whole current shoots were collected in with the formation of new cells beneath the early October, and were fixed with FAA after shoot apex (Foster, 1938; Cross, 1943; Eames separation into several pieces. They were em­ & Mac Daniels, 1947). Below this portion, as bedded in paraffin by ordinary procedure, and growth and development proceed, the vascular serial cross sections of IS J-Lm thick were cu t at cambium system becomes established in a appropriate intervals from the apex to the base shoot (Esau, 1943, 1965; Larson, 1982). Con­ of the shoot by using a rotary microtome. siderable controversy still exists in the literature These cross sections were double stained with as to when and where the cambium state begins safranin and light green, and observed with an and how this state can be identified. More in­ ordinary light microscope. Longitudinal radial formation still should be accumulated concern­ sections at several different heights along the ing the development of the vascular cambium shoot also were observed with a Nomarski dif­ system. ferential interference microscope. On the other hand, recovery of the normal vertical position in tilted young tree stems takes Results and Discussion place at the top first and gradually proceeds In the course of the righting movement in downward (Westing, 1961; Archer & Wilson, young trees inclined at 45°, the recovery of the 1973; Fukazawa, 1973). The first recovery stems initiated in the current shoot, and gradu­ seems to occur at the middle portion of the ally proceeded to the bottom thereafter, as in­ current shoots (Yoshizawa et aI., in press). This dicated by earlier observations (Westing, 1961; fact interestingly suggests a certain relationship Archer & Wilson, 1973; Fukazawa, 1973). The between the development of vascular cambium first recovery took place at the middle portion and the occurrence of compression wood in the of the current shoots shortly after inclination shoot. (Fig. I). The righting of the shoot proceeded Downloaded from Brill.com09/27/2021 08:41:26AM via free access 22 IAWA Bulletin n.s., Vol. 7 (1),1986 Fig. I. Righting movement in the shoot of Picea jezoensis var. hondoensis. - a: immediately after inclination; b: three days after inclination. Vigorous righting is found in the middle part of the current shoot (arrowhead). rapidly from the middle area of the shoot to morphological changes in the resin canal shape. the base within a few days. This phenomenon With the development of the procambium is of interest in relation to the occurrence of downward, the number of radial cell files in compression wood cells associated with the de­ each procambial bundle gradually increased. velopment of vascular cambium in shoots. Meanwhile, endarch development of primary Within the portion 0.2-0.5 em beneath the xylem, from protoxylem of spiral tracheids to apex, procambia consisting of cells with smaller metaxylem of pitted tracheids, proceeded (Fig. diameter appeared first in the periphery of the 5). However, the development and structure of central ground meristem, or future pith (Fig. 2). the primary tracheary elemen ts were similar ir­ Meanwhile, cell divisions in the procambium respective of the si te of the proeam bium which were initiated randomly accompanying proto­ produces such cells. It seems to be a general xylem formation in an endarch manner(Fig. 3). feature of primary traeheary elements that the In general, the procambia seem to show irregu­ primary wall remains unlignified (Kuo et aI., lar eell divisions with no tangential continuity 1974; O'Brien, 1981). No remarkable differ­ (Esau, 1977; O'Brien, 198 I; Larson, 1982). No ence in the intensity of lignification of the sec­ marked anatomical differences between the up­ ondary wall of the metaxylem cells, as well as per- and underside of the shoot could be ob­ in the number of primary xylem cells, was ob­ served in this part. The morphological differen­ served between the upper- and underside of ces between the upper- and underside were first the shoot. In the case of Picea jezoensis var. observed in the cortex (Fig. 4). As far down to hondoensis, the number of radially aligned pri­ the part 0.6 em below the apex, resin canals in mary xylem cells in each file was generally kept the cortex had irregular shapes irrespective of within a range of 2-5, by the gradual collapse upper- and underside of the shoot. However, of protoxylem cells inside and a corresponding further downward, resin canals became circular addition of metaxylem cells outside. or elliptic on the upperside although they retain­ Further down the shoot, the remaining cells ed irregular shapes on the underside. An altera­ between the procambial bundles became more tion in the internal pressure within the shoot meristematic, and periclinal divisions from fas­ due to minute movement might be the cause of cicular to in terfascicular regions seemed to pro- Downloaded from Brill.com09/27/2021 08:41:26AM via free access IAWA Bulletin n.s., Vol. 7(\),1986 23 Fig. 2. Transverse section of the shoot 0.2 cm below the apex. No difference in the upper- and underside of the shoot can be seen. - Fig. 3. Procambium viewed at a point 0.3 cm below the shoot apex. Note the protoxylem cells showing a more or less square outline in cross section (arrow­ head). Downloaded from Brill.com09/27/2021 08:41:26AM via free access 24 IAWA Bulletin n.s. , Vol. 7 (1),1986 @ Fig. 4. Transverse section of the shoot 0.6 cm below the apex. Note the irregular shape of resin canals on the underside (lower side of the photograph) of the shoot. ceed actively during primary growth (Esau, zones of periclinal divisions spread almost con­ 1943). In this way, many cells of the interfasci­ tinuous across the procambium bundles (Fig. 6). cular residual meristem were incorporated in No difference in the development of both fasci­ the developing and enlarging procambium bun­ cular and interfascicular cambium between the dles. Further downward, with the advance in upper- and underside of the shoot could be ob­ development of the interfascicular meristem, served (Fig. 6a & b). Downloaded from Brill.com09/27/2021 08:41:26AM via free access IAWA Bulletin n.s., Vol. 7 (1),1986 25 ness, 3) development of the helical ridges and cavities, 4) the rounding of the cross-sectional shape, and 5) occurrence of intercellular spaces. Although mild compression wood cells in the transition zone had normal cell shapes and wall thickness, they lacked the S 3 layer and had a slightly excessive lignification in the outer re­ gion of the S 2 layer at the cell corners. More­ over, they tended to lack helical cavities in the transition zone. These tendencies in the transi­ tion zone are in accordance with earlier obser­ vations (Saiki & Tachida, 1973; Fujita et aI., 1979; Yumoto et aI., 1982; Yoshizawa et aI., 1982). The first occurrence of these compres­ sion wood cells was located 2.5-3.5 cm below the apex, as suggested by Saiki and Tachida (1973). Thus, following the formation of such mild compression wood cells, typical compres­ sion wood cells appeared on the underside (Fig. 9), at which site the recovery of the shoots from the displacements first initiated. Summarising the above observations con­ cerning the occurrence of compression wood and the development of vascular cambium in the shoots, the conclusion can be drawn that compression wood cells are formed correspond­ Fig. 5. Longitudinal radial view of developing ing to the righting movement of the shoot a lit­ primary xylem.
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