SPIRAL GRAIN AND ANTICLINAL DIVISIONS IN THE CAMBIUM OF CONIFERS

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Abstract
In the cambium of mature trees a general conformity was found in the orientation of the partition in the pseudotransverse division of fusiform initials through out sectors of varying size. The proportion of divisions deviating from the preferred orientation varied with the tree, the locality, and the species, but was usually less than 10%. As radial accretion followed, periodic reversals occurred in the orientation of anticlinal divisions, the spacing between reversals, in terms of xylem increment, being related to the frequency of anticlinal division. The mean interval shortened as the rate of division rose according to the equation xy = k. While this overall relationship obtained, there was generally some inequality in the relative duration of leftward and rightward orientations of division. In species with grain in the outer wood slanted to the right, a rightward tilt of division persisted for longer periods than a leftward tilt. Conversely, in species with grain slanted to the left, thickness of xylem showing leftward orientation of multiplicative divisions exceeded that with rightward orientation. Changes associated with growth from sapling to adult were studied in Picea, in which genus grain is usually to the left in the inner wood and to the right in the outer wood. Here multiplicative divisions were usually inclined to the left in the growth of the first few years, whereas in the later growth orientation to the right endured for longer periods than to the left. Both orientation of pseudo-transverse division and direction of cell elongation after division seem to be under a general polar control. This apparently is an important element in the induction of spiral grain. Above-average net gain of fusiform initials in the turmoil of cell addition and loss, which accompanies circumferential expansion, may be a contributory factor. Serving to restrain the development of an excessive spirality are periodic reversals in direction of tilt in anticlinal division, and a high ratio of loss of cambial cells relative to the frequency of anticlinal division.