The Nature of Reaction Wood III. Cell Division and Cell Wall Formation in Conifer Stems

Cell division, the nature of extra-cambial readjustment, and the development of the secondary wall in the tracheids of conifer stems have been investigated in both compression wood and normal wood. It has been shown that the reduction in tracheid length, accompanying the development of compression wood and, in normal wood, increased radial growth after suppression, result from an increase in the number of anticlinal divisions in the cambium. From observations of bifurcated and otherwise distorted cell tips in mature tracheids, of small but distinct terminal canals connecting the lumen to the primary wall in the tips of mature tracheids, and of the presence of only primary wall at the tips of partly differentiated tracheids, and from the failure to observe remnants of the parent primary walls at the ends of differentiating tracheids, it has been concluded that extra-cambial readjustment of developing cells proceeds by tip or intrusive growth. It has been further concluded that the development of the secondary wall is progressive towards the cell tips, on the bases of direct observation of secondary wall formation in developing tracheids and of the increase found in the number of turns of the micellar helix per cell with increasing cell length. The significance of this in relation to the submicroscopic organization of the cell wall has been discussed. Results of X-ray examinations and of measurements of· tracheid length in successive narrow tangential zones from the cambium into the xylem have indicated that secondary wall formation begins before the dimensional changes of differentiation are complete.