A new study of cambium of Pinus sylvestris L., Tilia cordata Mill. and Wisteria floribunda (Willd.) DC provides fresh clues on the cambial dynamics, rejecting the hitherto held concept that intrusive growth of the fusiform initial occurs between the radial walls of adjacent initials. It demonstrates that intrusion of the elongating initial in fact takes place along tangential walls of adjacent fusiform initials and their immediate derivatives. It also suggests a new mechanism for ‘elimination of initials’. Intrusive growth of the fusiform initial was found to begin with development of characteristic slants, representing a transitional stage of the process of transformation of periclinal walls of fusiform initial cells into radial walls, as observed in transverse sections of active cambium. The gradually progressing event comprised (a) appearance of either a triangular microspace limited by two periclinal walls of a fusiform initial and its derivative and one radial wall of another fusiform initial in the adjacent radial file, or a rhomboidal microspace enclosed by four periclinal walls of two laterally adjacent fusiform initials and their immediate derivatives, (b) intrusion of elongating tip of fusiform initial from neighbouring file into the microspace thus formed, (c) symplastic growth of the cambial cell walls in radial direction, (d) unequal periclinal divisions of fusiform initial cells while growing intrusively, and (e) unequal periclinal divisions of derivative cells not growing intrusively. Intrusive growth between periclinal walls affected rearrangement of the fusiform initials but did not add to the cambial circumference. The existing concepts of (a) intrusion of the fusiform initial between radial walls of neighbouring initials and (b) elimination of fusiform initials from cambial surface have been reassessed and redefined.
This study on the cambium of Pinus sylvestris L. examines the intrusive growth of fusiform cambial initials and its possible contribution to the tangential and radial expansions of the cambial cylinder. The location and extent of intrusive growth of the fusiform initials were determined by microscopic observations and by mathematical modeling. In order to meet the required circumferential expansion of the cambial cylinder, the fusiform initials grow in groups by means of a symplastic rather than intrusive growth, leaving no room for the assumption that intrusive growth of the initials takes place between radial walls and has a direct role in the increase of the cambial circumference. Therefore, it is postulated that the fusiform initials grow intrusively between the tangential walls of the neighboring initials and their immediate derivatives and not between the radial walls of the adjacent initials as per common belief.
Currently it is believed that intrusive growth of fusiform cambial initials adds to the circumference of the cambial cylinder: the initial cells multiplied by anticlinal divisions are produced in excess, and the excess cells are later eliminated from the cambial surface. The present study, dealing with the intrusive growth of fusiform initials in the cambium of Laburnum anagyroides Medik, suggests that addition of a radial file of initials owing to intrusive growth, or elimination of any such file, has no visible effect on the tangential dimensions of a given cambial sector, and that intrusive growth of fusiform initials and the elimination of excess initials occur in unison. The two events complement each other, and the gain in size of the growing fusiform cell is accompanied by a reduction in size of its neighbour cell, thus keeping the tangential dimensions of the cambium unchanged. Our findings on L. anagyroides find support from illustrations of previous studies, which we have re-examined and re-interpreted. Our data suggest that increase in the cambial circumference is largely due to the symplastic growth of the fusiform initials in tangential direction.
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