Metabolic and physical control of cell elongation rate: in vivo studies in nitella

Several levels of control of elongation rate are revealed through the detailed study of responses of the Nitella internode to abrupt shifts in turgor. The immediate response, which apparently reflects the physical state of the cell, is approximately described by the equation r = (P - Y)m where r is rate, P is pressure, Y is the wall's yielding threshold, and m is related to the wall's apparent fluidity (reciprocal viscosity). Because P and Y are in the range 5 to 6 atmospheres, and (P - Y) is roughly 0.2 atmosphere, elongation rate is initially extremely sensitive to changes in P. A small step-down in turgor (0.7 atmosphere) stops growth, and a similar rise greatly accelerates it. These initial responses are, however, soon (15 minutes) compensated by changes in Y. An apparent metabolism-dependent reaction (azide-sensitive) lowers Y; strain hardening (azide-insensitive) raises it. These two opposing processes, acting on Y, serve as a governor on (P - Y), tending to maintain it at a given value despite changes in P. This ability to compensate is itself a function of turgor. Turgor step-downs are less and less well compensated, leading to lower rate, as turgor falls from 5 atmospheres to about 2 atmospheres where growth appears not to resume. This is the lowest attainable yield value, Y(1). The turgor dependency of compensation reflects a turgor requirement of the Y-lowering ("wall-softening") process. Thus the relation between steady state, r(s), and turgor is an indirect one, derived from time-dependent alterations of the cell wall. This relationship superficially resembles the instantaneously valid one in that, roughly, r(s) = (P - Y(1))m(s). Y(1) and m(s), however, have much lower values than Y and m. The duality of the elongation rate versus turgor relation and the prominent role of Y in regulating rate are the major features of growth control in Nitella.