Recently developed techniques have been used to reinvestigate the mechanism by which gibberellic acid (GA3) stimulates elongation of light-grown cucumber (Cucumis sativus L.) seedlings. Osmotic pressure and turgor pressure were slightly reduced in GA3-treated seedlings, which elongated 3.5 times faster than control seedlings. This indicated that GA3 enhancement of growth was not controlled by changes in the osmotic properties of the tissues. Stress/strain (Instron) analysis revealed that plastic extension of the cell walls of GA3-treated seedlings increased by up to 35% above the control values. Stress-relaxation measurements on frozen-thawed tissue showed that To, the minimum relaxation time, was reduced following application of GA3. In vivo wall relaxation (measured by the pressure block technique)showed that the wall yield coefficient was increased, and the yield threshold was slightly reduced. Thus GA3 affected both the mechanical (viscoelastic) and biochemical (chemorheological) properties of the cell walls of light-grown cucumber. The previous hypothesis, that GA3 stimulates cucumber hypocotyl growth by increasing osmotic pressure and cell turgor, is contradicted by our results.GA3 can reverse the inhibition of stem elongation caused by light in many plant species (1,12,17,18,21,23,24). The mechanism by which GA3 enhances the rate of elongation has been investigated by several groups, many of whom have concluded that GA3 acts on cell wall mechanical properties to increase growth (2,13,15,18,20).Cucumber is an often-cited exception to this general trend, in that GA3 is thought to stimulate hypocotyl elongation by causing increases in cell turgor pressure and osmotic pressure. This hypothesis, however, is based on meager direct evidence. Rather, it is indirectly supported by studies which found little effect of GA3 on the mechanical properties of growing walls (5, 14), and it assumes that wall growth properties are well correlated with wall mechanical properties (4). This assumption, however, is sometimes invalid. For example, a recent study (8) Therefore, this study was undertaken to reexamine the mechanism by which GA stimulated elongation in lightgrown cucumber. Osmotic and turgor pressures were measured directly, and in vivo wall relaxation properties were assessed in living tissue using the pressure block technique. We also assessed wall mechanical properties in frozen-thawed tissue by stress-relaxation and Instron analyses.
MATERIALS AND METHODS
Plant Material