Biomechanical study of the effect of a controlled bending on tomato stem elongation: global mechanical analysis.

An experiment was designed to apply a controlled bending to a tomato stem and simultaneously to measure its effect on stem elongation. Stem elongation was measured over 2 d until steady and equal rates were obtained for the control and the treated plants. Thereafter, the basal part of the stem was submitted to a transient controlled bending at constant displacement rate using a motorized dynamometer. After load removal, stem elongation was again measured for 2 d. The tested plants were mature (height visible internodes) and only the basal part of the stem, which had already finished elongation, was loaded (hypocotyl and the first three internodes). A few minutes after the application of bending, elongation stopped completely for 60 min. Thereafter it took 120-1000 min to recover a rate of elongation similar to the control. The growth response was exclusively due to the bending of the basal part of the stem. It was shown that the side mechanical perturbations on the roots and on the stem tissues interacting directly with the clamp were not significantly involved on the elongation response. These results give evidence for mechanical perception and plant signalling from the basal stem to the upper elongating zone. However, none of the variables characterizing the global mechanical state of the bent part of the stem (i.e. the maximal force, bending moment, inclination, mean curvature of the stem, stored mechanical energy) could quantitatively explain the variability of the growth response. A more local mechanical analysis is therefore needed to elucidate how the mechanical stimulus is perceived.

Biomechanical study of the effect of a controlled bending on tomato stem elongation: local strain sensing and spatial integration of the signal.

In a previous paper it has been demonstrated that tomato stems, submitted to a controlled basal bending, had a reduced terminal primary elongation, indicating mechanosensing and intra plant signalling. The 'intensity' of the growth response, as measured by the time to recover an elongation rate similar to the control, varied hugely between plants. However, no relation was found between the intensity of this response and the mechanical variables characterizing the global mechanical state of the stem. In this paper, a local analysis of mechanical state of each bent stem is performed in the context of beam theory. The spatial distributions of local variables all along the stem (curvature, bending moment, strains and stresses) are established. The validity of hypotheses underlying the mechanical analysis is demonstrated. To investigate the relationships between the mechanical stimulus and the growth response, a novel biomechanical analysis based on spatial integration of the mechanical stimulus is presented. It revealed that the mechanosensing is local and scattered through the stem and that the variability of the growth response is only explained by the integrals of the longitudinal strain field.

Morphological responses and molecular modifications in tomato plants after mechanical stimulation.

Study of the growth responses of Lycopersicon esculentum (Mill. cv. VFN8) to mechanical stimulation applied to a single young internode showed a rapid and sharp decrease in stem elongation and an inhibition of elongation of several internodes, indicative of information transmission in the plant. A new tomato cDNA partial clone encoding calmodulin was isolated and used to study the time course of the gene induction in response to the rubbing treatment. Northern blot analysis showed a maximum accumulation of calmodulin mRNA 2 h after mechanical stimulation, not only in the rubbed internode, but also in upper and lower internodes and in young leaves. Treatment of the plant with calcium and EGTA showed the involvement of calcium and, in particular, intracellular calcium in calmodulin gene expression and cellular response.