Restriction of nyctinastic movements and application of tensile forces to leaves affects diurnal patterns of expansion growth

Leaves of Ricinus communis L. show strong nyctinastic movements as well as diurnal variations of growth activity, combined with prominent basipetal gradients of relative growth rate. A novel technique, using digital image sequence processing, is able to resolve such spatio-temporal patterns of leaf growth with high resolution. In this paper we analyse the impact of prevention of nyctinastic movements, and the potential of tensile forces to overcome the retarding effects. Tensile forces affected leaf expansion and tissue expansibility in a dose-response relation. In a comparison with freely-growing leaves, an appropriate, optimal tensile force was identified that rebuilt the natural diurnal course of leaf expansion. With this tensile force, undisturbed patterns of temporal and spatial growth distribution as well as undisturbed concentrations of major cations, amino acids or soluble sugars were observed. However, diurnal fluctuations in starch content of almost fully-grown leaves were affected by the treatment and could not be compensated by tensile forces. This effect might point to a connection between diurnal growth variations of the leaf vein, biomechanical forces that synchronize growth within the vein, and metabolism of carbohydrates as growth substrates of the vein. We therefore hypothesize that interveinal tensions, which are produced during nyctinastic leaf movements, are (i) required for undisturbed leaf growth and (ii) can be simulated by application of adequate tensile forces.