Oscillations in stomatal conductance and plant functioning associated with stomatal conductance: Observations and a model.

Measurements of transpiration, leaf water content, and flux of water in a cotton plant exhibiting sustained oscillations, in stomatal conductance are presented, and a model of the mechanism causing this behaviour is developed. The dynamic elements, of the model are capacitors-representing the change of water content with water potential in mesophyll, subsidiary and guard cells-interconnected by resistances representing flow paths in the plant. Increase of water potential in guard cells causes an increase in stomatal conductance. Increase of water potential in the subsidiary cells has the opposite effect and provides the positive feed-back which can cause stomatal conductance to oscillate. The oscillations are shown to have many of the characteristics of free-running oscillations in real plants. The behaviour of the model has been examined, using an analogue computer, with constraints and perturbations representing some of those which could be applied to real plants in physiological experiments. Aspects of behaviour which have been simulated are (a) opening and closing of stomata under the influence of changes in illumination, (b) transient responses due to step changes in potential transpiration, root permeability and potential of water surrounding the roots, (c) the influence of these factors on the occurrence and shape of spontaneous oscillations, and (d) modulation of sustained oscillations due to a circadian rhythm in the permeability of roots.