Stomatal responses to changes in temperature at increasing water stress

Large areas of the lower epidermis of full-grown leaves of Polypodium vulgare (and Valerianella locusta) are normally separated from the mesophyll by an extensive subepidermal airspace. Epidermal stripes were prepared for experiments to simulate these conditions in order to investigate stomatal reactions. They were placed with their inner surface in contact with an airspace of uniformly high humidity. The outer surface was treated with air of varying degrees of humidity. The stomatal reactions were observed by microscope and the opening of the guard cells determined photographically.Treatment of the outer side of the epidermis with dry air led to a rapid closing of the stomata, whilst moist air caused opening. This induction of opening and closing movements could be repeated up to 15 times with the same stoma by changing the degree of humidity. Neighbouring groups of stomata showed different apertures according to their individual humidity conditions. The degree of aperture of the stomata depended on the water potential of the ambient air and also on the humidity conditions in the subepidermal airspace.The cause of this stomatal behaviour could lie in the "peristomatal transpiration". In this way, the guard cells are able to function as "humidity sensors" which "measure" the difference in water potential inside and outside the leaf. Their aperture thus is controlled by their individual transpiration conditions. This controlling mechanism could be very important for the water economy of plants. They would appear to be able to reduce their transpiration through an increase in diffusion resistance of the stomata during decreasing humidity in the ambient air, without changing the water status of the whole leaf.

The stomata of plants growing in the Negev Desert, namely the stomata of the mesomorphic leaves of Prunus armeniaca, the xeromorphic stems of Hammada scoparia, and the succulent leaves of Zygophyllum dumosum, respond to changes in air humidity. Under dry air conditions diffusion resistance increases. Under moist air conditions diffusion resistance decreases. When the stomata close at low air humidity the water content of the apricot leaves increases. The stomata open at high air humidity in spite of a decrease in leaf water content. This excludes a reaction via the water potential in the leaf tissue and proves that the stomatal aperture has a direct response to the evaporative conditions in the atmosphere. In all species the response to air humidity is maintained over a period of many hours also when the soil is considerably dry. The response is higher in plants with poor water supply then in well watered plants. Thus for field conditions and for morphologically different types of photosynthesizing organs the results confirm former experiments carried out with isolated epidermal strips.

When stomates of Zea mays open K and Cl migrate from the subsidiary cells into the guard cells; when the stomates close both elements return to the subsidiary cells. Subsidiary cells function as reservoirs for K and Cl. Import of K and Cl into the guard cells and loss of both elements from the guard cells become observable 1 or 2 min after light is turned on or off, both when histochemical methods and the electron-probe microanalyzer are used for detection. Each stomatal complex of maize contains on the average 10±3×10(-13) gram equivalents (eq) of K and 4±1×10(-13) eq of Cl. Guard cells accumulate K in the light and CO2-free air at an average rate of 10×10(-15) eq K per minute, and Cl at approximately half that rate.