Abstract. Peristomatal transpiration is defined as the relative high local rate of cuticular water loss from external and internal surfaces around the stomatal pore and its decisive role in the control of stomatal movement is re‐emphasized. As the resistance towards changes in air humidity is low in the pore surroundings, the state of turgor is particularly unsteady there. Due to the inherent instability the guard cell ‘senses’ fluctuations in the supply‐demand relationship of water and is thus the control unit proper. The environmental variables (supply and demand) are cross‐correlated within the subsidiary cell and the information is transmitted to the guard cell through the water potential gradient between the two cells. A conceptual segregation of a ‘humidity response’ by ‘passive’ stomatal movements is rejected. As ions always accumulate at the most distant point of the liquid path and as this point varies with pore width according to the prevailing water potential gradients, it is felt that the water stream is causing the characteristic pattern of ion distribution within the epidermis. Passive import of ions is attributed to local concentration gradients which are steepened by continuous supply and by water uptake into the guard cell in response to starch hydrolysis. A mechanistic model supplements the discussion.

This paper reviews research on the effects of ozone on stomatal behavior, and draws attention to the direct link between stomatal function and air pollution. Contradictory results concerning the effects of air pollutants on stomatal activity are attributed to the use of univariate statistical treatments of data. Alternative experimental methods that subject stomata to a functional test are described. These methods verify the hypothesis that ozone predisposes trees to drought stress. It is proposed that ozone directly attacks the walls of guard and subsidiary cells, leading to impairment of the sensory mechanism of the stomata.

Four experiments on the simulation of a persistent drought period were carried out with cloned Picea abies (L.) Karst. trees: two in the field under varying weather conditions and two in a climate chamber under variously manipulated humidity conditions. Patterns of diurnal dynamics in gas exchange rates and water potential were monitored and analyzed. The first phase of the drought was characterized by relatively high daily maxima for photosynthesis and transpiration. With decreasing humidity during the day, the values dropped steeply, and the declines were larger and occurred earlier on each passing day of the drought period. When soil water potential was lower than -2000 hPa, maximum stomatal aperture was greatly reduced despite a humid atmosphere. Under these conditions, rates of photosynthesis and transpiration decreased less steeply from the daily maxima and differences between maxima and minima were small. In the field, the daily sums of transpiration and photosynthesis were more dependent on atmospheric conditions than on soil water potential. In the growth chamber experiments, the daily sums of transpiration and photosynthesis decreased continuously as the soil dried, at first steeply until a soil water potential of -2000 hPa was reached, then slowly. Predawn water potential values fluctuated under field conditions, but tended to decrease with time, whereas needle osmotic potential increased slightly. Because relative humidities did not reach 100% in the growth chamber, predawn water potentials of plants in the growth chamber were never higher than -1.0 MPa although the soil was saturated. In the experiment with a high average air humidity during the daily stress period, relatively high predawn water potentials were maintained until lower soil water potentials of -8000 hPa were reached. Results were used to assess the importance of evaporative demand versus soil drying on stomatal responses within the context of current concepts of plant water relations. The observed trends in diurnal dynamics can be explained solely by the interdependency of leaf conductance and water potential. Stomata react directly to the ratio of water supply to demand. The central role of peristomatal transpiration in this system is emphasized.

Abstract. With the aid of specifically designed potometer experiments, it is shown that, after ozone fumigation, twigs transpiring in gas exchange chambers show poor water balance in decreasing humidity. The quotient of water uptake to water loss never falls below 0.9 in healthy material because of the control capacity of the stomata. In twigs from a tree fumigated with ozone irregular and delayed stomatal closure results in values of < 0.5 or even lower, depending on the degree of damage. As a result, in dry air, the transpiration rates of fumigated twigs often fall far below those of the control material, even if they were higher than the latter in humid air. In analogous experiments, the difference in behaviour between twigs of densely (‘healthy’) and sparsely needled (‘damaged’) trees from the natural stand is comparable to the difference between controls and ozone‐fumigated trees in most respects. In soil that is more or less dried out and after the best possible saturation of the twigs during the night, the transpiration rates of fumigated trees increase fairly strongly in the humid chamber air at dawn, but finally decrease more or less suddenly to lower values than in the controls. The results are placed in the context of the basic research on plant water relations and compared with histological changes in the stomatal apparatus after a period of fumigation as described earlier. Therefore, long‐term effects of pollution can be explained as a specific distrubance of hydroregulation.

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Summary. After fumigation with ozone, the exterior periclinal walls of the stomatal apparatus of Picea abies appear to be partially delignified, as in earlier experiments with SOy This shows up cytophotometrically as reduced UV absorption. Random samples from the stand named Wank in the Bavarian Alps clearly showed a relation between the degree of lignification of the stomatal cells and the grading of the respective trees in their damage classes. The significance of delignification for the regulatory capacity of the stomata is discussed, and a hypothesis is proposed for a specific disturbance of hydroregulation by ozone.

Reduced UV absorbance indicated partial delignification in the walls of guard and subsidiary cells in material damaged in the natural environment. After fumigation of healthy young plants with 0.3 ppm SO2 for 20 days the symptoms were basically the same. The loss of lignin is related to malfunetionings of the stomatal control system.

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