L. Kappen

Responses of stomata to changes in humidity

SummaryLarge 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.

Stomatal Responses to Changes in Humidity in Plants Growing in the Desert

SummaryThe 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.

The role of air humidity and leaf temperature in controlling stomatal resistance of Prunus armeniaca L. under desert conditions

SummaryThe gas exchange of the apricot (Prunus armeniaca L.) growing in the runoff farm at Avdat (Negev, Israel) was measured during its growing period using temperature- and humidity-controlled chambers. Water potentials of the xylem were measured with a pressure bomb, and the mesophyll internal CO2 concentration was calculated from simultaneous measurements of net photosynthesis and transpiration.The daily changes in water potential Ψ had only little effect on the daily course of stomatal resistance. The early morning peak of CO2 uptake was reached when Ψ had already dropped to very low values. On dry days, Ψ and the relative water content of the leaf were improved at noon during the time of stomatal closure. On humid days, Ψ dropped to very low values (43.5 bar) at a high transpiration rate without causing stomatal closure, as much as on the dry days when stomata where more closed at less water stress. The observed changing sensitivity of the stomata to changes in air humidity during the season is related to the water status in the plant. This change is possibly caused by a long-term effect of stress in this habitat.The daily changes in stomatal diffusion resistance did not consistently correlate with changes of the CO2 concentration in the intercellular air spaces. In the morning a decreasing internal CO2 concentration was even inversely correlated to the stomatal response. In the afternoon the effect of an increasing internal CO2 concentration and the effect of external climate on stomatal response could be additive. However, at the time, when CO2 uptake reached a second peak in the afternoon the same value of diffusion resistance is reached at very different levels of internal CO2 concentration as compared to the morning.For the regulation of the diffusion resistance in apricot under the natural conditions, the effects of plant internal control mechanisms are overruled and/or modified by the external climatic factors of air humidity and temperature. The significance of the climate-controlled stomatal response for the existence and cultivation of this plant species in an arid habitat is discussed.

Stomatal responses to changes in temperature at increasing water stress.

SummaryThe response of stomata to a gradual increase in temperature at increasing plant water stress was studied in a hot desert habitat (Negev, Israel) in the field, but under controlled temperature and humidity conditions. Four native species (Zygophyllum dumosum, Artemisia herba-alba, Hammada scoparia, Reaumuria negevensis) and one cultivated plant (Prunus armeniaca) were used in these studies. The stomatal response to temperature was compared with the response in well-irrigated plants of the same species.At low water stress, the diffusion resistance for water vapour decreased in response to a gradual increase in temperature. Transpiration increased accordingly. This response was reversible. All species responded in the same way. The opening of stomata with increasing temperature was apparently independent of the stomatal response regulated by atmospheric humidity. At high plant water stress, the stomatal response was reversed, i.e., the stomata closed when temperature was gradually increased. This stomatal closure was also independent of the closure regulated by atmospheric humidity. The plant water potential at which the stomatal response to temperature was reversed, differed among the species investigated.

Long-term effects of drought on wild and cultivated plants in the Negev desert. II. Diurnal patterns of net photosynthesis and daily carbon gain.

SummaryThe seasonal change in diurnal patterns of net photosynthesis and daily carbon gain is studied in relation to the plant water status of the irrigated and non-irrigated naturally growing desert species Hammada scoparia, Zygophyllum dumosum, Artemisia herba-alba and Reaumuria negevensis. Comparison is made to cultivated Prunus armeniaca. Under non-irrigated natural conditions Hammada scoparia, a C4 plant, showed one-peaked flat diurnal courses of CO2 uptake which changed into a pattern of a high morning peak of CO2 uptake or slightly two-speaked curves in the late dry season. In contrast, the C3 species Zygophyllum dumosum, Artemisia herba-alba and Prunus armeniaca changed from one-peaked to distinct two-peaked patterns. At the end of the dry season, non-irrigated plants showed respiration only. Reaumuria negevensis had one-peaked curves with a low level of CO2 uptake.There is no general relation between day-time CO2 gain and pre-dawn water potential for the investigated species. In order to characterize the effect of soil drought, the CO2 gain during day-time of non-irrigated plants is expressed as a percentage of the CO2 gain of the irrigated counterparts. After an initial period of minimal drought effect, the relative day-time CO2 gain decreases almost linearly with cumulative water stress as determined by the daily addition of pre-dawn water potentials for the non-irrigated plants since the last rainfall. The slope of decrease differs from species to species. The relation of daily CO2 gain to maximal net photosynthesis is discussed. Initially, at a good plant water status, the daily CO2 gain does not decrease in proportion to the maximal photosynthetic rates as a result of stomatal control at high photosynthetic activity. At increasing water stress the daily CO2 gain decreases more than proportionally to the decrease of the maximal rates.

Extreme water stress and photosynthetic activity of the desert plant Artemisia herba-alba asso

SummaryDuring the dry season in the Negev desert (Israel) Artemisia herbaalba in its natural habitat has a very low water content. It shows values of negative hydrostatic pressure in the xylem down to -163 bars and an extreme of osmotic potential in the leaves of -92 bars. The diurnal water stress does not decrease strongly in the night. Under these conditions Artemisia is still photosynthetically active for a few hours of the day during the whole dry period.

The temperature-related photosynthetic capacity of plants under desert conditions

SummaryAs described earlier, the native arido-active perennial Hammada scoparia and the cultivated Prunus armeniaca exhibit characteristic seasonal shifts of their temperature optimum of net photosynthesis (OP) under desert conditions in the Negev. In the present paper the OP values were compared with the actual tissue temperatures of the experimental plants.During the growing period from March to September the duration of optimal temperatures for net photosynthesis (OP±3°C) experienced by the plants was 32.2% of the total time at light saturation for P. armeniaca and 27.8% for H. scoparia. For optimal photosynthesis the branchlets of H. scoparia are too cold for 66.1% of the time span and too warm for 6.1% of the time. The respective values for the leaves of the apricot are 28.6% and 39.2%. Simulations at changed tissue temperature show, that for P. armeniaca neither a higher nor a lower temperature regime would lengthen the time span for optimal thermal conditions. For H. scoparia, however, an increase of the general temperature level by 6°C would considerably improve the temperature-related photosynthetic efficiency. The natural temperature responses of the plants were compared with simulations using OP values which are supposed not to shift but to stay constant from March through September at their spring minimum, their summer maximum, or at an intermediate value. For P. armeniaca such constant OP values would result in a shorter duration of optimal temperature conditions. With this plant the natural seasonal shift of the temperature characteristics appears to provide an advantage in respect to its photosynthetic capacity. Contrary to this, for H. scoparia a constant OP value at the low spring level or even at the intermediate level during all the season would result in a substantially prolonged period of favourable temperature conditions for photosynthesis. In this case the seasonal change of optimum temperature for photosynthesis with higher OP values in summer signifies a disadvantage with respect to the temperature-related photosynthetic capacity at the habitat in the central Negev. Apparently this C4 plant is adapted to higher temperatures than were present. It appears that “acclimations” of native plants are not always beneficial.

An empirical model of net photosynthesis for the desert plant Hammada scoparia (Pomel) Iljin

SummaryAn empirical model for describing daily courses of net photosynthesis in Hammada scoparia is being developed. The model is based on the functional relationships, by which various environmental factors affect the photosynthetic activity and which can be measured by experiment in the field. In a sequence of steady-states daily courses of net photosynthesis are predicted during a growing season considering the variability of the physiological states and the capacity for regulative adaptations. The rate of net photosynthesis at a certain date is calculated from the maximal rate of CO2 uptake being expected at that season and from the effects of light, temperature, and air humidity which are scaled from 0 to 1. All factors are connected multiplicatively. The light function accounts for the seasonal changes in the light curve, the temperature function is based on the seasonal shift of the temperature optimum, and the humidity function considers the increasing sensitivity of the stomatal humidity response at increasing water stress. The model is built to be a submodel of a general ecosystem model, where various other submodels (i.e. water stress model, phenology model) are supplied. The present model is tested by predicting daily courses at extreme climatic conditions during the year and by comparing the predicted values of gas exchange with values being measured in an independent experimental procedure. The result shows that the model is able to simulate the natural behaviour of Hammada scoparia during the growing and dry season of a desert habitat. The problems of incorporating the influence of water stress, the interaction of the various factors, and the phenological aspect of the photosynthetic activity is being discussed.

Distributional pattern of water relations and net photosynthesis of Hammada scoparia (Pomel) Iljin in a desert environment

SummaryIn the Central Negev, the arido-active Hammada scoparia (Chenopodiaceae) is mainly distributed in runnels and loessial plains of the wadis and exists with fewer and smaller individuals on slopes and tops of the neighboring hills. At the end of the dry season, water relations and chloride content of plants from these habitats and of artificially irrigated plants were determined and compared with plant shape and photosynthetic activity. The osmotic potentials and total water potentials of the plants differ characteristically with certain groups of stands. The simultaneously determined range of noon water potentials of the plants within a transect is as high as the annual amplitude of one plant individuum in the wadi (about 45 bars). Plants in the runnel of the wadi show, like the irrigated plants, the highest values of water potentials and their components but markedly lower chloride content than the irrigated ones. Total water and osmotic potentials of plants of the loessial wadi plains are extremely low. Their chloride content is not very high in contrast to that of plants of the hillslope and hilltop. Hill plants, although poorly developed and scarcely branched, have higher water and osmotic potentials than those of the plains.Net photosynthesis of a plant on a natural stand of the wadi plain is, in September, markedly depressed at noon but maximal at noon in an artificially irrigated plant. In contrast to irrigated and nonirrigated wadi plants, a plant of the hillslope shows, already in July, a midday depression of photosynthesis. Whether the relatively low water potentials of the big plants of the loessial plains are the results of a rapid biomass production in the rainy season which might have caused a very strong exhaustion of the soil water reserves for the late summer is discussed. Hill plants do not grow so well and obviously decrease their water exchange even in summer.