ALBERT

Notes: Summary The influence of climatic factors on net photosynthesis, dark respiration and transpiration was investigated in the Negev Desert at the end of the dry summer period when plant water stress was at a maximum. Species studied included: dominant species of the natural vegetation (Artemisia herba-alba, Hammada scoparia, Noaea mucronata, Reaumuria negevensis, Salsola inermis, Zygophyllum dumosum), cultivated plants receiving rainfall and run-off water during the winter season in the run-off farm Avdat (Prunus armeniaca, Vitis vinifera), and irrigated cultivated plants receiving additional water during the summer season (Citrullus colocynthis, Datura metel). 1. Light saturation of net photosynthesis was reached at 60–90 klx conforming to the high solar radiation intensities of the desert. 2. Maximum rates of CO2 uptake per unit of dry weight for the irrigated mesomorphic plants was ten times that of the wild plants. However, in comparison to the other species, maximal rates of CO2 uptake for wild plants were higher when calculated on a leaf area basis than when represented on a dry weight basis. Maximum rates of net photosynthesis per unit chlorophyll content for some of the wild plants (Salsola and Noaea) were comparable to those of the cultivated Vitis and irrigated Citrullus and Datura, Hammada exhibited even higher rates than Prunus. This demonstrates the great photosynthetic capacity of the wild plants even at the end of the dry season. 3. The upper temperature compensation point for net photosynthesis of the wild plants was unusually high as an adaptation to the temperatures of the habitat. Compensation points higher than 49°C exceed the maxima known so far for other flowering species. Maximum rates of net photosynthesis of Hammada were measured when the temperature of the photosynthetic organs was 37°C; at 49°C photosynthesis was only reduced by 50%. 4. Leaf temperature affects plant gas exchange by influencing stomatal aperture. Diffusion resistance of leaves to water vapour was reduced at low temperatures and increased at high temperatures. Reduction of net photosynthesis and transpiration of desert plants at midday may, therefore, be the result of temperature-induced stomatal closure. The possible influence of peristomatal transpiration on stomatal aperture is also discussed. Peristomatal transpiration is directly related to the vapour pressure gradient between the leaf mesophyll and the ambient air which increases with increasing temperatures. 5. Diffusion resistance to water vapour was reduced at high temperatures approaching the limits of heat resistance, due to increased stomatal aperture. This resulted in greater transpirational cooling. 6. Under conditions of increased leaf water stress, diffusion resistance increased, either by sudden stomatal closure at specific threshold values of water stress or through a continuous increase in resistance. This increased resistance is coupled with decreases in transpiration and photosynthesis. 7. In several plant species increased diffusion resistance during the course of the day caused decreased transpiration without a corresponding decrease in photosynthesis. Under these conditions, the ratio of CO2 uptake to transpiration became more favourable as the day progressed. The possibility that this favourable gas exchange response is the result of an increased mesophyll resistance to water vapour loss is discussed.

Notes: Summary Carbon dioxide exchange and transpiration measurements of various wild and cultivated plants were carried out during the dry summer period in 1967 in the Central Negev Desert (Israel). A mobile laboratory used for these investigations is described. Measurements were carried out with conditioned plant chambers which followed either the ambient temperature and humidity or else allowed the experiments to be carried out under constant conditions. The accuracy of the measurements was estimated. The mean error of the determination of the CO2 exchange rate amounts to ±0.07 mg CO2·g-1·h-1. Transpiration rate is measured with an error of ±0.15 g H2O·g-1·h-1. The response time of the instrumentation to reach 90% equilibrium after a change in photosynthesis or transpiration is 7 to 9 minutes. Errors which are caused by changes of quality of incident radiant energy and altered turbulence conditions for the leaves enclosed in the chamber, are discussed.

Notes: Summary The photosynthesizing branches of Hammada scoparia, one of the typical dwarf shrubs of the Negev desert, undergo a seasonal change from succulent to xeromorphic anatomy. This trend is accompanied by a marked decrease of water content and of total water Ψ plant and osmotic Ψ π plant potential. Irrigated plants do not show such transitions. The daily courses of Ψ plant and Ψ π plant showed minima around noon and a tendency for maxima before sunrise. Turgor pressure Ψ p plant reached minima around noon and became negative (until ca.-10 bars). Generally, Ψ plant decreases with increasing water vapour concentration difference between plant and air (WD) in the first half of the day, and in the second half the reversal of this trend occurs. Mostly smaller increments of Ψ plant were correlated with larger increases in WD which lead to the conclusion that stomates closed enough to maintain transpiration at a constant value. Non-irrigated and irrigated plants showed different hysteresis loops of relation between Ψ plant and WD. Regulatory reduction of transpiration appears largely independently of Ψ plant which is in spring and with irrigated plants on a high level, with non-irrigated plants in summer on a low level. In summer the continous but decreasing drop of Ψ plant with increasing WD was interpreted as caused by a change in soil or root resistance. Independent of the seasonal state and of the Ψ plant level, H. scoparia regulates its water status within limited ranges of Ψ p plant changes: the irrigated plants on a higher level, the non-irrigated on a lower level of Ψ p plant . The water contents of the tissues of H. scoparia are linearily related to Ψ plant as well as Ψ p plant . Steeper slopes with non-irrigated plants in summer than with spring palnts and with irrigated plants during the whole season signify that in the latter a certain increment in turgor pressure corresponds to a large gain in water content while in the non-irrigated summer plants it varies only little for an identical change in Ψ p plant . This behaviour of non-irrigated wild plants apparently is due to the change of the elastic properties of the tissues in the assimilating branches.

Notes: Summary An 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.

Notes: Summary Temperature dependence of net photosynthesis under conditions of light saturation and maximum air humidity was measured throughout the season in the Central Negev Desert (Israel). Experimental plants were the wild growing Hammada scoparia and Prunus armeniaca cultivated in the runoff farm of Avdat. The optimum temperature for net photosynthesis and the upper temperature compensation point of CO2 exchange showed a characteristic seasonal variation with low values in spring and fall and high values in mid-summer. This shift was exhibited by plants growing under conditions of normal soil-water stress as well as by irrigated plants. There was no general correlation between the changes in temperature dependence of net photosynthesis of the plants, their maximum photosynthetic capacity under the experimental conditions, their daily photosynthesis maximum under natural conditions, and their rate of dark respiration. The seasonal shift of the photosynthetic response to temperature cannot be explained by changes in the temperature sensitivity of the stomata. It may be caused by seasonal changes of biochemical and/or biophysical properties. A number of observations made on other wild plants also showed, in all cases, seasonal shifts of the upper temperature compensation point, with an amplitude of 6.0°C–13.7°C.

Notes: Summary Experiments with Prunus armeniaca were carried out under conditions of constant temperature but varying air humidity. Experiments were also contucted with a constant water vapor difference between the evaporating sites in a leaf and the air, but with varying leaf temperature. These served as a basis for predicting the daily course of total diffusion resistance under the natural climatic conditions of a desert. For the simulation, the rsults of the experiments at constant conditions with only one variable factor are fitted with empirical equations which serve as “calibration curves” to predict the change in diffusion resistance caused by a change in humidity and temperature calculated from the meteorological data of a desert day. The simulation shows that for P. armeniaca humidity and temperature are the dominating factors in controlling the daily course of diffusion resistance. For meteorologically very different days the simulation allows the increase in diffusion resistance in the morning to be predicted with an accuracy of 90%–105% as compared to directly observed measurements. In the afternoon, especially after extreme climatic conditions during the morning, the deviation between predicted and observed values of diffusion resistance may be greater, but not more than -20% to -30%. This possibly indicates the existence of an additional factor of significance which was not included in the simulation. The two peaked curves of net photosynthesis and transpiration characteristic of plants living under arid conditions can be explained in this species by the humidity-and temperature-controlled stomatal response. This stomatal regulation leads to a decreasing total daily transpirational water loss on a dry day as compared to a moist one. The significance of this controlling mechanism for the primary production and the water relations of P. armeniaca is discussed.

Notes: Summary In 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.

Notes: Summary The water relations characteristics of three grass species (Panicum maximum var. trichoglume, Cenchrus ciliaris, Heteropogon contortus), and a legume (Macroptilium atropurpureum) grown in the field were measured using both a modified pressure/volume technique with pressure bomb measurements on single leaves and a dewpoint hygrometry technique applied to fresh and to frozen and thawed leaf discs. The two techniques agreed well in the estimates of osmotic potential at full turgor and the water potential at zero turgor. However, for parameters such as the relative water content at zero turgor, bound water and bulk modulus of elasticity there was a poor correlation between the estimates from the two methods. The pressure/volume technique gave less variable results and is more convenient for field use than the hygrometry technique. The determination of the modulus of elasticity from various functions relating pressure potential to relative water content is discussed.

Notes: Summary A digital registration system used with temperature- and humidity-controlled cuvettes for net photosynthesis and transpiration measurements in the field is described. The associated errors of the measured parameters and calculated data are estimated. The digitalization is based on an analogue registration which is of primary importance in the control of experimental conditions in the cuvettes. The digital system is connected to the analogue registration in series. The error associated with digitalization is 0.1% across 70% of the scale. This error increases to 0.2% between 3 and 30% on the scale due to a minor lack of linearity. The reproducibility of the digitalization is ±0.024%. The error associated with data transfer in the digitalization and the errors of the analogue registration are estimated for temperature and humidity measurements (error of air and leaf temperature is ±0.1° C; error of the dew point temperature is ±1.1° C dew point). The effect of these errors on the calculation of relative humidity and the water vapour difference between the leaf and the air is determined using the progressive error law. At 30° C and 50% relative humidity, the error in relative humidity is ±7.4%, the error for the water vapour difference is ±6.6%. The dependence of these errors on temperature and humidity is shown. The instrument error of the net photosynthesis measurement is calculated to be ±4.2%. Transpiration measurements have an average inaccuracy of ±8.3%. The total diffusion resistance which is calculated from values of transpiration and the water vapour difference has an average error of ±10.9%. The sizeable influence of errors in humidity and temperature measurements on the calculated diffusion resistance is demonstrated. The additional influence of biological errors associated with field measurements is discussed.