ALBERT

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 During 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.

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.

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 A temperature- and humidity-controlled plant chamber for CO2 and H2O exchange measurements in the field is described in which the heat exchanger assembly and humidity controlling water vapour trap are separated from the plant cuvette. The shape and construction material of the plant cuvette can vary according to the demands of the experimental conditions and the size and growth form of the plant. The natural illumination field is only slightly altered in this plant cuvette. In the chamber, the temperature and humidity conditions can either be held constant throughout a wide range of conditions or can be programmed to track ambient condition. In this manner, not only temperature and absolute humidity are replicated, but it is also possible to reproduce the natural conditions of water vapour gradient between the evaporating surfaces in the mesophyll and the atmosphere, the relative humidity of the air, and the temperature difference between the leaf and the ambient air. Thus, the chamber appears to be an appropriate instrument to investigate with sufficient accuracy the reactions of individual plants in cultivation or in natural communities under field conditions.

Notes: Summary 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.

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 In a montane beech (Fagus sylvatica) forest the influence of the climatic factors, light and temperature, on net photosynthesis and on the CO2 balance of the ground vegetation was investigated. The total turnover of carbon was calculated. Species studied included: Athyrium filix-femina, Oxalis acetosella, Luzula luzuloides, Deschampsia flexuosa and young plants of Fagus sylvatica. 1. The light compensation point in all spp. is between 300 and 500 lux except for D. flexuosa where it is 2 klx. Light saturation is attained at 2–3 klx for A. filix-femina, at 5–6 klx for O. acetosella, and at 6–7 klx for L. luzuloides and F. sylvatica. The net photosynthesis of D. flexuosa increases linearly upto 12 klx. This plant, therefore, is more closely related to plants with high light requirements than all the other species under experiment. 2. The maximum rates of net photosynthesis in O. acetosella and A. filix-femina are higher than in all the other plants, independent of the reference system. Per unit dry weight they even attain rates of CO2 uptake (22–27 mg CO2/gdw·h) known from herbs under the much better light conditions of an open habitat. F. sylvatica and L. luzuloides exhibit per unit dry weight only 30% of this rate and D. flexuosa 25%. On a leaf surface area and chlorophyll content basis differences are smaller: F. sylvatics attains 75%, L. luzuloides reaches 50% and D. flexuosa only 30% of the maximal rates of net photosynthesis of O. acetosella and A. filix-femina. The higher CO2 uptake of O. acetosella and A. filix-femina points to a better adaptation of their photosynthetic apparatus in comparison to all the other species of the same habitat. 3. At light saturation the temperature optimum of A. filix-femina and O. acetosella covers a smaller range at lower temperatures than was found in the other species. These attain almost maximal rates of net photosynthesis over the whole range of temperatures of their natural habitat. At decreasing light intensities the temperature optimum of O. acetosella changes from 13–18° C at 8–12 klx to a range of even lower temperatures (9–12° C at 1 klx). 4. The respiration of the rhizome and the roots of O. acetosella is per unit dry weight 40% of the dark respiration rate in the above ground material. 5. The daily gain of net photosynthesis per unit dry weight of O. acetosella and A. filix-femina is 4 times as high as in L. luzuloides and in F. sylvatica and 7 times as high as in D. flexuosa. Per unit of surface area and chlorophyll content differences are smaller. The sequence in all cases remains the same. During the night D. flexuosa has the highest relative respiratory loss. Its CO2 gain over 24 hours is very small. 6. The importance of sun flecks on the CO2 balance is small in all species except D. flexuosa. More important is the mean light intensity and the rate of net photosynthesis which is attained under these conditions. The amount of CO2 photosynthetically bound in sun flecks is 6% of the daily balance in A. filix-femina, 16–19% of the daily balance in O. acetosella, L. luzuloides and F. sylvatica, and 27% of the daily balance in D. flexuosa. The existence of D. flexuosa is dependent on the occurrence of sun flecks on the forest floor. The ecological significance of the relative light intensity in the mosaic-like distribution of plants on the forest floor is discussed. 7. The varying success in adaptation to the conditions of the habitat becomes even more evident when compared with the primary production of the beech crown. The daily gain of net photosynthesis of O. acetosella and A. filix-femina per unit dry weight is much larger than in either the sun or shade leaves in the canopy of the same stand. Per unit surface area of the leaves they attain 18–20%, per unit chlorophyll content 32% (L. luzuloides and the young plants of F. sylvatica 16–27%, D. flexuosa 4%) of the gain of net photosynthesis in the beech sun leaves. 8. A comparison with a model of primary production (maximal rates of net photosynthesis under experimentally optimal conditions over the whole day = 100%) shows what effect the different climatic factors of the natural habitat have in limiting the CO2 balance, and to what extend the actual CO2 gain reaches the physiological optinum. On the forest floor the rate of net photosynthesis is reduced primarily through the intense shade of the beech canopy and by dawn and dusk (reduction of the maximal CO2 gain in O. acetosella and in A. filix-femina ca. 50%, in L. luzuloides and F. sylvatica ca. 60% and in D. flexuosa ca. 86%). The effect of additional clouds is smaller (reduction of the maximal CO2 gain in all species 4–6%, maximal 19%). The effect of temperature is very small for L. luzuloides, F. sylvatica and D. flexuosa. Corresponding to the low temperature optimum the influence of the prevailing temperatures is much higher in O. acetosella and A. filix-femina (4–6% reduction of the maximal CO2 gain through temperatures above optimum). 9. For an assessment of the competition potential of O. acetosella compared to L. luzuloides and F. sylvatica for the vegetation period of April to August, the carbon balances of the whole plants were estimated. These are compared with the dry weight increase. Among the three life forms of the rhizome geophyte (Oxalis), the hemicryptophyte (Luzula) and the phanerophyte (Fagus) there are striking differences in the use and in the distribution of the CO2 gain. L. luzuloides invests 65% of the net photosynthetic gain as dry weight increment (O. acetosella only 44%, F. sylvatica 40%). Moreover, the growth of L. luzuloides takes place primarily above ground with the establishment of new leaves. The relative proportion of the growth above ground to the total dry weight increment is for L. luzuloides 63%, as against 57% in F. sylvatica and only 42% in O. acetosella. In respect to the total carbon balance the better use of the CO2 gain gives L. luzuloides in this habitat a higher competition potential than the photosynthetically more active O. acetosella. The constitutional differences in the photosynthetic activity are compensated for by the distribution of the assimilates to shoot and root.