Seedlings of seven temperate tree species (Acer pseudoplatanus L., Betula pendula Roth, Fagus sylvatica L., Fraxinus excelsior L., Juglans regia L., Quercus petraea Matt. Liebl. and Quercus robur L.) were grown in a nursery under neutral filters transmitting 45% of incident global irradiance. During the second or third year of growth, leaf photosynthetic capacity (i.e., maximal carboxylation rate, Vcmax, maximal photosynthetic electron transport rate, Jmax, and dark respiration, Rd) was estimated for five leaves from each species at five or six leaf temperatures (10, 18, 25, 32, 36 and 40 degrees C). Values of Vcmax and Jmax were obtained by fitting the equations of the Farquhar model on response curves of net CO2 assimilation (A) to sub-stomatal CO2 mole fraction (ci), at high irradiance. Primary parameters describing the kinetic properties of Rubisco (specificity factor, affinity for CO2 and for O2, and their temperature responses) were taken from published data obtained with spinach and tobacco, and were used for all species. The temperature responses of Vcmax and Jmax, which were fitted to a thermodynamic model, differed. Mean values of Vcmax and Jmax at a reference temperature of 25 degrees C were 77.3 and 139 micromol m(-2) s(-1), respectively. The activation energy was higher for Vcmax than for Jmax (mean values of 73.1 versus 57.9 kJ mol(-1)) resulting in a decrease in Jmax/Vcmax ratio with increasing temperature. The mean optimal temperature was higher for Vcmax than for Jmax (38.9 versus 35.9 degrees C). In addition, differences in these temperature responses were observed among species. Temperature optima ranged between 35.9 and above 45 degrees C for Vcmax and between 31.7 and 43.3 degrees C for Jmax, but because of data scatter and the limited range of temperatures tested (10 to 40 degrees C), there were few statistically significant differences among species. The optimal temperature for Jmax was highest in Q. robur, Q. petraea and J. regia, and lowest in A. pseudoplatanus and F. excelsior. Measurements of chlorophyll a fluorescence revealed that the critical temperature at which basal fluorescence begins to increase was close to 47 degrees C, with no difference among species. These results should improve the parameterization of photosynthesis models, and be of particular interest when adapted to heterogeneous forests comprising mixtures of species with diverse ecological requirements.

The model RATP (radiation absorption, transpiration and photosynthesis) is presented. The model was designed to simulate the spatial distribution of radiation and leaf-gas exchanges within vegetation canopies as a function of canopy structure, canopy microclimate within the canopy and physical and physiological leaf properties. The model uses a three-dimensional (3D) representation of the canopy (i.e. an array of 3D cells, each characterized by a leaf area density). Radiation transfer is computed by a turbid medium analogy, transpiration by the leaf energy budget approach, and photosynthesis by the Farquhar model, each applied for sunlit and shaded leaves at the individual 3D cell-scale. The model typically operates at a 20-30 min time step. The RATP model was applied to an isolated, 20-yearold walnut tree grown in the field. The spatial distribution of wind speed, stomatal response to environmental variables, and light acclimation of leaf photosynthetic properties were taken into account. Model outputs were compared with data acquired in the field. The model was shown to simulate satisfactorily the intracrown distribution of radiation regime, transpiration and photosynthetic rates, at shoot or branch scales.

The variations of stem diameter as they can be accurately measured by Linear Variable Differential Transformer (LVDT) reflect the addition of four components: irreversible radial growth, reversible living-cell dehydration/rehydration, thermal expansion and contraction, and expansion of dead conducting elements due to the increase and relaxation of internal tensions. The correct interpretation of LVDT signals, with respect to the practical applications, should make an exact distinction between these four components. This paper describes a set of two experiments with potted hybrid walnut trees. Double girdling, water stress, and duration of the day versus night periods were used in the phytotron as experimental factors to induce variations of the carbon and water status of plant tissues. The latter were assessed, respectively, by water potential and transpiration, and by local stem respiration and carbohydrate content. The results are interpreted in terms of carbon or water limitation effects on stem diameter variations where radial growth and tissue elasticity could be distinguished. Moreover, they suggest no or very low involvement of CO2 originating from a distance, i.e. carried by the transpirational flux of xylem sap, in the total stem CO2 efflux rate.

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The biochemically based leaf photosynthesis model proposed by Farquhar et al. (1980) and the stomatal conductance model proposed by Jarvis (1976) were parameterized for walnut. Responses of photosynthesis to CO(2) and irradiance were used to determine the key parameters of the photosynthesis model. Concurrently, stomatal conductance responses to leaf irradiance (Q), leaf temperature (T(l)), water vapor pressure deficit at the leaf surface (D), and air CO(2) concentration at the leaf surface (C(s)) were used to parameterize the stomatal conductance model. To test the generality of the model parameters, measurements were made on leaves from a 20-year-old tree growing in the field, and from sunlit and shaded greenhouse-grown seedlings. The three key parameters of the photosynthesis model (maximum carboxylation rate V(cmax), electron transport capacity J(max), and dark respiration rate R(d)) and the key parameter of the conductance model (reference stomatal conductance, g(sref)) were linearly correlated with the amount of leaf nitrogen per unit leaf area. Unique relationships could be used to describe nitrogen effects on these parameters for leaves from both the tree and the seedlings. Our data allowed separation of the effects of increasing total photosynthetic apparatus per unit leaf area from the effects of partitioning nitrogen among different pools of this apparatus for foliage acclimation to leaf irradiance. Strong correlations were found between stomatal conductance g(s) and Q, D and C(s), whereas the relationship between g(s) and T(l) was weak. Based on these parameterizations, the model adequately predicted leaf photosynthesis and stomatal conductance when tested with an independent set of data obtained for the tree and seedlings. Total light-driven electron flows derived from chlorophyll fluorescence data obtained at different leaf temperatures were consistent with values computed by the model. The model was also tested with branch bag data acquired from a three-year-old potted walnut tree. Despite a relatively large variance between observed and simulated values, the model predicted stomatal conductance and photosynthesis reasonably well at the branch scale. The results indicate that the photosynthesis-conductance model developed here is robust and can be applied to walnut trees and seedlings under various environmental conditions where water is non-limiting.