Toward extension of a single tree functional–structural model of Scots pine to stand level: effect of the canopy of randomly distributed, identical trees on development of tree structure

We examined the effects of structural and physiological acclimation on the photosynthetic efficiency of Scots pine (Pinus sylvestris L.) shoots. We estimated daily light interception (DLI) and photosynthesis (DPHOT) of a number of sample shoots situated at different positions in the canopy. Photosynthetic efficiency (epsilon) was defined as the ratio of DPHOT to the potential daily light interception (DLI(ref)) defined as the photosynthetically active radiation (PAR) intercepted per unit area of a sphere at the shoot location. To calculate DLI(ref), DLI and DPHOT, the radiation field surrounding a shoot in the canopy was first modeled using simulated directional distributions of incoming PAR on a clear and an overcast day, and estimates of canopy gap fraction in different directions provided by hemispherical photographs. A model of shoot geometry and measured data on shoot structure and photosynthetic parameters were used to simulate the distribution of PAR irradiance on the needle surface area of the shoot. Photosynthetic efficiency (epsilon) was separated into light-interception efficiency (epsilon(I) = DLI/DLI(ref)) and conversion efficiency (epsilon(PHOT) = DPHOT/DLI). This allowed us to quantify separately the effect of structural acclimation on the efficiency of photosynthetic light capture (epsilon(l)), and the effect of physiological acclimation on conversion efficiency (epsilon(PHOT)). The value of epsilon increased from the top to the bottom of the canopy. The increase was largely explained by structural acclimation (higher epsilon(I)) of the shade shoots. The value of epsilon(PHOT) of shade foliage was similar to that of sun foliage. Given these efficiencies, the clear-day value of DPHOT for a sun shoot transferred to shade was only half that of a shade shoot at its original position. The method presented here provides a tool for quantitatively estimating the role of acclimation in total canopy photosynthesis.

Trees have been increasingly considered as modular organisms, with individual shoots forming autonomous units that respond semi-independently to their surrounding environment. However, there is evidence for fairly strict hormonal control of tree crown development. Studies on the hydraulic architecture of trees suggest a closer functional connection between shoots and crown development than is postulated by the theory of branch autonomy. We studied how shoot growth pattern influences growth and crown architecture in young Scots pine trees simulated by the LIGNUM model assuming that (a) the growth of a shoot mainly depends on its light climate and (b) the growth of a shoot is influenced by its position within the crown. We determined shoot position within the crown based on a recently developed vigor index. The vigor index compares the relative axis cross-sectional area from the base of the tree to each shoot and gives a value of 1 to the pathway of the greatest cross-sectional area. All other shoots attain values between 0 and 1 depending on their cross-sectional areas and the cross-sectional areas of the branches leading there from the main axis. The shoot light climate is characterized by annually intercepted photosynthetically active radiation. We compared the results from simulations (a) and (b) against an independent data set. The addition of a within-shoot position index (the vigor index) to our simulation (simulation b) resulted in a more realistic tree form than that obtained with simulation (a) alone. We discuss the functional significance of the results as well as the possibilities of using an index of shoot position in simulations of crown architecture.