Simulations of the effects of shoot structure and orientation on vertical gradients in intercepted light by conifer canopies.

Coniferous tree canopies typically carry more leaf area than is necessary to intercept most of the incoming light. I postulated that an excessively large leaf area will reduce net productivity at tree level, unless the net photosynthetic production of the most shaded shoots in the canopy remains positive. The hypothesis tested was that a coniferous tree canopy maintains a large productive leaf area by increasing the efficiency of light capture as the available light decreases. The light interception efficiency of a shoot was quantified by the ratio of shoot silhouette area to total needle surface area (STAR). The STAR depends on shoot geometry and varies with shoot orientation relative to the direction of light. Shade shoots have a larger STAR and, in particular, higher values of STAR(max) than sun shoots. In addition, shade shoots tend to be horizontally inclined, which may increase the advantage of a large STAR(max) in the lower canopy, where radiation is incident from angles closer to the zenith. Adaptation to shade (changes in STAR and shoot orientation) was described on the basis of empirical data for several coniferous species, and the vertical gradient of seasonal light interception by shoots was simulated assuming different adaptive strategies. Simulations were performed at two latitudes, to account for differences in the amount and directional distribution of light during the growing season. Results support the hypothesis that increases in STAR, shoot zenith angle and shoot asymmetry (flatness) with shading increase the efficiency of light interception by deeply shaded shoots. However, because competition for light among shoots increases progressively as soon as shade acclimation occurs, there cannot exist a deep layer of shade shoots, such that the net productivity of each shoot remains positive (i.e., irradiance is above the compensation point). Therefore, if maximization of productive leaf area is the goal, the optimal strategy is to maintain an inefficient deep canopy and to increase light interception efficiency only when shading becomes severe.