Variation in light-intercepting area and photosynthetic rate of sun and shade shoots of two Picea species in relation to the angle of incoming light.

We investigated the effects of sun- and shade-shoot architecture on the photosynthetic rates of two Picea species by applying light from various angles in the laboratory. Compared with sun shoots, shade shoots were characterized by lower mass allocation per light-intercepting area, less leaf mass per shoot mass, less mutual shading among leaves and more efficient allocation of chlorophyll to photosynthesis. The shoot silhouette to total leaf area ratio (STAR(ϕ)) decreased with increasing shoot inclination angle (ϕ, the shoot axis angle relative to the projection plane) and was consistently higher for the shade shoots. Morphological and physiological characteristics of the shade shoots resulted in maximum rates of net photosynthesis at ϕ = 0° (P(max,0)) similar to that of the sun shoots when expressed on a leaf mass, total leaf area and chlorophyll basis. When the angle of incoming light was varied, P(max,ϕ) per total leaf area (P(max,ϕ )/A(T)) of the shade shoots increased linearly with increasing STAR(ϕ), while P(max,ϕ) per shoot silhouette area did not change. In contrast, the response of the sun shoots was non-linear, and an optimum angle of incoming light was determined. Our results suggest that shade-shoot morphology is adaptive for utilizing diffuse light incoming from various angles, while sun-shoot morphology is adaptive for avoiding the negative effects of strong direct radiation and for enhancing light diffusion into the canopy. We propose that the angle of incoming light should be taken into account when estimating photosynthetic rates of sun shoots of conifer trees in the field.