Interannual variation in leaf photosynthetic capacity during summer in relation to nitrogen, leaf mass per area and climate within a Fagus crenata crown on Naeba Mountain, Japan.

During the summers (July and August) of 2002-2005, we measured interannual variation in maximum carboxylation rate (V(cmax)) within a Fagus crenata Blume crown in relation to climate variables such as air temperature, daytime vapor pressure deficit (VPD) and daily photosynthetic photon flux, leaf nitrogen per unit area (N(a)) and leaf mass per unit area (LMA). Climatic conditions in the summers of 2002-2004 differed markedly, with warm and dry atmospheric conditions in 2002, cool, humid and cloudy conditions in 2003, and warm clear conditions in 2004. Conditions in summer 2005 were intermediate between those of summers 2002 and 2003, and similar to recent (8-year) means. In July, marked interannual variation in V(cmax) was mainly observed in leaves in the high-light environment (relative photon flux > 50%) within the crown. At the crown top, V(cmax) was about twofold higher in 2002 than in 2003, and V(cmax) values in 2004 and 2005 were intermediate between those in 2002 and 2003. In August, although interannual variation in V(cmax) among the years 2003, 2004 and 2005 was less, marked variation between 2002 and the other study years was evident. Multiple regression analysis of V(cmax) against the climate variables revealed that VPD of the previous 10-30 days had a significant influence on variability in V(cmax). Neither N(a), LMA nor leaf CO(2) conductance from the stomata to the carboxylation site explained the variability in V(cmax). Our results indicate that the long-term climatic response of V(cmax) should be considered when estimating forest carbon gain across the year.

Vertical, horizontal and azimuthal variations in leaf photosynthetic characteristics within a Fagus crenata crown in relation to light acclimation.

An understanding of spatial variations in gas exchange parameters in relation to the light environment is crucial for modeling canopy photosynthesis. We measured vertical, horizontal and azimuthal (north and south) variations in photosynthetic capacity (i.e., the maximum rate of carboxylation: Vcmax), nitrogen content (N), leaf mass per area (LMA) and chlorophyll content (Chl) in relation to relative photosynthetic photon flux (rPPF) within a Fagus crenata Blume crown. The horizontal gradient of rPPF was similar in magnitude to the vertical gradient of rPPF from the upper to the lower crown. The rPPF in the north quadrant of the crown was slightly lower than in the south quadrant. Nitrogen content per area (Narea), LMA and Vcmax were strictly proportional to rPPF, irrespective of the vertical direction, horizontal direction and crown azimuth, whereas nitrogen content per dry mass, Chl per area and photosynthetic capacity per dry mass (Vm) were fairly constant. Statistical analyses separating vertical trends from horizontal and azimuthal trends indicated that, although horizontal and vertical light acclimation of leaf properties were similar, there were two significant azimuthal variations: (1) Vcmax was lower in north-facing leaves than in south-facing leaves for a given Narea, indicating low photosynthetic nitrogen-use efficiency (PNUE) of north-facing leaves; and (2) Vcmax was lower in north-facing leaves than in south-facing leaves for a given LMA, indicating low Vm of the north-facing leaves. With respect to the low PNUE of the north-facing leaves, there were no significant azimuthal variations in leaf CO2 conductance from the stomata to the carboxylation site. Biochemical analysis indicated that azimuthal variations in nitrogen allocation to ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) and in nitrogen allocation between carboxylation (Rubisco and other Calvin cycle enzymes) and light harvesting machinery (Chl pigment-protein complexes) were not the main contributor to the difference in PNUE between north- and south-facing leaves. Lower specific activity of Rubisco may be responsible for the low PNUE of the north-facing leaves. Anatomical analysis indicated that not only high leaf density, which is compatible with a greater fraction of non-photosynthetic tissue, but also thick photosynthetic tissue contributed to the low Vm in the north-facing leaves. These azimuthal variations may need to be considered when modeling canopy photosynthesis based on the Narea-Vcmax or LMA-Vcmax relationship.