Leaf shedding increases the photosynthetic rate of the canopy in N2-fixing and non-N2-fixing woody species

It has long been hypothesized that timing of leaf shedding is critical for plant fitness but there is little experimental evidence to support the hypothesis. According to an optimality theory, shedding of old leaves increases canopy photosynthesis despite some nitrogen (N) being lost as litterfall, when the ratio of daily photosynthesis to leaf N (N-use efficiency, epsilon) in old leaves, expressed as a fraction of epsilon in new leaves, becomes lower than the fraction of leaf N that is resorbed before shedding (R-N). This was shown to be true for N-poor plants but not for N-rich plants in a pot experiment; however, the use of planting pots imposes a variety of physical, chemical and biological constraints that could change the experimental results. Here we conducted a 3-year field survey in a cool temperate deciduous forest to examine whether Alnus sieboldiana Matsum. (N-2-fixing) and Carpinus tschonoskii Maxim. (non-N-2-fixing) shed their leaves to increase canopy photosynthesis in accord with the above criterion. These species often grow sympatrically and were chosen as representatives of N-rich and N-poor plants, respectively. Overall, daily photosynthesis decreased with leaf age, accompanied by small changes in leaf N, resulting in a decrease in epsilon. In both species, epsilon of leaves at shedding expressed as a fraction of epsilon in new leaves was nearly equal to R-N in all years, implying that the old leaves were shed to increase canopy photosynthesis. Our results, together with those of previous field surveys, suggested that the timing of leaf shedding is explained by N use in maximizing canopy photosynthesis across broad groups of species.