Dynamics of leaf area and nitrogen in the canopy of an annual herb, Xanthium canadense.

We studied leaf area and nitrogen dynamics in the canopy of stands of an annual herb Xanthium canadense, grown at a high (HN)- and a low-nitorgen (LN) availability. Standing leaf area increased continuously through the vegetative growth period in the LN stand, or leveled off in the later stage in the HN stand. When scaled against standing leaf area, both production and loss rates of leaf area increased but with different patterns: the production rate was retarded, while the loss rate was accelerated, implying an upper limit of standing leaf area of the canopy. The rate of leaf-area production was higher in the HN than in the LN stand, which was caused by the higher rate of leaf production per standing leaf area as well as the greater standing leaf area in the HN stand. Although the rate of leaf-area loss was higher in the HN than in the LN stand, it was not significantly different between the two stands when compared at a common standing leaf area, suggesting involvement of light climate in determination of the leaf-loss rate. On the other hand, the rate of leaf-area loss was positively correlated with nitrogen demand for leaf area development across the two stands, suggesting that leaf loss was caused by retranslocation of nitrogen for construction of new leaves. A simple simulation model of leaf and nitrogen dynamics in the canopy showed that, at steady state, where the rate of leaf-area loss becomes equal to the production rate, the standing leaf area was still greater in the HN than in the LN stand. Similarly, when the uptake and loss of nitrogen are equilibrated, the standing nitrogen was greater in the HN than in the LN stand. These results suggest that leaf-area production is strongly controlled by nitrogen availability, while both nitrogen and light climate determine leaf-loss rates in the canopy.