RESUMO
To examine the predictability of leaf physiology and biochemistry from light gradients within canopies, we measured photosynthetic light-response curves, leaf mass per area (LMA) and concentrations of nitrogen, phosphorus and chlorophyll at 15-20 positions within canopies of three conifer species with increasing shade tolerance, ponderosa pine [Pinus ponderosa (Laws.)], Douglas fir [Pseudotsuga menziesii (Mirb.) Franco], and western hemlock [Tsuga heterophylla (Raf.) Sarg.]. Adjacent to each sampling position, we continuously monitored photosynthetically active photon flux density (PPFD) over a 5-week period using quantum sensors. From these measurements we calculated FPAR: integrated PPFD at each sampling point as a fraction of full sun. From the shadiest to the brightest canopy positions, LMA increased by about 50% in ponderosa pine and 100% in western hemlock; Douglas fir was intermediate. Canopy-average LMA increased with decreasing shade tolerance. Most foliage properties showed more variability within and between canopies when expressed on a leaf area basis than on a leaf mass basis, although the reverse was true for chlorophyll. Where foliage biochemistry or physiology was correlated with FPAR, the relationships were non-linear, tending to reach a plateau at about 50% of full sunlight. Slopes of response functions relating physiology and biochemistry to ln(FPAR) were not significantly different among species except for the light compensation point, which did not vary in response to light in ponderosa pine, but did in the other two species. We used the physiological measurements for Douglas fir in a model to simulate canopy photosynthetic potential (daily net carbon gain limited only by PPFD) and tested the hypothesis that allocation of carbon and nitrogen is optimized relative to PPFD gradients. Simulated photosynthetic potential for the whole canopy was slightly higher (<10%) using the measured allocation of C and N within the canopy compared with no stratification (i.e., all foliage identical). However, there was no evidence that the actual allocation pattern was optimized on the basis of PPFD gradients alone; simulated net carbon assimilation increased still further when even more N and C were allocated to high-light environments at the canopy top.
RESUMO
The ecosystem-level carbon uptake and respiration were measured under different CO2 concentrations in the tropical rainforest and the coastal desert of Biosphere 2, a large enclosed facility. When the mesocosms were sealed and subjected to step-wise changes in atmospheric CO2 between daily means of 450 and 900 µmol mol-1, net ecosystem exchange (NEE) of CO2 was derived using the diurnal changes in atmospheric CO2 concentrations. The step-wise CO2 treatment was effectively replicated as indicated by the high repeatability of NEE measurements under similar CO2 concentrations over a 12-week period. In the rainforest mesocosm, daily NEE was increased significantly by the high CO2 treatments because of much higher enhancement of canopy CO2 assimilation relative to the increase in the nighttime ecosystem respiration under high CO2. Furthermore, the response of daytime NEE to increasing atmospheric CO2 in this mesocosm was not linear, with a saturation concentration of 750 µmol mol-1. In the desert mesocosm, a combination of a reduction in ecosystem respiration and a small increase in canopy CO2 assimilation in the high CO2 treatments also enhanced daily NEE. Although soil respiration was not affected by the short-term change in atmospheric CO2 in either mesocosm, plant dark respiration was increased significantly by the high CO2 treatments in the rainforest mesocosm while the opposite was found in the desert mesocosm. The high CO2 treatments increased the ecosystem light compensation points in both mesocosms. High CO2 significantly increased ecosystem radiation use efficiency in the rainforest mesocosm, but had a much smaller effect in the desert mesocosm. The desert mesocosm showed much lower absolute response in NEE to atmospheric CO2 than the rainforest mesocosm, probably because of the presence of C4 plants. This study illustrates the importance of large-scale experimental research in the study of complex global change issues.
