Down-regulation of photosynthesis and its relationship with changes in leaf N allocation and N availability after long-term exposure to elevated CO2 concentration

Abstract

Down-regulation of photosynthesis under elevated CO₂ (eCO₂) concentrations could be attributed to the depletion of nitrogen (N) availability after long-term exposure to eCO₂ (progressive nitrogen limitation, PNL) or leaf N dilutions due to excessive accumulation of nonstructural carbohydrates. To determine the mechanism underlying this down-regulation, we investigated N availability, photosynthetic characteristics, and N allocation in leaves of Pinus densiflora (shade-intolerant species, evergreen tree), Fraxinus rhynchophylla (intermediate shade-tolerant species, deciduous tree), and Sorbus alnifolia (shade-tolerant species, deciduous tree). The three species were grown under three different CO₂ concentrations in open-top chambers, i.e., ambient 400 ppm (aCO₂); ambient × 1.4, 560 ppm (eCO₂1.4); and ambient × 1.8, 720 ppm (eCO₂1.8), for 11 years. Unlike previous studies that addressed PNL, after 11 years of eCO₂ exposure, N availability remained higher under eCO₂1.8, and chlorophyll and photosynthetic N use efficiency increased under eCO₂. In the case of nonstructural carbohydrates, starch and soluble sugar showed significant increases under eCO₂. The maximum carboxylation rate, leaf N per mass (N_mass), and ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) were low under eCO₂1.8. The ratio of RuBP regeneration to the carboxylation rate as well as that of chlorophyll N to Rubisco N increased with CO₂ concentrations. Based on the reduction in N_mass (not in N_area) that was diluted by increase in nonstructural carbohydrate, down-regulation of photosynthesis was found to be caused by the dilution rather than PNL. The greatest increases in chlorophyll under eCO₂ were observed in S. alnifolia, which was the most shade-tolerant species. This study could help provide more detailed, mechanistically based processes to explain the down-regulation of photosynthesis by considering two hypotheses together and showed N allocation seems to be flexible against changes in CO₂ concentration.

Keywords: Elevated CO2; Leaf nitrogen content; Nitrogen allocation; Nonstructural carbohydrates; Photosynthesis; Rubisco.