20th century changes in carbon isotopes and water-use efficiency: tree-ring-based evaluation of the CLM4.5 and LPX-Bern models

Measurements of the stable carbon isotope ratio (δ¹³C) on annual tree rings offer new opportunities to evaluate mechanisms of variations in photosynthesis and stomatal conductance under changing CO₂ and climate conditions, especially in conjunction with process-based biogeochemical model simulations. The isotopic discrimination is indicative of the ratio between the CO₂ partial pressure in the intercellular cavities and the atmosphere (ci∕ca) and of the ratio of assimilation to stomatal conductance, termed intrinsic water-use efficiency (iWUE). We performed isotope-enabled simulations over the industrial period with the land biosphere module (CLM4.5) of the Community Earth System Model and the Land Surface Processes and Exchanges (LPX-Bern) dynamic global vegetation model. Results for C3 tree species show good agreement with a global compilation of δ¹³C measurements on leaves, though modeled ¹³C discrimination by C3 trees is smaller in arid regions than measured. A compilation of 76 tree-ring records, mainly from Europe, boreal Asia, and western North America, suggests on average small 20th century changes in isotopic discrimination and in ci∕ca and an increase in iWUE of about 27 % since 1900. LPX-Bern results match these century-scale reconstructions, supporting the idea that the physiology of stomata has evolved to optimize trade-offs between carbon gain by assimilation and water loss by transpiration. In contrast, CLM4.5 simulates an increase in discrimination and in turn a change in iWUE that is almost twice as large as that revealed by the tree-ring data. Factorial simulations show that these changes are mainly in response to rising atmospheric CO₂. The results suggest that the downregulation of ci∕ca and of photosynthesis by nitrogen limitation is possibly too strong in the standard setup of CLM4.5 or that there may be problems associated with the implementation of conductance, assimilation, and related adjustment processes on long-term environmental changes