Predicting leaf gas exchange and δ C responses to the past 30000 years of global environmental change.

Theoretical developments in our understanding of leaf gas exchange processes and carbon isotope composition (δ C) mean that it should now be possible to model their responses to global environmental change. Such a model would be of use for process-based interpretations of historical changes in leaf δ C and for understanding the global stable carbon isotope balance. This paper describes the development and validation of a model towards this aim. The resulting model is used to simulate changes in leaf photosynthesis, stomatal conductance and δ C of limber pine (Pinus flexilis) in response to the past 30000 y of global environmental change. The predictions of needle δ C are in line with reported measurements of δ C from fossilized Pinus flexilis needles preserved in packrat middens in western USA. Leaf gas exchange predictions show that the increased water use efficiency (WUE) of these trees growing in present-day environments, relative to the past, was brought about through an increase in photosynthetic rates and a decrease in stomatal conductance. This contrasts with the explanation of the recent (past 200 y) increase in the WUE of temperate and Mediterranean ecosystems inferred from δ C measurements which are predicted by the model to have arisen largely by a decrease in stomatal conductance in response to increases in the concentration of atmospheric CO since the pre-industrial era. The model as described offers the potential to contribute to our understanding of vegetation effects on the global carbon isotope balance during the glacial periods, and therefore to provide a further constraint on the carbon cycle models used to explain the low concentrations of atmospheric CO at these times.