Revisiting carbon isotope discrimination in C-3 plants shows respiration rules when photosynthesis is low

An updated carbon isotope discrimination model of diffusion of CO2 inside photosynthetic tissues is derived, treating the carbon pools independently. The modelled values for diffusion allow discussion of CO2 movement inside the mesophyll.Stable isotopes are commonly used to study the diffusion of CO2 within photosynthetic plant tissues. The standard method used to interpret the observed preference for the lighter carbon isotope in C-3 photosynthesis involves the model of Farquhar et al., which relates carbon isotope discrimination to physical and biochemical processes within the leaf. However, under many conditions the model returns unreasonable results for mesophyll conductance to CO2 diffusion (g(m)), especially when rates of photosynthesis are low. Here, we re-derive the carbon isotope discrimination model using modified assumptions related to the isotope effect of mitochondrial respiration. In particular, we treat the carbon pool associated with respiration as separate from the pool of primary assimilates. We experimentally test the model by comparing g(m) values measured with different CO2 source gases varying in their isotopic composition, and show that our new model returns matching g(m) values that are much more reasonable than those obtained with the previous model. We use our results to discuss CO2 diffusion properties within the mesophyll.