Mesophyll conductance and reaction-diffusion models for CO2 transport in C3 leaves; needs, opportunities and challenges

Abstract

One way to increase potential crop yield could be increasing mesophyll conductance g_m. This variable determines the difference between the CO₂ partial pressure in the intercellular air spaces (C_i) and that near Rubisco (C_c). Various methods can determine g_m from gas exchange measurements, often combined with measurements of chlorophyll fluorescence or carbon isotope discrimination. g_m lumps all biochemical and physical factors that cause the difference between C_c and C_i. g_m appears to vary with C_i. This variability indicates that g_m does not satisfy the physical definition of a conductance according to Fick's first law and is thus an apparent parameter. Uncertainty about the mechanisms that determine g_m can be limited to some extent by using analytical models that partition g_m into separate conductances. Such models are still only capable of describing the CO₂ diffusion pathway to a limited extent, as they make implicit assumptions about the position of mitochondria in the cells, which affect the re-assimilation of (photo)respired CO₂. Alternatively, reaction-diffusion models may be used. Rather than quantifying g_m, these models explicitly account for factors that affect the efficiency of CO₂ transport in the mesophyll. These models provide a better mechanistic description of the CO₂ diffusion pathways than mesophyll conductance models. Therefore, we argue that reaction-diffusion models should be used as an alternative to mesophyll conductance models, in case the aim of such a study is to identify traits that can be improved to increase g_m.

Keywords: 3D models; C(3) plants; CO(2); Mesophyll conductance; Photosynthesis; Reaction-diffusion models.