Abstract

The classical definition of mesophyll conductance (gm) represents an apparent parameter (gm,app) as it places (photo)respired CO2 at the same compartment where the carboxylation by Rubisco takes place. Recently, Tholen and co-workers developed a framework, in which gm better describes a physical diffusional parameter (gm,dif). They partitioned mesophyll resistance (rm,dif = 1/gm,dif) into two components, cell wall and plasmalemma resistance (rwp) and chloroplast resistance (rch), and showed that gm,app is sensitive to the ratio of photorespiratory (F) and respiratory (Rd) CO2 release to net CO2 uptake (A): gm,app = gm,dif/[1 + ω(F + Rd)/A], where ω is the fraction of rch in rm,dif. We herein extend the framework further by considering various scenarios for the intracellular arrangement of chloroplasts and mitochondria. We show that the formula of Tholen et al. implies either that mitochondria, where (photo)respired CO2 is released, locate between the plasmalemma and the chloroplast continuum or that CO2 in the cytosol is completely mixed. However, the model of Tholen et al. is still valid if ω is replaced by ω(1−σ), where σ is the fraction of (photo)respired CO2 that experiences rch (in addition to rwp and stomatal resistance) if this CO2 is to escape from being refixed. Therefore, responses of gm,app to (F + Rd)/A lie somewhere between no sensitivity in the classical method (σ =1) and high sensitivity in the model of Tholen et al. (σ =0).

Highlights

  • The biochemical C­ 3 photosynthesis model of Farquhar, von Caemmerer and Berry (1980), the FvCB model hereafter, has been widely used to interpret leaf physiology from gas exchange measurements

  • According to Fick’s diffusion law, gm can be expressed as follows: gm = A∕(Ci − Cc) where Ci is the partial pressure of ­CO2 at the intercellular air spaces

  • We introduced the parameter σ for defining the fraction of respired ­CO2 molecules that have to experience all rch, in addition to rwp and rsc, if these ­CO2 molecules are to escape from being refixed. σ has a value between 0 and 1, depending on the arrangement of organelles within mesophyll cells, i.e. (1) the relative position of chloroplasts and mitochondria and (2) the size of the gaps between chloroplasts

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Summary

Introduction

The biochemical C­ 3 photosynthesis model of Farquhar, von Caemmerer and Berry (1980), the FvCB model hereafter, has been widely used to interpret leaf physiology from gas exchange measurements. Tholen et al (2012) showed, based on their model framework, that the fraction of (photo)respired ­CO2 that is refixed by Rubisco can be quantified using the resistance components.

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