Summary
In the present chapter, we review the current state-of-the-art of knowledge on mesophyll (internal) CO2 diffusion conductance of photosynthetic tissues (for simplification, g m). We show that, despite concerns regarding the methodological approaches currently used for its estimation, a large and consistent body of evidence has accumulated showing that g m is finite and significantly limiting for photosynthesis, as well as being highly variable among photosynthetic organisms and in response to environmental changes. Part of this variation results from different anatomies of the photosynthetic tissues, with a particularly strong influence of chloroplast distribution and cell wall thickness. Besides these, it appears that a biochemical modulation of g m also occurs, likely involving aquaporins and, possibly, carbonic anhydrases and other metabolic components.
Further efforts are needed in the near future to improve CO2 diffusion models, both for the estimation of g m and for the precise physiological understanding of the CO2 assimilation process in different plants, as well as to increase our knowledge of the mechanistic base for g m and its regulation.
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Abbreviations
- a :
-
fractionation factor due to diffusion through the air in the stomatal pore and between photosynthetic tissues
- ā :
-
fractionation factor due to diffusion through stomata, between photosynthetic tissues and boundary layers
- a b :
-
fractionation factor associated with diffusion through the air in the boundary layers
- a i :
-
combined fractionation factor for dissolution of CO2 and diffusion through the liquid phase
- ā t :
-
fractionation factor for the diffusion pathway through stomata and boundary layers corrected for ternary effects
- a t i :
-
combined fractionation factor for dissolution of CO2 and diffusion through the liquid phase corrected for ternary effects
- A n :
-
net rate of photosynthesis
- AtPIP :
-
plasma membrane aquaporins of Arabidopsis thaliana
- ABA:
-
abscisic acid
- AQP:
-
aquaporin
- ATP:
-
Adenosine triphosphate
- α :
-
leaf absorptance
- b :
-
net fractionation factor associated with Rubisco and PEPC
- b 3 :
-
fractionation factor associated with Rubisco
- b 4 :
-
fractionation factor associated with PEPC
- b t :
-
net fractionation factor associated with Rubisco and PEPC corrected for ternary effects
- β :
-
fraction of photons absorbed by photosystem II
- C a :
-
ambient CO2 concentration
- C b :
-
CO2 concentration just outside the stomatal pore
- C 0 :
-
CO2 concentration after fixation by Rubisco, i.e. 0
- C3 :
-
three-carbon organic acids
- C4 :
-
four-carbon organic acids
- C c :
-
chloroplast stroma CO2 concentration
- C i :
-
sub-stomatal CO2 concentration
- C i * :
-
intercellular CO2 concentration when the carboxylation rate equals the photorespiration rate
- C mc :
-
cytosolic CO2 concentration of a mesophyll cell
- C s :
-
surface CO2 concentration
- CA:
-
carbonic anhydrase
- CAM:
-
Crassulacean acid metabolism
- CCMs:
-
CO2 concentrating mechanisms
- CRDS:
-
cavity ringdown spectroscopy
- D a :
-
diffusion coefficient for CO2 in the gas phase
- D w :
-
diffusion coefficient for CO2 in the aqueous phase
- Δ:
-
discrimination against 13CO2
- Δi :
-
expected amount of discrimination against 13CO2 by the tissue enclosed in a gas-exchange cuvette if mesophyll conductance is assumed infinite and in the absence of any (photo)respiratory fractionation
- Δo :
-
observed amount of discrimination against 13CO2 by the tissue enclosed in a gas-exchange cuvette
- δ13C-CO2 :
-
carbon isotopic composition of CO2
- e :
-
fractionation factor due to mitochondrial respiration in the light
- e t :
-
fractionation factor due to mitochondrial respiration in the light corrected for ternary effects
- E :
-
transpiration rate
- f :
-
fractionation factor for photorespiration
- f ias :
-
fraction of mesophyll volume occupied by intercellular air space
- f t :
-
fractionation factor for photorespiration corrected for ternary effects
- F :
-
photorespiration rate
- F s’:
-
steady state fluorescence in the light
- F m’:
-
maximal fluorescence in the light during a short saturating pulse of light
- ΦCO2 :
-
quantum yield derived from CO2 exchange in the light-limited region
- ΦPS II :
-
photochemical yield of photosystem II
- g ac :
-
total conductance to CO2 through stomata and boundary layers
- g b :
-
boundary layer conductance to CO2
- g i :
-
conductance of a given component of the diffusion pathway
- g ias :
-
gas-phase conductance between the sub-stomatal cavities and the outer surface of cell walls
- g liq :
-
liquid-phase conductance between the outer surface of the cell walls and the site of carboxylation in the chloroplast stroma
- g m :
-
mesophyll conductance to CO2
- g mA :
-
mesophyll conductance to CO2 estimate based on physical models and the anatomical properties of the leaves
- g s :
-
stomatal conductance to CO2
- g t :
-
total conductance to CO2
- ias:
-
intercellular air space
- γ:
-
molar proportion of carbon fixed by PEPC
- γi :
-
dimensionless factor accounting for decrease of diffusion conductance in the cytosol and in the stroma compared with free diffusion in water
- Г*:
-
chloroplastic CO2 concentration when the carboxylation rate equals the photorespiration rate
- H :
-
Henry’s law constant for dissolution of CO2 in water
- J :
-
electron transport rate
- J F :
-
whole chain electron transport rate derived from fluorescence measurements
- J C :
-
electron transport rate related to gas-exchange measurements
- J max :
-
maximum rate of electron transport
- K leaf :
-
leaf hydraulic conductance
- λ :
-
cyclic-pseudocyclic electron flow coefficient
- L i :
-
diffusion path length
- L ias :
-
diffusion path length in the gas phase
- NAD:
-
nicotinamide adenine dinucleotide
- NADPH:
-
reduced nicotinamide adenine dinucleotide phosphate
- OA-ICOS:
-
Off-Axis Integrated Cavity Output Spectroscopy
- p i :
-
coefficient related with the electron transport stoichiometry associated with the regeneration of RuBP
- PAM:
-
pulse amplitude modulation
- PEPC:
-
phosphoenolpyruvate carboxylase
- PIP:
-
plasma membrane intrinsic protein
- PNUE:
-
photosynthetic nitrogen use efficiency
- PPFD:
-
photosynthetically active photon flux density incident on the leaf
- PS:
-
photosystem
- Q10 :
-
scaling factor representing the relative increase in reaction rate over a 10°C temperature range at a particular temperature
- R :
-
gas constant
- r :
-
CO2 diffusion resistance (or sum of serial diffusion resistance)
- r b :
-
diffusion resistance of the boundary to CO2
- r ch :
-
diffusion resistance of the double membranes of the chloroplasts and the stroma
- r i :
-
diffusion resistance of a given component of the diffusion pathway
- r s :
-
diffusion resistance of the stomata to CO2
- r wp :
-
diffusion resistance of cell wall and plasma membrane
- R d :
-
rate of non-photorespiratory CO2 release in the light
- R n :
-
rate of CO2 emission in the dark
- RuBP:
-
ribulose-1,5-bisphosphate
- Rubisco:
-
ribulose-1,5-bisphosphate carboxylase-oxygenase
- S c :
-
chloroplast surface exposed to the intercellular airspace
- ς :
-
diffusion path tortuosity
- t :
-
correction factor to account for ternary effects
- T cw :
-
cell wall thickness
- T k :
-
absolute temperature
- TCAP:
-
tricarboxylic acid pathway
- T-DNA:
-
transfer DNA
- TDLAS:
-
tunable-diode laser absorption spectroscopy
- V c :
-
carboxylation rate
- V c,max :
-
maximum rate of carboxylation
- VPD:
-
vapor pressure deficit
- WUE:
-
water use efficiency
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Acknowledgments
This work was supported partially by the Plan Nacional, Spain (contract CTM2014-53902-C2-1-P from the Spanish Ministry of Economy and Competitiveness – MINECO – and the ERDF – FEDER) awarded to Jaume Flexas and by the Conselleria d’Educació, Cultura i Universitats (Govern de les Illes Balears) and European Social Fund, predoctoral fellowship FPI/1700/2014, awarded to Marc Carriquí. Dongliang Xiong thanks the China Scholarship Council (CSC) for the funding of joint PhD training. Francisco Javier Cano thanks funding by the Australian Research Council Centre of Excellence for Translational Photosynthesis (CE1401000015).
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Flexas, J. et al. (2018). CO2 Diffusion Inside Photosynthetic Organs. In: Adams III, W., Terashima, I. (eds) The Leaf: A Platform for Performing Photosynthesis. Advances in Photosynthesis and Respiration, vol 44. Springer, Cham. https://doi.org/10.1007/978-3-319-93594-2_7
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