Photosynthesis Research

, Volume 117, Issue 1–3, pp 45–59 | Cite as

Diffusional conductances to CO2 as a target for increasing photosynthesis and photosynthetic water-use efficiency

  • Jaume Flexas
  • Ülo Niinemets
  • Alexander Gallé
  • Margaret M. Barbour
  • Mauro Centritto
  • Antonio Diaz-Espejo
  • Cyril Douthe
  • Jeroni Galmés
  • Miquel Ribas-Carbo
  • Pedro L. Rodriguez
  • Francesc Rosselló
  • Raju Soolanayakanahally
  • Magdalena Tomas
  • Ian J. Wright
  • Graham D. Farquhar
  • Hipólito Medrano


A key objective for sustainable agriculture and forestry is to breed plants with both high carbon gain and water-use efficiency (WUE). At the level of leaf physiology, this implies increasing net photosynthesis (A N) relative to stomatal conductance (g s). Here, we review evidence for CO2 diffusional constraints on photosynthesis and WUE. Analyzing past observations for an extensive pool of crop and wild plant species that vary widely in mesophyll conductance to CO2 (g m), g s, and foliage A N, it was shown that both g s and g m limit A N, although the relative importance of each of the two conductances depends on species and conditions. Based on Fick’s law of diffusion, intrinsic WUE (the ratio A N/g s) should correlate on the ratio g m/g s, and not g m itself. Such a correlation is indeed often observed in the data. However, since besides diffusion A N also depends on photosynthetic capacity (i.e., V c,max), this relationship is not always sustained. It was shown that only in a very few cases, genotype selection has resulted in simultaneous increases of both A N and WUE. In fact, such a response has never been observed in genetically modified plants specifically engineered for either reduced g s or enhanced g m. Although increasing g m alone would result in increasing photosynthesis, and potentially increasing WUE, in practice, higher WUE seems to be only achieved when there are no parallel changes in g s. We conclude that for simultaneous improvement of A N and WUE, genetic manipulation of g m should avoid parallel changes in g s, and we suggest that the appropriate trait for selection for enhanced WUE is increased g m/g s.


Photosynthesis Water-use efficiency Stomatal conductance Mesophyll conductance Meta-analysis 



This work was partly supported by the Plan Nacional, Spain, contracts AGL2002-04525-CO2-01 (H.M.), BFU2008-1072-E/BFI and BFU2011-23294 (M.R.-C. and J.F.), AGL2009-07999 (J.G.), and MTM2009-07165 (F.R.); the Foundation for Research, Science and Technology, New Zealand, contract C09X0701 (M.M.B); the Australian Research Council, contract FT0992063 (M.M.B), FT100100910 (I.J.W), and DP1097276 (G.D.F.); the Estonian Ministry of Science and Education, (institutional grant IUT-8-3); the European Commission through the European Regional Fund (the Center of Excellence in Environmental Adaptation) (Ü.N.); and a collaboration project between the Estonian Academy of Sciences and the Spanish CSIC (H.M., Ü.N.).

Supplementary material

11120_2013_9844_MOESM1_ESM.xls (89 kb)
Data compilation in different species and conditions (see Online Resource 3 for a complete list of the references used). Supplementary material 1 (XLS 89 kb)
11120_2013_9844_MOESM2_ESM.doc (46 kb)
Complete list of references from which data in Online Resource 2 and 4 were compiled. Supplementary material 2 (DOC 46 kb)
11120_2013_9844_MOESM3_ESM.xls (34 kb)
Data compilation for specific genetic manipulations. Supplementary material 3 (XLS 34 kb)
11120_2013_9844_MOESM4_ESM.doc (524 kb)
The relationship between g m/g s and g m in a multi-species dataset. Data and symbols as in Fig. 1. Supplementary material 4 (DOC 523 kb)


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Copyright information

© Springer Science+Business Media Dordrecht 2013

Authors and Affiliations

  • Jaume Flexas
    • 1
  • Ülo Niinemets
    • 2
  • Alexander Gallé
    • 1
    • 3
  • Margaret M. Barbour
    • 4
  • Mauro Centritto
    • 5
  • Antonio Diaz-Espejo
    • 6
  • Cyril Douthe
    • 1
  • Jeroni Galmés
    • 1
  • Miquel Ribas-Carbo
    • 1
  • Pedro L. Rodriguez
    • 7
  • Francesc Rosselló
    • 8
  • Raju Soolanayakanahally
    • 9
  • Magdalena Tomas
    • 1
  • Ian J. Wright
    • 10
  • Graham D. Farquhar
    • 11
  • Hipólito Medrano
    • 1
  1. 1.Research Group on Plant Biology under Mediterranean Conditions, Departament de BiologiaUniversitat de les Illes BalearsPalma de MallorcaSpain
  2. 2.Institute of Agricultural and Environmental SciencesEstonian University of Life SciencesTartuEstonia
  3. 3.Bayer CropScience NVZwijnaardeBelgium
  4. 4.Faculty of Agriculture, Food and Natural ResourcesThe University of SydneyNarellanAustralia
  5. 5.Institute for Plant ProtectionNational Research CouncilSesto FiorentinoItaly
  6. 6.Instituto de Recursos Naturales y Agrobiología de Sevilla (IRNAS, CSIC)Irrigation and Crop Ecophysiology GroupSevillaSpain
  7. 7.Instituto de Biología Molecular y Celular de PlantasConsejo Superior de Investigaciones Científicas-Universidad Politécnica de ValenciaValenciaSpain
  8. 8.Computational Biology and Bioinformatics Research Group, Departament de Ciències Matemàtiques i InformàticaUniversitat de les Illes BalearsPalma de MallorcaSpain
  9. 9.Science and Technology BranchAgriculture and Agri-Food Canada, Indian HeadSaskatchewanCanada
  10. 10.Department of Biological SciencesMacquarie UniversityNorth RydeAustralia
  11. 11.Research School of BiologyThe Australian National UniversityCanberraAustralia

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