Chapter 4 C4 Photosynthesis: Kranz Forms and Single-Cell C4 in Terrestrial Plants

  • Gerald E. Edwards
  • Elena V. Voznesenskaya
Part of the Advances in Photosynthesis and Respiration book series (AIPH, volume 32)


Plants identified as having C4 photosynthesis have a C4 metabolic cycle with phosphoenolpyruvate carboxylase as the initial catalyst for fixation of atmospheric CO2, and a C4 acid decarboxylase (NADP-malic enzyme, NAD-malic enzyme, or phosphoenolpyruvate carboxykinase), which releases CO2 for fixation by the C3 cycle. Effective donation of CO2 to Rubisco minimizes competition by O2 and photorespiration, and thus increases photosynthesis under conditions where CO2 is limiting. To achieve this, fixation of atmospheric CO2 in the cytosol by phosphoenolpyruvate carboxylase must be separated from the donation of CO2 to Rubisco by the decarboxylation of C4 acids. In most documented C4 plants, this is accomplished through evolution of various forms of Kranz anatomy, with fixation of atmospheric CO2 in mesophyll cells and donation of CO2 from C4 acids to Rubisco in bundle sheath cells. In the family Chenopodiaceae, two alternative means of accomplishing this spatial separation evolved within individual photosynthetic cells, whereby one cytoplasmic compartment specializes in fixation of atmospheric CO2 in the carboxylation phase of the C4 cycle, and the other cytoplasmic compartment specializes in donating CO2 from C4 acids to Rubisco. In this chapter, biochemical and structural variations of Kranz anatomy in three major C4-containing families, Poaceae, Cyperaceae, and Chenopodiaceae, as well as other known forms for dicots, are summarized. Then, the phylogeny, biogeography, development, and structure-function relationships of the single-cell C4 systems are discussed in comparison to Kranz type C4 plants.


Bundle Sheath Bundle Sheath Cell Bundle Sheath Chloroplast Chlorenchyma Cell Mestome Sheath 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.



Bundle sheath(s);

Kranz cells

An inner layer of chlorenchyma cells specialized for C4 photosynthesis, irrespective of whether there is contact with vascular bundles (sometimes referred to as BS cells in C4 plants);




Mestome sheath(s);


NAD-malic enzyme;


NADP-malic enzyme;


Phosphoenolpyruvate carboxylase;


Phosphoenolpyruvate carboxykinase;


3-Phosphoglyceric acid;


Pyruvate,Pi dikinase;


Photosystem I;


Photosystem II;


Ribulose 1,5-bisphosphate;


Suberin lamella



This material is based upon work supported by the National Science Foundation under Grant Nos. IBN-0131098, IBN-0236959 and IBN-0641232, by Civilian Research and Development Foundation grants RB1-2502-ST-03 and RUB1-2829-ST-06, by Russian Foundation of Basic Research grant 05-04-49622 and 08-04-00936,and Bill and Melinda Gates Foundation to IRRI for C4 Rice Program. We also thank the Franceschi Microscopy and Imaging Center of Washington State University for use of their facilities and staff assistance, and C. Cody for plant growth management. The authors appreciate the helpful discussions with Dr. Nuria Koteyeva and the advice of two reviewers, especially in the sections devoted to grasses and sedges.


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© Springer Netherlands 2010

Authors and Affiliations

  1. 1.School of Biological SciencesWashington State UniversityPullmanUSA
  2. 2.Laboratory of Anatomy and MorphologyV.L. Komarov Botanical Institute of Russian Academy of SciencesSt. PetersburgRussia

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