Chapter 5 Single-Cell C4 Photosynthesis in Aquatic Plants

  • George Bowes
Part of the Advances in Photosynthesis and Respiration book series (AIPH, volume 32)


In water with low free [CO2] a common strategy of submersed plants is to use HCO 3 , but some ­species utilize a C4 photosynthetic system that surprisingly lacks the Kranz dual-cell compartmentation of most terrestrial C4 plants. Instead, the C4 and C3 cycles are in the same cell, with phosphoenolpyruvate ­carboxylase (PEPC) and ribulose bisphosphate carboxylase–oxygenase (rubisco) sequestered in the cytosol and chloroplasts, respectively. Malate decarboxylation by NADP malic enzyme (NADP-ME) in the chloroplasts produces a chloroplastic CO2 concentrating mechanism (CCM). It occurs in the ­submersed monocots Hydrilla verticillata and Egeria densa (Hydrocharitaceae), and in these species it is facultative because low [CO2] induces a metabolic shift in the leaves from C3 to single-cell C4 photosynthesis. ­Submersed leaves of other species also perform single-cell C4 photosynthesis, including Sagittaria ­subulata (Alismataceae), the grasses Orcuttia californica and O. viscida (Poaceae), and the sedge ­Eleocharis acicularis. A marine macroalga (Udotea flabellum, Chlorophyta) and a diatom (Thalassiosira weissflogii) likewise show evidence of its occurrence, so it is not restricted to higher plants. The change from C3 to C4 photosynthetic gas exchange and pulse-chase characteristics is well documented in Hydrilla, along with enzyme kinetics and localization; high internal [CO2], and improved growth. ­Multiple isoforms of PEPC, NADP-ME and pyruvate orthophosphate dikinase (PPDK) exist in Hydrilla and Egeria, but specific forms, including hvpepc4, hvme1 and hvppdk1are up-regulated in the C4 leaves of Hydrilla and encode proteins with C4 photosynthetic characteristics. Interestingly, the photosynthetic hvpepc4 differs from its terrestrial C4 counterparts in lacking a “C4-signature” serine near the carboxy terminus. The C3 leaf must maximize CO2 conductance to rubisco, but as the C4 system is induced, ­chloroplast conductance is probably minimized to reduce leakage from the CCM. Further study of the facultative system of Hydrilla could determine if down-regulation of chloroplast-envelope aquaporins is involved in reducing CO2 conductance. Hydrilla and Egeria are in the ancient Hydrocharitaceae ­family, and can give insights into early C4 photosynthesis, which likely originated in water prior to its advent on land.


PEPC Activity Bundle Sheath Chloroplast Submerse Plant Submerse Species Submerse Leave 
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.









Blue native polyacrylamide gel electrophoresis;


Crassulacean acid metabolism;


Carbonic anhydrase;


CO2 concentrating mechanism;


-CO2 compensation point;


Dry weight;


Fresh weight;




Malate dehydrogenase;


NADP malic enzyme;


NAD malic enzyme;






Phosphoenolpyruvate carboxylase;


Phosphoenolpyruvate carboxykinase;


Photosynthetic nitrogen use efficiency;


Pyruvate orthophosphate dikinase;


Photosystem II;


Real-time polymerase chain reaction;


Ribulose bisphosphate carboxylase–­oxygenase;




Water use efficiency



The credit for the research that emanated from my laboratory goes to the many colleagues and students with whom it has been my privilege and pleasure to work over the past 35 years. Financial support has been provided by the United States Department of Agriculture, National Research Initiatives Competitive Grants Program; the National Science Foundation, Integrative Plant Biology; Florida Department of Environmental Protection, Center for Aquatic and Invasive Plants, University of Florida; Florida Institute of Oceanography; and NATO, Scientific and Environmental Affairs Division, Collaborative Research Grants.


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Authors and Affiliations

  1. 1.Department of BiologyUniversity of FloridaGainesvilleUSA

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