Essential Information & explanations, latest texts & monographs on C4_photosynthesis.


C4 photosynthesis

C4 photosynthesis is a common metabolic pathway found in land plants living under arid conditions. CAM plants utilize a similar process. It is called "C4" because the product, oxaloacetate, contains four carbon atoms. It occurs in the mesophyll of the leaf. The chemical equation is: PEP carboxylase + PEP + CO2 → oxaloacetate The product is usually converted to malate, a simple organic compound that gives up its CO2 to the Calvin cycle after being shipped off to bundle sheath cells surronding a nearby vein. After losing the CO2, it becomes pyruvate, and can be phophorylated into PEP at the cost of a phosphorus group and one ATP. It can then be reused in the above equation.

The above article is adapted from from Wikipedia All Wikipedia article text is available under the terms of the GNU Free Documentation License



C3, C4: Mechanisms, Cellular and Environmental Regulation of Photosynthesis by Gerald Edwards

Energy, Plants and Man: An Introduction to Photosynthesis in C3, C4 and Cam Plants by David Walker

C3, C4: Mechanisms, and Cellular and Environmental Regulation of Photosynthesis by G. Edwards





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Recent C4_photosynthesis related patents

From USPTO:




from PUBMED
1:  Kubien DS, von Caemmerer S, Furbank RT, Sage RF. 
 C4 photosynthesis at low temperature. A study using transgenic plants with
reduced amounts of Rubisco.
Plant Physiol. 2003 Jul;132(3):1577-85. 
PMID: 12857837 

2:  Cousins AB, Adam NR, Wall GW, Kimball BA, Pinter PJ Jr, Ottman MJ, Leavitt
SW, Webber AN. 
 Development of C4 photosynthesis in sorghum leaves grown under free-air CO2
enrichment (FACE).
J Exp Bot. 2003 Aug;54(389):1969-75. Epub 2003 Jul 01. 
PMID: 12837815 

3:  Tamada Y, Imanari E, Kurotani K, Nakai M, Andreo CS, Izui K. 
 Effect of photooxidative destruction of chloroplasts on the expression of
nuclear genes for C4 photosynthesis and for chloroplast biogenesis in maize.
J Plant Physiol. 2003 Jan;160(1):3-8. 
PMID: 12685039 

4:  Gutierrez MV. 
 The scientific development of the physiology of plants in the American tropics.
Rev Biol Trop. 2002 Jun;50(2):429-38. Review. 
PMID: 12298276 

5:  Marshall JS, Stubbs JD, Chitty JA, Surin B, Taylor WC. 
 Expression of the C4 Me1 Gene from Flaveria bidentis Requires an Interaction
between 5[prime] and 3[prime] Sequences.
Plant Cell. 1997 Sep;9(9):1515-1525. 
PMID: 12237394 

6:  Brown RH, Byrd GT. 
 Estimation of Bundle Sheath Cell Conductance in C4 Species and O2 Insensitivity
of Photosynthesis.
Plant Physiol. 1993 Dec;103(4):1183-1188. 
PMID: 12232010 

7:  Dai Z, Ku M, Edwards GE. 
 C4 Photosynthesis (The CO2-Concentrating Mechanism and Photorespiration).
Plant Physiol. 1993 Sep;103(1):83-90. 
PMID: 12231916 

8:  Bakrim N, Prioul JL, Deleens E, Rocher JP, Arrio-Dupont M, Vidal J, Gadal P,
Chollet R. 
 Regulatory Phosphorylation of C4 Phosphoenolpyruvate Carboxylase (A Cardinal
Event Influencing the Photosynthesis Rate in Sorghum and Maize).
Plant Physiol. 1993 Mar;101(3):891-897. 
PMID: 12231740 

9:  Brown RH, Byrd GT, Bouton JH, Bassett CL. 
 Photosynthetic Characteristics of Segregates from Hybrids between Flaveria
brownii (C4 Like) and Flaveria linearis (C3-C4).
Plant Physiol. 1993 Mar;101(3):825-831. 
PMID: 12231732 

10:  Hatch MD, Agostino A, Jenkins C. 
 Measurement of the Leakage of CO2 from Bundle-Sheath Cells of Leaves during C4
Photosynthesis.
Plant Physiol. 1995 May;108(1):173-181. 
PMID: 12228461 

11:  Dai Z, Ku M, Edwards GE. 
 C4 Photosynthesis (The Effects of Leaf Development on the CO2-Concentrating
Mechanism and Photorespiration in Maize).
Plant Physiol. 1995 Mar;107(3):815-825. 
PMID: 12228406 

12:  Pfundel E, Nagel E, Meister A. 
 Analyzing the Light Energy Distribution in the Photosynthetic Apparatus of C4
Plants Using Highly Purified Mesophyll and Bundle-Sheath Thylakoids.
Plant Physiol. 1996 Nov;112(3):1055-1070. 
PMID: 12226432 

13:  Magnin NC, Cooley BA, Reiskind JB, Bowes G. 
 Regulation and Localization of Key Enzymes during the Induction of Kranz-Less,
C4-Type Photosynthesis in Hydrilla verticillata.
Plant Physiol. 1997 Dec;115(4):1681-1689. 
PMID: 12223888 

14:  Taniguchi M, Sugiyama T. 
 The Expression of 2-Oxoglutarate/Malate Translocator in the Bundle-Sheath
Mitochondria of Panicum miliaceum, a NAD-Malic Enzyme-Type C4 Plant, Is
Regulated by Light and Development.
Plant Physiol. 1997 May;114(1):285-293. 
PMID: 12223705 

15:  Pfundel E, Pfeffer M. 
 Modification of Photosystem I Light Harvesting of Bundle-Sheath Chloroplasts
Occurred during the Evolution of NADP-Malic Enzyme C4 Photosynthesis.
Plant Physiol. 1997 May;114(1):145-152. 
PMID: 12223694 

16:  Trevanion SJ, Furbank RT, Ashton AR. 
 NADP-Malate Dehydrogenase in the C4 Plant Flaveria bidentis (Cosense
Suppression of Activity in Mesophyll and Bundle-Sheath Cells and Consequences
for Photosynthesis).
Plant Physiol. 1997 Apr;113(4):1153-1165. 
PMID: 12223666 

17:  Cushman JC, Bohnert HJ. 
 Molecular Genetics of Crassulacean Acid Metabolism.
Plant Physiol. 1997 Mar;113(3):667-676. 
PMID: 12223634 

18:  Voznesenskaya EV, Franceschi VR, Kiirats O, Artyusheva EG, Freitag H,
Edwards GE. 
 Proof of C4 photosynthesis without Kranz anatomy in Bienertia cycloptera
(Chenopodiaceae).
Plant J. 2002 Sep;31(5):649-62. 
PMID: 12207654 

19:  Jeanneau M, Vidal J, Gousset-Dupont A, Lebouteiller B, Hodges M, Gerentes
D, Perez P. 
 Manipulating PEPC levels in plants.
J Exp Bot. 2002 Sep;53(376):1837-45. Review. 
PMID: 12177121 

20:  Blasing OE, Ernst K, Streubel M, Westhoff P, Svensson P. 
 The non-photosynthetic phosphoenolpyruvate carboxylases of the C4 dicot
Flaveria trinervia -- implications for the evolution of C4 photosynthesis.
Planta. 2002 Jul;215(3):448-56. Epub 2002 Apr 12. 
PMID: 12111227 

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Note again ... some material here is adapted from from Wikipedia All Wikipedia article text is available under the terms of the GNU Free Documentation License

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