Carbon assimilation C4 pathway (Zea mays)

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5-7Active RubiscoBundle sheath cell cytoplasmInactive RubiscoLEGENDMETABOLITEENZYMEMesophyll cell cytoplasmTRANSPORT REACTION (INPUT/OUTPUT)PlasmodesmataMesophyll cell chloroplastENZYME REACTION (INPUT/OUTPUT)Calvin cycleBundle sheath cell chloroplastBundle sheath cell mitochondrionBundle sheath cellMesophyll cellTRANSPORTERPO4ATPdihydroxy-acetone-phosphatePO4NADPHD-xylulose-5-phosphate3-phosphoglycerateD-glyceraldehyde-3-phosphateADPADPD-ribulose-1,5-bisphosphateD-ribulose-1,5-bisphosphateNADPHHCO3Oxaloacetic acidADPfructose 1,6-bisphosphataseCO2Triosephosphate isomerasePO4MalatePO4PO4PO4PO4PO4PO4Triosephosphate isomeraseH+D-sedophetulose-1,7-bisphosphateNADP-malic enzymePO4PyruvateH2ONADPphosphoenolpyruvate carboxykinase (ATP)Aspartate aminotransferaseRibose-5-phosphate isomeraseAspartate aminotransferaseCO2CO2CO2CO2MalateNADPHdihydroxy-acetone-phosphatePO4Phosphoglycerate kinaseNADPTransketolaseATPATPATPCO2Active Pyruvate, phosphate dikinaseInactive Pyruvate, phosphate dikinasePPDK-regulatory protein3-phosphoglycerateCarbonic anhydrasePO4D-fructose-6-phosphateADPADPADP1,3-diphosphateglyceratePO4PO4Ribulose-bisphosphate carboxylaseRibulose-bisphosphate carboxylaseRubisco activaseRibulose-bisphosphate carboxylaseL-GlutamateL-AspartateL-AspartateL-AspartateOxaloacetic acid2-Ketoglutaric acidOxaloacetic acidOxaloacetic acidL-Glutamate2-Ketoglutaric acidH+Mg2+Mg2+Phosphoenol Pyruvate (PEP) CarboxylaseATPD-glyceraldehyde-3-phosphateH+Sedoheptulose 1,7-bisphosphatase AMPAMPAldolaseNAD(P)-dependent Glyceraldehyde-3-phosphate dehydrogenasedihydroxy-acetone-phosphatePhosphoenolpyruvateATPD-ribose-5-phosphatefructose-bisphosphate aldolaseMalateNADPPyruvateH2Odihydroxy-acetone-phosphateD-glyceraldehyde-3-phosphatePO4Malate dehydrogenase (NADP)Ribulose phosphate 3-epimerase 3-phosphoglyceratePyruvateH+D-sedoheptulose-7-phosphatePyruvatePhosphoglycerate kinasePhosphoenolpyruvatePhosphoenolpyruvateMalateD-erythrose-4-phosphatePPiPPiD-fructose-1,6-phosphateNAD(P)-dependent Glyceraldehyde-3-phosphate dehydrogenaseD-ribulose-5-phosphatePhosphoribulokinase 3-phosphoglycerate1,3-diphosphateglyceratePi-triose phosphate antiporter2-oxoglutarate/malate translocatorPi-triose phosphate antiporterMEP32-oxoglutarate/malate translocatorPi-triose phosphate antiporterPi-triose phosphate antiporterMEP3MEP1Phosphoenolpyruvate transporter


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  1. Majeran W, Zybailov B, Ytterberg AJ, Dunsmore J, Sun Q, van Wijk KJ; ''Consequences of C4 differentiation for chloroplast membrane proteomes in maize mesophyll and bundle sheath cells.''; Mol Cell Proteomics, 2008 PubMed Europe PMC Scholia
  2. Friso G, Majeran W, Huang M, Sun Q, van Wijk KJ; ''Reconstruction of metabolic pathways, protein expression, and homeostasis machineries across maize bundle sheath and mesophyll chloroplasts: large-scale quantitative proteomics using the first maize genome assembly.''; Plant Physiol, 2010 PubMed Europe PMC Scholia
  3. Chastain CJ, Botschner M, Harrington GE, Thompson BJ, Mills SE, Sarath G, Chollet R; ''Further analysis of maize C(4) pyruvate,orthophosphate dikinase phosphorylation by its bifunctional regulatory protein using selective substitutions of the regulatory Thr-456 and catalytic His-458 residues.''; Arch Biochem Biophys, 2000 PubMed Europe PMC Scholia
  4. Ayala-Ochoa A, Vargas-Suárez M, Loza-Tavera H, León P, Jiménez-García LF, Sánchez-de-Jiménez E; ''In maize, two distinct ribulose 1,5-bisphosphate carboxylase/oxygenase activase transcripts have different day/night patterns of expression.''; Biochimie, 2004 PubMed Europe PMC Scholia
  5. Tausta SL, Coyle HM, Rothermel B, Stiefel V, Nelson T; ''Maize C4 and non-C4 NADP-dependent malic enzymes are encoded by distinct genes derived from a plastid-localized ancestor.''; Plant Mol Biol, 2002 PubMed Europe PMC Scholia
  6. Sánchez de Jiménez E, Medrano L, Martínez-Barajas E; ''Rubisco activase, a possible new member of the molecular chaperone family.''; Biochemistry, 1995 PubMed Europe PMC Scholia
  7. ''''; , PubMed Europe PMC Scholia
  8. Vargas-Suárez M, Ayala-Ochoa A, Lozano-Franco J, García-Torres I, Díaz-Quiñonez A, Ortíz-Navarrete VF, Sánchez-de-Jiménez E; ''Rubisco activase chaperone activity is regulated by a post-translational mechanism in maize leaves.''; J Exp Bot, 2004 PubMed Europe PMC Scholia
  9. Li P, Ponnala L, Gandotra N, Wang L, Si Y, Tausta SL, Kebrom TH, Provart N, Patel R, Myers CR, Reidel EJ, Turgeon R, Liu P, Sun Q, Nelson T, Brutnell TP; ''The developmental dynamics of the maize leaf transcriptome.''; Nat Genet, 2010 PubMed Europe PMC Scholia
  10. Majeran W, van Wijk KJ; ''Cell-type-specific differentiation of chloroplasts in C4 plants.''; Trends Plant Sci, 2009 PubMed Europe PMC Scholia
  11. Majeran W, Cai Y, Sun Q, van Wijk KJ; ''Functional differentiation of bundle sheath and mesophyll maize chloroplasts determined by comparative proteomics.''; Plant Cell, 2005 PubMed Europe PMC Scholia
  12. Chris J. Chastain, Raymond Chollet; ''Regulation of pyruvate, orthophosphate dikinase by ADP-/Pi-dependent reversible phosphorylation in C3 and C4 plants''; Plant Physiology and Biochemistry; Volume 41, Issues 6-7, June 2003, Pages 523-532, 2003
  13. Majeran W, Friso G, Ponnala L, Connolly B, Huang M, Reidel E, Zhang C, Asakura Y, Bhuiyan NH, Sun Q, Turgeon R, van Wijk KJ; ''Structural and metabolic transitions of C4 leaf development and differentiation defined by microscopy and quantitative proteomics in maize.''; Plant Cell, 2010 PubMed Europe PMC Scholia
  14. Chastain CJ, Lee ME, Moorman MA, Shameekumar P, Chollet R; ''Site-directed mutagenesis of maize recombinant C4-pyruvate,orthophosphate dikinase at the phosphorylatable target threonine residue.''; FEBS Lett, 1997 PubMed Europe PMC Scholia
  15. Slack CR, Hatch MD, Goodchild DJ; ''Distribution of enzymes in mesophyll and parenchyma-sheath chloroplasts of maize leaves in relation to the C4-dicarboxylic acid pathway of photosynthesis.''; Biochem J, 1969 PubMed Europe PMC Scholia
  16. Slack CR, Hatch MD; ''Comparative studies on the activity of carboxylases and other enzymes in relation to the new pathway of photosynthetic carbon dioxide fixation in tropical grasses.''; Biochem J, 1967 PubMed Europe PMC Scholia


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107275view14:36, 17 September 2019MaintBotChEBI identifier normalization
79802view16:23, 16 April 2015Mkutmonchanged interactions in legend to graphical lines
78447view10:27, 7 January 2015MaintBotadded missing graphIds
71957view23:26, 23 October 2013MaintBotremoved data source from nodes without identifier
70143view21:21, 12 July 2013MaintBotupdated to 2013 schema
57566view19:56, 29 January 2013Khanspersremoved transcript suffix from gene ids
57565view19:50, 29 January 2013Khanspersremoved transcript suffic from gene ids
48579view22:15, 5 June 2012Pjaiswaladded literature
48578view22:06, 5 June 2012PjaiswalModified description
48577view22:05, 5 June 2012PjaiswalModified description
48576view21:52, 5 June 2012PjaiswalOntology Term : 'carbon fixation pathway' added !
48575view21:51, 5 June 2012PjaiswalModified categories
48574view21:49, 5 June 2012Pjaiswaladded a lot of min catalyst line type. added geneproduct/gene IDs
48573view21:47, 5 June 2012PjaiswalPeriodical save, work in progress
48572view21:37, 5 June 2012PjaiswalPeriodical save, work in progress
48571view21:16, 5 June 2012PjaiswalPeriodical save, work in progress
40457view18:46, 1 March 2011KhanspersReverted to version '18:32, 1 March 2011' by Khanspers
40456view18:44, 1 March 2011Khansperstesting
40455view18:32, 1 March 2011MaintBotRemoved redundant pathway information and comments
38321view23:34, 31 August 2010KhanspersChanged cellulr shapes
38315view22:56, 31 August 2010KhanspersChanged OrganC to Ovals
36274view23:30, 16 March 2010AlexanderPicoModified categories
36270view17:55, 16 March 2010PjaiswalNew pathway

External references


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NameTypeDatabase referenceComment
1,3-diphosphateglycerateMetabolite1981-49-3 (CAS)
2-Ketoglutaric acidMetabolite30915 (ChEBI) 2-oxoglutaric acid
2-oxoglutarate/malate translocatorGeneProduct2-Oxoglutarate/malate translocator; DIT1
3-phosphoglycerateMetabolite17050 (ChEBI)
ADPMetabolite16761 (ChEBI)
AMPMetabolite16027 (ChEBI)
ATPMetabolite15422 (ChEBI)
Active Pyruvate, phosphate dikinaseGeneProduct2.7.9.1 (EC Number) Active PPDK found in light conditions has a phosphorylated Histitine and a hydroxy Threonine residue.
AldolaseGeneProduct4.1.2. (EC Number)
Aspartate aminotransferaseGeneProduct2.6.1.1 (EC Number) ASPTS
CO2Metabolite16526 (ChEBI)
Carbonic anhydraseGeneProduct4.2.1.1 (EC Number) CA
D-erythrose-4-phosphateMetabolite16897 (ChEBI)
D-fructose-1,6-phosphateMetabolite16905 (ChEBI)
D-fructose-6-phosphateMetabolite15946 (ChEBI)
D-glyceraldehyde-3-phosphateMetabolite29052 (ChEBI)
D-ribose-5-phosphateMetabolite16988 (ChEBI)
D-ribulose-1,5-bisphosphateMetabolite16710 (ChEBI)
D-ribulose-5-phosphateMetabolite17363 (ChEBI)
D-sedoheptulose-7-phosphateMetabolite15721 (ChEBI)
D-sedophetulose-1,7-bisphosphateMetabolite17969 (ChEBI)
D-xylulose-5-phosphateMetabolite16332 (ChEBI)
H+Metabolite15378 (ChEBI)
H2OMetabolite15377 (ChEBI)
HCO3Metabolite17544 (ChEBI)
Inactive Pyruvate, phosphate dikinaseGeneProduct2.7.9.1 (EC Number) Inactive PPDK found in dark conditions has a phosphorylated Histitine and a phosphorylated threonine residue.
L-AspartateMetabolite29993 (ChEBI) OAA
L-GlutamateMetabolite29988 (ChEBI)
Malate dehydrogenase (NADP)GeneProduct1.1.1.82 (EC Number)
MalateMetabolite15595 (ChEBI)
Mg2+Metabolite18420 (ChEBI)
NAD(P)-dependent Glyceraldehyde-3-phosphate dehydrogenaseGeneProduct1.2.1.59 (EC Number)
NADP-malic enzymeGeneProduct1.1.1.40 (EC Number)
NADPMetabolite18009 (ChEBI)
NADPHMetabolite16474 (ChEBI)
Oxaloacetic acidMetabolite30744 (ChEBI) OAA
PO4Metabolite18367 (ChEBI)
PPDK-regulatory proteinGeneProductPPDK
PPiMetabolite29888 (ChEBI)
Phosphoenol Pyruvate (PEP) CarboxylaseGeneProduct4.1.1.31 (EC Number) PEPCase
Phosphoenolpyruvate transporterGeneProductPEPT
PhosphoenolpyruvateMetabolite18021 (ChEBI) PEP
Phosphoglycerate kinaseGeneProduct2.7.2.3 (EC Number)
Phosphoribulokinase GeneProduct2.7.1.19 (EC Number) Also called as ribulose 5-phosphate kinase
Pi-triose phosphate antiporterGeneProductphosphate/triose-phosphate translocator
PyruvateMetabolite15361 (ChEBI)
Ribose-5-phosphate isomeraseGeneProduct5.3.1.6 (EC Number)
Ribulose phosphate 3-epimerase GeneProduct5.1.3.1 (EC Number)
Ribulose-bisphosphate carboxylaseComplex4.1.1.39 (EC Number)
Rubisco activaseComplex
Sedoheptulose 1,7-bisphosphatase GeneProduct3.1.3.37 (EC Number)
TransketolaseGeneProduct2.2.1.1 (EC Number)
Triosephosphate isomeraseGeneProduct5.3.1.1 (EC Number)
dihydroxy-acetone-phosphateMetabolite16108 (ChEBI)
fructose 1,6-bisphosphataseGeneProduct3.1.3.11 (EC Number)
fructose-bisphosphate aldolaseGeneProduct4.1.2.13 (EC Number)
phosphoenolpyruvate carboxykinase (ATP)GeneProduct4.1.1.49 (EC Number) PEPCK

Annotated Interactions

No annotated interactions

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