KEGG   PATHWAY: ko00720
Entry
ko00720                     Pathway                                
Name
Other carbon fixation pathways
Description
Carbon fixation is an important pathway for autotrophs living in various environments. Plants and cyanobacteria fix CO2 as organic compounds using solar energy mainly by the reductive pentose phosphate cycle (also called Calvin cycle, Calvin-Benson cycle, or Calvin-Benson-Bassham cycle) [MD:M00165]. There are, at least, five additional carbon fixation pathways known to exist in autotrophic bacteria and archaea, which differ in reducing compounds, energy source, and oxygen sensitivity of enzymes. (i) The reductive citric acid cycle (Arnon-Buchanan cycle) [MD:M00173] is found in microaerophiles and anaerobes, such as green sulfur bacteria. In one complete turn of this cycle, four molecules of CO2 are fixed by the enzymes that are sensitive to oxygen, resulting in the production of one molecule of oxaloacetate, which is itself an intermediate of the cycle. (ii) The reductive acetyl-CoA pathway (Wood-Ljungdahl pathway) [MD:M00377] is found in strictly anaerobic bacteria and archaea (Proteobacteria, Planctomycetes, Spirochaetes, and Euryarchaeota), some of which are methane-forming. A bifunctional enzyme, carbon monoxide dehydrogenase/acetyl-CoA synthase, catalyzes the reactions from CO2 to CO and from CO2 to a methyl group, and then to generate acetyl-CoA. (iii) The 3-hydroxypropionate bicycle [MD:M00376] is found in some green non-sulphur bacteria of the family Chloroflexaceae. In one complete turn of this bicycle, three molecules of bicarbonate are converted into one molecule of pyruvate. In addition, this bicycle provides the secondary benefit of useful intermediates for biosynthesis: acetyl-CoA, glyoxylate, and succinyl-CoA. (iv) The hydroxypropionate-hydroxybutyrate cycle [MD:M00375] is found in aerobic Crenarchaeota, Acidianus, Metallosphaera, and Sulfolobales. Some of the intermediates and the carboxylation reactions are the same as in the 3-hydroxypropionate bicycle. One complete turn of this cycle generates two molecules of acetyl-CoA, one of which is reutilized in the the cycle and the other is removed for cell material biosynthesis. (v) The dicarboxylate-hydroxybutyrate cycle [MD:M00374] was named after its intermediates: succinate (a kind of dicarboxylate) and hydroxybutyrate. This cycle has been found only in Ignicoccus hospitals, a strictly anaerobic hyperthermophilic archaea. Recent genome study suggests that this cycle may exist in Desulfurococcales (to which Ignicoccus belongs) and Thermoproteales (a taxon close to the origin of archaea). The first half of the cycle, from acetyl-CoA to succinate-CoA, corresponds to the reductive citric acid cycle and the latter half of the cycle, from succinate-CoA to two molecules of acetyl-CoA, corresponds to the hydroxypropionate-hydroxybutyrate cycle.
Class
Metabolism; Energy metabolism
Pathway map
ko00720  Other carbon fixation pathways
ko00720

Module
M00173  Reductive citrate cycle (Arnon-Buchanan cycle) [PATH:ko00720]
M00374  Dicarboxylate-hydroxybutyrate cycle [PATH:ko00720]
M00375  Hydroxypropionate-hydroxybutylate cycle [PATH:ko00720]
M00376  3-Hydroxypropionate bi-cycle [PATH:ko00720]
M00377  Reductive acetyl-CoA pathway (Wood-Ljungdahl pathway) [PATH:ko00720]
M00579  Phosphate acetyltransferase-acetate kinase pathway, acetyl-CoA => acetate [PATH:ko00720]
M00620  Incomplete reductive citrate cycle, acetyl-CoA => oxoglutarate [PATH:ko00720]
Other DBs
GO: 0015977
Orthology
K00024  mdh; malate dehydrogenase [EC:1.1.1.37]
K00031  IDH1, IDH2, icd; isocitrate dehydrogenase [EC:1.1.1.42]
K00169  porA; pyruvate ferredoxin oxidoreductase alpha subunit [EC:1.2.7.1]
K00170  porB; pyruvate ferredoxin oxidoreductase beta subunit [EC:1.2.7.1]
K00171  porD; pyruvate ferredoxin oxidoreductase delta subunit [EC:1.2.7.1]
K00172  porC, porG; pyruvate ferredoxin oxidoreductase gamma subunit [EC:1.2.7.1]
K00174  korA, oorA, oforA; 2-oxoglutarate/2-oxoacid ferredoxin oxidoreductase subunit alpha [EC:1.2.7.3 1.2.7.11]
K00175  korB, oorB, oforB; 2-oxoglutarate/2-oxoacid ferredoxin oxidoreductase subunit beta [EC:1.2.7.3 1.2.7.11]
K00176  korD, oorD; 2-oxoglutarate ferredoxin oxidoreductase subunit delta [EC:1.2.7.3]
K00177  korC, oorC; 2-oxoglutarate ferredoxin oxidoreductase subunit gamma [EC:1.2.7.3]
K00189  vorG, porG; 2-oxoisovalerate/pyruvate ferredoxin oxidoreductase gamma subunit [EC:1.2.7.7 1.2.7.1]
K00194  cdhD, acsD; acetyl-CoA decarbonylase/synthase, CODH/ACS complex subunit delta [EC:2.1.1.245]
K00196  cooF; anaerobic carbon-monoxide dehydrogenase iron sulfur subunit
K00197  cdhE, acsC; acetyl-CoA decarbonylase/synthase, CODH/ACS complex subunit gamma [EC:2.1.1.245]
K00198  cooS, acsA; anaerobic carbon-monoxide dehydrogenase catalytic subunit [EC:1.2.7.4]
K00233  frdA, fdrA; succinate dehydrogenase flavoprotein subunit
K00239  sdhA, frdA; succinate dehydrogenase flavoprotein subunit [EC:1.3.5.1]
K00240  sdhB, frdB; succinate dehydrogenase iron-sulfur subunit [EC:1.3.5.1]
K00241  sdhC, frdC; succinate dehydrogenase cytochrome b subunit
K00242  sdhD, frdD; succinate dehydrogenase membrane anchor subunit
K00244  frdA; succinate dehydrogenase flavoprotein subunit [EC:1.3.5.1]
K00245  frdB; succinate dehydrogenase iron-sulfur subunit [EC:1.3.5.1]
K00246  frdC; succinate dehydrogenase subunit C
K00247  frdD; succinate dehydrogenase subunit D
K00297  metF, MTHFR; methylenetetrahydrofolate reductase (NADH) [EC:1.5.1.54]
K00625  pta; phosphate acetyltransferase [EC:2.3.1.8]
K00626  ACAT, atoB; acetyl-CoA C-acetyltransferase [EC:2.3.1.9]
K00925  ackA; acetate kinase [EC:2.7.2.1]
K01006  ppdK; pyruvate, orthophosphate dikinase [EC:2.7.9.1]
K01007  pps, ppsA; pyruvate, water dikinase [EC:2.7.9.2]
K01491  folD; methylenetetrahydrofolate dehydrogenase (NADP+) / methenyltetrahydrofolate cyclohydrolase [EC:1.5.1.5 3.5.4.9]
K01500  fchA; methenyltetrahydrofolate cyclohydrolase
K01595  ppc; phosphoenolpyruvate carboxylase [EC:4.1.1.31]
K01648  ACLY; ATP citrate (pro-S)-lyase [EC:2.3.3.8]
K01675  fumD; fumarate hydratase D
K01676  E4.2.1.2A, fumA, fumB; fumarate hydratase, class I [EC:4.2.1.2]
K01677  E4.2.1.2AA, fumA; fumarate hydratase subunit alpha [EC:4.2.1.2]
K01678  E4.2.1.2AB, fumB; fumarate hydratase subunit beta [EC:4.2.1.2]
K01679  E4.2.1.2B, fumC, FH; fumarate hydratase, class II [EC:4.2.1.2]
K01681  ACO; aconitate hydratase [EC:4.2.1.3]
K01682  acnB; aconitate hydratase 2 / 2-methylisocitrate dehydratase [EC:4.2.1.3 4.2.1.99]
K01774  fumE; fumarate hydratase E
K01847  MUT; methylmalonyl-CoA mutase [EC:5.4.99.2]
K01848  E5.4.99.2A, mcmA1; methylmalonyl-CoA mutase, N-terminal domain [EC:5.4.99.2]
K01849  E5.4.99.2B, mcmA2; methylmalonyl-CoA mutase, C-terminal domain [EC:5.4.99.2]
K01895  ACSS1_2, acs; acetyl-CoA synthetase [EC:6.2.1.1]
K01902  sucD; succinyl-CoA synthetase alpha subunit [EC:6.2.1.5]
K01903  sucC; succinyl-CoA synthetase beta subunit [EC:6.2.1.5]
K01938  fhs; formate--tetrahydrofolate ligase [EC:6.3.4.3]
K01958  PC, pyc; pyruvate carboxylase [EC:6.4.1.1]
K01959  pycA; pyruvate carboxylase subunit A [EC:6.4.1.1]
K01960  pycB; pyruvate carboxylase subunit B [EC:6.4.1.1]
K01961  accC; acetyl-CoA carboxylase, biotin carboxylase subunit [EC:6.4.1.2 6.3.4.14]
K01962  accA; acetyl-CoA carboxylase carboxyl transferase subunit alpha [EC:6.4.1.2 2.1.3.15]
K01963  accD; acetyl-CoA carboxylase carboxyl transferase subunit beta [EC:6.4.1.2 2.1.3.15]
K01964  K01964; acetyl-CoA/propionyl-CoA carboxylase [EC:6.4.1.2 6.4.1.3]
K02160  accB, bccP; acetyl-CoA carboxylase biotin carboxyl carrier protein
K03737  por, nifJ; pyruvate-ferredoxin/flavodoxin oxidoreductase [EC:1.2.7.1 1.2.7.-]
K05299  fdhA; formate dehydrogenase (NADP+) alpha subunit [EC:1.17.1.10]
K05606  MCEE, epi; methylmalonyl-CoA/ethylmalonyl-CoA epimerase [EC:5.1.99.1]
K08691  mcl; malyl-CoA/(S)-citramalyl-CoA lyase [EC:4.1.3.24 4.1.3.25]
K09709  meh; 3-methylfumaryl-CoA hydratase [EC:4.2.1.153]
K13788  pta; phosphate acetyltransferase [EC:2.3.1.8]
K14138  acsB; acetyl-CoA synthase [EC:2.3.1.169]
K14449  mch, mcd; 2-methylfumaryl-CoA hydratase [EC:4.2.1.148]
K14465  K14465; succinate semialdehyde reductase (NADPH) [EC:1.1.1.-]
K14466  K14466; 4-hydroxybutyrate---CoA ligase (AMP-forming) [EC:6.2.1.40]
K14467  4hbl; 4-hydroxybutyrate---CoA ligase (AMP-forming) [EC:6.2.1.40]
K14468  mcr; malonyl-CoA reductase / 3-hydroxypropionate dehydrogenase (NADP+) [EC:1.2.1.75 1.1.1.298]
K14469  K14469; acrylyl-CoA reductase (NADPH) / 3-hydroxypropionyl-CoA dehydratase / 3-hydroxypropionyl-CoA synthetase [EC:1.3.1.84 4.2.1.116 6.2.1.36]
K14470  mct; 2-methylfumaryl-CoA isomerase [EC:5.4.1.3]
K14471  smtA1; succinyl-CoA:(S)-malate CoA-transferase subunit A [EC:2.8.3.22]
K14472  smtB; succinyl-CoA:(S)-malate CoA-transferase subunit B [EC:2.8.3.22]
K14534  abfD; 4-hydroxybutyryl-CoA dehydratase / vinylacetyl-CoA-Delta-isomerase [EC:4.2.1.120 5.3.3.3]
K15016  K15016; enoyl-CoA hydratase / 3-hydroxyacyl-CoA dehydrogenase [EC:4.2.1.17 1.1.1.35]
K15017  K15017; malonyl-CoA/succinyl-CoA reductase (NADPH) [EC:1.2.1.75 1.2.1.76]
K15018  K15018; 3-hydroxypropionyl-coenzyme A synthetase [EC:6.2.1.36]
K15019  K15019; 3-hydroxypropionyl-coenzyme A dehydratase [EC:4.2.1.116]
K15020  K15020; acryloyl-coenzyme A reductase [EC:1.3.1.84]
K15022  fdhB; formate dehydrogenase (NADP+) beta subunit [EC:1.17.1.10]
K15023  acsE; 5-methyltetrahydrofolate corrinoid/iron sulfur protein methyltransferase [EC:2.1.1.258]
K15024  K15024; putative phosphotransacetylase [EC:2.3.1.8]
K15036  K15036; acetyl-CoA/propionyl-CoA carboxylase [EC:6.4.1.2 6.4.1.3 2.1.3.15]
K15037  K15037; biotin carboxyl carrier protein
K15038  K15038; succinyl-CoA reductase [EC:1.2.1.76]
K15039  K15039; 3-hydroxypropionate dehydrogenase (NADP+) [EC:1.1.1.298]
K15052  K15052; propionyl-CoA carboxylase [EC:6.4.1.3 2.1.3.15]
K15230  aclA; ATP-citrate lyase alpha-subunit [EC:2.3.3.8]
K15231  aclB; ATP-citrate lyase beta-subunit [EC:2.3.3.8]
K15232  ccsA; citryl-CoA synthetase large subunit [EC:6.2.1.18]
K15233  ccsB; citryl-CoA synthetase small subunit
K15234  ccl; citryl-CoA lyase [EC:4.1.3.34]
K18209  tfrA; fumarate reductase (CoM/CoB) subunit A [EC:1.3.4.1]
K18210  tfrB; fumarate reductase (CoM/CoB) subunit B [EC:1.3.4.1]
K18556  frdA; NADH-dependent fumarate reductase subunit A [EC:1.3.1.6]
K18557  frdB; NADH-dependent fumarate reductase subunit B [EC:1.3.1.6]
K18558  frdC; NADH-dependent fumarate reductase subunit C
K18559  frdD; NADH-dependent fumarate reductase subunit D
K18560  frdE; NADH-dependent fumarate reductase subunit E
K18593  E6.2.1.56; 4-hydroxybutyrate---CoA ligase (ADP-forming) [EC:6.2.1.56]
K18594  K18594; 3-hydroxypropionyl-CoA synthetase (ADP-forming) [EC:6.2.1.-]
K18602  K18602; malonic semialdehyde reductase [EC:1.1.1.-]
K18603  K18603; acetyl-CoA/propionyl-CoA carboxylase [EC:6.4.1.2 6.4.1.3]
K18604  K18604; acetyl-CoA/propionyl-CoA carboxylase [EC:6.4.1.2 6.4.1.3 2.1.3.15]
K18605  K18605; biotin carboxyl carrier protein
K18859  sdhD, frdD; succinate dehydrogenase subunit D
K18860  sdhD, frdD; succinate dehydrogenase subunit D
K18861  K18861; 4-hydroxybutyrate---CoA ligase (AMP-forming) [EC:6.2.1.40]
K22015  fdhF; formate dehydrogenase (hydrogenase) [EC:1.17.98.4 1.17.98.-]
K25007  metV; methylenetetrahydrofolate reductase (NADH) small subunit
K25008  rnfC2; ion-translocating oxidoreductase complex subunit C
K25123  hydA2; iron hydrogenase HydA2
K25124  hycB; FeS-containing electron transfer protein
K25221  mtgA; [methyl-Co(III) glycine betaine-specific corrinoid protein]---tetrahydrofolate methyltransferase
K25774  K25774; 4-hydroxybutyrate---CoA ligase (AMP-forming)
K25915  mtvA; [methyl-Co(III) methoxylated-aromatic-compound-specific corrinoid protein]---tetrahydrofolate methyltransferase
K25995  frdB, fdrB; succinate dehydrogenase iron-sulfur subunit
K25996  frdC, fdrC; succinate dehydrogenase subunit C
K26318  fccA, fcc3; fumarate reductase (cytochrome)
K26930  mtqA; [methyl-Co(III) quaternary-amine-specific corrinoid protein]---tetrahydrofolate methyltransferase
K27802  acnA; aconitate hydratase A / 2-methylisocitrate dehydratase
Compound
C00011  CO2
C00022  Pyruvate
C00024  Acetyl-CoA
C00026  2-Oxoglutarate
C00033  Acetate
C00036  Oxaloacetate
C00042  Succinate
C00048  Glyoxylate
C00058  Formate
C00074  Phosphoenolpyruvate
C00083  Malonyl-CoA
C00091  Succinyl-CoA
C00100  Propanoyl-CoA
C00101  Tetrahydrofolate
C00122  Fumarate
C00143  5,10-Methylenetetrahydrofolate
C00149  (S)-Malate
C00158  Citrate
C00222  3-Oxopropanoate
C00227  Acetyl phosphate
C00232  Succinate semialdehyde
C00234  10-Formyltetrahydrofolate
C00237  CO
C00288  HCO3-
C00311  Isocitrate
C00332  Acetoacetyl-CoA
C00417  cis-Aconitate
C00440  5-Methyltetrahydrofolate
C00445  5,10-Methenyltetrahydrofolate
C00566  (3S)-Citryl-CoA
C00683  (S)-Methylmalonyl-CoA
C00877  Crotonoyl-CoA
C00894  Propenoyl-CoA
C00989  4-Hydroxybutanoic acid
C01011  (3S)-Citramalyl-CoA
C01013  3-Hydroxypropanoate
C01144  (S)-3-Hydroxybutanoyl-CoA
C01213  (R)-Methylmalonyl-CoA
C04348  L-Malyl-CoA
C05668  3-Hydroxypropionyl-CoA
C06027  L-erythro-3-Methylmalyl-CoA
C06028  2-Methylfumaryl-CoA
C11062  4-Hydroxybutyryl-CoA
C18323  3-Methylfumaryl-CoA
Reference
  Authors
Zarzycki J, Brecht V, Muller M, Fuchs G
  Title
Identifying the missing steps of the autotrophic 3-hydroxypropionate CO2 fixation cycle in Chloroflexus aurantiacus.
  Journal
Proc Natl Acad Sci U S A 106:21317-22 (2009)
DOI:10.1073/pnas.0908356106
Reference
  Authors
Teufel R, Kung JW, Kockelkorn D, Alber BE, Fuchs G
  Title
3-hydroxypropionyl-coenzyme A dehydratase and acryloyl-coenzyme A reductase, enzymes of the autotrophic 3-hydroxypropionate/4-hydroxybutyrate cycle in the Sulfolobales.
  Journal
J Bacteriol 191:4572-81 (2009)
DOI:10.1128/JB.00068-09
Reference
  Authors
Huber H, Gallenberger M, Jahn U, Eylert E, Berg IA, Kockelkorn D, Eisenreich W, Fuchs G
  Title
A dicarboxylate/4-hydroxybutyrate autotrophic carbon assimilation cycle in the hyperthermophilic Archaeum Ignicoccus hospitalis.
  Journal
Proc Natl Acad Sci U S A 105:7851-6 (2008)
DOI:10.1073/pnas.0801043105
Reference
  Authors
Hugler M, Wirsen CO, Fuchs G, Taylor CD, Sievert SM
  Title
Evidence for autotrophic CO2 fixation via the reductive tricarboxylic acid cycle by members of the epsilon subdivision of proteobacteria.
  Journal
J Bacteriol 187:3020-7 (2005)
DOI:10.1128/JB.187.9.3020-3027.2005
Reference
  Authors
Seravalli J, Kumar M, Ragsdale SW
  Title
Rapid kinetic studies of acetyl-CoA synthesis: evidence supporting the catalytic intermediacy of a paramagnetic NiFeC species in the autotrophic Wood-Ljungdahl pathway.
  Journal
Biochemistry 41:1807-19 (2002)
DOI:10.1021/bi011687i
Reference
PMID:3096193
  Authors
Ljungdahl LG
  Title
The autotrophic pathway of acetate synthesis in acetogenic bacteria.
  Journal
Annu Rev Microbiol 40:415-50 (1986)
DOI:10.1146/annurev.mi.40.100186.002215
Related
pathway
ko00710  Carbon fixation by Calvin cycle

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