Review Article

More Than 200 Genes Required for Methane Formation from H2 and CO2 and Energy Conservation Are Present in Methanothermobacter marburgensis and Methanothermobacter thermautotrophicus

Table 2

The ca. 200 identified genes required for methane formation and energy conservation in Methanothermobacter marburgensis and Methanothermobacter thermautotrophicus. The number of CDS that remain to be identified are given in parentheses. For functions, see Figure 2. H4MPT, tetrahydromethanopterin; MFR, methanofuran.

ProteinsGenes in M. marburgensis Genes in M. thermautotrophicus
(MTBMA_c)Number of CDS(MTH)

H2 activation

MvhADG: F420-non reducing [NiFe]-hydrogenase associated with15190–15160 (mvhDGAB) (MvhB is a polyferredoxin)41136–1133
HdrABC: heterodisulfide reductase17680, 04500, 0449031381, 1879, 1878
FrhABG: F420-reducing [NiFe]-hydrogenase; synthesis regulated by nickel16860–16830 (frhADGB) (FrhD is an endopeptidase)51300–1297
Hmd: [Fe]-hydrogenase (H2-forming methylene-H4MPT dehydrogenase); synthesis regulated by nickel1526011142
EhaA-T: Energy-converting [NiFe]-hydrogenase; membrane-associated; Na+-dependent; mainly anabolic function07840–0803020384–404
(EhaP twice)
EhbA-Q: Energy-converting [NiFe]-hydrogenase; membrane-associated; Na+-dependent; mainly anabolic function16230–16390171251–1235
Nickel-responsive transcriptional regulator NikR07330, 09830, 113403603, 739

CO2 reduction to methane

FwdA-DFGH: Tungsten-dependent formyl-MFR dehydrogenase; formed constitutively01390–01450 (fwdHFGDACB)71553–1559
FwdA/FmdBCE: Molybdenum-dependent formyl-MFR dehydrogenase; formed only in the presence of molybdate, involving DNA-binding protein Tfx13050–13070 (fmdECB)3917–919
130401916
Ftr: Formylmethanofuran: H4MPT formyltransferase164601403
Mch: Methenyl-H4MPT cyclohydrolase116901773
Mtd: F420-dependent methylene-H4MPT dehydrogenase; synthesis regulated by nickel0050011464
Mer: F420-dependent methylene-H4MPT reductase0327011752
MtrA-H: Methyl-H4MPT:coenzyme M methyltransferase; membrane-associated; Na+-dependent15400–15470 (mtrHGFABCDE)71156–1163
McrABG: Methyl-coenzyme M reductase isoenzyme I; contains 5 posttranslational modifications15480–15520 (mcrAGCDB)5 1164–1168
(McrCD of unknown function)(5)
MrtABG: Methyl-coenzyme M reductase isoenzyme II; contains 5 posttranslational modifications15120–15150 (mrtAGDB)41129–1132
(MrtD of unknown function)
AtwA: Mcr/Mrt-activating enzyme A213970, 060102151, 454, 1015
Putative transcriptional regulator of the mcr operon057601126

Electron transport

6[4Fe4S] polyferredoxin (EhaP); See above See above
8[4Fe4S] polyferredoxin (FwdF);
10[4Fe4S] polyferredoxin (EhaQ);
12[4Fe4S] polyferredoxin (MvhB);
14[4Fe4S] polyferredoxin (EhbK)
2[4Fe4S] ferredoxins00530, 03900, 07270, 1489041468, 1819, 278, 1106
6[4Fe4S] polyferredoxin080401405
8[4Fe4S] polyferredoxin1736011345

ADP phosphorylation via the Na+-motive force (generated in the MtrA-H-catalyzed reaction; see above)

AhaA-IK: A1A0 ATP synthase complex; membrane-associated; Na+-dependent?13380–13470 (ahaGDBAFCEKIH)10952–961
NhaA: Sodium ion/proton antiporter; pH regulation115401760

Synthesis of prosthetic groups of methanogenic enzymes

[NiFe] center of the [NiFe]-hydrogenases9
CarAB: Carbamoyl-phosphate synthase13790–13800996–998 (996+997 = CarB)
HypA-F: Hydrogenase maturation factors11790, 11780, 02320, 14600,06550 or 01080, 16720783, 782, 1649, 1072, 205 or 1525, 1287
FrhD and HycI: endopeptidasesSee above and 11320See above and 737
FeGP cofactor of Hmd8
HcgA-G: Hmd co-occurring proteins15270–15320, 15200 (hcgABCFED,G)1143–1148, 1137
HmdII and HmdIII00970, 089501512, 504
FeS-centers of hydrogenases, Fmd, Fwd, Hdr and ferredoxins3
IscS/SufS: Cysteine desulfurase177501389
SufB/D and SufC15340, 153301150, 1149
ApbC: Putative iron-sulfur cluster transfer proteinNot found1176
Molybdopterin cofactor of formyl-MFR dehydrogenase 10
MoaABCE01360, 04310, 12050, 059901550, 1861, 809, 149
MoeAB13850 or 17560, 015801003 or 1369, 1571
MobAB05930 or 09160, 01370143 or 528, 1551
B12 cofactor of methyltransferase MtrA-H4
HemA: Glutamyl-tRNA reductase139401012
HemB: δ-Aminolevulinic acid dehydratase11390744
HemC: Porphobilinogen deaminase12690874
HemD: Uroporphyrinogen-III synthase06170166
CysG1: Uroporphyrin-III C-methyltransferase (precorrin-2 synthesis)061801167
CbiX: Sirohydrochlorin cobalt chelatase17830141397
CbiL: Precorrin-2 C(20)-methyltransferase173801348
CbiH: Precorrin-3B C17 methylase179001403
CbiG: Cobalamin biosynthesis protein179501408
CbiF: Precorrin-4 methylase09820602
CbiD: Precorrin-6A synthase12040808
CbiE: Precorrin-6Y methylase009901514
CbiJ: Precorrin-6X reductase138401002
CbiT: Cobalt-precorrin-6Y C(15)-methyltransferase05960146
CbiC: Precorrin-8X methylmutase06760227
CbiA: Cobyrinic acid a,c-diamide synthase004601460
CobS: Cobalamin-5-phosphate synthase149601112
CbiB: Cobalamin biosynthesis protein179601409
CobN: Cobalamin biosynthesis protein06860 or 09040 or 17530237 or 514 or 1363
F430 cofactor of methyl-coenzyme reductases Biosynthesis starts from uroporphyrinogen-III, probably 7 reactions are involved?(7)?

Coenzyme biosynthesis

Coenzyme F4206
CofA: Lactaldehyde dehydrogenase13630978
CofB: L-Lactate kinase? (1)?
CofC: 2-phospho-L-lactate guanylyltransferase09930613
CofD: LPPG:FO 2-phospho-L-lactate transferase140001018
CofE: F420-0: γ-glutamyl ligase140101019
CofGH: FO synthase15760,121701198, 820
Methanofuran1
MfnA: L-Tyrosine decarboxylase150001116
and estimated 7 unknown proteins? (7)?
Methanopterin3
MptA: GTP cyclohydrolase157401196
MptB: Cyclic phosphodiesterase024601668
RFAP: Ribofuranosylaminobenzene 5′-P-synthase12280830
and estimated 7 unknown proteins? (7)?
Coenzyme M5
ComA: Phosphosulfolactate synthase025301674
ComB: 2-Phosphosulfolactate phosphatase155901182
ComC: Sulfolactate dehydrogenase158301205
ComDE: Sulfopyruvate decarboxylase15840,158501206, 1207
ComF: Sulfoacetaldehyde dehydrogenase? (1)?
Coenzyme B4
LeuA homolog: Isopropylmalate synthase00630 or 021501481 or 1630
LeuB homolog: Isopropylmalate dehydrogenase17740 or 063701388 or 184
LeuCD homolog: Isopropylmalate isomerase02160 or 17720, 12270 or 177301386 or 1631, 829 or 1387
and estimated 2 unknown proteins? (2)?

Ion transport systems

Ni2+ ABC transporter NikMNOQ (=CbiM1N1O1Q1)02830–0286041704–1707
Co2+ ABC transporter CbiM2N2O2Q205800–058304130–133
Fe2+ ABC transporter FeoAB17520, 1751021362, 1361
Zn2+ ABC transporter ZnuABC/ZupT09840–09860, 086604604–606, 473
Mg2+ transporter MgtE100101620
Ca2+ transporter (ATPase)1383011001
K+ transporter TrkAH16520, 1651021265, 1264
MoO42− ABC transporter ModA1B1C113080, 13090, 131103920, 921, 924
WO42− ABC transporter ModA2B2C208720–087403477–479
HPO42− ABC transport system PstABCS1S2/PhoU1U2
Putative transcriptional regulator of the pst/phu operon
03020–03070, 03090
03000
81727–1732, 1734
1724