TABLE 1.
Gene | Function | Cluster | Essential for C1 growtha | Metabolic moduleb | Previous name | Accession no. | Source or reference |
---|---|---|---|---|---|---|---|
Primary oxidation | |||||||
mxaW | Unknown | 1 | − | MOX | moxW | M31108 | 41 |
mxaF | Methanol dehydrogenase large subunit | 1 | + | MOX | moxF | M31108 | 1, 2, 26 |
mxaJ | Unknown | 1 | + | MOX | moxJ | M31108 | 1, 2, 26 |
mxaG | Cytochrome c1 | 1 | + | MOX | moxG | X07856 | 1, 2, 26, 30 |
mxaI | Methanol dehydrogenase small subunit | 1 | + | MOX | moxI | X15792 | 1, 26, 30, 31 |
mxaR | Unknown | 1 | + | MOX | moxR | Y07864 | 1, 26, 44 |
mxaS | Unknown | 1 | + | MOX | moxS | L41608 | 1, 26, 44 |
mxaA | Essential for Ca2+ insertion into MDH | 1 | + | MOX | moxA | L41608 | 1, 26, 29 |
mxaC | Essential for Ca2+ insertion into MDH | 1 | + | MOX | moxC | L41608 | 1, 26, 29, 44 |
mxaK | Essential for Ca2+ insertion into MDH | 1 | + | MOX | moxK | L41608 | 1, 26, 29 |
mxaL | Essential for Ca2+ insertion into MDH | 1 | + | MOX | moxL | L41608 | 1, 26, 29 |
mxaD | Essential for Ca2+ insertion into MDH | 1 | − | MOX | L41608 | 1, 26, 29, 44 | |
mxaE | Unknown | 1 | + | MOX | AF017434 | 41 | |
mxaH | Unknown | 1 | NM | MOX | AF017434 | 41 | |
mxaB | Transcriptional regulator | 1 | + | MOX | moxB | AF017434 | 41 |
mxbD | Sensor kinase | 2 | + | MOX | moxD | AF032114 | 42 |
mxbM | Transcriptional regulator | 2 | + | MOX | moxM | AF032114 | 42 |
pqqA | PQQ synthesis | 2 | − | PQQ-S | pqqD, moxT | L25889 | 26, 45, 46 |
pqqB | PQQ synthesis | 2 | + | PQQ-S | pqqG, moxO | U72662 | 26, 45 |
pqqC/D | PQQ synthesis | 2 | + | PQQ-S | pqqB, pqqC, moxV, moxT | L25889/U72662 | 26, 45 |
pqqE | PQQ synthesis | 2 | + | PQQ-S | pqqA, moxC | U72662 | 26, 45 |
mxcQ | Sensor kinase | 3 | + | MOX | moxQ | 26, 42 | |
mxcE | Transcriptional regulator | 3 | + | MOX | moxE | 26, 42 | |
pqqF | PQQ synthesis | 4 | + | PQQ-S | pqqE, moxH | L43135 | 26, 43 |
pqqG | PQQ synthesis | 4 | + | PQQ-S | pqqF, moxU | L43135 | 26, 43 |
mauF | Unknown | 5 | + | MAU | L26406 | 5 | |
mauB | Methylamine dehydrogenase large subunit | 5 | + | MAU | L26406 | 5 | |
mauE | Essential for small subunit maturation | 5 | + | MAU | L26406 | 5 | |
mauD | Essential for small subunit maturation | 5 | + | MAU | L26406 | 5 | |
mauA | Methylamine dehydrogenase small subunit | 5 | + | MAU | L26406 | 5 | |
mauC | Amicyanin | 5 | + | MAU | L26406 | 5 | |
mauJ | Unknown | 5 | − | MAU | L26406 | 5 | |
mauG | Unknown | 5 | + | MAU | L26406 | 5 | |
mauI | Unknown | 5 | + | MAU | L26406 | 5 | |
mauM | Unknown | 5 | − | MAU | L26406 | 5 | |
mauN | Unknown | 5 | − | MAU | L26406 | 5 | |
Formate oxidation | |||||||
fdh1A | Tungsten-dependent formate dehydrogenase, α subunit | 6 | − | FOX1 | AF489516 | 25 | |
fdh1B | Tungsten-dependent formate dehydrogenase, β subunit | 6 | − | FOX1 | AF489516 | 25 | |
fdh2A | Molybdenum-dependent formate dehydrogenase, α subunit | 7 | − | FOX2 | AY183757 | Unpublished | |
fdh2B | Molybdenum-dependent formate dehydrogenase, β subunit | 7 | − | FOX2 | AY183757 | Unpublished | |
fdh2C | Molybdenum-dependent formate dehydrogenase, γ subunit | 7 | − | FOX2 | AY183757 | Unpublished | |
fdh2D | Molybdenum-dependent formate dehydrogenase, δ subunit | 7 | − | FOX2 | AY183757 | Unpublished | |
fdh3A | Cytochrome-linked formate dehydrogenase, α subunit | 8 | − | FOX3 | AY181035 | Unpublished | |
fdh3B | Cytochrome-linked formate dehydrogenase, β subunit | 8 | − | FOX3 | AY181035 | Unpublished | |
fdh3C | Cytochrome-linked formate dehydrogenase, γ subunit | 8 | − | FOX3 | AY181035 | Unpublished | |
C1 transfer | |||||||
dmrA | Dihydromethanopterin reductase | 9 | + | H4MPT-T | AY093431 | 28 | |
fae | Formaldehyde activating enzyme | 2 | + | H4MPT-T | orf18 | AF032114 | 52 |
mtdB | Methylene-H4MPT dehydrogenase | 2 | + | H4MPT-T | orfX | AF032114 | 15, 18 |
mch | Methenyl-H4MPT cyclohydrolase | 2 | + | H4MPT-T | orfZ | AF032114 | 15, 36 |
fhcA | Formyltransferase/hydrolase complex, α subunit | 2 | + | H4MPT-T | orf1 | AF032114 | 15, 34, 35 |
fhcB | Formyltransferase/hydrolase complex, β subunit | 2 | + | H4MPT-T | orf2 | AF032114 | 15, 34, 35 |
fhcC | Formyltransferase/hydrolase complex, γ subunit | 2 | + | H4MPT-T | orf3 | AF032114 | 15, 34, 35 |
fhcD | Formyltransferase/hydrolase complex, δ subunit | 2 | + | H4MPT-T | ffsA | AF032114 | 15, 34, 35 |
orf4 | H4MPT biosynthesis | 2 | + | H4MPT-T | AF032114 | 15, 38 | |
orf5 | Unknown | 2 | − | H4MPT-T | AF032114 | 15 | |
orf7 | Cofactor biosynthesis | 2 | + | H4MPT-T | AF032114 | 15 | |
orf9 | Cofactor biosynthesis | 2 | + | H4MPT-T | AF032114 | 15 | |
orf17 | Cofactor biosynthesis | 2 | + | H4MPT-T | AF032114 | 15 | |
orf19 | Cofactor biosynthesis | 2 | + | H4MPT-T | AY117134 | Unpublished | |
orf20 | Cofactor biosynthesis | 2 | + | H4MPT-T | AY117134 | Unpublished | |
orf21 | Cofactor biosynthesis | 2 | + | H4MPT-T | AY117134 | Unpublished | |
orf22 | Cofactor biosynthesis | 2 | + | H4MPT-T | AY117134 | Unpublished | |
orfY | Unknown | 2 | − | H4MPT-T | AF032114 | 15 | |
mtdA | Methylene-H4MPT/Methylene-H4F dehydrogenase | 2 | + | H4F-T/H4MPT-T | L27235 | 9, 16, 17, 51 | |
fch | Methenyl-H4F cyclohydrolase | 2 | + | H4F-T | orf4 | L33465 | 36 |
ftfL | Formyl-H4F ligase | 10 | + | H4F-T | 28 | ||
folK | 2-Amino-4-hydroxy-6-hydroxymethyl-7,8-dihydropterin pyrophosphokinase | 2 | E | H4F-S | folA | U72662 | 13 |
folB | Dihydroneopterin aldolase | 2 | E | H4F-S | folB | U72662 | 13 |
folP | Dihydropteroate synthase | 2 | E | H4F-S | folC | U72662 | 13 |
folE | GTP cyclohydrolase | 11 | E | H4F-S | AY183759 | Unpublished | |
folC | Dihydrofolate synthase | 12 | E | H4F-S | AY183760 | Unpublished | |
dyr | Dihydrofolate reductase | 13 | NM | H4F-S | dfrA | AY093432 | 28 |
C1 assimilation | |||||||
sga | Serine glyoxylate aminotransferase | 2 | + | QSC | sgaA | L27235 | 9 |
hpr | Hydroxupyruvate reductase | 2 | + | QSC | hprA | L27235 | 7, 8, 9 |
ppc | PEP carboxylase | 2 | + | QSC | ppcA | U72662 | 4 |
mtkA | Malate thiokinase, α subunit | 2 | + | QSC/GRC | L33465 | 10 | |
mtkB | Malate thiokinase, β subunit | 2 | + | QSC/GRC | L33465 | 10 | |
mcl | Malyl-CoA lyase | 2 | + | QSC/GRC | mclA | U72662 | 4, 13 |
glyA | Serine hydroxymethyltrasferase | 14 | + | QSC | L33463 | 11 | |
gck | Glycerate kinase | 15 | + | QSC | gckA | U87316 | 14 |
eno | Enolase | 16 | E | QSC/GNG | AY181039 | Unpublished | |
mdh | Malate dehydrogenase | 17 | E | QSC/TCAC | AY181040 | Unpublished | |
qscR | Serine cycle transcriptional regulator | 18 | + | QSC | AF516903 | 20 | |
phaA | β-ketothiolase | 19 | + | GRC/PHBC | AF287907 | 22, 23, 24 | |
phaB | Acetoacetyl-CoA reductase | 19 | + | GRC/PHBC | AF287907 | 22, 23, 24 | |
phaR | Regulator of PHB synthesis and acetyl-CoA flux | 19 | + | GRC/PHBC | orf3 | AF287907 | 23, 24 |
croR | Crotonase | 20 | + | GRC | AF416777 | 22 | |
pccA | Propionyl-CoA carboxylase, α subunit | 21 | + | GRC | AY181038 | 22 | |
pccB | Propionyl-CoA carboxylase, β subunit | 22 | + | GRC | pccA | L48340 | 12, 22 |
crr | Crotonyl-CoA reductase | 22 | + | GRC | adhA | L48340 | 12, 22 |
meaA | A mutase | 22 | + | GRC | U28335 | 12, 22, 40 | |
meaB | Essential for methylmalonyl-CoA mutase reaction | 22 | + | GRC | AF416776 | 22 | |
meaC | Unknown | 23 | + | GRC | 28 | ||
meaD | Unknown | 24 | + | GRC | 28 | ||
ibd2 | Isobutyryl-CoA dehydrogenase | 25 | + | GRC | AY054980 | 22 | |
mcmA | Methylmalonyl-CoA mutase, α subunit | 26 | + | GRC | AY181037 | 22 | |
mcmB | Methylmalonyl-CoA mutase, β subunit | 27 | + | GRC | 22 | ||
epm | Epimerase | 28 | + | GRC | AY183758 | Unpublished | |
sdhA | Succinate dehydrogenase flavoprotein subunit | 29 | EG | GRC/TCAC | AY181036 | 48, 49 | |
sdhB | Succinate dehydrogenase iron-sulfur binding subunit | 29 | NM | GRC/TCAC | AY181036 | 48, 49 | |
sdhC | Succinate dehydrogenase cytochrome B556 subunit | 29 | NM | GRC/TCAC | AY181036 | 48, 49 | |
sdhD | Succinate dehydrogenase membrane anchor subunit | 29 | NM | GRC/TCAC | AY181036 | 48, 49 | |
fumA | Fumarase | 13 | EG | GRC/TCAC | AF497854 | 49 |
+, mutation impairs growth on a C1 compound; −, mutation does not impair growth on a C1 compound; NM, no mutant available: involvement in methylotrophy implicated by regulation, location, or phenotype of mutant in other subunit(s); E, essential for both methylotrophic and heterotrophic growth; EG, growth on C1 compounds rescued by glyoxylate.
Metabolic module designations: MOX, methanol oxidation; PQQ-S, PQQ biosynthesis; MAU, methylamine utilization; H4F-T, H4F-linked C1 transfer; H4F-S, H4F biosynthesis; H4MPT-TS, H4MPT-linked C1 transfer and H4MPT biosynthesis; FOX1, FOX2, and FOX3, three nonhomologous formate dehydrogenases; QSC, (Quayle) serine cycle; PHBC, PHB cycle; TCAC, TCA cycle.