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. 2020 Jan 24;120(12):4921–4968. doi: 10.1021/acs.chemrev.9b00489

Table 1. A. vinelandii Genes Involved in Mo-Dependent N2 Fixation and Roles of Their Products.

genea Nif phenotypeb identity/role(s)
nifH Fe protein. Obligate electron donor to the MoFe protein. Also required for FeMo-co biosynthesis and apo-MoFe maturation.
nifD MoFe protein α-subunit. FeMo-co binding subunit. Forms an α2β2 tetramer with NifK.
nifK MoFe protein β-subunit. Forms an α2β2 tetramer with NifD.
nifT + Unknown
nifY + Interacts with apo-MoFe protein prior to FeMo-co insertion.
nafA + Unknown function.
nifE Scaffold for NifB-co maturation into FeMo-co. Forms an α2β2 tetramer with NifN.
nifN Scaffold for NifB-co maturation into FeMo-co. Forms an α2β2 tetramer with NifE.
nifX + Binds NifB-co and VK-cluster. Interacts with apo-NifEN. Proposed carrier of FeMo-co precursors.
nafC + Unknown function.
feSII + Shethna protein II. Nitrogenase protection against O2.
nifIscA + Fe–S scaffolding Nif protein. Contains a [4Fe-4S] cluster.
nifU Supplies [Fe–S] clusters precursors for nitrogenase proteins and cofactors.
nifS ± Mobilizes S for [Fe–S] cluster synthesis on NifU.
nifV ± Homocitrate synthase. Involved in FeMo-co synthesis.
cysE1 + Serine O-acetyltransferase. Participates in cysteine formation.
nafH + Binds to apo-MoFe protein preceding P-cluster maturation.
nifW ± Associates with apo-MoFe protein preceding P-cluster maturation.
nifZ ± Involved in P-cluster maturation in combination with the Fe protein.
nifM Essential for the maturation of the Fe protein. Proposed peptidyl-prolyl cis–trans isomerases that would act over the Pro258 residue of NifH.
clpx2 + Regulates NifEN and NifB protein levels.
nifF + Flavodoxin. Physiological electron donor to the Fe protein.
nifL + Sensor protein. Acts as NifA antiactivator.
nifA Transcriptional activator. Forms a two-component regulatory system with NifL.
nifB Essential for FeMo-co synthesis. Catalyzes NifB-co formation, the first committed step in FeMo-co biosynthesis.
fdxN ± 2x[4Fe-4S] ferredoxin. Required for NifB in the formation of NifB-co.
nifO + Required for simultaneous activation of nitrogenase and nitrate reductase.
nifQ Processes Mo prior to its incorporation into FeMo-co.
nafW + Rhodanase-like protein. Also called rhdN. Unknown function.
nafZ + Glutaredoxin-like protein. Also called grx5nif. Unknown function.
rnfAc ± Transmembrane protein. Part of the Rnf complex, which is involved in supplying low potential electrons to the nitrogenase.
rnfBc ± Contains two [Fe–S] cluster binding sites. It is part of the Rnf complex.
rnfCc ± Contains [4Fe-4S] binding site. Electron transport component of the Rnf complex.
rnfDc ± Transmembrane protein of the Rnf complex. Includes a flavin binding motive.
rnfGc ± Last electron acceptor component of the Rnf complex. Contains FMN binding site.
rnfEc ± Transmembrane protein of the Rnf complex.
rnfHc ± Soluble protein of the rnf gene cluster.
nafY + In A. vinelandii, γ subunit of apo-MoFe protein. Stabilizes apo-MoFe protein prior to FeMo-co insertion. Also binds FeMo-co specifically.
mosA + Mo storage protein α subunit. Forms an α3β3 hexamer with MosB.
mosB + Mo storage protein β subunit.
a

Minimum set of essential Mo-nitrogenase genes is shown in bold.

b

+ Diazotrophic growth, ± slow diazotrophic growth, – no diazotrophic growth.

c

Phenotype of rnf genes was determined by deleting the entire rnf operon.