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Categories containing this gene/protein

carbon core metabolism, trigger enzyme, RNA binding regulators, most abundant proteins

This gene is a member of the following regulons

CcpA regulon, CcpC regulon, CodY regulon, FsrA regulon

The CitB regulon


  • Coordinates on the chromosome (coding sequence): 1,926,680 -> 1,929,409
  • Phenotypes of a mutant

  • glutamate auxotrophy and a defect in sporulation PubMed
  • The protein

    Catalyzed reaction/ biological activity

  • Citrate <=> isocitrate
  • Binding to iron responsive elements (IRE RNA) in the absence of the FeS cluster PubMed
  • Protein family

    Paralogous protein(s)

    Kinetic information




  • FeS cluster
  • Effectors of protein activity


  • 1L5J (E. coli)
  • Localization


  • Mdh-CitB PubMed
  • CitG-CitB PubMed
  • CitB-SufU (insertion of the Fe-S cluster) PubMed
  • Additional information

  • B. subtilis aconitase is both an enzyme and an RNA binding protein (moonlighting protein) PubMed
  • extensive information on the structure and enzymatic properties of CitB can be found at Proteopedia
  • Expression and Regulation


  • citB-yneN PubMed
  • Sigma factor

  • SigA PubMed
  • Regulation

  • repressed during growth in the presence of branched chain amino acids (CodY) PubMed
  • repressed in the presence of glucose and glutamate (CcpC) PubMed
  • expressed upon transition into the stationary phase (AbrB) PubMed, indirect negative regulation by AbrB PubMed
  • repressed by glucose (3.7-fold) (CcpA) PubMed
  • repression by glucose arginine (CcpC) PubMed
  • less expressed under conditions of extreme iron limitation (FsrA) PubMed
  • part of the iron sparing response (FsrA) PubMed
  • Regulatory mechanism

  • CodY: transcription repression PubMed
  • CcpC: transcription repression (molecular inducer: citrate) PubMed1 PubMed2
  • CcpA: transcription repression PubMed
  • FsrA: translational repression PubMed
  • Additional information

  • belongs to the 100 most abundant proteins PubMed
  • Biological materials


  • GP1275 (citB::erm), available in Jörg Stülke's lab
  • GP1441 (spc), available in Jörg Stülke's lab
  • 1A999 (citB::spec), PubMed, available at BGSC
  • GP2338 (citB::kan), Cre-recombinase is integrated in sacA, available in Jörg Stülke's lab
  • GP2339 (citB::lox72), Cre-recombinase is integrated in sacA, available in Jörg Stülke's lab
  • Expression vector

    lacZ fusion

  • pGP700 (in pAC5), available in Jörg Stülke's lab
  • GFP fusion

  • GP1434 (spc, based on pGP1870), available in Jörg Stülke's lab
  • two-hybrid system

  • B. pertussis adenylate cyclase-based bacterial two hybrid system (BACTH), available in Jörg Stülke's lab
  • FLAG-tag construct

  • GP1144 (spc, based on pGP1331), available in Jörg Stülke's lab
  • GP1145 (kan), available in Jörg Stülke's lab
  • Antibody

  • available in Linc Sonenshein's lab
  • Labs working on this gene/protein

  • Linc Sonenshein, Tufts University, Boston, MA, USA Homepage
  • Jörg Stülke, University of Göttingen, Germany Homepage
  • References


    Volz K

    The functional duality of iron regulatory protein 1

    Curr Opin Struct Biol. 2008 Feb;18(1):106-11. doi: 10.1016/ Epub 2008 Feb 7. Review. PubMed PMID: 18261896; PubMed Central PMCID: PMC2374851.
    Commichau FM, Stülke J

    Trigger enzymes: bifunctional proteins active in metabolism and in controlling gene expression

    Mol Microbiol. 2008 Feb;67(4):692-702. Epub 2007 Dec 11. Review. PubMed PMID: 18086213.
    Kiley PJ, Beinert H

    The role of Fe-S proteins in sensing and regulation in bacteria

    Curr Opin Microbiol. 2003 Apr;6(2):181-5. Review. PubMed PMID: 12732309.
    Switzer RL

    Non-redox roles for iron-sulfur clusters in enzymes

    Biofactors. 1989 Dec;2(2):77-86. Review. PubMed PMID: 2696478.

    Original publications

    Pechter KB, Meyer FM, Serio AW, Stülke J, Sonenshein AL

    Two roles for aconitase in the regulation of tricarboxylic acid branch gene expression in Bacillus subtilis

    J Bacteriol. 2013 Apr;195(7):1525-37. doi: 10.1128/JB.01690-12. Epub 2013 Jan 25. PubMed PMID: 23354745; PubMed Central PMCID: PMC3624536.
    Mittal M, Pechter KB, Picossi S, Kim HJ, Kerstein KO, Sonenshein AL

    Dual role of CcpC protein in regulation of aconitase gene expression in Listeria monocytogenes and Bacillus subtilis

    Microbiology. 2013 Jan;159(Pt 1):68-76. doi: 10.1099/mic.0.063388-0. Epub 2012 Nov 8. PubMed PMID: 23139400; PubMed Central PMCID: PMC3542727.
    Smaldone GT, Revelles O, Gaballa A, Sauer U, Antelmann H, Helmann JD

    A global investigation of the Bacillus subtilis iron-sparing response identifies major changes in metabolism

    J Bacteriol. 2012 May;194(10):2594-605. doi: 10.1128/JB.05990-11. Epub 2012 Mar 2. PubMed PMID: 22389480; PubMed Central PMCID: PMC3347201.
    Brown CT, Fishwick LK, Chokshi BM, Cuff MA, Jackson JM 4th, Oglesby T, Rioux AT, Rodriguez E, Stupp GS, Trupp AH, Woollcombe-Clarke JS, Wright TN, Zaragoza WJ, Drew JC, Triplett EW, Nicholson WL

    Whole-genome sequencing and phenotypic analysis of Bacillus subtilis mutants following evolution under conditions of relaxed selection for sporulation

    Appl Environ Microbiol. 2011 Oct;77(19):6867-77. doi: 10.1128/AEM.05272-11. Epub 2011 Aug 5. PubMed PMID: 21821766; PubMed Central PMCID: PMC3187084.
    Albrecht AG, Landmann H, Nette D, Burghaus O, Peuckert F, Seubert A, Miethke M, Marahiel MA

    The frataxin homologue Fra plays a key role in intracellular iron channeling in Bacillus subtilis

    Chembiochem. 2011 Sep 5;12(13):2052-61. doi: 10.1002/cbic.201100190. Epub 2011 Jul 8. PubMed PMID: 21744456.
    Gao W, Dai S, Liu Q, Xu H, Bai Y, Qiao M

    Effect of site-directed mutagenesis of citB on the expression and activity of Bacillus subtilis aconitase

    Mikrobiologiia. 2010 Nov-Dec;79(6):774-8. PubMed PMID: 21446632.
    Gao W, Dai S, Liu Q, Xu H, Qiao M

    CitB mutation increases the alkaline protease productivity in Bacillus subtilis

    J Gen Appl Microbiol. 2010 Oct;56(5):403-7. PubMed PMID: 21099137.
    Meyer FM, Gerwig J, Hammer E, Herzberg C, Commichau FM, Völker U, Stülke J

    Physical interactions between tricarboxylic acid cycle enzymes in Bacillus subtilis: evidence for a metabolon

    Metab Eng. 2011 Jan;13(1):18-27. doi: 10.1016/j.ymben.2010.10.001. Epub 2010 Oct 8. PubMed PMID: 20933603.
    Chumsakul O, Takahashi H, Oshima T, Hishimoto T, Kanaya S, Ogasawara N, Ishikawa S

    Genome-wide binding profiles of the Bacillus subtilis transition state regulator AbrB and its homolog Abh reveals their interactive role in transcriptional regulation

    Nucleic Acids Res. 2011 Jan;39(2):414-28. doi: 10.1093/nar/gkq780. Epub 2010 Sep 3. PubMed PMID: 20817675; PubMed Central PMCID: PMC3025583.
    Albrecht AG, Netz DJ, Miethke M, Pierik AJ, Burghaus O, Peuckert F, Lill R, Marahiel MA

    SufU is an essential iron-sulfur cluster scaffold protein in Bacillus subtilis

    J Bacteriol. 2010 Mar;192(6):1643-51. doi: 10.1128/JB.01536-09. Epub 2010 Jan 22. PubMed PMID: 20097860; PubMed Central PMCID: PMC2832514.
    Gaballa A, Antelmann H, Aguilar C, Khakh SK, Song KB, Smaldone GT, Helmann JD

    The Bacillus subtilis iron-sparing response is mediated by a Fur-regulated small RNA and three small, basic proteins

    Proc Natl Acad Sci U S A. 2008 Aug 19;105(33):11927-32. doi: 10.1073/pnas.0711752105. Epub 2008 Aug 12. PubMed PMID: 18697947; PubMed Central PMCID: PMC2575260.
    Serio AW, Sonenshein AL

    Expression of yeast mitochondrial aconitase in Bacillus subtilis

    J Bacteriol. 2006 Sep;188(17):6406-10. PubMed PMID: 16923908; PubMed Central PMCID: PMC1595382.
    Serio AW, Pechter KB, Sonenshein AL

    Bacillus subtilis aconitase is required for efficient late-sporulation gene expression

    J Bacteriol. 2006 Sep;188(17):6396-405. Erratum in: J Bacteriol. 2007 Jul;189(14):5403. PubMed PMID: 16923907; PubMed Central PMCID: PMC1595401.
    Blencke HM, Reif I, Commichau FM, Detsch C, Wacker I, Ludwig H, Stülke J

    Regulation of citB expression in Bacillus subtilis: integration of multiple metabolic signals in the citrate pool and by the general nitrogen regulatory system

    Arch Microbiol. 2006 Mar;185(2):136-46. Epub 2006 Jan 5. PubMed PMID: 16395550.
    Eymann C, Dreisbach A, Albrecht D, Bernhardt J, Becher D, Gentner S, Tam le T, Büttner K, Buurman G, Scharf C, Venz S, Völker U, Hecker M

    A comprehensive proteome map of growing Bacillus subtilis cells

    Proteomics. 2004 Oct;4(10):2849-76. PubMed PMID: 15378759.
    Blencke HM, Homuth G, Ludwig H, Mäder U, Hecker M, Stülke J

    Transcriptional profiling of gene expression in response to glucose in Bacillus subtilis: regulation of the central metabolic pathways

    Metab Eng. 2003 Apr;5(2):133-49. PubMed PMID: 12850135.
    Kim HJ, Kim SI, Ratnayake-Lecamwasam M, Tachikawa K, Sonenshein AL, Strauch M

    Complex regulation of the Bacillus subtilis aconitase gene

    J Bacteriol. 2003 Mar;185(5):1672-80. PubMed PMID: 12591885; PubMed Central PMCID: PMC148081.
    Jourlin-Castelli C, Mani N, Nakano MM, Sonenshein AL

    CcpC, a novel regulator of the LysR family required for glucose repression of the citB gene in Bacillus subtilis

    J Mol Biol. 2000 Jan 28;295(4):865-78. PubMed PMID: 10656796.
    Alén C, Sonenshein AL

    Bacillus subtilis aconitase is an RNA-binding protein

    Proc Natl Acad Sci U S A. 1999 Aug 31;96(18):10412-7. PubMed PMID: 10468622; PubMed Central PMCID: PMC17902.
    Craig JE, Ford MJ, Blaydon DC, Sonenshein AL

    A null mutation in the Bacillus subtilis aconitase gene causes a block in Spo0A-phosphate-dependent gene expression

    J Bacteriol. 1997 Dec;179(23):7351-9. PubMed PMID: 9393699; PubMed Central PMCID: PMC179685.
    Fisher SH, Magasanik B

    2-Ketoglutarate and the regulation of aconitase and histidase formation in Bacillus subtilis

    J Bacteriol. 1984 Apr;158(1):379-82. PubMed PMID: 6143742; PubMed Central PMCID: PMC215431.
    Rosenkrantz MS, Dingman DW, Sonenshein AL

    Bacillus subtilis citB gene is regulated synergistically by glucose and glutamine

    J Bacteriol. 1985 Oct;164(1):155-64. PubMed PMID: 2413006; PubMed Central PMCID: PMC214224.
    Fouet A, Jin SF, Raffel G, Sonenshein AL

    Multiple regulatory sites in the Bacillus subtilis citB promoter region

    J Bacteriol. 1990 Sep;172(9):5408-15. PubMed PMID: 2118511; PubMed Central PMCID: PMC213206.
    Fouet A, Sonenshein AL

    A target for carbon source-dependent negative regulation of the citB promoter of Bacillus subtilis

    J Bacteriol. 1990 Feb;172(2):835-44. PubMed PMID: 2105305; PubMed Central PMCID: PMC208513.
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