Decoupling filamentous phage uptake and energy of the TolQRA motor in Escherichia coli. Samire P, Serrano B, Duche D, Lemarie E, Lloubes R, Houot L. J Bacteriol. 2019 Oct 21. doi: 10.1128/JB.00428-19.

Cryo-EM structure of the bacterial Ton motor subcomplex ExbB-ExbD provides information on structure and stoichiometry. Celia H, Botos I, Ni X, Fox T, De Val N, Lloubes R, Jiang J, Buchanan SK. Nature Commun Biol. 2019 Oct 4;2:358.

Tol Energy-Driven Localization of Pal and Anchoring to the Peptidoglycan Promote Outer-Membrane Constriction. Petiti M, Serrano B, Faure L, Lloubes R, Mignot T, Duché D. J Mol Biol. 431(17):3275-3288, 2019

Connected partner-switches control the life style of Pseudomonas aeruginosa through RpoS regulation. Bouillet S., Bab M., Houot L., Iobbi-Nivola C. and Bordi C. Sci Rep. 9(1):6496, 2019

Similarities and Differences between Colicin and Filamentous Phage Uptake by Bacterial Cells. Duché D. et Houot L. EcoSal Plus. 8(2), 2019



Active transport across the bacterial outer membrane: the Ton motor complex. S. K. Buchanan, C. Hervé, N. Noinaj, S. D. Zakharov, E. Bordignon, I. Botos, M. Santamaria, T. J. Barnard, W. A. Cramer and R. Lloubes. Acta Cryst. 2018. A74, a79


Cell Fractionation. Petiti M, Houot L, Duché D. Methods Mol Biol. 2017;1615:59-64.

Large Complexes: Cloning Strategy, Production, and Purification. Durand E, Lloubes R. Methods Mol Biol. 2017;1615:299-309.

Electrostatic interactions between the CTX phage minor coat protein and the bacterial host receptor TolA drive the pathogenic conversion of Vibrio cholerae. Houot L , Navarro R, Nouailler M, Duché D , Guerlesquin F, Lloubes R. J Biol Chem. 2017 Aug 18;292(33):13584-13598.

Host cell interactions of outer membrane vesicle-associated virulence factors of enterohemorrhagic Escherichia coli O157: Intracellular delivery, trafficking and mechanisms of cell injury. Bielaszewska M, Rüter C, Bauwens A, Greune L, Jarosch KA, Steil D, Zhang W, He X, Lloubes R , Fruth A, Kim KS, Schmidt MA, Dobrindt U, Mellmann A, Karch H. PLoS Pathog. 2017 Feb 3;13(2):e1006159.



Structural insight into the role of the Ton complex in energy transduction. Célia H , Noinaj N, Zakharov SD, Bordignon E, Botos I, Santamaria M, Cramer WA, Lloubes R and Buchanan SK. Nature. 2016 Oct 6;538(7623):60-65.

1H, 15N and 13C resonance assignments of the C-terminal domain of Vibrio cholerae TolA protein. Navarro R, Bornet O, Houot L , Lloubes R , Guerlesquin F Nouailler M. Biomol NMR Assign. 2016 Oct;10(2):311-3.



Virulence from vesicles: Novel mechanisms of host cell injury by Escherichia coli O104:H4 outbreak strain. Kunsmann L, Rüter C, Bauwens A, Greune L, Glüder M, Kemper B, Fruth A, Wai SN, He X, Lloubes R , Schmidt MA, Dobrindt U, Mellmann A, Karch H, Bielaszewska M. Sci Rep. 2015 5: 13252.


Type VI secretion and bacteriophage tail tubes share a common assembly pathway. Brunet YR, Hénin J, Celia H , Cascales E . EMBO Rep. 15(3):315-21.


Non classical secretion systems. Lloubes R, Bernadac A, Houot L , Pommier S . Res Microbiol. 2013 Jul-Aug;164(6):655-63. Review.

Mannitol and the mannitol-specific enzyme IIB subunit activate Vibrio cholerae biofilm formation. Ymele-Leki P, Houot L , Watnick PI. Appl Environ Microbiol. 2013 Aug;79(15):4675-83.



A bacterial two-hybrid genome fragment library for deciphering regulatory networks of the opportunistic pathogen Pseudomonas aeruginosa. Houot L , Fanni A, de Bentzmann S, Bordi C. Microbiology. 2012 Aug;158(Pt 8):1964-71.

Colicin M, a peptidoglycan lipid-II-degrading enzyme: potential use for antibacterial means? Touzé T, Barreteau H, El Ghachi M, Bouhss A, Barnéoud-Arnoulet A , Patin D, Sacco E, Blanot D, Arthur M, Duché D , Lloubès R , Mengin-Lecreulx D. Biochem Soc Trans. 2012 Dec 1;40(6):1522-7.

Energetics of colicin import revealed by genetic cross-complementation between the Tol and Ton systems. Lloubes R, Goemaere E, Zhang X, Cascales E, Duché D. Biochem Soc Trans.2012, 40(6):1480-5.

The C-tail anchored TssL subunit, an essential protein of the enteroaggregative Escherichia coli Sci-1 Type VI secretion system, is inserted by YidC. Aschtgen MS, Zoued A, Lloubes R , Journet L, Cascales E. Microbiologyopen. 2012, 1(1):71-82.

Characterization of colicin M and its orthologs targeting bacterial cell wall peptidoglycan biosynthesis. Barreteau H, El Ghachi M, Barnéoud-Arnoulet A , Sacco E, Touzé T, Duché D , Gérard F, Brooks M, Patin D, Bouhss A, Blanot D, van Tilbeurgh H, Arthur M, Lloubes R , Mengin-Lecreulx D. Microb Drug Resist. 2012, 18(3):222-9.

Prominent role of the three Synechocystis PchR-like regulators in the defense against metal and oxidative stresses. Soni B, Houot L , Cassier-Chauvat C, Chauvat F. Open Journal of Biochemistry. 2012, 1, 1.



An epigenetic switch involving overlapping fur and DNA methylation optimizes expression of a type VI secretion gene cluster. Brunet YR, Bernard CS, Gavioli M , Lloubes R , Cascales E. PLoS Genet. 7(7):e1002205.

Regulation of type VI secretion gene clusters by sigma54 and cognate enhancer binding proteins. Bernard CS, Brunet YR, Gavioli M , Lloubes R , Cascales E. J Bacteriol. 193(9):2158-67.

Mapping the interactions between Escherichia coli TolQ transmembrane segments. Zhang XY, Goemaere EL, Seddiki N, Célia, Gavioli M, Cascales E, Lloubes R. J Biol Chem. 286(13):11756-64.

High water solubility and fold in amphipols of proteins with large hydrophobic regions: oleosins and caleosin from seed lipid bodies. Gohon Y, Vindigni JD, Pallier A, Wien F, Celia H , Giuliani A, Tribet C, Chardot T, Briozzo P. Biochim Biophys Acta. 1808(3):706-16.



Channel domain of colicin A modifies the dimeric organization of its immunity protein. Zhang XY, Lloubes R, Duché D. J Biol Chem. 285(49):38053-61.

Interaction of the colicin K bactericidal toxin with components of its import machinery in the periplasm of Escherichia coli. Barnéoud-Arnoulet A, Gavioli M, Lloubes R, Cascales E. J Bacteriol. 192(22):5934-42.

Toxicity of the colicin M catalytic domain exported to the periplasm is FkpA independent. Barnéoud-Arnoulet A , Barreteau H, Touzé T, Mengin-Lecreulx D, Lloubes R , Duché D. J Bacteriol 192(19):5212-9.

The SciZ protein anchors the enteroaggregative Escherichia coli Type VI secretion system to the cell wall. Aschtgen MS , Gavioli M , Dessen A, Lloubes R , Cascales E. Mol Microbiol. 275(4):886-99.

Deciphering the catalytic domain of colicin M, a peptidoglycan lipid II-degrading enzyme. Barreteau H, Bouhss A, Gérard F, Duché D , Boussaid B, Blanot D, Lloubes R, Mengin-Lecreulx D, Touzé T. J Biol Chem. 285(16):12378-89.

The PTS regulates Vibrio cholerae biofilm formation through multiple independent pathways. Houot L. , Chang S., Absalon C., Pickering B., and Watnick PI. J. Bacteriol 192(12) :3055-67.

Vibrio cholerae PTS control of carbohydrate transport, biofilm formation, and colonization of the germ-free mouse intestine. Houot L. , Chang S., Absalon C., and Watnick P.I. Infect and Immunity 78(4):1482-94.

Determinants of the proton selectivity of the colicin A channel. S. Slatin, D. Duché and D. Baty. Biochemistry. 49; 4786-93.

Components of E. coli Energy-Transducing Complexes, ExbB and TolQ, Display Ion Channels. Zakharov S, Célia H , Datsenko K, Santamaria M, Whitelegge J, Lloubes R , Cramer W. Biophys J. vol. 98, issue 3, pp. 540a.



Mapping the interactions between escherichia coli tol subunits: rotation of the TolR transmembrane helix. Zhang XY, Goemaere EL, Thomé R, Gavioli M, Cascales E, Lloubes R. J Biol Chem. 284(7):4275-82.

Immunity protein protects colicin E2 from OmpT protease. Duché D, Issouf M, Lloubes R. J Biochem. 145(1):95-101.

FpvA bound to non-cognate pyoverdines: molecular basis of siderophore recognition by an iron transporter. Greenwald J, Nader M, Celia H , Gruffaz C, Geoffroy V, Meyer JM, Schalk IJ, Pattus F. Mol Microbiol. 72(5):1246-59.


SciN is an outer membrane lipoprotein required for type VI secretion in enteroaggregative Escherichia coli. Aschtgen MS, Bernard CS, De Bentzmann S, Lloubes R , Cascales E. J Bacteriol. 190(22):7523-31.

Visualization of interactions between siderophore transporters and the energizing protein TonB by native PAGE. Choul-Li S, Adams H, Pattus F, Celia H. Electrophoresis. 29(6):1333-8.

The metal dependence of pyoverdine interactions with its outer membrane receptor FpvA. Greenwald J, Zeder-Lutz G, Hagege A, Celia H , Pattus F. J Bacteriol. 190(20):6548-58.

Carbon nanotubes as templates for polymerized lipid assemblies. Thauvin C, Rickling S, Schultz P, Célia H , Meunier S, Mioskowski C. Nat Nanotechnol. 3(12):743-8.


Movements of the TolR C-terminal domain depend on TolQR ionizable key residues and regulate activity of the Tol complex

Goemaere EL, Devert A, Lloubes R, Cascales E. J Biol Chem. 282(24):17749-57.

Colicin biology. Cascales E, Buchanan SK, Duché D, Kleanthous C, Lloubes R, Postle K, Riley M, Slatin S, Cavard D. Microbiol Mol Biol Rev. 71(1):158-229.

Mutational analyses define helix organization and key residues of a bacterial membrane energy-transducing complex.

Goemaere EL, Cascales E, Lloubes R. J Mol Biol. 366(5):1424-36.

Colicin E2 is Still in Contact with its Receptor and Import Machinery when its Nuclease Domain enters the Cytoplasm.

D. Duché. J. Bacteriol. 189, 4217−22.


Release of immunity protein requires functional endonuclease colicin import machinery. Duché D, Frenkian A, Prima V, Lloubes R. J Bacteriol. 188(24):8593-600.



Tol-dependent macromolecule import through the Escherichia coli cell envelope requires the presence of an exposed TolA binding motif.

Pommier S, Gavioli M, Cascales E, Lloubes R. J Bacteriol. 187(21):7526-34.

Solution structure of the E.coli TolA C-terminal domain reveals conformational changes upon binding to the phage g3p N-terminal domain.

Deprez C, Lloubes R , Gavioli M , Marion D, Guerlesquin F, Blanchard L. J Mol Biol. 346(4):1047-57.

Combining high-field EPR with site-directed spin labeling reveals unique information on proteins in action. K. Möbius, A. Savitsky, C. Wegener, M. Plato, M. Fuchs, A Schnegg, A. A. Dubinskii, Y. A. Grishin, I. A. Grigorev, M Kühn, D. Duché , H Zimmermann and H.J. Steinhoff. Magn. Reson. Chem. 43, S4-S19.


Improved methods for producing outer membrane vesicles in Gram-negative bacteria. Henry T, Pommier S , Journet L , Bernadac A, Gorvel JP, Lloubes R. Res Microbiol. 155(6):437-46.

Deletion analyses of the peptidoglycan-associated lipoprotein Pal reveals three independent binding sequences including a TolA box. Cascales E, Lloubes R. Mol Microbiol. 51(3):873-85.

Gating movements of colicin A and colicin Ia are different. S.L. Slatin, D. Duché , P.K. Kienker and D. Baty. J. Memb. Biol. 202, 73-83.

Spontaneous refolding of the pore-forming Colicin A toxin upon membrane association as studied by X-band and W-band high-field EPR spectroscopy.

Savitsky, M. Kühn, D. Duché , K. Möbtus and H-J. Steinhoff . J. Phys. Chem. B. 108 , 9541-9548.


Analysis of the Escherichia coli Tol-Pal and TonB systems by periplasmic production of Tol, TonB, colicin, or phage capsid soluble domains. Bouveret E, Journet L , Walburger A, Cascales E , Bénédetti H, Lloubes R. Biochimie. 84(5-6):413-21.

Macromolecular import into Escherichia coli: the TolA C-terminal domain changes conformation when interacting with the colicin A toxin. Deprez C, Blanchard L, Guerlesquin F, Gavioli M , Simorre JP, Lazdunski C, Marion D, Lloubes R. Biochemistry. 2002 Feb 26;41(8):2589-98.

Pal lipoprotein of Escherichia coli plays a major role in outer membrane integrity. Cascales E , Bernadac A, Gavioli M , Lazzaroni JC, Lloubes R. J Bacteriol. 184(3):754-9.

Translocation of a functional protein by a voltage-dependent ion channel. S.L. Slatin, A. Nardi, K. Jakes, D. Baty and D. Duché. Proc. Natl. Acad. Sci., USA, 99, 1286-91.

The pore-forming domain of colicin A fused to a signal peptide : a tool for studying pore-formation and inhibition. D. Duché. Biochimie. 84, 455-464.


The TolQ-TolR proteins energize TolA and share homologies with the flagellar motor proteins MotA-MotB. Cascales E , Lloubes R , Sturgis JN. Mol Microbiol. 42(3):795-807.

The C-terminal half of the colicin A pore-forming domain is active in vivo and in vitro. Nardi, S. L. Slatin, D. Baty and D. Duché. J. Mol. Biol. 307 : 1293-1303.

Colicin A immunity protein interacts with the hydrophobic helical hairpin of the colicin A channel domain in the Escherichia coli inner membrane. Nardi, Y. Corda, D. Baty and D. Duché. J. Bacteriol. 183 ; 6721-6725.

Import of colicins across the outer membrane of Escherichia coli involves multiple protein interactions in the periplasm. Journet L , Bouveret E, Rigal A, Lloubes R , Lazdunski C, Bénédetti H. Mol Microbiol 42(2):331-44.

The Tol-Pal proteins of the Escherichia coli cell envelope: an energized system required for outer membrane integrity? Lloubes R, Cascales E , Walburger A, Bouveret E, Lazdunski C, Bernadac A, Journet L. Res Microbiol. 152(6):523-9.

Crystallization and preliminary crystallographic study of the periplasmic domain of the Escherichia coli TolR protein. Abergel C, Journet L , Chenivesse S, Gavioli M , Lloubes R. Acta Crystallogr D Biol Crystallogr. 57(Pt 2):323-5.


Impairment of bile salt-dependent lipase secretion in human pancreatic tumoral SOJ-6 cells. Caillol N, Pasqualini E, Lloubes R , Lombardo D. J Cell Biochem. 79(4):628-47.

Phosphorylation of the rat pancreatic bile-salt-dependent lipase by casein kinase II is essential for secretion. Pasqualini E, Caillol N, Valette A, Lloubes R , Verine A, Lombardo D. Biochem J. 1;345 Pt 1:121-8.

Proton motive force drives the interaction of the inner membrane TolA and outer membrane pal proteins in Escherichia coli.

Cascales E , Gavioli M , Sturgis JN, Lloubes R. Mol Microbiol. 38(4):904-15.

Colicin import into Escherichia coli cells requires the proximity of the inner and outer membranes and other factors.

Lazdunski C, Bouveret E, Rigal A, Journet L , Lloubes R , Bénédetti H. Int J Med Microbiol. 290(4-5):337-44.

Assignment of the 1H, 15N and 13C resonances of the C-terminal domain of the TolA protein of Escherichia coli, involved in cell envelope integrity.

Deprez C, Blanchard L, Simorre JP, Gavioli M , Guerlesquin F, Lazdunski C, Lloubes R , Marion D. J Biomol NMR. 18(2):179-80.


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