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Escherichia coli K-12 substr. MG1655 Transporter: mechanosensitive channel MscL



Gene: mscL Accession Numbers: EG11180 (EcoCyc), b3291, ECK3277

Synonyms: yhdC

Regulation Summary Diagram: ?

Subunit composition of mechanosensitive channel MscL = [MscL]5

Summary:
MscL is the mechanosensitive channel of large conductance that is involved in the transduction of membrane stress (mechanical or osmotic) into an electrochemical response [Oakley99]. Upon osmotic downshift in the cell's environment, water rapidly flows in, increasing pressure on the membrane. MscL responds to this membrane tension by opening a large pore that allows solutes to exit the cell. In this way MscL acts as a pressure release valve, preventing lysis due to increased osmotic pressure.

The protein was originally discovered by patch clamp studies on protein fractions reconstituted into liposomes [Sukharev94, Sukharev97]. The export of ions is fairly nonspecific, with slight selectivity for cations over anions, and unsaturable (up to 2M KCl) [Sukharev99]. MscL has also been shown to transfer cytoplasmic proteins including thioredoxin, elongation factor Tu (EF-Tu), and DnaK to the periplasm upon shock [Ajouz98, Berrier00]. MscL will allow passage of macromolecules smaller than 6.5 kDa [vandenBogaart07]. Gating of MscL is induced by lateral tension within the lipid bilayer [Sukharev93, Moe05, Tang06]. It has also been suggested that gating is pH-dependent due to its effect on the charged cluster of amino acids within the cytoplasmic helices of MscL leading to the alteration of the conformation of those domains [Kloda06]. Reconstituted functional MscL activity has been studied on a tethered membrane surface. Results support evidence for MscL being gated in response to stress in the lipid membrane as opposed to pressure build-up and also present the possibility of using MscL as a release valve for engineered membrane devices [Andersson08]. Concentration of anionic lipids within the membrane effect the rate of transition between the fully open and subconductance states as well as the rate and amplitute of release of calcein [Powl08]. Mutational studies indicate that the action of MscL is directly modulated by interaction with the surrounding, annular phospholipids that contact the protein in the membrane [Wiriyathanawudh09, Powl08a]. Hierarchical simulation framework studies [Chen08a, Tang08] and disulfide-trapping pattern and functional studies [Iscla08] have been done to study regions involved in the gating mechanism of MscL. Structural aspects of the lipid bilayer have been suggested to be important for MscL gating [Turner04].

Parabens have been shown to induce MscL activity which allows leakage of cell contents, suggesting a mechanism for their capability as antimicrobial agents [Nguyen05].

MscL, located within the cytoplasmic membrane, is a homopentamer. Each monomer consits of two transmembrane segments connected by an external loop that acts as a spring to resist opening of the channel and a cytoplasmic helix connected to the second TM segment through a linker [Blount96, Folgering05]. The cytoplasmic helices form a stable bundle creating a basket-like sieve with pores between the linkers on the cytoplasmic face of the channel so that only small molecules can pass through [Anishkin03]. MscL is also capable of forming complexes composed of two, three, four, or six subunits [Blount96], though extensive cross-linking studies show the pentameric form is the primary functional complex [Sukharev99a]. Two-dimensional crystallization of MscL resulted in a projection map of the hexameric protein at a resolution of 15 Å [Saint98]. The structure of the pentameric MscL homolog from Mycobacterium tuberculosis has been determined by X-ray crystallography to a resolution of 3.5 Å [Chang98]. Experiments using FRET spectroscopy show the diameter of MscL increases by 16 Å upon activation [Corry05]. The extramembranous domains of the channel, including the external loop, have been shown by patch clamp experiments to resist its opening and facilitate its closure when it is open [Ajouz00]. Extensive analyses of the gating mechanism of MscL have been performed [Blount96a, Hase97, Gu98, Ou98, Sukharev99b, Yoshimura99, Sukharev01, Gullingsrud01, Yoshimura01, Sukharev01a, Bilston02, Batiza02, Kong02, Perozo02, Betanzos02, Perozo02a, Maurer03, Valadie03, Gullingsrud03, Wiggins04, Yoshimura04, Chiang04, Levin04, Bartlett04, Shapovalov04, Anishkin05, Khan05, Chiang05, Bartlett06, Iscla07, Birkner12, Gracia13].

mscL expression is induced by RpoS upon entry into stationary phase and by osmotic stress [Stokes03]. MscL is targeted to the membrane by the SRP pathway and is inserted independently of the Sec translocon [Facey07, Pop09]. Conflicting results on the requirement of YidC for membrane insertion of MscL monomers have been reported [Facey07, Pop09] and a role for YidC in the assembly of homopentameric MscL has been suggested [Pop09].

Disruption of mscL resulted in loss of MscL activity which is restored by expression of the gene on a plasmid [Sukharev94]. mscS mscL double deletion mutants exhibit a lysis phenotype upon osmotic downshock [Levina99]. Reconstituted MscL is fully functional after purification of MscL-GST fusions and removal of GST or of histidine tagged MscL [Hase95, Blount96].

Seven mechanosensitive ion channels have been characterised in E. coli K-12: MscL, MscS, MscK, MscM, YnaI, YbiO and YbdG. A strain lacking all seven of these proteins fails to demonstrate channel opening in patch clamp experiments [Edwards12].

mscL: mechanosensitive channel of large conductance

Reviews: [Sukharev96, Blount96b, Sukharev97, Blount99, Wood99, Booth99, Sukharev99, Batiza99, Spencer99, Blount99a, Oakley99, Rees00, Ajouz00, Kubalski00, Martinac01, Biggin01, Blount97, Koprowski01, Roux02, Spencer02, Pivetti03, Strop03, Perozo03, Clarkson04, Edwards04, Sukharev04, Kung04]

Citations: [Hase97a, Cruickshank97, Blount97a, Arkin98, Moe98, Nakamaru99, Sukharev00, Maurer00, Konforti01, Perozo01, Maurer01, Okada02, Kloda02, Sachs02, Bezanilla02, Powl03, Park04, Clayton04, Buurman04, Li04, Folgering04, Wiggins05, Tsai05, Norman05, Hughes05, Ewis05, Meyer06, Ursell07, Carney07, Li09a]

Gene Citations: [Kawano05]

Locations: inner membrane

Map Position: [3,436,046 -> 3,436,456] (74.06 centisomes)
Length: 411 bp / 136 aa

Molecular Weight of Polypeptide: 14.957 kD (from nucleotide sequence), 17.0 kD (experimental) [Blount96 ]

Unification Links: ASAP:ABE-0010790 , CGSC:35417 , DIP:DIP-29827N , EchoBASE:EB1167 , EcoGene:EG11180 , EcoliWiki:b3291 , Mint:MINT-8071627 , ModBase:P0A742 , OU-Microarray:b3291 , PortEco:mscL , PR:PRO_000023296 , Pride:P0A742 , Protein Model Portal:P0A742 , RefSeq:NP_417749 , RegulonDB:EG11180 , SMR:P0A742 , String:511145.b3291 , UniProt:P0A742

Relationship Links: InterPro:IN-FAMILY:IPR001185 , InterPro:IN-FAMILY:IPR019823 , PDB:Structure:1KYK , PDB:Structure:1KYL , PDB:Structure:1KYM , Pfam:IN-FAMILY:PF01741 , Prints:IN-FAMILY:PR01264 , Prosite:IN-FAMILY:PS01327

Gene-Reaction Schematic: ?

GO Terms:

Biological Process: GO:0006811 - ion transport Inferred from experiment Inferred by computational analysis [UniProtGOA11, Sukharev94]
GO:0009992 - cellular water homeostasis Inferred from experiment [Levina99]
GO:0034220 - ion transmembrane transport Inferred by computational analysis Inferred from experiment [Sukharev94, GOA01]
GO:0006810 - transport Inferred by computational analysis [UniProtGOA11]
GO:0055085 - transmembrane transport Inferred by computational analysis [GOA06]
Molecular Function: GO:0008381 - mechanically-gated ion channel activity Inferred from experiment [Sukharev94]
GO:0042802 - identical protein binding Inferred from experiment [Price11, Lasserre06]
GO:0005216 - ion channel activity Inferred by computational analysis [GOA01]
GO:0015267 - channel activity Inferred by computational analysis [GOA06]
Cellular Component: GO:0005886 - plasma membrane Inferred from experiment Inferred by computational analysis [UniProtGOA11a, UniProtGOA11, GOA06, DiazMejia09, Zhang07, Daley05, Sukharev94]
GO:0016020 - membrane Inferred from experiment Inferred by computational analysis [UniProtGOA11, Lasserre06]
GO:0016021 - integral component of membrane Inferred from experiment Inferred by computational analysis [UniProtGOA11, GOA01, Blount96]

MultiFun Terms: cell processes adaptations other (mechanical, nutritional, oxidative stress)
cell structure membrane
transport Channel-type Transporters alpha-type channels

Essentiality data for mscL knockouts: ?

Growth Medium Growth? T (°C) O2 pH Osm/L Growth Observations
LB Lennox Yes 37 Aerobic 7   Yes [Baba06, Comment 1]
M9 medium with 1% glycerol Yes 37 Aerobic 7.2 0.35 Yes [Joyce06, Comment 2]
MOPS medium with 0.4% glucose Yes 37 Aerobic 7.2 0.22 Yes [Baba06, Comment 1]

Credits:
Created 03-Oct-2007 by Johnson A , JCVI
Last-Curated ? 03-Oct-2007 by Johnson A , JCVI


Enzymatic reaction of: mechanosensitive channel MscL


Sequence Features

Feature Class Location Common Name Citations
Transmembrane-Region 19 -> 38 M1
[Blount96]
Transmembrane-Region 70 -> 96 M2
[Blount96]


Gene Local Context (not to scale): ?

Transcription Unit:

Notes:

History:
10/20/97 Gene b3291 from Blattner lab Genbank (v. M52) entry merged into EcoCyc gene EG11180; confirmed by SwissProt match.


References

Ajouz00: Ajouz B, Berrier C, Besnard M, Martinac B, Ghazi A (2000). "Contributions of the different extramembranous domains of the mechanosensitive ion channel MscL to its response to membrane tension." J Biol Chem 2000;275(2);1015-22. PMID: 10625640

Ajouz98: Ajouz B, Berrier C, Garrigues A, Besnard M, Ghazi A (1998). "Release of thioredoxin via the mechanosensitive channel MscL during osmotic downshock of Escherichia coli cells." J Biol Chem 273(41);26670-4. PMID: 9756908

Andersson08: Andersson M, Okeyo G, Wilson D, Keizer H, Moe P, Blount P, Fine D, Dodabalapur A, Duran RS (2008). "Voltage-induced gating of the mechanosensitive MscL ion channel reconstituted in a tethered lipid bilayer membrane." Biosens Bioelectron 23(6);919-23. PMID: 17996439

Anishkin03: Anishkin A, Gendel V, Sharifi NA, Chiang CS, Shirinian L, Guy HR, Sukharev S (2003). "On the conformation of the COOH-terminal domain of the large mechanosensitive channel MscL." J Gen Physiol 121(3);227-44. PMID: 12601086

Anishkin05: Anishkin A, Chiang CS, Sukharev S (2005). "Gain-of-function mutations reveal expanded intermediate states and a sequential action of two gates in MscL." J Gen Physiol 125(2);155-70. PMID: 15684093

Arkin98: Arkin IT, Sukharev SI, Blount P, Kung C, Brunger AT (1998). "Helicity, membrane incorporation, orientation and thermal stability of the large conductance mechanosensitive ion channel from E. coli." Biochim Biophys Acta 1369(1);131-40. PMID: 9528681

Baba06: Baba T, Ara T, Hasegawa M, Takai Y, Okumura Y, Baba M, Datsenko KA, Tomita M, Wanner BL, Mori H (2006). "Construction of Escherichia coli K-12 in-frame, single-gene knockout mutants: the Keio collection." Mol Syst Biol 2;2006.0008. PMID: 16738554

Bartlett04: Bartlett JL, Levin G, Blount P (2004). "An in vivo assay identifies changes in residue accessibility on mechanosensitive channel gating." Proc Natl Acad Sci U S A 101(27);10161-5. PMID: 15226501

Bartlett06: Bartlett JL, Li Y, Blount P (2006). "Mechanosensitive channel gating transitions resolved by functional changes upon pore modification." Biophys J 91(10);3684-91. PMID: 16935962

Batiza02: Batiza AF, Kuo MM, Yoshimura K, Kung C (2002). "Gating the bacterial mechanosensitive channel MscL invivo." Proc Natl Acad Sci U S A 99(8);5643-8. PMID: 11960017

Batiza99: Batiza AF, Rayment I, Kung C (1999). "Channel gate! Tension, leak and disclosure." Structure 7(5);R99-103. PMID: 10378264

Berrier00: Berrier C, Garrigues A, Richarme G, Ghazi A (2000). "Elongation factor Tu and DnaK are transferred from the cytoplasm to the periplasm of Escherichia coli during osmotic downshock presumably via the mechanosensitive channel mscL." J Bacteriol 2000;182(1);248-51. PMID: 10613892

Betanzos02: Betanzos M, Chiang CS, Guy HR, Sukharev S (2002). "A large iris-like expansion of a mechanosensitive channel protein induced by membrane tension." Nat Struct Biol 9(9);704-10. PMID: 12172538

Bezanilla02: Bezanilla F, Perozo E (2002). "Structural biology. Force and voltage sensors in one structure." Science 298(5598);1562-3. PMID: 12446894

Biggin01: Biggin PC, Sansom MS (2001). "Channel gating: twist to open." Curr Biol 11(9);R364-6. PMID: 11369249

Bilston02: Bilston LE, Mylvaganam K (2002). "Molecular simulations of the large conductance mechanosensitive (MscL) channel under mechanical loading." FEBS Lett 512(1-3);185-90. PMID: 11852077

Birkner12: Birkner JP, Poolman B, Kocer A (2012). "Hydrophobic gating of mechanosensitive channel of large conductance evidenced by single-subunit resolution." Proc Natl Acad Sci U S A 109(32);12944-9. PMID: 22826215

Blount96: Blount P, Sukharev SI, Moe PC, Schroeder MJ, Guy HR, Kung C (1996). "Membrane topology and multimeric structure of a mechanosensitive channel protein of Escherichia coli." EMBO J 15(18);4798-805. PMID: 8890153

Blount96a: Blount P, Sukharev SI, Schroeder MJ, Nagle SK, Kung C (1996). "Single residue substitutions that change the gating properties of a mechanosensitive channel in Escherichia coli." Proc Natl Acad Sci U S A 93(21);11652-7. PMID: 8876191

Blount96b: Blount P, Sukharev SI, Moe PC, Nagle SK, Kung C (1996). "Towards an understanding of the structural and functional properties of MscL, a mechanosensitive channel in bacteria." Biol Cell 87(1-2);1-8. PMID: 9004483

Blount97: Blount P, Sukharev S, Kung C (1997). "A mechanosensitive channel protein and its gene in E. coli." Gravit Space Biol Bull 10(2);43-7. PMID: 11540118

Blount97a: Blount P, Schroeder MJ, Kung C (1997). "Mutations in a bacterial mechanosensitive channel change the cellular response to osmotic stress." J Biol Chem 272(51);32150-7. PMID: 9405414

Blount99: Blount P, Sukharev SI, Moe PC, Martinac B, Kung C (1999). "Mechanosensitive channels of bacteria." Methods Enzymol 294;458-82. PMID: 9916243

Blount99a: Blount P, Moe PC (1999). "Bacterial mechanosensitive channels: integrating physiology, structure and function." Trends Microbiol 7(10);420-4. PMID: 10498951

Booth99: Booth IR, Louis P (1999). "Managing hypoosmotic stress: aquaporins and mechanosensitive channels in Escherichia coli." Curr Opin Microbiol 2(2);166-9. PMID: 10322175

Buurman04: Buurman ET, McLaggan D, Naprstek J, Epstein W (2004). "Multiple paths for nonphysiological transport of K+ in Escherichia coli." J Bacteriol 186(13);4238-45. PMID: 15205426

Carney07: Carney J, East JM, Lee AG (2007). "Penetration of lipid chains into transmembrane surfaces of membrane proteins: studies with MscL." Biophys J 92(10);3556-63. PMID: 17307828

Chang98: Chang G, Spencer RH, Lee AT, Barclay MT, Rees DC (1998). "Structure of the MscL homolog from Mycobacterium tuberculosis: a gated mechanosensitive ion channel." Science 282(5397);2220-6. PMID: 9856938

Chen08a: Chen X, Cui Q, Tang Y, Yoo J, Yethiraj A (2008). "Gating mechanisms of mechanosensitive channels of large conductance, I: a continuum mechanics-based hierarchical framework." Biophys J 95(2);563-80. PMID: 18390626

Chiang04: Chiang CS, Anishkin A, Sukharev S (2004). "Gating of the large mechanosensitive channel in situ: estimation of the spatial scale of the transition from channel population responses." Biophys J 86(5);2846-61. PMID: 15111402

Chiang05: Chiang CS, Shirinian L, Sukharev S (2005). "Capping transmembrane helices of MscL with aromatic residues changes channel response to membrane stretch." Biochemistry 44(37);12589-97. PMID: 16156671

Clarkson04: Clarkson S (2004). "Regulating osmosensing." Nat Rev Microbiol 2(2);86-7. PMID: 15040250

Clayton04: Clayton D, Shapovalov G, Maurer JA, Dougherty DA, Lester HA, Kochendoerfer GG (2004). "Total chemical synthesis and electrophysiological characterization of mechanosensitive channels from Escherichia coli and Mycobacterium tuberculosis." Proc Natl Acad Sci U S A 101(14);4764-9. PMID: 15041744

Corry05: Corry B, Rigby P, Liu ZW, Martinac B (2005). "Conformational changes involved in MscL channel gating measured using FRET spectroscopy." Biophys J 89(6);L49-51. PMID: 16199508

Cruickshank97: Cruickshank CC, Minchin RF, Le Dain AC, Martinac B (1997). "Estimation of the pore size of the large-conductance mechanosensitive ion channel of Escherichia coli." Biophys J 73(4);1925-31. PMID: 9336188

Daley05: Daley DO, Rapp M, Granseth E, Melen K, Drew D, von Heijne G (2005). "Global topology analysis of the Escherichia coli inner membrane proteome." Science 308(5726);1321-3. PMID: 15919996

DiazMejia09: Diaz-Mejia JJ, Babu M, Emili A (2009). "Computational and experimental approaches to chart the Escherichia coli cell-envelope-associated proteome and interactome." FEMS Microbiol Rev 33(1);66-97. PMID: 19054114

Edwards04: Edwards MD, Booth IR, Miller S (2004). "Gating the bacterial mechanosensitive channels: MscS a new paradigm?." Curr Opin Microbiol 7(2);163-7. PMID: 15063854

Edwards12: Edwards MD, Black S, Rasmussen T, Rasmussen A, Stokes NR, Stephen TL, Miller S, Booth IR (2012). "Characterization of three novel mechanosensitive channel activities in Escherichia coli." Channels (Austin) 6(4). PMID: 22874652

Ewis05: Ewis HE, Lu CD (2005). "Osmotic shock: a mechanosensitive channel blocker can prevent release of cytoplasmic but not periplasmic proteins." FEMS Microbiol Lett 253(2);295-301. PMID: 16288836

Facey07: Facey SJ, Neugebauer SA, Krauss S, Kuhn A (2007). "The mechanosensitive channel protein MscL is targeted by the SRP to the novel YidC membrane insertion pathway of Escherichia coli." J Mol Biol 365(4);995-1004. PMID: 17113597

Folgering04: Folgering JH, Kuiper JM, de Vries AH, Engberts JB, Poolman B (2004). "Lipid-mediated light activation of a mechanosensitive channel of large conductance." Langmuir 20(17);6985-7. PMID: 15301476

Folgering05: Folgering JH, Wolters JC, Poolman B (2005). "Engineering covalent oligomers of the mechanosensitive channel of large conductance from Escherichia coli with native conductance and gating characteristics." Protein Sci 14(12);2947-54. PMID: 16322576

GOA01: GOA, DDB, FB, MGI, ZFIN (2001). "Gene Ontology annotation through association of InterPro records with GO terms."

GOA06: GOA, SIB (2006). "Electronic Gene Ontology annotations created by transferring manual GO annotations between orthologous microbial proteins."

Gracia13: Gracia E, Farre D, Cortes A, Ferrer-Costa C, Orozco M, Mallol J, Lluis C, Canela EI, McCormick PJ, Franco R, Fanelli F, Casado V (2013). "The catalytic site structural gate of adenosine deaminase allosterically modulates ligand binding to adenosine receptors." FASEB J 27(3);1048-61. PMID: 23193172

Gu98: Gu L, Liu W, Martinac B (1998). "Electromechanical coupling model of gating the large mechanosensitive ion channel (MscL) of Escherichia coli by mechanical force." Biophys J 74(6);2889-902. PMID: 9635742

Gullingsrud01: Gullingsrud J, Kosztin D, Schulten K (2001). "Structural determinants of MscL gating studied by molecular dynamics simulations." Biophys J 80(5);2074-81. PMID: 11325711

Gullingsrud03: Gullingsrud J, Schulten K (2003). "Gating of MscL studied by steered molecular dynamics." Biophys J 85(4);2087-99. PMID: 14507677

Hase95: Hase CC, Le Dain AC, Martinac B (1995). "Purification and functional reconstitution of the recombinant large mechanosensitive ion channel (MscL) of Escherichia coli." J Biol Chem 270(31);18329-34. PMID: 7543101

Hase97: Hase CC, Le Dain AC, Martinac B (1997). "Molecular dissection of the large mechanosensitive ion channel (MscL) of E. coli: mutants with altered channel gating and pressure sensitivity." J Membr Biol 157(1);17-25. PMID: 9141355

Hase97a: Hase CC, Minchin RF, Kloda A, Martinac B (1997). "Cross-linking studies and membrane localization and assembly of radiolabelled large mechanosensitive ion channel (MscL) of Escherichia coli." Biochem Biophys Res Commun 232(3);777-82. PMID: 9126353

Hughes05: Hughes S, El Haj AJ, Dobson J, Martinac B (2005). "The influence of static magnetic fields on mechanosensitive ion channel activity in artificial liposomes." Eur Biophys J 34(5);461-8. PMID: 15959773

Iscla07: Iscla I, Levin G, Wray R, Blount P (2007). "Disulfide trapping the mechanosensitive channel MscL into a gating-transition state." Biophys J 92(4);1224-32. PMID: 17114217

Iscla08: Iscla I, Wray R, Blount P (2008). "On the structure of the N-terminal domain of the MscL channel: helical bundle or membrane interface." Biophys J 95(5);2283-91. PMID: 18515388

Joyce06: Joyce AR, Reed JL, White A, Edwards R, Osterman A, Baba T, Mori H, Lesely SA, Palsson BO, Agarwalla S (2006). "Experimental and computational assessment of conditionally essential genes in Escherichia coli." J Bacteriol 188(23);8259-71. PMID: 17012394

Kawano05: Kawano M, Reynolds AA, Miranda-Rios J, Storz G (2005). "Detection of 5'- and 3'-UTR-derived small RNAs and cis-encoded antisense RNAs in Escherichia coli." Nucleic Acids Res 33(3);1040-50. PMID: 15718303

Khan05: Khan RN, Martinac B, Madsen BW, Milne RK, Yeo GF, Edeson RO (2005). "Hidden Markov analysis of mechanosensitive ion channel gating." Math Biosci 193(2);139-58. PMID: 15748727

Kloda02: Kloda A, Martinac B (2002). "Mechanosensitive channels of bacteria and archaea share a common ancestral origin." Eur Biophys J 31(1);14-25. PMID: 12046893

Kloda06: Kloda A, Ghazi A, Martinac B (2006). "C-terminal charged cluster of MscL, RKKEE, functions as a pH sensor." Biophys J 90(6);1992-8. PMID: 16387769

Konforti01: Konforti B (2001). "Picture story. Leak proof channel." Nat Struct Biol 8(3);202. PMID: 11224560

Kong02: Kong Y, Shen Y, Warth TE, Ma J (2002). "Conformational pathways in the gating of Escherichia coli mechanosensitive channel." Proc Natl Acad Sci U S A 99(9);5999-6004. PMID: 11972047

Koprowski01: Koprowski P, Kubalski A (2001). "Bacterial ion channels and their eukaryotic homologues." Bioessays 23(12);1148-58. PMID: 11746234

Kubalski00: Kubalski A, Koprowski P (2000). "[Bacterial ion canals]." Postepy Hig Med Dosw 54(3);317-27. PMID: 10941266

Kung04: Kung C, Blount P (2004). "Channels in microbes: so many holes to fill." Mol Microbiol 53(2);373-80. PMID: 15228520

Lasserre06: Lasserre JP, Beyne E, Pyndiah S, Lapaillerie D, Claverol S, Bonneu M (2006). "A complexomic study of Escherichia coli using two-dimensional blue native/SDS polyacrylamide gel electrophoresis." Electrophoresis 27(16);3306-21. PMID: 16858726

Levin04: Levin G, Blount P (2004). "Cysteine scanning of MscL transmembrane domains reveals residues critical for mechanosensitive channel gating." Biophys J 86(5);2862-70. PMID: 15111403

Levina99: Levina N, Totemeyer S, Stokes NR, Louis P, Jones MA, Booth IR (1999). "Protection of Escherichia coli cells against extreme turgor by activation of MscS and MscL mechanosensitive channels: identification of genes required for MscS activity." EMBO J 1999;18(7);1730-7. PMID: 10202137

Li04: Li Y, Wray R, Blount P (2004). "Intragenic suppression of gain-of-function mutations in the Escherichia coli mechanosensitive channel, MscL." Mol Microbiol 53(2);485-95. PMID: 15228529

Li09a: Li Y, Wray R, Eaton C, Blount P (2009). "An open-pore structure of the mechanosensitive channel MscL derived by determining transmembrane domain interactions upon gating." FASEB J 23(7);2197-204. PMID: 19261722

Martinac01: Martinac B (2001). "Mechanosensitive channels in prokaryotes." Cell Physiol Biochem 11(2);61-76. PMID: 11275684

Maurer00: Maurer JA, Elmore DE, Lester HA, Dougherty DA (2000). "Comparing and contrasting Escherichia coli and Mycobacterium tuberculosis mechanosensitive channels (MscL). New gain of function mutations in the loop region." J Biol Chem 275(29);22238-44. PMID: 10801868

Maurer01: Maurer JA, Dougherty DA (2001). "A high-throughput screen for MscL channel activity and mutational phenotyping." Biochim Biophys Acta 1514(2);165-9. PMID: 11557017

Maurer03: Maurer JA, Dougherty DA (2003). "Generation and evaluation of a large mutational library from the Escherichia coli mechanosensitive channel of large conductance, MscL: implications for channel gating and evolutionary design." J Biol Chem 278(23);21076-82. PMID: 12670944

Meyer06: Meyer GR, Gullingsrud J, Schulten K, Martinac B (2006). "Molecular dynamics study of MscL interactions with a curved lipid bilayer." Biophys J 91(5);1630-7. PMID: 16751236

Moe05: Moe P, Blount P (2005). "Assessment of potential stimuli for mechano-dependent gating of MscL: effects of pressure, tension, and lipid headgroups." Biochemistry 44(36);12239-44. PMID: 16142922

Moe98: Moe PC, Blount P, Kung C (1998). "Functional and structural conservation in the mechanosensitive channel MscL implicates elements crucial for mechanosensation." Mol Microbiol 28(3);583-92. PMID: 9632260

Nakamaru99: Nakamaru Y, Takahashi Y, Unemoto T, Nakamura T (1999). "Mechanosensitive channel functions to alleviate the cell lysis of marine bacterium, Vibrio alginolyticus, by osmotic downshock." FEBS Lett 444(2-3);170-2. PMID: 10050752

Nguyen05: Nguyen T, Clare B, Guo W, Martinac B (2005). "The effects of parabens on the mechanosensitive channels of E. coli." Eur Biophys J 34(5);389-95. PMID: 15770478

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Other References Related to Gene Regulation

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