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Escherichia coli K-12 substr. MG1655 Transporter: osmolyte:H+ symporter ProP

Gene: proP Accession Numbers: EG11612 (EcoCyc), b4111, ECK4104

Synonyms: ProP proline/betaine MFS transporter, PPII, proline permease II, ProP osmosensory MFS transporter

Regulation Summary Diagram: ?

Regulation summary diagram for proP

Subunit composition of osmolyte:H+ symporter ProP = [ProP]2

ProP is an osmoprotectant/proton symporter capable of transporting proline and glycine betaine, and mediates the uptake of osmoprotectants to adapt to increases in osmotic pressure. ProP is a member of the major facilitator superfamily (MFS) of transporters [Griffith92] and transports its zwitterionic substrates with protons through a symport mechanism [MacMillan99]. ProP has also been suggested to transport the osmoprotectants taurine [McLaggan91], ectoine [Jebbar92], carnitine [Verheul98], 3,4-dehydroproline [Stalmach83], azetidine-2-carboxylate [Csonka81], 5-hydroxy-L-pipecolic acid [Gowrishankar86], pipecolate, proline betaine, N,N-dimethylglycine, 1-carboxymethylpyridinium [MacMillan99], dimethylsulfoniopropionate, and dimethylsulfonioacetate [Cosquer99]. Choline, alanine, trigonelline, and sarcosine are all inhibitors of proline import, but are not transport substrates [MacMillan99].

Studies in whole cells and membrane vesicles have shown that ProP-mediated proline transport is inhibited by protonophores [Milner88] and ProP binds proline with a low affinity (Km = 0.3 μM) [Anderson80]. ProP activity is dependent upon exogenous K+ and stimulates K+ release from the cell through an unknown mechanism [MacMillan99] although other results suggest that ProP activity is not specifically K+-dependent with K+ being required for respiration rather than for direct involvement in transport mediated by ProP [Culham08]. ProP has been overproduced, purified and reconstituted in liposomes as an active transporter. ProP-mediated transport in liposomes required both the membrane potential and an osmotic upshift, suggesting it functions as both a transporter and an osmosensor [Milner88, Racher99]. Studies using purified ProP-His6 proteoliposomes suggest that ProP senses both membrane potential (ΔΨ) and osmotic pressure, switching among "off", "on", and "osmolality-sensitive" states due to changes in membrane potential [Culham08]. Conformational change of ProP (as measured by the accessibility of cysteine residues to alkylation) has been observed in response to osmolality [Culham08a].

The three dimensional structure of ProP, modeled after GlpT, has been substantiated with LacZ-PhoA fusions [Wood05]. ProP is a dimer in the membrane [Zoetewey03]. Characterisation of purified, reconstituted ProP using infrared spectroscopy shows that it is a highly α helical membrane protein [Sayeed09]. The N-terminal six-helix bundle is predicted to contain a proton pathway and to be involved in osmotic activation [Wood05]. The C-terminal coiled-coil and the transmembrane segment XII of ProP are also involved in osmosensing by influencing the osmolality required to activate ProP [Culham, Hillar05, Tsatskis05, Liu07c]. Polar and septal localization of ProP is dependent upon cardiolipin localization [Romantsov07].

Expression of proP is increased by osmotic upshock, amino acid limitation, and stationary phase [Mellies95]. Expression is activated by the activities of Fis, CRP-cAMP, and EσS when they are all preset in the cell during late exponential and early stationary phases by binding at promoter P2 which responds to stationary phase signals [Xu95c, Xu97b, McLeod99, Landis99, McLeod00, Cheng00, Typas07]. Binding of CRP-cAMP at P1, which responds to osmotic signals, prevents expression from this promoter [Mellies95, Xu97c]. Over-expression of the EvgA/EvgS two component system increased expression of proP through the PhoP/PhoQ system, but over-expression of PhoP/PhoQ could not accomplish this alone [Eguchi04]. CspC and CspE have also been shown to be involved in increasing expression of proP through EσS [Phadtare01].

An uncharacterized interaction with ProQ is important for activation of ProP [Milner89, Kunte99, Smith04a].

Expression of proP from a multicopy plasmid allowed growth of K+ transport deficient mutants to grow at lower K+ concentrations [Buurman04].

Reviews: [Mileykovskaya07, Wood06, Wood01, Wood99a]

Citations: [Hillar03, Culham03, Culham03a, McLeod02, Aiyar02, Racher01, Randall95a, Gauthier90, Dinnbier88, May86, Wood80]

Locations: inner membrane

Map Position: [4,328,525 -> 4,330,027] (93.29 centisomes, 336°)
Length: 1503 bp / 500 aa

Molecular Weight of Polypeptide: 54.846 kD (from nucleotide sequence), 42.0 kD (experimental) [Culham93 ]

Unification Links: ASAP:ABE-0013462 , CGSC:361 , EchoBASE:EB1569 , EcoGene:EG11612 , EcoliWiki:b4111 , ModBase:P0C0L7 , OU-Microarray:b4111 , PortEco:proP , PR:PRO_000023599 , Protein Model Portal:P0C0L7 , RefSeq:NP_418535 , RegulonDB:EG11612 , SMR:P0C0L7 , String:511145.b4111 , UniProt:P0C0L7

Relationship Links: InterPro:IN-FAMILY:IPR004736 , InterPro:IN-FAMILY:IPR005828 , InterPro:IN-FAMILY:IPR005829 , InterPro:IN-FAMILY:IPR015041 , InterPro:IN-FAMILY:IPR020846 , PDB:Structure:1r48 , PDB:Structure:1Y8S , Pfam:IN-FAMILY:PF00083 , Pfam:IN-FAMILY:PF08946 , Prosite:IN-FAMILY:PS00216 , Prosite:IN-FAMILY:PS50850

In Paralogous Gene Group: 17 (37 members)

Gene-Reaction Schematic: ?

Gene-Reaction Schematic

Genetic Regulation Schematic: ?

Genetic regulation schematic for proP

GO Terms:

Biological Process: GO:0006865 - amino acid transport Inferred from experiment [Racher99]
GO:0015824 - proline transport Inferred from experiment [Gowrishankar86]
GO:0031460 - glycine betaine transport Inferred from experiment [Milner88]
GO:0042538 - hyperosmotic salinity response Inferred from experiment [Gowrishankar86]
GO:0006810 - transport Inferred by computational analysis [UniProtGOA11, GOA01]
GO:0055085 - transmembrane transport Inferred by computational analysis [GOA01]
Molecular Function: GO:0005297 - hydrogen:proline symporter activity Inferred from experiment [Milner88]
GO:0015653 - glycine betaine:proton symporter activity Inferred from experiment [Milner88]
GO:0005215 - transporter activity Inferred by computational analysis [GOA01]
GO:0015293 - symporter activity Inferred by computational analysis [UniProtGOA11]
GO:0022857 - transmembrane transporter activity Inferred by computational analysis [GOA01]
Cellular Component: GO:0005886 - plasma membrane Inferred from experiment Inferred by computational analysis [UniProtGOA11a, UniProtGOA11, DiazMejia09, Zhang07, Daley05, Milner88, Racher99]
GO:0005887 - integral component of plasma membrane Inferred from experiment Inferred by computational analysis [Culham93, Racher99]
GO:0016021 - integral component of membrane Inferred from experiment Inferred by computational analysis [UniProtGOA11, GOA01, Racher99]
GO:0016020 - membrane Inferred by computational analysis [UniProtGOA11]

MultiFun Terms: cell processes adaptations osmotic pressure
cell structure membrane
metabolism biosynthesis of building blocks amino acids proline
transport Electrochemical potential driven transporters Porters (Uni-, Sym- and Antiporters)

Essentiality data for proP knockouts: ?

Growth Medium Growth? T (°C) O2 pH Osm/L Growth Observations
LB enriched Yes 37 Aerobic 6.95   Yes [Gerdes03, Comment 1]
LB Lennox Yes 37 Aerobic 7   Yes [Baba06, Comment 2]
M9 medium with 1% glycerol Yes 37 Aerobic 7.2 0.35 Yes [Joyce06, Comment 3]
MOPS medium with 0.4% glucose Yes 37 Aerobic 7.2 0.22 Yes [Baba06, Comment 2]
Yes [Feist07, Comment 4]

Created 31-Oct-2007 by Johnson A , JCVI

Enzymatic reaction of: glycine betaine:H+ symporter (osmolyte:H+ symporter ProP)

Transport reaction diagram for glycine betaine:H+ symporter

Enzymatic reaction of: L-proline:H+ symporter (osmolyte:H+ symporter ProP)

Transport reaction diagram for L-proline:H+ symporter

Alternative Products for L-proline: dimethylsulfonioacetate [Cosquer99 ] , taurine [McLaggan91 ] , K+ [Buurman04 ] , dimethylsulfoniopropanoate [Cosquer99 ] , ectoine [Jebbar92 ] , 1-carboxymethylpyridinium [MacMillan99 ] , L-pipecolate [Gouesbet94 ] , 3,4-dehydroproline [Stalmach83 ] , azetidine-2-carboxylate [Csonka81 ] , dimethylglycine [MacMillan99 ] , L-proline betaine [Haardt95 ] , D-carnitine [Verheul98 ] , L-carnitine [Verheul98 ] , 5-hydroxy-pipecolate [Gowrishankar86 ]

Inhibitors (Unknown Mechanism): choline [MacMillan99] , L-alanine [MacMillan99] , methylnicotinate [MacMillan99] , sarcosine [MacMillan99]

Sequence Features

Protein sequence of ProP with features indicated

Feature Class Location Citations Comment
Transmembrane-Region 38 -> 58
UniProt: Helical; Name=1;; Non-Experimental Qualifier: potential;
Transmembrane-Region 66 -> 86
UniProt: Helical; Name=2;; Non-Experimental Qualifier: potential;
Transmembrane-Region 98 -> 118
UniProt: Helical; Name=3;; Non-Experimental Qualifier: potential;
Transmembrane-Region 122 -> 142
UniProt: Helical; Name=4;; Non-Experimental Qualifier: potential;
Transmembrane-Region 170 -> 190
UniProt: Helical; Name=5;; Non-Experimental Qualifier: potential;
Transmembrane-Region 195 -> 215
UniProt: Helical; Name=6;; Non-Experimental Qualifier: potential;
Transmembrane-Region 261 -> 281
UniProt: Helical; Name=7;; Non-Experimental Qualifier: potential;
Transmembrane-Region 298 -> 318
UniProt: Helical; Name=8;; Non-Experimental Qualifier: potential;
Transmembrane-Region 326 -> 346
UniProt: Helical; Name=9;; Non-Experimental Qualifier: potential;
Transmembrane-Region 351 -> 371
UniProt: Helical; Name=10;; Non-Experimental Qualifier: potential;
Transmembrane-Region 391 -> 411
UniProt: Helical; Name=11;; Non-Experimental Qualifier: potential;
Transmembrane-Region 417 -> 437
UniProt: Helical; Name=12;; Non-Experimental Qualifier: potential;

Gene Local Context (not to scale): ?

Gene local context diagram

Transcription Units:

Transcription-unit diagram

Transcription-unit diagram


10/20/97 Gene b4111 from Blattner lab Genbank (v. M52) entry merged into EcoCyc gene EG11612; confirmed by SwissProt match.


Aiyar02: Aiyar SE, McLeod SM, Ross W, Hirvonen CA, Thomas MS, Johnson RC, Gourse RL (2002). "Architecture of Fis-activated transcription complexes at the Escherichia coli rrnB P1 and rrnE P1 promoters." J Mol Biol 316(3);501-16. PMID: 11866514

Anderson80: Anderson RR, Menzel R, Wood JM (1980). "Biochemistry and regulation of a second L-proline transport system in Salmonella typhimurium." J Bacteriol 1980;141(3);1071-6. PMID: 6988401

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

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

Cheng00: Cheng YS, Yang WZ, Johnson RC, Yuan HS (2000). "Structural analysis of the transcriptional activation on Fis: crystal structures of six Fis mutants with different activation properties." J Mol Biol 302(5);1139-51. PMID: 11183780

Cosquer99: Cosquer A, Pichereau V, Pocard JA, Minet J, Cormier M, Bernard T (1999). "Nanomolar levels of dimethylsulfoniopropionate, dimethylsulfonioacetate, and glycine betaine are sufficient to confer osmoprotection to Escherichia coli." Appl Environ Microbiol 65(8);3304-11. PMID: 10427011

Csonka81: Csonka LN (1981). "The role of proline in osmoregulation in Salmonella typhimurium and Escherichia coli." Basic Life Sci 18;533-42. PMID: 7023463

Culham: Culham DE, Tripet B, Racher KI, Voegele RT, Hodges RS, Wood JM "The role of the carboxyl terminal alpha-helical coiled-coil domain in osmosensing by transporter ProP of Escherichia coli." J Mol Recognit 13(5);309-22. PMID: 10992293

Culham03: Culham DE, Hillar A, Henderson J, Ly A, Vernikovska YI, Racher KI, Boggs JM, Wood JM (2003). "Creation of a fully functional cysteine-less variant of osmosensor and proton-osmoprotectant symporter ProP from Escherichia coli and its application to assess the transporter's membrane orientation." Biochemistry 42(40);11815-23. PMID: 14529293

Culham03a: Culham DE, Henderson J, Crane RA, Wood JM (2003). "Osmosensor ProP of Escherichia coli responds to the concentration, chemistry, and molecular size of osmolytes in the proteoliposome lumen." Biochemistry 42(2);410-20. PMID: 12525168

Culham08: Culham DE, Romantsov T, Wood JM (2008). "Roles of K+, H+, H2O, and DeltaPsi in solute transport mediated by major facilitator superfamily members ProP and LacY." Biochemistry 47(31);8176-85. PMID: 18620422

Culham08a: Culham DE, Vernikovska Y, Tschowri N, Keates RA, Wood JM, Boggs JM (2008). "Periplasmic loops of osmosensory transporter ProP in Escherichia coli are sensitive to osmolality." Biochemistry 47(51);13584-93. PMID: 19049385

Culham93: Culham DE, Lasby B, Marangoni AG, Milner JL, Steer BA, van Nues RW, Wood JM (1993). "Isolation and sequencing of Escherichia coli gene proP reveals unusual structural features of the osmoregulatory proline/betaine transporter, ProP." J Mol Biol 229(1);268-76. PMID: 8421314

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

Dinnbier88: Dinnbier U, Limpinsel E, Schmid R, Bakker EP (1988). "Transient accumulation of potassium glutamate and its replacement by trehalose during adaptation of growing cells of Escherichia coli K-12 to elevated sodium chloride concentrations." Arch Microbiol 150(4);348-57. PMID: 3060036

Eguchi04: Eguchi Y, Okada T, Minagawa S, Oshima T, Mori H, Yamamoto K, Ishihama A, Utsumi R (2004). "Signal transduction cascade between EvgA/EvgS and PhoP/PhoQ two-component systems of Escherichia coli." J Bacteriol 186(10);3006-14. PMID: 15126461

Feist07: Feist AM, Henry CS, Reed JL, Krummenacker M, Joyce AR, Karp PD, Broadbelt LJ, Hatzimanikatis V, Palsson BO (2007). "A genome-scale metabolic reconstruction for Escherichia coli K-12 MG1655 that accounts for 1260 ORFs and thermodynamic information." Mol Syst Biol 3;121. PMID: 17593909

Gauthier90: Gauthier MJ, Le Rudulier D (1990). "Survival in seawater of Escherichia coli cells grown in marine sediments containing glycine betaine." Appl Environ Microbiol 56(9);2915-8. PMID: 2275537

Gerdes03: Gerdes SY, Scholle MD, Campbell JW, Balazsi G, Ravasz E, Daugherty MD, Somera AL, Kyrpides NC, Anderson I, Gelfand MS, Bhattacharya A, Kapatral V, D'Souza M, Baev MV, Grechkin Y, Mseeh F, Fonstein MY, Overbeek R, Barabasi AL, Oltvai ZN, Osterman AL (2003). "Experimental determination and system level analysis of essential genes in Escherichia coli MG1655." J Bacteriol 185(19);5673-84. PMID: 13129938

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

Gouesbet94: Gouesbet G, Jebbar M, Talibart R, Bernard T, Blanco C (1994). "Pipecolic acid is an osmoprotectant for Escherichia coli taken up by the general osmoporters ProU and ProP." Microbiology 140 ( Pt 9);2415-22. PMID: 7952193

Gowrishankar86: Gowrishankar J (1986). "proP-mediated proline transport also plays a role in Escherichia coli osmoregulation." J Bacteriol 166(1);331-3. PMID: 3082857

Griffith92: Griffith JK, Baker ME, Rouch DA, Page MG, Skurray RA, Paulsen IT, Chater KF, Baldwin SA, Henderson PJ (1992). "Membrane transport proteins: implications of sequence comparisons." Curr Opin Cell Biol 1992;4(4);684-95. PMID: 1419050

Haardt95: Haardt M, Kempf B, Faatz E, Bremer E (1995). "The osmoprotectant proline betaine is a major substrate for the binding-protein-dependent transport system ProU of Escherichia coli K-12." Mol Gen Genet 246(6);783-6. PMID: 7898450

Hillar03: Hillar A, Tripet B, Zoetewey D, Wood JM, Hodges RS, Boggs JM (2003). "Detection of alpha-helical coiled-coil dimer formation by spin-labeled synthetic peptides: a model parallel coiled-coil peptide and the antiparallel coiled coil formed by a replica of the ProP C-terminus." Biochemistry 42(51);15170-8. PMID: 14690427

Hillar05: Hillar A, Culham DE, Vernikovska YI, Wood JM, Boggs JM (2005). "Formation of an antiparallel, intermolecular coiled coil is associated with in vivo dimerization of osmosensor and osmoprotectant transporter ProP in Escherichia coli." Biochemistry 44(30);10170-80. PMID: 16042394

Jebbar92: Jebbar M, Talibart R, Gloux K, Bernard T, Blanco C (1992). "Osmoprotection of Escherichia coli by ectoine: uptake and accumulation characteristics." J Bacteriol 1992;174(15);5027-35. PMID: 1629159

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

Kunte99: Kunte HJ, Crane RA, Culham DE, Richmond D, Wood JM (1999). "Protein ProQ influences osmotic activation of compatible solute transporter ProP in Escherichia coli K-12." J Bacteriol 181(5);1537-43. PMID: 10049386

Landis99: Landis L, Xu J, Johnson RC (1999). "The cAMP receptor protein CRP can function as an osmoregulator of transcription in Escherichia coli." Genes Dev 13(23);3081-91. PMID: 10601034

Liu07c: Liu F, Culham DE, Vernikovska YI, Keates RA, Boggs JM, Wood JM (2007). "Structure and function of transmembrane segment XII in osmosensor and osmoprotectant transporter ProP of Escherichia coli." Biochemistry 46(19);5647-55. PMID: 17441691

MacMillan99: MacMillan SV, Alexander DA, Culham DE, Kunte HJ, Marshall EV, Rochon D, Wood JM (1999). "The ion coupling and organic substrate specificities of osmoregulatory transporter ProP in Escherichia coli." Biochim Biophys Acta 1420(1-2);30-44. PMID: 10446288

May86: May G, Faatz E, Villarejo M, Bremer E (1986). "Binding protein dependent transport of glycine betaine and its osmotic regulation in Escherichia coli K12." Mol Gen Genet 205(2);225-33. PMID: 2949137

McLaggan91: McLaggan D, Epstein W (1991). "Escherichia coli accumulates the eukaryotic osmolyte taurine at high osmolarity." FEMS Microbiol Lett 1991;81(2);209-13. PMID: 1884995

McLeod00: McLeod SM, Xu J, Johnson RC (2000). "Coactivation of the RpoS-dependent proP P2 promoter by fis and cyclic AMP receptor protein." J Bacteriol 182(15);4180-7. PMID: 10894725

McLeod02: McLeod SM, Aiyar SE, Gourse RL, Johnson RC (2002). "The C-terminal domains of the RNA polymerase alpha subunits: contact site with Fis and localization during co-activation with CRP at the Escherichia coli proP P2 promoter." J Mol Biol 316(3);517-29. PMID: 11866515

McLeod99: McLeod SM, Xu J, Cramton SE, Gaal T, Gourse RL, Johnson RC (1999). "Localization of amino acids required for Fis to function as a class II transcriptional activator at the RpoS-dependent proP P2 promoter." J Mol Biol 294(2);333-46. PMID: 10610762

Mellies95: Mellies J, Wise A, Villarejo M (1995). "Two different Escherichia coli proP promoters respond to osmotic and growth phase signals." J Bacteriol 1995;177(1);144-51. PMID: 8002611

Mileykovskaya07: Mileykovskaya E (2007). "Subcellular localization of Escherichia coli osmosensory transporter ProP: focus on cardiolipin membrane domains." Mol Microbiol 64(6);1419-22. PMID: 17555431

Milner88: Milner JL, Grothe S, Wood JM (1988). "Proline porter II is activated by a hyperosmotic shift in both whole cells and membrane vesicles of Escherichia coli K12." J Biol Chem 1988;263(29);14900-5. PMID: 3049595

Milner89: Milner JL, Wood JM (1989). "Insertion proQ220::Tn5 alters regulation of proline porter II, a transporter of proline and glycine betaine in Escherichia coli." J Bacteriol 171(2);947-51. PMID: 2536686

Phadtare01: Phadtare S, Inouye M (2001). "Role of CspC and CspE in regulation of expression of RpoS and UspA, the stress response proteins in Escherichia coli." J Bacteriol 183(4);1205-14. PMID: 11157932

Racher01: Racher KI, Culham DE, Wood JM (2001). "Requirements for osmosensing and osmotic activation of transporter ProP from Escherichia coli." Biochemistry 40(24);7324-33. PMID: 11401581

Racher99: Racher KI, Voegele RT, Marshall EV, Culham DE, Wood JM, Jung H, Bacon M, Cairns MT, Ferguson SM, Liang WJ, Henderson PJ, White G, Hallett FR (1999). "Purification and reconstitution of an osmosensor: transporter ProP of Escherichia coli senses and responds to osmotic shifts." Biochemistry 1999;38(6);1676-84. PMID: 10026245

Randall95a: Randall K, Lever M, Peddie BA, Chambers ST (1995). "Competitive accumulation of betaines by Escherichia coli K-12 and derivative strains lacking betaine porters." Biochim Biophys Acta 1245(1);116-20. PMID: 7654759

Romantsov07: Romantsov T, Helbig S, Culham DE, Gill C, Stalker L, Wood JM (2007). "Cardiolipin promotes polar localization of osmosensory transporter ProP in Escherichia coli." Mol Microbiol 64(6);1455-65. PMID: 17504273

Sayeed09: Sayeed WM, Baenziger JE (2009). "Structural characterization of the osmosensor ProP." Biochim Biophys Acta 1788(5);1108-15. PMID: 19366597

Smith04a: Smith MN, Crane RA, Keates RA, Wood JM (2004). "Overexpression, purification, and characterization of ProQ, a posttranslational regulator for osmoregulatory transporter ProP of Escherichia coli." Biochemistry 43(41);12979-89. PMID: 15476391

Stalmach83: Stalmach ME, Grothe S, Wood JM (1983). "Two proline porters in Escherichia coli K-12." J Bacteriol 156(2);481-6. PMID: 6355059

Tsatskis05: Tsatskis Y, Khambati J, Dobson M, Bogdanov M, Dowhan W, Wood JM (2005). "The osmotic activation of transporter ProP is tuned by both its C-terminal coiled-coil and osmotically induced changes in phospholipid composition." J Biol Chem 280(50);41387-94. PMID: 16239220

Typas07: Typas A, Stella S, Johnson RC, Hengge R (2007). "The -35 sequence location and the Fis-sigma factor interface determine sigmas selectivity of the proP (P2) promoter in Escherichia coli." Mol Microbiol 63(3);780-96. PMID: 17302803

UniProt10: UniProt Consortium (2010). "UniProt version 2010-07 released on 2010-06-15 00:00:00." Database.

UniProtGOA11: UniProt-GOA (2011). "Gene Ontology annotation based on manual assignment of UniProtKB keywords in UniProtKB/Swiss-Prot entries."

UniProtGOA11a: UniProt-GOA (2011). "Gene Ontology annotation based on the manual assignment of UniProtKB Subcellular Location terms in UniProtKB/Swiss-Prot entries."

Verheul98: Verheul A, Wouters JA, Rombouts FM, Abee T (1998). "A possible role of ProP, ProU and CaiT in osmoprotection of Escherichia coli by carnitine." J Appl Microbiol 1998;85(6);1036-46. PMID: 9871325

Wood01: Wood JM, Bremer E, Csonka LN, Kraemer R, Poolman B, van der Heide T, Smith LT (2001). "Osmosensing and osmoregulatory compatible solute accumulation by bacteria." Comp Biochem Physiol A Mol Integr Physiol 130(3);437-60. PMID: 11913457

Wood05: Wood JM, Culham DE, Hillar A, Vernikovska YI, Liu F, Boggs JM, Keates RA (2005). "A structural model for the osmosensor, transporter, and osmoregulator ProP of Escherichia coli." Biochemistry 44(15);5634-46. PMID: 15823022

Wood06: Wood JM (2006). "Osmosensing by bacteria." Sci STKE 2006(357);pe43. PMID: 17047223

Wood80: Wood JM, Zadworny D (1980). "Amplification of the put genes and identification of the put gene products in Escherichia coli K12." Can J Biochem 58(10);787-96. PMID: 7006756

Wood99a: Wood JM (1999). "Osmosensing by bacteria: signals and membrane-based sensors." Microbiol Mol Biol Rev 63(1);230-62. PMID: 10066837

Xu95c: Xu J, Johnson RC (1995). "Fis activates the RpoS-dependent stationary-phase expression of proP in Escherichia coli." J Bacteriol 1995;177(18);5222-31. PMID: 7545153

Xu97b: Xu J, Johnson RC (1997). "Activation of RpoS-dependent proP P2 transcription by the Fis protein in vitro." J Mol Biol 270(3);346-59. PMID: 9237902

Xu97c: Xu J, Johnson RC (1997). "Cyclic AMP receptor protein functions as a repressor of the osmotically inducible promoter proP P1 in Escherichia coli." J Bacteriol 1997;179(7);2410-7. PMID: 9079929

Zhang07: Zhang N, Chen R, Young N, Wishart D, Winter P, Weiner JH, Li L (2007). "Comparison of SDS- and methanol-assisted protein solubilization and digestion methods for Escherichia coli membrane proteome analysis by 2-D LC-MS/MS." Proteomics 7(4);484-93. PMID: 17309111

Zoetewey03: Zoetewey DL, Tripet BP, Kutateladze TG, Overduin MJ, Wood JM, Hodges RS (2003). "Solution structure of the C-terminal antiparallel coiled-coil domain from Escherichia coli osmosensor ProP." J Mol Biol 334(5);1063-76. PMID: 14643666

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Please cite the following article in publications resulting from the use of EcoCyc: Nucleic Acids Research 41:D605-12 2013
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