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Escherichia coli K-12 substr. MG1655 Protein: TorS sensory histidine kinase



Gene: torS Accession Numbers: G6514 (EcoCyc), b0993, ECK0984

Synonyms: yccI

Regulation Summary Diagram: ?

Subunit composition of TorS sensory histidine kinase = [TorS]2
         TorS monomer = TorS

Alternative forms of TorS monomer:
TorS sensory histidine kinase - his850 phosphorylated (summary available)
TorS sensory histidine kinase - asp723 phosphorylated (summary available)
TorS sensory histidine kinase - his443 phosphorylated (summary available)

Summary:
TorS is the sensor partner of the periplasmic TorT protein and the TorR response regulator, responding to changes in the concentration of TMAO. TorS consists of an N-terminal detector region followed by a transmitter domain with a histidine phosphorylation site and an ATP binding motif, a receiver domain with an aspartate phosphorylation site, and a C-terminal alternative transmitter domain with a phosphorylation site. All three phosphorylation sites are essential for function. The C-terminal alternative transmitter domain contains the phosphodonor site for TorR [Jourlin97]. TorR is thus transphosphorylated by a four-step phosphorelay, beginning with autophosphorylation of TorS at His443, transfer of the phosphate group to Asp723 and finally His850, which then transfers the phosphate to TorR.

TorS may also be able to dephosphorylate TorR via a reverse phosphorelay when the TMAO inducer is removed [Ansaldi01]. The C-terminal domain of the immature form of TorC, a c-type cytochrome, can interact with the periplasmic detector region of TorS and downregulate its own expression, presumably via dephosphorylation of TorR [Ansaldi99, Gon01].

Certain mutations in torS cause TMAO-independent expression of the tor operon. One allele appears to affect the TMAO binding site, while two alleles localize to the linker region between the sensor and transmitter domains. A torS null mutation abolishes tor operon expression under all tested growth conditions. Overexpression of TorR bypasses the requirement for TorS, and TorT is dispensable in a torS constitutive mutant [Jourlin96a]. Complementation between torS mutants defective at different sites suggests that TorS forms multimers [Jourlin97].

Locations: inner membrane

Map Position: [1,052,657 <- 1,055,401] (22.69 centisomes)
Length: 2745 bp / 914 aa

Molecular Weight of Polypeptide: 101.02 kD (from nucleotide sequence)

Unification Links: ASAP:ABE-0003354 , DIP:DIP-11017N , EchoBASE:EB2501 , EcoGene:EG12617 , EcoliWiki:b0993 , ModBase:P39453 , OU-Microarray:b0993 , PortEco:torS , PR:PRO_000024090 , Pride:P39453 , Protein Model Portal:P39453 , RegulonDB:G6514 , SMR:P39453 , String:511145.b0993 , UniProt:P39453

Relationship Links: InterPro:IN-FAMILY:IPR001789 , InterPro:IN-FAMILY:IPR003594 , InterPro:IN-FAMILY:IPR003660 , InterPro:IN-FAMILY:IPR003661 , InterPro:IN-FAMILY:IPR004358 , InterPro:IN-FAMILY:IPR005467 , InterPro:IN-FAMILY:IPR008207 , InterPro:IN-FAMILY:IPR009082 , InterPro:IN-FAMILY:IPR011006 , InterPro:IN-FAMILY:IPR014302 , PDB:Structure:3I9W , Pfam:IN-FAMILY:PF00072 , Pfam:IN-FAMILY:PF00512 , Pfam:IN-FAMILY:PF00672 , Pfam:IN-FAMILY:PF01627 , Pfam:IN-FAMILY:PF02518 , Prints:IN-FAMILY:PR00344 , Prosite:IN-FAMILY:PS50109 , Prosite:IN-FAMILY:PS50110 , Prosite:IN-FAMILY:PS50885 , Prosite:IN-FAMILY:PS50894 , Smart:IN-FAMILY:SM00073 , Smart:IN-FAMILY:SM00304 , Smart:IN-FAMILY:SM00387 , Smart:IN-FAMILY:SM00388 , Smart:IN-FAMILY:SM00448

In Paralogous Gene Group: 121 (40 members) , 122 (29 members)

Reactions known to consume the compound:

TorSR Two-Component Signal Transduction System, TMAO dependent :
ATP + TorS sensory histidine kinase[inner membrane] → ADP + TorS sensory histidine kinase - his443 phosphorylated[inner membrane]

Reactions known to produce the compound:

TorSR Two-Component Signal Transduction System, TMAO dependent :
TorS sensory histidine kinase - his850 phosphorylated[inner membrane] + TorR → TorS sensory histidine kinase[inner membrane] + TorR-Pasp

Gene-Reaction Schematic: ?

GO Terms:

Biological Process: GO:0000160 - phosphorelay signal transduction system Inferred from experiment Inferred by computational analysis [UniProtGOA11a, GOA01a, Jourlin97]
GO:0009061 - anaerobic respiration Inferred from experiment [Jourlin96a]
GO:0016311 - dephosphorylation Inferred from experiment [Ansaldi01]
GO:0023014 - signal transduction by phosphorylation Inferred by computational analysis Inferred from experiment [Jourlin96a, GOA01a]
GO:0046777 - protein autophosphorylation Inferred from experiment [Jourlin97]
GO:0071310 - cellular response to organic substance Inferred from experiment [Jourlin96a]
GO:0007165 - signal transduction Inferred by computational analysis [GOA01a]
GO:0016310 - phosphorylation Inferred by computational analysis [UniProtGOA11a, GOA01a]
GO:0018106 - peptidyl-histidine phosphorylation Inferred by computational analysis [GOA01, GOA01a]
Molecular Function: GO:0000155 - phosphorelay sensor kinase activity Inferred from experiment Inferred by computational analysis [GOA01a, Jourlin96a]
GO:0004721 - phosphoprotein phosphatase activity Inferred from experiment [Ansaldi01]
GO:0005515 - protein binding Inferred from experiment [Gon01]
GO:0016772 - transferase activity, transferring phosphorus-containing groups Inferred from experiment Inferred by computational analysis [GOA01a, Jourlin97]
GO:0016868 - intramolecular transferase activity, phosphotransferases Inferred from experiment [Jourlin97]
GO:0042803 - protein homodimerization activity Inferred from experiment [Jourlin97]
GO:0000166 - nucleotide binding Inferred by computational analysis [UniProtGOA11a]
GO:0004673 - protein histidine kinase activity Inferred by computational analysis [GOA01, GOA01a]
GO:0004871 - signal transducer activity Inferred by computational analysis [GOA01a]
GO:0005524 - ATP binding Inferred by computational analysis [UniProtGOA11a]
GO:0016301 - kinase activity Inferred by computational analysis [UniProtGOA11a]
GO:0016740 - transferase activity Inferred by computational analysis [UniProtGOA11a]
Cellular Component: GO:0005886 - plasma membrane Inferred from experiment Inferred by computational analysis [UniProtGOA11, UniProtGOA11a, DiazMejia09, Daley05, Jourlin96a]
GO:0005887 - integral component of plasma membrane Inferred by computational analysis [Jourlin96a]
GO:0016020 - membrane Inferred by computational analysis [UniProtGOA11a, GOA01a]
GO:0016021 - integral component of membrane Inferred by computational analysis [UniProtGOA11a, GOA01a]

MultiFun Terms: cell structure membrane
information transfer protein related posttranslational modification
metabolism energy metabolism, carbon anaerobic respiration
regulation type of regulation posttranscriptional covalent modification, demodification, maturation
regulation type of regulation transcriptional level complex regulation two component regulatory systems (external signal)

Essentiality data for torS 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]

Sequence Features

Feature Class Location Citations Comment State
Transmembrane-Region 9 -> 29
[UniProt10a]
UniProt: Helical;; Non-Experimental Qualifier: potential;
 
Extrinsic-Sequence-Variant 295 -> 297
[UniProt10]
Alternate sequence: AQF → missing; UniProt: (in torS726; constitutively active);
 
Transmembrane-Region 333 -> 353
[UniProt10a]
UniProt: Helical;; Non-Experimental Qualifier: potential;
 
Conserved-Region 354 -> 407
[UniProt09]
UniProt: HAMP;
 
Extrinsic-Sequence-Variant 408
[UniProt10]
Alternate sequence: H → L; UniProt: (in torS729; partial activation);
 
Extrinsic-Sequence-Variant 414
[UniProt10]
Alternate sequence: A → P; UniProt: (in torS13; partial activation);
 
Conserved-Region 450 -> 664
[UniProt09]
UniProt: Histidine kinase;
 
Phosphorylation-Modification 453
[UniProt13]
UniProt: Phosphohistidine; by autocatalysis; Non-Experimental Qualifier: probable.
Unmodified
Mutagenesis-Variant 453
[Jourlin97, UniProt11a]
Alternate sequence: H → Q; UniProt: Loss of activity.
 
Conserved-Region 683 -> 798
[UniProt09]
UniProt: Response regulatory;
 
Mutagenesis-Variant 733
[Jourlin97, UniProt11a]
Alternate sequence: D → E; UniProt: Loss of activity.
Alternate sequence: D → A; UniProt: Loss of activity.
 
4-aspartylphosphate-Modification 733
[UniProt11a]
UniProt: 4-aspartylphosphate; Non-Experimental Qualifier: probable.
Unmodified
Sequence-Conflict 771 -> 772
[Jourlin96a, UniProt10]
Alternate sequence: QR → HG; UniProt: (in Ref. 1; CAA63920);
 
Conserved-Region 821 -> 914
[UniProt09]
UniProt: HPt;
 
Phosphorylation-Modification 860
[UniProt13]
UniProt: Phosphohistidine; Non-Experimental Qualifier: probable.
Unmodified
Mutagenesis-Variant 860
[Jourlin97, UniProt11a]
Alternate sequence: H → Q; UniProt: Decrease in activity.
 


Gene Local Context (not to scale): ?

Transcription Unit:

Notes:

History:
Peter D. Karp on Wed Jan 18, 2006:
Gene right-end position adjusted based on analysis performed in the 2005 E. coli annotation update [Riley06 ].
Markus Krummenacker on Tue Oct 14, 1997:
Gene object created from Blattner lab Genbank (v. M52) entry.


References

Ansaldi01: Ansaldi M, Jourlin-Castelli C, Lepelletier M, Theraulaz L, Mejean V (2001). "Rapid dephosphorylation of the TorR response regulator by the TorS unorthodox sensor in Escherichia coli." J Bacteriol 2001;183(8);2691-5. PMID: 11274133

Ansaldi99: Ansaldi M, Bordi C, Lepelletier M, Mejean V (1999). "TorC apocytochrome negatively autoregulates the trimethylamine N-oxide (TMAO) reductase operon in Escherichia coli." Mol Microbiol 33(2);284-95. PMID: 10411745

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

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

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, MGI (2001). "Gene Ontology annotation based on Enzyme Commission mapping." Genomics 74;121-128.

GOA01a: GOA, DDB, FB, MGI, ZFIN (2001). "Gene Ontology annotation through association of InterPro records with GO terms."

Gon01: Gon S, Jourlin-Castelli C, Theraulaz L, Mejean V (2001). "An unsuspected autoregulatory pathway involving apocytochrome TorC and sensor TorS in Escherichia coli." Proc Natl Acad Sci U S A 98(20);11615-20. PMID: 11562502

Jourlin96a: Jourlin C, Bengrine A, Chippaux M, Mejean V (1996). "An unorthodox sensor protein (TorS) mediates the induction of the tor structural genes in response to trimethylamine N-oxide in Escherichia coli." Mol Microbiol 1996;20(6);1297-306. PMID: 8809780

Jourlin97: Jourlin C, Ansaldi M, Mejean V (1997). "Transphosphorylation of the TorR response regulator requires the three phosphorylation sites of the TorS unorthodox sensor in Escherichia coli." J Mol Biol 1997;267(4);770-7. PMID: 9135110

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

Riley06: Riley M, Abe T, Arnaud MB, Berlyn MK, Blattner FR, Chaudhuri RR, Glasner JD, Horiuchi T, Keseler IM, Kosuge T, Mori H, Perna NT, Plunkett G, Rudd KE, Serres MH, Thomas GH, Thomson NR, Wishart D, Wanner BL (2006). "Escherichia coli K-12: a cooperatively developed annotation snapshot--2005." Nucleic Acids Res 34(1);1-9. PMID: 16397293

UniProt09: UniProt Consortium (2009). "UniProt version 15.8 released on 2009-10-01 00:00:00." Database.

UniProt10: UniProt Consortium (2010). "UniProt version 2010-11 released on 2010-11-02 00:00:00." Database.

UniProt10a: UniProt Consortium (2010). "UniProt version 2010-07 released on 2010-06-15 00:00:00." Database.

UniProt11a: UniProt Consortium (2011). "UniProt version 2011-11 released on 2011-11-22 00:00:00." Database.

UniProt13: UniProt Consortium (2013). "UniProt version 2013-08 released on 2013-08-01 00:00:00." Database.

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

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


Report Errors or Provide Feedback
Please cite the following article in publications resulting from the use of EcoCyc: Nucleic Acids Research 41:D605-12 2013
Page generated by SRI International Pathway Tools version 18.5 on Fri Dec 19, 2014, BIOCYC14A.