Escherichia coli K-12 substr. MG1655 Transporter: nucleoside channel; receptor of phage T6 and colicin K

Gene: tsx Accession Numbers: EG11035 (EcoCyc), b0411, ECK0405

Synonyms: T6rec, nupA

Regulation Summary Diagram: ?

Regulation summary diagram for tsx

Tsx is an outer membrane porin which is responsible for the uptake of nucleosides and deoxynucleosides at sub-uM concentrations [Hantke76, Maier88]. Reconstitution of purified Tsx into lipid bilayer membranes suggests that Tsx has a greater affinity for deoxynucleosides than for nucleosides and that it does not play a role in the transport of nucleobases [Benz88]. Tsx forms a monomeric βbarrel consisting of 12 β strands [Ye04].

In addition to being a channel for (deoxy)nucleosides Tsx functions as a receptor for bacteriophage and colicins and also transports the antibiotic, albicidin [Hantke76, Birch90]. Tsx has been shown to localize to the cellular poles [Lai04a].

Gene Citations: [Bremer90]

Locations: outer membrane

Map Position: [430,353 <- 431,237] (9.28 centisomes, 33°)
Length: 885 bp / 294 aa

Molecular Weight of Polypeptide: 33.589 kD (from nucleotide sequence)

Unification Links: ASAP:ABE-0001425 , CGSC:62 , EchoBASE:EB1028 , EcoGene:EG11035 , EcoliWiki:b0411 , OU-Microarray:b0411 , PortEco:tsx , PR:PRO_000024135 , Pride:P0A927 , Protein Model Portal:P0A927 , RefSeq:NP_414945 , RegulonDB:EG11035 , SMR:P0A927 , String:511145.b0411 , UniProt:P0A927

Relationship Links: InterPro:IN-FAMILY:IPR003055 , InterPro:IN-FAMILY:IPR018013 , PDB:Structure:1TLW , PDB:Structure:1TLY , PDB:Structure:1TLZ , Pfam:IN-FAMILY:PF03502 , Prints:IN-FAMILY:PR01277

Gene-Reaction Schematic: ?

Gene-Reaction Schematic

Genetic Regulation Schematic: ?

Genetic regulation schematic for tsx

GO Terms:

Biological Process: GO:0015858 - nucleoside transport Inferred from experiment [Benz88]
GO:0046718 - viral entry into host cell Inferred from experiment [Hantke76]
GO:1901642 - nucleoside transmembrane transport Inferred from experiment Inferred by computational analysis [GOA01, Benz88]
GO:0006810 - transport Inferred by computational analysis [UniProtGOA11a]
GO:0006811 - ion transport Inferred by computational analysis [UniProtGOA11a]
GO:0055085 - transmembrane transport Inferred by computational analysis [UniProtGOA11a]
Molecular Function: GO:0015471 - nucleoside-specific channel forming porin activity Inferred from experiment [Benz88]
GO:0005337 - nucleoside transmembrane transporter activity Inferred by computational analysis [GOA01]
GO:0015288 - porin activity Inferred by computational analysis [UniProtGOA11a]
Cellular Component: GO:0009279 - cell outer membrane Inferred from experiment Inferred by computational analysis [UniProtGOA11, UniProtGOA11a, GOA01, Benz88]
GO:0046930 - pore complex Inferred from experiment Inferred by computational analysis [UniProtGOA11a, Ye04]
GO:0016020 - membrane Inferred by computational analysis [UniProtGOA11a]
GO:0016021 - integral component of membrane Inferred by computational analysis [UniProtGOA11a]

MultiFun Terms: extrachromosomal colicin related
extrachromosomal prophage genes and phage related functions
metabolism central intermediary metabolism nucleotide and nucleoside conversions
transport Channel-type Transporters Beta barrel porins (The Outer Membrane Porin (OMP) Functional Superfamily)

Essentiality data for tsx 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]

Last-Curated ? 09-May-2011 by Mackie A , Macquarie University

Enzymatic reaction of: outer membrane (deoxy)nucleoside channel (nucleoside channel; receptor of phage T6 and colicin K)

Transport reaction diagram for outer membrane (deoxy)nucleoside channel

Sequence Features

Protein sequence of nucleoside channel; receptor of phage T6 and colicin K with features indicated

Feature Class Location Citations Comment
Signal-Sequence 1 -> 22
UniProt: Non-Experimental Qualifier: potential;
Chain 23 -> 294
UniProt: Nucleoside-specific channel-forming protein tsx;
Extrinsic-Sequence-Variant 276
UniProt: In phage resistant mutant..

Gene Local Context (not to scale): ?

Gene local context diagram

Transcription Units:

Transcription-unit diagram

Transcription-unit diagram


10/20/97 Gene b0411 from Blattner lab Genbank (v. M52) entry merged into EcoCyc gene EG11035; confirmed by SwissProt match.


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

Benz88: Benz R, Schmid A, Maier C, Bremer E (1988). "Characterization of the nucleoside-binding site inside the Tsx channel of Escherichia coli outer membrane. Reconstitution experiments with lipid bilayer membranes." Eur J Biochem 176(3);699-705. PMID: 2458926

Birch90: Birch RG, Pemberton JM, Basnayake WV (1990). "Stable albicidin resistance in Escherichia coli involves an altered outer-membrane nucleoside uptake system." J Gen Microbiol 136(1);51-8. PMID: 2191080

Bremer90: Bremer E, Middendorf A, Martinussen J, Valentin-Hansen P (1990). "Analysis of the tsx gene, which encodes a nucleoside-specific channel-forming protein (Tsx) in the outer membrane of Escherichia coli." Gene 1990;96(1);59-65. PMID: 2265760

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

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."

Hantke76: Hantke K (1976). "Phage T6--colicin K receptor and nucleoside transport in Escherichia coli." FEBS Lett 70(1);109-12. PMID: 791677

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

Lai04a: Lai EM, Nair U, Phadke ND, Maddock JR (2004). "Proteomic screening and identification of differentially distributed membrane proteins in Escherichia coli." Mol Microbiol 52(4);1029-44. PMID: 15130122

Maier88: Maier C, Bremer E, Schmid A, Benz R (1988). "Pore-forming activity of the Tsx protein from the outer membrane of Escherichia coli. Demonstration of a nucleoside-specific binding site." J Biol Chem 1988;263(5);2493-9. PMID: 3276691

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

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

UniProt15: UniProt Consortium (2015). "UniProt version 2015-01 released on 2015-01-16 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."

Ye04: Ye J, Van Den Berg B (2004). "Crystal structure of the bacterial nucleoside transporter Tsx." EMBO J 23(16);3187-95. PMID: 15272310

Other References Related to Gene Regulation

Bremer88: Bremer E, Gerlach P, Middendorf A (1988). "Double negative and positive control of tsx expression in Escherichia coli." J Bacteriol 170(1);108-16. PMID: 2826388

Brikun96: Brikun I, Suziedelis K, Stemmann O, Zhong R, Alikhanian L, Linkova E, Mironov A, Berg DE (1996). "Analysis of CRP-CytR interactions at the Escherichia coli udp promoter." J Bacteriol 1996;178(6);1614-22. PMID: 8626289

Gavigan99: Gavigan SA, Nguyen T, Nguyen N, Senear DF (1999). "Role of multiple CytR binding sites on cooperativity, competition, and induction at the Escherichia coli udp promoter." J Biol Chem 1999;274(23);16010-9. PMID: 10347150

Gerlach91: Gerlach P, Sogaard-Andersen L, Pedersen H, Martinussen J, Valentin-Hansen P, Bremer E (1991). "The cyclic AMP (cAMP)-cAMP receptor protein complex functions both as an activator and as a corepressor at the tsx-p2 promoter of Escherichia coli K-12." J Bacteriol 1991;173(17);5419-30. PMID: 1715855

Gogol11: Gogol EB, Rhodius VA, Papenfort K, Vogel J, Gross CA (2011). "Small RNAs endow a transcriptional activator with essential repressor functions for single-tier control of a global stress regulon." Proc Natl Acad Sci U S A 108(31);12875-80. PMID: 21768388

Holst92: Holst B, Sogaard-Andersen L, Pedersen H, Valentin-Hansen P (1992). "The cAMP-CRP/CytR nucleoprotein complex in Escherichia coli: two pairs of closely linked binding sites for the cAMP-CRP activator complex are involved in combinatorial regulation of the cdd promoter." EMBO J 11(10);3635-43. PMID: 1327747

Holt10: Holt AK, Senear DF (2010). "An unusual pattern of CytR and CRP binding energetics at Escherichia coli cddP suggests a unique blend of class I and class II mediated activation." Biochemistry 49(3);432-42. PMID: 20000490

Jorgensen98: Jorgensen CI, Kallipolitis BH, Valentin-Hansen P (1998). "DNA-binding characteristics of the Escherichia coli CytR regulator: a relaxed spacing requirement between operator half-sites is provided by a flexible, unstructured interdomain linker." Mol Microbiol 27(1);41-50. PMID: 9466254

Meibom00: Meibom KL, Kallipolitis BH, Ebright RH, Valentin-Hansen P (2000). "Identification of the subunit of cAMP receptor protein (CRP) that functionally interacts with CytR in CRP-CytR-mediated transcriptional repression." J Biol Chem 275(16);11951-6. PMID: 10766824

Pedersen97: Pedersen H, Valentin-Hansen P (1997). "Protein-induced fit: the CRP activator protein changes sequence-specific DNA recognition by the CytR repressor, a highly flexible LacI member." EMBO J 16(8);2108-18. PMID: 9155036

Perini96: Perini LT, Doherty EA, Werner E, Senear DF (1996). "Multiple specific CytR binding sites at the Escherichia coli deoP2 promoter mediate both cooperative and competitive interactions between CytR and cAMP receptor protein." J Biol Chem 271(52);33242-55. PMID: 8969182

SogaardAndersen90: Sogaard-Andersen L, Mollegaard NE, Douthwaite SR, Valentin-Hansen P (1990). "Tandem DNA-bound cAMP-CRP complexes are required for transcriptional repression of the deoP2 promoter by the CytR repressor in Escherichia coli." Mol Microbiol 4(9);1595-601. PMID: 1962841

SogaardAndersen90a: Sogaard-Andersen L, Martinussen J, Mollegaard NE, Douthwaite SR, Valentin-Hansen P (1990). "The CytR repressor antagonizes cyclic AMP-cyclic AMP receptor protein activation of the deoCp2 promoter of Escherichia coli K-12." J Bacteriol 1990;172(10);5706-13. PMID: 2170326

SogaardAndersen91b: Sogaard-Andersen L, Pedersen H, Holst B, Valentin-Hansen P (1991). "A novel function of the cAMP-CRP complex in Escherichia coli: cAMP-CRP functions as an adaptor for the CytR repressor in the deo operon." Mol Microbiol 5(4);969-75. PMID: 1649947

ValentinHansen96: Valentin-Hansen P, Sogaard-Andersen L, Pedersen H (1996). "A flexible partnership: the CytR anti-activator and the cAMP-CRP activator protein, comrades in transcription control." Mol Microbiol 20(3);461-6. PMID: 8736525

Zheng04: Zheng D, Constantinidou C, Hobman JL, Minchin SD (2004). "Identification of the CRP regulon using in vitro and in vivo transcriptional profiling." Nucleic Acids Res 32(19);5874-93. PMID: 15520470

Zolotukhina02: Zolotukhina MA, Ovcharova IV, Eremina SIu, Errais LL, Mironov AS (2002). "[Functional interrelationship between elements of the Escherichia coli udp gene promotor responsible for binding regulatory proteins CytR, CRP, and RNA polymerase]." Genetika 38(9);1223-34. PMID: 12391883

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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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