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Escherichia coli K-12 substr. MG1655 Transporter: N-acetylmuramic acid PTS permease

Synonyms: Enzyme IINAcMur, EIINAcMur

Subunit composition of N-acetylmuramic acid PTS permease = [MurP][Crr]
         N-acetylmuramic acid PTS permease / anhydro-N-acetylmuramic acid permease = MurP (summary available)
         Enzyme IIAGlc; Crr = Crr (extended summary available)

Summary:
MurP, the N-acetylmuramic acid PTS permease, belongs to the functional superfamily of the phosphoenolpyruvate (PEP)-dependent, sugar transporting phosphotransferase system (PTSsugar). The PTSsugar transports and simultaneously phosphorylates its sugar substrates in a process called group translocation (reviewed in [Postma93].

murP encodes a protein containing Enyme IIB (EIIB) and Enzyme IIC (EIIC) domains. Strains lacking murP are unable to grown on N-acetylmuramic acid as the sole source of carbon. Strains lacking crr, which encodes an Enzyme IIA (EIIA) protein are also unable to grow on N-acetylmuramic acid indicating that MurP, which lacks an EIIA domain requires Crr for activity. Sequence analysis suggests that a conserved cysteine residue at position 29 is the site of phosphotransfer [Dahl04]. MurP/Crr is a member of the PTS Glucose-Glucoside family of transporters [Saier14]. The product of MurP transport, N-acetyl-β-muramate 6-phosphate, is further degraded by the action of the MurQ etherase.

The overall PTS-mediated phosphoryl transfer reaction, requiring the two general energy coupling proteins of the PTSsugar, PtsI (Enzyme I) and PtsH (HPr), as well as the Crr protein and the two domains of the MurP protein can be represented as:

PEP → Enzyme I-Phis189 → HPr-Phis15 → Enzyme IIAglc-Phis90 → Enzyme IIB-Pcys29 - (Enzyme IIC) → N-acetylmuramic acid-6-P.

Gene-Reaction Schematic: ?

GO Terms:

Biological Process: GO:0009401 - phosphoenolpyruvate-dependent sugar phosphotransferase system Inferred by computational analysis Inferred from experiment [Dahl04]
Molecular Function: GO:0090588 - protein-phosphocysteine-N-acetylmuramate phosphotransferase system transporter activity Inferred from experiment Inferred by computational analysis [Dahl04]

Credits:
Last-Curated ? 16-Apr-2014 by Mackie A , Macquarie University


Enzymatic reaction of: N-acetylmuramic acid PTS permease

Synonyms: Transport of N-acetylmuramate

pH(opt): 6 [BRENDA14, Saier80], 6.5 [BRENDA14, Jacobson83], 7 [BRENDA14, Jacobson79], 7.5 [BRENDA14, Sutrina87], 8.5 [BRENDA14, Jacobson83], 9 [BRENDA14, Jacobson79], 9.5 [BRENDA14, Saier80]


Component enzyme of N-acetylmuramic acid PTS permease : N-acetylmuramic acid PTS permease / anhydro-N-acetylmuramic acid permease

Synonyms: MurP, YfeV, EIIBCNAcMur, Enzyme IIBCNAcMur

Gene: murP Accession Numbers: G7264 (EcoCyc), b2429, ECK2424

Locations: inner membrane

Sequence Length: 474 AAs

Molecular Weight: 49.802 kD (from nucleotide sequence)

Molecular Weight: 37 kD (experimental) [Dahl04]

GO Terms:

Biological Process: GO:0009401 - phosphoenolpyruvate-dependent sugar phosphotransferase system Inferred from experiment Inferred by computational analysis [UniProtGOA11a, GOA01a, Dahl04]
GO:0034219 - carbohydrate transmembrane transport Inferred from experiment [Dahl04]
GO:0006810 - transport Inferred by computational analysis [UniProtGOA11a]
GO:0008643 - carbohydrate transport Inferred by computational analysis [UniProtGOA11a]
GO:0016310 - phosphorylation Inferred by computational analysis [UniProtGOA11a]
Molecular Function: GO:0090588 - protein-phosphocysteine-N-acetylmuramate phosphotransferase system transporter activity Inferred by computational analysis Inferred from experiment [Dahl04]
GO:0008982 - protein-N(PI)-phosphohistidine-sugar phosphotransferase activity Inferred by computational analysis [GOA01, GOA01a]
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]
GO:0016020 - membrane Inferred from experiment Inferred by computational analysis [UniProtGOA11a, GOA01a, Dahl04]
GO:0005887 - integral component of plasma membrane Inferred by computational analysis [Dahl04]
GO:0016021 - integral component of membrane Inferred by computational analysis [UniProtGOA11a]

MultiFun Terms: metabolism carbon utilization carbon compounds
transport Group Translocators Phosphotransferase Systems (PEP-dependent PTS)

Unification Links: EcoliWiki:b2429 , ModBase:P77272 , PR:PRO_000023320 , Pride:P77272 , Protein Model Portal:P77272 , RefSeq:NP_416924 , SMR:P77272 , String:511145.b2429 , UniProt:P77272

Relationship Links: InterPro:IN-FAMILY:IPR001996 , InterPro:IN-FAMILY:IPR003352 , InterPro:IN-FAMILY:IPR013013 , InterPro:IN-FAMILY:IPR018113 , Pfam:IN-FAMILY:PF00367 , Pfam:IN-FAMILY:PF02378 , Prosite:IN-FAMILY:PS01035 , Prosite:IN-FAMILY:PS51098 , Prosite:IN-FAMILY:PS51103

Catalyzes:
1,6-anhydro-N-acetyl-β-muramate[periplasmic space] → 1,6-anhydro-N-acetyl-β-muramate[cytosol]

Summary:
murP encodes the permease component of the N-acetylmuramic acid PTS transport system. MurP contains PTS Enzyme IIB and IIC domains [Dahl04].

MurP is required for the uptake of anhydrous N-acetylmuramic (anhMurNAc) acid from the medium. MurP transports but does not phosphorylate anhMurNAc. The anmK encoded anhydro-N-acetylmuramic acid kinase is required to convert imported anhMurNAc to MurNAc-P. E. coli K-12 cannot use anhMurNAc as the sole source of carbon [Uehara05, Uehara06].

Essentiality data for murP 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]

Subunit of N-acetylmuramic acid PTS permease: Enzyme IIAGlc; Crr

Synonyms: TreD, Tgs, Iex, Gsr, EIIAGlc, Enzyme IIIGlc, EIIIGlc, IIIGlc

Gene: crr Accession Numbers: EG10165 (EcoCyc), b2417, ECK2412

Locations: inner membrane, cytosol

Sequence Length: 169 AAs

Molecular Weight: 18.251 kD (from nucleotide sequence)

Molecular Weight: 20.0 kD (experimental) [Dorschug84]

GO Terms:

Biological Process: GO:0009401 - phosphoenolpyruvate-dependent sugar phosphotransferase system Inferred from experiment Inferred by computational analysis [UniProtGOA11a, GOA01a, Buhr94, Dahl04, Boos90]
GO:0043610 - regulation of carbohydrate utilization Inferred from experiment [Saier76]
GO:0045912 - negative regulation of carbohydrate metabolic process Inferred from experiment [Dean90, Bao13, Novotny85, Nelson83]
GO:0006810 - transport Inferred by computational analysis [UniProtGOA11a]
GO:0008643 - carbohydrate transport Inferred by computational analysis [UniProtGOA11a]
GO:0016310 - phosphorylation Inferred by computational analysis [UniProtGOA11a]
Molecular Function: GO:0016301 - kinase activity Inferred from experiment Inferred by computational analysis [UniProtGOA11a, Dorschug84, Presper89, Wang00, Cai03]
GO:0016740 - transferase activity Inferred by computational analysis [UniProtGOA11a]
GO:0046872 - metal ion binding Inferred by computational analysis [UniProtGOA11a]
Cellular Component: GO:0005829 - cytosol Inferred from experiment Inferred by computational analysis [DiazMejia09, Ishihama08, LopezCampistrou05, Lasserre06]
GO:0016020 - membrane Inferred from experiment [Lasserre06]
GO:0031234 - extrinsic component of cytoplasmic side of plasma membrane Inferred from experiment [Wang00a, Chen13]
GO:0005737 - cytoplasm Inferred by computational analysis [UniProtGOA11, UniProtGOA11a]

MultiFun Terms: metabolism carbon utilization carbon compounds
transport Group Translocators Phosphotransferase Systems (PEP-dependent PTS)

Unification Links: DIP:DIP-31863N , EcoliWiki:b2417 , PR:PRO_000022334 , Pride:P69783 , Protein Model Portal:P69783 , RefSeq:NP_416912 , SMR:P69783 , String:511145.b2417 , UniProt:P69783

Relationship Links: InterPro:IN-FAMILY:IPR001127 , InterPro:IN-FAMILY:IPR011055 , PDB:Structure:1F3G , PDB:Structure:1F3Z , PDB:Structure:1GGR , PDB:Structure:1GLA , PDB:Structure:1GLB , PDB:Structure:1GLC , PDB:Structure:1GLD , PDB:Structure:1GLE , PDB:Structure:1O2F , PDB:Structure:2F3G , PDB:Structure:4JBW , Pfam:IN-FAMILY:PF00358 , Prosite:IN-FAMILY:PS00371 , Prosite:IN-FAMILY:PS51093

Summary:
EIIAGlc is an intermediate phosphotransfer protein in the uptake and phosphorylation of glucose [Buhr94], N-acetylmuramic acid [Dahl04] and trehalose [Boos90] and also probably interacts with MalX - a PTS permease whose physiological substrate is unclear [Reidl91]. EIIAGlc accepts a phosphoryl group from PtsH and transfers it to the the EIIB domain of the sugar PTS permeases.

EIIAGlc is phosphorylated by phosphoenolpyruvate (PEP) in a reaction requiring PtsH and PtsI. EIIAGlc is phosphorylated at the N3 position of His90 [Dorschug84, Presper89]. Solution structures of the complex between PtsH and EIIAGlc and between EIIAGlc and EIIBGlc have been determined. The binding surface on EIIAGlc for EIIBGlc and PtsH overlap. There are both similarities and differences between the binding interaction of EIIAGlc with PtsH and EIIBGlc [Wang00, Cai03].

Two forms of EIIAGlc have been purified from E.coli K-12 and identified on the basis of electrophoretic mobility as EIIAGlcfast and EIIAGlcslow. EIIAGlcfast lacks an N-terminal heptapeptide region due to proteolytic activity which appears to reside in the outer membrane. The two forms are equally able to act as phosphoacceptors but EIIAGlcfast is an inefficient phosphodonor [Meadow86, Meadow82].

Crr or Enzyme IIAGlc also plays an important role in the regulation of carbon metabolism in E. coli K-12 ([Saier76] and reviewed in [Postma93] and [Deutscher06]). EIIAGlc is a negative regulator of several enzymes involved in the transport and metabolism of non-PTS carbohydrates such as glycerol, lactose, melibiose and maltose. Purified, unphosphorylated EIIAGlc binds to, and inhibits galactoside transport by the lactose permease in vitro [Nelson83, Osumi82]. Purified, unphosphorylated EIIAGlc binds to, and inhibits glycerol kinase in vitro [Novotny85, deBoer86].

EIIAGlc inhibits the activity of the maltose ABC transporter reconstituted in proteopliposomes [Dean90]. EIIAGlc interacts with the nucleotide binding domain and the C-terminal regulatory domain of the MalK dimer and inhibits the ATPase activity of the maltose transporter reconstituted in proteoliposomes. The stoichiometry of interaction between EIIAGlc and MalFGK2 is 2:1 [Bao13]. A crystal structure of EIIAGlc in complex with the maltose ABC transporter has been obtained. Two molecules of EIIAGlc interact with the MalK subunits and stabilise an inward facing conformation of the transporter. This interaction would prevent the structural rearrangements required for ATP hydrolysis and maltose transport. The phosphorylation site, His90, is located at the EIIAGlc-MalK interface, and is in its unphosphorylated form. Phosphorylation of the active histidine is predicted to disrupt the interface [Chen13].

The N-terminal domain of EIIAGlc (residues 1-18) adopts a helical conformation in detergent micelles and in the presence of E. coli lipids. The N-terminal domain probably acts as a membrane anchor and allows EIIAGlc to shuttle between the cytoplasm and the inner membrane [Wang00a, Wang03]. Full-length EIIAGlc is concentrated near the inner surface of the cytoplasmic membrane via its N-terminal region [Chen13].

EIIAGlc is also involved in the regulation of adenylate cyclase activity although the precise mechanism remains unclear [Saier76a, Feucht80, Daniel84, Liberman86, Peterkofsky88, Levy90, Takahashi98]. A direct interaction between membrane tethered EIIAGlc (both phosphorylated and unphosphorylated) and adenylate cyclase has been demonstrated in vitro [Park06a]. Adenylate cyclase activity is not affected by either phosphorylated or dephosphorylated EIIAGlc however phosphorylated EIIAGlc activates adenylate cyclase in the presence of a crude E. coli extract implying that regulation depends upon an additional unidentified factor [Park06a]. Phosphorylated EIIAGlc is a positive effector of adenylate cyclase activity while EIIAGlc does not affect basal activity [Reddy98].

crr insertion mutants were identified in a genetic screen for genes that are important for survival of exposure to ionizing radiation (IR). A crr deletion mutant has a moderate decrease in IR survival [Byrne14].

Reviews: [Chen93b, Peterkofsky89]

crr: carbohydrate repression resistant

Citations: [Pelton92, Pelton91, Pelton93, Worthylake91, Meadow06]

Essentiality data for crr 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 No 37 Aerobic 7.2 0.35 No [Joyce06]
MOPS medium with 0.4% glucose Yes 37 Aerobic 7.2 0.22 Yes [Baba06, Comment 2]
Yes [Feist07, Comment 4]

References

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Uehara05: Uehara T, Suefuji K, Valbuena N, Meehan B, Donegan M, Park JT (2005). "Recycling of the anhydro-N-acetylmuramic acid derived from cell wall murein involves a two-step conversion to N-acetylglucosamine-phosphate." J Bacteriol 187(11);3643-9. PMID: 15901686

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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."

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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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