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Escherichia coli K-12 substr. MG1655 Transporter: mannitol PTS permease



Gene: mtlA Accession Numbers: EG10615 (EcoCyc), b3599, ECK3588

Synonyms: mtlC, EIIMtl

Regulation Summary Diagram: ?

Subunit composition of mannitol PTS permease = [MtlA]2

Summary:
MtlA, the mannitol 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]). MtlA takes up exogenous mannitol, releasing the phosphate ester, mannitol-1-P, into the cell cytoplasm in preparation for oxidation to fructose-6-P by the NAD-dependent mannitol-P dehydrogenase (MtlD) and subsequent metabolism via glycolysis (mannitol degradation I). Purified MtlA transports D-glucitol and D-arabitol with low efficiency [Jacobson79, Jacobson83].

MtlA, the Enzyme IIMtl complex, possesses three domains in a single polypeptide chain with the domain order IIC-IIB-IIA. MtlA consists of an N-terminal hydrophobic domain (IIC) that resides in the inner membrane and a C-terminal, hydrophilic domain (IIBA) that is located on the cytoplasmic side of the membrane [Lee83a, Jacobson83a]. Histidine 554 and cysteine 384 located in the hydrophilic domain are the phosphotransfer sites. His554 accepts a phosphoryl group from the cytoplasmic HPr protein; Cys384 is the site of phosphotransfer to D-mannitol [Pas88, Pas88a, White90, vanWeeghel91, vanWeeghel91a]. The N-terminal domain of MtlA contains 6 transmembrane α-helical segments [Sugiyama91] and is responsible for binding and translocation of mannitol [Lolkema90].

MtlA functions as a homodimer [Pas87, Boer94, Boer96]. MtlA is an asymmetric dimer [Vos09, Opa12] MtlA contains one high affinity mannitol binding site and one low affinity binding site [Pas88a]. Unphosphorylated MtlA contains one high affinity mannitol binding site [Veldhuis05]. The mannitol binding site is located in the middle of the membrane - this is also the site where mannitol is phosphorylated [Opacic10]. Phosphotransfer from His554 to Cys384 occurs between subunits in a permease dimer [Stephan89]. MtlA is a member of the PTS Fructose-Mannitol (Fru) family of transporters [Saier14].

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

PEP → Enzyme I-Phis189 → HPr-Phis15 → Enzyme IIA-Phis554 → Enzyme IIB-Pcys384 - (Enzyme IIC) → mannitol-1-P.

MtlA catalyses facilitated diffusion of mannitol at a low rate in vitro [Lolkema90, Elferink90]. MtlA catalyses vectorial and non-vectorial mannitol - mannitol-1-phosphate transphosphorylation in intact cells and membrane vesicles [Saier81, Leonard83]. MtlA catalyses an HPr / HPr-P phospho exchange reaction in vitro [Sutrina87].

The mtl operon (mtlADR) is inducible (~~20x) by growth of wild type E. coli K12 in the presence of mannitol. The MtlR protein is a negative transcriptional regulator of the operon [Figge94]. The operon is also positively controlled by the cyclic AMP-cyclic AMP receptor protein (CRP) complex and negatively by the catabolite repressor/activator (Cra) protein.

Reviews: [Jacobson93, Robillard89, Jacobson, Robillard, Jacobson89, Jacobson92]

Citations: [Vos08a, Grisafi89, Lolkema93, Kroon93, Dijkstra97, Boer95, SaraceniRichard97, SaraceniRichard97a, Weng93, Stephan86, Grenier86, Yamada91]

Locations: inner membrane

Map Position: [3,770,304 -> 3,772,217] (81.26 centisomes)
Length: 1914 bp / 637 aa

Molecular Weight of Polypeptide: 67.972 kD (from nucleotide sequence)

Unification Links: ASAP:ABE-0011761 , CGSC:481 , DIP:DIP-10267N , EchoBASE:EB0610 , EcoGene:EG10615 , EcoliWiki:b3599 , Mint:MINT-245984 , ModBase:P00550 , OU-Microarray:b3599 , PortEco:mtlA , PR:PRO_000023303 , Pride:P00550 , Protein Model Portal:P00550 , RefSeq:NP_418056 , RegulonDB:EG10615 , SMR:P00550 , String:511145.b3599 , UniProt:P00550

Relationship Links: InterPro:IN-FAMILY:IPR002178 , InterPro:IN-FAMILY:IPR003352 , InterPro:IN-FAMILY:IPR003501 , InterPro:IN-FAMILY:IPR004718 , InterPro:IN-FAMILY:IPR013011 , InterPro:IN-FAMILY:IPR013014 , InterPro:IN-FAMILY:IPR016152 , PDB:Structure:1A3A , PDB:Structure:1J6T , PDB:Structure:1VKR , PDB:Structure:1VRV , PDB:Structure:2FEW , Pfam:IN-FAMILY:PF00359 , Pfam:IN-FAMILY:PF02302 , Pfam:IN-FAMILY:PF02378 , Prosite:IN-FAMILY:PS00372 , Prosite:IN-FAMILY:PS51094 , Prosite:IN-FAMILY:PS51099 , Prosite:IN-FAMILY:PS51104

In Paralogous Gene Group: 187 (14 members)

Gene-Reaction Schematic: ?

Genetic Regulation Schematic: ?

GO Terms:

Biological Process: GO:0009401 - phosphoenolpyruvate-dependent sugar phosphotransferase system Inferred from experiment Inferred by computational analysis [UniProtGOA11a, GOA01a, Jacobson83a, Grisafi89]
GO:0015795 - sorbitol transport Inferred from experiment [Lengeler75a]
GO:0015797 - mannitol transport Inferred from experiment [Jacobson83a, Grisafi89]
GO:0034219 - carbohydrate transmembrane transport Inferred by computational analysis Inferred from experiment [Jacobson83a, Pas88, GOA01, GOA01a]
GO:0006810 - transport Inferred by computational analysis [UniProtGOA11a, GOA01a]
GO:0008643 - carbohydrate transport Inferred by computational analysis [UniProtGOA11a]
GO:0016310 - phosphorylation Inferred by computational analysis [UniProtGOA11a]
Molecular Function: GO:0090565 - protein-phosphocysteine-mannitol phosphotransferase system transporter activity Inferred from experiment [Jacobson83a, Pas88]
GO:0005215 - transporter activity Inferred by computational analysis [GOA01a]
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]
GO:0022872 - protein-N(PI)-phosphohistidine-mannitol phosphotransferase system transmembrane transporter activity Inferred by computational analysis [GOA01a]
Cellular Component: GO:0005886 - plasma membrane Inferred from experiment Inferred by computational analysis [UniProtGOA11, UniProtGOA11a, DiazMejia09, Zhang07, Daley05, LopezCampistrou05]
GO:0005887 - integral component of plasma membrane Inferred from experiment [Jacobson83b, Sugiyama91]
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
metabolism carbon utilization carbon compounds
transport Group Translocators Phosphotransferase Systems (PEP-dependent PTS)

Essentiality data for mtlA 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]

Credits:
Last-Curated ? 13-Mar-2014 by Mackie A , Macquarie University


Enzymatic reaction of: transport and phosphorylation of D-mannitol (mannitol PTS permease)

Inhibitors (Unknown Mechanism): D-sorbitol [Jacobson83]

Kinetic Parameters:

Substrate
Km (μM)
Citations
D-mannitol
10.0
[Jacobson83]


Enzymatic reaction of: transport and phosphorylation of D-sorbitol (mannitol PTS permease)

Summary:
MtlA transports D-sorbitol with low efficiency [Lengeler75a]


Sequence Features

Feature Class Location Citations Comment
Conserved-Region 12 -> 341
[UniProt09]
UniProt: PTS EIIC type-2;
Transmembrane-Region 25 -> 44
[UniProt10a]
UniProt: Helical;; Non-Experimental Qualifier: probable;
Transmembrane-Region 51 -> 69
[UniProt10a]
UniProt: Helical;; Non-Experimental Qualifier: probable;
Transmembrane-Region 135 -> 154
[UniProt10a]
UniProt: Helical;; Non-Experimental Qualifier: probable;
Transmembrane-Region 166 -> 184
[UniProt10a]
UniProt: Helical;; Non-Experimental Qualifier: probable;
Transmembrane-Region 274 -> 291
[UniProt10a]
UniProt: Helical;; Non-Experimental Qualifier: probable;
Transmembrane-Region 314 -> 333
[UniProt10a]
UniProt: Helical;; Non-Experimental Qualifier: probable;
Conserved-Region 378 -> 473
[UniProt09]
UniProt: PTS EIIB type-2;
Active-Site 384
[Pas88, UniProt11]
UniProt: Phosphocysteine intermediate; for EIIB activity.
Conserved-Region 494 -> 636
[UniProt09]
UniProt: PTS EIIA type-2;
Active-Site 554
[Pas88, UniProt11]
UniProt: Tele-phosphohistidine intermediate; for EIIA activity.


Gene Local Context (not to scale): ?

Transcription Unit:

Notes:

History:
10/20/97 Gene b3599 from Blattner lab Genbank (v. M52) entry merged into EcoCyc gene EG10615; confirmed by SwissProt match.


References

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

Boer94: Boer H, ten Hoeve-Duurkens RH, Schuurman-Wolters GK, Dijkstra A, Robillard GT (1994). "Expression, purification, and kinetic characterization of the mannitol transport domain of the phosphoenolpyruvate-dependent mannitol phosphotransferase system of Escherichia coli. Kinetic evidence that the E. coli mannitol transport protein is a functional dimer." J Biol Chem 269(27);17863-71. PMID: 8027040

Boer95: Boer H, ten Hoeve-Duurkens RH, Lolkema JS, Robillard GT (1995). "Phosphorylation site mutants of the mannitol transport protein enzyme IImtl of Escherichia coli: studies on the interaction between the mannitol translocating C-domain and the phosphorylation site on the energy-coupling B-domain." Biochemistry 34(10);3239-47. PMID: 7880818

Boer96: Boer H, ten Hoeve-Duurkens RH, Robillard GT (1996). "Relation between the oligomerization state and the transport and phosphorylation function of the Escherichia coli mannitol transport protein: interaction between mannitol-specific enzyme II monomers studied by complementation of inactive site-directed mutants." Biochemistry 35(39);12901-8. PMID: 8841134

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

Dijkstra97: Dijkstra DS, Broos J, Visser AJ, van Hoek A, Robillard GT (1997). "Dynamic fluorescence spectroscopy on single tryptophan mutants of EII(mtl) in detergent micelles. Effects of substrate binding and phosphorylation on the fluorescence and anisotropy decay." Biochemistry 36(16);4860-6. PMID: 9125506

Elferink90: Elferink MG, Driessen AJ, Robillard GT (1990). "Functional reconstitution of the purified phosphoenolpyruvate-dependent mannitol-specific transport system of Escherichia coli in phospholipid vesicles: coupling between transport and phosphorylation." J Bacteriol 172(12);7119-25. PMID: 2123863

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

Figge94: Figge RM, Ramseier TM, Saier MH (1994). "The mannitol repressor (MtlR) of Escherichia coli." J Bacteriol 1994;176(3);840-7. PMID: 8300537

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."

Grenier86: Grenier FC, Waygood EB, Saier MH (1986). "The bacterial phosphotransferase system: kinetic characterization of the glucose, mannitol, glucitol, and N-acetylglucosamine systems." J Cell Biochem 31(2);97-105. PMID: 3015992

Grisafi89: Grisafi PL, Scholle A, Sugiyama J, Briggs C, Jacobson GR, Lengeler JW (1989). "Deletion mutants of the Escherichia coli K-12 mannitol permease: dissection of transport-phosphorylation, phospho-exchange, and mannitol-binding activities." J Bacteriol 171(5);2719-27. PMID: 2496114

Jacobson: Jacobson GR "Coupling of energy to D-mannitol transport in Escherichia coli." Res Microbiol 141(3);365-8. PMID: 2126390

Jacobson79: Jacobson GR, Lee CA, Saier MH (1979). "Purification of the mannitol-specific enzyme II of the Escherichia coli phosphoenolpyruvate:sugar phosphotransferase system." J Biol Chem 254(2);249-52. PMID: 368051

Jacobson83: Jacobson GR, Tanney LE, Kelly DM, Palman KB, Corn SB (1983). "Substrate and phospholipid specificity of the purified mannitol permease of Escherichia coli." J Cell Biochem 23(1-4);231-40. PMID: 6427236

Jacobson83a: Jacobson GR, Lee CA, Leonard JE, Saier MH (1983). "Mannitol-specific enzyme II of the bacterial phosphotransferase system. I. Properties of the purified permease." J Biol Chem 258(17);10748-56. PMID: 6350293

Jacobson83b: Jacobson GR, Kelly DM, Finlay DR (1983). "The intramembrane topography of the mannitol-specific enzyme II of the Escherichia coli phosphotransferase system." J Biol Chem 258(5);2955-9. PMID: 6338009

Jacobson89: Jacobson GR, Stephan MM (1989). "Structural and functional domains of the mannitol-specific enzyme II of the E. coli phosphoenolpyruvate-dependent phosphotransferase system." FEMS Microbiol Rev 5(1-2);25-34. PMID: 2517400

Jacobson92: Jacobson GR (1992). "Interrelationships between protein phosphorylation and oligomerization in transport and chemotaxis via the Escherichia coli mannitol phosphotransferase system." Res Microbiol 143(1);113-6. PMID: 1641509

Jacobson93: Jacobson GR, Saraceni-Richards C (1993). "The Escherichia coli mannitol permease as a model for transport via the bacterial phosphotransferase system." J Bioenerg Biomembr 25(6);621-6. PMID: 8144490

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

Kroon93: Kroon GJ, Grotzinger J, Dijkstra K, Scheek RM, Robillard GT (1993). "Backbone assignments and secondary structure of the Escherichia coli enzyme-II mannitol A domain determined by heteronuclear three-dimensional NMR spectroscopy." Protein Sci 1993;2(8);1331-41. PMID: 8401218

Lee83a: Lee CA, Saier MH (1983). "Mannitol-specific enzyme II of the bacterial phosphotransferase system. III. The nucleotide sequence of the permease gene." J Biol Chem 1983;258(17);10761-7. PMID: 6309813

Lengeler75a: Lengeler J (1975). "Nature and properties of hexitol transport systems in Escherichia coli." J Bacteriol 124(1);39-47. PMID: 1100608

Leonard83: Leonard JE, Saier MH (1983). "Mannitol-specific enzyme II of the bacterial phosphotransferase system. II. Reconstitution of vectorial transphosphorylation in phospholipid vesicles." J Biol Chem 258(17);10757-60. PMID: 6350294

Lolkema90: Lolkema JS, Dijkstra DS, ten Hoeve-Duurkens RH, Robillard GT (1990). "The membrane-bound domain of the phosphotransferase enzyme IImtl of Escherichia coli constitutes a mannitol translocating unit." Biochemistry 29(47);10659-63. PMID: 2125492

Lolkema93: Lolkema JS, Kuiper H, ten Hoeve-Duurkens RH, Robillard GT (1993). "Mannitol-specific enzyme II of the phosphoenolpyruvate-dependent phosphotransferase system of Escherichia coli: physical size of enzyme IImtl and its domains IIBA and IIC in the active state." Biochemistry 32(6);1396-400. PMID: 8431419

LopezCampistrou05: Lopez-Campistrous A, Semchuk P, Burke L, Palmer-Stone T, Brokx SJ, Broderick G, Bottorff D, Bolch S, Weiner JH, Ellison MJ (2005). "Localization, annotation, and comparison of the Escherichia coli K-12 proteome under two states of growth." Mol Cell Proteomics 4(8);1205-9. PMID: 15911532

Opacic10: Opacic M, Vos EP, Hesp BH, Broos J (2010). "Localization of the substrate-binding site in the homodimeric mannitol transporter, EIImtl, of Escherichia coli." J Biol Chem 285(33);25324-31. PMID: 20522557

Opa12: Opačić M, Hesp BH, Fusetti F, Dijkstra BW, Broos J (2012). "Structural investigation of the transmembrane C domain of the mannitol permease from Escherichia coli using 5-FTrp fluorescence spectroscopy." Biochim Biophys Acta 1818(3);861-8. PMID: 22100747

Pas87: Pas HH, Ellory JC, Robillard GT (1987). "Bacterial phosphoenolpyruvate-dependent phosphotransferase system: association state of membrane-bound mannitol-specific enzyme II demonstrated by inactivation." Biochemistry 26(21);6689-96. PMID: 3322385

Pas88: Pas HH, Robillard GT (1988). "S-phosphocysteine and phosphohistidine are intermediates in the phosphoenolpyruvate-dependent mannitol transport catalyzed by Escherichia coli EIIMtl." Biochemistry 27(16);5835-9. PMID: 3142516

Pas88a: Pas HH, ten Hoeve-Duurkens RH, Robillard GT (1988). "Bacterial phosphoenolpyruvate-dependent phosphotransferase system: mannitol-specific EII contains two phosphoryl binding sites per monomer and one high-affinity mannitol binding site per dimer." Biochemistry 27(15);5520-5. PMID: 3140890

Postma93: Postma PW, Lengeler JW, Jacobson GR (1993). "Phosphoenolpyruvate:carbohydrate phosphotransferase systems of bacteria." Microbiol Rev 57(3);543-94. PMID: 8246840

Robillard: Robillard GT, Pas HH, Gage D, Elferink MG "The redox state and the phosphorylation state of the mannitol-specific carrier of the E. coli phosphoenolpyruvate-dependent phosphotransferase system." Mol Cell Biochem 82(1-2);113-8. PMID: 3141776

Robillard89: Robillard GT, Pas HH, ten Hoeve-Duurkens RH, Elferink MG (1989). "Molecular details of Escherichia coli EIImtl catalyzed mannitol transport and phosphorylation." FEMS Microbiol Rev 5(1-2);135-42. PMID: 2517397

Roossien84: Roossien FF, Blaauw M, Robillard GT (1984). "Kinetics and subunit interaction of the mannitol-specific enzyme II of the Escherichia coli phosphoenolpyruvate-dependent phosphotransferase system." Biochemistry 23(21);4934-9. PMID: 6437444

Saier14: Saier MH, Reddy VS, Tamang DG, Vastermark A (2014). "The transporter classification database." Nucleic Acids Res 42(1);D251-8. PMID: 24225317

Saier81: Saier MH, Schmidt MR (1981). "Vectorial and nonvectorial transphosphorylation catalyzed by enzymes II of the bacterial phosphotransferase system." J Bacteriol 145(1);391-7. PMID: 6780516

SaraceniRichard97: Saraceni-Richards CA, Jacobson GR (1997). "Subunit and amino acid interactions in the Escherichia coli mannitol permease: a functional complementation study of coexpressed mutant permease proteins." J Bacteriol 179(16);5171-7. PMID: 9260961

SaraceniRichard97a: Saraceni-Richards CA, Jacobson GR (1997). "A conserved glutamate residue, Glu-257, is important for substrate binding and transport by the Escherichia coli mannitol permease." J Bacteriol 179(4);1135-42. PMID: 9023195

Stephan86: Stephan MM, Jacobson GR (1986). "Membrane disposition of the Escherichia coli mannitol permease: identification of membrane-bound and cytoplasmic domains." Biochemistry 25(25);8230-4. PMID: 3545289

Stephan89: Stephan MM, Khandekar SS, Jacobson GR (1989). "Hydrophilic C-terminal domain of the Escherichia coli mannitol permease: phosphorylation, functional independence, and evidence for intersubunit phosphotransfer." Biochemistry 28(19);7941-6. PMID: 2692705

Sugiyama91: Sugiyama JE, Mahmoodian S, Jacobson GR (1991). "Membrane topology analysis of Escherichia coli mannitol permease by using a nested-deletion method to create mtlA-phoA fusions." Proc Natl Acad Sci U S A 1991;88(21);9603-7. PMID: 1946374

Sutrina87: Sutrina SL, Waygood EB, Grenier FC, Saier MH (1987). "HPr/HPr-P phosphoryl exchange reaction catalyzed by the mannitol specific enzyme II of the bacterial phosphotransferase system." J Biol Chem 262(6);2636-41. PMID: 3102473

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

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

UniProt11: UniProt Consortium (2011). "UniProt version 2011-06 released on 2011-06-30 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."

vanWeeghel91: van Weeghel RP, Meyer G, Pas HH, Keck W, Robillard GT (1991). "Cytoplasmic phosphorylating domain of the mannitol-specific transport protein of the phosphoenolpyruvate-dependent phosphotransferase system in Escherichia coli: overexpression, purification, and functional complementation with the mannitol binding domain." Biochemistry 30(39);9478-85. PMID: 1909895

vanWeeghel91a: van Weeghel RP, Meyer GH, Keck W, Robillard GT (1991). "Phosphoenolpyruvate-dependent mannitol phosphotransferase system of Escherichia coli: overexpression, purification, and characterization of the enzymatically active C-terminal domain of enzyme IImtl equivalent to enzyme IIImtl." Biochemistry 30(7);1774-9. PMID: 1993192

Veldhuis05: Veldhuis G, Broos J, Poolman B, Scheek RM (2005). "Stoichiometry and substrate affinity of the mannitol transporter, EnzymeIImtl, from Escherichia coli." Biophys J 89(1);201-10. PMID: 15879478

Vos08a: Vos EP, Ter Horst R, Poolman B, Broos J (2008). "Domain complementation studies reveal residues critical for the activity of the mannitol permease from Escherichia coli." Biochim Biophys Acta. PMID: 19013424

Vos09: Vos EP, Bokhove M, Hesp BH, Broos J (2009). "Structure of the cytoplasmic loop between putative helices II and III of the mannitol permease of Escherichia coli: a tryptophan and 5-fluorotryptophan spectroscopy study." Biochemistry 48(23);5284-90. PMID: 19402710

Weng93: Weng QP, Jacobson GR (1993). "Role of a conserved histidine residue, His-195, in the activities of the Escherichia coli mannitol permease." Biochemistry 32(41);11211-6. PMID: 8218185

White90: White DW, Jacobson GR (1990). "Molecular cloning of the C-terminal domain of Escherichia coli D-mannitol permease: expression, phosphorylation, and complementation with C-terminal permease deletion proteins." J Bacteriol 172(3);1509-15. PMID: 2407724

Yamada91: Yamada Y, Chang YY, Daniels GA, Wu LF, Tomich JM, Yamada M, Saier MH (1991). "Insertion of the mannitol permease into the membrane of Escherichia coli. Possible involvement of an N-terminal amphiphilic sequence." J Biol Chem 266(27);17863-71. PMID: 1917927

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

Other References Related to Gene Regulation

Davis88: Davis T, Yamada M, Elgort M, Saier MH (1988). "Nucleotide sequence of the mannitol (mtl) operon in Escherichia coli." Mol Microbiol 1988;2(3);405-12. PMID: 3135464

GonzalezGil96: Gonzalez-Gil G, Bringmann P, Kahmann R (1996). "FIS is a regulator of metabolism in Escherichia coli." Mol Microbiol 22(1);21-9. PMID: 8899705

Hale94: Hale WB, van der Woude MW, Low DA (1994). "Analysis of nonmethylated GATC sites in the Escherichia coli chromosome and identification of sites that are differentially methylated in response to environmental stimuli." J Bacteriol 1994;176(11);3438-41. PMID: 8195106

MedinaRivera11: Medina-Rivera A, Abreu-Goodger C, Thomas-Chollier M, Salgado H, Collado-Vides J, van Helden J (2011). "Theoretical and empirical quality assessment of transcription factor-binding motifs." Nucleic Acids Res 39(3);808-24. PMID: 20923783

Ramseier95: Ramseier TM, Saier MH (1995). "cAMP-cAMP receptor protein complex: five binding sites in the control region of the Escherichia coli mannitol operon." Microbiology 141 ( Pt 8);1901-7. PMID: 7551052

Ramseier95a: Ramseier TM, Bledig S, Michotey V, Feghali R, Saier MH (1995). "The global regulatory protein FruR modulates the direction of carbon flow in Escherichia coli." Mol Microbiol 1995;16(6);1157-69. PMID: 8577250

Tan09b: Tan K, Borovilos M, Zhou M, Horer S, Clancy S, Moy S, Volkart LL, Sassoon J, Baumann U, Joachimiak A (2009). "The Mannitol Operon Repressor MtlR Belongs to a New Class of Transcription Regulators in Bacteria." J Biol Chem. PMID: 19840941

Webster94: Webster NJ, Kong Y, Cameron KE, Resnik JL (1994). "An upstream element from the human insulin receptor gene promoter contains binding sites for C/EBP beta and NF-1." Diabetes 1994;43(2);305-12. PMID: 8288055


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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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