WORKLIST ENTRIES (1):

GSTRNSFRASEM View alignment View Structure    Mu-class glutathione S-transferase signature
 Type of fingerprint: COMPOUND with 4  elements
Links:
   PRINTS; PR01266 GSTRNSFRASEA; PR01268 GSTRNSFRASEP; PR01269 SCRYSTALLIN
   PRINTS; PR01625 GSTRNSFRASEO
   PDB; 1GTU 3Dinfo
   SCOP; 1GSB
   CATH; 1GTU

 Creation date 21-DEC-1999

   1. ALLARDYCE, C.S., MCDONAGH, P.D., LIAN, L-Y., WOLF, R. AND ROBERTS, G.C.K.
   The role of tyrosine-9 and the C-terminal helix in the catalytic mechanism
   of alpha-class glutathione S-transferases.
   BIOCHEM.J. 343 525-531 (1999).

   2. NUCCETELLI, M.N., MAZZETTI, A.P., ROSSJOHN, J., PARKER, M.W., BOARD, P.,
   CACCURI, A.M., FEDERICI, G., RICCI, G. AND LO BELLO, M.
   Shifting substrate specificity of human glutathione transferase (from class
   pi to class alpha) by a single point mutation.
   BIOCHEM.BIOPHYS.RES.COMMUN. 252(1) 184-189 (1998).

   3. DIRR, H., REINEMER, P. AND HUBER, R.
   X-ray crystal structures of cytosolic glutathione S-transferases.
   Implications for protein architecture, substrate recognition and catalytic
   function.
   EUR.J.BIOCHEM. 220 645-661 (1994).

   4. TAKAHASHI, Y., CAMBELL, E.A., HIRATA, Y., TAKAYAMA, T. AND LISTOWSKY, I.
   A basis for differentiating among the multiple human mu-glutathione
   S-transferases and molecular cloning of brain GSTM5.
   J.BIOL.CHEM. 268(12) 8893-8 (1993).

   5. HANSSON, L.O., BOLTON-GROB, R., MASSOUD, T. AND MANNERVIK, B.
   Evolution of differential substrate specificities in mu class glutathione
   transferases probed by DNA shuffling.
   J.MOL.BIOL. 287 265-276 (1999).

   Glutathione S-transferases (GSTs) are a range of dimeric proteins that
   catalyse the conjugation of glutathione to a wide range of hydrophobic
   compounds through the formation of a thioether bond with their
   electrophilic centre. Based on amino acid sequence identity, there are at
   least seven major classes of GST (designated alpha, kappa, mu, pi, sigma, 
   theta and zeta). Pi-, mu-, alpha- and theta-class crystal structures have
   been elucidated; all possess a similar GSH-binding site (G subsite), but 
   the hydrophobic substrate-binding site (H subsite) is subject to variation 
   across the classes [1]. Whilst most of the GSTs share common substrates,
   there are distinct differences in substrate preference between subfamilies.
   Sequence similarity between classes is rather low, ranging between 20-30%.
   However, a single point mutation in the H-subsite region is enough to shift
   substrate specificity from class pi to alpha [2].
  
   These enzymes have evolved as a cellular protection system against a range
   of xenobiotics, oxidative metabolism by-products, and in particular are
   known to metabolise a number of environmental carcinogens. The wide range
   of GST isoforms present in the various subfamilies provides cells with an
   efficient way of scavenging the huge number of potentially toxic compounds
   encountered. Genetic differences in GST expression have been implicated in
   individual susceptibility to certain types of cancer. Conversely, over- 
   expression of GSTs is thought to be involved in the phenomenon of multi-drug
   resistance to cancer chemotherapy.
  
   In spite of relatively low sequence identity, the GSTs exhibit a high degree
   of structural similarity. The structure comprises 2 domains: domain I is the
   smaller of the two and is formed from the N-terminal region of the sequence
   - it possesses an alpha/beta-type core structure comprising a central 
   4-stranded beta-pleated sheet, flanked on one side by two alpha-helices and
   on the other by a single helix; domain II is the larger of the domains and
   occurs towards the C-terminal region of the sequence - it contains a
   predominantly all-alpha-type core comprising 5 amphipathic alpha-helices,
   arranged in a right-handed spiral. The active site is situated near the
   subunit interface. G-subsite molecular recognition is attributable mostly 
   to residues in domain I of one subunit and 1 or 2 residues in domain II of
   the other subunit. Residues contributing to H-subsite specificity are found
   within domains I and II of the same subunit [3].
  
   At present, human mu-class GSTs can be subdivided into 5 isoforms based on
   differing substrate specificities [4]. Mu-class GSTs are thought to be
   involved in the detoxification of reactive oxygen species (cyclised
   o-quinones) produced via oxidative metabolism of catecholamines. These
   toxins are thought to be involved in neurological disorders of the
   nigrostriatal and mesolimbic systems (Parkinsons and Schizophrenia,
   respectively). Indeed, mu-class GSTs are expressed in the substantia nigra
   and have preferential substrate specificity for the cyclised o-quinones
   formed by catecholamine metabolism [5]. Mu-class GSTs possess the so-called
   "mu-loop", which occurs between strand beta-2 and helix alpha-3. This is a
   consequence of an insertion in the primary sequence and the loop allows the
   overall domain I topology to remain [3].  
  
   GSTRNSFRASEM is a 4-element fingerprint that provides a signature for mu-
   class glutathione S-transferases. The fingerprint was derived from an
   initial alignment of 12 sequences: the motifs were drawn from conserved 
   regions spanning virtually the full alignment length - motif 1 includes the
   C-terminal region of beta-strand 2 and the loop between strand 2 and 
   alpha-helix 3; motif 2 spans helix 2 and the following loop; motif 3
   includes the N-terminal region of helix 5; and motif 4 includes the 
   C-terminal region of helix 6 and the following loop. Three iterations on 
   SPTR37_10f were required to reach convergence, at which point a true set 
   comprising 22 sequences was identified. Three partial matches were also 
   found: GTM1_DERPT and O16058 are mu-class glutathione transferases from 
   Dermatophagoides pteronyssinus (house-dust mite) and Echinococcus 
   granulosus, respectively, which match motifs 1 and 2; and Q27653 is 
   an unclassified GST that again matches motifs 1 and 2.

  SUMMARY INFORMATION
     22 codes involving  4 elements
      0 codes involving  3 elements
      3 codes involving  2 elements

   COMPOSITE FINGERPRINT INDEX
  
    4|  22   22   22   22  
    3|   0    0    0    0  
    2|   3    3    0    0  
   --+---------------------
     |   1    2    3    4  

True positives..
 GTM5_HUMAN     GTM3_RAT       GTM1_HUMAN     GTMU_CRILO     
 GTM4_HUMAN     Q05465         GTM2_MOUSE     GTM1_RAT       
 GTM2_RAT       GTM1_MOUSE     GTM3_MOUSE     O35660         
 GTMU_MESAU     GTMU_CAVPO     O60550         GTM2_HUMAN     
 GTM3_HUMAN     GTMU_RABIT     Q9Z1B2         GTM5_MOUSE     
 GTM2_CHICK     O97117         
Subfamily:  Codes involving 2 elements
 Subfamily True positives..
 GTM1_DERPT     Q27653         O16058         


  PROTEIN TITLES
   GTM5_HUMAN       GLUTATHIONE S-TRANSFERASE MU 5 (EC 2.5.1.18) (GSTM5-5) (GST 
   GTM3_RAT         GLUTATHIONE S-TRANSFERASE YB3 (EC 2.5.1.18) (CHAIN 4) (GST C
   GTM1_HUMAN       GLUTATHIONE S-TRANSFERASE MU 1 (EC 2.5.1.18) (GSTM1-1) (HB S
   GTMU_CRILO       GLUTATHIONE S-TRANSFERASE Y1 (EC 2.5.1.18) (CHAIN 3) (GST CL
   GTM4_HUMAN       GLUTATHIONE S-TRANSFERASE MU 4 (EC 2.5.1.18) (GSTM4-4) (GTS-
   Q05465           GLUTATHIONE S-TRANSFERASE (EC 2.5.1.18) (CLASS-MU) - HOMO SA
   GTM2_MOUSE       GLUTATHIONE S-TRANSFERASE 5 (EC 2.5.1.18) (GST 5-5) (GST CLA
   GTM1_RAT         GLUTATHIONE S-TRANSFERASE YB1 (EC 2.5.1.18) (CHAIN 3) (GST M
   GTM2_RAT         GLUTATHIONE S-TRANSFERASE YB2 (EC 2.5.1.18) (CHAIN 4) (GST C
   GTM1_MOUSE       GLUTATHIONE S-TRANSFERASE GT8.7 (EC 2.5.1.18) (GST 1-1) (GST
   GTM3_MOUSE       GLUTATHIONE S-TRANSFERASE GT9.3 (EC 2.5.1.18) (GST CLASS-MU)
   O35660           GLUTATHIONE-S-TRANSFERASE CLASS M5 (EC 2.5.1.18) (GLUTATHION
   GTMU_MESAU       GLUTATHIONE S-TRANSFERASE (EC 2.5.1.18) (GST CLASS-MU) - MES
   GTMU_CAVPO       GLUTATHIONE S-TRANSFERASE B (EC 2.5.1.18) (GST B) (GST CLASS
   O60550           GLUTATHIONE S-TRANSFERASE MU 3 - HOMO SAPIENS (HUMAN).
   GTM2_HUMAN       GLUTATHIONE S-TRANSFERASE MU 2 (EC 2.5.1.18) (GSTM2-2) (GST 
   GTM3_HUMAN       GLUTATHIONE S-TRANSFERASE MU 3 (EC 2.5.1.18) (GSTM3-3) (GST 
   GTMU_RABIT       GLUTATHIONE S-TRANSFERASE MU 1 (EC 2.5.1.18) (GST MU I) (GST
   Q9Z1B2           GLUTATHIONE S-TRANSFERASE M5 (EC 2.5.1.18) - RATTUS NORVEGIC
   GTM5_MOUSE       GLUTATHIONE S-TRANSFERASE MU 5 (EC 2.5.1.18) (GSTM5-5) (GST 
   GTM2_CHICK       GLUTATHIONE S-TRANSFERASE 2 (EC 2.5.1.18) (GST-CL2) (GST CLA
   O97117           GLUTATHIONE S-TRANSFERASE - BOOPHILUS MICROPLUS (CATTLE TICK
 
   GTM1_DERPT       GLUTATHIONE S-TRANSFERASE (EC 2.5.1.18) (GST CLASS-MU) (MAJO
   Q27653           GLUTATHIONE TRANSFERASE (EC 2.5.1.18) (GLUTATHIONE S-ALKYLTR
   O16058           GLUTATHIONE S-TRANSFERASE - ECHINOCOCCUS GRANULOSUS.

SCAN HISTORY SPTR37_10f 3 90 NSINGLE INITIAL MOTIF SETS GSTRNSFRASEM1 Length of motif = 13 Motif number = 1 Mu-class glutathione S-transferase motif I - 1 PCODE ST INT KKYTMGDAPDYDR GTM1_HUMAN 30 30 KRYAMGDAPDYDR GTM1_RAT 30 30 KRYTMGDAPDFDR GTM1_MOUSE 30 30 KKYTMGDAPDYDR GTM2_HUMAN 30 30 KKYSMGDAPDYDR GTM2_RAT 30 30 KKYTMGDAPDYDR GTM2_MOUSE 30 30 KRYTCGEAPDYDR GTM3_HUMAN 35 35 KRYTMGDAPDFDR GTM3_RAT 30 30 KRYVMGDAPNFDR GTM3_MOUSE 30 30 KKYTMGDAPDYDR GTM4_HUMAN 31 31 KKYTMGDAPDYDR GTM5_HUMAN 30 30 KRYICGEAPDYDR GTM5_MOUSE 34 34 GSTRNSFRASEM2 Length of motif = 13 Motif number = 2 Mu-class glutathione S-transferase motif II - 1 PCODE ST INT SQWLNEKFKLGLD GTM1_HUMAN 43 0 SQWLNEKFKLGLD GTM1_RAT 43 0 SQWLNEKFKLGLD GTM1_MOUSE 43 0 SQWLNEKFKLGLD GTM2_HUMAN 43 0 SQWLSEKFKLGLD GTM2_RAT 43 0 SQWLSEKFKLGLD GTM2_MOUSE 43 0 SQWLDVKFKLDLD GTM3_HUMAN 48 0 SQWLNEKFKLGLD GTM3_RAT 43 0 SQWLSEKFNLGLD GTM3_MOUSE 43 0 SQWLNEKFKLGLD GTM4_HUMAN 44 0 SQWLNEKFKLGLD GTM5_HUMAN 43 0 SQWLDVKFKLDLD GTM5_MOUSE 47 0 GSTRNSFRASEM3 Length of motif = 12 Motif number = 3 Mu-class glutathione S-transferase motif III - 1 PCODE ST INT CGETEEEKIRVD GTM1_HUMAN 86 30 CGETEEERIRAD GTM1_RAT 86 30 DGETEEERIRAD GTM1_MOUSE 86 30 CGESEKEQIRED GTM2_HUMAN 86 30 CGETEEERIRVD GTM2_RAT 86 30 CGETEEERIRVD GTM2_MOUSE 86 30 CGETEEEKIRVD GTM3_HUMAN 91 30 CGETEEERIRVD GTM3_RAT 86 30 CGETEEERIRVD GTM3_MOUSE 86 30 CGETEEEKIRVD GTM4_HUMAN 87 30 CGETEEEKIRVD GTM5_HUMAN 86 30 CGDTEEEKIRVD GTM5_MOUSE 90 30 GSTRNSFRASEM4 Length of motif = 14 Motif number = 4 Mu-class glutathione S-transferase motif IV - 1 PCODE ST INT SEFLGKRPWFAGNK GTM1_HUMAN 138 40 SEFLGKRPWFAGDK GTM1_RAT 138 40 SEFLGKRPWFAGDK GTM1_MOUSE 138 40 SQFLGKQPWFLGDK GTM2_HUMAN 138 40 SEFLGKQPWFAGNK GTM2_RAT 138 40 SEFLGKQPWFAGNK GTM2_MOUSE 138 40 SMFLWKFSWFAGEK GTM3_HUMAN 143 40 SEFLGKRPWFAGDK GTM3_RAT 138 40 SEFLGKRPWFAGDK GTM3_MOUSE 138 40 SQFLGKRPWFVGDK GTM4_HUMAN 139 40 SEFLGKRPWFAGDK GTM5_HUMAN 138 40 SLFLGKFTWFAGEK GTM5_MOUSE 142 40 FINAL MOTIF SETS GSTRNSFRASEM1 Length of motif = 13 Motif number = 1 Mu-class glutathione S-transferase motif I - 3 PCODE ST INT KKYTMGDAPDYDR GTM5_HUMAN 30 30 KRYTMGDAPDFDR GTM3_RAT 30 30 KKYTMGDAPDYDR GTM1_HUMAN 30 30 KKYTMGDAPDSDR GTMU_CRILO 30 30 KKYTMGDAPDYDR GTM4_HUMAN 31 31 KKYTMGDAPDYDR Q05465 31 31 KKYTMGDAPDYDR GTM2_MOUSE 30 30 KRYAMGDAPDYDR GTM1_RAT 30 30 KKYSMGDAPDYDR GTM2_RAT 30 30 KRYTMGDAPDFDR GTM1_MOUSE 30 30 KRYVMGDAPNFDR GTM3_MOUSE 30 30 RRYAMGDAPDYDR O35660 31 31 KKYTMGDAPNFDR GTMU_MESAU 30 30 KRYNMGDAPDYDR GTMU_CAVPO 30 30 KRYTCGEAPDYDR O60550 35 35 KKYTMGDAPDYDR GTM2_HUMAN 30 30 KRYTCGEAPDYDR GTM3_HUMAN 35 35 KKYTMGDAPNYDQ GTMU_RABIT 30 30 KQYTCGEAPDYDR Q9Z1B2 35 35 KRYICGEAPDYDR GTM5_MOUSE 34 34 RRYKAGPAPDFDP GTM2_CHICK 30 30 KRYTCGPPPDFDR O97117 31 31 GSTRNSFRASEM2 Length of motif = 13 Motif number = 2 Mu-class glutathione S-transferase motif II - 3 PCODE ST INT SQWLNEKFKLGLD GTM5_HUMAN 43 0 SQWLNEKFKLGLD GTM3_RAT 43 0 SQWLNEKFKLGLD GTM1_HUMAN 43 0 SQWLNEKFKLGLD GTMU_CRILO 43 0 SQWLNEKFKLGLD GTM4_HUMAN 44 0 SQWLNEKFKLGLD Q05465 44 0 SQWLSEKFKLGLD GTM2_MOUSE 43 0 SQWLNEKFKLGLD GTM1_RAT 43 0 SQWLSEKFKLGLD GTM2_RAT 43 0 SQWLNEKFKLGLD GTM1_MOUSE 43 0 SQWLSEKFNLGLD GTM3_MOUSE 43 0 SQWLNDKFKLXLD O35660 44 0 SQWLNEKFKLGLD GTMU_MESAU 43 0 SQWLNEKFKLGLD GTMU_CAVPO 43 0 SQWLDVKFKLDLD O60550 48 0 SQWLNEKFKLGLD GTM2_HUMAN 43 0 SQWLDVKFKLDLD GTM3_HUMAN 48 0 SKWLSEKFTLGLD GTMU_RABIT 43 0 SQWLDVKFKLDLD Q9Z1B2 48 0 SQWLDVKFKLDLD GTM5_MOUSE 47 0 SDWTNEKEKLGLD GTM2_CHICK 43 0 SSWLNEKTKLGLE O97117 44 0 GSTRNSFRASEM3 Length of motif = 12 Motif number = 3 Mu-class glutathione S-transferase motif III - 3 PCODE ST INT CGETEEEKIRVD GTM5_HUMAN 86 30 CGETEEERIRVD GTM3_RAT 86 30 CGETEEEKIRVD GTM1_HUMAN 86 30 CGETEEERIRVD GTMU_CRILO 86 30 CGETEEEKIRVD GTM4_HUMAN 87 30 CGETEEEKIRVD Q05465 87 30 CGETEEERIRVD GTM2_MOUSE 86 30 CGETEEERIRAD GTM1_RAT 86 30 CGETEEERIRVD GTM2_RAT 86 30 DGETEEERIRAD GTM1_MOUSE 86 30 CGETEEERIRVD GTM3_MOUSE 86 30 CGETEEERIRVD O35660 87 30 CGETEEERIQLD GTMU_MESAU 86 30 CGVTEEETIRMD GTMU_CAVPO 86 30 CGETEEEKIRVD O60550 91 30 CGESEKEQIRED GTM2_HUMAN 86 30 CGETEEEKIRVD GTM3_HUMAN 91 30 CGETEEERIRVD GTMU_RABIT 86 30 CGDTEEEKIRVD Q9Z1B2 91 30 CGDTEEEKIRVD GTM5_MOUSE 90 30 CGETEVEKQRVD GTM2_CHICK 86 30 EGKTEAEKQRVD O97117 87 30 GSTRNSFRASEM4 Length of motif = 14 Motif number = 4 Mu-class glutathione S-transferase motif IV - 3 PCODE ST INT SEFLGKRPWFAGDK GTM5_HUMAN 138 40 SEFLGKRPWFAGDK GTM3_RAT 138 40 SEFLGKRPWFAGNK GTM1_HUMAN 138 40 SEFLGKRPWFAGDK GTMU_CRILO 138 40 SQFLGKRPWFVGDK GTM4_HUMAN 139 40 SQFLGKRPWFVGDK Q05465 139 40 SEFLGKQPWFAGNK GTM2_MOUSE 138 40 SEFLGKRPWFAGDK GTM1_RAT 138 40 SEFLGKQPWFAGNK GTM2_RAT 138 40 SEFLGKRPWFAGDK GTM1_MOUSE 138 40 SEFLGKRPWFAGDK GTM3_MOUSE 138 40 SEFLGKQPWFAGDK O35660 139 40 SEFLGKRSWFAGDK GTMU_MESAU 138 40 SQFLGKLPWFAGNK GTMU_CAVPO 138 40 SMFLGKFSWFAGEK O60550 143 40 SQFLGKQPWFLGDK GTM2_HUMAN 138 40 SMFLWKFSWFAGEK GTM3_HUMAN 143 40 SQFLGSLPWFAGDK GTMU_RABIT 138 40 SLFLGKFTWFAGEK Q9Z1B2 143 40 SLFLGKFTWFAGEK GTM5_MOUSE 142 40 SRFLGSRSWFVGDK GTM2_CHICK 138 40 SDYLGTHKFFAGDN O97117 139 40

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