DB code: S00417

CATH domain 3.40.718.10 : Isopropylmalate Dehydrogenase Catalytic domain
E.C. 1.1.1.85
CSA 1a05
M-CSA 1a05
MACiE

CATH domain Related DB codes (homologues)
3.40.718.10 : Isopropylmalate Dehydrogenase S00416

Uniprot Enzyme Name
UniprotKB Protein name Synonyms Pfam
P12010 3-isopropylmalate dehydrogenase
EC 1.1.1.85
Beta-IPM dehydrogenase
IMDH
3-IPM-DH
PF00180 (Iso_dh)
[Graphical View]
P61495 3-isopropylmalate dehydrogenase
EC 1.1.1.85
Beta-IPM dehydrogenase
IMDH
3-IPM-DH
PF00180 (Iso_dh)
[Graphical View]
Q56268 3-isopropylmalate dehydrogenase
EC 1.1.1.85
Beta-IPM dehydrogenase
IMDH
3-IPM-DH
PF00180 (Iso_dh)
[Graphical View]

KEGG enzyme name
3-isopropylmalate dehydrogenase
beta-isopropylmalic enzyme
beta-isopropylmalate dehydrogenase
threo-Ds-3-isopropylmalate dehydrogenase
3-carboxy-2-hydroxy-4-methylpentanoate:NAD+ oxidoreductase

UniprotKB: Accession Number Entry name Activity Subunit Subcellular location Cofactor
P12010 LEU3_BACCO (2R,3S)-3-isopropylmalate + NAD(+) = 4-methyl- 2-oxopentanoate + CO(2) + NADH. Homodimer. Cytoplasm. Binds 1 magnesium or manganese ion per subunit.
P61495 LEU3_THETH (2R,3S)-3-isopropylmalate + NAD(+) = 4-methyl- 2-oxopentanoate + CO(2) + NADH. Homodimer. Cytoplasm. Binds 1 magnesium or manganese ion per subunit.
Q56268 LEU3_THIFE (2R,3S)-3-isopropylmalate + NAD(+) = 4-methyl- 2-oxopentanoate + CO(2) + NADH. Homodimer. Cytoplasm. Binds 1 magnesium or ion per subunit.

KEGG Pathways
Map code Pathways E.C.
MAP00290 Valine, leucine and isoleucine biosynthesis

Compound table
Cofactors Substrates Products Intermediates
KEGG-id C00305 C04411 C00003 C04236 C00004 C00233 C00011
E.C.
Compound Magnesium 3-Isopropylmalate NAD+ 3-Carboxy-4-methyl-2-oxopentanoate NADH 4-Methyl-2-oxopentanoate CO2
Type divalent metal (Ca2+, Mg2+) carbohydrate,carboxyl group amide group,amine group,nucleotide carbohydrate,carboxyl group amide group,amine group,nucleotide carbohydrate,carboxyl group others
ChEBI 18420
 18420
 		43468
 43468
 		15846
 15846
 		1467
 1467
 		16908
 16908
 		48430
 48430
 		16526
 16526
PubChem 888
 888
 		5462261
 5462261
 		5893
 5893
 		5462259
 5462259
 		439153
 439153
 		70
 70
 		280
 280
1a05A Bound:_MG Bound:IPM Unbound Unbound Unbound Unbound Unbound
1a05B Bound:_MG Bound:IPM Unbound Unbound Unbound Unbound Unbound
1ayqA Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1ayqB Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1dpzA Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1dpzB Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1dr0A Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1dr0B Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1dr8A Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1dr8B Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1g2uA Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1gc8A Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1gc8B Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1gc9A Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1hexA Unbound Unbound Bound:NAD Unbound Unbound Unbound Unbound
1idmA Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1ipdA Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1osiA Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1osiB Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1osiC Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1osiD Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1osjA Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1osjB Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1xaaA Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1xabA Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1xacA Unbound Unbound Unbound Unbound Unbound Unbound Unbound
1xadA Unbound Unbound Unbound Unbound Unbound Unbound Unbound
2ayqA Unbound Unbound Unbound Unbound Unbound Unbound Unbound
2ayqB Unbound Unbound Unbound Unbound Unbound Unbound Unbound

Reference for Active-site residues
resource references E.C.
Literature [5], [12] & [15]

Active-site residues
PDB Catalytic residues Cofactor-binding residues Modified residues Main-chain involved in catalysis Comment
1a05A TYR 140;LYS 190;ASP 222 ASP 222;ASP 246;ASP 250(Magnesium binding)
1a05B TYR 140;LYS 190;ASP 222 ASP 222;ASP 246;ASP 250(Magnesium binding)
1ayqA TYR 141;LYS 190;ASP 222 ASP 222;ASP 246;ASP 250(Magnesium binding)
1ayqB TYR 141;LYS 190;ASP 222 ASP 222;ASP 246;ASP 250(Magnesium binding)
1dpzA TYR 139;LYS 185;ASP 217 ASP 217;ASP 241;ASP 245(Magnesium binding)
1dpzB TYR 139;LYS 185;ASP 217 ASP 217;ASP 241;ASP 245(Magnesium binding)
1dr0A TYR 139;LYS 185;ASP 217 ASP 217;ASP 241;ASP 245(Magnesium binding)
1dr0B TYR 139;LYS 185;ASP 217 ASP 217;ASP 241;ASP 245(Magnesium binding)
1dr8A TYR 139;LYS 185;ASP 217 ASP 217;ASP 241;ASP 245(Magnesium binding)
1dr8B TYR 139;LYS 185;ASP 217 ASP 217;ASP 241;ASP 245(Magnesium binding)
1g2uA TYR 139;LYS 185;ASP 217 ASP 217;ASP 241;ASP 245(Magnesium binding) mutant A172V
1gc8A TYR 1139;LYS 1185;ASP 1217 ASP 1217;ASP 1241;ASP 1245(Magnesium binding) mutant A1172F
1gc8B TYR 139;LYS 185;ASP 217 ASP 217;ASP 241;ASP 245(Magnesium binding) mutant A172F
1gc9A TYR 139;LYS 185;ASP 217 ASP 217;ASP 241;ASP 245(Magnesium binding) mutant A172G
1hexA TYR 139;LYS 185;ASP 217 ASP 217;ASP 241;ASP 245(Magnesium binding)
1idmA TYR 137;LYS 183;ASP 215 ASP 215;ASP 239;ASP 243(Magnesium binding) mutant R82;K83
1ipdA TYR 139;LYS 185;ASP 217 ASP 217;ASP 241;ASP 245(Magnesium binding)
1osiA TYR 139;LYS 185;ASP 217 ASP 217;ASP 241;ASP 245(Magnesium binding)
1osiB TYR 139;LYS 185;ASP 217 ASP 217;ASP 241;ASP 245(Magnesium binding)
1osiC TYR 139;LYS 185;ASP 217 ASP 217;ASP 241;ASP 245(Magnesium binding)
1osiD TYR 139;LYS 185;ASP 217 ASP 217;ASP 241;ASP 245(Magnesium binding)
1osjA TYR 139;LYS 185;ASP 217 ASP 217;ASP 241;ASP 245(Magnesium binding) mutant A172L
1osjB TYR 139;LYS 185;ASP 217 ASP 217;ASP 241;ASP 245(Magnesium binding)
1xaaA TYR 139;LYS 185;ASP 217 ASP 217;ASP 241;ASP 245(Magnesium binding)
1xabA TYR 139;LYS 185;ASP 217 ASP 217;ASP 241;ASP 245(Magnesium binding)
1xacA TYR 139;LYS 185;ASP 217 ASP 217;ASP 241;ASP 245(Magnesium binding) mutant S82R
1xadA TYR 139;LYS 185;ASP 217 ASP 217;ASP 241;ASP 245(Magnesium binding) mutant S82R
2ayqA TYR 141;LYS 190;ASP 222 ASP 222;ASP 246;ASP 250(Magnesium binding)
2ayqB TYR 141;LYS 190;ASP 222 ASP 222;ASP 246;ASP 250(Magnesium binding)

References for Catalytic Mechanism
References Sections No. of steps in catalysis
[5]
Fig.7, p.8675-8676
[7]
[9]
Fig.1
[15]
p.2164-2165
[26]
p.976-980

References
[1]
Resource
Comments
Medline ID
PubMed ID 3069841
Journal J Biochem (Tokyo)
Year 1988
Volume 104
Pages 679-80
Authors Katsube Y, Tanaka N, Takenaka A, Yamada T, Oshima T
Title Crystallization and preliminary X-ray data for 3-isopropylmalate dehydrogenase of Thermus thermophilus.
Related PDB
Related UniProtKB
[2]
Resource
Comments X-ray crystallography
Medline ID
PubMed ID 2277037
Journal J Biochem (Tokyo)
Year 1990
Volume 108
Pages 449-56
Authors Yamada T, Akutsu N, Miyazaki K, Kakinuma K, Yoshida M, Oshima T
Title Purification, catalytic properties, and thermal stability of threo-Ds-3-isopropylmalate dehydrogenase coded by leuB gene from an extreme thermophile, Thermus thermophilus strain HB8.
Related PDB
Related UniProtKB
[3]
Resource
Comments
Medline ID
PubMed ID 1901851
Journal J Biochem (Tokyo)
Year 1991
Volume 109
Pages 1-2
Authors Onodera K, Moriyama H, Takenaka A, Tanaka N, Akutsu N, Muro M, Oshima T, Imada K, Sato M, Katsube Y
Title Crystallization and preliminary X-ray studies of a Bacillus subtilis and Thermus thermophilus HB8 chimeric 3-isopropylmalate dehydrogenase.
Related PDB
Related UniProtKB
[4]
Resource
Comments X-RAY CRYSTALLOGRAPHY (2.2 ANGSTROMS)
Medline ID 92085286
PubMed ID 1748999
Journal J Mol Biol
Year 1991
Volume 222
Pages 725-38
Authors Imada K, Sato M, Tanaka N, Katsube Y, Matsuura Y, Oshima T
Title Three-dimensional structure of a highly thermostable enzyme, 3-isopropylmalate dehydrogenase of Thermus thermophilus at 2.2 A resolution.
Related PDB 1ipd
Related UniProtKB P61495
[5]
Resource
Comments Homologous enzyme
Medline ID
PubMed ID 1888729
Journal Biochemistry
Year 1991
Volume 30
Pages 8671-8
Authors Hurley JH, Dean AM, Koshland DE Jr, Stroud RM
Title Catalytic mechanism of NADP(+)-dependent isocitrate dehydrogenase: implications from the structures of magnesium-isocitrate and NADP+ complexes.
Related PDB
Related UniProtKB
[6]
Resource
Comments
Medline ID
PubMed ID 1400259
Journal J Biochem (Tokyo)
Year 1992
Volume 112
Pages 173-4
Authors Sakurai M, Onodera K, Moriyama H, Matsumoto O, Tanaka N, Numata K, Imada K, Sato M, Katsube Y, Oshima T
Title Crystallization and preliminary X-ray studies of a Bacillus subtilis and Thermus thermophilus HB8 chimeric 3-isopropylmalate dehydrogenase and thermostable mutants of it.
Related PDB
Related UniProtKB
[7]
Resource
Comments
Medline ID
PubMed ID 8405446
Journal FEBS Lett
Year 1993
Volume 332
Pages 37-8
Authors Miyazaki K, Oshima T
Title Tyr-139 in Thermus thermophilus 3-isopropylmalate dehydrogenase is involved in catalytic function.
Related PDB
Related UniProtKB
[8]
Resource
Comments
Medline ID
PubMed ID 8029202
Journal Protein Eng
Year 1994
Volume 7
Pages 453-9
Authors Onodera K, Sakurai M, Moriyama H, Tanaka N, Numata K, Oshima T, Sato M, Katsube Y
Title Three-dimensional structures of chimeric enzymes between Bacillus subtilis and Thermus thermophilus 3-isopropylmalate dehydrogenases.
Related PDB
Related UniProtKB
[9]
Resource
Comments X-ray crystallography
Medline ID
PubMed ID 7881901
Journal Structure
Year 1994
Volume 2
Pages 1007-16
Authors Hurley JH, Dean AM
Title Structure of 3-isopropylmalate dehydrogenase in complex with NAD+: ligand-induced loop closing and mechanism for cofactor specificity.
Related PDB 1hex
Related UniProtKB
[10]
Resource
Comments
Medline ID
PubMed ID 8140100
Journal Protein Eng
Year 1994
Volume 7
Pages 99-102
Authors Miyazaki K, Kadono S, Sakurai M, Moriyama H, Tanaka N, Oshima T
Title Chemical modification and site-directed mutagenesis of Tyr36 of 3-isopropylmalate dehydrogenase from Thermus thermophilus HB8.
Related PDB
Related UniProtKB
[11]
Resource
Comments
Medline ID
PubMed ID 7590327
Journal Gene
Year 1995
Volume 164
Pages 85-7
Authors Kryger G, Wallon G, Lovett ST, Ringe D, Petsko GA
Title Revision of the amino-acid sequence of 3-isopropylmalate dehydrogenase from Salmonella typhimurium by means of X-ray crystallography.
Related PDB
Related UniProtKB
[12]
Resource
Comments
Medline ID
PubMed ID 8576088
Journal J Biochem (Tokyo)
Year 1995
Volume 118
Pages 745-52
Authors Kadono S, Sakurai M, Moriyama H, Sato M, Hayashi Y, Oshima T, Tanaka N
Title Ligand-induced changes in the conformation of 3-isopropylmalate dehydrogenase from Thermus thermophilus.
Related PDB
Related UniProtKB
[13]
Resource
Comments
Medline ID
PubMed ID 7608131
Journal J Biochem (Tokyo)
Year 1995
Volume 117
Pages 408-13
Authors Moriyama H, Onodera K, Sakurai M, Tanaka N, Kirino-Kagawa H, Oshima T, Katsube Y
Title The crystal structures of mutated 3-isopropylmalate dehydrogenase from Thermus thermophilus HB8 and their relationship to the thermostability of the enzyme.
Related PDB
Related UniProtKB
[14]
Resource
Comments
Medline ID
PubMed ID 8637845
Journal Protein Eng
Year 1995
Volume 8
Pages 763-7
Authors Sakurai M, Moriyama H, Onodera K, Kadono S, Numata K, Hayashi Y, Kawaguchi J, Yamagishi A, Oshima T, Tanaka N
Title The crystal structure of thermostable mutants of chimeric 3-isopropylmalate dehydrogenase, 2T2M6T.
Related PDB 1xac 1xad
Related UniProtKB
[15]
Resource
Comments
Medline ID
PubMed ID 8535253
Journal Protein Sci
Year 1995
Volume 4
Pages 2156-67
Authors Dean AM, Dvorak L
Title The role of glutamate 87 in the kinetic mechanism of Thermus thermophilus isopropylmalate dehydrogenase.
Related PDB
Related UniProtKB
[16]
Resource
Comments
Medline ID
PubMed ID 7773180
Journal Protein Sci
Year 1995
Volume 4
Pages 84-92
Authors Zhang T, Koshland DE Jr
Title Modeling substrate binding in Thermus thermophilus isopropylmalate dehydrogenase.
Related PDB
Related UniProtKB
[17]
Resource
Comments X-ray crystallography
Medline ID
PubMed ID
Journal Acta Crystallogr D Biol Crystallogr
Year 1996
Volume 52
Pages 124-8
Authors Sakurai M, Ohzeki M, Miyazaki K, Moriyama H, Sato M, Tanaka N, Oshima T
Title Structure of a loop-deleted variant of 3-isopropylmalate dehydrogenase from Thermus thermophilus: An internal reprieve tolerance mechanism.
Related PDB 1idm
Related UniProtKB
[18]
Resource
Comments
Medline ID
PubMed ID 8875643
Journal Protein Eng
Year 1996
Volume 9
Pages 663-70
Authors Magyar C, Szilagyi A, Zavodszky P
Title Relationship between thermal stability and 3-D structure in a homology model of 3-isopropylmalate dehydrogenase from Escherichia coli.
Related PDB
Related UniProtKB
[19]
Resource
Comments
Medline ID
PubMed ID 15299625
Journal Acta Crystallogr D Biol Crystallogr
Year 1996
Volume 52
Pages 623-30
Authors Nagata C
Title Cryocrystallography of 3-Isopropylmalate Dehydrogenase from Thermus thermophilus and its Chimeric Enzyme.
Related PDB 1xaa 1xab
Related UniProtKB
[20]
Resource
Comments X-RAY CRYSTALLOGRAPHY (3.0 ANGSTROMS)
Medline ID 98158316
PubMed ID 9498551
Journal J Biochem (Tokyo)
Year 1997
Volume 122
Pages 1092-104
Authors Tsuchiya D, Sekiguchi T, Takenaka A
Title Crystal structure of 3-isopropylmalate dehydrogenase from the moderate facultative thermophile, Bacillus coagulans: two strategies for thermostabilization of protein structures.
Related PDB 1ayq 2ayq
Related UniProtKB P12010
[21]
Resource
Comments X-RAY CRYSTALLOGRAPHY (2.27 ANGSTROMS)
Medline ID 97240818
PubMed ID 9086278
Journal J Mol Biol
Year 1997
Volume 266
Pages 1016-31
Authors Wallon G, Kryger G, Lovett ST, Oshima T, Ringe D, Petsko GA
Title Crystal structures of Escherichia coli and Salmonella typhimurium 3-isopropylmalate dehydrogenase and comparison with their thermophilic counterpart from Thermus thermophilus.
Related PDB
Related UniProtKB P61495
[22]
Resource
Comments X-ray crystallography
Medline ID
PubMed ID 9051733
Journal Protein Eng
Year 1997
Volume 10
Pages 45-52
Authors Qu C, Akanuma S, Moriyama H, Tanaka N, Oshima T
Title A mutation at the interface between domains causes rearrangement of domains in 3-isopropylmalate dehydrogenase.
Related PDB 1osi 1osj
Related UniProtKB
[23]
Resource
Comments
Medline ID
PubMed ID 9278279
Journal Protein Eng
Year 1997
Volume 10
Pages 665-72
Authors Wallon G, Lovett ST, Magyar C, Svingor A, Szilagyi A, Zavodszky P, Ringe D, Petsko GA
Title Sequence and homology model of 3-isopropylmalate dehydrogenase from the psychrotrophic bacterium Vibrio sp. I5 suggest reasons for thermal instability.
Related PDB
Related UniProtKB
[24]
Resource
Comments
Medline ID
PubMed ID 9761923
Journal Acta Crystallogr D Biol Crystallogr
Year 1998
Volume 54
Pages 444-5
Authors Suzuki T, Moriyama H, Hirose R, Sakurai M, Tanaka N, Oshima T
Title Crystallization and preliminary X-ray studies on the hyperstable 3-isopropylmalate dehydrogenase from the thermoacidophilic archaeon Sulfolobus sp. strain 7.
Related PDB
Related UniProtKB
[25]
Resource
Comments
Medline ID
PubMed ID 9636162
Journal Proc Natl Acad Sci U S A
Year 1998
Volume 95
Pages 7406-11
Authors Zavodszky P, Kardos J, Svingor, Petsko GA
Title Adjustment of conformational flexibility is a key event in the thermal adaptation of proteins.
Related PDB
Related UniProtKB
[26]
Resource
Comments X-RAY CRYSTALLOGRAPHY (2.0 ANGSTROMS)
Medline ID 98416693
PubMed ID 9739088
Journal Structure
Year 1998
Volume 6
Pages 971-82
Authors Imada K, Inagaki K, Matsunami H, Kawaguchi H, Tanaka H, Tanaka N, Namba K
Title Structure of 3-isopropylmalate dehydrogenase in complex with 3-isopropylmalate at 2.0 A resolution: the role of Glu88 in the unique substrate-recognition mechanism.
Related PDB 1a05
Related UniProtKB Q56268
[27]
Resource
Comments
Medline ID
PubMed ID 10683439
Journal FEBS Lett
Year 2000
Volume 468
Pages 48-52
Authors Nemeth A, Svingor A, Pocsik M, Dobo J, Magyar C, Szilagyi A, Gal P, Zavodszky P
Title Mirror image mutations reveal the significance of an intersubunit ion cluster in the stability of 3-isopropylmalate dehydrogenase.
Related PDB
Related UniProtKB
[28]
Resource
Comments
Medline ID
PubMed ID 10989439
Journal Methods Enzymol
Year 2000
Volume 324
Pages 301-22
Authors Hayashi-Iwasaki Y, Oshima T
Title Purification and characterization of recombinant 3-isopropylmalate dehydrogenases from Thermus thermophilus and other microorganisms.
Related PDB
Related UniProtKB
[29]
Resource
Comments
Medline ID
PubMed ID 10964981
Journal Protein Eng
Year 2000
Volume 13
Pages 527-33
Authors Hori T, Moriyama H, Kawaguchi J, Hayashi-Iwasaki Y, Oshima T, Tanaka N
Title The initial step of the thermal unfolding of 3-isopropylmalate dehydrogenase detected by the temperature-jump Laue method.
Related PDB
Related UniProtKB
[30]
Resource
Comments X-RAY CRYSTALLOGRAPHY (2.2 ANGSTROMS)
Medline ID 20272095
PubMed ID 10810156
Journal Protein Eng
Year 2000
Volume 13
Pages 253-8
Authors Nurachman Z, Akanuma S, Sato T, Oshima T, Tanaka N
Title Crystal structures of 3-isopropylmalate dehydrogenases with mutations at the C-terminus: crystallographic analyses of structure-stability relationships.
Related PDB 1dpz 1dr0 1dr8
Related UniProtKB P61495
[31]
Resource
Comments X-ray crystallography
Medline ID
PubMed ID 11173468
Journal Acta Crystallogr D Biol Crystallogr
Year 2001
Volume 57
Pages 225-32
Authors Qu C, Akanuma S, Tanaka N, Moriyama H, Oshima T
Title Design, X-ray crystallography, molecular modelling and thermal stability studies of mutant enzymes at site 172 of 3-isopropylmalate dehydrogenase from Thermus thermophilus.
Related PDB 1g2u 1gc8 1gc9
Related UniProtKB
[32]
Resource
Comments
Medline ID
PubMed ID 11369782
Journal J Biol Chem
Year 2001
Volume 276
Pages 28121-5
Authors Svingor A, Kardos J, Hajdu I, Nemeth A, Zavodszky P
Title A better enzyme to cope with cold. Comparative flexibility studies on psychrotrophic, mesophilic, and thermophilic IPMDHs.
Related PDB
Related UniProtKB
[33]
Resource
Comments
Medline ID
PubMed ID 11297665
Journal Protein Eng
Year 2001
Volume 14
Pages 81-4
Authors Hirose R, Suzuki T, Moriyama H, Sato T, Yamagishi A, Oshima T, Tanaka N
Title Crystal structures of mutants of Thermus thermophilus IPMDH adapted to low temperatures.
Related PDB
Related UniProtKB
[34]
Resource
Comments
Medline ID
PubMed ID 11297666
Journal Protein Eng
Year 2001
Volume 14
Pages 85-91
Authors Suzuki T, Yasugi M, Arisaka F, Yamagishi A, Oshima T
Title Adaptation of a thermophilic enzyme, 3-isopropylmalate dehydrogenase, to low temperatures.
Related PDB
Related UniProtKB

Comments
This enzyme catalyzes two reactions; firstly the dehydrogenation of 3-carboxy-2-hydroxy-4-methylpentanoate, secondly decarboxylation of the dehydrogenated product.

Created Updated
2004-07-22 2009-02-26