hierarchical classification of glycoside hydrolases daniil g. … · 2007-07-06 · hierarchical...

Hierarchical Classification of Glycoside HydrolasesDaniil G. Naumoff

Laboratory of BioinformaticsState Institute for Genetics and Selection of Industrial Microorganisms

Moscow, Russiahttp://bioinform.genetika.ru/members/Naumoff/index.htm

Gos NII Genetika

Moscow, Russia

Иерархическая классификация гликозил- гидролаз

Даниил Геннадиевич НаумовЛаборатория биоинформатики

Государственный Научно-Исследовательский Институт Генетики и СелекцииПромышленных Микроорганизмов, Москва

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Enzyme Classification (IUBMB)EC 1 – OxidoreductasesEC 2 – Transferases

EC 2.4 – Glycosyltransferases (including transglycosidases)EC 2.4.1 – Hexosyltransferases

EC 2.4.1.10 – Levansucrase (β-fructosyltransferase)EC 2.4.1.67 – Galactinol-raffinose galactosyltransferase

EC 2.4.2 – PentosyltransferasesEC 3 – Hydrolases

EC 3.2 – Glycosylases (glycoside hydrolases)EC 3.2.1 – O- and S-Glycosylases

EC 3.2.1.20 – Maltase (a-glucosidase)EC 3.2.1.22 – Melibiase (a-galactosidase)EC 3.2.1.26 – Sucrase (invertase, β-fructofuranosidase)

EC 3.2.2 – N-GlycosylasesEC 4 – LyasesEC 5 – IsomerasesEC 6 – Ligases

Glycoside Hydrolases = EC 3.2.1.X; X=1…>200

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Progress in the Sequence Based Classification ofGlycoside Hydrolases and Their Homologues

CAZy = the Carbohydrate-Active enZYmes server (http://www.cazy.org/CAZY)

Number ofproteins

Number of families Reference

300 36 Henrissat. Biochem. J. 1991, 280:309-316

480 45 Henrissat & Bairoch. Biochem. J. 1993, 293:781-788

950 57 (17 => 5 clans) Henrissat & Bairoch. Biochem. J. 1996, 316:695-696

… 62 (19 => 5 clans) Henrissat & Davies. Curr. Opin. Struct. Biol. 1997, 7:637-644

2.200 70 (…) Coutinho & Henrissat. CAZy server. 1998

>20.000 101 (46 => 14 clans) Coutinho & Henrissat. CAZy server. 2005

~30.000 106 (46 => 14 clans) Coutinho & Henrissat. CAZy server. 2006

>30.000 110 (46 => 14 clans) Coutinho & Henrissat. CAZy server. 2007

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Sequence Based Classification of Glycoside Hydrolasesthe Carbohydrate-Active Enzymes (CAZy) server (www.cazy.org/CAZY/)

• 30.000 protein sequences => 105 families (GH1-GH110, except GH21, GH40, GH41, GH60, and GH69)• 46 glycoside hydrolase families => 14 clans (GH-A–GH-N)

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(β)3-solenoidinversion (axial orientation)28, 49GH-N(a/a)6inversion (equatorial orientation)8, 48GH-M(a/a)6inversion (axial orientation)15, 65GH-L(β/a)8 -barrelretention (equatorial orientation)18, 20GH-K5-fold β-propellerretention (β-furanoside)32, 68GH-Ja+βinversion (equatorial orientation)24, 46, 80GH-I(β/a)8 -barrelretention (axial orientation)13, 70, 77GH-HNot knowninversion (axial orientation)37, 63GH-G5-fold β-propellerinversion (equatorial orientation)43, 62GH-F6-fold β-propellerretention (equatorial orientation)33, 34, 83GH-E(β/a)8 -barrelretention (axial orientation)27, 36GH-Dβ-jelly rollretention (equatorial orientation)11, 12GH-Cβ-jelly rollretention (equatorial orientation)7, 16GH-B

(β/a)8 -barrelretention (equatorial orientation)1, 2, 5, 10, 17, 26, 30, 35, 39,42, 50, 51, 53, 59, 72, 79, 86

GH-A

Tertiary StructureOptical ConfigurationFamilies (GH)Clan

Clans of Glycoside HydrolasesClic

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Common name: α-galactosidase

Reaction: Hydrolysis of terminal, non-reducing α-D-galactose residues in α-D-galactosides,including galactose oligosaccharides, galactomannans, and galactolipids

Molecular mechanism of hydrolyzing reaction: double displacement with overall retention ofthe anomeric configuration of the axial glycosidic bond

Other names: melibiase; α-D-galactosidase; α-galactosidase A; α-D-galactosidegalactohydrolase; α-galactoside galactohydrolase

Systematic name: α-D-galactoside galactohydrolase

Comments: Also hydrolyses α-D-fucosides

EC 3.2.1.22

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GH4 – α-galactosidase from Escherichia coli and bifunctional enzymes with α-galactosidase andα-glucosidase (EC 3.2.1.20) activities from Thermotoga maritime and Thermotoga neapolitana

+ 6-phospho-α-glucosidases (EC 3.2.1.122) from Eubacteria+ 6-phospho-β-glucosidases (EC 3.2.1.86) from Eubacteria

GH27 – mainly α-galactosidases from Eukaryota (animals, plants, fungi, etc.) and some Eubacteria+ α-N-acetylgalactosaminidases (EC 3.2.1.49) from Eukaryota (animals and fungi)+ isomalto-dextranase (EC 3.2.1.94) from Arthrobacter globiformis

GH36 – mainly α-galactosidases from Eubacteria and Eukaryota (fungi and plants)+ α-galactosyltransferases (EC 2.4.1.67 and EC 2.4.1.82) from plants+ α-N-acetylgalactosaminidase from Clostridium perfringens+ ORFs from Sulfolobus solfataricus and Sulfolobus tokodaii

GH57 – α-galactosidases from Pyrococcus furiosus and Thermococcus alcaliphilus+ α-amylases (EC 3.2.1.1), amylopullulanase (EC 3.2.1.41), and

4-α-glucanotransferases (EC 2.4.1.x) from Eubacteria and ArchaeaGH110 – α-galactosidase from Bacteroides fragilis

Glycoside hydrolase families, which include α-galactosidases

clanGH-D

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Subfamily 27a:

– plant α-galactosidases: Arabidopsis, Carica, Coffea, Cyamopsis, Glycine, Helianthus, Hordeum, Lycopersicon, Oryza, Petunia,Phaseolus, and Senna

– α-galactosyltransferase from Ajuga reptans

– animal α-galactosidases and α-N-acetylgalactosaminidases: Anopheles, Apis, Ateles, Brugia, Caenorhabditis, Ciona, Danio,Drosophila, Gallus, Homo, Mus, Rattus, Takifugu, and Tetraodon

– fungal and yeast α-galactosidases: Aspergillus, Ganoderma, Gibberella, Magnaporthe, Mortierella, Penicillium, Phanerochaete,Saccharomyces, Schizosaccharomyces, Thermomyces, Torulaspora, Trichoderma, Ustilago, and Zygosaccharomyces

– α-N-acetylgalactosaminidase from Acremonium sp.

– ORFs from Dictyostelium discoideum and Toxoplasma gondii

– bacterial α-galactosidases: Cellvibrio mixtus, Clostridium josui, Pseudomonas fluorescens, Saccharopolyspora erythraea, andStreptomyces coelicolor

– bacterial ORFs: Bacteroides, Fibrobacter, Microbulbifer, Porphyromonas, and Streptomyces

Family GH27

Subfamily 27b:

– plant ORFs: Arabidopsis thaliana and Oryza sativa

– bacterial ORFs: Bacillus halodurans, Bifidobacterium longum, Kineococcus radiotolerans, and Ruminococcus albus

Subfamily 27c:

– α-galactosidase from Trichoderma reesei

– ORF from Aspergillus nidulans

Unclassified proteins

– isomalto-dextranase from Arthrobacter globiformis

– bacterial ORFs: Bacteroides thetaiotaomicron and Streptomyces avermitilis

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Tertiary Structure of GH27 Glycosidases(Trichoderma reesei α-galactosidase)

C- terminal domain(β-sandwich fold,Greek key motif)

N-terminal catalytic(β/α)8-domain(TIM barrel-type structure)

GH27N

GH27C

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GH27N GH27C

GH27N

GH27N GH27C CBM13

GH27N GH27C CBM6

GH27N GH27C CBM6 CBM13

GH27N CBM13 GH27C

NEW1 GH27N CBM13 GH27C

NEW1 GH27N GH27C

NEW2 NEW1 GH27N GH27C

GH27N GH27C NEW3 NEW2

GH27N GH27C NEW3

GH27N GH27C Dockerin

GH27N GH27C CBM1 CE1N-terminal domain of GH27 family

C -terminal domain of GH27 family

CE1 domain of carbohydrate esterases

Carbohydrate-binding module CBM1



Dockerin I domain

Uncharacterized domain

Uncharacterized domain (NPCBM)

Uncharacterized domain

CBM13

CBM6

Dockerin

NEW1

NEW2

NEW3

CBM1

CE1

GH27C

GH27N

Domain structure of proteins of the GH27 familyaccording to Naumoff D.G. Phylogenetic analysis of α-galactosidases of the GH27 family. Molecular Biology (Engl Transl), 2004, 38(3):388-399.

PDF: http://bioinform.genetika.ru/members/Naumoff/MB2004E.pdf

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Family GH36

Subfamily GH36A:– bacterial α-galactosidases: Azotobacter, Bacillus, Geobacillus, Bifidobacterium, Carnobacterium, Clostridium, Enterococcus,

Erwinia, Escherichia, Geobacillus, Klebsiella, Lactobacillus, Lactococcus, Leuconostoc, Novosphingobium, Oenococcus,

Pediococcus, Ruminococcus, Streptococcus, Streptomyces, Thermoanaerobacter, Treponema, Vibrio, and Yersinia

– fungal α-galactosidases: Absidia, Aspergillus, Gibberella, Penicillium, and Trichoderma

Subfamily GH36B:– bacterial α-galactosidases: Burkholderia, Leptospira, Streptomyces, Thermotoga, Thermus, and Vibrio

Subfamily GH36C:– plant alkaline α-galactosidases or seed imbibition proteins: Arabidopsis, Brassica, Cicer, Cucumis, Hordeum, Lycopersicon,

Malus, Oryza, and Persea

– plant α-galactosyltransferases: Alonsoa, Arabidopsis, Cucumis, Medicago, Oryza, Pisum, Stachys, and Vigna

– fungal ORFs: Aspergillus, Coccidioides, Cryptococcus, Gibberella, Magnaporthe, Neurospora, and Ustilago

– ORF from Toxoplasma gondii

– α-galactosidase from Bifidobacterium breve

– bacterial ORFs: Bifidobacterium longum, Bacteroides fragilis, and Bacteroides thetaiotaomicron

– ORFs from Sulfolobus solfataricus and Sulfolobus tokodaii

Subfamily GH36D:– α-N-acetylgalactosaminidase from Clostridium perfringens

– bacterial ORFs: Aeromonas, Clostridium, Escherichia, Photorhabdus, and Streptococcus

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MEL1_YEAST 65 GYKYIILDDCWS 141 NRVDYLKYDNCY 204 WRMSGDV 258 WNDLDNLAGAL_PHAVU 106 GYQYINIDDCWG 184 WGIDYLKYDNCE 242 WRTTGDI 274 WNDPDMLNAGA_HUMAN 71 GYTYLNIDDCWI 148 WKVDMLKLDGCF 212 WRNYDDI 248 WNDPDMLAGL1_BIFLO 48 GWDTLVIDIDWY 178 WGLDFLKVDDMQ 240 WRISDDL 272 WADADMVAGL3_HYPJE 234 GYDLCSLDSGWQ 339 WGVDMLKLDFLT 407 MRTDQDL 451 YPDMDALIMD_ARTGO 100 GYDIACTD-GWI 219 LGVPYLRIDFLS 290 VRINADA 338 ILDGDFM

AGAL_LACPL 367 GIEMFVLDDGWF 478 VPIDYIKWDMNR 551 QSWPSDN 584 GTSPDELAGL2_HYPJE 375 GIKLFVLDDGWF 496 ASISYVKWDNNR 561 HIWTSDD 594 SAVPNGQAGL7_ASPFU 413 GAGYFVIDAGWY 524 YGVGYFKFDYNI 595 LQSSSDQ 626 WAYPQPA

AGAL_THEMA 213 PFEVFQIDDAYE 319 MGYRYFKIDFLF 382 MRIGPDT 423 LNDPDCLAGAL_VIBPA 227 DLEWVLLDDGYQ 341 WGVELFKLDANY 404 MRVSDDV 436 QIDPDCAAGL3_STRCO 329 GLKWAVLDDGWQ 438 WGYEGLKIDGQH 512 QYPSSDP 540 SYSGDHV

AGAL_SULSO 256 RLNWVIIDDGWQ 360 RDFDLVKVDNQW 421 MRNSIDY 454 YPDYDMFAGAL_BIFBR 339 PVSWVLIDDGWS 458 AGVDFVKVDSQS 523 TRTSDDF 556 HCDWDMFGALT_VIGAN 251 APRFVVIDDGWQ 476 TGVTGVKIDVIH 544 GRVGDDF 585 QPDWDMF

NAGA_CLOPE 253 TLDAFVVDDGWA 355 YDISYWKIDGML 429 IQTSQDV ...ORF1_CLOPE 515 PIDSYVVDDGWH 634 FDIDYWKLDGFA 715 IQNSQDT ...ORF1_ECOLI 297 ALDAFLLDDGWD 393 EHITSFKLDGMG 458 WRQGDDI ...

ORF2_CLOPE 112 PKGIIMIDDGWS 220 YGVDGFKFDAGD 286 HSWEYNG 344 ALMPMMQXYLQ_LACPE 297 PLDVFHFDCFWQ 406 MGVDSFKTDFGE 474 IQYTGAA 534 LLSSHSRXYLS_SULSO 250 PLDVIVLDWRYW 345 LGIDAYWLDASE 423 ISWSGDV 483 TFCPILRLYAG_HUMAN 397 PLDVQWNDLDYM 510 VPFDGMWIDMNE 611 GHWTGDV 666 AFYPFMR

GH27

GH36A

GH36B

GH36C

GH36D

GH31

Four conserved regions in proteins of the α-galactosidase superfamily, containing Asp residues

Nucleophile Donor H+

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The neighbor-joining phylogenetic tree of the α-galactosidase superfamily

GH31

XYLS SULSOAGL2 BACTQ

AGLU ACIACSUIS HUMANc

SUIS HUMANnLYAG HUMAN

5572

3859

89

XYLQ LACPEORF1 THEMA

ORF1 BACHAYICI ECOLIORF1 CLOAC

4270

4077

86

ORF1 CHLAUORF2 CLOPE

ORF1 MOUSEORF1 DROME

8036

43

69

ORF1 AERHYORF1 ECOLI

93

NAGA CLOPEORF1 STRPNORF1 CLOPE

3992

98

25

AGL3 STRCOAGL2 STRCO

AGAL THETHAGAL THET2 77

AGAL THEMAAGAL LEPIN 72

3757

24

AGAL VIBCHAGAL VIBPA 99

20

30

AGA2 PEDPEAGA1 PEDPE

AGAL LACPLAGAL STRMUAGL2 RUMAL 40

39

AGL5 BACFRAGL6 BACFR

94

54

49

AGL6 ASPFUAGLC ASPNGAGL2 HYPJE

6979

21

AGAL ABSCOAGL2 BIFLORAFA ECOLI 51

31

39

86

AGL3 RUMALAGL7 ASPFU

99

65

AGAL PORGI

MEL2 ARATHAGAL CYATEAGAL PHAVU

10061

AGAL SACERAGAL PSEFLAGAL MICDE

9716

9

AGL1 STRCOAGL2 ASPFU

AGAL FIBSUAGAL CLOJO

6716

5

AGLB ASPNGMEL1 YEASTMELA PHACH

2733

6

AGLA ASPNGNAGA ACRSP

98

MEL1 CAEELMEL1 DROME

NAGA HUMANAGAL HUMAN56

8046

48

AGL3 BACFRAGL2 BACFRAGL1 BACFR

10012

7

21

AGL3 HYPJEIMD ARTGO

MEL4 ARATHMEL5 ORYSA

94

AGL1 BIFLOAGAL BACHAAGL1 RUMAL

6247

49

3972

84

AGAL SULTOAGAL SULSO93

AGL4 BACFRAGAL BIFBR76

ORF2 ARATHSTAS PISSAGALT VIGAN67

45

ORF1 ARATHSIP CICAR

SIP HORVU5358

94

36

89

45

57

GH27

GH36C

GH36A

GH36B

GH36D

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AGAL SULSOAGL5 BACTH

AGAL BIFBRORF1 NEUCR

STAS STAAFAGA1 CUCME713

623670

519995

AGL4 STRAVIMD ARTGO

AGL8 ASPNIAGL3 HYPJE983

879532

MEL4 ARATHMEL5 ORYSA999

AGL1 BIFLOAGAL BACHAAGL1 RUMAL862

905903

410

AGL3 STRAVAGL3 BACFR

AGL4 BACTHAGL3 BACTH

996988

AGL5 BACTHNAGA ACRSP

AGL4 ASPFUAGLA ASPNG

999993

AGL2 DICDIAGAL HUMAN

MEL1 CAEELNAGA HUMANMEL1 DROME

794479996

525

616

AGL1 STRAVAGL2 STRAVAGL1 STRCO1000

906

AGAL MICDEAGAL PSEFL998

AGAL SACERAGAL FIBSUAGAL CLOJO376

479378

124

AGL1 DICDIMEL6 ORYSA

MEL2 ORYSAMEL1 ORYSA767

990265

AGAL PORGIAGL1 BACFR

AGL1 BACTHAGL2 BACFR973

1000580

AGL2 BACTHAGL2 ASPNIAGL2 ASPFU1000

AGL1 USTMAAGL2 GIBZE

AGL3 ASPFUAGL3 ASPNI1000

885689

827

MELA PHACHMEL1 UMBVI

MEL1 SCHPOAGL1 ASPFU

MEL5 YEASTMEL1 SACKL1000

515250

386975

MEL2 UMBVIAGL1 GIBZE

AGL1 HYPJEAGL1 PENPU

AGL5 ASPNIAGL1 PENSI

814554

987866

994374

421

342

466

143

76

103

57

265

372

721

513

AGLB ASPNG

27b27c

27a

1000

AGL5 ASPFU

GH36CPhylogenetic tree of

GH27 family

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Phylogenetic tree ofGH27 family

27cIMD ARTGOAGL8 ASPNIAGL3 HYPJE1000

AGL4 STRAV MEL5 ORYSAMEL4 ARATH999AGL1 RUMAL

AGL1 BIFLOAGL1 BACHA555953

944

AGL3 STRAVAGL3 BACFR

AGL3 BACTHAGL4 BACTH1000991

AGL5 BACTHNAGA ACRSP

AGL4 ASPFUAGLA ASPNG999985

AGL2 DICDIAGAL DANRE

AGAL TAKRUAGAL MOUSEAGAL HUMAN10006681000

NAGA CIOINNAGA DANRENAGA TAKRU976NAGA GALGANAGA HUMAN

NAGA RATNAGA MOUSE10001000923

1000809

MEL1 BRUMAMEL1 CAEBRMEL1 CAEEL1000981

MEL2 DROMEMEL2 ANOGA782

MEL1 ANOGAMEL1 DROME1000

913839

802

999

613

552

AGAL CLOJOAGAL FIBSU220

AGAL PSEFLAGAL MICDE1000

AGAL SACERAGL1 STRAV

AGL1 STRCOAGL2 STRAV1000874

42480

51

AGL1 DICDI MEL2 ARATHMEL6 ORYSAMEL4 ORYSA10001000

MEL3 ORYSAMEL2 ORYSA

AGAL CARPAMEL3 ARATH1000897746

MEL1 ORYSAAGAL HELAN

MEL1 ARATHAGAL LYCESAGAL COFAR399

AGAL SENOCAGAL CYATE

AGAL GLYMAAGAL PHAVU1000597

977233

373732988

579

999

404

AGL2 BACTHAGAL PORGI

AGL1 BACFRAGL1 BACTHAGL2 BACFR8311000514334

AGL2 ASPNIAGL2 ASPFU1000

AGL1 USTMAAGL2 GIBZE

AGL3 ASPFUAGL3 ASPNI1000955

663578

AGL1 GIBZEAGL1 HYPJE

AGL1 PENPUAGL1 PENSI

AGL5 ASPNIAGL5 ASPFUAGLB ASPNG763529

997939

1000727

MEL2 UMBVIMELA PHACH

MEL1 UMBVIMEL1 SCHPO

AGL1 ASPFUMEL4 SACKLMEL1 SACKL1000

MELC YEASTMELR YEAST

MELT YEASTMELP YEASTMEL5 YEAST782MELJ YEAST

MELD YEASTMELX YEAST958842

998630

978696

1000958

556388

950

474

508

170

250

343

352

919

397

6441000

50

27b

I

II

III

IV

27a

468

630

1000

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Families of the α-galactosidase superfamily and family GH97Family GH27 GH31 GH36A GH36B GH36C GH36D GH97Clan GH-D None GH-D GH-D GH-D GH-D None

COG1501KOG1065

EC 2.4.1.x EC 2.4.1.x EC 2.4.1.67EC 3.2.1.22 EC 3.2.1.3 EC 2.4.1.82EC 3.2.1.49 EC 3.2.1.10 EC 3.2.1.22EC 3.2.1.94 EC 3.2.1.20

EC 3.2.1.48EC 4.2.2.13

Molecularmechanism

Retaining Retaining Retaining Not known Not known Not known Not known

Eukaryota: Eukaryota: Eukaryota: Eubacteria: Eukaryota: Eubacteria: Eukaryota:Alveolata Alveolata Fungi Acidobacteria Alveolata Firmicutes Metazoa (?)

FungiEntamoebidae

Eubacteria:

Proteobacteria

Fungi Proteobacteria Eubacteria:

MetazoaEuglenozoa

Actinobacteria

Spirochaetes

Viridiplantae Acidobacteria

MycetozoaFungi

Bacteroidetes

Thermotogales

Eubacteria:BacteroidetesViridiplantae

MetazoaFirmicutes

Thermus

Actinobacteria

PlanctomycetesEubacteria:

Mycetozoa

Proteobacteria Bacteroidetes

ProteobacteriaAcidobacteria

Rhodophyta

Spirochaetes Archaea:

Archaea:Actinobacteria

Viridiplantae

Crenarchaeota

EuryarchaeotaBacteroidetes

Eubacteria:

FibrobacteresActinobacteria

FirmicutesBacteroidetes

ProteobacteriaCyanobacteriaFirmicutesProteobacteriaSpirochaetesThermotogales

Archaea:CrenarchaeotaEuryarchaeota

COG3345 COG3345 None

Origin

KOG2366 None None

Knownenzymaticactivities

EC 3.2.1.22 EC 3.2.1.22 EC 3.2.1.49 EC 3.2.1.20

COG/KOG

Firmicutes

Actinobacteria

ActinobacteriaDeinococcus

Acidobacteria

Thermus

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(β/α)8-barrel-type retaining α-D-glycopyranosidase families

GH66?

α-glucosidasesuperfamily

clan GH-H

GH13

GH70

GH77

COG0296

COG0366

COG1523

COG3280

COG1649

α-galactosidasesuperfamily

clan GH-D

GH27

GH36

GH36A

GH36B

GH36C

GH36D

GH31

family II

family III

GH38

GH57?

GH97

GH27a

GH27b

GH27c

GH97c

GH97d

GH97e

GH97a

GH97b ?

?

?

??

?

– [EC 3.2.1.22]

– [EC 3.2.1.49]

– [EC 3.2.1.20]

– ORF?

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Rigden DJ. Iterative database searches demonstrate that glycoside hydrolase families 27, 31, 36,and 66 share a common evolutionary origin with family 13. FEBS Lett. 2002, 523(1-3):17-22.

clans

GH-D

GH-H

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Nagano N, Porter CT, Thornton JM. The (β/α)8 glycosidases: sequence and structureanalyses suggest distant evolutionary relationships. Protein Eng. 2001, 14(11):845-855.

clans: GH-H GH-A GH-K?

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GH14 GH44 GH71

GH17 GH26 GH30 GH35 GH39

GH42 GH50 GH51 GH53 GH59 GH72 GH79 GH86

clan GH-AGH29

type I

GH1 GH2 GH5 GH10

GH67 GH89

type IV

GH13GH36 GH70GH31

COG1649

GH27

GH36A GH36B GH36C GH36D

GH77

clan GH-D clan GH-H

type II

type III

GH66 GH97

GH84

GH20

GH85GH25

GH18

clan GH-K

(β/α)8-barrel fold

GH56GH3

α-galactosidasesuperfamily

GH6 GH38 GH57

(β/α)7-barrel fold

TIM-barrelfold type

A hierarchical classification of the(β/α)8-type glycosyl hydrolases

α-glucosidasesuperfamily

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GH-A


Clans of Glycoside HydrolasesClic

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COG2152NoneNoneCOG3507COG1621 / KOG0228COG / KOG

Eukaryota:FungiMetazoaViridiplantae

Eubacteria:ActinobacteriaAquificalesCyanobacteriaFirmicutesProteobacteriaThermotogales

Archaea:CrenarchaeotaEuryarchaeota

Eubacteria:ActinobacteriaFirmicutesProteobacteria

Archaea:Euryarchaeota

Eukaryota:Fungi

Eubacteria:ActinobacteriaProteobacteria

Eukaryota:FungiViridiplantae

Eubacteria:ActinobacteriaBacteroidetesFirmicutesProteobacteriaThermotogales

Eukaryota:EuglenozoaFungiParabasalideaViridiplantae

Eubacteria:ActinobacteriaBacteroidetesChloroflexiFirmicutesFusobacteriaProteobacteriaThermotogales

Archaea:Euryarchaeota

Origin

Not knownRetainingNot knownInvertingRetainingMolecularmechanism

Not knownEC 2.4.1.xEC 2.4.1.9EC 2.4.1.10EC 3.2.1.26

EC 3.2.1.55EC 3.2.1.8EC 3.2.1.37EC 3.2.1.55EC 3.2.1.99

EC 2.4.1.99EC 2.4.1.100EC 2.4.1.xEC 3.2.1.7EC 3.2.1.26EC 3.2.1.65EC 3.2.1.80

Knownenzymaticactivities

Not knownGH-JGH-FGH-FGH-JClan

GHLPGH68GH62GH43GH32Family

b-fructosidase (furanosidase) superfamilyClic

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A hierarchical structure of the b-fructosidase (furanosidase) superfamily

furanosidasesuperfamily

GH32

GH68

GH43

GH62

GHLP

clan GH-J

clan GH-F

GH32a

GH32b

GH32c

GH32d

GH68a

GH68b

GH43a

GH43b

GH43c

GH43d

GH43e

GH43f

GH43g

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GH-A


Clans of Glycoside Hydrolases

Strohmeieret al.(2004)

?

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Thank you for the attention!

Please, visit my web-site to find a list of publications:http://bioinform.genetika.ru/members/Naumoff/index.htm

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hierarchical classification of glycoside hydrolases daniil g. … · 2007-07-06 · hierarchical...

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