Clans of Peptidases
A clan contains all the modern-day peptidases that have arisen from a single evolutionary origin of peptidases. It represents one or more families that show evidence of their evolutionary relationship by their similar tertiary structures, or when structures are not available, by the order of catalytic-site residues in the polypeptide chain and often by common sequence motifs around the catalytic residues. Each clan is identified with two letters, the first representing the catalytic type of the families included in the clan (with the letter "P" being used for a clan containing families of more than one of the catalytic types serine, threonine and cysteine). Some families cannot yet be assigned to clans, and when a formal assignment is required, such a family is described as belonging to clan A-, C-, M-, S-, T- or U-, according to the catalytic type. Some clans are divided into subclans because there is evidence of a very ancient divergence within the clan, for example MA(E), the gluzincins, and MA(M), the metzincins.
Aspartic, Cysteine, Glutamic, Metallo, Asparagine, Mixed, Serine, Threonine, Unknown,Clans of Aspartic Peptidases | |||
---|---|---|---|
CLAN | FAMILY | TYPE PEPTIDASE | STRUCTURE |
AA | A2 | HIV-1 retropepsin (human immunodeficiency virus 1) | Yes |
A3 | cauliflower mosaic virus-type peptidase (cauliflower mosaic virus) | - | |
A9 | spumapepsin (human spumaretrovirus) | Yes | |
A11 | Copia transposon peptidase (Drosophila melanogaster) | - | |
A28 | DNA-damage inducible protein 1 (Saccharomyces cerevisiae) | Yes | |
A32 | PerP peptidase (Caulobacter crescentus) | Yes | |
AC | A8 | signal peptidase II (Escherichia coli) | Yes |
AD | A22 | presenilin 1 (Homo sapiens) | Yes |
A24 | type 4 prepilin peptidase 1 (Pseudomonas aeruginosa) | Yes | |
AE | A25 | gpr peptidase (Bacillus megaterium) | Yes |
A31 | HybD peptidase (Escherichia coli) | Yes | |
AF | A26 | omptin (Escherichia coli) | Yes |
unassigned | A5 | thermopsin (Sulfolobus acidocaldarius) | - |
A36 | sporulation factor SpoIIGA (Bacillus subtilis) | - | |
A37 | sso1175 g.p. (Sulfolobus solfataricus) | - | |
A39 | TseI g.p. (Aeromonas dhakensis) | - | |
Clans of Cysteine Peptidases | |||
CLAN | FAMILY | TYPE PEPTIDASE | STRUCTURE |
CA | C1 | papain (Carica papaya) | Yes |
C2 | calpain-2 (Homo sapiens) | Yes | |
C6 | potato virus Y-type helper component peptidase (potato virus Y) | Yes | |
C10 | streptopain (Streptococcus pyogenes) | Yes | |
C12 | ubiquitinyl hydrolase-L1 (Homo sapiens) | Yes | |
C16 | murine hepatitis coronavirus papain-like peptidase 1 (murine hepatitis virus) | Yes | |
C19 | ubiquitin-specific peptidase 14 (Homo sapiens) | Yes | |
C21 | tymovirus peptidase (turnip yellow mosaic virus) | Yes | |
C28 | foot-and-mouth disease virus L-peptidase (foot-and-mouth disease virus) | Yes | |
C31 | porcine reproductive and respiratory syndrome arterivirus-type cysteine peptidase alpha (lactate-dehydrogenase-elevating virus) | Yes | |
C32 | equine arteritis virus-type cysteine peptidase (porcine reproductive and respiratory syndrome virus) | Yes | |
C33 | equine arteritis virus Nsp2-type cysteine peptidase (equine arteritis virus) | Yes | |
C39 | bacteriocin-processing peptidase (Pediococcus acidilactici) | Yes | |
C47 | staphopain A (Staphylococcus aureus) | Yes | |
C51 | D-alanyl-glycyl peptidase (Staphylococcus aureus) | Yes | |
C54 | autophagin-1 (Homo sapiens) | Yes | |
C58 | YopT peptidase (Yersinia pestis) | Yes | |
C64 | Cezanne peptidase (Homo sapiens) | Yes | |
C65 | otubain-1 (Homo sapiens) | Yes | |
C66 | IdeS peptidase (Streptococcus pyogenes) | Yes | |
C67 | CylD peptidase (Homo sapiens) | Yes | |
C70 | AvrRpt2 peptidase (Pseudomonas syringae) | - | |
C71 | pseudomurein endoisopeptidase Pei (Methanobacterium phage psiM2) | - | |
C76 | viral tegument protein deubiquitinylating peptidase (human herpesvirus 1) | Yes | |
C78 | UfSP1 peptidase (Mus musculus) | Yes | |
C83 | gamma-glutamylcysteine dipeptidyltranspeptidase (Nostoc sp. PCC 7120) | Yes | |
C85 | OTUD5 peptidase (Homo sapiens) | Yes | |
C86 | ataxin-3 (Homo sapiens) | Yes | |
C87 | nairovirus deubiquitinylating peptidase (Crimean-Congo hemorrhagic fever virus) | Yes | |
C93 | LapG peptidase (Pseudomonas fluorescens) | Yes | |
C96 | McjB peptidase (Escherichia coli) | - | |
C98 | USPL1 peptidase (Homo sapiens) | - | |
C100 | agglutinin peptidase (Marasmius oreades) | Yes | |
C101 | OTULIN peptidase (Homo sapiens) | Yes | |
C102 | GtgE peptidase (Salmonella enterica) | Yes | |
C103 | HopX1 peptidase } (Pseudomonas syringae) | - | |
C104 | PlyC phage lysin (Streptococcus phage C1) | Yes | |
C105 | papain-like peptidase 1 alpha (simian hemorrhagic fever virus) | - | |
C110 | kyphoscoliosis peptidase (Mus musculus) | - | |
C111 | coagulation factor XIIIa (Homo sapiens) | Yes | |
C113 | IgdE peptidase (Streptococcus suis) | - | |
C115 | MINDY-1 protein (Homo sapiens) | - | |
C117 | SpvD g.p. (Salmonella typhimurium) | Yes | |
C119 | LotA g.p. (Legionella pneumophila) | - | |
C121 | MINDY-4 peptidase (Homo sapiens) | - | |
C124 | Lem27 (Legionella pneumophila) | - | |
CD | C11 | clostripain (Hathewaya histolytica) | Yes |
C13 | legumain (Canavalia ensiformis) | Yes | |
C14 | caspase-1 (Rattus norvegicus) | Yes | |
C25 | gingipain RgpA (Porphyromonas gingivalis) | Yes | |
C50 | separase (Saccharomyces cerevisiae) | Yes | |
C80 | RTX self-cleaving toxin (Vibrio cholerae) | Yes | |
C84 | prtH peptidase (Tannerella forsythia) | - | |
C123 | RARP2 (Rickettsia rickettsii) | - | |
CE | C5 | adenain (human adenovirus type 2) | Yes |
C48 | Ulp1 peptidase (Saccharomyces cerevisiae) | Yes | |
C55 | YopJ protein (Yersinia pseudotuberculosis) | Yes | |
C57 | vaccinia virus I7L processing peptidase (Vaccinia virus) | - | |
C63 | African swine fever virus processing peptidase (African swine fever virus) | - | |
C79 | ElaD peptidase (Escherichia coli) | Yes | |
C122 | SdeA (Legionella pneumophila) | Yes | |
CF | C15 | pyroglutamyl-peptidase I (Bacillus amyloliquefaciens) | Yes |
CL | C60 | sortase A (Staphylococcus aureus) | Yes |
C82 | L,D-transpeptidase (Enterococcus faecium) | Yes | |
CM | C18 | hepatitis C virus peptidase 2 (hepatitis C virus) | Yes |
CN | C9 | sindbis virus-type nsP2 peptidase (Sindbis virus) | Yes |
CO | C40 | dipeptidyl-peptidase VI (Lysinibacillus sphaericus) | Yes |
CP | C97 | desumoylating isopeptidase 1 (Mus musculus) | Yes |
CQ | C53 | pestivirus Npro peptidase (classical swine fever virus) | Yes |
CR | C108 | Prp peptidase (Staphylococcus aureus) | Yes |
unassigned | C7 | chestnut blight fungus virus p29 peptidase (Cryphonectria hypovirus) | - |
C8 | chestnut blight fungus virus p48 peptidase (Cryphonectria hypovirus 1) | - | |
C23 | carlavirus peptidase (apple stem pitting virus) | - | |
C27 | rubella virus peptidase (Rubella virus) | - | |
C36 | beet necrotic yellow vein furovirus-type papain-like peptidase (beet necrotic yellow vein virus) | - | |
C42 | beet yellows virus-type papain-like peptidase (beet yellows virus) | - | |
C49 | Strawberry mottle virus 3C-like peptidase (strawberry mottle virus) | - | |
C75 | AgrB peptidase (Staphylococcus aureus) | - | |
C106 | archaeosortase A (Haloferax volcanii) | - | |
C116 | dermonecrotic toxin (Pasteurella multocida) | Yes | |
C118 | EspL g.p. (Escherichia coli) | - | |
C120 | mavirus processing peptidase (Maverick-related virus strain Spezl) | - | |
Clans of Glutamic Peptidases | |||
CLAN | FAMILY | TYPE PEPTIDASE | STRUCTURE |
GA | G1 | scytalidoglutamic peptidase (Scytalidium lignicolum) | Yes |
GB | G2 | pre-neck appendage protein (bacteriophage phi-29) | Yes |
GC | G4 | Tiki1 peptidase (Homo sapiens) | - |
G6 | Ras/Rap1-specific peptidase (Vibrio vulnificus) | Yes | |
unassigned | G3 | strawberry mottle virus glutamic peptidase (strawberry mottle virus) | - |
G5 | RCE1 peptidase (Saccharomyces cerevisiae) | Yes | |
Clans of Metallo Peptidases | |||
CLAN | FAMILY | TYPE PEPTIDASE | STRUCTURE |
MA | M1 | aminopeptidase N (Homo sapiens) | Yes |
M2 | angiotensin-converting enzyme peptidase unit 1 (Homo sapiens) | Yes | |
M3 | thimet oligopeptidase (Rattus norvegicus) | Yes | |
M4 | thermolysin (Bacillus thermoproteolyticus) | Yes | |
M5 | mycolysin (Streptomyces cacaoi) | - | |
M6 | immune inhibitor A peptidase (Bacillus thuringiensis) | Yes | |
M7 | snapalysin (Streptomyces lividans) | Yes | |
M8 | leishmanolysin (Leishmania major) | Yes | |
M9 | collagenase V (Vibrio alginolyticus) | Yes | |
M10 | matrix metallopeptidase-1 (Homo sapiens) | Yes | |
M11 | gametolysin (Chlamydomonas reinhardtii) | - | |
M12 | astacin (Astacus astacus) | Yes | |
M13 | neprilysin (Homo sapiens) | Yes | |
M26 | IgA1-specific metallopeptidase (Streptococcus sanguinis) | - | |
M27 | tentoxilysin (Clostridium tetani) | Yes | |
M30 | hyicolysin (Staphylococcus hyicus) | - | |
M32 | carboxypeptidase Taq (Thermus aquaticus) | Yes | |
M34 | anthrax lethal factor (Bacillus anthracis) | Yes | |
M35 | deuterolysin (Aspergillus flavus) | Yes | |
M36 | fungalysin (Aspergillus fumigatus) | Yes | |
M41 | FtsH peptidase (Escherichia coli) | Yes | |
M43 | cytophagalysin (Cytophaga sp.) | Yes | |
M48 | Ste24 peptidase (Saccharomyces cerevisiae) | Yes | |
M49 | dipeptidyl-peptidase III (Rattus norvegicus) | Yes | |
M54 | archaelysin (Methanocaldococcus jannaschii) | Yes | |
M56 | BlaR1 peptidase (Staphylococcus aureus) | Yes | |
M57 | prtB g.p. (Myxococcus xanthus) | - | |
M60 | enhancin (Lymantria dispar nucleopolyhedrovirus) | Yes | |
M61 | glycyl aminopeptidase (Sphingomonas capsulata) | Yes | |
M64 | IgA peptidase (Clostridium ramosum) | Yes | |
M66 | StcE peptidase (Escherichia coli) | Yes | |
M72 | peptidyl-Asp metallopeptidase (Pseudomonas aeruginosa) | - | |
M76 | Atp23 peptidase (Homo sapiens) | - | |
M78 | ImmA peptidase (Bacillus subtilis) | Yes | |
M80 | Wss1 peptidase (Saccharomyces cerevisiae) | Yes | |
M84 | MpriBi peptidase (Bacillus intermedius) | - | |
M85 | NleC peptidase (Escherichia coli) | Yes | |
M87 | chloride channel accessory protein 1 (Homo sapiens) | - | |
M88 | IMPa peptidase (Pseudomonas aeruginosa) | Yes | |
M90 | MtfA peptidase (Escherichia coli) | Yes | |
M91 | NleD peptidase (Escherichia coli) | - | |
M95 | abylysin (Pyrococcus abyssi) | Yes | |
M97 | EcxAB peptidase (Escherichia coli) | Yes | |
M98 | YghJ g.p. (Escherichia coli) | - | |
M100 | spartan peptidase (Homo sapiens) | Yes | |
M101 | flagellinolysin (Clostridium haemolyticum) | - | |
M102 | DA1 peptidase (Arabidopsis thaliana) | - | |
M105 | MAP1 peptidase (Myxococcus xanthus) | - | |
M106 | FP0506 g.p. (Flavobacterium psychrophilum) | - | |
MC | M14 | carboxypeptidase A1 (Homo sapiens) | Yes |
M86 | PghP gamma-polyglutamate hydrolase (Bacillus phage phiNIT1) | Yes | |
M99 | Csd4 peptidase (Helicobacter pylori) | Yes | |
MD | M15 | zinc D-Ala-D-Ala carboxypeptidase (Streptomyces albus) | Yes |
M74 | murein endopeptidase (Escherichia coli) | Yes | |
ME | M16 | pitrilysin (Escherichia coli) | Yes |
M44 | pox virus metallopeptidase (Vaccinia virus) | - | |
MF | M17 | leucyl aminopeptidase 3 (Bos taurus) | Yes |
MG | M24 | methionyl aminopeptidase 1 (Escherichia coli) | Yes |
MH | M18 | aminopeptidase I (Saccharomyces cerevisiae) | Yes |
M20 | glutamate carboxypeptidase (Pseudomonas sp.) | Yes | |
M28 | aminopeptidase S (Streptomyces griseus) | Yes | |
M42 | glutamyl aminopeptidase (Lactococcus lactis) | Yes | |
MJ | M19 | membrane dipeptidase (Homo sapiens) | Yes |
M38 | isoaspartyl dipeptidase (Escherichia coli) | Yes | |
MM | M50 | site 2 peptidase (Homo sapiens) | Yes |
MN | M55 | D-aminopeptidase DppA (Bacillus subtilis) | Yes |
MO | M23 | beta-lytic metallopeptidase (Achromobacter lyticus) | Yes |
MP | M67 | PSMD14 peptidase (Saccharomyces cerevisiae) | Yes |
MQ | M29 | aminopeptidase T (Thermus aquaticus) | Yes |
MS | M75 | imelysin (Pseudomonas aeruginosa) | Yes |
MT | M81 | microcystinase MlrC (Sphingomonas sp. ACM-3962) | Yes |
MU | M103 | TldD peptidase (Escherichia coli) | Yes |
unassigned | M73 | camelysin (Bacillus cereus) | Yes |
Clans of Asparagine Peptide Lyases | |||
CLAN | FAMILY | TYPE PEPTIDE LYASE | STRUCTURE |
NA | N1 | nodavirus peptide lyase (flock house virus) | Yes |
N2 | tetravirus coat protein (Nudaurelia capensis omega virus) | Yes | |
N8 | poliovirus capsid VP0-type self-cleaving protein (human poliovirus 1) | Yes | |
NB | N6 | YscU protein (Yersinia pseudotuberculosis) | Yes |
NC | N7 | reovirus type 1 coat protein (Mammalian orthoreovirus 1) | Yes |
ND | N4 | Tsh-associated self-cleaving domain and similar (Escherichia coli) | Yes |
NE | N5 | picobirnavirus self-cleaving protein (Human picobirnavirus) | Yes |
Clans of Mixed (C, S, T) Catalytic Type | |||
CLAN | FAMILY | TYPE PEPTIDASE | STRUCTURE |
PA | C3 | poliovirus-type picornain 3C (human poliovirus 1) | Yes |
C4 | nuclear-inclusion-a peptidase (plum pox virus) | Yes | |
C24 | rabbit hemorrhagic disease virus 3C-like peptidase (rabbit hemorrhagic disease virus) | Yes | |
C30 | porcine transmissible gastroenteritis virus-type main peptidase (transmissible gastroenteritis virus) | Yes | |
C37 | calicivirin (Southampton virus) | Yes | |
C62 | gill-associated virus 3C-like peptidase (gill-associated virus) | - | |
C74 | pestivirus NS2 peptidase (bovine viral diarrhea virus 1) | Yes | |
C99 | iflavirus processing peptidase (Ectropis obliqua picorna-like virus) | - | |
C107 | alphamesonivirus 3C-like peptidase (Cavally virus) | Yes | |
C125 | protease (Sacbrood virus) | - | |
S1 | chymotrypsin A (Bos taurus) | Yes | |
S3 | togavirin (Sindbis virus) | Yes | |
S6 | IgA1-specific serine peptidase (Neisseria gonorrhoeae) | Yes | |
S7 | flavivirin (yellow fever virus) | Yes | |
S29 | hepacivirin (hepatitis C virus) | Yes | |
S30 | potyvirus P1 peptidase (plum pox virus) | - | |
S31 | pestivirus NS3 polyprotein peptidase (bovine viral diarrhea virus 1) | Yes | |
S32 | equine arteritis virus serine peptidase (equine arteritis virus) | Yes | |
S39 | sobemovirus peptidase (cocksfoot mottle virus) | Yes | |
S46 | dipeptidyl-peptidase 7 (Porphyromonas gingivalis) | Yes | |
S55 | SpoIVB peptidase (Bacillus subtilis) | - | |
S64 | Ssy5 peptidase (Saccharomyces cerevisiae) | - | |
S65 | picornain-like serine peptidase (Breda virus) | - | |
S75 | White bream virus serine peptidase (White bream virus) | - | |
PB | C44 | amidophosphoribosyltransferase precursor (Homo sapiens) | Yes |
C45 | acyl-coenzyme A:6-aminopenicillanic acid acyl-transferase precursor (Penicillium chrysogenum) | Yes | |
C59 | penicillin V acylase precursor (Lysinibacillus sphaericus) | Yes | |
C69 | dipeptidase A (Lactobacillus helveticus) | - | |
C89 | acid ceramidase precursor (Homo sapiens) | Yes | |
C95 | lysosomal 66.3 kDa protein (Mus musculus) | Yes | |
P2 | EGF-like module containing mucin-like hormone receptor-like 2 (Homo sapiens) | Yes | |
S45 | penicillin G acylase precursor (Escherichia coli) | Yes | |
T1 | archaean proteasome, beta component (Thermoplasma acidophilum) | Yes | |
T2 | glycosylasparaginase precursor (Homo sapiens) | Yes | |
T3 | gamma-glutamyltransferase 1 (Escherichia coli) | Yes | |
T7 | CwpV self-cleaving threonine peptidase (Peptoclostridium difficile) | - | |
PC | C26 | gamma-glutamyl hydrolase (Rattus norvegicus) | Yes |
C56 | PfpI peptidase (Pyrococcus furiosus) | Yes | |
S51 | dipeptidase E (Escherichia coli) | Yes | |
PD | C46 | hedgehog protein (Drosophila melanogaster) | Yes |
N9 | intein-containing V-type proton ATPase catalytic subunit A (Saccharomyces cerevisiae) | Yes | |
N10 | intein-containing replicative DNA helicase precursor (Synechocystis sp. PCC 6803) | Yes | |
N11 | intein-containing chloroplast ATP-dependent peptide lyase (Chlamydomonas eugametos) | - | |
PE | P1 | DmpA aminopeptidase (Ochrobactrum anthropi) | Yes |
T5 | ornithine acetyltransferase precursor (Saccharomyces cerevisiae) | Yes | |
Clans of Serine Peptidases | |||
CLAN | FAMILY | TYPE PEPTIDASE | STRUCTURE |
SB | S8 | subtilisin Carlsberg (Bacillus licheniformis) | Yes |
S53 | sedolisin (Pseudomonas sp. 101) | Yes | |
SC | S9 | prolyl oligopeptidase (Sus scrofa) | Yes |
S10 | carboxypeptidase Y (Saccharomyces cerevisiae) | Yes | |
S15 | Xaa-Pro dipeptidyl-peptidase (Lactococcus lactis) | Yes | |
S28 | lysosomal Pro-Xaa carboxypeptidase (Homo sapiens) | Yes | |
S33 | prolyl aminopeptidase (Neisseria gonorrhoeae) | Yes | |
S37 | PS-10 peptidase (Streptomyces lividans) | - | |
S82 | autocrine proliferation repressor protein A (Dictyostelium discoideum) | - | |
SE | S11 | D-Ala-D-Ala carboxypeptidase A (Geobacillus stearothermophilus) | Yes |
S12 | D-Ala-D-Ala carboxypeptidase B (Streptomyces lividans) | Yes | |
S13 | D-Ala-D-Ala peptidase C (Escherichia coli) | Yes | |
SF | S24 | repressor LexA (Escherichia coli) | Yes |
S26 | signal peptidase I (Escherichia coli) | Yes | |
SH | S21 | cytomegalovirus assemblin (human herpesvirus 5) | Yes |
S73 | gpO peptidase (Enterobacteria phage P2) | - | |
S77 | prohead peptidase gp21 (Enterobacteria phage T4) | Yes | |
S78 | prohead peptidase (Enterobacteria phage HK97) | - | |
S80 | prohead peptidase gp175 (Pseudomonas phage phiKZ) | - | |
SJ | S16 | Lon-A peptidase (Escherichia coli) | Yes |
S50 | infectious pancreatic necrosis birnavirus Vp4 peptidase (infectious pancreatic necrosis virus) | Yes | |
S69 | Tellina virus 1 VP4 peptidase (Tellina virus 1) | Yes | |
SK | S14 | peptidase Clp (Escherichia coli) | Yes |
S41 | C-terminal processing peptidase-1 (Escherichia coli) | Yes | |
S49 | signal peptide peptidase A (Escherichia coli) | Yes | |
SO | S74 | Escherichia coli phage K1F endosialidase CIMCD self-cleaving protein (Enterobacteria phage K1F) | Yes |
SP | S59 | nucleoporin 145 (Homo sapiens) | Yes |
SR | S60 | lactoferrin (Homo sapiens) | Yes |
SS | S66 | murein tetrapeptidase LD-carboxypeptidase (Pseudomonas aeruginosa) | Yes |
ST | S54 | rhomboid-1 (Drosophila melanogaster) | Yes |
unassigned | S48 | HetR putative peptidase (Anabaena variabilis) | Yes |
S62 | influenza A PA peptidase (influenza A virus) | Yes | |
S68 | PIDD auto-processing protein unit 1 (Homo sapiens) | - | |
S71 | MUC1 self-cleaving mucin (Homo sapiens) | Yes | |
S72 | dystroglycan (Homo sapiens) | Yes | |
S79 | CARD8 self-cleaving protein (Homo sapiens) | Yes | |
S81 | destabilase (Hirudo medicinalis) | Yes | |
S85 | small protease (Pseudomonas aeruginosa) | Yes | |
Clans of Threonine Peptidases | |||
CLAN | FAMILY | TYPE PEPTIDASE | STRUCTURE |
unassigned | T8 | HopB1 g.p. (Pseudomonas syringae) | - |
Clans of Peptidases of Unknown Catalytic Type | |||
CLAN | FAMILY | TYPE PEPTIDASE | STRUCTURE |
unassigned | U32 | collagenase (Porphyromonas gingivalis) | Yes |
U40 | protein P5 murein endopeptidase (bacteriophage phi-6) | Yes | |
U49 | Lit peptidase (Escherichia coli) | - | |
U56 | homomultimeric peptidase (Thermotoga maritima) | Yes | |
U57 | yabG protein (Bacillus subtilis) | - | |
U69 | AIDA-I self-cleaving autotransporter protein (Escherichia coli) | Yes | |
U72 | Dop isopeptidase (Mycobacterium tuberculosis) | Yes | |
U74 | neprosin (Nepenthes ventrata) | - |