Thioesterase

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(Difference between revisions)

Revision as of 11:49, 18 February 2020

Human ubiquitin esterase 2 (deepskyblue) complex with ubiquitin (green) and zinc+2 ion (grey) (PDB code 2hd5).

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1 Function
2 Disease
3 Structural highlights
4 3D structures of thioesterase

Function

Thioesterase (TE) catalyzes the break of an ester bond to produce acid and alcohol at a thiol group. TEs are substrate-specific.

Palmitoyl protein TE removes fatty acids like palmitate from modified cysteine residues during lysosomal degradation^[1]. For details see Palmitoyl protein thioesterase.
4-hydroxybenzoyl-CoA TE converts 4-hydroxybenzoyl-CoA to 4-hydroxybenzoate and CoA^[2].
Acyl-CoA TE hydrolyzes acyl-CoA to the fatty acid and CoA and is involved in lipid metabolism^[3]. See also YbgC.
Fluoroacetyl-CoA TE from Streptomyces cattleya hydrolyzes fluoroacetyl-CoA thus preventing it from being metabolized to the lethal 4-hydroxy-trans-aconitate^[4].
Ubiquitin TE or ubiquitin carboxyl-terminal hydrolase (USP) removes conjugated ubiquitin (UB) from proteins thus regulating protein level by preventing their degradation. USP hydrolyze the peptide bond at the C-terminal glycine of ubiquitin. The USPs are involved in the processing of poly-UB precursors and of ubiquitinated proteins^[5]. USP contains catalytic domain surrounded several domains: Ub-like (UBL); Ub-associated (UBA); zinc finger-Ub-specific protease domain (UBP or DUSP); TRAF homology domain.

USP-L1, USP25 hydrolyze C-terminal adducts of UB.
USP-L3 hydrolyzes C-terminal adducts of UB and NEDD8.
USP5 cleaves multiubiquitin polymers.
USP6 has ATP-independent isopeptidase activity.
USP7, USP4, USP13, USP15 deubiquitinate several proteins.
USP8 removes conjugated ubiquitin from proteins thus preventing protein degradation. USP8 is involved in cell proliferation and is active in the M phase of proliferation.
USP11, USP14 are proteasome-associated.
USP12 stabilizes T-cell complexes^[6].
USP16, USP21 deubiquitinate histone H2A.
USP18 is a down regulator of the type I interferon signaling pathway^[7].
USP28 deubiquitinates proteins of the DNA damage pathway.
USP33 regulates centrosome duplication.
USP37 deubiquitinates cyclin A.
USP46 deubiquitinates AMPA receptor^[8].

Disease

Mutations in palmiotoyl protein TE cause neuronal ceroid lipocfuscinosis^[9]^[10].

Structural highlights

. Ubiquitin thioesterase 2 active site contains the . The metal-binding enzyme contains a . The ^[11].

3D structures of thioesterase

Thioesterase 3D structures

References

↑ Cho S, Dawson G. Palmitoyl protein thioesterase 1 protects against apoptosis mediated by Ras-Akt-caspase pathway in neuroblastoma cells. J Neurochem. 2000 Apr;74(4):1478-88. PMID:10737604
↑ Zhuang Z, Gartemann KH, Eichenlaub R, Dunaway-Mariano D. Characterization of the 4-hydroxybenzoyl-coenzyme A thioesterase from Arthrobacter sp. strain SU. Appl Environ Microbiol. 2003 May;69(5):2707-11. PMID:12732540
↑ Hunt MC, Alexson SE. The role Acyl-CoA thioesterases play in mediating intracellular lipid metabolism. Prog Lipid Res. 2002 Mar;41(2):99-130. PMID:11755680
↑ Weeks AM, Coyle SM, Jinek M, Doudna JA, Chang MC. Structural and Biochemical Studies of a Fluoroacetyl-CoA-Specific Thioesterase Reveal a Molecular Basis for Fluorine Selectivity. Biochemistry. 2010 Oct 11. PMID:20836570 doi:10.1021/bi101102u
↑ Jagannathan M, Nguyen T, Gallo D, Luthra N, Brown GW, Saridakis V, Frappier L. A role for USP7 in DNA replication. Mol Cell Biol. 2014 Jan;34(1):132-45. doi: 10.1128/MCB.00639-13. Epub 2013 Nov 4. PMID:24190967 doi:http://dx.doi.org/10.1128/MCB.00639-13
↑ Jahan AS, Lestra M, Swee LK, Fan Y, Lamers MM, Tafesse FG, Theile CS, Spooner E, Bruzzone R, Ploegh HL, Sanyal S. Usp12 stabilizes the T-cell receptor complex at the cell surface during signaling. Proc Natl Acad Sci U S A. 2016 Feb 9;113(6):E705-14. doi:, 10.1073/pnas.1521763113. Epub 2016 Jan 25. PMID:26811477 doi:http://dx.doi.org/10.1073/pnas.1521763113
↑ Malhotra S, Morcillo-Suarez C, Nurtdinov R, Rio J, Sarro E, Moreno M, Castillo J, Navarro A, Montalban X, Comabella M. Roles of the ubiquitin peptidase USP18 in multiple sclerosis and the response to interferon-beta treatment. Eur J Neurol. 2013 Oct;20(10):1390-7. doi: 10.1111/ene.12193. Epub 2013 May 22. PMID:23700969 doi:http://dx.doi.org/10.1111/ene.12193
↑ Huo Y, Khatri N, Hou Q, Gilbert J, Wang G, Man HY. The deubiquitinating enzyme USP46 regulates AMPA receptor ubiquitination and trafficking. J Neurochem. 2015 Sep;134(6):1067-80. doi: 10.1111/jnc.13194. Epub 2015 Jul 16. PMID:26077708 doi:http://dx.doi.org/10.1111/jnc.13194
↑ Vesa J, Hellsten E, Verkruyse LA, Camp LA, Rapola J, Santavuori P, Hofmann SL, Peltonen L. Mutations in the palmitoyl protein thioesterase gene causing infantile neuronal ceroid lipofuscinosis. Nature. 1995 Aug 17;376(6541):584-7. PMID:7637805 doi:http://dx.doi.org/10.1038/376584a0
↑ van Diggelen OP, Thobois S, Tilikete C, Zabot MT, Keulemans JL, van Bunderen PA, Taschner PE, Losekoot M, Voznyi YV. Adult neuronal ceroid lipofuscinosis with palmitoyl-protein thioesterase deficiency: first adult-onset patients of a childhood disease. Ann Neurol. 2001 Aug;50(2):269-72. PMID:11506414
↑ Renatus M, Parrado SG, D'Arcy A, Eidhoff U, Gerhartz B, Hassiepen U, Pierrat B, Riedl R, Vinzenz D, Worpenberg S, Kroemer M. Structural basis of ubiquitin recognition by the deubiquitinating protease USP2. Structure. 2006 Aug;14(8):1293-302. PMID:16905103 doi:10.1016/j.str.2006.06.012

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Michal Harel, Alexander Berchansky, Joel L. Sussman

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Category: Topic Page

Thioesterase

From Proteopedia

Revision as of 11:49, 18 February 2020

Contents

Function

Disease

Structural highlights

3D structures of thioesterase

References

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@@ Line 33: / Line 33: @@
 </StructureSection>
-==3D structures of thioesterase==
-Updated on {{REVISIONDAY2}}-{{MONTHNAME|{{REVISIONMONTH}}}}-{{REVISIONYEAR}}
-{{#tree:id=OrganizedByTopic|openlevels=0|
-*Maristoyl-ACP-specific thioesterase
-**[[1tht]] – TE – ''Vibrio harveyi''
-*4-hydroxybenzoyl-CoA thioesterase
-**[[1bvq]] – PsTE – ''Pseudomonas''<br />
-**[[1lo7]] – PsTE + 4-hydroxyphenyl CoA<br />
-**[[1lo9]] - PsTE (mutant) + 4-hydroxybenzoyl CoA<br />
-**[[1lo8]] - PsTE + 4-hydroxybenzyl CoA<br />
-**[[1q4s]] - ArTE + 4-hydroxybenzoate – '''Arthrobacte''r'<br />
-**[[1q4t]] - ArTE + 4-hydroxyphenyl CoA<br />
-**[[1q4u]] - ArTE + 4-hydroxybenzyl CoA<br />
-**[[3r32]], [[3r35]], [[3r37]], [[3r38]], [[3r3b]], [[3r3c]], [[3r3d]], [[3r3f]], [[3tea]] - ArTE (mutant) + 4-hydroxyphenacyl CoA<br />
-**[[3r36]] - ArTE (mutant) + 4-hydroxybenzoate<br />
-**[[3r3a]] - ArTE (mutant) + 4-hydroxybenzoate + CoA<br />
-**[[3r34]] - ArTE (mutant) + CoA<br />
-**[[6fdg]] – SaTE  – ''Staphylococcus aureus''<br />
-**[[5wh9]] – TE  – ''Bacillus halodurans''<br />
-*Palmitoyl protein thioesterase
-**[[3gro]] - hTEI - human<br />
-**[[1pja]] – hTEII <br />
-**[[1eh5]] – bTE1 + Palmitate – bovine<br />
-**[[1ei9]] – bTE1<br />
-**[[1exw]] – bTE1 + hexadecylsulfonyl fluoride<br />
-*Acyl-CoA thioesterase or acyl-CoA thioester hydrolase
-**[[3hlk]] – hTE2<br />
-**[[3k2i]] – hTE4<br />
-**[[2qq2]] - hTE7 C terminal<br />
-**[[3fo5]] – hTE11 START domain<br />
-**[[3b7k]], [[4moc]] – hTE12<br />
-**[[4mob]] – hTE12 + ADP<br />
-**[[2v1o]] – mTE7 hotdog domain – mouse<br />
-**[[2q2b]] – mTE7 C terminal<br />
-**[[6vfy]] – mTE7 42-367 + acyl-CoA TE hydrolase + CoA<br />
-**[[1tbu]] – TEI N terminal (peroximal) – yeast<br />
-**[[2pzh]] - TE - ''Helicobacter pylori''<br />
-**[[3rd7]] – TE – ''Mycobacterium avium''<br />
-**[[5hz4]], [[5hwf]], [[5egk]] – SaTE (mutant)  – ''Staphylococcus aureus''<br />
-**[[4egj]] – SaTE + CoA<br />
-**[[1ivn]], [[5tic]] – EcTEI – ''Escherichia coli''<br />
-**[[1jrl]] – EcTEI (mutant) <br />
-**[[1j00]] – EcTEI + diethyl phosphono derivative<br />
-**[[1v2g]], [[1u8u]] - EcTEI + octanoic acid<br />
-**[[5tid]], [[5tie]], [[5tif]] - EcTEI (mutant) + octanoic acid<br />
-**[[5tid]], [[5tie]], [[5tif]] - EcTEI (mutant) + octanoic acid<br />
-**[[5t06]] - EcTEI (mutant) + hexanoyl-CoA<br />
-**[[5t07]] - EcTEI (mutant) + decanoyl-CoA<br />
-**[[1c8u]] – EcTEII <br />
-**[[3u0a]] – TEII – ''Mycobacterium marinum''<br />
-**[[4qfw]] – YpTE II – ''Yersinia pestis''<br />
-**[[4r4u]] – YpTE II + CoA<br />
-**[[5szv]], [[5szu]] – NmTE  + CoA – ''Neisseria meningitis''<br />
-**[[5szz]], [[5szy]] – NmTE  + CoA + GDP<br />
-**[[5v3a]] – NmTE  + CoA + GDP <br />
-**[[5t02]] – NmTE (mutant)  + CoA + GDP <br />
-*Acyl protein thioesterase
-**[[5sym]], [[6bje]] – hTE 1 + inhibitor<br />
-**[[6qgs]] – hTE 1 + palmitate<br />
-**[[6qgq]], [[6qgo]], [[6qgn]] – hTE 1 (mutant) + palmitate<br />
-**[[5syn]] – hTE 2 + inhibitor<br />
-**[[6avy]] – TE 2 - maize<br />
-**[[1fj2]] – EcTEI<br />
-*Acyl-ACP thioesterase
-**[[2ess]] – TE – ''Bacterioides thetaiotaomicron''<br />
-**[[4gak]] – TE – ''Spirosoma linguale''<br />
-*ACP-polyene thioesterase
-**[[4i4j]] – TE – ''Streptomyces globisporus''<br />
-*Fluoroacetyl-CoA thioesterase
-**[[3kuv]], [[3kuw]], [[3kvi]] – ScTE (mutant) + acetate derivative – ''Streptomyces cattleya''<br />
-**[[3kv7]], [[3kv8]], [[3p2r]], [[3p2s]] – ScTE + acetate derivative<br />
-**[[3kvu]] – ScTE + acetyl-CoA<br />
-**[[3kvz]], [[3kw1]] – ScTE + FAcOPan<br />
-**[[3kx7]], [[3kx8]], [[3p2q]] – ScTE <br />
-**[[3p3f]] – ScTE (mutant) <br />
-**[[3p3i]] - ScTE (mutant) + acetate derivative + CoA<br />
-*Dihydroxynaphthoyl-CoA thioesterase
-**[[4k00]] – TE – ''Synechocystis''<br />
-**[[4k02]] – AtTE – ''Arabidopsis thaliana''<br />
-*RedJ thioesterase
-**[[3qmv]], [[3qmw]] – TE –'' Streptomyces coelicolor''
-*Orf6 thioesterase
-**[[4i45]] – TE – ''Photobacterium profundum''<br />
-*thioesterase
-**[[2av9]], [[2o5u]], [[2o6b]], [[2o6t]], [[2o6u]], [[3qy3]], [[4qd7]] – PaTE – ''Pseudomonas aeruginosa''<br />
-**[[4qda]], [[4qdb]] – PaTE (mutant)<br />
-**[[4qd8]] – PaTE + phenacyl-CoA<br />
-**[[4qd9]] – PaTE + benzoyl-CoA<br />
-*Ubiquitin thioesterase or ubiquitin carboxyl-terminal hydrolase
-*Ubiquitin thioesterase 2
-**[[2zfy]] – hUSP <BR />
-**[[2hd5]], [[3nhe]], [[3v6c]], [[3v6e]] – hUSP + Ub   <BR />
-**[[1tff]] – hUSP (mutant)  <BR />
-**[[2ibi]], [[5xve]] – hUSP (mutant) + Ub <BR />
-**[[5xu8]] – hUSP + Ub + thioguanine <BR />
-*Ubiquitin thioesterase 3
-**[[2qiy]] – yUSP + USP-associated protein  <BR />
-*Ubiquitin thioesterase 4
-**[[2y6e]] – hUSP catalytic domain <br />
-**[[5ctr]] – hUSP DUSP-UBL domain + SART3<br />
-**[[3jyu]] – mUSP N terminal domain <br />
-*Ubiquitin thioesterase 5
-**[[2dag]] – hUSP UBA domain 1 - NMR<br />
-**[[2dak]] – hUSP UBA domain 2 - NMR<br />
-**[[2g43]], [[6dxh]], [[6dxt]] – hUSP zinc finger USP domain<br />
-**[[2g45]] – hUSP zinc finger USP domain + Ub<br />
-**[[6p9g]], [[6nft]] – hUSP zinc finger USP domain + quinazoline derivative<br />
-**[[3ihp]] – hUSP + Ub  <BR />
-*Ubiquitin thioesterase 7; domains - TRAF 63-205; catalytic 207-560; UBL1 537-664; UBL2 665-865
-**[[2f1z]], [[5j7t]] – hUSP<br />
-**[[1nb8]], [[4m5w]], [[4m5x]] – hUSP catalytic domain <br />
-**[[5kyb]] – hUSP catalytic domain (mutant) <br />
-**[[2kvr]] – hUSP UBL domain - NMR<br />
-**[[4pyz]], [[4wph]], [[4wpi]] – hUSP UBL domains 1+2<br />
-**[[5fwi]], [[5j7t]] – hUSP catalytic + UBL domains 1+2<br />
-**[[1yze]], [[2f1w]] – hUSP TRAF domain <br />
-**[[2ylm]] – hUSP C terminal domain <br />
-**[[2f1x]], [[2f1y]], [[2foj]], [[2foo]], [[2fop]] – hUSP TRAF domain/peptide <br />
-**[[4zv3]] – mUSP N terminal domain <br />
-*Ubiquitin thioesterase 7 complexes
-**[[1nbf]] – hUSP catalytic domain + Ub aldehyde<br />
-**[[5jtj]], [[5jtv]] – hUSP catalytic domain + polyubiquitin <br />
-**[[5whc]], [[5n9r]], [[5n9t]], [[5uqv]], [[5uqx]], [[6f5h]], [[5vsk]], [[5vsb]], [[5vs6]], [[5ngf]], [[5nge]] – hUSP catalytic domain + inhibitor<br />
-**[[5kyc]], [[5kyd]], [[5kye]], [[5kyf]] – hUSP catalytic domain (mutant) + polyubiquitin <br />
-**[[1yy6]] – hUSP TRAF domain + EBNA1 peptide<br />
-**[[2xxn]] – hUSP TRAF domain + VIRF-4 peptide<br />
-**[[3mqr]] – hUSP TRAF domain + HDMX peptide<br />
-**[[3mqs]] – hUSP TRAF domain + HDM2 peptide<br />
-**[[2fop]] – hUSP TRAF domain + MDM2 peptide<br />
-**[[4jjq]] – hUSP TRAF domain + E2 peptide<br />
-**[[5gg4]] – hUSP TRAF domain + E3 peptide<br />
-**[[4kg9]] – hUSP TRAF domain + MCM-BP peptide<br />
-**[[4ysi]], [[2foj]], [[2foo]] – hUSP TRAF domain + peptide<br />
-**[[4yoc]], [[4z96]], [[4z97]] – hUSP residues 560-1102 + DNMT1<br />
-**[[5c6d]] – hUSP UBL2 domain + UHRF1<br />
-**[[5c56]] – hUSP residues 560-1102 + Ub E3 ligase Icp0444w<br />
-*Ubiquitin thioesterase 8
-**[[1whb]] – hUSP rhodanase domain - NMR<br />
-**[[2gwf]] – hUSP rhodanase domain + NRDP1br />
-**[[2a9u]] – hUSP N terminal domain <br />
-**[[2gfo]] – hUSP catalytic domain <br />
-*Ubiquitin thioesterase 8 complexes
-**[[2gwf]] – hUSP rhodanase domain + ring finger protein 41 USP8 interaction domain<br />
-**[[3n3k]] – hUSP catalytic domain + Ub<br />
-**[[3mhh]], [[3m99]], [[4fip]], [[4fjc]], [[4f5k]], [[4fk5]], [[4wa6]] – yUSP + SUS1 + SGF11 +SGF73 <br />
-**[[3mhs]], [[6aqr]] – yUSP + SUS1 + SGF11 +SGF73 + Ub<br />
-**[[4zux]] – yUSP + SUS1 + SGF11 +SGF73 + Ub + Histones H3,2, H4, H2A, H2B + DNA<br />
-*Ubiquitin thioesterase 9
-**[[5vbd]] – hUSP 9x UBL domain <br />
-*Ubiquitin thioesterase 11
-**[[4mel]] – hUSP DUSP+UBL domains <br />
-**[[5ok6]] – hUSP DUSP+UBL domains + peptide<br />
-**[[4mem]] – rUSP DUSP+UBL domains - rat<br />
-*Ubiquitin thioesterase 12
-**[[5k16]] – hUSP <br />
-**[[5k1a]], [[5k1b]], [[5k1c]] – hUSP + WD repeat-containing protein<br />
-**[[5l8w]] – hUSP + WD repeat-containing protein + polyubiquitin<br />
-*Ubiquitin thioesterase 13
-**[[2l80]] – hUSP zinc finger domain - NMR<br />
-**[[2lbc]] – hUSP UBA domain - NMR<br />
-*Ubiquitin thioesterase 14
-**[[2ayn]] – hUSP catalytic domain<br />
-**[[2ayo]] – hUSP catalytic domain + Ub aldehyde<br />
-**[[6iin]], [[6iim]], [[6iil]], [[6iik]] – hUSP catalytic domain + inhibitor<br />
-**[[5gjq]] – hUSP in proteasome – Cryo EM<br />
-**[[1wiv]] – AtUSP UBA domain - NMR<br />
-**[[1wgg]] – mUSP N terminal domain - NMR<br />
-*Ubiquitin thioesterase 15; domains – DUSP 1-133; UBL 134-223; catalytic 255-439+757-919
-**[[3lmn]] – hUSP DUSP domain  <BR />
-**[[1w6v]] – hUSP DUSP domain - NMR<br />
-**[[3ppa]], [[3pv1]], [[3t9l]], [[4a3o]], [[4a3p]] – hUSP DUSP+UBL domains <br />
-**[[6dj9]] – hUSP DUSP domain + Ub<BR />
-**[[6gha]] – hUSP catalytic domain  <BR />
-**[[5ctr]], [[5jjw]] – hUSP DUSP+UBL domains + SART HAT domain <br />
-**[[6ml1]], [[6crn]], [[6cpm]] – hUSP catalytic domain + Ub <BR />
-**[[6gh9]] – hUSP catalytic domain + quinone derivative <BR />
-*Ubiquitin thioesterase 16
-**[[2i50]] – hUSP zinc finger domain - NMR  <BR />
-*Ubiquitin thioesterase 18
-**[[5cht]] – mUSP   <BR />
-**[[5chv]] – mUSP + Ub-like  <BR />
-*USP 20
-**[[6kcz]] – hUSP zinc finger + UBP domains - NMR  <BR />
-*Ubiquitin thioesterase 21
-**[[3i3t]], [[2y5b]] – hUSP + Ub  <BR />
-**[[3mtn]] – hUSP + Ub inhibitor <BR />
-*Ubiquitin thioesterase 25
-**[[5o71]] – hUSP <BR />
-**[[6hem]], [[6h4k]] – hUSP C terminal  <BR />
-**[[6hel]], [[6h4j]] – hUSP residues 157-714 <BR />
-**[[1vdl]] – mUSP catalytic domain <br />
-*Ubiquitin thioesterase 28;; domains – catalytic 149-399
-**[[6hej]] – hUSP residues 149-703<BR />
-**[[2lva]], [[2muu]], [[2mux]] – hUSP N terminal - NMR  <BR />
-**[[6h4i]] – hUSP catalytic domain <BR />
-**[[6heh]] – hUSP catalytic domain (mutant)<BR />
-**[[6hek]] – hUSP residues 149-703 + Ub  <BR />
-**[[6hei]], [[6h4h]] – hUSP catalytic domain (mutant) + Ub  <BR />
-*Ubiquitin thioesterase 30
-**[[5ohp]] – hUSP (mutant) + diubiquitin <BR />
-**[[5ohk]], [[5ohn]] – hUSP (mutant) + polyubiquitin <BR />
-*Ubiquitin thioesterase 33
-**[[2uzg]] – hUSP zinc finger domain - NMR  <BR />
-*Ubiquitin thioesterase 35
-**[[5txk]] – hUSP (mutant) + polyubiquitin <BR />
-*Ubiquitin thioesterase 37
-**[[3ihr]] – hUSP (mutant)  <BR />
-**[[3a7s]] – hUSP catalytic domain (mutant)  <BR />
-**[[3u12]] – hUSP pleckstrin domain <br />
-*Ubiquitin thioesterase 38
-**[[4rxx]] – hUSP N terminal  <BR />
-*Ubiquitin thioesterase 46
-**[[5cvm]], [[5l8h]] – hUSP + Ub  <BR />
-**[[5cvn]], [[5cvo]] – hUSP + WDR48 + Ub<br />
-**[[6jlq]] – hUSP + WDR48 + WDR20<br />
-*Ubiquitin thioesterase Cyld
-**[[2vhf]] – hUSP  USP domain <BR />
-*Ubiquitin thioesterase L1
-**[[2etl]] – hUSP-L1  <br />
-**[[3irt]], [[4jkj]] – hUSP  (mutant) <BR />
-**[[2len]] – hUSP  (mutant) - NMR<BR />
-**[[3ifw]], [[3kvf]], [[3kw5]] – hUSP  (mutant) + Ub<BR />
-**[[4dm9]] – hUSP + peptide inhibitor<BR />
-*Ubiquitin thioesterase L3 or UCH-L3
-**[[1uch]] – hUSP-L3  <br />
-**[[1xd3]] – hUSP-L3 + UBC<br />
-**[[6isu]] – hUSP-L3 + Ub <br />
-**[[2we6]] – PfUSP-L3 – ''Plasmodium falciparum''<br />
-**[[2wdt]] – PfUSP-L3 + Ub <br />
-*Ubiquitin thioesterase L5
-**[[3ris]] – hUSP catalytic domain  <BR />
-**[[3rii]], [[3a7s]] – hUSP catalytic domain (mutant)  <BR />
-**[[3tb3]] – hUSP UCH domain (mutant)  <BR />
-**[[3ihr]] – hUSP-L5 (mutant)  <br />
-**[[4uel]] – hUSP-L5 + polyUb + proteasomal Ub receptor Deubad domain  <br />
-**[[4uem]] – hUSP-L5 + proteasomal Ub receptor Deubad domain  <br />
-**[[4uf5]], [[4wlp]] – hUSP-L5 + nuclear factor Deubad domain  <br />
-**[[4uf6]] – hUSP-L5 + polyUb + nuclear factor Deubad domain  <br />
-**[[4wlq]] – mUSP-L5 + proteasomal Ub receptor C terminal  <br />
-**[[4wlr]] – mUSP-L5 + polyUb + proteasomal Ub receptor C terminal  <br />
-*Ubiquitin thioesterase ZranB1
-**[[3zrh]] – hUSP   <BR />
-**[[4s1z]] – bUSP  zinc finger + Ub <BR />
-*Ubiquitin thioesterase OTUB1
-**[[3von]] – hUSP + E2 <BR />
-**[[4ddg]], [[4ddi]] – hUSP + Ub<BR />
-**[[4i6l]] – hUSP (mutant) + Ub<BR />
-**[[4dhz]], [[4ldt]] – hUSP + E2 + Ub<BR />
-**[[4boq]] - hUSP OTU domain <BR />
-**[[4boz]] - hUSP OTU domain (mutant) + Ub <BR />
-**[[4bos]] - hUSP OTU domain (mutant) + Ub + OTUD2 peptide<BR />
-**[[4dhi]] – nUSP + E2 <BR />
-**[[2kzr]] – mUSP UBX-like domain – NMR<br />
-**[[4dhj]] – nUSP + E2 + Ub - nematode<BR />
-**[[4kdi]], [[4kdl]] – yUSP UBX-like domain + transitional endoplasmic reticulum ATPase<br />
-**[[3by4]], [[3c0r]] - yUSP OTU domain + Ub<BR />
-*USP OTUB2
-**[[5qiz]], [[5qiy]], [[5qix]], [[5qiw]], [[5qiu]], [[5qiv]], [[5qis]], [[5qit]], [[5qio]], [[5qip]], [[5qiq]], [[5qir]] – hUSP OTUB2 + ligand <BR />
-**[[4fjv]] - hUSP OTUB2 + Ub <BR />
-*USP OTULIN; Domains – OUT 80-345
-**[[6i9c]] – hUSP OTU domain (mutant) <BR />
-**[[5oe7]] – hUSP OUT domain + Ub <BR />
-**[[6sak]] – hUSP OUT domain (mutant) + sorting nexin-27<BR />
-*Ubiquitin thioesterase
-**[[3znh]] – USP OTU domain + Ub – Crimean-Congo hemorrhagic fever virus<BR />
-*Pseudomonas aeruginosa TE
-**[[2av9]], [[2o5u]], [[2o6b]], [[2o6t]], [[2o6u]], [[3qy3]] – PaTE – ''Pseudomonas aeruginosa''<br />
-**[[4qd7]] – PaTE hotdog domain<br />
-**[[4qda]], [[4qdb]] – PaTE hotdog domain (mutant)<br />
-**[[4qd8]], [[4dq9]] – PaTE hotdog domain + acyl-CoA derivatives<br />
-*Thioesterase
-**[[5vpj]] – TE – ''Actinomadura verrucosospora''<br />
-**[[5dio]], [[5byu]] – TE – ''Legionella pneumophila''<br />
-**[[6fvj]] – TE – ''Mycobacterium tuberculosis''<br />
-}}
 == References ==
 <references/>
 [[Category:Topic Page]]