Gene Details | GeDiPNet - Complete Gene Info, Disease Associations & Pathways

Gene Gene information from NCBI Gene database. Gene miRNA Gene ontology Other IDs Protein Pathways Associated diseases
Entrez ID	51377
Gene name	Ubiquitin C-terminal hydrolase L5
Gene symbol	UCHL5
Synonyms (NCBI Gene)	CGI-70INO80RUCH-L5UCH37
Chromosome	1
Chromosome location	1q31.2

194

Show/Hide all (194)

miRTarBase ID	miRNA	Experiments	Reference
MIRT665556	hsa-miR-1-5p	HITS-CLIP	23824327
MIRT665555	hsa-miR-6512-5p	HITS-CLIP	23824327
MIRT665554	hsa-miR-454-5p	HITS-CLIP	23824327
MIRT665553	hsa-miR-485-3p	HITS-CLIP	23824327
MIRT665552	hsa-miR-539-3p	HITS-CLIP	23824327
MIRT665551	hsa-miR-6883-3p	HITS-CLIP	23824327
MIRT665550	hsa-miR-4755-5p	HITS-CLIP	23824327
MIRT665549	hsa-miR-5006-3p	HITS-CLIP	23824327
MIRT665548	hsa-miR-4446-5p	HITS-CLIP	23824327
MIRT665547	hsa-miR-2681-5p	HITS-CLIP	23824327
MIRT456377	hsa-miR-5584-5p	PAR-CLIP	23592263
MIRT456376	hsa-miR-6750-5p	PAR-CLIP	23592263
MIRT456375	hsa-miR-6822-5p	PAR-CLIP	23592263
MIRT456374	hsa-miR-9500	PAR-CLIP	23592263
MIRT456373	hsa-miR-1322	PAR-CLIP	23592263
MIRT456372	hsa-miR-6872-3p	PAR-CLIP	23592263
MIRT456371	hsa-miR-3185	PAR-CLIP	23592263
MIRT456370	hsa-miR-3665	PAR-CLIP	23592263
MIRT456369	hsa-miR-657	PAR-CLIP	23592263
MIRT456368	hsa-miR-5581-3p	PAR-CLIP	23592263
MIRT665556	hsa-miR-1-5p	HITS-CLIP	23824327
MIRT665555	hsa-miR-6512-5p	HITS-CLIP	23824327
MIRT665554	hsa-miR-454-5p	HITS-CLIP	23824327
MIRT665553	hsa-miR-485-3p	HITS-CLIP	23824327
MIRT665552	hsa-miR-539-3p	HITS-CLIP	23824327
MIRT665551	hsa-miR-6883-3p	HITS-CLIP	23824327
MIRT665550	hsa-miR-4755-5p	HITS-CLIP	23824327
MIRT665549	hsa-miR-5006-3p	HITS-CLIP	23824327
MIRT665548	hsa-miR-4446-5p	HITS-CLIP	23824327
MIRT665547	hsa-miR-2681-5p	HITS-CLIP	23824327
MIRT456377	hsa-miR-5584-5p	PAR-CLIP	23592263
MIRT456376	hsa-miR-6750-5p	PAR-CLIP	23592263
MIRT456375	hsa-miR-6822-5p	PAR-CLIP	23592263
MIRT456374	hsa-miR-9500	PAR-CLIP	23592263
MIRT456373	hsa-miR-1322	PAR-CLIP	23592263
MIRT456372	hsa-miR-6872-3p	PAR-CLIP	23592263
MIRT456371	hsa-miR-3185	PAR-CLIP	23592263
MIRT456370	hsa-miR-3665	PAR-CLIP	23592263
MIRT456369	hsa-miR-657	PAR-CLIP	23592263
MIRT456368	hsa-miR-5581-3p	PAR-CLIP	23592263
MIRT1472400	hsa-let-7a	CLIP-seq
MIRT1472401	hsa-let-7b	CLIP-seq
MIRT1472402	hsa-let-7c	CLIP-seq
MIRT1472403	hsa-let-7d	CLIP-seq
MIRT1472404	hsa-let-7e	CLIP-seq
MIRT1472405	hsa-let-7f	CLIP-seq
MIRT1472406	hsa-let-7g	CLIP-seq
MIRT1472407	hsa-let-7i	CLIP-seq
MIRT1472408	hsa-miR-1183	CLIP-seq
MIRT1472409	hsa-miR-1193	CLIP-seq
MIRT1472410	hsa-miR-1229	CLIP-seq
MIRT1472411	hsa-miR-1265	CLIP-seq
MIRT1472412	hsa-miR-1273d	CLIP-seq
MIRT1472413	hsa-miR-1273f	CLIP-seq
MIRT1472414	hsa-miR-1294	CLIP-seq
MIRT1472415	hsa-miR-1324	CLIP-seq
MIRT1472416	hsa-miR-143	CLIP-seq
MIRT1472417	hsa-miR-1912	CLIP-seq
MIRT1472418	hsa-miR-192	CLIP-seq
MIRT1472419	hsa-miR-200b	CLIP-seq
MIRT1472420	hsa-miR-200c	CLIP-seq
MIRT1472421	hsa-miR-202	CLIP-seq
MIRT1472422	hsa-miR-2113	CLIP-seq
MIRT1472423	hsa-miR-215	CLIP-seq
MIRT1472424	hsa-miR-224	CLIP-seq
MIRT1472425	hsa-miR-2277-3p	CLIP-seq
MIRT1472426	hsa-miR-2392	CLIP-seq
MIRT1472427	hsa-miR-27a	CLIP-seq
MIRT1472428	hsa-miR-27b	CLIP-seq
MIRT1472429	hsa-miR-297	CLIP-seq
MIRT1472430	hsa-miR-29a	CLIP-seq
MIRT1472431	hsa-miR-29b	CLIP-seq
MIRT1472432	hsa-miR-29c	CLIP-seq
MIRT1472433	hsa-miR-3120-5p	CLIP-seq
MIRT1472434	hsa-miR-3144-3p	CLIP-seq
MIRT1472435	hsa-miR-3160-5p	CLIP-seq
MIRT1472436	hsa-miR-323b-3p	CLIP-seq
MIRT1472437	hsa-miR-329	CLIP-seq
MIRT1472438	hsa-miR-34b	CLIP-seq
MIRT1472439	hsa-miR-362-3p	CLIP-seq
MIRT1472440	hsa-miR-3647-3p	CLIP-seq
MIRT1472441	hsa-miR-3660	CLIP-seq
MIRT1472442	hsa-miR-3670	CLIP-seq
MIRT1472443	hsa-miR-3671	CLIP-seq
MIRT1472444	hsa-miR-371-5p	CLIP-seq
MIRT1472445	hsa-miR-371b-5p	CLIP-seq
MIRT1472446	hsa-miR-3941	CLIP-seq
MIRT1472447	hsa-miR-3973	CLIP-seq
MIRT1472448	hsa-miR-409-5p	CLIP-seq
MIRT1472449	hsa-miR-412	CLIP-seq
MIRT1472450	hsa-miR-4272	CLIP-seq
MIRT1472451	hsa-miR-4275	CLIP-seq
MIRT1472452	hsa-miR-4278	CLIP-seq
MIRT1472453	hsa-miR-4289	CLIP-seq
MIRT1472454	hsa-miR-429	CLIP-seq
MIRT1472455	hsa-miR-4307	CLIP-seq
MIRT1472456	hsa-miR-4316	CLIP-seq
MIRT1472457	hsa-miR-4458	CLIP-seq
MIRT1472458	hsa-miR-4500	CLIP-seq
MIRT1472459	hsa-miR-4502	CLIP-seq
MIRT1472460	hsa-miR-4519	CLIP-seq
MIRT1472461	hsa-miR-4526	CLIP-seq
MIRT1472462	hsa-miR-4652-3p	CLIP-seq
MIRT1472463	hsa-miR-4668-3p	CLIP-seq
MIRT1472464	hsa-miR-4677-3p	CLIP-seq
MIRT1472465	hsa-miR-4679	CLIP-seq
MIRT1472466	hsa-miR-4699-3p	CLIP-seq
MIRT1472467	hsa-miR-4708-5p	CLIP-seq
MIRT1472468	hsa-miR-4716-3p	CLIP-seq
MIRT1472469	hsa-miR-4719	CLIP-seq
MIRT1472470	hsa-miR-4735-5p	CLIP-seq
MIRT1472471	hsa-miR-4760-5p	CLIP-seq
MIRT1472472	hsa-miR-4762-3p	CLIP-seq
MIRT1472473	hsa-miR-4770	CLIP-seq
MIRT1472474	hsa-miR-4775	CLIP-seq
MIRT1472475	hsa-miR-4789-5p	CLIP-seq
MIRT1472476	hsa-miR-4795-3p	CLIP-seq
MIRT1472477	hsa-miR-487a	CLIP-seq
MIRT1472478	hsa-miR-500a	CLIP-seq
MIRT1472479	hsa-miR-513a-3p	CLIP-seq
MIRT1472480	hsa-miR-513a-5p	CLIP-seq
MIRT1472481	hsa-miR-513c	CLIP-seq
MIRT1472482	hsa-miR-514b-5p	CLIP-seq
MIRT1472483	hsa-miR-520g	CLIP-seq
MIRT1472484	hsa-miR-520h	CLIP-seq
MIRT1472485	hsa-miR-548ac	CLIP-seq
MIRT1472486	hsa-miR-548ae	CLIP-seq
MIRT1472487	hsa-miR-548aj	CLIP-seq
MIRT1472488	hsa-miR-548am	CLIP-seq
MIRT1472489	hsa-miR-548c-3p	CLIP-seq
MIRT1472490	hsa-miR-548d-3p	CLIP-seq
MIRT1472491	hsa-miR-548p	CLIP-seq
MIRT1472492	hsa-miR-548x	CLIP-seq
MIRT1472493	hsa-miR-548z	CLIP-seq
MIRT1472494	hsa-miR-556-3p	CLIP-seq
MIRT1472495	hsa-miR-556-5p	CLIP-seq
MIRT1472496	hsa-miR-567	CLIP-seq
MIRT1472497	hsa-miR-584	CLIP-seq
MIRT1472498	hsa-miR-587	CLIP-seq
MIRT1472499	hsa-miR-590-3p	CLIP-seq
MIRT1472500	hsa-miR-603	CLIP-seq
MIRT1472501	hsa-miR-616	CLIP-seq
MIRT1472502	hsa-miR-624	CLIP-seq
MIRT1472503	hsa-miR-634	CLIP-seq
MIRT1472504	hsa-miR-651	CLIP-seq
MIRT1472505	hsa-miR-660	CLIP-seq
MIRT1472506	hsa-miR-671-5p	CLIP-seq
MIRT1472507	hsa-miR-758	CLIP-seq
MIRT1472508	hsa-miR-875-3p	CLIP-seq
MIRT1472509	hsa-miR-875-5p	CLIP-seq
MIRT1472510	hsa-miR-889	CLIP-seq
MIRT1472511	hsa-miR-98	CLIP-seq
MIRT2142311	hsa-miR-3136-5p	CLIP-seq
MIRT2142312	hsa-miR-4439	CLIP-seq
MIRT2142313	hsa-miR-4513	CLIP-seq
MIRT2142314	hsa-miR-4760-3p	CLIP-seq
MIRT2142315	hsa-miR-943	CLIP-seq
MIRT2142316	hsa-miR-630	CLIP-seq
MIRT2362772	hsa-miR-1825	CLIP-seq
MIRT2362773	hsa-miR-548u	CLIP-seq
MIRT1472501	hsa-miR-616	CLIP-seq
MIRT2462832	hsa-miR-3121-3p	CLIP-seq
MIRT2462833	hsa-miR-339-5p	CLIP-seq
MIRT2462834	hsa-miR-3607-3p	CLIP-seq
MIRT1472411	hsa-miR-1265	CLIP-seq
MIRT2656886	hsa-miR-1282	CLIP-seq
MIRT2656887	hsa-miR-132	CLIP-seq
MIRT1472422	hsa-miR-2113	CLIP-seq
MIRT2656888	hsa-miR-212	CLIP-seq
MIRT1472426	hsa-miR-2392	CLIP-seq
MIRT2656889	hsa-miR-298	CLIP-seq
MIRT2656890	hsa-miR-3177-5p	CLIP-seq
MIRT1472440	hsa-miR-3647-3p	CLIP-seq
MIRT2656891	hsa-miR-3944-5p	CLIP-seq
MIRT2656892	hsa-miR-455-5p	CLIP-seq
MIRT2656893	hsa-miR-4705	CLIP-seq
MIRT1472475	hsa-miR-4789-5p	CLIP-seq
MIRT2656894	hsa-miR-518a-5p	CLIP-seq
MIRT2656895	hsa-miR-527	CLIP-seq
MIRT2656896	hsa-miR-577	CLIP-seq
MIRT1472498	hsa-miR-587	CLIP-seq
MIRT1472499	hsa-miR-590-3p	CLIP-seq
MIRT1472504	hsa-miR-651	CLIP-seq
MIRT1472411	hsa-miR-1265	CLIP-seq
MIRT1472426	hsa-miR-2392	CLIP-seq
MIRT2656889	hsa-miR-298	CLIP-seq
MIRT1472440	hsa-miR-3647-3p	CLIP-seq
MIRT2656891	hsa-miR-3944-5p	CLIP-seq
MIRT2656892	hsa-miR-455-5p	CLIP-seq
MIRT2656893	hsa-miR-4705	CLIP-seq
MIRT1472475	hsa-miR-4789-5p	CLIP-seq
MIRT2656896	hsa-miR-577	CLIP-seq
MIRT1472498	hsa-miR-587	CLIP-seq
MIRT1472504	hsa-miR-651	CLIP-seq

Show/Hide all (54)

GO ID	Ontology	Definition	Evidence	Reference
GO:0000502	Component	Proteasome complex	IEA
GO:0000723	Process	Telomere maintenance	IEA
GO:0000723	Process	Telomere maintenance	ISO
GO:0003723	Function	RNA binding	HDA	22658674
GO:0004843	Function	Cysteine-type deubiquitinase activity	IBA
GO:0004843	Function	Cysteine-type deubiquitinase activity	IDA	18922472
GO:0004843	Function	Cysteine-type deubiquitinase activity	IEA
GO:0004843	Function	Cysteine-type deubiquitinase activity	IMP	28992318
GO:0004866	Function	Endopeptidase inhibitor activity	IMP	18162577
GO:0005515	Function	Protein binding	IPI	11163772, 16990800, 17139257, 18922472, 19490896, 19615732, 21303910, 21516116, 25416956, 27107012, 31515488, 32296183, 33961781, 35271311
GO:0005634	Component	Nucleus	IDA	18922472
GO:0005634	Component	Nucleus	IEA
GO:0005654	Component	Nucleoplasm	IDA
GO:0005654	Component	Nucleoplasm	TAS
GO:0005730	Component	Nucleolus	IDA
GO:0005737	Component	Cytoplasm	IBA
GO:0005737	Component	Cytoplasm	IEA
GO:0005829	Component	Cytosol	IDA	18922472
GO:0005829	Component	Cytosol	TAS
GO:0006275	Process	Regulation of DNA replication	IMP	25016522
GO:0006281	Process	DNA repair	IEA
GO:0006282	Process	Regulation of DNA repair	IEA
GO:0006282	Process	Regulation of DNA repair	ISO
GO:0006310	Process	DNA recombination	IEA
GO:0006338	Process	Chromatin remodeling	IDA	21303910
GO:0006508	Process	Proteolysis	IEA
GO:0006511	Process	Ubiquitin-dependent protein catabolic process	IEA
GO:0006974	Process	DNA damage response	IEA
GO:0008233	Function	Peptidase activity	IEA
GO:0008234	Function	Cysteine-type peptidase activity	IEA
GO:0016579	Process	Protein deubiquitination	IDA	18922472
GO:0016579	Process	Protein deubiquitination	IEA
GO:0016787	Function	Hydrolase activity	IEA
GO:0021670	Process	Lateral ventricle development	IEA
GO:0030901	Process	Midbrain development	IEA
GO:0031011	Component	Ino80 complex	IDA	18922472, 21303910
GO:0031011	Component	Ino80 complex	IEA
GO:0032435	Process	Negative regulation of proteasomal ubiquitin-dependent protein catabolic process	IMP	23770237
GO:0033044	Process	Regulation of chromosome organization	IBA
GO:0033044	Process	Regulation of chromosome organization	IMP	26340092
GO:0045739	Process	Positive regulation of DNA repair	IEA
GO:0045739	Process	Positive regulation of DNA repair	ISO
GO:0045880	Process	Positive regulation of smoothened signaling pathway	IMP	28992318
GO:0045893	Process	Positive regulation of DNA-templated transcription	IMP	27641337
GO:0045995	Process	Regulation of embryonic development	IEA
GO:0045995	Process	Regulation of embryonic development	ISO
GO:0048853	Process	Forebrain morphogenesis	IEA
GO:0051726	Process	Regulation of cell cycle	IMP	26340092
GO:0060382	Process	Regulation of DNA strand elongation	IMP	25016522
GO:0061136	Process	Regulation of proteasomal protein catabolic process	IMP	18162577
GO:0070628	Function	Proteasome binding	IDA	18162577, 18922472
GO:0070628	Function	Proteasome binding	IEA
GO:1904507	Process	Positive regulation of telomere maintenance in response to DNA damage	IEA
GO:1904507	Process	Positive regulation of telomere maintenance in response to DNA damage	ISO

MIM	HGNC	e!Ensembl
610667	19678	ENSG00000116750

Q9Y5K5

Protein name

Ubiquitin carboxyl-terminal hydrolase isozyme L5 (UCH-L5) (EC 3.4.19.12) (Ubiquitin C-terminal hydrolase UCH37) (Ubiquitin thioesterase L5)

Protein function

Protease that specifically cleaves 'Lys-48'-linked polyubiquitin chains. Deubiquitinating enzyme associated with the 19S regulatory subunit of the 26S proteasome. Putative regulatory component of the INO80 complex; however is inactive in the INO

PDB

3A7S , 3IHR , 3RII , 3RIS , 3TB3 , 4UEL , 4UEM , 4UF5 , 4UF6 , 4WLP

Family and domains

Pfam

Accession	ID	Position in sequence	Description	Type
PF01088	Peptidase_C12	8 → 209	Ubiquitin carboxyl-terminal hydrolase, family 1	Domain
PF18031	UCH_C	264 → 309	Ubiquitin carboxyl-terminal hydrolases	Domain

Sequence

MTGNAGEWCLMESDPGVFTELIKGFGCRGAQVEEIWSLEPENFEKLKPVHGLIFLFKWQP
GEEPAGSVVQDSRLDTIFFAKQVINNACATQAIVSVLLNCTHQDVHLGETLSEFKEFSQS
FDAAMKGLALSNSDVIRQVHNSFARQQMFEFDTKTSAKEEDAFHFVSYVPVNGRLYELDG
LREGPIDLGACNQDDWISAVRPVIEKRIQKYSEGEIRFNLMAIVSDRKMIYEQKIAELQR
QLAEEEPMDTDQGNSMLSAIQSEVAKNQMLIEEEVQKLKRYKIENIRRKHNYLPFIMELL
KTLAEHQQLIPLVEKAKEKQNAKKAQETK

Sequence length

329

Interactions

View interactions

	KEGG		Reactome
	ATP-dependent chromatin remodeling		UCH proteinases

Evidence Score: ★☆☆☆☆ Gene-disease association found in Text Mining only ★★☆☆☆ Found in Text Mining and Unknown/Other Associations ★★★☆☆ Reported in Unknown/Other Associations across ≥2 Sources ★★★★☆ ClinVar: Pathogenic/Likely Pathogenic (<5 Variants) ★★★★★ ClinVar: Pathogenic/Likely Pathogenic (≥5 Variants)

Show/Hide Unknown Diseases (1)

Phenotype Name	Clinical Significance	Source	Evidence Score
Unknown / Other Associations ClinVar entries with uncertain/conflicting evidence, and associations from other databases (OMIM, Orphanet, GWAS, etc.) where the gene is not established as causal.
CANNABIS DEPENDENCE	—	GWAS catalog	★★★★★ ★★☆☆☆ Found in Text Mining + Unknown/Other Associations

Show/Hide Text Mining Associations (28)

Disease Name	Disease (Merged)	Source	PMID	Relationship Type	Evidence Score
Associations from Text Mining Disease associations identified through text mining
Adenocarcinoma of lung (disorder)	Lung adenocarcinoma	BEFREE	30527624		★★★★★ ★☆☆☆☆ Found in Text Mining only
Alzheimer Disease	Alzheimer disease	Pubtator	24348898	Associate	★★★★★ ★☆☆☆☆ Found in Text Mining only
Carcinogenesis	Carcinogenesis	Pubtator	23220010	Associate	★★★★★ ★☆☆☆☆ Found in Text Mining only
Carcinoma Hepatocellular	Hepatocellular carcinoma	Pubtator	23220010	Associate	★★★★★ ★☆☆☆☆ Found in Text Mining only
Carcinoma Hepatocellular	Hepatocellular carcinoma	Pubtator	32486284	Stimulate	★★★★★ ★☆☆☆☆ Found in Text Mining only
Carcinoma Ovarian Epithelial	Epithelial ovarian carcinoma	Pubtator	25123264	Associate	★★★★★ ★☆☆☆☆ Found in Text Mining only
Carcinoma Pancreatic Ductal	Pancreatic ductal carcinoma	Pubtator	28653876, 29474458	Associate	★★★★★ ★☆☆☆☆ Found in Text Mining only
Carcinoma, Ovarian Epithelial	Ovarian Epithelial carcinoma	BEFREE	31666925		★★★★★ ★☆☆☆☆ Found in Text Mining only
Colon Carcinoma	Colon Carcinoma	BEFREE	30149015		★★★★★ ★☆☆☆☆ Found in Text Mining only
Colorectal Carcinoma	Colorectal Cancer	BEFREE	28681694		★★★★★ ★☆☆☆☆ Found in Text Mining only
Diffuse Large B-Cell Lymphoma	Diffuse Lymphoma	BEFREE	27301795		★★★★★ ★☆☆☆☆ Found in Text Mining only
Leukemia Myeloid Acute	Myeloid leukemia	Pubtator	39408703	Associate	★★★★★ ★☆☆☆☆ Found in Text Mining only
Liver carcinoma	Liver carcinoma	BEFREE	23220010, 29377617		★★★★★ ★☆☆☆☆ Found in Text Mining only
Malignant neoplasm of ovary	Ovarian cancer	BEFREE	31666925		★★★★★ ★☆☆☆☆ Found in Text Mining only
Malignant Neoplasms	Malignant Neoplasm	BEFREE	22615012, 28598414, 28653876, 29698671, 31666925		★★★★★ ★☆☆☆☆ Found in Text Mining only
Malignant tumor of colon	Colonic Neoplasms	BEFREE	30149015		★★★★★ ★☆☆☆☆ Found in Text Mining only
Multiple Myeloma	Multiple myeloma	BEFREE	24319254, 31639138		★★★★★ ★☆☆☆☆ Found in Text Mining only
Multiple Myeloma	Multiple myeloma	Pubtator	33531688	Associate	★★★★★ ★☆☆☆☆ Found in Text Mining only
Neoplasms	Neoplasms	BEFREE	22057347		★★★★★ ★☆☆☆☆ Found in Text Mining only
Non-Small Cell Lung Carcinoma	Lung carcinoma	BEFREE	30165348		★★★★★ ★☆☆☆☆ Found in Text Mining only
ovarian neoplasm	Ovarian neoplasm	BEFREE	31666925		★★★★★ ★☆☆☆☆ Found in Text Mining only
Pancreatic Ductal Adenocarcinoma	Pancreatic adenocarcinoma	BEFREE	28653876, 29474458		★★★★★ ★☆☆☆☆ Found in Text Mining only
Rectal Carcinoma	Rectal Carcinoma	BEFREE	28681694, 29474458		★★★★★ ★☆☆☆☆ Found in Text Mining only
Squamous cell carcinoma of esophagus	Esophagus Neoplasm	BEFREE	22615012, 31342239		★★★★★ ★☆☆☆☆ Found in Text Mining only
Stomach Neoplasms	Stomach neoplasms	Pubtator	29474458, 32351584	Associate	★★★★★ ★☆☆☆☆ Found in Text Mining only
Uterine Cervical Neoplasm	Uterine cervix neoplasm	LHGDN	16402389		★★★★★ ★☆☆☆☆ Found in Text Mining only
Waldenstrom Macroglobulinemia	Waldenstrom`s Macroglobulinemia	BEFREE	25691154, 27813535, 27825468		★★★★★ ★☆☆☆☆ Found in Text Mining only
Waldenstrom Macroglobulinemia	Waldenstrom macroglobulinemia	Pubtator	25691154	Associate	★★★★★ ★☆☆☆☆ Found in Text Mining only

UCHL5 (ubiquitin C-terminal hydrolase L5)