Eukaryotic NAT genes

PROTISTS: See table 1

FUNGI: See table 2

PLANTS: No NAT genes identified to date

ANIMALS: See table 3

Table 1: Annotation of NAT genes in sequenced genomes of protists^(a),(b)

Taxon	Organism scientific name^(c)	Organism identification code	Taxonomy identifier	Gene	ORF (nt)	Protein (aa)	Exon span	Number of introns	Locus tag	TPA accession number
Rhizaria (1)	None
Cryptophyta (2)	None
Heterolobosea (1)	Naegleria gruberi NEG-M	NAEGR	5762	NAT1	969	322	1 (1-969)	0	NAEGRDRAFT_80760	BN001457
				NAT2	990	329	1 (1-990)	0	NAEGRDRAFT_56390	BN001458
Amoebozoa (5)	Entamoebidae (3)
	None
	Dictyosteliida (2)
	Dictyostelium discoideum AX4	DICDI	352472	NAT1	921	306	1 (1-310) 2 (410-1020)	1	DDB_G0288509	BN001450
				NAT2	978	325	1 (1-301) 2 (415-1091)	1	DDB_G0288503	BN001451
				NAT3	900	299	1 (1-310) 2 (402-991)	1	DDB_G0288507	BN001452
				NAT4	837	278	1 (1-837)	0	DDB_G0287121	BN001453
				NAT5	870	289	1 (1-870)	0	DDB_G0276993	BN001454
				NAT6	1464	487	1 (1-256) 2 (324-784) 3 (1959-2220) 4 (2321-2776) 5 (2805-2833)	4	DDB_G0277549	BN001455
	Polysphondylium pallidum PN500	POLPA	670386	NAT1	822	273	1 (1-822)	0	PPL_12417	BN001460
				NAT2	777	258	1 (1-562) 2 (623-837)	1	PPL_12418	BN001461
Parabasalia (1)	None
Stramenopiles (7)	Oomycetes (5)
	Saprolegnia parasitica CBS 223.65	SAPPC	695850	NAT1	951	316	1 (1-951)	0		BN001462
				NAT2	933	310	1 (1-933)	0		BN001463
	Bacillariophyta (diatoms) (2)
	Phaeodactylum tricornutum CCAP 1055/1	PHATR	556484	NAT1	981	326	1 (1-981)	0		BN001459
Alveolata (18)	Perkinsea (1)
	None
	Apicomplexa (15)
	None
	Ciliophora (2)
	Paramecium tetraurelia d4-2	PARTE	412030	NAT1	861	286	1 (1-861)	0	GSPATT00024900001	BN001464
				NAT2	861	286	1 (1-861)	0	GSPATT00029589001	BN001465
				NAT3	867	288	1 (1-867)	0	GSPATT00029590001	BN001466
				NAT4	441	146	1 (441)	0	GSPATT00029592001	BN001467
	Tetrahymena thermophila SB210	TETTH	312017	NAT1	858	285	1 (1-858)	0	TTHERM_00616140	BN001468
				NAT2	873	290	1 (1-873)	0	TTHERM_00408820	BN001469
				NAT3	885	294	1 (1-491) 2 (605-758) 3 (1905-2001) 4 (2276-2418)	3	TTHERM_00693000	BN001470
				NAT4	861	286	1 (1-861)	0	TTHERM_01222540	BN001471
				NAT5	858	285	1 (1-858)	0	TTHERM_00888050	BN001472
				NAT6	843	280	1 (1-843)	0	TTHERM_00335770	BN001473
Fornicata (1)	None
Euglenozoa (5)	None
Ichthyosporea (1)	None
Choanoflagellida (1)	Monosiga brevicollis MX1	MONBE	431895	NAT1	1029	342	1 (1-1029)	0	MONBRDRAFT_7289	BN001456
Chlorophyta (2)	Chlamydomonas reinhardtii CC-503 cw92 mt+	CHLRE	3055	NAT1	924 (partial 3' end)	308 (partial C-terminus)	1 (1-99) 2 (390-470) 3 (757-822) 4 (1639-1698) 5 (1862-1942) 6 (2429-2491) 7 (3163-3255) 8 (3760-3821) 9 (4573-4704) 10 (5094-5118) 11 (5911-6051) 12 (6339-6504) 13 (7099-7145)	12	CHLREDRAFT_187400	BN001479

Footnotes, Table 1:

^(a) The details and provisional taxonomic classification of the sequenced genomes of protist species are available on http://www.ncbi.nlm.nih.gov/genomes/leuks.cgi. The term "protists" is used to describe a diverse range of lower eukaryotic organisms and does not refer to any particular monophyletic taxonomic group (http://www.discoverlife.org/mp/20m?tree=Protists&res=1000). The appropriate reference sequence for each taxonomic group of protists should be requested from the NAT Gene Nomenclature Committee.

^(b) Column key – Columns 1, 2: The numbers in brackets are sequenced genomes of protist species accessible via BLAST (2010); Columns 2-4: The scientific name, official identification code and unique taxonomy identifier for each organism are available from the UniProt Taxonomy Browser (http://www.uniprot.org/taxonomy/); Columns 5-9: The retrieval and annotation of NAT-homologous sequences (ORF span and exon-intron structure) were carried out as described [41], and the gene symbols were assigned according to the guidelines of the NAT Gene Nomenclature Committee. The sequence length of open reading frames (ORF) is shown in nucleotides (nt) and of deduced proteins in amino acids (aa); Column 10: Where available, the locus tags are provided to anchor each annotated NAT gene to the corresponding genomic map; Column 11: Third-party annotation (TPA) accession numbers were assigned by the EMBL database and can be used to retrieve the complete details of NAT genes in the table.

^(c) Search of the EST database additionally identified partial NAT-homologous sequences in the haptophyte Emiliania huxleyi (dbEST ID: FP203643), the alveolate Karenia brevis (dbEST ID: EX962146), the amoebozoon Acanthamoeba castellanii (dbEST ID: EC098681) and the choanoflagellate Monosiga ovata (dbEST ID: DC473432).

Table 2: Annotation of NAT genes in sequenced genomes of fungi^(a),(b)

Phylum	Organism scientific name	Organism identification code	Taxonomy identifier	Gene	ORF (nt)	Protein (aa)	Exon span	Number of introns	Locus tag	TPA accession number
Microsporidia (4)	NONE
Chytridiomycota (2)	Batrachochytrium dendrobatidis	BATDE	109871	NAT1	897	298	1 (1-891) 2 (981-986)	1	BDEG_00384	BN001378
Blastocladiomycota (1)	NONE
Zygomycota (1)	NONE
Basidiomycota (16)	Pucciniomycotina (2)
	Puccinia graminis f. sp. tritici CRL 75-36-700-3	PUCGR	418459	ΝΑΤ1	1008	335	1 (1-345) 2 (430-837) 3 (986-1240)	2	PGTG_03435	BN001423
	Ustilaginomycotina (4)
	Malassezia globosa CBS 7966	MALGO	425265	ΝΑΤ1	834	277	1 (1-834)	0	MGL_4060	BN001438
	Ustilago maydis 521	USTMA	237631	ΝΑΤ1	1119	372	1 (1-102) 2 (177-1193)	1	UM01576	BN001424
	Agaricomycotina (10)
	Coprinus cinereus (Coprinopsis cinerea) okayama 7-130	COPC7	240176	ΝΑΤ1	1089	362	1 (1-233) 2 (285-414) 3 (464-725) 4 (784-1247)	3	CC1G_07781	BN001421
				ΝΑΤ2	1089	362	1 (1-230) 2 (287-415) 3 (493-749) 4 (831-1303)	3	CC1G_07803	BN001422
	Moniliophthora perniciosa FA553	CRIPE	554373	NAT1	459 (partial 3' end)	153 (partial C-terminus)	1 (1-459)	0		BN001437
Ascomycota (122)	Taphrinomycotina (4)
	NONE
	Saccharomycotina (40)
	NONE
	Pezizomycotina (78)
	Ajellomyces capsulata G186AR	AJECG	447093	ΝΑΤ1	963	320	1 (1-402) 2 (534-580) 3 (644-1157)	2		BN001360
	Ajellomyces capsulata H88	AJEC8	544711	ΝΑΤ1	963	320	1 (1-402) 2 (534-580) 3 (645-1158)	2	HCEG_02416	BN001361
	Ajellomyces capsulata H143	AJECH	544712	ΝΑΤ1	963	320	1 (1-402) 2 (534-580) 3 (645-1158)	2	HCDG_05981	BN001362
	Ajellomyces capsulata G217B	AJECA	447094	ΝΑΤ1	963	320	1 (1-402) 2 (534-580) 3 (645-1158)	2		BN001442
	Ajellomyces capsulata NAm1	AJECN	339724	ΝΑΤ1	963	320	1 (1-402) 2 (534-580) 3 (645-1158)	2	HCAG_01594	BN001443
	Ajellomyces dermatitidis SLH14081	AJEDS	559298	ΝΑΤ1	963	320	1 (1-402) 2 (538-584) 3 (659-1172)	2	BDBG_03471	BN001364
	Ajellomyces dermatitidis ER-3	AJEDR	559297	ΝΑΤ1	963	320	1 (1-402) 2 (538-584) 3 (659-1172)	2	BDCG_08161	BN001363
	Alternaria brassicicola ATCC 96836	ALTBR	622360	ΝΑΤ1	924	307	1 (1-924)	0		BN001365
	Aspergillus clavatus NRRL 1	ASPCL	344612	ΝΑΤ1	963	320	1 (1-410) 2 (462-507) 3 (581-1087)	2	ACLA_050060	BN001369
	Aspergillus flavus NRRL-3357	ASPFN	332952	ΝΑΤ1	1017	338	1 (1-14) 2 (863-1865)	1		BN001444
				ΝΑΤ2	981	326	1 (1-426) 2 (482-528) 3 (588-1095)	2	AFL2G_01915	BN001445
				ΝΑΤ3	957	318	1 (1-396) 2 (450-1010)	1	AFL2G_11316	BN001446
				ΝΑΤ4	641	possible pseudogene	1 (1-92) 2 (144-692)	1		BN001447
	Aspergillus fumigatus CEA 10 / Af293	ASPFC	451804	ΝΑΤ1	990	329	1 (1-410) 2 (440-514) 3 (574-1078)	2	Afu4g10930	BN001370
	Aspergillus niger CBS 513.88 / ATCC 1015	ASPNC	425011	ΝΑΤ1	1047	348	1 (1-1047)	0	fge1_pg_C_12000355	BN001371
				ΝΑΤ2	990	329	1 (1-441) 2 (508-1056)	1	fge1_pm_C_7000250	BN001372
				ΝΑΤ3	948	315	1 (1-397) 2 (491-536) 3 (592-1096)	2	fge1_pg_C_17000131	BN001373
				ΝΑΤ4	1050	349	1 (1-539) 2 (600-1110)	1	fge1_pg_C_12000261	BN001374
				ΝΑΤ5	975	324	1 (1-975)	0		BN001375
	Aspergillus terreus NIH 2624	ASPTN	341663	NAT1	957	318	1 (1-411) 2 (493-539) 3 (593-1091)	2	ATEG_03982	BN001376
				NAT2	969	322	1 (1-393) 2 (448-1023)	1	ATEG_05747	BN001377
	Botryotinia fuckeliana B05.10	BOTFB	332648	NAT1	981	326	1 (1-380) 2 (470-1070)	1		BN001379
	Chaetomium globosum CBS 148.51	CHAGB	306901	NAT1	927	308	1 (1-348) 2 (417-995)	1	CHGG_09458	BN001380
				NAT2	984	327	1 (1-984)	0	CHGG_06975	BN001381
	Coccidioides immitis H538.4	COCIT	396776	NAT1	984	327	1 (1-411) 2 (473-1045)	1	CIHG_06736	BN001383
	Coccidioides immitis RS	COCIM	246410	NAT1	984	327	1 (1-411) 2 (473-1045)	1	CIMG_01522	BN001384
	Coccidioides immitis RMSCC 3703	COCIT	454286	NAT1	984	327	1 (1-411) 2 (473-1045)	1	CISG_01266	BN001385
	Coccidioides immitis RMSCC 2394	COCIT	404692	NAT1	984	327	1 (1-411) 2 (473-1045)	1	CIRG_01630	BN001382
	Coccidioides posadasii C735	COCP7	222929	NAT1	984	327	1 (1-411) 2 (473-1045)	1		BN001440
	Coccidioides posadasii RMSCC 3488	COCPO	454284	NAT1	984	327	1 (1-411) 2 (473-1045)	1	CPAG_01321	BN001441
	Coccidioides posadasii RMSCC 2133	COCPO	469470	NAT1	984	327	1 (1-411) 2 (473-1045)	1		BN001386
	Coccidioides posadasii CPA 0020	COCPO	490068	NAT1	984	327	1 (1-411) 2 (473-1045)	1		BN001387
	Coccidioides posadasii RMSCC 1038	COCPO	490066	NAT1	984	327	1 (1-411) 2 (473-1045)	1		BN001388
	Coccidioides posadasii CPA 0001	COCPO	469472	NAT1	984	327	1 (1-411) 2 (473-1045)	1		BN001389
	Coccidioides posadasii RMSCC 1037	COCPO	490065	NAT1	984	327	1 (1-411) 2 (473-1045)	1		BN001390
	Coccidioides posadasii str. Silveira	COCPS	443226	NAT1	984	327	1 (1-411) 2 (473-1045)	1	CPSG_01649	BN001391
	Emericella nidulans (Aspergillus nidulans) FGSC A4	EMENI	227321	ΝΑΤ1	960	319	1 (1-407) 2 (463-510) 3 (568-1072)	2	ANID_10723	BN001425
	Fusarium oxysporum f. sp. Lycopersici 4287	FUSO4	426428	ΝΑΤ1	1053	350	1 (1-1053)	0	FOXG_15318	BN001426
				ΝΑΤ2	957	318	1 (1-372) 2 (424-1008)	1	FOXG_06095	BN001427
				ΝΑΤ3	999	332	1 (1-999)	0	FOXG_03795	BN001428
				ΝΑΤ4	963	320	1 (1-366) 2 (549-1145)	1		BN001429
	Gibberella moniliformis (Fusarium verticillioides) 7600	GIBM7	334819	ΝΑΤ1	1038	345	1 (1-1038)	0	FVEG_12636	BN001430
				ΝΑΤ2	957	318	1 (1-957)	0	FVEG_03961	BN001431
				ΝΑΤ3	978	325	1 (1-978)	0	FVEG_12062	BN001432
				ΝΑΤ4	1011	336	1 (1-295) 2 (628-1024) 3 (1079-1397)	2		BN001433
	Gibberella zeae (Fusarium graminearum) RRL31084	GIBZE	229533	ΝΑΤ1	1032	343	1 (1-1032)	0	FGSG_00080	BN001434
				ΝΑΤ2	957	318	1 (1-372) 2 (431-1015)	1	FGSG_09400	BN001435
				ΝΑΤ3	960	319	1 (1-960)	0	FGSG_07888	BN001436
	Glomerella graminicola M1.001	COLGM	645133	NAT1	927	308	1 (1-360) 2 (425-991)	1		BN001392
	Magnaporthe grisea 70-15	MAGO7	242507	ΝΑΤ1	963	320	1 (1-354) 2 (454-1062)	1	MGG_10560	BN001393
				ΝΑΤ2	1113	370	1 (1-1113)	0		BN001394
	Microsporum canis CBS 113480	ARTOC	554155	ΝΑΤ1	1002	333	1 (1-422) 2 (486- 533) 3 (589-1120)	2	MCYG_06389	BN001368
	Microsporum gypseum CBS 118893	ARTGP	535722	ΝΑΤ1	942	313	1 (1-416) 2 (585-1110)	1	MGYG_06842	BN001366
				ΝΑΤ2	1008	335	1 (1-414) 2 (534-1127)	1	MGYG_02694	BN001367
	Nectria haematococca mpVI 77-13-4	NECH7	660122	NAT1	891	296	1 (1-331) 2 (403-962)	1	NECHADRAFT_33380	BN001395
				NAT2	966	321	1 (1-381) 2 (435-1019)	1	NECHADRAFT_75515	BN001396
				NAT3	897	298	1 (1-344) 2 (401-953)	1	NECHADRAFT_87499	BN001397
				NAT4	1008	335	1 (1-1008)	0	NECHADRAFT_34602	BN001398
				NAT5	930	309	1 (1-366) 2 (418-563) 3 (588-1005)	2	NECHADRAFT_44961	BN001399
NAT6				960	319	1 (1-366) 2 (413-1006)	1	NECHADRAFT_51017	BN001400
Neosartorya fischeri (Aspergillus fischerianus) NRRL 181	NEOFI	331117	ΝΑΤ1	963	320	1 (1-410) 2 (467-514) 3 (574-1078)	2	NFIA_104890	BN001401
			ΝΑΤ2	948	315	1 (1-378) 2 (436-1005)	1	NFIA_004340	BN001402
Neurospora crassa OR74A	NEUCR	367110	ΝΑΤ1	933	310	1 (1-333) 2 (402-1001)	1	NCU09784	BN001448
Paracoccidioides brasiliensis Pb01	PARBA	502779	ΝΑΤ1	951	possible pseudogene	1 (1-402) 2 (527-573) 3 (641-1142)	2		BN001404
Paracoccidioides brasiliensis Pb03	PARBP	482561	ΝΑΤ1	951	316	1 (1-402) 2 (528-574) 3 (642-1143)	2	PABG_03148	BN001403
Paracoccidioides brasiliensis Pb18	PARBD	502780	ΝΑΤ1	951	316	1 (1-402) 2 (527-573) 3 (641-1142)	2	PADG_01689	BN001405
Penicillium chrysogenum Wisconsin 54-1255	PENCW	500485	NAT1	942	313	1 (1-396) 2 (522-1067)	1	Pc20g04880	BN001474
			NAT2	972	323	1 (1-401) 2 (497-1067)	1	Pc22g06840	BN001475
			NAT3	948	315	1 (1-396) 2 (447-998)	1	Pc12g02960	BN001476
Phaeosphaeria nodorum SN15	PHANO	321614	ΝΑΤ1	933	310	1 (1-933)	0	SNOG_06352	BN001406
			ΝΑΤ2	1053	350	1 (1-1053)	0	SNOG_06959	BN001407
Podospora anserina DSM980	PODAN	515849	NAT1	912	303	1 (1-342) 2 (401-970)	1	PODANSg3401
			NAT2	1002	333	1(1-1002)	0	PODANSg9179
Pyrenophora tritici-repentis Pt-1C-BFP	PYRTR	426418	ΝΑΤ1	1029	342	1 (1-1029)	0	PTRG_06864	BN001408
			ΝΑΤ2	924	307	1 (1-924)	0	PTRG_02465	BN001409
Sordaria macrospora k-hell	SORMA	5147	NAT1	1101	366	1 (1-477) 2 (536-1159)	1		BN001439
Talaromyces stipitatus ATCC 10500	TALSN	441959	ΝΑΤ1	993	330	1 (1-438) 2 (555-1109)	1	TSTA_053130	BN001410
Trichoderma atroviride IMI 206040	HYPAI	452589	ΝΑΤ1	915	304	1 (1-339) 2 (417-992)	1		BN001411
Trichoderma reesei (Hypocrea jecorina) QM6a	HYPJQ	431241	ΝΑΤ1	918	305	1 (1-336) 2 (430-1011)	1		BN001412
Trichoderma virens (Hypocrea virens) Gv29-8	HYPVG	413071	ΝΑΤ1	918	305	1 (1-333) 2 (426-1010)	1		BN001413
			ΝΑΤ2	1026	341	1 (1-420) 2 (490-1095)	1		BN001414
Trichophyton equinum CBS 127.97	TRIEC	559882	ΝΑΤ1	981	326	1 (1-416) 2 (481-528) 3 (593-1109)	2	TEQG_03758	BN001415
Trichophyton rubrum CBS 118892	TRIRC	559305	ΝΑΤ1	981	326	1 (1-416) 2 (481-528) 3 (593-1109)	2	TERG_07215	BN001416
Trichophyton tonsurans CBS 112818	TRIT1	647933	ΝΑΤ1	981	326	1 (1-416) 2 (481-528) 3 (593-1109)	2	TESG_05215	BN001417
Uncinocarpus reesii UAMH 1704	UNCRE	336963	ΝΑΤ1	990	329	1 (1-417) 2 (467-1039)	1	UREG_05260	BN001418
Verticillium albo-atrum VaMs.102	VERA1	526221	ΝΑΤ1	762	253	1 (1-360) 2 (484-708) 3 (832-1008)	2		BN001419
Verticillium dahliae VdLs.17	VERDV	498257	ΝΑΤ1	951	316	1 (1-951)	0	VDAG_09857	BN001420

Footnotes, Table 2:

^(a) The details and provisional taxonomic classification of the sequenced fungal genomes are available on http://www.ncbi.nlm.nih.gov/genomes/leuks.cgi and http://www.broadinstitute.org/science/projects/fungal-genome-initiative/current-fgi-sequence-projects. These fungal sequences have been described in detail in [41,42,43,44]. The reference sequence for annotation of fungal NATs is the NAT1 protein of the ascomycetes Fusarium verticillioides (Gibberella moniliformis), GenBank ID: EU552489 [5,43].

^(b) Column key – Colums 1, 2: The numbers in brackets are sequenced fungal genomes accessible via BLAST (2010). Search of the EST database additionally retrieved a number of partial NAT-homologous sequences (data not shown), all of which belong to either basidiomycetes or ascomycetes (Pezizomycotina); Columns 2-4: The scientific name, official identification code and unique taxonomy identifier for each organism are available from the UniProt Taxonomy Browser (http://www.uniprot.org/taxonomy/); Columns 5-9: The retrieval and annotation of NAT-homologous sequences (ORF span and exon-intron structure) were carried out as described [41], and the gene symbols were assigned according to the guidelines of the NAT Gene Nomenclature Committee. The sequence length of open reading frames (ORF) is shown in nucleotides (nt) and of deduced proteins in amino acids (aa); Column 10: Where validated, the locus tags are provided to anchor each annotated NAT gene to the corresponding genomic map; Column 11: Third-party annotation (TPA) accession numbers were assigned by the EMBL database and can be used to retrieve the complete details of the new NAT genes described in this study.

Table 3: Annotation of NAT genes in sequenced genomes of animals^(a)

METAZOA^(b)
PLACOZOA
NONE
EUMETAZOA
Cnidaria
Acropora millepora [mRNA sequence bearing the complete ORF of a NAT-homologous gene (GenBank ID: EZ029316)]
Bilateria
Acoelomata (Platyelminthes)
NONE
Pseudocoelomata (Nematoda)
NONE
Coelomata
Protostomia
Mollusca
NONE
Panarthropoda
Arthropoda
Hexapoda (insects)
NONE
Organism scientific name^(c)	Organism identification code^(c)	Taxonomy identifier^(c)	Gene (alias)^(c)	ORF (nt) ^(c)	Protein (aa) ^(c)	Identities (%)	Positives (%)	Literature
Deuterostomia
Echinozoa
Strongylocentrotus purpuratus (purple sea urchin)	STRPU	7668	NAT1	933	310	27	44	1
			NAT2	654	217	24	42
			NAT3	933	310	23	43
Hemichordata
Saccoglossus kowalevskii (Acorn worm)	SACKO	10224	NAT1	903	300	20	38	41
Chordata
Tunicata
Ciona intestinalis (Transparent sea squirt)	CIOIN	7719	NAT1	789 ORF consists of 3 exons Exon 1: 1-335 Exon 2: 651-838 Exon 3: 1472-1737 (Gene size: 1737nt)	262	28	45	41, BN001483
			NAT2	942 ORF consists of 5 exons Exon 1: 1-335 Exon 2: 685-866 Exon 3: 1863-1968 Exon 4: 3067-3157 Exon 5: 3624-3851 (Gene size: 3851nt)	313	25	47	41, BN001484
Cephalochordata
Branchiostoma floridae (Florida lancelet)	BRAFL	7739	NAT1	780 ORF consists of 4 exons Exon 1: 1-119 Exon 2: 639-957 Exon 3: 1238-1277 Exon 4: 1347-1648 (Gene size: 1648nt)	259	30	48	41, BN001480
			NAT2	864 ORF consists of 3 exons Exon 1: 1-119 Exon 2: 1713-2037 Exon 3: 2282-2701 (Gene size: 2701nt)	287	27	48	41, BN001481
			NAT3	909 ORF consists of 3 exons Exon 1: 1-176 Exon 2: 779-906 Exon 3: 4565-5169 (Gene size: 5169nt)	302	23	42	41, BN001482
Vertebrata
Chondrichthyes
NONE
Euteleostomi
Actinopterygii
Danio rerio (zebrafish)	DANRE	7955	NAT1	867	288	43	62	1-4
			NAT2	867	288	42	62
			NAT3	867	288	42	61
			NAT4	873	290	40	59
Gasterosteus aculeatus (three-spined stickleback)	GASAC	69293	NAT1	912	303	42	62	1
			NAT2	870	289	41	61
Oryzias latipes (medaka fish)	ORYLA	8090	NAT1	873	290	44	61	1
			NAT2	873	290	40	61
			NAT3	846	281	35	52
Takifugu (Fugu) rubripes (Japanese pufferfish)	TAKRU	31033	NAT1	846	281	33	50	1,2
Tetraodon nigroviridis (green spotted pufferfish)	TETNG	99883	NAT1	843	280	34	49	1,2
Sarcopterygii
Amniota
Mammalia ^(d)
Primates^(e)
Allenopithecus nigroviridis (Allen’s swamp monkey)	ALLNI	54135	NAT1	873	290	94	98	46
			NAT2	873	290	82	90
Cebus apella (Brown-capped capuchin monkey)	CEBAP	9515	NAT1	873	290	94	98	46
			NAT2	873	290	81	90
Cercopithecus diana (Diana monkey)	CERDI	36224	NAT1	873	290	95	98	46
			NAT2	873	290	82	90
Colobus guereza (black and white colobus monkey)	COLGU	33548	NAT1	873	290	95	99	46
			NAT2	873	290	82	89
Chlorocebus tantalus (tantalus monkey)	CERTA	60712	NAT1	873	290	96	99	46
			NAT2	873	290	82	89
Erythrocebus patas (Red guenon)	ERYPA	9538	NAT1	873	290	96	99	46
			NAT2	873	290	82	90
Gorilla gorilla (Lowland gorilla)	GORGO	9595	NAT1	873	290	98	99	45,46
			NAT2	873	290	82	89
Homo sapiens (human)	HUMAN	9606	NAT1	873	290	100	100	9,10
			NAT2	873	290	81	89
Hylobates lar (White handed gibbon)	HYLLA	9580	NAT1	873	290	96	99	46
			NAT2	873	290	81	88
Presbytis cristatus (silvered-leaf monkey)	TRACR	122765	NAT2	873	290	96	99	46
			NAT1	873	290	82	90
Macaca fascicularis (Crab-eating macaque)	MACFA	9541	NAT1	873	290	94	98	45
			NAT2	873	290	82	89
Macaca mulatta (Rhesus macaque)	MACMU	9544	NAT1	873	290	94	99	11
			NAT2	873	290	82	90
Macaca sylvanus (Barbary ape)	MACSY	9546	NAT1	873	290	95	99	46
			NAT2	873	290	82	90
Mandrillus sphinx (Mandrill)	MANSP	9561	NAT1	873	290	95	99	46
			NAT2	873	290	82	89
Microcebus murinus (gray mouse lemur)	MICMU	30608	NAT1	873	290	80	90	1
			NAT2	873	290	74	85
Nomascus gabriellae (Red cheeked gibbon)	NOMGA	61852	NAT1	873	290	96	99	46
			NAT2	873	290	82	89
Otolemur garnettii (small-eared galago)	OTOGA	30611	NAT1	873	290	82	93	1
Pan paniscus (Pygmy chimpanzee)	PANPA	9597	NAT1	873	290	98	100	46
			NAT2	873	290	81	89
Pan troglodytes (chimpanzee)	PANTR	9598	NAT1	873	290	97	100	1
			NAT2	873	290	81	89	1
Papio anubis (Olive baboon)	PAPAN	9555	NAT1	873	290	95	99	45
			NAT2	873	290	82	89
Pithecia pithecia (White faced saki)	PITPI	43777	NAT1	873	290	95	99	46
			NAT2	873	290	82	91
Pongo abelii (Sumatran orangutan)	PONAB	9601	NAT1	873	290	96	99	45
			NAT2	873	290	81	89
Saimiri boliviensis boliviensis (Bolivian squirrel monkey)	SAIBB	39432	NAT1	873	290	93	97
Other mammals
Bos taurus (domestic cattle)	BOVIN	9913	NAT1	873	290	82	93	1,4,8
Cavia porcellus (domestic Guinea pig)	CAVPO	10141	Nat1	870	289	79	91	1
			Nat2	873	290	81	93
			Nat3	873	290	71	86
Cricetulus griseus (Chinese hamster)	CRIGR	10029	Nat2	873	290	80	92	AJ289858
Echinops telfairi (lesser hedgehog tenrec)	ECHTE	9371	NAT1	873	290	80	88	1
Equus caballus (domestic horse)	HORSE	9796	NAT1	873	290	82	92	1
Loxodonta Africana (African bush elephant)	LOXAF	9785	NAT1	873	290	84	93	1
			NAT2	873	290	82	91
			NAT3	873	290	77	87
Mesocricetus auratus (Syrian hamster)	MESAU	10036	Nat1	873	290	71	83	12-16
			Nat2	873	290	81	92
Monodelphis domestica (gray short-tailed opossum)	MONDO	13616	NAT1	870	289	68	85	1
Mus musculus (house mouse)	MOUSE	10090	Nat1	873	290	74	85	17,18
			Nat2	873	290	82	94
			Nat3	873	290	68	81
Myotis lucifugus (little brown bat)	MYOLU	59463	NAT1	873	290	82	92	1
			NAT2	873	290	73	86
Ornithorhynchus anatinus (platypus)	ORNAN	9258	NAT1	870	289	56	73	1
Oryctolagus cuniculus (European rabbit)	RABIT	9986	NAT1	873	290	75	87	19-21
			NAT2	873	290	74	87
			NAT2	870	289	81	88
Rattus norvegicus (brown rat)	RAT	10116	Nat1	873	290	76	87	22,23
			Nat2	873	290	81	93
			Nat3	873	290	68	81
Sus scrofa (pig)	PIG	9823	NAT1	873	290	84	92	1,4
Sauria
Lepidosauria
Anolis carolinensis (Carolina green anole)	ANOCA	28377	NAT1	867	288	54	72	1
			NAT2	867	288	53	72
			NAT3	816	271	35	50
Archosauria
Aves
Gallus gallus (chicken)	CHICK	9031	NAT1	873	290	57	76	1,24,25
			NAT2	873	290	53	72
			NAT3	873	290	53	72
			NAT4	852	283	48	71
Taeniopygia guttata (Zebra finch)	TAEGU	59729	NAT1	873	290	59	76	41
			NAT2	873	290	53	71
			NAT3	867	288	52	70
			NAT4	891	296	50	70

Footnotes, Table 3

(a) The reference sequence for annotation of animal NATs is the NAT1 protein of Homo sapiens (Human), GenBank ID: X17059 [9,10]. Amino acid sequence homology (identity and similarity) is calculated relative to this reference sequence.

(b) Search of the EST database identified partial NAT sequences from additional metazoa, including the rotifer Brachionus plicatilis (dbEST ID: FM940182), the mollusc Lottia gigantea (dbEST ID: FC756210), the crustacean Litopenaeus vannamei (dbEST ID: FE155086), the insect Adelphocoris lineolatus (dbEST ID: GR483776), the asterozoon Patiria pectinifera (dbEST ID: DB395455), the salamander Cynops pyrrhogaster (dbEST ID: FS311170), and the frogs Xenopus laevis (dbEST ID: EB646873) and Xenopus (Silurana) tropicalis (dbEST ID: EL679494) [41]. An additional sequence has been reported in Felis silvestris catus [26]. NAT activity has also been detected in the domestic pigeon Columba livia, but no gene sequence is yet available [27,28]. NAT genes are absent in the genome of the domestic dog Canis familiaris [29]. NAT activity has been reported in some nematodes [6,7], but the results are not verified as yet.

(c) Official organism scientific names, identification codes and taxonomy identifiers are from the NEWT UniProt Taxonomy Browser. Taxonomic presentation is according to the NCBI Taxonomy Database, which provides a platform for the classification of all organisms represented in the GenBank database. Please note that these databases are not a primary source for taxonomic or phylogenetic information and should not be used as such. Abbreviations: ORF – open reading frame; nt – nucleotides; aa – amino acids.

(d) A number of upstream non-coding exons have been reported in the literature for the NAT genes of the primates [11,30-36], rodents [12,15,16,37-39] and chicken [2]. The complex structure of vertebrate NAT genes is reviewed in references [2,40].

(e) Pseudogenes with homology to NAT genes (designated NATP1) have been identified in the primates Homo sapiens, Colobus guereza, Presbytis cristatus, Allenopithecus nigroviridis, Cercopithecus diana, Chlorocebus tantalus, Erythrocebus patas, Mandrillus sphinx, Macaca sylvanus, Macaca mulatta, Nomascus gabriellae, Gorilla gorilla, Pan troglodytes, Pan paniscus [46].

All reported sequences are stored in electronic format and can be requested from the curator of this website. Please note that many of the ORFs described in the above tables were reconstructed from draft low homology genomic sequences retrieved during our database searches. Therefore, these sequences represent only putative NAT genes and each ORF should be checked for possible inaccuracies in length and sequence before any further study is undertaken.

If you wish to acknowledge this website as a source of data in future publications, please cite references [1 and 41].

For new submissions or enquiries, please contact Dr. Sotiria Boukouvala (sboukouv@mbg.duth.gr).

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Literature

[1] Vagena, E.; Fakis, G. and Boukouvala, S. (2008) Arylamine N-acetyltransferases in prokaryotic and eukaryotic genomes: A survey of public databases. Curr. Drug Metab. 9(7), 628-660.

[2] Boukouvala, S. and Fakis, G. (2005) Arylamine N-acetyltransferases: what we learn from genes and genomes. Drug Metab. Rev. 37(3), 511-564.

[3] Sandy, J.; Mushtaq, A.; Holton, S.J.; Schartau, P.; Noble, M.E.M. and Sim, E. (2005) Investigation of the catalytic triad of arylamine N-acetyltransferases: essential residues required for acetyl transfer to arylamines. Biochem. J. 390(Pt. 1), 115-123.

[4] Rodrigues-Lima, F. and Dupret, J.M. (2002) In silico sequence analysis of arylamine N-acetyltransferases: evidence for an absence of lateral gene transfer from bacteria to vertebrates and first description of paralogs in bacteria. Biochem. Biophys. Res. Commun. 293(2), 783-792.

[5] Glenn, A.E.; Meredith, F.I.; Morrison, W.H. III and Bacon, C.W. (2003) Identification of intermediate and branch metabolites resulting from biotransformation of 2-benzoxazolinone by Fusarium verticillioides. Appl. Environ. Microbiol. 69(6), 3165-3169.

[6] Chung, J.G. (1999) Purification and characterization of an arylamine N-acetyltransferase in the nematode Enterobius vermicularis. Microbios. 98(389), 15-25.

[7] Chung, J.G.; Kuo, H.M.; Lin, T.H.; Ho, C.C.; Lee, J.H.; Lai, J.M.; Levy, G.N. and Weber, W.W. (1996) Evidence for arylamine N-acetyltransferase in the nematode Anisakis simplex. Cancer Lett. 106(1), 1-8.

[8] Upton, A. and Sim, E. (1999) Presence of an arylamine N-acetyltransferase gene homologue and N-acetylation activity in domestic cattle. Biochemical Society Meeting, Cork, Ireland.

[9] Blum, M.; Grant, D.M.; McBride, W.; Heim, M. and Meyer, U.A. (1990) Human arylamine N-acetyltransferase genes: isolation, chromosomal localization, and functional expression. DNA Cell Biol. 9(3), 193-203.

[10] Ohsako, S. and Deguchi, T. (1990) Cloning and expression of cDNAs for polymorphic and monomorphic arylamine N-acetyltransferases from human liver. J. Biol. Chem. 265(8), 4630-4634.

[11] Fakis, G.; Boukouvala, S.; Kawamura, A. and Kennedy, S. (2007) Description of a novel polymorphic gene encoding for arylamine N-acetyltransferase in the rhesus macaque (Macaca mulatta), a model animal for endometriosis. Pharmacogenet. Genomics 17(3), 181-188.

[12] Abu-Zeid, M.; Nagata, K.; Miyata, M.; Ozawa, S.; Fukuhara, M.; Yamazoe, Y. and Kato, R. (1991) An arylamine acetyltransferase (AT-1) from Syrian golden hamster liver: Cloning, complete nucleotide sequence and expression in mammalian cells. Mol. Carcinogen. 4(1), 81-88.

[13] Ferguson, R.J., Doll, M.A., Rustan, T.D., Baumstark, B.R. and Hein, D.W. (1994) Syrian hamster monomorphic N-acetyltransferase (NAT1) alleles: Amplification, cloning, sequencing, and expression in E. coli. Pharmacogenetics 4(2), 82-90.

[14] Ferguson, R.J.; Doll, M.A.; Rustan, T.D. and Hein, D.W. (1996) Cloning, expression and functional characterisation of rapid and slow acetylator polymorphic N-acetyltransferase encoding genes of the Syrian hamster. Pharmacogenetics 6(1), 55-66.

[15] Land, S.J.; Jones, R.E. and King, C.M. (1994) Biochemical and genetic analysis of two acetyltransferases from hamster tissues that can metabolise aromatic amine derivatives. Carcinogenesis 15(8), 1585-1595.

[16] Nagata, K.; Ozawa, S.; Miyata, M.; Shimada, M.; Yamazoe, Y. and Kato, R. (1994) Primary structure and molecular basis of polymorphic appearance of an acetyltransferase (AT-II) in hamsters. Pharmacogenetics 4(2), 91-100.

[17] Martell, K.J.; Vatsis, K.P. and Weber, W.W. (1991) Molecular genetic basis of rapid and slow acetylation in mice. Mol. Pharmacol. 40(2), 218-227.

[18] Kelly, S.L. and Sim, E. (1994) Arylamine N-acetyltransferase in Balb/c mice: identification of a novel mouse isoenzyme by cloning and expression in vitro. Biochem. J. 302(2), 347-353.

[19] Blum, M.; Heim, M. and Meyer, U.A. (1990) Nucleotide sequence of rabbit NAT1 encoding monomorphic arylamine N-acetyltransferase. Nucl. Acids Res. 18(17), 5287.

[20] Blum, M.; Heim, M. and Meyer, U.A. (1990) Nucleotide sequence of rabbit NAT2 encoding polymorphic liver arylamine N-acetyltransferase (NAT). Nucl. Acids Res. 18(17), 5295.

[21] Sasaki, Y.; Ohsako, S. and Deguchi, T. (1991) Molecular and genetic analyses of arylamine N-acetyltransferase polymorphism of rabbit liver. J. Biol. Chem. 266(20), 13243-13250.

[22] Doll, M.A. and Hein, D.W. (1995) Cloning, sequencing and expression of NAT1 and NAT2 encoding genes from rapid and slow acetylator inbred rats. Pharmacogenetics 5(4), 247-251.

[23] Walraven, J.M.; Doll, M.A. and Hein, D.W. (2006) Identification and characterization of functional rat arylamine N-acetyltransferase 3: comparisons with rat arylamine N-acetyltransferases 1 and 2. J. Pharmacol. Exp. Ther. 319(1), 369-375.

[24] Ohsako, S.; Ohtomi, M.; Sakamoto, Y.; Uyemura, K. and Deguchi, T. (1988) Arylamine N-acetyltransferase from chicken liver II. Cloning of cDNA and expression in Chinese hamster ovary cells. J. Biol. Chem. 263(16), 7534-7538.

[25] Ohtomi, M.; Sasaki, M. and Deguchi, T. (1989) Two arylamine N-acetyltransferases from chicken pineal gland as identified by cDNA cloning. Eur. J. Biochem. 185(2), 253-261.

[26] Trepanier, L.A.; Cribb, A.; Spielberg, S.P. and Ray, K. (1998) Deficiency of cytosolic arylamine N-acetylation in the domestic cat and wild felids caused by the presence of a single NAT1-like gene. Pharmacogenetics 8(2), 169-179.

[27] Andres, H.H.; Kolb, H.J.; Schreiber, R.J. and Weiss, L. (1983) Characterization of the active site, substrate specificity and kinetic properties of acetyl-CoA:arylamine N-acetyltransferase from pigeon liver. Biochem. Biophys. Acta 746(3), 193-201.

[28] Andres, H.H.; Kolb, H.J. and Weiss, L. (1983) Purification and physical-chemical properties of acetyl-CoA:arylamine N-acetyltransferase from pigeon liver. Biochim. Biophys. Acta 746(3), 182-192.

[29] Trepanier, L.A.; Ray, K.; Winand, N.J.; Spielberg, S.P. and Cribb, A.E. (1997) Cytosolic arylamine N-acetyltransferase (NAT) deficiency in the dog and other canids due to an absence of NAT genes. Biochem. Pharmacol. 54(1), 73-80.

[30] Ebisawa, T. and Deguchi, T. (1991) Structure and restriction fragment length polymorphism of genes for human liver arylamine N-acetyltransferases. Biochem. Biophys. Res. Commun. 177(3), 1252-1257.

[31] Husain, A.; Barker, D.F.; States, J.C.; Doll, M.A. and Hein, D.W. (2004) Identification of the major promoter and non-coding exons of the human arylamine N-acetyltransferase 1 gene (NAT1). Pharmacogenetics 14(7), 397-406.

[32] Boukouvala, S. and Sim, E. (2005) Structural analysis of the genes for human arylamine N-acetyltransferases and characterisation of alternative transcripts. Basic Clin. Pharmacol. Toxicol. 96(5), 343-351.

[33] Butcher, N.J.; Arulpragasam, A.; Goh, H.L.; Davey, T. and Minchin, R.F. (2005) Genomic organization of human arylamine N-acetyltransferase Type I reveals alternative promoters that generate different 5'-UTR splice variants with altered translational activities. Biochem. J. 387(1), 119-127.

[34] Barker, D.F.; Husain, A.; Neale, J.R.; Martini, B.D.; Zhang, X.; Doll, M.A.; States, J.C. and Hein, D.W. (2006) Functional properties of an alternative, tissue-specific promoter for human arylamine N-acetyltransferase 1. Pharmacogenet. Genomics 16(7), 515-525.

[35] Husain, A.; Zhang, X.; Doll, M.A.; States, J.C.; Barker, D.F. and Hein, D.W. (2007) Functional analysis of the human N-acetyltransferase 1 major promoter: Quantitation of tissue expression and identification of critical sequence elements. Drug Metab. Dispos. 35(9), 1649-1656.

[36] Husain, A.; Zhang, X.; Doll, M.A.; States, J.C.; Barker, D.F. and Hein, D.W. (2007) Identification of N-acetyltransferase 2 (NAT2) transcription start sites and quantitation of NAT2-specific mRNA in human tissues. Drug Metab. Dispos. 35(5), 721-727.

[37] Fakis, G.; Boukouvala, S.; Buckle, V.; Payton, M.; Denning, C. and Sim, E. (2000) Chromosome mapping of the genes for murine arylamine N-acetyltransferases (NATs), enzymes involved in the metabolism of carcinogens: identification of a novel upstream noncoding exon for murine Nat2. Cytogenet. Cell Genet. 90(1-2), 134-138.

[38] Boukouvala, S.; Price, N.; Plant, K.E. and Sim, E. (2003) Structure and transcriptional regulation of the Nat2 gene encoding for the drug-metabolizing enzyme arylamine N-acetyltransferase type 2 in mice. Biochem. J. 375(3), 593-602.

[39] Ebisawa, T.; Sasaki, Y. and Deguchi, T. (1995) Complementary DNAs for two arylamine N-acetyltransferases with identical 5¢ non-coding regions from rat pineal gland. Eur. J. Biochem. 228(1), 129-137.

[40] Sim, E.; Walters, K. and Boukouvala, S. (2008) Arylamine N-acetyltransferases: From structure to function. Drug Metab. Rev. 40(3), 479-510.

[41] Glenn, A.E.; Karagianni, E.P.; Ulndreaj, A. and Boukouvala, S. (2010) Comparative genomic and phylogenetic investigation of the xenobiotic metabolizing arylamine N-acetyltransferase enzyme family. FEBS Lett. 584(14), 3158-3164.

[42] Martins, M.; Dairou, J.; Rodrigues-Lima, F.; Dupret, J.M. and Silar, P. (2010) Insights into the phylogeny or arylamine N-acetyltransferases in fungi. J. Mol. Evol. 71(2), 141-152.

[43] Glenn, A.E. and Bacon, C.W. (2009) FDB2 encodes a member of the arylamine N-acetyltransferase family and is necessary for biotransformation of benzoxazolinones by Fusarium verticillioides. J. Appl. Microbiol. 107(2), 657-671.

[44] Martins, M.; Rodrigues-Lima, F.; Dairou, J.; Lamouri, A.; Malagnac, F.; Silar, P. and Dupret, J.M. (2009) An acetyltransferase conferring tolerance to toxic aromatic amine chemicals: molecular and functional studies. J. Biol. Chem. 284(28), 18726-18733.

[45] Sim, E.; Fakis, G.; Laurieri, N. and Boukouvala, S. (2012) Arylamine N-acetyltransferases--from drug metabolism and pharmacogenetics to identification of novel targets for pharmacological intervention. Adv. Pharmacol. 63, 169-205.

[46] Sabbagh, A.; Marin, J.; Veyssiere, C.; Lecompte, E.; Boukouvala, S.; Paloni, E.S.; Darlu, P. and Crouau-Roy, B. (2013) Rapid birth-and-death evolution of the xenobiotic metabolizing NAT gene family in vertebrates with evidence of adaptive selection. BMC Evol. Biol. 13:62.

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