Reviewed, UniProtKB/Swiss-Prot Q9UKV0 (HDAC9_HUMAN)

Names and origin

Protein names

Recommended name:
    Histone deacetylase 9
      Short name=HD9
    EC=3.5.1.98
Alternative name(s):
    HD7B
      Short name=HD7
    Histone deacetylase-related protein
    MEF2-interacting transcription repressor MITR

Gene names

Name:	HDAC9
Synonyms:	HDAC7, HDAC7B, HDRP, KIAA0744, MITR

Organism

Homo sapiens (Human)

Taxonomic identifier

9606 [NCBI]

Taxonomic lineage

Eukaryota › Metazoa › Chordata › Craniata › Vertebrata › Euteleostomi › Mammalia › Eutheria › Euarchontoglires › Primates › Haplorrhini › Catarrhini › Hominidae › Homo

Protein attributes

Sequence length	1011 AA.
Sequence status	Complete.
Sequence processing	The displayed sequence is not processed.
Protein existence	Evidence at protein level.

General annotation (Comments)

Function	Responsible for the deacetylation of lysine residues on the N-terminal part of the core histones (H2A, H2B, H3 and H4). Histone deacetylation gives a tag for epigenetic repression and plays an important role in transcriptional regulation, cell cycle progression and developmental events. Represses MEF2-dependent transcription. Ref.1 Ref.3 Ref.8 Ref.9 Ref.13 Isoform 3, called MITR/HDRP, lacks active site residues and therefore is catalytically inactive. Represses MEF2-dependent transcription by recruiting HDAC1 and/or HDAC3. Seems to inhibit skeletal myogenesis and to be involved in heart development. Protects neurons from apoptosis, both by inhibiting c-Jun phosphorylation by MAPK10 and by repressing c-Jun transcription via HDAC1 recruitment to c-Jun promoter. Ref.1 Ref.3 Ref.8 Ref.9 Ref.13
Catalytic activity	Hydrolysis of an N(6)-acetyl-lysine residue of a histone to yield a deacetylated histone.
Enzyme regulation	Inhibited by Trichostatin A (TSA) and suberoylanilide hydroxamic acid. Ref.1
Subunit structure	Homodimer. Interacts with CTBP1. The phosphorylated form interacts with 14-3-3 By similarity. Interacts with HDAC1 and HDAC3, and probably with HDAC4 and HDAC5. Interacts with MEF2, MAPK10, ETV6, NCOR1 and BCL6.
Subcellular location	Nucleus By similarity.
Tissue specificity	Broadly expressed, with highest levels in brain, heart, muscle and testis. Isoform 3 is present in human bladder carcinoma cells (at protein level). Ref.1 Ref.3 Ref.9 Ref.2
Post-translational modification	Phosphorylated on Ser-220 and Ser-450; which promotes 14-3-3-binding, impairs interaction with MEF2, and antagonizes antimyogenic activity. Phosphorylated on Ser-240; which impairs nuclear accumulation By similarity. Isoform 7 is phosphorylated on Tyr-1010. Sumoylated. Ref.3 Ref.11
Involvement in disease	A chromosomal aberration involving HDAC9 is found in a family with Peters anomaly [MIM:604229]. Translocation t(1;7)(q41;p21) with TGFB2 resulting in lack of HDAC9 protein. Peters anomaly consists of a central corneal leukoma, absence of the posterior corneal stroma and Descemet membrane, and a variable degree of iris and lenticular attachments to the central aspect of the posterior cornea.
Sequence similarities	Belongs to the histone deacetylase family. Type 2 subfamily.

Ontologies

Keywords
Biological process	Transcription Transcription regulation
Cellular component	Nucleus
Coding sequence diversity	Alternative splicing Chromosomal rearrangement
Molecular function	Chromatin regulator Hydrolase Repressor
PTM	Phosphoprotein Ubl conjugation
Gene Ontology (GO)
Biological process	B cell differentiation Traceable author statement. Source: UniProtKB histone deacetylation Ref.7 Non-traceable author statement. Source: UniProtKB inflammatory response Traceable author statement. Source: UniProtKB negative regulation of striated muscle development Traceable author statement. Source: UniProtKB regulation of transcription, DNA-dependent Inferred from electronic annotation. Source: UniProtKB-KW transcription Inferred from electronic annotation. Source: UniProtKB-KW
Cellular component	cytoplasm Traceable author statement. Source: UniProtKB histone deacetylase complex Traceable author statement. Source: UniProtKB
Molecular function	histone deacetylase activity Ref.7 Non-traceable author statement. Source: UniProtKB specific transcriptional repressor activity Traceable author statement. Source: UniProtKB transcription factor binding Traceable author statement. Source: UniProtKB
Complete GO annotation...

Binary interactions

With	Entry	#Exp.	IntAct	Notes
BCL6	P41182	1	EBI-765444,EBI-765407
BCL6	P41182	1	EBI-1372717,EBI-765407
BCL6	P41182	1	EBI-765476,EBI-765407
ETV6	P41212	1	EBI-1372713,EBI-1372759
ETV6	P41212	2	EBI-765476,EBI-1372759
ETV6	P41212	1	EBI-765444,EBI-1372759
ETV6	P41212	1	EBI-1372717,EBI-1372759
HDAC3	O15379	1	EBI-1372717,EBI-607682
HDAC3	O15379	1	EBI-765476,EBI-607682
HDAC4	P56524	1	EBI-1372717,EBI-308629
HDAC4	P56524	1	EBI-765476,EBI-308629
NCOR1	O75376	1	EBI-765444,EBI-347233
Ncor1	Q60974	1	EBI-1372713,EBI-349004	From a different organism.
Ncor1	Q60974	2	EBI-765476,EBI-349004	From a different organism.
Ncor1	Q60974	2	EBI-1372717,EBI-349004	From a different organism.
Sin3a	Q60520	1	EBI-1372717,EBI-349034	From a different organism.
Sin3a	Q60520	1	EBI-765476,EBI-349034	From a different organism.
Sin3b	Q62141	1	EBI-1372717,EBI-591450	From a different organism.
Sin3b	Q62141	1	EBI-765476,EBI-591450	From a different organism.

Alternative products

This entry describes 7 isoforms produced by alternative splicing. [Align] [Select]

Note: Additional isoforms seem to exist.

Isoform 1 (identifier: Q9UKV0-1)

This isoform has been chosen as the 'canonical' sequence. All positional information in this entry refers to it. This is also the sequence that appears in the downloadable versions of the entry.

Isoform 2 (identifier: Q9UKV0-2)

The sequence of this isoform differs from the canonical sequence as follows:
     487-574: Missing.

Isoform 3 (identifier: Q9UKV0-3)

Also known as: HDRP; MITR;

The sequence of this isoform differs from the canonical sequence as follows:
     575-590: PFLEPTHTRALSVRQA → VIGKDLAPGFVIKVII
     591-1011: Missing.

Note: Major form in most tissues. Inactive due to lack of active site residues.

Isoform 4 (identifier: Q9UKV0-4)

Also known as: HDAC9a;

The sequence of this isoform differs from the canonical sequence as follows:
     861-879: GTGLGEGYNINIAWTGGLD → RFISLEPHFYLYLSGNCIA
     880-1011: Missing.

Isoform 5 (identifier: Q9UKV0-5)

Also known as: HDAC9b; HDAC9fl;

The sequence of this isoform differs from the canonical sequence as follows:
     1006-1011: MSLKFS → KYWKSVRMVAVPRGCALAGAQLQEETETVSALASLTVDVEQPFAQEDSRTAGEPMEEEPAL

Note: Phosphorylated on Tyr-1007 (Probable).

Isoform 6 (identifier: Q9UKV0-6)

The sequence of this isoform differs from the canonical sequence as follows:
     88-88: K → KLQQ
     218-261: Missing.
     1006-1011: MSLKFS → KYWKSVRMVAVPRGCALAGAQLQEETETVSALASLTVDVEQPFAQEDSRTAGEPMEEEPAL

Note: Ref.6 (AAI11736) sequence differs from that shown due to a frameshift in position 1021. Phosphorylated on Tyr-966 (Probable). Excluded from the nucleus. Does not interact with ETV6..

Isoform 7 (identifier: Q9UKV0-7)

The sequence of this isoform differs from the canonical sequence as follows:
     88-88: K → KLQQ
     1006-1011: MSLKFS → KYWKSVRMVAVPRGCALAGAQLQEETETVSALASLTVDVEQPFAQEDSRTAGEPMEEEPAL

Note: Phosphorylated on Tyr-1010.

Sequence annotation (Features)

	Feature key	Position(s)	Length	Description	Graphical view	Feature identifier
Molecule processing
	Chain	1 – 1011	1011	Histone deacetylase 9		PRO_0000114710
Regions
	Region	23 – 27	5	Interaction with CTBP1 By similarity
	Region	136 – 154	19	Interaction with MEF2 By similarity
	Region	175 – 343	169	Interaction with MAPK10 By similarity
	Region	218 – 261	44	Interaction with ETV6
	Region	631 – 978	348	Histone deacetylase
Sites
	Active site	783	1	By similarity
Amino acid modifications
	Modified residue	220	1	Phosphoserine By similarity
	Modified residue	240	1	Phosphoserine By similarity
	Modified residue	451	1	Phosphoserine By similarity
Natural variations
	Alternative sequence	88	1	K → KLQQ in isoform 6 and isoform 7.		VSP_023766
	Alternative sequence	218 – 261	44	Missing in isoform 6.		VSP_023767
	Alternative sequence	487 – 574	88	Missing in isoform 2.		VSP_002082
	Alternative sequence	575 – 590	16	PFLEP…SVRQA → VIGKDLAPGFVIKVII in isoform 3.		VSP_002083
	Alternative sequence	591 – 1011	421	Missing in isoform 3.		VSP_002084
	Alternative sequence	861 – 879	19	GTGLG…TGGLD → RFISLEPHFYLYLSGNCIA in isoform 4.		VSP_002085
	Alternative sequence	880 – 1011	132	Missing in isoform 4.		VSP_002086
	Alternative sequence	1006 – 1011	6	MSLKFS → KYWKSVRMVAVPRGCALAGA QLQEETETVSALASLTVDVE QPFAQEDSRTAGEPMEEEPA L in isoform 5, isoform 6 and isoform 7.		VSP_023768
Experimental info
	Sequence conflict	99	1	L → I Ref.7
	Sequence conflict	437	1	T → P Ref.7
	Sequence conflict	644 – 647	4	HQCV → KPNS Ref.7
	Sequence conflict	746	1	H → R in AAO27363. Ref.3

Sequences

	Sequence		Length	Mass (Da)
	Isoform 1 [UniParc]. Last modified October 18, 2001. Version 2. Checksum: 43ED2785E73CD924	FASTA	1,011	111,297
	10 20 30 40 50 60 MHSMISSVDV KSEVPVGLEP ISPLDLRTDL RMMMPVVDPV VREKQLQQEL LLIQQQQQIQ 70 80 90 100 110 120 KQLLIAEFQK QHENLTRQHQ AQLQEHIKEL LAIKQQQELL EKEQKLEQQR QEQEVERHRR 130 140 150 160 170 180 EQQLPPLRGK DRGRERAVAS TEVKQKLQEF LLSKSATKDT PTNGKNHSVS RHPKLWYTAA 190 200 210 220 230 240 HHTSLDQSSP PLSGTSPSYK YTLPGAQDAK DDFPLRKTAS EPNLKVRSRL KQKVAERRSS 250 260 270 280 290 300 PLLRRKDGNV VTSFKKRMFE VTESSVSSSS PGSGPSSPNN GPTGSVTENE TSVLPPTPHA 310 320 330 340 350 360 EQMVSQQRIL IHEDSMNLLS LYTSPSLPNI TLGLPAVPSQ LNASNSLKEK QKCETQTLRQ 370 380 390 400 410 420 GVPLPGQYGG SIPASSSHPH VTLEGKPPNS SHQALLQHLL LKEQMRQQKL LVAGGVPLHP 430 440 450 460 470 480 QSPLATKERI SPGIRGTHKL PRHRPLNRTQ SAPLPQSTLA QLVIQQQHQQ FLEKQKQYQQ 490 500 510 520 530 540 QIHMNKLLSK SIEQLKQPGS HLEEAEEELQ GDQAMQEDRA PSSGNSTRSD SSACVDDTLG 550 560 570 580 590 600 QVGAVKVKEE PVDSDEDAQI QEMESGEQAA FMQQPFLEPT HTRALSVRQA PLAAVGMDGL 610 620 630 640 650 660 EKHRLVSRTH SSPAASVLPH PAMDRPLQPG SATGIAYDPL MLKHQCVCGN STTHPEHAGR 670 680 690 700 710 720 IQSIWSRLQE TGLLNKCERI QGRKASLEEI QLVHSEHHSL LYGTNPLDGQ KLDPRILLGD 730 740 750 760 770 780 DSQKFFSSLP CGGLGVDSDT IWNELHSSGA ARMAVGCVIE LASKVASGEL KNGFAVVRPP 790 800 810 820 830 840 GHHAEESTAM GFCFFNSVAI TAKYLRDQLN ISKILIVDLD VHHGNGTQQA FYADPSILYI 850 860 870 880 890 900 SLHRYDEGNF FPGSGAPNEV GTGLGEGYNI NIAWTGGLDP PMGDVEYLEA FRTIVKPVAK 910 920 930 940 950 960 EFDPDMVLVS AGFDALEGHT PPLGGYKVTA KCFGHLTKQL MTLADGRVVL ALEGGHDLTA 970 980 990 1000 1010 ICDASEACVN ALLGNELEPL AEDILHQSPN MNAVISLQKI IEIQSMSLKF S
	Isoform 2. Checksum: 095F634F623A684F	FASTA	923	101,805
	10 20 30 40 50 60 MHSMISSVDV KSEVPVGLEP ISPLDLRTDL RMMMPVVDPV VREKQLQQEL LLIQQQQQIQ 70 80 90 100 110 120 KQLLIAEFQK QHENLTRQHQ AQLQEHIKEL LAIKQQQELL EKEQKLEQQR QEQEVERHRR 130 140 150 160 170 180 EQQLPPLRGK DRGRERAVAS TEVKQKLQEF LLSKSATKDT PTNGKNHSVS RHPKLWYTAA 190 200 210 220 230 240 HHTSLDQSSP PLSGTSPSYK YTLPGAQDAK DDFPLRKTAS EPNLKVRSRL KQKVAERRSS 250 260 270 280 290 300 PLLRRKDGNV VTSFKKRMFE VTESSVSSSS PGSGPSSPNN GPTGSVTENE TSVLPPTPHA 310 320 330 340 350 360 EQMVSQQRIL IHEDSMNLLS LYTSPSLPNI TLGLPAVPSQ LNASNSLKEK QKCETQTLRQ 370 380 390 400 410 420 GVPLPGQYGG SIPASSSHPH VTLEGKPPNS SHQALLQHLL LKEQMRQQKL LVAGGVPLHP 430 440 450 460 470 480 QSPLATKERI SPGIRGTHKL PRHRPLNRTQ SAPLPQSTLA QLVIQQQHQQ FLEKQKQYQQ 490 500 510 520 530 540 QIHMNKPFLE PTHTRALSVR QAPLAAVGMD GLEKHRLVSR THSSPAASVL PHPAMDRPLQ 550 560 570 580 590 600 PGSATGIAYD PLMLKHQCVC GNSTTHPEHA GRIQSIWSRL QETGLLNKCE RIQGRKASLE 610 620 630 640 650 660 EIQLVHSEHH SLLYGTNPLD GQKLDPRILL GDDSQKFFSS LPCGGLGVDS DTIWNELHSS 670 680 690 700 710 720 GAARMAVGCV IELASKVASG ELKNGFAVVR PPGHHAEEST AMGFCFFNSV AITAKYLRDQ 730 740 750 760 770 780 LNISKILIVD LDVHHGNGTQ QAFYADPSIL YISLHRYDEG NFFPGSGAPN EVGTGLGEGY 790 800 810 820 830 840 NINIAWTGGL DPPMGDVEYL EAFRTIVKPV AKEFDPDMVL VSAGFDALEG HTPPLGGYKV 850 860 870 880 890 900 TAKCFGHLTK QLMTLADGRV VLALEGGHDL TAICDASEAC VNALLGNELE PLAEDILHQS 910 920 PNMNAVISLQ KIIEIQSMSL KFS
	Isoform 3 (HDRP) (MITR). Checksum: 02931057CF0E4C25	FASTA	590	65,887
	10 20 30 40 50 60 MHSMISSVDV KSEVPVGLEP ISPLDLRTDL RMMMPVVDPV VREKQLQQEL LLIQQQQQIQ 70 80 90 100 110 120 KQLLIAEFQK QHENLTRQHQ AQLQEHIKEL LAIKQQQELL EKEQKLEQQR QEQEVERHRR 130 140 150 160 170 180 EQQLPPLRGK DRGRERAVAS TEVKQKLQEF LLSKSATKDT PTNGKNHSVS RHPKLWYTAA 190 200 210 220 230 240 HHTSLDQSSP PLSGTSPSYK YTLPGAQDAK DDFPLRKTAS EPNLKVRSRL KQKVAERRSS 250 260 270 280 290 300 PLLRRKDGNV VTSFKKRMFE VTESSVSSSS PGSGPSSPNN GPTGSVTENE TSVLPPTPHA 310 320 330 340 350 360 EQMVSQQRIL IHEDSMNLLS LYTSPSLPNI TLGLPAVPSQ LNASNSLKEK QKCETQTLRQ 370 380 390 400 410 420 GVPLPGQYGG SIPASSSHPH VTLEGKPPNS SHQALLQHLL LKEQMRQQKL LVAGGVPLHP 430 440 450 460 470 480 QSPLATKERI SPGIRGTHKL PRHRPLNRTQ SAPLPQSTLA QLVIQQQHQQ FLEKQKQYQQ 490 500 510 520 530 540 QIHMNKLLSK SIEQLKQPGS HLEEAEEELQ GDQAMQEDRA PSSGNSTRSD SSACVDDTLG 550 560 570 580 590 QVGAVKVKEE PVDSDEDAQI QEMESGEQAA FMQQVIGKDL APGFVIKVII
	Isoform 4 (HDAC9a). Checksum: D3BFC3CE99E817DB	FASTA	879	97,471
	10 20 30 40 50 60 MHSMISSVDV KSEVPVGLEP ISPLDLRTDL RMMMPVVDPV VREKQLQQEL LLIQQQQQIQ 70 80 90 100 110 120 KQLLIAEFQK QHENLTRQHQ AQLQEHIKEL LAIKQQQELL EKEQKLEQQR QEQEVERHRR 130 140 150 160 170 180 EQQLPPLRGK DRGRERAVAS TEVKQKLQEF LLSKSATKDT PTNGKNHSVS RHPKLWYTAA 190 200 210 220 230 240 HHTSLDQSSP PLSGTSPSYK YTLPGAQDAK DDFPLRKTAS EPNLKVRSRL KQKVAERRSS 250 260 270 280 290 300 PLLRRKDGNV VTSFKKRMFE VTESSVSSSS PGSGPSSPNN GPTGSVTENE TSVLPPTPHA 310 320 330 340 350 360 EQMVSQQRIL IHEDSMNLLS LYTSPSLPNI TLGLPAVPSQ LNASNSLKEK QKCETQTLRQ 370 380 390 400 410 420 GVPLPGQYGG SIPASSSHPH VTLEGKPPNS SHQALLQHLL LKEQMRQQKL LVAGGVPLHP 430 440 450 460 470 480 QSPLATKERI SPGIRGTHKL PRHRPLNRTQ SAPLPQSTLA QLVIQQQHQQ FLEKQKQYQQ 490 500 510 520 530 540 QIHMNKLLSK SIEQLKQPGS HLEEAEEELQ GDQAMQEDRA PSSGNSTRSD SSACVDDTLG 550 560 570 580 590 600 QVGAVKVKEE PVDSDEDAQI QEMESGEQAA FMQQPFLEPT HTRALSVRQA PLAAVGMDGL 610 620 630 640 650 660 EKHRLVSRTH SSPAASVLPH PAMDRPLQPG SATGIAYDPL MLKHQCVCGN STTHPEHAGR 670 680 690 700 710 720 IQSIWSRLQE TGLLNKCERI QGRKASLEEI QLVHSEHHSL LYGTNPLDGQ KLDPRILLGD 730 740 750 760 770 780 DSQKFFSSLP CGGLGVDSDT IWNELHSSGA ARMAVGCVIE LASKVASGEL KNGFAVVRPP 790 800 810 820 830 840 GHHAEESTAM GFCFFNSVAI TAKYLRDQLN ISKILIVDLD VHHGNGTQQA FYADPSILYI 850 860 870 SLHRYDEGNF FPGSGAPNEV RFISLEPHFY LYLSGNCIA
	Isoform 5 (HDAC9b) (HDAC9fl). Checksum: EE7D908DF45D20CB	FASTA	1,066	117,208
	10 20 30 40 50 60 MHSMISSVDV KSEVPVGLEP ISPLDLRTDL RMMMPVVDPV VREKQLQQEL LLIQQQQQIQ 70 80 90 100 110 120 KQLLIAEFQK QHENLTRQHQ AQLQEHIKEL LAIKQQQELL EKEQKLEQQR QEQEVERHRR 130 140 150 160 170 180 EQQLPPLRGK DRGRERAVAS TEVKQKLQEF LLSKSATKDT PTNGKNHSVS RHPKLWYTAA 190 200 210 220 230 240 HHTSLDQSSP PLSGTSPSYK YTLPGAQDAK DDFPLRKTAS EPNLKVRSRL KQKVAERRSS 250 260 270 280 290 300 PLLRRKDGNV VTSFKKRMFE VTESSVSSSS PGSGPSSPNN GPTGSVTENE TSVLPPTPHA 310 320 330 340 350 360 EQMVSQQRIL IHEDSMNLLS LYTSPSLPNI TLGLPAVPSQ LNASNSLKEK QKCETQTLRQ 370 380 390 400 410 420 GVPLPGQYGG SIPASSSHPH VTLEGKPPNS SHQALLQHLL LKEQMRQQKL LVAGGVPLHP 430 440 450 460 470 480 QSPLATKERI SPGIRGTHKL PRHRPLNRTQ SAPLPQSTLA QLVIQQQHQQ FLEKQKQYQQ 490 500 510 520 530 540 QIHMNKLLSK SIEQLKQPGS HLEEAEEELQ GDQAMQEDRA PSSGNSTRSD SSACVDDTLG 550 560 570 580 590 600 QVGAVKVKEE PVDSDEDAQI QEMESGEQAA FMQQPFLEPT HTRALSVRQA PLAAVGMDGL 610 620 630 640 650 660 EKHRLVSRTH SSPAASVLPH PAMDRPLQPG SATGIAYDPL MLKHQCVCGN STTHPEHAGR 670 680 690 700 710 720 IQSIWSRLQE TGLLNKCERI QGRKASLEEI QLVHSEHHSL LYGTNPLDGQ KLDPRILLGD 730 740 750 760 770 780 DSQKFFSSLP CGGLGVDSDT IWNELHSSGA ARMAVGCVIE LASKVASGEL KNGFAVVRPP 790 800 810 820 830 840 GHHAEESTAM GFCFFNSVAI TAKYLRDQLN ISKILIVDLD VHHGNGTQQA FYADPSILYI 850 860 870 880 890 900 SLHRYDEGNF FPGSGAPNEV GTGLGEGYNI NIAWTGGLDP PMGDVEYLEA FRTIVKPVAK 910 920 930 940 950 960 EFDPDMVLVS AGFDALEGHT PPLGGYKVTA KCFGHLTKQL MTLADGRVVL ALEGGHDLTA 970 980 990 1000 1010 1020 ICDASEACVN ALLGNELEPL AEDILHQSPN MNAVISLQKI IEIQSKYWKS VRMVAVPRGC 1030 1040 1050 1060 ALAGAQLQEE TETVSALASL TVDVEQPFAQ EDSRTAGEPM EEEPAL
	Isoform 6. Checksum: 3BEFD662BF24E677	FASTA	1,025	112,451
	10 20 30 40 50 60 MHSMISSVDV KSEVPVGLEP ISPLDLRTDL RMMMPVVDPV VREKQLQQEL LLIQQQQQIQ 70 80 90 100 110 120 KQLLIAEFQK QHENLTRQHQ AQLQEHIKLQ QELLAIKQQQ ELLEKEQKLE QQRQEQEVER 130 140 150 160 170 180 HRREQQLPPL RGKDRGRERA VASTEVKQKL QEFLLSKSAT KDTPTNGKNH SVSRHPKLWY 190 200 210 220 230 240 TAAHHTSLDQ SSPPLSGTSP SYKYTLPGAQ DAKDDFPLRK TESSVSSSSP GSGPSSPNNG 250 260 270 280 290 300 PTGSVTENET SVLPPTPHAE QMVSQQRILI HEDSMNLLSL YTSPSLPNIT LGLPAVPSQL 310 320 330 340 350 360 NASNSLKEKQ KCETQTLRQG VPLPGQYGGS IPASSSHPHV TLEGKPPNSS HQALLQHLLL 370 380 390 400 410 420 KEQMRQQKLL VAGGVPLHPQ SPLATKERIS PGIRGTHKLP RHRPLNRTQS APLPQSTLAQ 430 440 450 460 470 480 LVIQQQHQQF LEKQKQYQQQ IHMNKLLSKS IEQLKQPGSH LEEAEEELQG DQAMQEDRAP 490 500 510 520 530 540 SSGNSTRSDS SACVDDTLGQ VGAVKVKEEP VDSDEDAQIQ EMESGEQAAF MQQPFLEPTH 550 560 570 580 590 600 TRALSVRQAP LAAVGMDGLE KHRLVSRTHS SPAASVLPHP AMDRPLQPGS ATGIAYDPLM 610 620 630 640 650 660 LKHQCVCGNS TTHPEHAGRI QSIWSRLQET GLLNKCERIQ GRKASLEEIQ LVHSEHHSLL 670 680 690 700 710 720 YGTNPLDGQK LDPRILLGDD SQKFFSSLPC GGLGVDSDTI WNELHSSGAA RMAVGCVIEL 730 740 750 760 770 780 ASKVASGELK NGFAVVRPPG HHAEESTAMG FCFFNSVAIT AKYLRDQLNI SKILIVDLDV 790 800 810 820 830 840 HHGNGTQQAF YADPSILYIS LHRYDEGNFF PGSGAPNEVG TGLGEGYNIN IAWTGGLDPP 850 860 870 880 890 900 MGDVEYLEAF RTIVKPVAKE FDPDMVLVSA GFDALEGHTP PLGGYKVTAK CFGHLTKQLM 910 920 930 940 950 960 TLADGRVVLA LEGGHDLTAI CDASEACVNA LLGNELEPLA EDILHQSPNM NAVISLQKII 970 980 990 1000 1010 1020 EIQSKYWKSV RMVAVPRGCA LAGAQLQEET ETVSALASLT VDVEQPFAQE DSRTAGEPME EEPAL
	Isoform 7. Checksum: 5DFBB8DD4C6547C5	FASTA	1,069	117,577
	10 20 30 40 50 60 MHSMISSVDV KSEVPVGLEP ISPLDLRTDL RMMMPVVDPV VREKQLQQEL LLIQQQQQIQ 70 80 90 100 110 120 KQLLIAEFQK QHENLTRQHQ AQLQEHIKLQ QELLAIKQQQ ELLEKEQKLE QQRQEQEVER 130 140 150 160 170 180 HRREQQLPPL RGKDRGRERA VASTEVKQKL QEFLLSKSAT KDTPTNGKNH SVSRHPKLWY 190 200 210 220 230 240 TAAHHTSLDQ SSPPLSGTSP SYKYTLPGAQ DAKDDFPLRK TASEPNLKVR SRLKQKVAER 250 260 270 280 290 300 RSSPLLRRKD GNVVTSFKKR MFEVTESSVS SSSPGSGPSS PNNGPTGSVT ENETSVLPPT 310 320 330 340 350 360 PHAEQMVSQQ RILIHEDSMN LLSLYTSPSL PNITLGLPAV PSQLNASNSL KEKQKCETQT 370 380 390 400 410 420 LRQGVPLPGQ YGGSIPASSS HPHVTLEGKP PNSSHQALLQ HLLLKEQMRQ QKLLVAGGVP 430 440 450 460 470 480 LHPQSPLATK ERISPGIRGT HKLPRHRPLN RTQSAPLPQS TLAQLVIQQQ HQQFLEKQKQ 490 500 510 520 530 540 YQQQIHMNKL LSKSIEQLKQ PGSHLEEAEE ELQGDQAMQE DRAPSSGNST RSDSSACVDD 550 560 570 580 590 600 TLGQVGAVKV KEEPVDSDED AQIQEMESGE QAAFMQQPFL EPTHTRALSV RQAPLAAVGM 610 620 630 640 650 660 DGLEKHRLVS RTHSSPAASV LPHPAMDRPL QPGSATGIAY DPLMLKHQCV CGNSTTHPEH 670 680 690 700 710 720 AGRIQSIWSR LQETGLLNKC ERIQGRKASL EEIQLVHSEH HSLLYGTNPL DGQKLDPRIL 730 740 750 760 770 780 LGDDSQKFFS SLPCGGLGVD SDTIWNELHS SGAARMAVGC VIELASKVAS GELKNGFAVV 790 800 810 820 830 840 RPPGHHAEES TAMGFCFFNS VAITAKYLRD QLNISKILIV DLDVHHGNGT QQAFYADPSI 850 860 870 880 890 900 LYISLHRYDE GNFFPGSGAP NEVGTGLGEG YNINIAWTGG LDPPMGDVEY LEAFRTIVKP 910 920 930 940 950 960 VAKEFDPDMV LVSAGFDALE GHTPPLGGYK VTAKCFGHLT KQLMTLADGR VVLALEGGHD 970 980 990 1000 1010 1020 LTAICDASEA CVNALLGNEL EPLAEDILHQ SPNMNAVISL QKIIEIQSKY WKSVRMVAVP 1030 1040 1050 1060 RGCALAGAQL QEETETVSAL ASLTVDVEQP FAQEDSRTAG EPMEEEPAL

References

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[1]	"Cloning and characterization of a histone deacetylase, HDAC9." Zhou X., Marks P.A., Rifkind R.A., Richon V.M. Proc. Natl. Acad. Sci. U.S.A. 98:10572-10577(2001) [PubMed: 11535832] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [MRNA] (ISOFORMS 1 AND 4), TISSUE SPECIFICITY, INTERACTION WITH MEF2, FUNCTION, ENZYME REGULATION. Tissue: Brain.
[2]	"Molecular characterization of a familial translocation implicates disruption of HDAC9 and possible position effect on TGFbeta2 in the pathogenesis of Peters' anomaly." David D., Cardoso J., Marques B., Marques R., Silva E.D., Santos H., Boavida M.G. Genomics 81:489-503(2003) [PubMed: 12706107] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [MRNA] (ISOFORM 5), TISSUE SPECIFICITY, CHROMOSOMAL TRANSLOCATION WITH TGFB2. Tissue: Lens.
[3]	"The histone deacetylase 9 gene encodes multiple protein isoforms." Petrie K., Guidez F., Howell L., Healy L., Waxman S., Greaves M., Zelent A. J. Biol. Chem. 278:16059-16072(2003) [PubMed: 12590135] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [MRNA] (ISOFORM 7), TISSUE SPECIFICITY, ALTERNATIVE SPLICING (ISOFORMS 3; 6 AND 7), INTERACTION WITH ETV6; NCOR1 AND BCL6, FUNCTION, SUBCELLULAR LOCATION, SUMOYLATION. Tissue: Brain.
[4]	"Prediction of the coding sequences of unidentified human genes. XI. The complete sequences of 100 new cDNA clones from brain which code for large proteins in vitro." Nagase T., Ishikawa K., Suyama M., Kikuno R., Miyajima N., Tanaka A., Kotani H., Nomura N., Ohara O. DNA Res. 5:277-286(1998) [PubMed: 9872452] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] (ISOFORM 3). Tissue: Brain.
[5]	"The DNA sequence of human chromosome 7." Hillier L.W., Fulton R.S., Fulton L.A., Graves T.A., Pepin K.H., Wagner-McPherson C., Layman D., Maas J., Jaeger S., Walker R., Wylie K., Sekhon M., Becker M.C., O'Laughlin M.D., Schaller M.E., Fewell G.A., Delehaunty K.D., Miner T.L. , Nash W.E., Cordes M., Du H., Sun H., Edwards J., Bradshaw-Cordum H., Ali J., Andrews S., Isak A., Vanbrunt A., Nguyen C., Du F., Lamar B., Courtney L., Kalicki J., Ozersky P., Bielicki L., Scott K., Holmes A., Harkins R., Harris A., Strong C.M., Hou S., Tomlinson C., Dauphin-Kohlberg S., Kozlowicz-Reilly A., Leonard S., Rohlfing T., Rock S.M., Tin-Wollam A.-M., Abbott A., Minx P., Maupin R., Strowmatt C., Latreille P., Miller N., Johnson D., Murray J., Woessner J.P., Wendl M.C., Yang S.-P., Schultz B.R., Wallis J.W., Spieth J., Bieri T.A., Nelson J.O., Berkowicz N., Wohldmann P.E., Cook L.L., Hickenbotham M.T., Eldred J., Williams D., Bedell J.A., Mardis E.R., Clifton S.W., Chissoe S.L., Marra M.A., Raymond C., Haugen E., Gillett W., Zhou Y., James R., Phelps K., Iadanoto S., Bubb K., Simms E., Levy R., Clendenning J., Kaul R., Kent W.J., Furey T.S., Baertsch R.A., Brent M.R., Keibler E., Flicek P., Bork P., Suyama M., Bailey J.A., Portnoy M.E., Torrents D., Chinwalla A.T., Gish W.R., Eddy S.R., McPherson J.D., Olson M.V., Eichler E.E., Green E.D., Waterston R.H., Wilson R.K. Nature 424:157-164(2003) [PubMed: 12853948] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
[6]	"The status, quality, and expansion of the NIH full-length cDNA project: the Mammalian Gene Collection (MGC)." The MGC Project Team Genome Res. 14:2121-2127(2004) [PubMed: 15489334] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] OF 2-1011 (ISOFORM 6).
[7]	"HDAC4, a human histone deacetylase related to yeast HDA1, is a transcriptional corepressor." Wang A.H., Bertos N.R., Vezmar M., Pelletier N., Crosato M., Heng H.H., Th'ng J., Han J., Yang X.-J. Mol. Cell. Biol. 19:7816-7827(1999) [PubMed: 10523670] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [MRNA] OF 99-650 (ISOFORM 2). Tissue: Brain.
[8]	"MEF-2 function is modified by a novel co-repressor, MITR." Sparrow D.B., Miska E.A., Langley E., Reynaud-Deonauth S., Kotecha S., Towers N., Spohr G., Kouzarides T., Mohun T.J. EMBO J. 18:5085-5098(1999) [PubMed: 10487760] [Abstract] Cited for: IDENTIFICATION (ISOFORM 3), INTERACTION WITH HDAC1, FUNCTION.
[9]	"Identification of a transcriptional repressor related to the noncatalytic domain of histone deacetylases 4 and 5." Zhou X., Richon V.M., Rifkind R.A., Marks P.A. Proc. Natl. Acad. Sci. U.S.A. 97:1056-1061(2000) [PubMed: 10655483] [Abstract] Cited for: IDENTIFICATION (ISOFORM 3), TISSUE SPECIFICITY, INTERACTION WITH HDAC1 AND HDAC3, FUNCTION.
[10]	"HDAC4 deacetylase associates with and represses the MEF2 transcription factor." Miska E.A., Karlsson C., Langley E., Nielsen S.J., Pines J., Kouzarides T. EMBO J. 18:5099-5107(1999) [PubMed: 10487761] [Abstract] Cited for: INTERACTION WITH MEF2.
[11]	"The SUMO E3 ligase RanBP2 promotes modification of the HDAC4 deacetylase." Kirsh O., Seeler J.-S., Pichler A., Gast A., Mueller S., Miska E., Mathieu M., Harel-Bellan A., Kouzarides T., Melchior F., Dejean A. EMBO J. 21:2682-2691(2002) [PubMed: 12032081] [Abstract] Cited for: SUMOYLATION.
[12]	"Global, in vivo, and site-specific phosphorylation dynamics in signaling networks." Olsen J.V., Blagoev B., Gnad F., Macek B., Kumar C., Mortensen P., Mann M. Cell 127:635-648(2006) [PubMed: 17081983] [Abstract] Cited for: PHOSPHORYLATION [LARGE SCALE ANALYSIS] AT TYR-1010 (ISOFORM 7), MASS SPECTROMETRY. Tissue: Epithelium.
[13]	"Neuroprotection by histone deacetylase-related protein." Morrison B.E., Majdzadeh N., Zhang X., Lyles A., Bassel-Duby R., Olson E.N., D'Mello S.R. Mol. Cell. Biol. 26:3550-3564(2006) [PubMed: 16611996] [Abstract] Cited for: FUNCTION.
+	Additional computationally mapped references.

Cross-references

Sequence databases
EMBLGenBankDDBJ	AY032737 mRNA. Translation: AAK66821.1. AY032738 mRNA. Translation: AAK66822.1. AJ459808 mRNA. Translation: CAD30851.1. AY197371 mRNA. Translation: AAO27363.1. AB018287 mRNA. Translation: BAA34464.2. Different initiation. AC004994 Genomic DNA. Translation: AAC78618.2. BC111735 mRNA. Translation: AAI11736.1. Frameshift. AF124924 mRNA. Translation: AAF04254.1.
IPI	IPI00215728. IPI00215729. IPI00215730. IPI00232519. IPI00328781. IPI00829837. IPI00829883.
RefSeq	NP_055522.1. NP_478056.1. NP_478057.1. NP_848510.1. NP_848512.1.
UniGene	Hs.196054
3D structure databases
SMR	Q9UKV0. Positions 633-1005.
ModBase	Search...
Protein-protein interaction databases
IntAct	Q9UKV0. 19 interactions.
Proteomic databases
PRIDE	Q9UKV0.
Genome annotation databases
Ensembl	ENSG00000048052. Homo sapiens. [Contig view]
GeneID	9734.
KEGG	hsa:9734.
Organism-specific databases
GeneCards	GC07P018501.
HGNC	HGNC:14065. HDAC9.
MIM	604229. phenotype. 606543. gene.
Orphanet	708. Peters anomaly.
PharmGKB	PA38377.
HUGE	Search...
GenAtlas	Search...
Phylogenomic databases
HOGENOM	Q9UKV0.
HOVERGEN	Q9UKV0.
OMA	Q9UKV0. MFEVTES.
Enzyme and pathway databases
Pathway_Interaction_DB	hdac_classi_pathway. Signaling events mediated by HDAC Class I. hdac_classii_pathway. Signaling events mediated by HDAC Class II.
Reactome	REACT_71. Gene Expression.
Gene expression databases
ArrayExpress	Q9UKV0.
Bgee	Q9UKV0.
CleanEx	HS_HDAC7. HS_HDAC9.
GermOnline	ENSG00000048052. Homo sapiens.
Family and domain databases
InterPro	IPR000286. His_deacetylse. IPR017320. Histone_deAcase_II_euk. [Graphical view]
Gene3D	G3DSA:3.40.800.20. His_deacetylse. 1 hit.
PANTHER	PTHR10625. His_deacetylse. 1 hit.
Pfam	PF00850. Hist_deacetyl. 1 hit. [Graphical view]
PIRSF	PIRSF037911. HDAC_II_euk. 1 hit.
PRINTS	PR01270. HDASUPER.
ProtoNet	Search...
Other Resources
DrugBank	DB00313. Valproic Acid.
NextBio	36620.
SOURCE	Search...

Entry information

Entry name

HDAC9_HUMAN

Accession

Primary (citable) accession number: Q9UKV0
Secondary accession number(s): O94845 , O95028, Q2M2R6, Q86SL1, Q86US3

Entry history

Integrated into UniProtKB/Swiss-Prot:	December 1, 2000
Last sequence update:	October 18, 2001
Last modified:	June 16, 2009
This is version 88 of the entry and version 2 of the sequence. [Complete history]

Entry status

Reviewed (UniProtKB/Swiss-Prot)

Annotation project

HPI (Human Proteome Initiative)