IPO9

Summary

IPO9 (importin 9, HGNC:19425) is a protein-coding gene on chromosome 1q32.1, encoding Importin-9 (Q96P70). Nuclear transport receptor that mediates nuclear import of proteins, such as histones, proteasome and actin. It is a selective cancer dependency (DepMap: 81.7% of cell lines).

Enables histone binding activity; histone chaperone activity; and nuclear import signal receptor activity. Involved in proteasome localization and protein import into nucleus. Located in cytosol and nucleoplasm.

Source: NCBI Gene 55705 — RefSeq curated summary.

At a glance

• GWAS associations: 4
• Clinical variants (ClinVar): 126 total
• Druggable target: yes
• Cancer dependency (DepMap): dependent in 81.7% of screened cell lines
• MANE Select transcript: NM_018085

Identifiers

Gene identifiers

Field	Value
HGNC ID	HGNC:19425
Approved symbol	IPO9
Name	importin 9
Location	1q32.1
Locus type	gene with protein product
Status	Approved
Aliases	Imp9, FLJ10402
Ensembl gene	ENSG00000198700
Ensembl biotype	protein_coding
OMIM	620893
Entrez	55705

Gene structure

Transcript identifiers

Ensembl transcripts: 20 — 18 protein_coding, 2 protein_coding_CDS_not_defined

ENST00000361565, ENST00000456707, ENST00000464348, ENST00000479374, ENST00000886559, ENST00000886560, ENST00000886561, ENST00000886562, ENST00000886563, ENST00000926156, ENST00000926157, ENST00000926158, ENST00000926159, ENST00000926160, ENST00000926161, ENST00000926162, ENST00000926163, ENST00000926164, ENST00000926165, ENST00000926166

RefSeq mRNA: 1 — MANE Select: NM_018085 NM_018085

CCDS: CCDS1415

Canonical transcript exons

ENST00000361565 — 24 exons

Exon	Start	End
ENSE00001434838	201854823	201854923
ENSE00001434876	201872828	201872961
ENSE00001434913	201875152	201875228
ENSE00001434944	201874832	201874936
ENSE00001435320	201874250	201874372
ENSE00001435402	201871161	201871327
ENSE00001435474	201853011	201853097
ENSE00001435626	201855783	201855934
ENSE00001435653	201855124	201855182
ENSE00001435727	201854595	201854714
ENSE00001435758	201857096	201857194
ENSE00001435798	201863448	201863607
ENSE00001435853	201858447	201858553
ENSE00001435931	201870583	201870858
ENSE00001435957	201869590	201869718
ENSE00001435980	201868648	201868796
ENSE00001436031	201866733	201866959
ENSE00001436244	201858855	201858994
ENSE00001444083	201875944	201884291
ENSE00001821710	201829157	201829372
ENSE00003548972	201848393	201848594
ENSE00003611153	201847279	201847340
ENSE00003643024	201847552	201847638
ENSE00003656006	201852104	201852192

Expression profiles

Bgee: expression breadth ubiquitous, 288 present calls, max score 96.29.

FANTOM5 (CAGE): breadth ubiquitous, TPM avg 18.0554 / max 325.6449, expressed in 1798 samples.

FANTOM5 promoters (5 alternative TSS)

Promoter ID	TPM avg	Samples expressed
7756	16.3347	1796
7758	0.6991	288
7760	0.4455	187
7759	0.4337	160
7755	0.1423	35

Top tissues by expression

294 total, by Bgee expression score (0-100, higher = more expressed):

Tissue	Anatomy ID	Expression score	Quality
nipple	UBERON:0002030	96.29	gold quality
ventricular zone	UBERON:0003053	95.67	gold quality
pylorus	UBERON:0001166	95.62	gold quality
parietal lobe	UBERON:0001872	94.91	gold quality
ganglionic eminence	UBERON:0004023	94.89	gold quality
postcentral gyrus	UBERON:0002581	94.87	gold quality
ventral tegmental area	UBERON:0002691	94.87	gold quality
entorhinal cortex	UBERON:0002728	94.75	gold quality
embryo	UBERON:0000922	94.67	gold quality
cardia of stomach	UBERON:0001162	94.58	gold quality
inferior vagus X ganglion	UBERON:0005363	94.37	gold quality
Brodmann (1909) area 46	UBERON:0006483	94.31	gold quality
superior vestibular nucleus	UBERON:0007227	94.25	gold quality
substantia nigra pars reticulata	UBERON:0001966	94.17	gold quality
lateral globus pallidus	UBERON:0002476	94.16	gold quality
lateral nuclear group of thalamus	UBERON:0002736	94.04	gold quality
medulla oblongata	UBERON:0001896	94.01	gold quality
trigeminal ganglion	UBERON:0001675	94.00	gold quality
stromal cell of endometrium	CL:0002255	93.94	gold quality
subthalamic nucleus	UBERON:0001906	93.90	gold quality
caput epididymis	UBERON:0004358	93.85	gold quality
orbitofrontal cortex	UBERON:0004167	93.81	gold quality
dorsal root ganglion	UBERON:0000044	93.76	gold quality
cauda epididymis	UBERON:0004360	93.76	gold quality
renal medulla	UBERON:0000362	93.66	gold quality
urethra	UBERON:0000057	93.65	gold quality
trabecular bone tissue	UBERON:0002483	93.41	gold quality
lower lobe of lung	UBERON:0008949	93.36	gold quality
substantia nigra pars compacta	UBERON:0001965	93.35	gold quality
dorsal plus ventral thalamus	UBERON:0001897	93.32	gold quality

Single-cell (SCXA)

Detected in 1 experiment(s), a significant marker in 1.

Experiment	Marker?	Max mean expression
E-ANND-3	yes	9.72

Regulation

Is transcription factor: no

miRNA regulators (miRDB)

288 targeting IPO9, top 30 by miRDB confidence (max_score; target_count = how many genes the miRNA targets in total — lower means more specific):

miRNA	Max score	Avg score	miRNA target_count
HSA-MIR-6867-5P	100.00	82.21	3464
HSA-MIR-3120-5P	100.00	65.56	965
HSA-MIR-518D-5P	100.00	67.51	979
HSA-MIR-518E-5P	100.00	67.66	954
HSA-MIR-518F-5P	100.00	67.51	979
HSA-MIR-519A-5P	100.00	67.66	954
HSA-MIR-519B-5P	100.00	67.66	954
HSA-MIR-519C-5P	100.00	67.66	954
HSA-MIR-520C-5P	100.00	67.51	979
HSA-MIR-522-5P	100.00	67.66	954
HSA-MIR-523-5P	100.00	67.66	954
HSA-MIR-526A-5P	100.00	67.51	979
HSA-MIR-3064-3P	100.00	70.09	1254
HSA-MIR-29A-3P	100.00	73.11	1835
HSA-MIR-29B-3P	100.00	73.18	1833
HSA-MIR-29C-3P	100.00	73.15	1833
HSA-MIR-6833-3P	100.00	70.63	3197
HSA-MIR-150-5P	99.99	66.69	1976
HSA-MIR-548AW	99.99	72.57	3559
HSA-MIR-3667-3P	99.99	67.17	1636
HSA-MIR-10401-5P	99.99	65.79	948
HSA-MIR-4500	99.99	72.72	2367
HSA-MIR-520G-5P	99.99	66.76	658
HSA-MIR-8068	99.98	73.85	2376
HSA-MIR-4715-3P	99.98	66.03	670
HSA-MIR-6888-3P	99.97	65.95	1170
HSA-MIR-3148	99.97	75.06	6478
HSA-MIR-6778-3P	99.96	67.29	2693
HSA-MIR-302E	99.96	70.74	2669
HSA-MIR-548AJ-3P	99.96	73.38	5345

Functional genomics

DepMap (CRISPR cell-line fitness): dependent in 81.7% of screened cell lines.

Literature-anchored findings (GeneRIF, showing 4)

• specific interaction between a member of the importin beta/karyopherin beta superfamily, importin 9, and the A subunit of PP2A (PR65) (PMID:12670497)
• IPO9 associates with 2 stable stem-loop structures of the IFN-epsilon 5’UTR. IPO9 overexpression decreased, and IPO9 silencing increased basal IFN-epsilon expression. (PMID:23851686)
• CFL1 itself does not translocate actin into the cell nucleus but this transport requires the functional expression of IPO9. (PMID:26934847)
• Like many histone chaperones, which prevent inappropriate non-nucleosomal interactions, Importin-9 also sequesters H2A-H2B from DNA. Importin-9 appears to act as a storage chaperone for H2A-H2B while escorting it to the nucleus. (PMID:30855230)

Cross-species orthologs

4 orthologs

Organism	Symbol	Gene ID
danio_rerio	ipo9	ENSDARG00000016753
mus_musculus	Ipo9	ENSMUSG00000041879
rattus_norvegicus	Ipo9	ENSRNOG00000007418
drosophila_melanogaster	Ipo9	FBGN0037894

Paralogs (4): IPO11 (ENSG00000086200), CSE1L (ENSG00000124207), IPO8 (ENSG00000133704), IPO7 (ENSG00000205339)

Protein

Protein identifiers

Importin-9 — Q96P70 (reviewed: Q96P70)

Alternative names: Ran-binding protein 9

All UniProt accessions (2): Q96P70, Q5SVH5

UniProt curated annotations — full annotation on UniProt →

Function. Nuclear transport receptor that mediates nuclear import of proteins, such as histones, proteasome and actin. Serves as receptor for nuclear localization signals (NLS) in cargo substrates. Is thought to mediate docking of the importin/substrate complex to the nuclear pore complex (NPC) through binding to nucleoporin and the complex is subsequently translocated through the pore by an energy requiring, Ran-dependent mechanism. At the nucleoplasmic side of the NPC, Ran binds to the importin, the importin/substrate complex dissociates and importin is re-exported from the nucleus to the cytoplasm where GTP hydrolysis releases Ran. The directionality of nuclear import is thought to be conferred by an asymmetric distribution of the GTP- and GDP-bound forms of Ran between the cytoplasm and nucleus. Mediates the import of pre-assembled proteasomes into the nucleus; AKIRIN2 acts as a molecular bridge between IPO9 and the proteasome complex. Mediates the nuclear import of histones H2A, H2B, H4 and H4. In addition to nuclear import, also acts as a chaperone for histones by preventing inappropriate non-nucleosomal interactions. Mediates the nuclear import of actin.

Subunit / interactions. Interacts with histones H2A, H2B, H3 and H4. The binding is coupled to RanGTP cycles. Interacts with AKIRIN2; promoting association with pre-assembled proteasomes. Associates with pre-assembled proteasomes; interaction is indirect and mediated via interaction with AKIRIN2. Interacts with PPP2R1A and PPP2R1B.

Subcellular location. Cytoplasm. Nucleus.

Similarity. Belongs to the importin beta family.

RefSeq proteins (1): NP_060555* (*=MANE)

Domains & families (InterPro)

ID	Name	Type
IPR001494	Importin-beta_N	Domain
IPR011989	ARM-like	Homologous_superfamily
IPR016024	ARM-type_fold	Homologous_superfamily
IPR056840	HEAT_IPO9_central	Domain
IPR058669	TPR_IPO7/11-like	Domain

Pfam: PF03810, PF25018, PF25758

UniProt features (83 total): helix 59, turn 8, strand 6, sequence conflict 4, initiator methionine 1, chain 1, domain 1, region of interest 1, compositionally biased region 1, modified residue 1

Structure

Experimental structures (PDB)

7 structures.

PDB	Method	Resolution (Å)
6N1Z	X-RAY DIFFRACTION	2.7
9QON	ELECTRON MICROSCOPY	3.2
9QOO	ELECTRON MICROSCOPY	3.3
10SM	ELECTRON MICROSCOPY	3.5
9QOP	ELECTRON MICROSCOPY	3.7
8F7A	ELECTRON MICROSCOPY	3.78
9QNO	ELECTRON MICROSCOPY	4.2

Predicted structure (AlphaFold)

Model	pLDDT	Fraction very-high
AF-Q96P70-F1	88.37	0.73

Functional residue map

Curated UniProt residues grouped by drug-discovery relevance — catalytic, ligand-binding, modification, and mutation-validated positions. Source: UniProtKB sequence features.

Post-translational modifications (1): 2

Function

Pathways and Gene Ontology

Reactome pathways

0 pathways

MSigDB gene sets: 190 (showing top): TGCGCANK_UNKNOWN, GCM_GSPT1, GOBP_INTRACELLULAR_PROTEIN_TRANSPORT, STARK_PREFRONTAL_CORTEX_22Q11_DELETION_DN, MORF_BRCA1, GOBP_ESTABLISHMENT_OF_PROTEIN_LOCALIZATION_TO_ORGANELLE, GOMF_GTPASE_BINDING, MORF_RAD51L3, GOBP_NUCLEAR_TRANSPORT, ONKEN_UVEAL_MELANOMA_UP, MORF_CTSB, MORF_IL4, MORF_PRKCA, MORF_THPO, DODD_NASOPHARYNGEAL_CARCINOMA_UP

GO Biological Process (5): protein import into nucleus (GO:0006606), proteasome localization (GO:0031144), intracellular protein transport (GO:0006886), proteasomal protein catabolic process (GO:0010498), protein transport (GO:0015031)

GO Molecular Function (5): small GTPase binding (GO:0031267), histone binding (GO:0042393), nuclear import signal receptor activity (GO:0061608), histone chaperone activity (GO:0140713), protein binding (GO:0005515)

GO Cellular Component (6): nuclear envelope (GO:0005635), nucleoplasm (GO:0005654), cytoplasm (GO:0005737), cytosol (GO:0005829), membrane (GO:0016020), nucleus (GO:0005634)

GO top-level categories

Rollup of top GO terms by namespace:

Category	Terms
cellular anatomical structure	4
import into nucleus	2
intracellular protein localization	2
intracellular protein transport	1
protein localization to nucleus	1
establishment of protein localization to organelle	1
protein-containing complex localization	1
protein transport	1
intracellular transport	1
protein catabolic process	1
transport	1
establishment of protein localization	1
GTPase binding	1
protein binding	1
nucleocytoplasmic carrier activity	1
histone binding	1
protein carrier activity	1
binding	1
nucleus	1
endomembrane system	1
organelle envelope	1
nuclear lumen	1
intracellular anatomical structure	1
cytoplasm	1
intracellular membrane-bounded organelle	1

Protein interactions and networks

STRING

1260 interactions, top by confidence (×1000):

Protein A	Protein B	Partner UniProt	Score
IPO9	TNPO2	O14787	874
IPO9	IPO7	O95373	840
IPO9	XPO6	Q96QU8	828
IPO9	IPO8	O15397	787
IPO9	TNPO1	Q92973	778
IPO9	PFN4	Q8NHR9	770
IPO9	IPO13	O94829	759
IPO9	IPO4	Q8TEX9	742
IPO9	CFL1	P23528	729
IPO9	CFL2	Q9Y281	725
IPO9	PFN3	P60673	718
IPO9	XPOT	O43592	704
IPO9	PFN1	P07737	693
IPO9	IPO5	O00410	674
IPO9	XPO1	O14980	652

IntAct

222 interactions, top by confidence:

A	B	Type	Score
ANP32E	H2AZ1	psi-mi:“MI:0915”(physical association)	0.770
H2AZ1	ZNHIT1	psi-mi:“MI:0914”(association)	0.770
H2AX	PPM1G	psi-mi:“MI:0914”(association)	0.730
KLHDC2	CUL2	psi-mi:“MI:0914”(association)	0.730
CFTR	ESYT2	psi-mi:“MI:0914”(association)	0.710
CFTR	ESYT2	psi-mi:“MI:2364”(proximity)	0.710
H2AC4	PPM1G	psi-mi:“MI:0914”(association)	0.670
H2BC1	PPM1G	psi-mi:“MI:0914”(association)	0.640
TNFSF13B	IPO8	psi-mi:“MI:0914”(association)	0.640
MAPK7	PFDN6	psi-mi:“MI:0914”(association)	0.640
SLC16A3	CASK	psi-mi:“MI:0914”(association)	0.590
BROX	TCEA1	psi-mi:“MI:0914”(association)	0.560
YWHAG	SHTN1	psi-mi:“MI:0914”(association)	0.560

BioGRID (304): IPO9 (Affinity Capture-MS), IPO9 (Affinity Capture-MS), IPO9 (Affinity Capture-MS), IPO9 (Affinity Capture-MS), IPO9 (Affinity Capture-MS), IPO9 (Affinity Capture-MS), IPO9 (Affinity Capture-MS), IPO9 (Affinity Capture-MS), IPO9 (Affinity Capture-RNA), IPO7 (Co-fractionation), IPO9 (Co-fractionation), IPO9 (Co-fractionation), IPO9 (Co-fractionation), IPO9 (Co-fractionation), IPO9 (Co-fractionation)

ESM2 similar proteins: A1CAU2, A1DEK2, A2QMS5, A3LWK3, A4RMB1, A5DLM5, A5E7I7, A6RVT8, A7EPT5, A8NU66, A8Q513, B0DAD3, B0Y4D6, B2AXG6, G5ED41, O13671, O59809, O74476, O94258, P0CN64, P0CN65, P40069, P46970, P53067, Q02932, Q06142, Q0CIL3, Q0V6W0, Q12754, Q1DY99, Q2H6R9, Q2U3V3, Q4PC84, Q4WUV9, Q5ASE3, Q5BAH2, Q6C715, Q6GMY9, Q753A0, Q7PC79

Diamond homologs: Q91YE6, Q96P70, Q9VGP4

SIGNOR signaling

1 interactions.

A	Effect	B	Mechanism
ATM	“up-regulates activity”	IPO9	phosphorylation

Enriched among interaction partners

Reactome pathways and GO biological processes over-represented among this gene’s 215 IntAct physical interaction partners (hypergeometric vs the genome-wide background, BH-FDR, gene-set size 15–500, ranked by fold). A functional readout of the neighbourhood — distinct from this gene’s own memberships above, and biased toward well-studied / hub proteins, so read it as themes rather than proof.

Reactome pathways:

Pathway	Partners	Fold	FDR
Activation of BAD and translocation to mitochondria	7	35.3×	3e-07
Chk1/Chk2(Cds1) mediated inactivation of Cyclin B:Cdk1 complex	6	26.7×	3e-06
SARS-CoV-1 targets host intracellular signalling and regulatory pathways	6	26.7×	3e-06
Activation of BH3-only proteins	7	23.0×	2e-06
Attachment and Entry	5	19.9×	6e-05
Respiratory syncytial virus (RSV) attachment and entry	5	16.4×	2e-04
RHO GTPases activate PKNs	7	14.7×	1e-05
Intrinsic Pathway for Apoptosis	7	13.6×	1e-05

GO biological processes:

GO term	Partners	Fold	FDR
heterochromatin formation	9	11.8×	9e-05

Disease & clinical

Clinical variants and AI predictions

ClinVar

126 variants total. Per-class counts are floors (≥ shown; pagination cap):

Classification	Count (floor)
Pathogenic	0
Likely pathogenic	0
Uncertain significance	90
Likely benign	1
Benign	1

Top pathogenic / likely-pathogenic (0)

SpliceAI

3456 predictions. Top by Δscore:

Variant	Effect	Δscore
1:201829276:G:GT	donor_gain	1.0000
1:201829321:G:GT	donor_gain	1.0000
1:201829498:G:GT	donor_gain	1.0000
1:201847275:TCA:T	acceptor_loss	1.0000
1:201847276:CA:C	acceptor_loss	1.0000
1:201847277:A:AG	acceptor_gain	1.0000
1:201847277:A:AT	acceptor_loss	1.0000
1:201847278:G:GC	acceptor_gain	1.0000
1:201847278:GA:G	acceptor_gain	1.0000
1:201847278:GAAT:G	acceptor_gain	1.0000
1:201847339:AGGTA:A	donor_loss	1.0000
1:201847341:G:A	donor_loss	1.0000
1:201847342:T:G	donor_loss	1.0000
1:201847550:AGCT:A	acceptor_gain	1.0000
1:201847551:GCT:G	acceptor_gain	1.0000
1:201847551:GCTG:G	acceptor_gain	1.0000
1:201848387:TTACA:T	acceptor_loss	1.0000
1:201848388:TACAG:T	acceptor_loss	1.0000
1:201848389:ACAGG:A	acceptor_loss	1.0000
1:201848390:CAGG:C	acceptor_loss	1.0000
1:201848391:A:AC	acceptor_loss	1.0000
1:201848391:A:AG	acceptor_gain	1.0000
1:201848392:G:A	acceptor_loss	1.0000
1:201848392:G:GG	acceptor_gain	1.0000
1:201848392:GGCAA:G	acceptor_gain	1.0000
1:201852099:TGTAG:T	acceptor_loss	1.0000
1:201852101:TA:T	acceptor_loss	1.0000
1:201852102:A:AG	acceptor_gain	1.0000
1:201852103:G:GC	acceptor_loss	1.0000
1:201852103:G:GG	acceptor_gain	1.0000

AlphaMissense

6858 scored. Top likely-pathogenic:

Variant	Protein change	am_pathogenicity
1:201847591:T:A	W89R	1.000
1:201847591:T:C	W89R	1.000
1:201847593:G:C	W89C	1.000
1:201847593:G:T	W89C	1.000
1:201848466:C:A	A129D	1.000
1:201848487:C:A	A136D	1.000
1:201848574:G:A	G165E	1.000
1:201848589:T:C	L170P	1.000
1:201871278:T:A	W843R	1.000
1:201871278:T:C	W843R	1.000
1:201829295:T:C	L29P	0.999
1:201829358:T:C	L50P	0.999
1:201847281:T:C	F56L	0.999
1:201847283:T:A	F56L	0.999
1:201847283:T:G	F56L	0.999
1:201847294:T:G	L60W	0.999
1:201847336:G:C	R74P	0.999
1:201847553:T:C	L76P	0.999
1:201847556:C:A	A77E	0.999
1:201847562:T:A	V79D	0.999
1:201847588:C:G	H88D	0.999
1:201847592:G:C	W89S	0.999
1:201848454:G:C	R125P	0.999
1:201848459:A:C	S127R	0.999
1:201848461:T:A	S127R	0.999
1:201848461:T:G	S127R	0.999
1:201848480:G:C	A134P	0.999
1:201848486:G:C	A136P	0.999
1:201848498:T:A	W140R	0.999
1:201848498:T:C	W140R	0.999

dbSNP variants (sampled 300 via entrez): RS1000094957 (1:201827513 A>G), RS1000135102 (1:201871638 C>G), RS1000153396 (1:201831987 C>T), RS1000156459 (1:201856210 A>G), RS1000187189 (1:201873814 C>T), RS1000265725 (1:201853776 G>C), RS1000340985 (1:201840698 G>A,C), RS1000373568 (1:201836189 G>A,C), RS1000418632 (1:201856534 T>C,G), RS1000486207 (1:201871876 G>A), RS1000579726 (1:201866578 T>G), RS1000618567 (1:201867226 G>A), RS1000633569 (1:201866244 GC>G,GCC), RS1000645972 (1:201830184 C>G), RS1000653600 (1:201853539 G>A)

Disease associations

OMIM: gene MIM:620893 | disease phenotypes:

GenCC curated gene-disease

Mondo (0):

Orphanet (0):

HPO phenotypes

0 total (0 of 0 shown, HPO-id order):

GWAS associations

4 associations (top):

Study	Trait	p-value
GCST007559_12	Sleep duration (short sleep)	2.000000e-09
GCST010659_6	Waist circumference	2.000000e-16
GCST010988_266	Adult body size	3.000000e-25
GCST011365_20	Myocardial infarction	4.000000e-08

Drugs & pharmacology

Drug and pharmacology data

Is drug target: yes

ChEMBL targets (1): CHEMBL6067150 (SINGLE PROTEIN)

PharmGKB: 1 entry (VIP=true, CPIC=false)

ChEMBL bioactivities

2 potent at pChembl≥5 of 4 total, top 2 by pChembl (potency: 10 = 0.1 nM, 6 = 1 µM).

pChembl	Type	Value	Unit	Molecule
8.43	Kd	3.754	nM	CHEMBL5653589
8.43	ED50	3.754	nM	CHEMBL5653589

PubChem BioAssay actives

1 with measured affinity, of 4 total; 1 most potent distinct compounds. Largely complementary to BindingDB; screening values are coarse (µM, 4 dp), so sub-nM hits tie at the floor.

Compound	Assay	Type	Value	Unit
4-methyl-3-[(2-methyl-6-pyridin-3-ylpyrazolo[3,4-d]pyrimidin-4-yl)amino]-N-[3-(trifluoromethyl)phenyl]benzamide	2148599: Binding affinity to human IPO9 incubated for 45 mins by Kinobead based pull down assay	kd	0.0038	uM

CTD chemical–gene interactions

47 total (human), top 30 by PubMed support.

Chemical	Actions (top 5)	PubMed papers
bisphenol A	affects cotreatment, decreases methylation, decreases expression, increases expression	3
trichostatin A	affects cotreatment, decreases expression	2
cobaltous chloride	decreases expression	2
4-(5-benzo(1,3)dioxol-5-yl-4-pyridin-2-yl-1H-imidazol-2-yl)benzamide	affects cotreatment, increases expression, decreases expression	2
Resveratrol	affects cotreatment, increases expression	2
Air Pollutants	affects expression, increases abundance, increases expression	2
Benzo(a)pyrene	affects methylation, decreases expression, increases methylation	2
Rotenone	decreases expression	2
dicrotophos	increases expression	1
methylmercuric chloride	decreases expression	1
triphenyl phosphate	affects expression	1
tetrahydropalmatine	decreases expression	1
sodium arsenite	decreases expression	1
butyraldehyde	decreases expression	1
CGP 52608	affects binding, increases reaction	1
ICG 001	decreases expression	1
dorsomorphin	affects cotreatment, decreases expression	1
bisphenol S	increases expression	1
LDN 193189	affects cotreatment, increases expression	1
4-(4-((5-(4,5-dimethyl-2-nitrophenyl)-2-furanyl)methylene)-4,5-dihydro-3-methyl-5-oxo-1H-pyrazol-1-yl)benzoic acid	decreases expression	1
Fulvestrant	affects cotreatment, decreases methylation	1
Acetaminophen	increases expression	1
Atrazine	decreases expression	1
Calcitriol	decreases expression	1
Carbamazepine	affects expression	1
Carcinogens	decreases expression	1
Coumestrol	affects cotreatment, increases expression	1
Dimethyl Sulfoxide	increases expression	1
Dinitrochlorobenzene	affects binding	1
Doxorubicin	decreases expression	1

ChEMBL screening assays

1 unique, capped per target: 1 binding

Representative assays (with source publication via chembl_document):

Assay ID	Type	Description	Source paper
CHEMBL5651641	Binding	Binding affinity to human IPO9 incubated for 45 mins by Kinobead based pull down assay	NVP-BHG712: Effects of Regioisomers on the Affinity and Selectivity toward the EPHrin Family. — ChemMedChem

Cellosaurus cell lines

1 cell lines: 1 cancer cell line

First 10 cell lines (id-ordered, not curated):

Cellosaurus	Name	Category	Sex
CVCL_B1UM	Abcam HeLa IPO9 KO	Cancer cell line	Female

Clinical trials (associated diseases)

0 trials via MONDO — disease-level, not drug-specific.