TSSK3

gene
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Also known as SPOGA3

Summary

TSSK3 (testis specific serine kinase 3, HGNC:15473) is a protein-coding gene on chromosome 1p35.1, encoding Testis-specific serine/threonine-protein kinase 3 (Q96PN8). Serine/threonine protein kinase required for spermatid development and male fertility.

This gene encodes a kinase expressed exclusively in the testis that is thought to play a role in either germ cell differentiation or mature sperm function.

Source: NCBI Gene 81629 — RefSeq curated summary.

At a glance

  • GWAS associations: 1
  • Clinical variants (ClinVar): 49 total
  • Druggable target: yes — 1 molecules with ChEMBL bioactivity
  • MANE Select transcript: NM_052841

Identifiers

Gene identifiers

FieldValue
HGNC IDHGNC:15473
Approved symbolTSSK3
Nametestis specific serine kinase 3
Location1p35.1
Locus typegene with protein product
StatusApproved
AliasesSPOGA3
Ensembl geneENSG00000162526
Ensembl biotypeprotein_coding
OMIM607660
Entrez81629

Gene structure

Transcript identifiers

Ensembl transcripts: 4 — 2 protein_coding, 2 protein_coding_CDS_not_defined

ENST00000373534, ENST00000421153, ENST00000574315, ENST00000574792

RefSeq mRNA: 1 — MANE Select: NM_052841 NM_052841

CCDS: CCDS362

Canonical transcript exons

ENST00000373534 — 2 exons

ExonStartEnd
ENSE000014607833236359532364312
ENSE000014607853236253932362846

Expression profiles

Bgee: expression breadth ubiquitous, 159 present calls, max score 94.69.

FANTOM5 (CAGE): breadth ubiquitous, TPM avg 2.6814 / max 113.6734, expressed in 1112 samples.

FANTOM5 promoters (4 alternative TSS)

Promoter IDTPM avgSamples expressed
19841.3580764
19851.2438671
19870.06626
19860.01343

Top tissues by expression

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

TissueAnatomy IDExpression scoreQuality
spermCL:000001994.69gold quality
right testisUBERON:000453492.39gold quality
left testisUBERON:000453392.15gold quality
testisUBERON:000047389.23gold quality
male germ line stem cell (sensu Vertebrata) in testisCL:0000089 ∩ UBERON:000047387.67gold quality
right ovaryUBERON:000211879.68gold quality
left ovaryUBERON:000211979.16gold quality
sural nerveUBERON:001548878.87gold quality
stromal cell of endometriumCL:000225578.52gold quality
right uterine tubeUBERON:000130278.50gold quality
vena cavaUBERON:000408778.36gold quality
body of uterusUBERON:000985378.27gold quality
cerebellar hemisphereUBERON:000224578.15gold quality
cerebellar cortexUBERON:000212978.08gold quality
right hemisphere of cerebellumUBERON:001489077.95gold quality
mucosa of stomachUBERON:000119977.83gold quality
subthalamic nucleusUBERON:000190677.58gold quality
left uterine tubeUBERON:000130377.50gold quality
endocervixUBERON:000045877.43gold quality
cerebellumUBERON:000203777.34gold quality
right lobe of thyroid glandUBERON:000111977.32gold quality
apex of heartUBERON:000209876.60gold quality
right lungUBERON:000216776.54gold quality
granulocyteCL:000009476.47gold quality
left lobe of thyroid glandUBERON:000112076.44gold quality
epithelium of nasopharynxUBERON:000195176.07gold quality
thyroid glandUBERON:000204675.99gold quality
left adrenal gland cortexUBERON:003582575.75gold quality
dorsal plus ventral thalamusUBERON:000189775.74gold quality
gastrocnemiusUBERON:000138875.65gold quality

Single-cell (SCXA)

Detected in 1 experiment(s), a significant marker in 0.

ExperimentMarker?Max mean expression
E-ANND-3no1.07

Regulation

Is transcription factor: no

Literature-anchored findings (GeneRIF, showing 3)

  • human homolog of mouse stk22d; It was cloned and found it to be expressed exclusively in the testis. (PMID:11597141)
  • PDK1-induced phosphorylation increased in vitro TSSK3 kinase activity, suggesting that TSSK3 can be regulated in the same way as AGC kinase family members. The peptide sequence RRSSSY containing Ser5 is identified as a target for TSSK3 phosphorylation (PMID:16336268)
  • TSSK3, a novel target for male contraception, is required for spermiogenesis. (PMID:34623009)

Cross-species orthologs

9 orthologs

OrganismSymbolGene ID
danio_rerionim1kbENSDARG00000079031
mus_musculusTssk3ENSMUSG00000000411
rattus_norvegicusTssk3ENSRNOG00000008693
drosophila_melanogasterSnrkFBGN0033915
drosophila_melanogasterNuakFBGN0262617
caenorhabditis_elegansWBGENE00012638
caenorhabditis_elegansZK524.4WBGENE00013994
caenorhabditis_eleganstag-344WBGENE00015230
caenorhabditis_elegansWBGENE00044388

Paralogs (17): NUAK1 (ENSG00000074590), PRKAA1 (ENSG00000132356), TSSK4 (ENSG00000139908), HUNK (ENSG00000142149), SIK1 (ENSG00000142178), BRSK1 (ENSG00000160469), SIK3 (ENSG00000160584), PRKAA2 (ENSG00000162409), NUAK2 (ENSG00000163545), SNRK (ENSG00000163788), MELK (ENSG00000165304), SIK2 (ENSG00000170145), BRSK2 (ENSG00000174672), NIM1K (ENSG00000177453), TSSK6 (ENSG00000178093), TSSK2 (ENSG00000206203), TSSK1B (ENSG00000212122)

Protein

Protein identifiers

Testis-specific serine/threonine-protein kinase 3Q96PN8 (reviewed: Q96PN8)

Alternative names: Serine/threonine-protein kinase 22C

All UniProt accessions (2): Q96PN8, I3L1X4

UniProt curated annotations — full annotation on UniProt →

Function. Serine/threonine protein kinase required for spermatid development and male fertility.

Subcellular location. Cell projection. Cilium. Flagellum.

Post-translational modifications. Autophosphorylated at Ser-166. Phosphorylation at Thr-168 by PDPK1 activates the serine/threonine protein kinase activity.

Activity regulation. Activated by phosphorylation on Thr-168 by PDPK1.

Cofactor. Can use both Mg(2+) and Mn(2+) as cofactor, with a preference for Mn(2+) in vitro.

Similarity. Belongs to the protein kinase superfamily. CAMK Ser/Thr protein kinase family.

RefSeq proteins (1): NP_443073* (*=MANE)

Domains & families (InterPro)

IDNameType
IPR000719Prot_kinase_domDomain
IPR011009Kinase-like_dom_sfHomologous_superfamily
IPR017441Protein_kinase_ATP_BSBinding_site
IPR042709TSSK3_STKcDomain

Pfam: PF00069

Catalyzed reactions (Rhea), 2 shown:

  • L-seryl-[protein] + ATP = O-phospho-L-seryl-[protein] + ADP + H(+) (RHEA:17989)
  • L-threonyl-[protein] + ATP = O-phospho-L-threonyl-[protein] + ADP + H(+) (RHEA:46608)

UniProt features (10 total): sequence variant 3, binding site 2, modified residue 2, chain 1, domain 1, active site 1

Structure

Experimental structures (PDB)

0 structures.

Predicted structure (AlphaFold)

ModelpLDDTFraction very-high
AF-Q96PN8-F193.650.83

Functional residue map

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

Catalytic / active sites (1): 134 (proton acceptor)

Ligand- & substrate-binding residues (2): 16–24; 39

Post-translational modifications (2): 166, 168

Function

Pathways and Gene Ontology

Reactome pathways

0 pathways

MSigDB gene sets: 106 (showing top): AAGCAAT_MIR137, MAZ_Q6, TGACCTY_ERR1_Q2, GOBP_MALE_GAMETE_GENERATION, AAAYRNCTG_UNKNOWN, GATA1_01, GRE_C, GOBP_CELLULAR_PROCESS_INVOLVED_IN_REPRODUCTION_IN_MULTICELLULAR_ORGANISM, HIF1_Q3, GOBP_DEVELOPMENTAL_PROCESS_INVOLVED_IN_REPRODUCTION, CDPCR3HD_01, YY1_01, DBP_Q6, AP2_Q6_01, AP2GAMMA_01

GO Biological Process (4): protein phosphorylation (GO:0006468), spermatogenesis (GO:0007283), spermatid development (GO:0007286), cell differentiation (GO:0030154)

GO Molecular Function (11): magnesium ion binding (GO:0000287), protein serine/threonine kinase activity (GO:0004674), ATP binding (GO:0005524), manganese ion binding (GO:0030145), protein serine kinase activity (GO:0106310), nucleotide binding (GO:0000166), protein kinase activity (GO:0004672), protein binding (GO:0005515), kinase activity (GO:0016301), transferase activity (GO:0016740), metal ion binding (GO:0046872)

GO Cellular Component (4): sperm flagellum (GO:0036126), cilium (GO:0005929), motile cilium (GO:0031514), cell projection (GO:0042995)

GO top-level categories

Rollup of top GO terms by namespace:

CategoryTerms
protein kinase activity2
phosphorylation1
protein modification process1
developmental process involved in reproduction1
male gamete generation1
germ cell development1
spermatid differentiation1
cellular developmental process1
metal ion binding1
adenyl ribonucleotide binding1
purine ribonucleoside triphosphate binding1
transition metal ion binding1
nucleoside phosphate binding1
heterocyclic compound binding1
kinase activity1
phosphotransferase activity, alcohol group as acceptor1
catalytic activity, acting on a protein1
binding1
transferase activity, transferring phosphorus-containing groups1
catalytic activity1
cation binding1
9+2 motile cilium1
intraciliary transport particle1
membrane-bounded organelle1
plasma membrane bounded cell projection1
cilium1
cellular anatomical structure1

Protein interactions and networks

STRING

1581 interactions, top by confidence (×1000):

Protein AProtein BPartner UniProtScore
TSSK3TSKSQ9UJT2720
TSSK3CALYQ9NYX4594
TSSK3KLHL10Q6JEL2514
TSSK3CRABP1P29762454
TSSK3SIGLEC1Q9BZZ2452
TSSK3SSMEM1Q8WWF3417
TSSK3TNP1P09430401
TSSK3IZUMO1Q8IYV9397
TSSK3FAM170BA6NMN3392
TSSK3TXNDC2Q86VQ3383
TSSK3SPATC1Q76KD6382
TSSK3CCDC70Q6NSX1380
TSSK3PRM2P04554364
TSSK3SOX30O94993354
TSSK3TCP11Q8WWU5351

IntAct

216 interactions, top by confidence:

ABTypeScore
TSSK3LNX1psi-mi:“MI:0915”(physical association)0.780
LNX1TSSK3psi-mi:“MI:0915”(physical association)0.780
GOLGA2TSSK3psi-mi:“MI:0915”(physical association)0.720
EFHC2TSSK3psi-mi:“MI:0915”(physical association)0.720
TSSK3VPS52psi-mi:“MI:0915”(physical association)0.720
TSSK3RBCK1psi-mi:“MI:0915”(physical association)0.720
IKZF3TSSK3psi-mi:“MI:0915”(physical association)0.720
TSSK3GOLGA2psi-mi:“MI:0915”(physical association)0.720
TSSK3EFHC2psi-mi:“MI:0915”(physical association)0.720
VPS52TSSK3psi-mi:“MI:0915”(physical association)0.720
TSSK3IKZF3psi-mi:“MI:0915”(physical association)0.720
DDIT4LTSSK3psi-mi:“MI:0915”(physical association)0.600
TCF4TSSK3psi-mi:“MI:0915”(physical association)0.560
TSSK3ZBTB25psi-mi:“MI:0915”(physical association)0.560
TSSK3RELpsi-mi:“MI:0915”(physical association)0.560
TLE5TSSK3psi-mi:“MI:0915”(physical association)0.560

BioGRID (88): TSSK3 (Two-hybrid), TSSK3 (Two-hybrid), TSSK3 (Two-hybrid), TSSK3 (Two-hybrid), TSSK3 (Two-hybrid), TSSK3 (Two-hybrid), TSSK3 (Two-hybrid), TSSK3 (Two-hybrid), TSSK3 (Two-hybrid), TSSK3 (Two-hybrid), LNX1 (Two-hybrid), TRIM9 (Two-hybrid), TSSK3 (Two-hybrid), TSSK3 (Two-hybrid), TSSK3 (Biochemical Activity)

ESM2 similar proteins: A0A5B9GBF0, A1IVT7, A3B529, B2MVY4, O65554, O70150, O70444, P05128, P11802, P23111, P27638, P29618, P30285, P34112, P35426, P48609, P52389, P58750, P79432, P93101, Q05006, Q0U4L8, Q15759, Q1DUU8, Q2PQN9, Q2U469, Q32KY4, Q38772, Q38HL5, Q41639, Q5JLD8, Q683C9, Q69Q47, Q6ERS4, Q6P2M8, Q6SA08, Q80YP0, Q86V86, Q8C1R0, Q8TGA9

Diamond homologs: A2XFF4, A3B529, A8WYE4, B8BBT7, O08678, O08679, O22932, O22971, O54863, O60285, O65554, O94168, P06782, P27448, P54645, P57059, P92937, P92958, Q00372, Q02723, Q03141, Q05512, Q0D4B2, Q0JI49, Q10LQ2, Q12263, Q13131, Q21017, Q28GW8, Q2QY53, Q2QYM3, Q2RAX3, Q2RBF0, Q38997, Q3SZW1, Q54DF2, Q54TA3, Q54YF2, Q5EG47, Q5JLD8

SIGNOR signaling

2 interactions.

AEffectBMechanism
TSSK3“up-regulates activity”TSSK3phosphorylation
PDPK1“up-regulates activity”TSSK3phosphorylation

Disease & clinical

Clinical variants and AI predictions

ClinVar

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

ClassificationCount (floor)
Pathogenic0
Likely pathogenic0
Uncertain significance46
Likely benign0
Benign0

Top pathogenic / likely-pathogenic (0)

SpliceAI

435 predictions. Top by Δscore:

VariantEffectΔscore
1:32362417:GGC:Gdonor_gain1.0000
1:32361725:CTTA:Cdonor_loss0.9900
1:32361726:TTACC:Tdonor_loss0.9900
1:32361727:TACC:Tdonor_loss0.9900
1:32361728:A:Tdonor_loss0.9900
1:32361729:C:CAdonor_loss0.9900
1:32362418:GC:Gdonor_gain0.9900
1:32362419:C:Gdonor_gain0.9900
1:32363585:A:AGacceptor_gain0.9900
1:32363591:TCAG:Tacceptor_gain0.9900
1:32363593:A:ACacceptor_loss0.9900
1:32363593:A:AGacceptor_gain0.9900
1:32363594:G:GGacceptor_gain0.9900
1:32363594:G:Tacceptor_loss0.9900
1:32363594:GA:Gacceptor_gain0.9900
1:32363594:GAGT:Gacceptor_gain0.9900
1:32361729:CCTGA:Cdonor_gain0.9800
1:32361731:TG:Tdonor_gain0.9800
1:32362416:GGGC:Gdonor_gain0.9800
1:32362430:G:GAdonor_gain0.9800
1:32362433:G:GAdonor_gain0.9800
1:32362435:T:TAdonor_gain0.9800
1:32362436:A:AAdonor_gain0.9800
1:32363591:TCAGA:Tacceptor_gain0.9800
1:32363592:CAG:Cacceptor_gain0.9800
1:32363592:CAGAG:Cacceptor_gain0.9800
1:32363593:AGA:Aacceptor_gain0.9800
1:32363594:GAG:Gacceptor_gain0.9800
1:32363594:GAGTT:Gacceptor_gain0.9800
1:32361728:A:ACdonor_gain0.9700

AlphaMissense

1772 scored. Top likely-pathogenic:

VariantProtein changeam_pathogenicity
1:32362818:A:CK39N0.999
1:32362818:A:TK39N0.999
1:32363847:G:CR133P0.998
1:32363850:A:CD134A0.998
1:32363850:A:TD134V0.998
1:32363901:A:TD151V0.998
1:32363902:C:AD151E0.998
1:32363902:C:GD151E0.998
1:32364077:T:CF210L0.998
1:32364079:T:AF210L0.998
1:32364079:T:GF210L0.998
1:32363850:A:GD134G0.997
1:32363851:C:AD134E0.997
1:32363851:C:GD134E0.997
1:32363857:A:CK136N0.997
1:32363857:A:TK136N0.997
1:32364029:T:AW194R0.997
1:32364029:T:CW194R0.997
1:32363609:T:CF54L0.996
1:32363611:C:AF54L0.996
1:32363611:C:GF54L0.996
1:32363855:A:GK136E0.996
1:32363856:A:TK136I0.996
1:32363901:A:CD151A0.996
1:32363963:A:CS172R0.996
1:32363965:T:AS172R0.996
1:32363965:T:GS172R0.996
1:32362811:C:AA37E0.995
1:32362816:A:GK39E0.995
1:32363913:C:AA155D0.995

dbSNP variants (sampled 300 via entrez): RS1000068694 (1:32362322 A>G), RS1000652777 (1:32361200 G>A), RS1000982482 (1:32360986 C>T), RS1001722336 (1:32360696 G>A), RS1002459355 (1:32361849 G>A), RS1002472014 (1:32360794 A>G), RS1003416754 (1:32364449 CCTT>C), RS1003728868 (1:32363037 C>A,G,T), RS1004007596 (1:32364338 T>C), RS1004249861 (1:32364560 C>T), RS1004800162 (1:32361860 C>T), RS1004872163 (1:32362080 G>A,C), RS1005879120 (1:32363011 G>A,T), RS1006441808 (1:32361803 A>C), RS1006569479 (1:32362698 C>A,T)

Disease associations

OMIM: gene MIM:607660 | disease phenotypes:

GenCC curated gene-disease

Mondo (0):

Orphanet (0):

HPO phenotypes

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

GWAS associations

1 associations (top):

StudyTraitp-value
GCST008758_22Pre-treatment viral load in HIV-1 infection4.000000e-16

EFO canonical traits (1, from GWAS)

EFO IDTrait name
EFO:0010125viral load

Drugs & pharmacology

Drug and pharmacology data

Is drug target: yes

ChEMBL targets (1): CHEMBL3414416 (SINGLE PROTEIN)

Molecules with ChEMBL bioactivity

1 molecules (phase ≥1), by development phase (incl. off-target/promiscuous compounds). Patent mentions across the top 20 by phase: 1,538 (via chembl_molecule»patent_compound — counts attach to the compound, not the gene–compound relationship, so off-target/promiscuous molecules can dominate).

MoleculeNamePhasePatents
CHEMBL1232461MOLIBRESIB21,538

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

GtoPdb / IUPHAR curated pharmacology

(IUPHAR/BPS Guide to Pharmacology — expert-curated)

Target class: enzyme — Testis specific kinase (TSSK) family

ChEMBL bioactivities

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

pChemblTypeValueUnitMolecule
8.49IC503.21nMSTAUROSPORINE
8.15IC507.01nMSTAUROSPORINE
7.83IC5014.9nMSTAUROSPORINE
7.04IC5091nMCHEMBL2334586
6.48Kd330nMCHEMBL4576489
6.42Kd380nMCHEMBL4465866
6.38IC50420nMCHEMBL3415608
6.38IC50420nMCHEMBL3943181
6.00IC501000nMTP-030n
5.00IC501e+04nMMOLIBRESIB

PubChem BioAssay actives

9 with measured affinity, of 67 total; 7 most potent distinct compounds. Largely complementary to BindingDB; screening values are coarse (µM, 4 dp), so sub-nM hits tie at the floor.

CompoundAssayTypeValueUnit
(2S,3R,4R,6R)-3-methoxy-2-methyl-4-(methylamino)-29-oxa-1,7,17-triazaoctacyclo[12.12.2.12,6.07,28.08,13.015,19.020,27.021,26]nonacosa-8,10,12,14,19,21,23,25,27-nonaen-16-one1715136: Inhibition of human TSSK3 using KKKVSRSGLYRSPSMPENLNRPR as substrate by [gamma-33P]-ATP assayic500.0032uM
4-(4-amino-3,5,12-triazatetracyclo[9.7.0.02,7.013,18]octadeca-1(11),2,4,6,13(18),14,16-heptaen-16-yl)-2-methylbut-3-yn-2-ol1856574: Inhibition of wild-type human full length TSSK3 (M1 to T268 residues) expressed in mammalian expression system by Kinomescan methodic500.0910uM
3-(2,2-difluoro-10,12-dimethyl-1-aza-3-azonia-2-boranuidatricyclo[7.3.0.03,7]dodeca-3,5,7,9,11-pentaen-4-yl)-N-[2-[2-[2-[2-[[(2S,3R,4R,6R)-3-methoxy-2-methyl-16-oxo-29-oxa-1,7,17-triazaoctacyclo[12.12.2.12,6.07,28.08,13.015,19.020,27.021,26]nonacosa-8,10,12,14,19,21,23,25,27-nonaen-4-yl]-methylamino]ethoxy]ethoxy]ethoxy]ethyl]propanamide1526261: Binding affinity to recombinant full-length N-terminal His-FLAG-GST-tagged TSSK3 (unknown origin) expressed in baculovirus infected Sf9 insect cells incubated for 1 hr by TR-FRET assaykd0.3300uM
3-(2,2-difluoro-10,12-dimethyl-1-aza-3-azonia-2-boranuidatricyclo[7.3.0.03,7]dodeca-3,5,7,9,11-pentaen-4-yl)-N-[2-[2-[2-[2-[[(2S,3R,4R,6R)-3-methoxy-2-methyl-16-oxo-29-oxa-1,7,17-triazaoctacyclo[12.12.2.12,6.07,28.08,13.015,19.020,27.021,26]nonacosa-8,10,12,14,19,21,23,25,27-nonaen-4-yl]amino]ethoxy]ethoxy]ethoxy]ethyl]propanamide1526261: Binding affinity to recombinant full-length N-terminal His-FLAG-GST-tagged TSSK3 (unknown origin) expressed in baculovirus infected Sf9 insect cells incubated for 1 hr by TR-FRET assaykd0.3800uM
5-[[(1R,2S)-2-aminocyclohexyl]amino]-3-(1H-benzimidazol-2-yl)pyridine-2-carboxamide1199576: Inhibition of TSSK3 (unknown origin)ic500.4200uM
5-[[(1R,2S)-2-aminocyclohexyl]amino]-3-(1H-benzimidazol-2-yl)pyrazine-2-carboxamide1326206: Inhibition of TSSK3 (unknown origin)ic500.4200uM
2-[(4S)-6-(4-chlorophenyl)-8-methoxy-1-methyl-4H-[1,2,4]triazolo[4,3-a][1,4]benzodiazepin-4-yl]-N-ethylacetamide2178641: Inhibition of TSSK3 (unknown origin) incubated for 1 hr by colloidal coomassie staining based LC-MS/MS analysisic5010.0000uM

CTD chemical–gene interactions

11 total (human), top 11 by PubMed support.

ChemicalActions (top 5)PubMed papers
aminomethylphosphonic acid (AMPA)decreases expression1
tris(1,3-dichloro-2-propyl)phosphateincreases expression1
jinfukangincreases expression, affects cotreatment1
Air Pollutantsincreases expression, increases abundance1
Cisplatinaffects cotreatment, increases expression1
Estradioldecreases expression1
Folic Acidincreases expression1
Valproic Acidincreases methylation1
2,4-Dichlorophenoxyacetic Aciddecreases expression1
Okadaic Aciddecreases expression1
Particulate Matterincreases abundance, increases expression1

ChEMBL screening assays

83 unique, capped per target: 83 binding

Representative assays (with source publication via chembl_document):

Assay IDTypeDescriptionSource paper
CHEMBL3418524BindingInhibition of TSSK3 (unknown origin)Overcoming mutagenicity and ion channel activity: optimization of selective spleen tyrosine kinase inhibitors. — J Med Chem

Cellosaurus cell lines

2 cell lines: 2 cancer cell line

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

CellosaurusNameCategorySex
CVCL_B2JVAbcam HeLa TSSK3 KOCancer cell lineFemale
CVCL_TV00HAP1 TSSK3 (-)Cancer cell lineMale

Clinical trials (associated diseases)

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

No linked Atlas pages yet — the cross-entity mesh grows as the corpus expands.