GLUL

Summary

GLUL (glutamate-ammonia ligase, HGNC:4341) is a protein-coding gene on chromosome 1q25.3, encoding Glutamine synthetase (P15104). Glutamine synthetase that catalyzes the ATP-dependent conversion of glutamate and ammonia to glutamine.

The protein encoded by this gene belongs to the glutamine synthetase family. It catalyzes the synthesis of glutamine from glutamate and ammonia in an ATP-dependent reaction. This protein plays a role in ammonia and glutamate detoxification, acid-base homeostasis, cell signaling, and cell proliferation. Glutamine is an abundant amino acid, and is important to the biosynthesis of several amino acids, pyrimidines, and purines. Mutations in this gene are associated with congenital glutamine deficiency, and overexpression of this gene was observed in some primary liver cancer samples. There are six pseudogenes of this gene found on chromosomes 2, 5, 9, 11, and 12. Alternative splicing results in multiple transcript variants.

Source: NCBI Gene 2752 — RefSeq curated summary.

At a glance

• Gene–disease (curated): congenital brain dysgenesis due to glutamine synthetase deficiency (Definitive, GenCC) — +2 more curated relationships
• GWAS associations: 50
• Clinical variants (ClinVar): 274 total — 5 pathogenic, 5 likely-pathogenic
• Phenotypes (HPO): 48
• Druggable target: yes — 1 molecules with ChEMBL bioactivity
• Dosage sensitivity (ClinGen): haploinsufficiency autosomal recessive, triplosensitivity no evidence
• MANE Select transcript: NM_001033044

Identifiers

Gene identifiers

Field	Value
HGNC ID	HGNC:4341
Approved symbol	GLUL
Name	glutamate-ammonia ligase
Location	1q25.3
Locus type	gene with protein product
Status	Approved
Ensembl gene	ENSG00000135821
Ensembl biotype	protein_coding
OMIM	138290
Entrez	2752

Gene structure

Transcript identifiers

Ensembl transcripts: 11 — 5 protein_coding, 5 retained_intron, 1 protein_coding_CDS_not_defined

ENST00000311223, ENST00000331872, ENST00000339526, ENST00000417584, ENST00000461447, ENST00000462444, ENST00000463851, ENST00000475808, ENST00000489818, ENST00000491322, ENST00000642379

RefSeq mRNA: 3 — MANE Select: NM_001033044 NM_001033044, NM_001033056, NM_002065

CCDS: CCDS1344

Canonical transcript exons

ENST00000331872 — 7 exons

Exon	Start	End
ENSE00000000279	182378098	182384723
ENSE00003520164	182387131	182387292
ENSE00003541539	182386256	182386402
ENSE00003616388	182388572	182388750
ENSE00003650736	182385760	182385887
ENSE00003686762	182385357	182385556
ENSE00003902209	182391679	182391790

Expression profiles

Bgee: expression breadth ubiquitous, 309 present calls, max score 99.98.

FANTOM5 (CAGE): breadth ubiquitous, TPM avg 104.3890 / max 2825.2504, expressed in 1808 samples.

FANTOM5 promoters (10 alternative TSS)

Promoter ID	TPM avg	Samples expressed
16149	93.8107	1799
16147	5.7904	1351
16148	2.7983	782
16150	0.6022	303
16140	0.6010	323
16146	0.3362	186
16144	0.2387	9
16151	0.1006	34
16145	0.0718	5
16143	0.0392	23

Top tissues by expression

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

Tissue	Anatomy ID	Expression score	Quality
endothelial cell	CL:0000115	99.98	gold quality
medial globus pallidus	UBERON:0002477	99.95	gold quality
globus pallidus	UBERON:0001875	99.93	gold quality
palpebral conjunctiva	UBERON:0001812	99.92	gold quality
amniotic fluid	UBERON:0000173	99.91	gold quality
buccal mucosa cell	CL:0002336	99.90	gold quality
mucosa of paranasal sinus	UBERON:0005030	99.90	gold quality
eye	UBERON:0000970	99.89	gold quality
epithelium of nasopharynx	UBERON:0001951	99.89	gold quality
nasopharynx	UBERON:0001728	99.88	gold quality
choroid plexus epithelium	UBERON:0003911	99.87	gold quality
ventral tegmental area	UBERON:0002691	99.85	gold quality
occipital lobe	UBERON:0002021	99.84	gold quality
germinal epithelium of ovary	UBERON:0001304	99.83	gold quality
trigeminal ganglion	UBERON:0001675	99.83	gold quality
primary visual cortex	UBERON:0002436	99.83	gold quality
lateral globus pallidus	UBERON:0002476	99.83	gold quality
dorsal motor nucleus of vagus nerve	UBERON:0002870	99.83	gold quality
inferior vagus X ganglion	UBERON:0005363	99.82	gold quality
Brodmann (1909) area 23	UBERON:0013554	99.82	gold quality
nucleus accumbens	UBERON:0001882	99.81	gold quality
substantia nigra pars reticulata	UBERON:0001966	99.81	gold quality
right frontal lobe	UBERON:0002810	99.81	gold quality
left testis	UBERON:0004533	99.81	gold quality
right testis	UBERON:0004534	99.81	gold quality
parietal pleura	UBERON:0002400	99.80	gold quality
putamen	UBERON:0001874	99.79	gold quality
medulla oblongata	UBERON:0001896	99.79	gold quality
superior vestibular nucleus	UBERON:0007227	99.79	gold quality
cardia of stomach	UBERON:0001162	99.78	gold quality

Single-cell (SCXA)

Detected in 35 experiment(s), a significant marker in 33.

Experiment	Marker?	Max mean expression
E-GEOD-137537	yes	10515.84
E-MTAB-7316	yes	7899.61
E-ANND-2	yes	3177.06
E-MTAB-6308	yes	3025.91
E-MTAB-10137	yes	2122.17
E-MTAB-7407	yes	2059.69
E-MTAB-8530	yes	1974.78
E-HCAD-1	yes	97.28
E-CURD-122	yes	80.40
E-MTAB-6701	yes	72.71
E-MTAB-8410	yes	57.60
E-HCAD-10	yes	54.99
E-CURD-88	yes	46.17
E-GEOD-135922	yes	45.71
E-MTAB-8142	yes	44.73

Regulation

Is transcription factor: no

Upstream regulators (CollecTRI, top): CTNNB1, FOXO1, HNF4A, MYB, NR3C1, RBPJ, REST, STAT5A, STAT5B

miRNA regulators (miRDB)

219 targeting GLUL, 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-4481	100.00	66.42	1669
HSA-MIR-3646	100.00	73.56	5283
HSA-MIR-1252-5P	100.00	69.80	2774
HSA-MIR-3163	100.00	77.23	8605
HSA-MIR-12118	100.00	65.88	1270
HSA-MIR-4283	100.00	66.42	2097
HSA-MIR-1193	100.00	65.93	529
HSA-MIR-548C-3P	99.99	74.01	7587
HSA-MIR-3662	99.99	73.82	5684
HSA-MIR-371B-5P	99.99	75.34	4759
HSA-MIR-196A-1-3P	99.99	72.15	2772
HSA-MIR-4534	99.99	66.58	1907
HSA-MIR-6077	99.99	68.04	2299
HSA-MIR-4745-5P	99.98	65.95	1028
HSA-MIR-4775	99.98	75.00	6394
HSA-MIR-373-5P	99.98	75.36	4753
HSA-MIR-616-5P	99.98	75.58	4775
HSA-MIR-8068	99.98	73.85	2376
HSA-MIR-6891-5P	99.98	66.53	1372
HSA-MIR-3173-3P	99.98	66.49	1217
HSA-MIR-512-3P	99.97	67.35	1049
HSA-MIR-3148	99.97	75.06	6478
HSA-MIR-3658	99.96	73.87	4379
HSA-MIR-548AA	99.96	70.64	3753
HSA-MIR-548AP-3P	99.96	70.64	3753
HSA-MIR-548T-3P	99.96	70.64	3753
HSA-MIR-590-3P	99.96	74.34	6478
HSA-MIR-302E	99.96	70.74	2669
HSA-MIR-548AJ-3P	99.96	73.38	5345
HSA-MIR-548X-3P	99.96	73.38	5345

Functional genomics

ClinGen dosage: haploinsufficiency 30 (autosomal recessive), triplosensitivity 0 (no evidence). ClinGen Gene Dosage Map

Literature-anchored findings (GeneRIF, showing 40)

• 2D fingerprinting & mass spectrometry reveal specific targets of protein oxidation in Alzheimer’s disease brain, including glutamine synthase, suggesting involvement of oxidatively modified proteins in neurodegeneration. (PMID:12160938)
• The detection of a significantly reduced enzymatic activity in the epileptic amygdala supports the assumption that the enzyme defect is localized to the epileptic mesial temporal lobe of corresponding patients. (PMID:16095760)
• Liver tissue samples from HCV-HCC cancerous tissues and corresponding non-cancerous tissues from patients showed three protein spots of the same molecular mass (42 kDa), whose expression increased in well-differentiated cancerous tissues. (PMID:16609938)
• In myocytes and hepatomas, but not in adipocytes, glutamine acts to moderate glutamine synthetase induction by glucocorticoids. (PMID:17197094)
• We confirmed the induction of GS expression by dexamethasone published previously. Components of the glutamatergic system may play a role in bone pathophysiology (PMID:17627080)
• Crystal structure of GLUL illustrates substrate-induced conformational change and provides opportunities for drug and herbicide design. (PMID:18005987)
• in solid pseudopapillary neoplasm of the pancreas glutamine synthetase expression is highly correlated with Wnt/beta-catenin activation, demonstrating its faithfulness as a Wnt target gene (PMID:18192886)
• glutamine synthetase has a role in control of glutamate signalling through Wnt3A and steroid pathways in osteoblastic cells (PMID:18555765)
• Glutamine synthetase can be found during early human fetal stages when it displays a significant effect on cell proliferation (PMID:18662667)
• These results suggest that GAD2 and GLUL do not play a major role in schizophrenia pathogenesis and there is no gene-gene interaction between the eight genes in the Japanese population. (PMID:19125103)
• This study suggests that seizures in GBM are coupled with a highly localized glutamine synthetase deficiency. (PMID:19183851)
• GS localization was investigated using immunohistochemistry and double-labeling of young and adult human and rat skin sections as well as skin cells in culture. (PMID:19204801)
• A reliable marker for the detection of hepatocellular carcinoma in liver biopsies. (PMID:19231003)
• GS mRNA and protein expression was not different between normal and fetal growth restriction pregnancy (PMID:19500843)
• Standard histology and glutamine synthase status identify a hepatocellular carcinoma subset with distinct clinical and pathologic features. (PMID:20233882)
• This protein has been found differentially expressed in thalami from patients with schizophrenia. (PMID:20471030)
• Glutamine synthetase was up-regulated in hepatocellular carcinoma and may be a novel marker for early diagnosis. (PMID:20525558)
• The Glul can serve as novel target in diabetes mellitus genetic therapy. (PMID:20536387)
• This study suggests that serum glutamine synthetase levels have no diagnostic value for Alzheimer’s disease. (PMID:21597934)
• glutamine synthetase expression can be used to distinguish astrocytic from oligodendroglial tumors and may play a role in the pathogenesis of astrocytomas. (PMID:21682567)
• Data show that the acidification followed by neutralisation (AFBN) pre-treatment protocol for cerebrospinal fluid (CSF) significantly enhances the measurement of glutamine synthetase (GS) and myelin basic protein (MBP) in CSF. (PMID:22542401)
• The presence of GS in the post-acrosomal region of the perinuclear theca suggests that human sperm can carry out in glutamine synthesis. (PMID:22777743)
• these results suggest that GLUL contributes to pancreatic regeneration. (PMID:22930410)
• Glutamine synthetase expression is increased in regenerating hepatocytes and in early hepatocyte progenitor cells prior to morphological evidence of hepatocellular differentiation. (PMID:23362937)
• High Glutamine synthetase expression is associated with epilepsy in newly diagnosed glioblastoma multiforme. (PMID:23410662)
• Results highlight the diagnostic errors that can be caused by variant patterns of staining with glutamine synthetase and serum amyloid-associated protein in inflammatory hepatocellular adenoma and focal nodular hyperplasia. (PMID:23807780)
• through a 3-stage genome-wide association study in 4188 type 2 diabetic patients, identified a novel susceptibility locus for coronary heart disease in the region of the GLUL gene (PMID:23982368)
• findings point to SNP rs10911021 of GLULas an independent modulator of mortality in patients with type 2 diabetes (PMID:25677913)
• Data indicate that the triple stain of reticulin, glypican-3, and glutamine synthetae is useful in the differentiation of hepatocellular carcinoma, hepatic adenoma, and focal nodular hyperplasia. (PMID:25822763)
• GLUL rs10911021 is associated prospectively with adjudicated cardiovascular composite end points among overweight/obese individuals with type 2 diabetes. (PMID:26395743)
• Data show that glutamine synthetase (GS) produces glutamine (Gln) from tricarboxylic acid (TCA)-cycle-derived carbons. (PMID:26595383)
• This study demonstrated that Influence of glutamine synthetase gene polymorphisms on the development of hyperammonemia during valproic acid-based therapy. (PMID:26599579)
• Studied molecular mechanisms of glutamine synthetase mutations that lead to clinically relevant pathologies. (PMID:26836257)
• GS is acetylated at lysines 11 and 14, yielding a degron that is necessary and sufficient for binding and ubiquitylation by CRL4(CRBN) and degradation by the proteasome. (PMID:26990986)
• GLUL knockdown markedly inhibited the p38 MAPK and ERK1/ERK2 signaling pathways in cultured breast cancer cells and reduces their proliferation. (PMID:27791265)
• co-targeting glutamine synthetase in stroma and glutaminase in cancer cells reduces tumor weight, nodules, and metastasis. (PMID:27829138)
• genome-wide association study to identify genetic factors for familial hepatitis B virus-related hepatocellular carcinoma; results identified 2 large effect susceptible haplotypes located at GLUL and SLC13A2/FOXN1 (PMID:28662289)
• The data identify GS activity as mediator of the proangiogenic, immunosuppressive, and pro-metastatic function of M2-like macrophages and highlight the possibility of targeting this enzyme in the treatment of cancer metastasis. (PMID:28813676)
• Data show that 1,25-dihydroxyvitamin D (1,25D) downregulation of glutamine synthetase (GLUL; GS)) was sufficient to reduce abundance and enzyme activity of GS. (PMID:29029014)
• The SNP rs10911021 near glutamate-ammonia ligase is associated with oxidative stress in coronary heart disease patients from Pakistan. (PMID:29304826)

Cross-species orthologs

9 orthologs

Organism	Symbol	Gene ID
danio_rerio	glula	ENSDARG00000099776
danio_rerio	glulb	ENSDARG00000100003
mus_musculus	Glul	ENSMUSG00000026473
rattus_norvegicus	Glul	ENSRNOG00000049560
drosophila_melanogaster	Gs2	FBGN0001145
caenorhabditis_elegans		WBGENE00001602
caenorhabditis_elegans		WBGENE00001604
caenorhabditis_elegans		WBGENE00001606
caenorhabditis_elegans		WBGENE00001607

Paralogs (1): LGSN (ENSG00000146166)

Protein

Protein identifiers

Glutamine synthetase — P15104 (reviewed: P15104)

Alternative names: Glutamate–ammonia ligase, Palmitoyltransferase GLUL

All UniProt accessions (3): A0A2R8YDT1, A8YXX4, P15104

UniProt curated annotations — full annotation on UniProt →

Function. Glutamine synthetase that catalyzes the ATP-dependent conversion of glutamate and ammonia to glutamine. Its role depends on tissue localization: in the brain, it regulates the levels of toxic ammonia and converts neurotoxic glutamate to harmless glutamine, whereas in the liver, it is one of the enzymes responsible for the removal of ammonia. Plays a key role in ammonium detoxification during erythropoiesis: the glutamine synthetase activity is required to remove ammonium generated by porphobilinogen deaminase (HMBS) during heme biosynthesis to prevent ammonium accumulation and oxidative stress. Essential for proliferation of fetal skin fibroblasts. Independently of its glutamine synthetase activity, required for endothelial cell migration during vascular development: acts by regulating membrane localization and activation of the GTPase RHOJ, possibly by promoting RHOJ palmitoylation. May act as a palmitoyltransferase for RHOJ: able to autopalmitoylate and then transfer the palmitoyl group to RHOJ. Plays a role in ribosomal 40S subunit biogenesis. Through the interaction with BEST2, inhibits BEST2 channel activity by affecting the gating at the aperture in the absence of intracellular L-glutamate, but sensitizes BEST2 to intracellular L-glutamate, which promotes the opening of BEST2 and thus relieves its inhibitory effect on BEST2.

Subunit / interactions. Decamer; composed of two pentamers. Interacts with PALMD. Interacts with RHOJ. Interacts with BEST2; this interaction tethers a fraction of GLUL to the membrane, causing a decrease of cytosolic glutamine synthase (GS) activity and inhibits the chloride channel activity of BEST2 by affecting the gating at the aperture in the absence of intracellular glutamate.

Subcellular location. Cytoplasm. Cytosol. Microsome. Mitochondrion. Cell membrane.

Tissue specificity. Expressed in endothelial cells.

Post-translational modifications. Acetylated by EP300/p300; acetylation is stimulated by increased glutamine levels and promotes ubiquitin-mediated proteasomal degradation. Palmitoylated; undergoes autopalmitoylation. Ubiquitinated by ZNRF1. Ubiquitinated by the DCX (DDB1-CUL4-X-box) E3 ubiquitin-protein ligase complex called CRL4(CRBN), leading to proteasomal degradation.

Disease relevance. Glutamine deficiency, congenital (GLND) [MIM:610015] An autosomal recessive disorder characterized by variable brain malformations, encephalopathy, severe developmental delay, seizures, and decreased glutamine levels in bodily fluids. Death in early infancy may occur. The disease is caused by variants affecting the gene represented in this entry. Developmental and epileptic encephalopathy 116 (DEE116) [MIM:620806] A form of epileptic encephalopathy, a heterogeneous group of early-onset epilepsies characterized by refractory seizures, neurodevelopmental impairment, and poor prognosis. Development is normal prior to seizure onset, after which cognitive and motor delays become apparent. DEE116 is autosomal dominant form characterized by severe developmental delay, seizures, and white matter abnormalities. The disease is caused by variants affecting the gene represented in this entry. DEE116 is caused by variants that disrupt the canonical translation start codon in GLUL resulting in initiation of translation at Met-18. The resulting protein is enzymatically competent but insensitive to negative feedback regulation via glutamine-induced degradation.

Activity regulation. Glutamine synthetase activity is inhibited by methionine sulfoximine (MSO).

Induction. By glucocorticoids. Vitamin D and the Wnt signaling pathway inhibit its expression and activity. Glutamine synthase (GS) levels are regulated by a negative feedback mechanism where elevated glutamine levels stimulate GS ubiquitin-mediated proteasomal degradation.

Similarity. Belongs to the glutamine synthetase family.

RefSeq proteins (3): NP_001028216, NP_001028228, NP_002056 (=MANE)

Domains & families (InterPro)

ID	Name	Type
IPR008146	Gln_synth_cat_dom	Domain
IPR008147	Gln_synt_N	Domain
IPR014746	Gln_synth/guanido_kin_cat_dom	Homologous_superfamily
IPR027302	Gln_synth_N_conserv_site	Conserved_site
IPR027303	Gln_synth_gly_rich_site	Conserved_site
IPR036651	Gln_synt_N_sf	Homologous_superfamily
IPR050292	Glutamine_Synthetase	Family

Pfam: PF00120, PF03951

Enzyme classification (BRENDA):

• EC 6.3.1.2 — glutamine synthetase (BRENDA: 154 organisms, 215 substrates, 506 inhibitors, 335 Km, 63 kcat entries)

Substrate kinetics (BRENDA)

17 substrates with measured Km, best-characterized 15. Km ranges are aggregated across organisms/conditions.

Substrate	Km (mM)	Measurements
ATP	0.05–28	87
NH4+	0.004–33	55
L-GLUTAMATE	0.2–104	44
HYDROXYLAMINE	0.35–62.5	30
L-GLU	0.0025–50	27
ADP	0.0001–3.8	15
GLU	0.9–14	15
L-GLUTAMINE	0.0067–32.6	13
NH3	0.17–120	10
GLN	4.9–48.6	6
GLUTAMATE	1.4–21.6	6
L-GLN	11.1–60	6
ETHYLAMINE	500–700	2
METHYLAMINE	20–100	2
3-AMINOPENTANEDIOATE	175	1

Catalyzed reactions (Rhea), 2 shown:

• L-glutamate + NH4(+) + ATP = L-glutamine + ADP + phosphate + H(+) (RHEA:16169)
• L-cysteinyl-[protein] + hexadecanoyl-CoA = S-hexadecanoyl-L-cysteinyl-[protein] + CoA (RHEA:36683)

UniProt features (82 total): strand 22, binding site 15, helix 14, mutagenesis site 8, sequence conflict 6, modified residue 5, turn 5, domain 2, sequence variant 2, initiator methionine 1, chain 1, region of interest 1

Structure

Experimental structures (PDB)

12 structures.

PDB	Method	Resolution (Å)
9NM5	X-RAY DIFFRACTION	1.85
9OTP	ELECTRON MICROSCOPY	1.95
9OTO	ELECTRON MICROSCOPY	2.03
2OJW	X-RAY DIFFRACTION	2.05
9OTN	ELECTRON MICROSCOPY	2.11
9OTM	ELECTRON MICROSCOPY	2.19
9OTQ	ELECTRON MICROSCOPY	2.27
9NLR	X-RAY DIFFRACTION	2.3
2QC8	X-RAY DIFFRACTION	2.6
8DNU	ELECTRON MICROSCOPY	2.73
9NR3	X-RAY DIFFRACTION	2.93
7EVT	X-RAY DIFFRACTION	2.95

Predicted structure (AlphaFold)

Model	pLDDT	Fraction very-high
AF-P15104-F1	97.58	0.97

Functional residue map

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

Ligand- & substrate-binding residues (15): 203; 246–247; 253; 255–257; 319; 319; 324; 336–338; 338; 340; 134; 134 …

Post-translational modifications (5): 2, 11, 14, 104, 343

Mutagenesis-validated functional residues (8):

Position	Phenotype
2–17	is stable in high glutamine conditions and does not undergo glutamine-induced degradation.
11	increased ubiquitination and increased proteasomal degradation; when associated with a-14.
11	decreased glutamine-induced acetylation; when associated with r-14. decreased ubiquitination and decreased proteasomal d
14	increased ubiquitination and increased proteasomal degradation; when associated with a-11.
14	decreased glutamine-induced acetylation; when associated with r-11. decreased ubiquitination and decreased proteasomal d
209	reduced ability to mediate autopalmitoylation.
299	loss of glutamine synthase activity. does not affect interaction with best2.
324	decreases ribosomal 40s subunit synthesis. loss of nucleolar location of bysl.

Function

Pathways and Gene Ontology

Reactome pathways

2 pathways

ID	Pathway
R-HSA-210455	Astrocytic Glutamate-Glutamine Uptake And Metabolism
R-HSA-8964539	Glutamate and glutamine metabolism

MSigDB gene sets: 519 (showing top): GSE45365_NK_CELL_VS_CD8_TCELL_DN, GOBP_MYELOID_CELL_DIFFERENTIATION, MODULE_92, GOBP_RIBOSOME_BIOGENESIS, DAVIES_MULTIPLE_MYELOMA_VS_MGUS_DN, KOBAYASHI_EGFR_SIGNALING_24HR_UP, GOBP_GLUTAMINE_FAMILY_AMINO_ACID_BIOSYNTHETIC_PROCESS, WANG_CLIM2_TARGETS_UP, GOBP_POSITIVE_REGULATION_OF_ERYTHROCYTE_DIFFERENTIATION, GOBP_MYELOID_CELL_HOMEOSTASIS, GOBP_LIPOPROTEIN_METABOLIC_PROCESS, GOBP_ALPHA_AMINO_ACID_METABOLIC_PROCESS, GOBP_GLUTAMATE_METABOLIC_PROCESS, GOBP_ERYTHROCYTE_HOMEOSTASIS, GRAESSMANN_APOPTOSIS_BY_DOXORUBICIN_UP

GO Biological Process (13): angiogenesis (GO:0001525), L-glutamate catabolic process (GO:0006538), obsolete glutamine biosynthetic process (GO:0006542), cell population proliferation (GO:0008283), cellular response to starvation (GO:0009267), response to glucose (GO:0009749), regulation of endothelial cell migration (GO:0010594), protein palmitoylation (GO:0018345), ribosome biogenesis (GO:0042254), positive regulation of erythrocyte differentiation (GO:0045648), intracellular ammonium homeostasis (GO:0097275), regulation of sprouting angiogenesis (GO:1903670), regulation of protein localization to nucleolus (GO:1904749)

GO Molecular Function (10): glutamine synthetase activity (GO:0004356), ATP binding (GO:0005524), protein-cysteine S-palmitoyltransferase activity (GO:0019706), identical protein binding (GO:0042802), metal ion binding (GO:0046872), nucleotide binding (GO:0000166), catalytic activity (GO:0003824), protein binding (GO:0005515), transferase activity (GO:0016740), ligase activity (GO:0016874)

GO Cellular Component (10): nucleus (GO:0005634), cytoplasm (GO:0005737), mitochondrion (GO:0005739), cytosol (GO:0005829), plasma membrane (GO:0005886), cell body (GO:0044297), extracellular exosome (GO:0070062), glial cell projection (GO:0097386), endoplasmic reticulum (GO:0005783), membrane (GO:0016020)

Reactome top-level categories

Rollup of top-2 pathways:

Category	Pathways
Neurotransmitter uptake and metabolism In glial cells	1
Metabolism of amino acids and derivatives	1

GO top-level categories

Rollup of top GO terms by namespace:

Category	Terms
cellular anatomical structure	4
intracellular membrane-bounded organelle	3
cytoplasm	3
catalytic activity	2
blood vessel morphogenesis	1
anatomical structure formation involved in morphogenesis	1
glutamate metabolic process	1
dicarboxylic acid catabolic process	1
L-amino acid catabolic process	1
proteinogenic amino acid catabolic process	1
cellular process	1
cellular response to nutrient levels	1
cellular response to stress	1
response to starvation	1
response to hexose	1
regulation of cell migration	1
endothelial cell migration	1
protein lipidation	1
protein acylation	1
ribonucleoprotein complex biogenesis	1
erythrocyte differentiation	1
positive regulation of myeloid cell differentiation	1
regulation of erythrocyte differentiation	1
intracellular chemical homeostasis	1
ammonium homeostasis	1
sprouting angiogenesis	1
regulation of angiogenesis	1
regulation of protein localization to nucleus	1
protein localization to nucleolus	1
ammonia ligase activity	1
adenyl ribonucleotide binding	1
purine ribonucleoside triphosphate binding	1
palmitoyltransferase activity	1
protein-cysteine S-acyltransferase activity	1
protein binding	1
cation binding	1
nucleoside phosphate binding	1
heterocyclic compound binding	1
molecular_function	1
binding	1

Protein interactions and networks

STRING

4714 interactions, top by confidence (×1000):

Protein A	Protein B	Partner UniProt	Score
GLUL	LGSN	Q5TDP6	940
GLUL	HSPD1	P10809	845
GLUL	SLC1A2	P43004	817
GLUL	SLC1A3	P43003	812
GLUL	PRDX4	Q13162	804
GLUL	GLUD1	P00367	797
GLUL	RIPK3	Q9Y572	747
GLUL	ASNS	P08184	733
GLUL	PC	P11498	728
GLUL	H7C2H4	H7C2H4	722
GLUL	GFAP	P14136	717
GLUL	GLS	O94925	717
GLUL	GLS2	Q9UI32	714
GLUL	PLG	P00747	706
GLUL	P0DN79	P0DN79	695

IntAct

113 interactions, top by confidence:

A	B	Type	Score
GLUL	GLUL	psi-mi:“MI:0915”(physical association)	0.850
AMPH	BIN1	psi-mi:“MI:0914”(association)	0.740
CCNC	MED19	psi-mi:“MI:0914”(association)	0.640
RNASEH2C	RNASEH2A	psi-mi:“MI:0914”(association)	0.640
TAT	GLUL	psi-mi:“MI:0915”(physical association)	0.560
NUDT18	GLUL	psi-mi:“MI:0915”(physical association)	0.550
C1orf94	USO1	psi-mi:“MI:0914”(association)	0.550
GLUL	NUDT18	psi-mi:“MI:0915”(physical association)	0.550
TBC1D22B	A2ML1	psi-mi:“MI:0914”(association)	0.530
GLUL	DMXL2	psi-mi:“MI:0914”(association)	0.530
SSBP2	CLEC18A	psi-mi:“MI:0914”(association)	0.530
CREB3	MYO9A	psi-mi:“MI:0914”(association)	0.530
DHDH	ATRN	psi-mi:“MI:0914”(association)	0.530
PNMA2	CCDC85C	psi-mi:“MI:0914”(association)	0.530
PB2	GLUL	psi-mi:“MI:0915”(physical association)	0.510
GLUL	PB2	psi-mi:“MI:0915”(physical association)	0.510
RHOJ	GLUL	psi-mi:“MI:0915”(physical association)	0.510
RHOJ	GLUL	psi-mi:“MI:0915”(physical association)	0.400
PA	GLUL	psi-mi:“MI:0915”(physical association)	0.370

BioGRID (179): GLUL (Two-hybrid), GLUL (Affinity Capture-MS), NME7 (Affinity Capture-MS), DMXL2 (Affinity Capture-MS), TCHP (Affinity Capture-MS), UBAP2 (Affinity Capture-MS), ASPSCR1 (Affinity Capture-MS), PLEKHF2 (Two-hybrid), GLUL (Two-hybrid), NUDT18 (Two-hybrid), GLUL (Affinity Capture-MS), GLUL (Two-hybrid), ZNRF1 (Affinity Capture-Western), GLUL (Affinity Capture-Western), GLUL (Affinity Capture-MS)

ESM2 similar proteins: O00088, O22504, O82560, P00965, P04078, P04770, P08282, P0CN84, P0CN85, P12424, P14654, P15103, P15104, P16580, P23712, P24099, P32288, P32289, P38559, P38560, P38561, P38562, P46410, P51118, P51119, P52782, Q09179, Q12613, Q42899, Q43785, Q4R7U3, Q4W8D0, Q56WN1, Q6B4U7, Q6C3E0, Q6FMT6, Q75BT9, Q86ZF9, Q86ZU6, Q874T6

Diamond homologs: O00088, O04867, O22504, O22506, O82560, P00965, P04078, P04770, P04771, P04772, P04773, P07694, P08281, P08282, P09606, P0CN84, P0CN85, P11600, P12424, P13564, P14636, P14654, P14655, P14656, P15102, P15103, P15104, P15105, P16580, P19432, P20477, P20478, P20805, P22878, P23712, P24099, P25462, P32288, P32289, P34497

SIGNOR signaling

4 interactions.

A	Effect	B	Mechanism
GLUL	“down-regulates quantity”	L-glutamate(1-)	“chemical modification”
GLUL	“down-regulates quantity”	ammonium	“chemical modification”
GLUL	“up-regulates quantity”	“L-glutamine zwitterion”	“chemical modification”
dexamethasone	“up-regulates quantity by expression”	GLUL

Enriched among interaction partners

Reactome pathways and GO biological processes over-represented among this gene’s 112 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
SUMOylation of transcription factors	5	43.3×	5e-05
Formation of Incision Complex in GG-NER	5	19.2×	2e-03

Disease & clinical

Clinical variants and AI predictions

ClinVar

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

Classification	Count (floor)
Pathogenic	5
Likely pathogenic	5
Uncertain significance	146
Likely benign	65
Benign	31

Top pathogenic / likely-pathogenic (10)

Variant ID	HGVS	Classification
1048524	NM_001033044.4(GLUL):c.415del (p.Leu139fs)	Pathogenic
16083	NM_001033044.4(GLUL):c.970C>T (p.Arg324Cys)	Pathogenic
16084	NM_001033044.4(GLUL):c.1021C>T (p.Arg341Cys)	Pathogenic
29734	NM_001033044.4(GLUL):c.970C>A (p.Arg324Ser)	Pathogenic
430447	NM_001033044.4(GLUL):c.1A>T (p.Met1Leu)	Pathogenic
1048523	NM_001033044.4(GLUL):c.1075T>G (p.Cys359Gly)	Likely pathogenic
3899941	NM_001033044.4(GLUL):c.-13-1G>C	Likely pathogenic
3899942	NM_001033044.4(GLUL):c.604T>C (p.Trp202Arg)	Likely pathogenic
4277364	NM_001033044.4(GLUL):c.602A>C (p.Gln201Pro)	Likely pathogenic
4294439	NM_001033044.4(GLUL):c.2T>C (p.Met1Thr)	Likely pathogenic

SpliceAI

935 predictions. Top by Δscore:

Variant	Effect	Δscore
1:182384719:TGTAC:T	acceptor_gain	1.0000
1:182384720:GTAC:G	acceptor_gain	1.0000
1:182384721:TAC:T	acceptor_gain	1.0000
1:182384722:AC:A	acceptor_gain	1.0000
1:182384723:CC:C	acceptor_gain	1.0000
1:182384724:C:CC	acceptor_gain	1.0000
1:182384725:T:G	acceptor_loss	1.0000
1:182384729:G:GC	acceptor_gain	1.0000
1:182385348:GGTAC:G	donor_loss	1.0000
1:182385349:GTAC:G	donor_loss	1.0000
1:182385350:TAC:T	donor_loss	1.0000
1:182385351:AC:A	donor_loss	1.0000
1:182385352:CT:C	donor_loss	1.0000
1:182385353:TCA:T	donor_loss	1.0000
1:182385354:CACTT:C	donor_loss	1.0000
1:182385355:A:AC	donor_gain	1.0000
1:182385356:C:CA	donor_gain	1.0000
1:182385356:C:G	donor_loss	1.0000
1:182385356:CT:C	donor_gain	1.0000
1:182385356:CTT:C	donor_gain	1.0000
1:182385358:TCAG:T	donor_gain	1.0000
1:182385552:TCCCA:T	acceptor_gain	1.0000
1:182385553:CCCA:C	acceptor_gain	1.0000
1:182385553:CCCAC:C	acceptor_gain	1.0000
1:182385554:CCA:C	acceptor_gain	1.0000
1:182385554:CCAC:C	acceptor_gain	1.0000
1:182385555:CA:C	acceptor_gain	1.0000
1:182385555:CAC:C	acceptor_gain	1.0000
1:182385557:C:CC	acceptor_gain	1.0000
1:182385755:CTTA:C	donor_loss	1.0000

AlphaMissense

0 scored. Top likely-pathogenic:

dbSNP variants (sampled 300 via entrez): RS1000190852 (1:182391224 A>G), RS1000298455 (1:182392378 G>C), RS1000655502 (1:182384806 G>A,T), RS1000868450 (1:182378941 C>T), RS1001215764 (1:182391413 G>A), RS1001245873 (1:182389180 A>C,G), RS1001302224 (1:182378586 G>A), RS1001540947 (1:182385181 T>A,C), RS1001636991 (1:182391662 G>A), RS1001697285 (1:182391484 G>A,T), RS1001905585 (1:182379758 A>T), RS1001948770 (1:182380055 C>T), RS1002073592 (1:182384760 C>T), RS1002183824 (1:182390640 G>A,T), RS1002276094 (1:182380076 G>A,C)

Disease associations

OMIM: gene MIM:138290 | disease phenotypes: MIM:610015, MIM:620806

GenCC curated gene-disease

Disease	Classification	Inheritance
congenital brain dysgenesis due to glutamine synthetase deficiency	Definitive	Autosomal recessive
developmental and epileptic encephalopathy 116	Strong	Autosomal dominant

ClinGen Gene-Disease Validity (2)

Expert-panel classifications — Definitive > Strong > Moderate > Limited > Disputed > Refuted.

Disease	Classification	Inheritance
congenital brain dysgenesis due to glutamine synthetase deficiency	Moderate	AR
genetic developmental and epileptic encephalopathy	Moderate	AD

Mondo (2): congenital brain dysgenesis due to glutamine synthetase deficiency (MONDO:0012393), developmental and epileptic encephalopathy 116 (MONDO:0970945)

Orphanet (1): Congenital brain dysgenesis due to glutamine synthetase deficiency (Orphanet:71278)

HPO phenotypes

48 total (30 of 48 shown, HPO-id order):

HPO	Term
HP:0000006	Autosomal dominant inheritance
HP:0000007	Autosomal recessive inheritance
HP:0000233	Thin vermilion border
HP:0000369	Low-set ears
HP:0000431	Wide nasal bridge
HP:0000463	Anteverted nares
HP:0001250	Seizure
HP:0001252	Hypotonia
HP:0001290	Generalized hypotonia
HP:0001298	Encephalopathy
HP:0001371	Flexion contracture
HP:0001662	Bradycardia
HP:0001987	Hyperammonemia
HP:0002069	Bilateral tonic-clonic seizure
HP:0002079	Hypoplasia of the corpus callosum
HP:0002104	Apnea
HP:0002121	Generalized non-motor (absence) seizure
HP:0002197	Generalized-onset seizure
HP:0002205	Recurrent respiratory infections
HP:0002266	Focal clonic seizure
HP:0002395	Lower limb hyperreflexia
HP:0002416	Subependymal cysts
HP:0002643	Neonatal respiratory distress
HP:0002983	Micromelia
HP:0003196	Short nose
HP:0003429	CNS hypomyelination
HP:0003593	Infantile onset
HP:0003811	Neonatal death
HP:0005280	Depressed nasal bridge
HP:0006956	Lateral ventricle dilatation

GWAS associations

50 associations (top):

Study	Trait	p-value
GCST002153_1	Cardiovascular heart disease in diabetics	2.000000e-08
GCST007565_115	Morning person	2.000000e-25
GCST007565_144	Morning person	3.000000e-16
GCST010242_213	HDL cholesterol levels	7.000000e-09
GCST010796_2274	Electrocardiogram morphology (amplitude at temporal datapoints)	4.000000e-08
GCST010796_2275	Electrocardiogram morphology (amplitude at temporal datapoints)	3.000000e-08
GCST010796_2276	Electrocardiogram morphology (amplitude at temporal datapoints)	2.000000e-08
GCST010796_2277	Electrocardiogram morphology (amplitude at temporal datapoints)	5.000000e-09
GCST010796_2278	Electrocardiogram morphology (amplitude at temporal datapoints)	4.000000e-09
GCST010796_2279	Electrocardiogram morphology (amplitude at temporal datapoints)	1.000000e-09
GCST010796_2280	Electrocardiogram morphology (amplitude at temporal datapoints)	8.000000e-10
GCST010796_2281	Electrocardiogram morphology (amplitude at temporal datapoints)	3.000000e-10
GCST010796_2282	Electrocardiogram morphology (amplitude at temporal datapoints)	8.000000e-10
GCST010796_2283	Electrocardiogram morphology (amplitude at temporal datapoints)	4.000000e-09
GCST010796_2284	Electrocardiogram morphology (amplitude at temporal datapoints)	3.000000e-09
GCST010796_2285	Electrocardiogram morphology (amplitude at temporal datapoints)	7.000000e-10
GCST010796_2286	Electrocardiogram morphology (amplitude at temporal datapoints)	2.000000e-09
GCST010796_2287	Electrocardiogram morphology (amplitude at temporal datapoints)	4.000000e-09
GCST010796_2288	Electrocardiogram morphology (amplitude at temporal datapoints)	4.000000e-09
GCST010796_2289	Electrocardiogram morphology (amplitude at temporal datapoints)	3.000000e-08
GCST010796_2290	Electrocardiogram morphology (amplitude at temporal datapoints)	4.000000e-08
GCST010796_2291	Electrocardiogram morphology (amplitude at temporal datapoints)	5.000000e-11
GCST010796_2292	Electrocardiogram morphology (amplitude at temporal datapoints)	4.000000e-13
GCST010796_2293	Electrocardiogram morphology (amplitude at temporal datapoints)	4.000000e-11
GCST010796_2294	Electrocardiogram morphology (amplitude at temporal datapoints)	1.000000e-08
GCST010796_2295	Electrocardiogram morphology (amplitude at temporal datapoints)	2.000000e-09
GCST010796_2296	Electrocardiogram morphology (amplitude at temporal datapoints)	3.000000e-10
GCST010796_2297	Electrocardiogram morphology (amplitude at temporal datapoints)	2.000000e-09
GCST010796_2298	Electrocardiogram morphology (amplitude at temporal datapoints)	2.000000e-10
GCST010796_2299	Electrocardiogram morphology (amplitude at temporal datapoints)	9.000000e-13

EFO canonical traits (3, from GWAS)

EFO ID	Trait name
EFO:0008328	chronotype measurement
EFO:0004612	high density lipoprotein cholesterol measurement
EFO:0004327	electrocardiography

MeSH disease descriptors (1)

Descriptor	Name	Tree numbers
C536832	Glutamine deficiency, congenital (supp.)

Drugs & pharmacology

Drug and pharmacology data

Is drug target: yes

ChEMBL targets (1): CHEMBL4612 (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: 140,604 (via chembl_molecule»patent_compound — counts attach to the compound, not the gene–compound relationship, so off-target/promiscuous molecules can dominate).

Molecule	Name	Phase	Patents
CHEMBL131	PREDNISOLONE	4	140,604

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

Binding affinities (BindingDB)

6 measured of 6 human assays (6 total across all organisms); most potent 6 below. Values come from heterogeneous assays and are not directly comparable.

Ligand	Measure	Value
Phosphinothricin analog, 1 (L Isomer)	KI	0.22 nM
Phosphinothricin analog, 4 (D,L Isomer)	KI	0.47 nM
Phosphinothricin analog, 6 (D,L Isomer)	KI	5 nM
Phosphinothricin analog, 3 (L Isomer)	KI	45 nM
Phosphinothricin analog, 2 (D,L mixture)	KI	140 nM
Phosphinothricin analog, 5 (D,L Isomer)	KI	310 nM

ChEMBL bioactivities

9 potent at pChembl≥5 of 10 total, top 9 by pChembl (potency: 10 = 0.1 nM, 6 = 1 µM).

pChembl	Type	Value	Unit	Molecule
7.85	EC50	14.2	nM	CHEMBL310570
7.50	EC50	32	nM	PREDNISOLONE
7.42	EC50	37.7	nM	CHEMBL83110
7.27	EC50	53.7	nM	CHEMBL82546
7.18	EC50	66	nM	CHEMBL309243
7.18	EC50	66	nM	CHEMBL312399
6.89	EC50	129.2	nM	CHEMBL311662
6.74	EC50	181.6	nM	CHEMBL312041
6.07	Ki	850	nM	CHEMBL3142429

PubChem BioAssay actives

9 with measured affinity, of 118 total; 9 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
(1S)-1-[(4aR,5S)-1-(4-fluorophenyl)-4a-methyl-5,6,7,8-tetrahydro-4H-benzo[f]indazol-5-yl]-1-thiophen-3-ylethanol	74963: Ability to induce human glutamine synthetase in skeletal muscle cells	ec50	0.0142	uM
Prednisolone	74963: Ability to induce human glutamine synthetase in skeletal muscle cells	ec50	0.0320	uM
(R)-[(4aR,5S)-1-(4-fluorophenyl)-4a-methyl-5,6,7,8-tetrahydro-4H-benzo[f]indazol-5-yl]-(4-fluoro-3-methylphenyl)methanol	74963: Ability to induce human glutamine synthetase in skeletal muscle cells	ec50	0.0377	uM
(1S)-1-[(4aR,5S)-1-(4-fluorophenyl)-4a-methyl-5,6,7,8-tetrahydro-4H-benzo[f]indazol-5-yl]-2-(4-fluorophenyl)ethanol	74963: Ability to induce human glutamine synthetase in skeletal muscle cells	ec50	0.0537	uM
(1S)-1-[(4aR,5S)-1-(4-fluorophenyl)-4a-methyl-5,6,7,8-tetrahydro-4H-benzo[f]indazol-5-yl]but-3-en-1-ol	74963: Ability to induce human glutamine synthetase in skeletal muscle cells	ec50	0.0660	uM
(1S)-1-[(4aR,5S)-1-(4-fluorophenyl)-4a-methyl-5,6,7,8-tetrahydro-4H-benzo[f]indazol-5-yl]-3-cyclopropylpropan-1-ol	74963: Ability to induce human glutamine synthetase in skeletal muscle cells	ec50	0.0660	uM
(1S)-1-[(4aR,5S)-1-(4-fluorophenyl)-4a-methyl-5,6,7,8-tetrahydro-4H-benzo[f]indazol-5-yl]-1-(4-fluorophenyl)ethanol	74963: Ability to induce human glutamine synthetase in skeletal muscle cells	ec50	0.1292	uM
2-[(4aR,5S)-1-(4-fluorophenyl)-4a-methyl-5,6,7,8-tetrahydro-4H-benzo[f]indazol-5-yl]propan-2-ol	74963: Ability to induce human glutamine synthetase in skeletal muscle cells	ec50	0.1816	uM
tert-butyl N-[(2S)-1-[[(2S)-1-[[(2S,3S,5R)-1-cyclohexyl-3-hydroxy-7-methyl-5-(methylcarbamoyl)octan-2-yl]amino]-3-(1H-imidazol-5-yl)-1-oxopropan-2-yl]amino]-1-oxo-3-phenylpropan-2-yl]carbamate	75127: Compound was tested for binding affinity against Glutamine synthetase	ki	0.8500	uM

CTD chemical–gene interactions

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

Chemical	Actions (top 5)	PubMed papers
Benzo(a)pyrene	decreases expression, decreases methylation, increases expression	4
Tobacco Smoke Pollution	decreases expression, increases expression, affects expression	4
Tretinoin	increases expression, decreases expression, affects cotreatment	4
Air Pollutants	increases abundance, increases oxidation, affects expression, decreases expression, affects cotreatment	3
Cyclosporine	decreases expression	3
bisphenol A	increases methylation, increases expression	2
trichostatin A	affects cotreatment, increases expression	2
sodium arsenite	increases expression, decreases expression, increases abundance	2
4-(5-benzo(1,3)dioxol-5-yl-4-pyridin-2-yl-1H-imidazol-2-yl)benzamide	affects cotreatment, increases expression	2
Arsenic Trioxide	affects binding, decreases reaction, increases expression, affects cotreatment	2
Acetaminophen	decreases expression	2
Copper	affects binding, decreases expression, increases expression	2
Dexamethasone	increases expression, affects cotreatment	2
Doxorubicin	decreases expression, increases expression, affects reaction	2
Estradiol	affects cotreatment, decreases expression, increases expression	2
Indomethacin	decreases expression, affects cotreatment, increases expression	2
Nickel	increases expression	2
Ozone	affects cotreatment, increases oxidation, increases abundance, affects expression	2
Silicon Dioxide	decreases expression	2
Valproic Acid	affects expression, increases expression	2
aristolochic acid I	increases expression	1
bisphenol F	increases expression	1
sotorasib	decreases expression, affects cotreatment	1
4-oxoretinoic acid	increases expression	1
bufotalin	decreases expression	1
methyleugenol	decreases expression	1
triphenyl phosphate	affects expression	1
alpha-pinene	increases oxidation, increases abundance, affects cotreatment	1
uranyl acetate	affects expression	1
lead acetate	decreases expression	1

ChEMBL screening assays

45 unique, capped per target: 45 binding

Representative assays (with source publication via chembl_document):

Assay ID	Type	Description	Source paper
CHEMBL682550	Binding	Tested for the inhibition of Glutamine synthetase (Escherichia coli) activity using biosynthetic assay [50 nM Hepes (pH 7), 5 mM MgCl2] at a concentration of 0.1 mM	Design and synthesis of phosphonate inhibitors of glutamine synthetase. — J Med Chem

Cellosaurus cell lines

11 cell lines: 9 cancer cell line, 2 transformed cell line

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

Cellosaurus	Name	Category	Sex
CVCL_8560	HepG2-GS-CYP3A4	Cancer cell line	Male
CVCL_D1MV	Abcam K-562 GLUL KO	Cancer cell line	Female
CVCL_D1ST	Abcam U-87MG GLUL KO	Cancer cell line	Male
CVCL_D2JF	Abcam Raji GLUL KO	Cancer cell line	Male
CVCL_E0UB	Ubigene Hep G2 GLUL KO	Cancer cell line	Male
CVCL_E3KD	HEK293SF-3F6 GLUL KO	Transformed cell line	Female
CVCL_H214	HepG2-GS	Cancer cell line	Male
CVCL_JZ97	HEK293_GS-KO	Transformed cell line	Female
CVCL_SQ11	HAP1 GLUL (-) 1	Cancer cell line	Male
CVCL_UQ62	Abcam Jurkat GLUL KO	Cancer cell line	Male

Clinical trials (associated diseases)

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