FUT4

Summary

FUT4 (fucosyltransferase 4, HGNC:4015) is a protein-coding gene on chromosome 11q21, encoding Alpha-(1,3)-fucosyltransferase 4 (P22083). Catalyzes alpha(1->3) linkage of fucosyl moiety transferred from GDP-beta-L-fucose to N-acetyl glucosamine (GlcNAc) within type 2 lactosamine (LacNAc, Gal-beta(1->4)GlcNAc) glycan attached to N- or O-linked glycoproteins.

The product of this gene transfers fucose to N-acetyllactosamine polysaccharides to generate fucosylated carbohydrate structures. It catalyzes the synthesis of the non-sialylated antigen, Lewis x (CD15).

Source: NCBI Gene 2526 — RefSeq curated summary.

At a glance

• GWAS associations: 5
• Clinical variants (ClinVar): 91 total — 1 likely-pathogenic
• Druggable target: yes — 1 molecules with ChEMBL bioactivity
• MANE Select transcript: NM_002033

Identifiers

Gene identifiers

Field	Value
HGNC ID	HGNC:4015
Approved symbol	FUT4
Name	fucosyltransferase 4
Location	11q21
Locus type	gene with protein product
Status	Approved
Aliases	FUC-TIV
Ensembl gene	ENSG00000196371
Ensembl biotype	protein_coding
OMIM	104230
Entrez	2526

Gene structure

Transcript identifiers

Ensembl transcripts: 1 — 1 protein_coding

ENST00000358752

RefSeq mRNA: 1 — MANE Select: NM_002033 NM_002033

CCDS: CCDS8301

Canonical transcript exons

ENST00000358752 — 1 exons

Exon	Start	End
ENSE00001416465	94543921	94549895

Expression profiles

Bgee: expression breadth ubiquitous, 214 present calls, max score 91.00.

FANTOM5 (CAGE): breadth ubiquitous, TPM avg 9.8920 / max 838.6499, expressed in 1726 samples.

FANTOM5 promoters (5 alternative TSS)

Promoter ID	TPM avg	Samples expressed
116262	3.6571	1191
116261	2.9927	1191
116260	1.5827	865
116263	1.4353	541
206416	0.2241	58

Top tissues by expression

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

Tissue	Anatomy ID	Expression score	Quality
mucosa of sigmoid colon	UBERON:0004993	91.00	gold quality
bone marrow	UBERON:0002371	89.53	gold quality
ileal mucosa	UBERON:0000331	89.42	gold quality
colonic mucosa	UBERON:0000317	88.69	gold quality
trabecular bone tissue	UBERON:0002483	87.04	gold quality
monocyte	CL:0000576	86.30	gold quality
mononuclear cell	CL:0000842	85.98	gold quality
bone marrow cell	CL:0002092	85.97	gold quality
jejunal mucosa	UBERON:0000399	85.65	gold quality
leukocyte	CL:0000738	85.32	gold quality
duodenum	UBERON:0002114	84.95	gold quality
primordial germ cell in gonad	CL:0000670 ∩ UBERON:0000991	82.08	gold quality
rectum	UBERON:0001052	79.51	gold quality
tendon of biceps brachii	UBERON:0008188	79.01	gold quality
mucosa of transverse colon	UBERON:0004991	78.98	gold quality
granulocyte	CL:0000094	76.16	gold quality
tendon	UBERON:0000043	76.10	gold quality
tibia	UBERON:0000979	75.44	gold quality
male germ line stem cell (sensu Vertebrata) in testis	CL:0000089 ∩ UBERON:0000473	75.29	gold quality
islet of Langerhans	UBERON:0000006	75.21	gold quality
large intestine	UBERON:0000059	74.60	gold quality
transverse colon	UBERON:0001157	74.58	gold quality
intestine	UBERON:0000160	74.52	gold quality
colon	UBERON:0001155	74.24	gold quality
small intestine	UBERON:0002108	74.17	gold quality
pancreas	UBERON:0001264	74.05	gold quality
vermiform appendix	UBERON:0001154	73.68	gold quality
body of stomach	UBERON:0001161	73.53	gold quality
calcaneal tendon	UBERON:0003701	73.46	gold quality
body of pancreas	UBERON:0001150	73.33	gold quality

Single-cell (SCXA)

Detected in 3 experiment(s), a significant marker in 3.

Experiment	Marker?	Max mean expression
E-MTAB-9067	yes	13.72
E-MTAB-9801	yes	4.25
E-ANND-3	yes	3.96

Regulation

Is transcription factor: no

Upstream regulators (CollecTRI, top): ELK1, HSF1, SP1

miRNA regulators (miRDB)

151 targeting FUT4, 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-5011-5P	100.00	83.46	5820
HSA-MIR-3613-3P	100.00	76.36	7965
HSA-MIR-5692A	100.00	74.40	6850
HSA-MIR-1277-5P	100.00	73.95	5056
HSA-MIR-190A-3P	100.00	80.35	5520
HSA-MIR-200B-3P	100.00	73.31	2693
HSA-MIR-200C-3P	100.00	73.35	2685
HSA-MIR-429	100.00	73.44	2698
HSA-MIR-513A-5P	100.00	69.77	2465
HSA-MIR-5692B	100.00	71.32	2622
HSA-MIR-5692C	100.00	71.32	2622
HSA-MIR-30A-5P	100.00	76.31	3233
HSA-MIR-30D-5P	100.00	76.32	3233
HSA-MIR-30E-5P	100.00	76.32	3242
HSA-MIR-371B-5P	99.99	75.34	4759
HSA-MIR-548AW	99.99	72.57	3559
HSA-MIR-548C-3P	99.99	74.01	7587
HSA-MIR-3662	99.99	73.82	5684
HSA-MIR-33A-5P	99.99	68.62	1055
HSA-MIR-33B-5P	99.99	68.58	1062
HSA-MIR-19A-3P	99.98	75.33	2762
HSA-MIR-19B-3P	99.98	75.44	2754
HSA-MIR-373-5P	99.98	75.36	4753
HSA-MIR-616-5P	99.98	75.58	4775
HSA-MIR-548P	99.98	72.25	3784
HSA-MIR-1468-3P	99.96	72.74	3797
HSA-MIR-590-3P	99.96	74.34	6478
HSA-MIR-548AT-5P	99.96	70.83	2666
HSA-MIR-548AJ-3P	99.96	73.38	5345
HSA-MIR-548X-3P	99.96	73.38	5345

Literature-anchored findings (GeneRIF, showing 40)

• Strong activation of Ras represents a major pathway for induction of FucT-VII gene expression in T cells. (PMID:12738675)
• induction of FUT (mainly FUT4), the gene expression of which is mediated by signals downstream of caspase 3, increases Le(X) and Le(Y) expression in apoptotic cells (PMID:15182935)
• human FucT-IV and -VII both contribute and cooperate in regulating L-selectin-, P-selectin-, and E-selectin-dependent rolling on PSGL-1, with FucT-VII playing a predominant role in conferring selectin binding activity to PSGL-1 (PMID:15579466)
• interaction of sialyl Lewis X antigen with lectin-like receptors on NK cells induces cytotoxicity that is mediated through a tyrosine-phosphorylated 17-kDa protein (PMID:16842926)
• Granzyme treatment increased cell surface FUT4 expression. (PMID:17409497)
• Fucosyltransferase IV overexpression augments Lewis Y expression to trigger neoplastic cell proliferation. (PMID:17556010)
• Expression is frequenty decreased in chronic myelogenous leukemia. (PMID:17662271)
• Suppressing the expression of FUT1/4 by RNAi technology reduces the synthesis of LeY and inhibits cancer growth. (PMID:18023290)
• CD13 was expressed in 73% of acute myeloid leukemia patients, CD15 was expressed in 43% of patients, CD33 was expressed in 64% of patients, and CD34 was expressed in 66% of patients. (PMID:18085638)
• Data show that tumors from a model of medulloblastoma, the Patched mutant mouse, are propagated not by CD133(+) cells but by cells expressing the progenitor markers Math1 and CD15/SSEA-1. (PMID:19185848)
• Data show that CD45, CD11b, CD15, and CD56 were diagnostic parameters with flow cytometry. (PMID:19546439)
• Findings suggest the important role of the CD11b+/CD14/CD15+/CD33+ MDSCs in mediating immunosuppression in NSCLC. (PMID:19572148)
• Dynamic changes in surface expression of the identified antigens CD15, CD24, and CD29 in combination can be exploited for the experimental separation of key neural cell populations. (PMID:19725119)
• CD133, CD15/SSEA-1, and CD34 do not resume tumor-initiating properties of long-term cultured malignant glio-neuronal stem cells (PMID:20181261)
• The presence of nanog, Oct-4, SSEA-1, and SSEA-4 suggests that periodontal ligament mesenchymal stem cells are less differentiated than bone marrow-derived MSCs, and that the frizzled-9/Wnt pathway is important in proliferation and differentiation. (PMID:20458727)
• These studies suggest that FUT4 regulates A431 cell growth through controlling cell cycle progression via MAPK and PI3K/Akt signaling pathways. (PMID:20506505)
• In endometrial tissues, knockdown of FUT4 expression reduces expression of Lewis Y antigen but not of integrin alphavbeta3. (PMID:20605574)
• The biosynthesis of the selectin-ligand sialyl Lewis x in colorectal cancer tissues is regulated by fucosyltransferase VI and can be inhibited by an RNA interference-based approach. (PMID:20965272)
• FUT4 inhibited cell apoptosis through decreasing the expression of apoptotic proteins and increasing the expression of anti-apoptotic protein Bcl-2 (PMID:21337384)
• CD15-expressing nodular lymphocyte-predominant Hodgkin lymphoma (PMID:21457163)
• NEU4 plays an important role in control of sialyl Lewis antigen expression and its impairment in colon cancer (PMID:21521691)
• Data indicate that among MDS cases, CD15+ and CD19+ cell TLs were positively correlated, and PBL TL was was not associated with hTERT genotype. (PMID:21635204)
• Data show that tumours induced by transformed fibroblasts are hierarchically organized, and the cells that act as CSCs to initiate and maintain tumour growth are marked by the stage-specific embryonic antigen SSEA-1. (PMID:21857669)
• MiR-34a targeting of Notch ligand delta-like 1 impairs CD15+/CD133+ tumor-propagating cells and supports neural differentiation in medulloblastoma (PMID:21931765)
• Resting natural killer (NK) cells that have been coincubated with NK-resistant CD15-positive tumor cells lyse Raji cells and are capable of lysing a variety of NK-resistant tumor cells of different lineages. (PMID:22084431)
• the expression of Lewis Y and FUT4 correlates with endometrial receptivity (PMID:22145955)
• FUT4 expression is negatively correlated with the methylation degree of a CpG island in the FUT4 promoter. (PMID:22287018)
• High CD15 expression is associated with medulloblastoma. (PMID:22411914)
• TNF increases the expression of alpha2,3-sialyltransferase gene ST3GAL4 (PMID:22691873)
• Lewis x appeared to be a new, reliable marker that can be used to clearly distinguish invasive hydatidiform moles from choriocarcinomas. (PMID:23681114)
• The expression of ST3GAL4 leads to SLe(x) antigen expression in gastric cancer cells which in turn induces an increased invasive phenotype through the activation of c-Met (PMID:23799130)
• FUT4 is a target gene for HSF1 and Sp1 that is required for cell cycle progression in breast cancer epithelial cells.HSF1 and Sp1 regulate FUT4 gene expression and cell proliferation in breast cancer cells. (PMID:23959823)
• AQP9 expressing glioma cells were negative for the brain tumor stem cell marker CD15. (PMID:24086629)
• MMP-2 and sLe(x) were negative prognostic markers for survival in these head and neck squamous cell carcinoma patients. (PMID:24171785)
• Results suggest that FUT4-, FUT6- or FUT8-mediated MDR in human HCC is associated with the activation of the PI3K/Akt pathway and the expression of MRP1. (PMID:24232099)
• The correlation of sLex overexpression in primary tumors and metastatic lymph nodes, the discrimination among the normal, adenoma, and CRC groups based on sLex expression. (PMID:24425323)
• Physiologically immature placentas and pathologically immature term placentas were characterized by marked endothelial CD15-immunostaining. Significant loss of CD15-positive endothelium was associated with physiological and accelerated villous maturity. (PMID:25043745)
• Titration of the monomeric DC-SIGN CRD with Le(X) monitored by 2D NMR revealed significant perturbations of DC-SIGN cross-peak positions in (1)H-(15)N heteronuclear single quantum coherence (HSQC) spectra and identified residues near the binding site. (PMID:25121780)
• level of expression in the macro- and microvasculature reflects the degree of pathological placental villous immaturity (PMID:25149387)
• The lower expression of FUT4 in HaCaT cells is correlated with the methylation of CpG island in FUT4 promoter. (PMID:25608813)

Cross-species orthologs

15 orthologs

Organism	Symbol	Gene ID
mus_musculus	Fut4	ENSMUSG00000049307
rattus_norvegicus	Fut4	ENSRNOG00000009157
drosophila_melanogaster	FucTD	FBGN0035217
caenorhabditis_elegans		WBGENE00001505
caenorhabditis_elegans		WBGENE00001507
caenorhabditis_elegans		WBGENE00004010
caenorhabditis_elegans		WBGENE00006402
caenorhabditis_elegans		WBGENE00007211
caenorhabditis_elegans		WBGENE00012922
caenorhabditis_elegans		WBGENE00016163
caenorhabditis_elegans		WBGENE00017343
caenorhabditis_elegans		WBGENE00043986
caenorhabditis_elegans		WBGENE00044383
caenorhabditis_elegans		WBGENE00194870
caenorhabditis_elegans		WBGENE00206359

Paralogs (7): FUT5 (ENSG00000130383), FUT6 (ENSG00000156413), FUT3 (ENSG00000171124), FUT9 (ENSG00000172461), FUT10 (ENSG00000172728), FUT7 (ENSG00000180549), FUT11 (ENSG00000196968)

Protein

Protein identifiers

Alpha-(1,3)-fucosyltransferase 4 — P22083 (reviewed: P22083)

Alternative names: 4-galactosyl-N-acetylglucosaminide 3-alpha-L-fucosyltransferase, ELAM-1 ligand fucosyltransferase, Fucosyltransferase 4, Fucosyltransferase IV, Galactoside 3-L-fucosyltransferase

All UniProt accessions (1): P22083

UniProt curated annotations — full annotation on UniProt →

Function. Catalyzes alpha(1->3) linkage of fucosyl moiety transferred from GDP-beta-L-fucose to N-acetyl glucosamine (GlcNAc) within type 2 lactosamine (LacNAc, Gal-beta(1->4)GlcNAc) glycan attached to N- or O-linked glycoproteins. Robustly fucosylates nonsialylated distal LacNAc unit of the polylactosamine chain to form Lewis X antigen (CD15), a glycan determinant known to mediate important cellular functions in development and immunity. Fucosylates with lower efficiency sialylated LacNAc acceptors to form sialyl Lewis X and 6-sulfo sialyl Lewis X determinants that serve as recognition epitopes for C-type lectins. Together with FUT7 contributes to SELE, SELL and SELP selectin ligand biosynthesis and selectin-dependent lymphocyte homing, leukocyte migration and blood leukocyte homeostasis. In a cell type specific manner, may also fucosylate the internal LacNAc unit of the polylactosamine chain to form VIM-2 antigen that serves as recognition epitope for SELE. Does not generate Lewis X antigens.

Subcellular location. Golgi apparatus. Golgi stack membrane.

Tissue specificity. Expressed at low levels in bone marrow-derived mesenchymal stem cells. Expressed in cord blood immature promyelocytes and in peripheral blood myeloid and lymphoid cell populations.

Pathway. Protein modification; protein glycosylation.

Similarity. Belongs to the glycosyltransferase 10 family.

Isoforms (2)

UniProt ID	Names	Canonical?
P22083-1	Long, ELFT-L	yes
P22083-2	Short, ELFT

RefSeq proteins (1): NP_002024* (*=MANE)

Domains & families (InterPro)

ID	Name	Type
IPR001503	Glyco_trans_10	Family
IPR031481	Glyco_tran_10_N	Domain
IPR038577	GT10-like_C_sf	Homologous_superfamily
IPR055270	Glyco_tran_10_C	Domain

Pfam: PF00852, PF17039

Enzyme classification (BRENDA):

• EC 2.4.1.152 — 4-galactosyl-N-acetylglucosaminide 3-alpha-L-fucosyltransferase (BRENDA: 24 organisms, 171 substrates, 55 inhibitors, 103 Km, 24 kcat entries)

Substrate kinetics (BRENDA)

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

Substrate	Km (mM)	Measurements
GDP-BETA-L-FUCOSE	0.0001–1.12	18
N-ACETYLLACTOSAMINE	0.0026–0.61	9
D-LACTOSE	72–543	6
GDP-FUCOSE	0.0016–0.06	5
GDP-L-FUCOSE	0.0055–0.062	5
FUCALPHA(1,2)GALBETA(1,4)GLCNAC	0.7–3.9	4
GALBETA1,4GLCNAC-O(CH2)8CO2CH3	0.37–1.7	4
GALBETA(1,4)GLCNAC	1.4–23	3
GDP-BETA-FUCOSE	0.021–6.448	3
FUCALPHA(1,2)GALBETA(1,4)GLC	6.7–20	2
LACTO-N-NEOTETRAOSE	0.3–0.76	2
LACTOSE	3.729–83	2
NEUACALPHA(2,3)GALBETA(1,4)GLCNAC	0.2–0.9	2
3’-SIALYL N-ACETYL LACTOSAMINE	0.65	1
6-DEOXY-6-N-(2-NAPHTHALENE-2-YL-ACETAMIDE)-BETA-	0.0009	1

Catalyzed reactions (Rhea), 4 shown:

• a beta-D-galactosyl-(1->4)-N-acetyl-beta-D-glucosaminyl derivative + GDP-beta-L-fucose = a beta-D-galactosyl-(1->4)-[alpha-L-fucosyl-(1->3)]-N-acetyl-beta-D-glucosaminyl derivative + GDP + H(+) (RHEA:14257)
• an N-acetyl-alpha-neuraminyl-(2->3)-beta-D-galactosyl-(1->4)-N-acetyl-beta-D-glucosaminyl derivative + GDP-beta-L-fucose = an alpha-Neu5Ac-(2->3)-beta-D-Gal-(1->4)-[alpha-L-Fuc-(1->3)]-beta-D-GlcNAc derivative + GDP + H(+) (RHEA:56076)
• an alpha-Neu5Ac-(2->3)-beta-D-Gal-(1->4)-beta-D-GlcNAc6S derivative + GDP-beta-L-fucose = an alpha-Neu5Ac-(2->3)-beta-D-Gal-(1->4)-[alpha-L-Fuc-(1->3)]-beta-D-GlcNAc6S derivative + GDP + H(+) (RHEA:62004)
• an alpha-Neu5Ac-(2->3)-beta-D-Gal-(1->4)-beta-D-GlcNAc-(1->3)-beta-D-Gal-(1->4)-beta-D-GlcNAc derivative + GDP-beta-L-fucose = an alpha-Neu5Ac-(2->3)-beta-D-Gal-(1->4)-beta-D-GlcNAc-(1->3)-beta-D-Gal-(1->4)-[alpha-L-Fuc-(1->3)]-beta-D-GlcNAc derivative + GDP + H(+) (RHEA:68044)

UniProt features (13 total): topological domain 2, sequence conflict 2, region of interest 2, glycosylation site 2, chain 1, sequence variant 1, transmembrane region 1, compositionally biased region 1, splice variant 1

Structure

Experimental structures (PDB)

0 structures.

Predicted structure (AlphaFold)

Model	pLDDT	Fraction very-high
AF-P22083-F1	74.21	0.53

Functional residue map

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

Glycosylation sites (2): 216, 315

Function

Pathways and Gene Ontology

Reactome pathways

1 pathways

ID	Pathway
R-HSA-9037629	Lewis blood group biosynthesis

MSigDB gene sets: 238 (showing top): GSE37336_LY6C_POS_VS_NEG_NAIVE_CD4_TCELL_UP, GOBP_OLIGOSACCHARIDE_METABOLIC_PROCESS, BUYTAERT_PHOTODYNAMIC_THERAPY_STRESS_DN, BENPORATH_ES_WITH_H3K27ME3, GOBP_MYELOID_LEUKOCYTE_MIGRATION, GOBP_INFLAMMATORY_RESPONSE, GOBP_GLYCOLIPID_BIOSYNTHETIC_PROCESS, GOBP_PROTEIN_N_LINKED_GLYCOSYLATION, IVANOVA_HEMATOPOIESIS_LATE_PROGENITOR, GOCC_CELL_SURFACE, GOBP_MONOSACCHARIDE_CATABOLIC_PROCESS, NAGASHIMA_NRG1_SIGNALING_UP, GOBP_REGULATION_OF_LEUKOCYTE_MIGRATION, GOBP_CARBOHYDRATE_DERIVATIVE_METABOLIC_PROCESS, GOBP_POSITIVE_REGULATION_OF_CELL_ADHESION

GO Biological Process (17): carbohydrate metabolic process (GO:0005975), protein N-linked glycosylation (GO:0006487), protein O-linked glycosylation (GO:0006493), sphingolipid metabolic process (GO:0006665), glycosphingolipid biosynthetic process (GO:0006688), inflammatory response (GO:0006954), glycoprotein biosynthetic process (GO:0009101), oligosaccharide metabolic process (GO:0009311), oligosaccharide biosynthetic process (GO:0009312), L-fucose catabolic process (GO:0042355), lymphocyte migration into lymph node (GO:0097022), Lewis x epitope biosynthetic process (GO:0106402), positive regulation of neutrophil migration (GO:1902624), regulation of leukocyte cell-cell adhesion (GO:1903037), positive regulation of leukocyte tethering or rolling (GO:1903238), obsolete protein glycosylation (GO:0006486), obsolete fucosylation (GO:0036065)

GO Molecular Function (5): fucosyltransferase activity (GO:0008417), 4-galactosyl-N-acetylglucosaminide 3-alpha-L-fucosyltransferase activity (GO:0017083), alpha-(1->3)-fucosyltransferase activity (GO:0046920), transferase activity (GO:0016740), glycosyltransferase activity (GO:0016757)

GO Cellular Component (7): Golgi membrane (GO:0000139), Golgi apparatus (GO:0005794), trans-Golgi network (GO:0005802), cell surface (GO:0009986), membrane (GO:0016020), Golgi cisterna membrane (GO:0032580), cell periphery (GO:0071944)

Reactome top-level categories

Rollup of top-1 pathways:

Category	Pathways
Blood group systems biosynthesis	1

GO top-level categories

Rollup of top GO terms by namespace:

Category	Terms
cellular anatomical structure	3
glycoprotein biosynthetic process	2
carbohydrate derivative biosynthetic process	2
primary metabolic process	1
lipid metabolic process	1
glycosphingolipid metabolic process	1
glycolipid biosynthetic process	1
sphingolipid biosynthetic process	1
defense response	1
macromolecule biosynthetic process	1
glycoprotein metabolic process	1
carbohydrate metabolic process	1
oligosaccharide metabolic process	1
carbohydrate biosynthetic process	1
hexose catabolic process	1
L-fucose metabolic process	1
lymphocyte migration into lymphoid organs	1
positive regulation of leukocyte migration	1
regulation of neutrophil migration	1
neutrophil migration	1
leukocyte cell-cell adhesion	1
regulation of cell-cell adhesion	1
leukocyte tethering or rolling	1
regulation of leukocyte tethering or rolling	1
positive regulation of leukocyte adhesion to vascular endothelial cell	1
hexosyltransferase activity	1
alpha-(1->3)-fucosyltransferase activity	1
fucosyltransferase activity	1
catalytic activity	1
transferase activity	1
Golgi apparatus	1
bounding membrane of organelle	1
cytoplasm	1
endomembrane system	1
intracellular membrane-bounded organelle	1
Golgi apparatus subcompartment	1
organelle membrane	1
Golgi cisterna	1

Protein interactions and networks

STRING

1940 interactions, top by confidence (×1000):

Protein A	Protein B	Partner UniProt	Score
FUT4	SELE	P16111	979
FUT4	FUT1	P19526	899
FUT4	CD19	P15391	889
FUT4	CD33	P20138	879
FUT4	TNFRSF8	P28908	872
FUT4	ANPEP	P15144	872
FUT4	CD7	P09564	865
FUT4	FUT2	Q10981	865
FUT4	CD34	P28906	861
FUT4	CD2	P06729	852
FUT4	FCGR3B	O75015	841
FUT4	FCGR3A	P08637	841
FUT4	MME	P08473	824
FUT4	CD5	P06127	819
FUT4	CD4	P01730	807

IntAct

14 interactions, top by confidence:

A	B	Type	Score
HLA-DPA1	TYW5	psi-mi:“MI:0914”(association)	0.530
AOC3	AOC2	psi-mi:“MI:0914”(association)	0.530
HLA-DPA1	GXYLT2	psi-mi:“MI:0914”(association)	0.350
AKAP10	HAX1	psi-mi:“MI:0914”(association)	0.350
BSCL2	TMEM223	psi-mi:“MI:0914”(association)	0.350
CTLA4	TMEM120B	psi-mi:“MI:0914”(association)	0.350
LRP3	TMEM131L	psi-mi:“MI:0914”(association)	0.350
PCDHGB5	FAM171A2	psi-mi:“MI:0914”(association)	0.350
HFE	PODXL	psi-mi:“MI:0914”(association)	0.350
SFTPC	CYTH3	psi-mi:“MI:0914”(association)	0.350
SLC14A2	SMCHD1	psi-mi:“MI:0914”(association)	0.350
SLC30A10	GOLIM4	psi-mi:“MI:0914”(association)	0.350
SLC5A7	FUT4	psi-mi:“MI:0914”(association)	0.350

BioGRID (14): FUT4 (Affinity Capture-MS), FUT4 (Affinity Capture-MS), FUT4 (Affinity Capture-MS), FUT4 (Affinity Capture-MS), FUT4 (Affinity Capture-MS), FUT4 (Affinity Capture-MS), FUT4 (Affinity Capture-MS), FUT4 (Affinity Capture-MS), FUT4 (Affinity Capture-MS), FUT7 (Negative Genetic), FUT4 (Affinity Capture-MS), FUT4 (Affinity Capture-MS), FUT4 (Affinity Capture-MS), FUT4 (Affinity Capture-MS)

ESM2 similar proteins: A0A0U1RR11, A0A0U1RRI6, A6NCS6, A6NJG2, B0BN44, D3YXK1, E9PY61, E9Q0B3, F5H4A9, O00220, O00221, P09038, P0DPI3, P22083, P98077, Q08AU9, Q2M2W7, Q2M3V2, Q2TBI2, Q5F267, Q5FW56, Q5IS69, Q5R866, Q5T4W7, Q5TM52, Q5U4P2, Q5VTJ3, Q659K9, Q673H1, Q69ZB3, Q6AYE8, Q6IPT2, Q6PJ61, Q7RTU4, Q7TSX9, Q7YR31, Q80SU3, Q86SH2, Q86Y97, Q8NBR0

Diamond homologs: G3MZR2, G5EDR5, O19058, O88819, P21217, P22083, P51993, P56433, P56434, P83088, Q11126, Q11127, Q11128, Q11130, Q11131, Q62994, Q659K9, Q659L0, Q659L1, Q712G6, Q8HYJ3, Q8HYJ4, Q8HYJ5, Q8HYJ6, Q8HYJ7, Q8HZR2, Q8HZR3, Q99JB3, Q9FX97, Q9GKU6, Q9JIG1, Q9LJK1, Q9VUL9, Q9Y231, G5EE06, G5EEE1, Q5F2N3, Q5F2L1, Q5F2N2, Q68FV3

SIGNOR signaling

0 interactions.

Disease & clinical

Clinical variants and AI predictions

ClinVar

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

Classification	Count (floor)
Pathogenic	0
Likely pathogenic	1
Uncertain significance	76
Likely benign	12
Benign	2

Top pathogenic / likely-pathogenic (1)

Variant ID	HGVS	Classification
2582900	GRCh37/hg19 11q21(chr11:94126651-94331081)x1	Likely pathogenic

SpliceAI

259 predictions. Top by Δscore:

Variant	Effect	Δscore
11:94545815:GGGA:G	donor_gain	0.9800
11:94545816:G:GT	donor_gain	0.9800
11:94545817:GA:G	donor_gain	0.9800
11:94545819:G:GG	donor_gain	0.9800
11:94544017:G:T	donor_gain	0.9700
11:94545816:GGA:G	donor_gain	0.9700
11:94545817:GAG:G	donor_gain	0.9700
11:94545755:G:T	donor_gain	0.9500
11:94545814:TGGGA:T	donor_gain	0.9400
11:94545815:GGGAG:G	donor_gain	0.9400
11:94545816:G:T	donor_gain	0.9400
11:94546141:T:A	acceptor_gain	0.9400
11:94544039:G:GT	donor_gain	0.9300
11:94545818:AG:A	donor_loss	0.8600
11:94545819:G:C	donor_loss	0.8600
11:94545820:T:A	donor_loss	0.8600
11:94544044:C:T	donor_gain	0.8500
11:94545821:AA:A	donor_loss	0.8500
11:94545822:AGTT:A	donor_loss	0.8500
11:94545823:G:C	donor_loss	0.8500
11:94546715:T:A	acceptor_gain	0.8500
11:94548261:G:GC	acceptor_gain	0.8500
11:94544016:G:GT	donor_gain	0.8400
11:94544061:G:GG	donor_gain	0.8400
11:94545742:G:GT	donor_gain	0.8200
11:94543991:TCTC:T	donor_gain	0.8100
11:94544040:A:T	donor_gain	0.7700
11:94546558:T:G	acceptor_gain	0.7400
11:94546145:CCA:C	acceptor_gain	0.7300
11:94544060:A:AG	donor_gain	0.7100

AlphaMissense

3358 scored. Top likely-pathogenic:

Variant	Protein change	am_pathogenicity
11:94545021:C:A	N296K	0.996
11:94545021:C:G	N296K	0.996
11:94545358:T:C	F409L	0.996
11:94545359:T:C	F409S	0.996
11:94545360:C:A	F409L	0.996
11:94545360:C:G	F409L	0.996
11:94545374:A:T	E414V	0.996
11:94545412:T:A	W427R	0.996
11:94545412:T:C	W427R	0.996
11:94545009:G:C	W292C	0.995
11:94545009:G:T	W292C	0.995
11:94545208:T:A	W359R	0.995
11:94545208:T:C	W359R	0.995
11:94545503:T:A	I457N	0.995
11:94544843:T:C	F237S	0.994
11:94545378:C:A	N415K	0.994
11:94545378:C:G	N415K	0.994
11:94545404:A:T	E424V	0.994
11:94545499:T:C	F456L	0.994
11:94545501:C:A	F456L	0.994
11:94545501:C:G	F456L	0.994
11:94545548:T:C	L472P	0.994
11:94545007:T:A	W292R	0.993
11:94545007:T:C	W292R	0.993
11:94545597:G:C	W488C	0.993
11:94545597:G:T	W488C	0.993
11:94545646:T:A	C505S	0.993
11:94545647:G:C	C505S	0.993
11:94545656:G:A	C508Y	0.993
11:94545013:T:A	W294R	0.992

dbSNP variants (sampled 300 via entrez): RS1000023336 (11:94542140 T>A,C), RS1000694446 (11:94544104 G>A), RS1000879031 (11:94546404 A>C), RS1000907456 (11:94544314 G>A), RS1001172907 (11:94542803 A>G), RS1001205539 (11:94542670 A>C), RS1002126221 (11:94544460 C>T), RS1002422029 (11:94547239 T>A), RS1002759571 (11:94547188 C>T), RS1002875971 (11:94543870 C>A,T), RS1003221646 (11:94547474 T>G), RS1003802067 (11:94545940 C>T), RS1003869475 (11:94544501 T>A,C), RS1004059336 (11:94544078 C>A,G), RS1004133708 (11:94548519 G>A)

Disease associations

OMIM: gene MIM:104230 | disease phenotypes:

GenCC curated gene-disease

Mondo (0):

Orphanet (0):

HPO phenotypes

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

GWAS associations

5 associations (top):

Study	Trait	p-value
GCST001518_1	Response to angiotensin II receptor blocker therapy (opposite direction w/ diuretic therapy)	4.000000e-06
GCST001520_9	Response to angiotensin II receptor blocker therapy	9.000000e-07
GCST004746_25	Small cell lung carcinoma	8.000000e-06
GCST009391_1127	Metabolite levels	3.000000e-06
GCST009391_1942	Metabolite levels	8.000000e-06

EFO canonical traits (2, from GWAS)

EFO ID	Trait name
EFO:0010456	allantoin measurement
EFO:0009766	asparagine measurement

Drugs & pharmacology

Drug and pharmacology data

Is drug target: yes

ChEMBL targets (1): CHEMBL4996 (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: 103,233 (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
CHEMBL288114	GALLIC ACID	2	103,233

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

ChEMBL bioactivities

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

pChembl	Type	Value	Unit	Molecule
5.75	IC50	1800	nM	CHEMBL338988

CTD chemical–gene interactions

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

Chemical	Actions (top 5)	PubMed papers
Valproic Acid	increases methylation, affects cotreatment, increases expression, affects expression	8
Estradiol	affects binding, increases expression, affects cotreatment	3
entinostat	increases expression, affects cotreatment	2
Panobinostat	affects cotreatment, increases expression	2
Formaldehyde	decreases expression	2
Tetrachlorodibenzodioxin	decreases reaction, increases expression	2
Tobacco Smoke Pollution	decreases expression	2
Tretinoin	increases expression	2
aminomethylphosphonic acid (AMPA)	decreases expression	1
bisphenol A	decreases expression, decreases reaction	1
lead acetate	affects cotreatment, decreases expression	1
trichostatin A	increases expression	1
zinc protoporphyrin	affects cotreatment, decreases expression	1
sodium arsenite	increases expression	1
oligomycin A	increases expression	1
2,3,4,7,8-pentachlorodibenzofuran	increases expression	1
endotoxin, Escherichia coli	decreases reaction, decreases expression	1
tamibarotene	increases expression	1
di-n-butylphosphoric acid	affects expression	1
cagA protein, Helicobacter pylori	increases expression, decreases reaction	1
ginsenoside Rg3	decreases reaction, increases expression, decreases expression	1
4-(5-benzo(1,3)dioxol-5-yl-4-pyridin-2-yl-1H-imidazol-2-yl)benzamide	affects cotreatment, increases expression	1
abrine	decreases expression	1
dorsomorphin	affects cotreatment, increases expression	1
N2-(1H-indazole-5-yl)-N6-methyl-3-nitropyridine-2,6-diamine	decreases reaction, increases expression	1
Air Pollutants	increases abundance, increases expression	1
Ethanol	increases expression, decreases reaction	1
Arsenic	affects methylation	1
Benzo(a)pyrene	increases methylation	1
Calcitriol	decreases expression	1

ChEMBL screening assays

6 unique, capped per target: 6 binding

Representative assays (with source publication via chembl_document):

Assay ID	Type	Description	Source paper
CHEMBL3598451	Binding	Inhibition of FucT-4 (unknown origin)	Beyond substrate analogues: new inhibitor chemotypes for glycosyltransferases — Medchemcomm

Clinical trials (associated diseases)

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