From "Physical Barrier" to "Signaling Hub": How Mucin MUC1 Drives Tumor Progression

1. Structural Background and Physiological Functions of MUC1

The mucin family comprises several members, including transmembrane types (such as MUC1, MUC4, MUC16), secreted types, and soluble mucins. Among these, MUC1 is the best-characterized transmembrane mucin, localized to the apical surface of epithelial cells ^[20], where it participates in various physiological processes including epithelial protection, repair, cell adhesion, and immune response.

Structurally, MUC1 is a heterodimer composed of a non-covalently linked N-terminal (MUC1-N) and C-terminal (MUC1-C) subunit ^[21]. MUC1-N is extracellular and contains a variable number of 20-amino acid tandem repeats (VNTR), a region rich in O-glycosylation sites. The highly glycosylated MUC1-N extends outward from the cell surface, forming a physical barrier on the mucosal surface that protects epithelial cells from external damage. MUC1-C consists of an extracellular domain, a transmembrane domain, and a cytoplasmic tail, serving as the key component for signal transduction. Mediated by the SEA structural domain, MUC1 undergoes autoproteolytic cleavage; upon receiving inflammatory signals (such as IFN-γ, TNF-α), MUC1-N is released, subsequently activating MUC1-C and initiating downstream signaling pathways.

In normal tissues, MUC1 exhibits strict polarized expression, localized exclusively to the apical cell membrane, maintaining epithelial barrier integrity and preventing aberrant signal activation. Under infectious or inflammatory conditions, MUC1 can bind to viruses (e.g., limiting influenza A infection) through sialic acid structures at the termini of its glycan chains, while simultaneously modulating inflammatory intensity to prevent immune damage. Thus, under physiological conditions, MUC1 possesses both barrier functions and immunomodulatory roles.

However, in the context of chronic inflammation, hypoxia, or genetic susceptibility, MUC1 loses its polarized expression pattern and becomes aberrantly overexpressed across the entire cell surface, transforming from a "physical defender" into a "signaling hub molecule" and becoming a key driver in the development of various malignant diseases.

Fig. Schematic representation of the MUC1 structure both in physiological conditions (Normal MUC1) and in cancer (tMUC1) ^[27]

2. Genetic Susceptibility and Epigenetic Regulation

The abnormal expression of MUC1 is closely related to genetic background. Genome-wide association studies have identified the 1q22 region as a significant susceptibility locus for gastric cancer, with the functional single nucleotide polymorphism rs4072037 receiving widespread attention ^[1,2]. Multiple meta-analyses have shown that the G allele significantly reduces the risk of gastric cancer, with its effect size (OR) typically ranging between 0.70 and 0.79 ^[3-5]. In East Asian populations, the population attributable risk at this locus is relatively high ^[6,7], indicating its epidemiological significance. This polymorphism not only influences the overall risk of gastric cancer but is also closely associated with the prognosis of diffuse-type gastric cancer ^[10] and demonstrates relevance even in the precancerous lesion stage ^[11].

At the epigenetic level, the methylation status of the MUC1 promoter is a key factor regulating its expression. In pancreatic ductal adenocarcinoma, promoter hypomethylation is significantly associated with high mRNA expression and indicates poor prognosis ^[15]. Similar epigenetic abnormalities have also been validated in lung adenocarcinoma ^[16]. Additionally, functional variants such as rs4072037 and rs12411216 are associated with metabolic and renal function indicators ^[17], reflecting the multi-system pleiotropic effects of MUC1.

Overall, the genetic and epigenetic background of MUC1 provides a molecular basis for its abnormal expression, but its pathogenic effects still need to be realized through downstream signaling networks.

3. Abnormal Glycosylation and Key Signaling Pathway Mechanisms

The functional transformation of MUC1 in tumors primarily relies on abnormal glycosylation and activation of the cytoplasmic tail signaling. In tumor cells, aberrant expression of C1GALT1 and Cosmc leads to the exposure of Tn and sTn antigens ^[19]. This truncation of glycan chains alters the conformation of MUC1 and exposes new ligand-binding sites.

Galectin-3 can bind to abnormally glycosylated MUC1, inducing MUC1-C phosphorylation and activating the ERK1/2 and Akt signaling pathways ^[21]. Simultaneously, YBX1 enhances MUC1 expression at the transcriptional level ^[20]. In terms of spatial regulation, ADAM10/17 and γ-secretase mediate the hydrolysis and release of MUC1-C, allowing it to enter the nucleus ^[26], where it forms a complex with β-Catenin to regulate the expression of EMT and stemness-related genes ^[9]. Furthermore, MUC1-C enhances PD-L1 expression and the formation of an immunosuppressive environment through the JAK/STAT3 and NF-κB signaling pathways ^[18].

Therefore, the transformation of MUC1 from a membrane-bound structural molecule into a nuclear transcriptional co-regulator is a critical step in driving malignant transformation.

4. The Role and Clinical Significance of MUC1 in Diseases

Abnormal expression of MUC1 is present throughout the entire process of tumor initiation, progression, and immune evasion, and it holds clear pathological and clinical significance in various solid tumors.

In gastric cancer, the rs4072037 polymorphism not only influences overall disease risk but is also closely associated with recurrence and disease-related mortality in diffuse-type gastric cancer ^[10]. Studies indicate that abnormal MUC1 expression emerges as early as the stages of severe intestinal metaplasia and atypical hyperplasia ^[11], suggesting its early driving role in the transition from inflammation to carcinogenesis. The synergistic effect with Helicobacter pylori infection further reinforces this continuum model ^{[12, 13]}. The combination of genetic susceptibility and chronic infection positions MUC1 as a crucial molecular bridge linking inflammation to the development of gastric cancer.

In pancreatic ductal adenocarcinoma (PDAC), MUC1 is identified as one of the core genes in the protein interaction network ^[22]. Histopathological studies have shown that its expression in cancerous tissues is significantly higher than in normal ductal tissues ^[25]. The detection method based on pancreatic juice DNA methylation can distinguish PDAC from IPMN with high specificity and sensitivity ^[23]. It demonstrates its potential application value in early differential diagnosis. Notably, MUC1 is not only a diagnostic marker, but its cytoplasmic tail region maintains the survival advantage of tumor cells by continuously activating the PI3K/Akt signaling pathway, indicating its important role in therapeutic targeting.

In breast cancer, overexpression of MUC1 has been confirmed as an independent predictor of poor prognosis.[8]Its aberrant glycosylation enhances interactions with E-selectin and ICAM-1 ^[24], promoting the "rolling" and adhesion of circulating tumor cells on vascular endothelium, thereby accelerating distant metastasis. Additionally, MUC1 constructs an immunosuppressive microenvironment by enhancing PD-L1 expression and NF-κB activation, enabling tumor cells to evade immune clearance.

In non-small cell lung cancer, EIF4G1-driven high expression of MUC1 is significantly correlated with PD-L1 levels.[14]and enhances immune escape capability through the JAK/STAT3 pathway. Its high enrichment in exosomes ^[40] further suggests its potential application in liquid biopsy.

In addition to solid tumors, functional variants of MUC1 are also associated with cardiovascular metabolic indicators and renal function parameters ^[17], suggesting its potential role in systemic inflammation and metabolic regulation. Although its mechanisms in non-neoplastic diseases are not yet fully elucidated, its pleiotropic characteristics indicate that it may be involved in chronic inflammation and tissue fibrosis processes.

Overall, the role of MUC1 in various diseases exhibits common characteristics: loss of expression polarity, abnormal glycosylation, enhanced nuclear translocation, and dysregulation of immune modulation. These mechanisms constitute its "shared pathogenic framework" across multiple cancer types.

5. Latest Research Progress on MUC1-Targeted Drugs

Currently, the development of MUC1-targeted drugs has entered a diversified clinical stage dominated by immunotherapy and precision conjugation technologies. In terms of drug types, cell therapies (CAR-T) and antibody-drug conjugates (ADCs) constitute the dual main lines of research and development, with the most advanced candidates primarily in Phase II clinical trials. The indications are mainly focused on advanced solid tumors such as prostate cancer, pancreatic cancer, and non-small cell lung cancer. Notably, CAR-T therapy development is particularly active, with domestic biotechnology companies such as Guangzhou Anjie, Shenzhen Institute of Immunogene Therapy, Shanghai Sidansai, and Guangdong Zhaotai having pipelines entering clinical trials, demonstrating China's deep involvement in the field of MUC1-targeted cell therapy. Some of the pipelines under development are summarized in the following table:

Drug	Mechanism of action	Drug Type	Indications Under Research	Research Institutions	Highest R&D Stage
Anti-MUC1 CAR T cell(Guangzhou Anjie Biomedical Technology Co., Ltd.)	MUC1 Modulator \| T Lymphocyte Substitute	CAR-T	Prostate cancer	Guangzhou Anjie Biomedical Technology Co., Ltd.	Phase 2 Clinical Trial
GO-2032c	MUC1 inhibitor	Synthetic Polypeptide	Acute Myeloid Leukemia	Dana-Farber Cancer Institute, Inc. \| Genus Oncology LLC	Phase 2 Clinical Trial
Anti-MUC1 monoclonal antibody(AltaRex Corp.)	MUC1 inhibitor	Monoclonal antibody	Pancreatic cancer	HYULIM A-TECH Co., Ltd. \| ViRexx Medical Corp. \| Shenzhen Hepalink Pharmaceutical Group Co., Ltd.	Phase 2 Clinical Trial
Cancer vaccine(Therion Biologics Corp.)	CEACAM5 antagonist \| MUC1 inhibitor \| Immunostimulant	Recombinant Vector Vaccine \| Therapeutic Vaccine	Small Intestine Cancer \| Colorectal Cancer \| Non-Small Cell Lung Cancer \| Prostate CancerWait	National Cancer Institute \| Bavarian Nordic, Inc.	Phase 2 Clinical Trial
Adenovirus 5 CEA/MUC1/Brachyury Vaccine(National Cancer Institute)	CEA Inhibitor \| MUC1 Inhibitor \| TBXT Inhibitor \| Immunostimulant	Recombinant Vector Vaccine \| Therapeutic Vaccine	Hereditary Nonpolyposis Colorectal Cancer	National Cancer Institute	Phase 2 Clinical Trial
CV-9202	5T4 inhibitor \| MAGEC1 antagonist \| MAGEC2 antagonist \| MUC1 inhibitor \| NY-ESO-1 inhibitor \| survivin inhibitor	Therapeutic Vaccine \| mRNA Vaccine	Non-Small Cell Lung Cancer	Ludwig Institute for Cancer Research Ltd. \| C. H. Boehringer Sohn AG & Co. KG	Phase 2 Clinical Trial
Gemcitabine Liposome	DNA Polymerase Inhibitors \| MUC1 Inhibitors	Small Molecule Chemical Drugs \| Liposomal Drugs	Advanced Non-Small Cell Lung Cancer \| Advanced Malignant Solid Tumors \| Advanced Urothelial CarcinomaWait	FUJIFILM Pharmaceuticals USA, Inc. \| FUJIFILM Holdings America Corp.	Phase 2 Clinical Trial
Cancer vaccine-MUC 1(Vaxil Biotherapeutics)	MUC1 inhibitor	Therapeutic vaccine	Breast Cancer \| Multiple Myeloma	Vaxil Biotherapeutics Ltd. \| Hadassah Foundation	Phase 2 Clinical Trial
MUC1-Poly-ICLC	MUC1 modulator	Therapeutic vaccine	Non-Small Cell Lung Cancer	University of Pittsburgh Medical Center	Phase 2 Clinical Trial
Monoclonal antibody HMFG1(Cancer Research UK)	MUC1 Inhibitor \| Immunostimulant	Monoclonal antibody	Locally Advanced Breast Cancer \| Metastatic Breast Cancer	Antisoma Research Ltd.	Phase 2 Clinical Trial
Tipapkinogene sovacivec	E7 Inhibitor \| HPV E6 Inhibitor \| MUC1 Modulator	Recombinant Vector Vaccines \| Universal Antigen Vaccines	HPV-related anal squamous cell carcinoma \| Human papillomavirus-associated cervical cancerWait	Transgene SA	Phase 1/2 Clinical
BM7PE	MUC1 inhibitor	Antibody-drug conjugate	Metastatic Colorectal Cancer \| Metastatic Tumor	Oslo universitetssykehus HF	Phase 1/2 Clinical
Mucin gene (MUC1) transfected dendritic cells vaccine(Free University of Berlin)	MUC1 modulator	Therapeutic vaccine	Advanced Breast Cancer \| Pancreatic Cancer	Freie Universität Berlin	Phase 1/2 Clinical
DS-3939	MUC1 Modulator \| TOP1 Inhibitor	ADC	Advanced Breast Cancer \| Recurrent Breast Cancer \| Ductal Carcinoma of the Breast \| Colorectal CancerWait	Daiichi Sankyo, Inc. \| Daiichi Sankyo Co., Ltd. \| Glycotope GmbH	Phase 1/2 Clinical
M0324	CD40 Agonist \| MUC1 Inhibitor	Bispecific antibody	Advanced Malignant Solid Tumor \| Tumor	Merck & Co., Inc. \| EMD Serono Research & Development Institute, Inc. \| Merck KGaA \| EMD Serono, Inc.	Phase 1 Clinical Trial
Recombinant Humanized Anti-Muc1 Monoclonal Antibody-Tub201 Conjugate (Hangzhou DAC Biotechnology)	MUC1 inhibitor	ADC	Advanced Malignant Solid Tumors \| Colorectal Cancer \| Pancreatic Cancer	Hangzhou Duoxi Biotechnology Co., Ltd.	Phase 1 Clinical Trial
huMNC2-CAR44 T cells(Minerva Biotechnologies)	MUC1 modulator	Autologous CAR-T	Advanced Breast Cancer \| HER2-Positive Metastatic Breast Cancer	Minerva Biotechnologies Corp.	Phase 1 Clinical Trial
huMNC2-CAR22(Minerva Biotechnologies)	MUC1 inhibitor	Autologous CAR-T	Advanced Breast Cancer \| HER2-Positive Metastatic Breast Cancer	Minerva Biotechnologies Corp.	Phase 1 Clinical Trial
SBO-154	MUC1 inhibitor	ADC	Advanced Malignant Solid Tumors	Sun Pharma Advanced Research Co. Ltd. \| Sun Pharma Advanced Research Company Ltd.	Phase 1 Clinical Trial
MUC1-TILs(Guangdong Zhaotai InVivo Biomedicine)	HPK1 Inhibitor \| MUC1 Inhibitor \| PD-1 Inhibitor \| Immune Cytotoxicity \| Lymphocyte Substitute	Tumor-infiltrating lymphocytes	Advanced Malignant Solid Tumors \| Brain Cancer \| Breast Cancer \| Colorectal Cancer \| Liver Cancer \| Lung Cancer	Guangdong Zhaotai In Vivo Biomedical Technology Co., Ltd.	Phase 1 Clinical Trial
DAA/TAA Peptide Vaccine(University of Pittsburgh)	MUC1 modulator	Synthetic Peptide Vaccine \| Therapeutic Vaccine	Ductal carcinoma	University of Pittsburgh	Phase 1 Clinical Trial
Anti-MUC1 CAR-T (Shenzhen Geno-Immune Medical)	MUC1 inhibitor	Autologous CAR-T	Acute Lymphoblastic Leukemia \| Refractory Anemia with Excess Blasts \| Refractory Leukemia	Shenzhen Institute of Immunotherapy and Gene Therapy	Phase 1 Clinical Trial
Anti MUC 1 chimeric antigen receptor T cell therapy(Innovative Cellular Therapeutics Co., Ltd.)	MUC1 Inhibitor \| Immune Cytotoxicity \| T Lymphocyte Substitutes	CAR-T	Breast Cancer \| Lung Cancer \| Pancreatic Cancer	Shanghai Sidansai Biotechnology Co., Ltd.	Phase 1 Clinical Trial
MUC1-CAR-T Cells(Guangdong Zhaotai InVivo Biomedicine)	HPK1 Inhibitor \| MUC1 Inhibitor \| PD-1 Inhibitor \| Immune Cytotoxicity \| T Lymphocyte Substitute	Autologous CAR-T	Advanced Cancer \| Lung Cancer	Guangdong Zhaotai In Vivo Biomedical Technology Co., Ltd.	Phase 1 Clinical Trial
MUC1-CAR-TILs(Guangdong Zhaotai InVivo Biomedicine)	HPK1 Inhibitor \| MUC1 Inhibitor \| PD-1 Inhibitor \| Immune Cytotoxicity \| Lymphocyte Substitute	Autologous CAR-T	Advanced Malignant Solid Tumors \| Brain Cancer \| Breast Cancer \| Colorectal Cancer \| Liver Cancer \| Lung Cancer	Guangdong Zhaotai In Vivo Biomedical Technology Co., Ltd.	Phase 1 Clinical Trial
DM-002 (Xadcera Biopharma)	HER3 Antagonist \| MUC1 Inhibitor \| TOP1 Inhibitor	ADC	Colorectal Cancer \| Endometrial Cancer \| Ovarian Cancer \| Prostate CancerWait	Sidao Pharmaceutical Technology (Suzhou) Co., Ltd. \| Doma Pharmaceutical Technology (Suzhou) Co., Ltd. \| Biocytogen (Beijing) Pharmaceutical Technology Co., Ltd.	Phase 1 Clinical Trial
Autologous MUC1 activated T cells(Mayo Clinic)	MUC1 modulator	T-cell therapy	Ovarian Carcinosarcoma \| Platinum-Resistant Ovarian Cancer \| Platinum-Resistant Primary Peritoneal CarcinomaWait	Mayo Clinic	Phase 1 Clinical Trial

(Data as of February 27, 2026, sourced from Synapse)

6. MUC1-Related Research Tools

Regarding the broad research prospects of MUC1 from basic mechanisms to clinical translation,CUSABIO offering a comprehensive suite of tools to meet the research needs mentioned above, including recombinant proteins, antibodies, and ELISA kits, to support your studies in genetic regulation, signaling pathways, drug screening, and liquid biopsy.

● MUC1 Recombinant Protein

Recombinant Human Mucin-1 (MUC1), partial; Code: CSB-MP015215HU2

Recombinant Human Mucin-1(MUC1),partial,Biotinylated;
CSB-MP015215HU3-B

Recombinant Mouse Mucin-1 (Muc1), partial;
CSB-EP312543MO

● MUC1 Antibody

MUC1 Recombinant Monoclonal Antibody; CSB-RA049212A0HU

● MUC1 ELISA Kit

Human mammary carcinoma marker CA15-3 (CA15-3) ELISA kit; CSB-E04772h

► View all MUC1-related products

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