Affinage

MAN2A1

Alpha-mannosidase 2 · UniProt Q16706

Length
1144 aa
Mass
131.1 kDa
Annotated
2026-06-10
54 papers in source corpus 15 papers cited in narrative 15 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 5/5 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

MAN2A1 encodes Golgi alpha-mannosidase II, a medial-Golgi-resident enzyme whose catalytic activity is required to convert hybrid-type N-glycans into the precursors of complex-type N-glycans; loss of its activity causes accumulation of hybrid-type structures and failure of complex N-glycan maturation (PMID:19465480, PMID:33878161). Deficiency in N-glycan processing from reduced MAN2A1 underlies a form of congenital dyserythropoietic anemia type II (HEMPAS) (PMID:2217175). Correct medial-Golgi positioning of MAN2A1 is maintained by retrograde trafficking machinery, depending on the COG tethering complex (PMID:21421995) and on Arf1-GTPase-driven COPI vesicle formation (PMID:27748959), as well as on SAC1-controlled Golgi PI(4)P distribution (PMID:20573065). Through its control of N-glycan maturation, MAN2A1 governs the cell-surface trafficking and glycosylation of client glycoproteins (PMID:31455607) and shapes physiological and pathological outcomes: intestinal-epithelial MAN2A1 promotes inflammation by supporting neutrophil-attracting chemokine expression (PMID:29343654), and MAN2A1-dependent N-glycan maturation on tumor cells suppresses cytotoxic T-cell killing and anti-PD-L1 responses, an effect reversed by the inhibitor swainsonine (PMID:32723834, PMID:25797317). Independently, a MAN2A1-FER chromosomal fusion exploits the MAN2A1 Golgi-targeting signal to deliver a constitutively active tyrosine kinase to the Golgi, where it phosphorylates the extracellular domains of EGFR, PDGFRA, MET, AXL, and N-cadherin to drive BRAF/MEK/AKT oncogenic signaling (PMID:28245430, PMID:39082961).

Mechanistic history

Synthesis pass · year-by-year structured walk · 11 steps
  1. 1990 High

    Established that MAN2A1 is required for N-linked oligosaccharide maturation in vivo and that its deficiency causes human disease, framing the gene as a glycan-processing enzyme with physiological consequence.

    Evidence Cell-surface labeling, FAB mass spectrometry, enzyme activity assay, and Northern blot in a HEMPAS patient

    PMID:2217175

    Open questions at the time
    • Did not define the precise catalytic step or substrate specificity
    • Causal mutation in MAN2A1 not molecularly resolved
  2. 2009 High

    Pinpointed the specific catalytic role of MAN2A1 as the enzyme converting hybrid- to complex-type N-glycans, using active-site mutation plus complementation rescue.

    Evidence Sequencing of Lec36 alleles, nano-ESI-MS glycan analysis, and wild-type vs. mutant complementation

    PMID:19465480

    Open questions at the time
    • Did not address redundancy with the MAN2A2 isozyme
    • No structural basis for substrate selectivity
  3. 2010 Medium

    Addressed how MAN2A1 is retained in the medial-Golgi by linking its localization to phosphoinositide control.

    Evidence SAC1 RNAi knockdown with immunofluorescence, ultrastructure, and glycosylation readouts

    PMID:20573065

    Open questions at the time
    • Single lab
    • Mechanism connecting PI(4)P levels to enzyme retention not resolved
  4. 2011 High

    Showed that MAN2A1 Golgi residence requires active retrograde transport via the COG tethering complex, distinguishing localization maintenance from biosynthetic delivery.

    Evidence COG subunit siRNA depletion with BFA/Sar1-DN redistribution assays and glycan analysis

    PMID:21421995

    Open questions at the time
    • Direct physical interaction between MAN2A1 and COG not demonstrated
    • Cargo recognition mechanism unknown
  5. 2016 Medium

    Extended the trafficking model by demonstrating that medial-Golgi positioning depends on Arf1 GTPase activity and COPI vesicle formation.

    Evidence 3D-SIM, beta-COP siRNA, dominant-negative Arf1(T31N), and electron microscopy under ethanol perturbation

    PMID:27748959

    Open questions at the time
    • Single lab
    • Did not establish whether COPI directly packages MAN2A1
  6. 2017 High

    Revealed an oncogenic gain-of-function in which the MAN2A1 Golgi-targeting signal mislocalizes a hyperactive FER kinase that phosphorylates EGFR to drive proliferation and tumorigenesis.

    Evidence Golgi fractionation, kinase assays, phospho-analysis, KO/OE in cell lines, and xenografts

    PMID:28245430

    Open questions at the time
    • Mechanism of extracellular-domain phosphorylation by a Golgi-luminal kinase not fully explained
    • Prevalence of the fusion across tumor types not defined here
  7. 2017 Medium

    Demonstrated that the MAN2A1-FER genomic breakpoint is transcriptionally active and can be exploited as a tumor-selective therapeutic target.

    Evidence CRISPR-Cas9 nickase HDR insertion of HSV1-tk at the breakpoint with ganciclovir killing and xenografts

    PMID:28459452

    Open questions at the time
    • Single lab
    • Delivery efficiency to clinical tumors untested
  8. 2018 High

    Connected MAN2A1 N-glycan processing to inflammation, showing tissue-intrinsic enzyme activity drives chemokine-mediated neutrophil recruitment.

    Evidence IEC-specific conditional knockout mice in DSS colitis with histology, flow cytometry, and expression analysis

    PMID:29343654

    Open questions at the time
    • Specific glycoproteins mediating chemokine induction not identified
    • Role of alpha-MIIx compensation in other tissues unresolved
  9. 2019 Medium

    Provided a specific client example by showing MAN2A1-dependent glycosylation is needed for plasma-membrane trafficking of the sodium iodide symporter.

    Evidence MAN2A1 RNAi with immunofluorescence and glycosylation profiling in MCF-7 cells

    PMID:31455607

    Open questions at the time
    • Single substrate, single lab
    • Glycan site responsible for NIS mistrafficking not mapped
  10. 2020 High

    Established MAN2A1-driven N-glycan maturation as a mechanism of tumor immune evasion and a druggable target with checkpoint inhibitors.

    Evidence CRISPR KO, T-cell co-culture killing, syngeneic tumor models, and swainsonine inhibition with anti-PD-L1

    PMID:32723834

    Open questions at the time
    • Specific glycoproteins shielding tumor cells from T cells not identified
    • Swainsonine specificity beyond MAN2A1 not delimited
  11. 2024 Medium

    Broadened the MAN2A1-FER oncogenic mechanism to multiple receptor substrates and validated a phospho-epitope-directed therapeutic antibody.

    Evidence Phospho-analysis of PDGFRA/MET/AXL/N-cadherin, monoclonal antibody specificity, and ADC xenografts with isogenic KO controls

    PMID:39082961

    Open questions at the time
    • Single lab
    • Structural basis of ectopic extracellular-domain phosphorylation unresolved

Open questions

Synthesis pass · forward-looking unresolved questions
  • How MAN2A1-FER, a Golgi-localized kinase, gains access to and phosphorylates the extracellular domains of cell-surface receptors remains mechanistically unexplained.
  • Topological mechanism of extracellular-domain phosphorylation unknown
  • Structural model of the fusion protein lacking

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0016787 hydrolase activity 3 GO:0140098 catalytic activity, acting on RNA 2
Localization
GO:0005794 Golgi apparatus 4
Pathway
R-HSA-392499 Metabolism of proteins 3 R-HSA-5653656 Vesicle-mediated transport 2

Evidence

Reading pass · 15 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1990 Alpha-mannosidase II (MAN2A1) deficiency in a HEMPAS patient caused incomplete processing of asparagine-linked oligosaccharides in erythrocyte membranes, as demonstrated by cell-surface labeling, FAB mass spectrometry of glycopeptides, and enzyme activity assays showing decreased alpha-ManII activity; reduced MAN2A1 mRNA expression (<10% of normal) was detected by Northern blot, establishing that MAN2A1 is required for the maturation of N-linked oligosaccharides and that its defect underlies at least one form of congenital dyserythropoietic anemia type II (HEMPAS). Cell-surface labeling, FAB mass spectrometry, enzyme activity assay, Northern blot Proceedings of the National Academy of Sciences of the United States of America High 2217175
2009 Mutations in both alleles of MAN2A1 in the HEK293T-derived Lec36 cell line result in accumulation of hybrid-type N-glycans instead of complex-type N-glycans; one allele carried an active-site point mutation and the other an in-frame 12-nucleotide deletion. Expression of wild-type but not mutant MAN2A1 in Lec36 cells restored processing of a reporter glycoprotein to complex-type glycosylation, demonstrating that MAN2A1 catalytic activity is required for conversion of hybrid- to complex-type N-glycans. Nucleotide sequencing, mass spectrometry (negative ion nano-ESI-MS, CID), complementation assay with wild-type vs. mutant MAN2A1 expression The Journal of biological chemistry High 19465480
2017 The MAN2A1-FER fusion protein retains the MAN2A1 signal peptide for Golgi localization and translocates from the cytoplasm to the Golgi in cancer cell lines. The fusion protein has tyrosine kinase activity approximately 4-fold higher than wild-type FER and phosphorylates EGFR at tyrosine 88 in its N-terminus, activating downstream BRAF, MEK, and AKT signaling, thereby driving proliferation, invasion, and tumor formation. Tagged protein expression and Golgi fractionation, kinase activity assay, phosphorylation analysis, KO/OE in multiple cell lines, xenograft mouse model Gastroenterology High 28245430
2011 MAN2A1 is severely mislocalized upon siRNA-mediated depletion of COG complex subunits, and COG-depleted cells are deficient in Brefeldin A- and Sar1 DN-induced redistribution of MAN2A1 to the ER, establishing that the conserved oligomeric Golgi (COG) complex is required for retrograde trafficking and maintenance of MAN2A1 at its correct Golgi location. siRNA knockdown, immunofluorescence localization, BFA redistribution assay, lectin staining, MALDI-TOF glycan analysis Glycobiology High 21421995
2010 RNAi-mediated knockdown of the lipid phosphatase SAC1 caused mislocalization of MAN2A1 (mannosidase-II) from its normal medial-Golgi position to aberrant intracellular structures and to the cell surface, accompanied by specific defects in N- and O-linked glycosylation, establishing that SAC1-controlled PI(4)P distribution is required for correct MAN2A1 Golgi localization. RNAi knockdown, immunofluorescence, ultrastructural analysis, glycosylation assay Traffic (Copenhagen, Denmark) Medium 20573065
2009 Raising Golgi luminal pH with chloroquine did not induce mislocalization of MAN2A1 (MANII) at moderate pH increases (0.2 units) that were sufficient to mislocalize the alpha(2,3)-sialyltransferase ST3, indicating that MAN2A1 Golgi localization is relatively insensitive to moderate pH elevations compared to late-acting glycosyltransferases. Chloroquine pH manipulation, immunofluorescence localization of Golgi enzymes, glycosylation reporter assay Journal of cellular physiology Medium 19277980
2016 Ethanol treatment induces medial-to-cis relocation of MAN2A1 (Man-II) within the Golgi, an effect mimicked by silencing beta-COP or expressing GDP-bound mutant Arf1(T31N), demonstrating that MAN2A1 correct medial-Golgi positioning depends on functional Arf1 GTPase activity and intact COPI vesicle formation. 3D structured illumination microscopy, beta-COP siRNA, dominant-negative Arf1 expression, electron microscopy Alcoholism, clinical and experimental research Medium 27748959
2018 IEC-specific deletion of MAN2A1 (alpha-MII) in mice impaired N-glycan maturation specifically in intestinal epithelial cells (where the isozyme alpha-MIIx does not compensate), reduced neutrophil infiltration in colonic mucosa, and downregulated neutrophil-attracting chemokine gene expression, establishing that IEC-intrinsic MAN2A1 promotes intestinal inflammation by facilitating chemokine expression. Conditional (IEC-specific) knockout mouse, DSS-induced colitis model, histology, flow cytometry, gene expression analysis Cell structure and function High 29343654
2019 RNAi-mediated knockdown of MAN2A1 impaired correct plasma membrane trafficking of the sodium iodide symporter (NIS) in MCF-7 breast cancer cells, establishing that MAN2A1-dependent N-glycan processing is required for correct subcellular localization of NIS. RNAi knockdown, immunofluorescence, glycosylation RT-Profiler array Journal of cell science Medium 31455607
2020 Loss of Man2a1 in cancer cells increased their sensitivity to T-cell-mediated killing and enhanced response to anti-PD-L1 treatment with higher cytotoxic T-cell infiltration; pharmacologic inhibition of MAN2A1 with swainsonine synergized with anti-PD-L1 in syngeneic melanoma and lung cancer models, establishing that MAN2A1-dependent N-glycan maturation in cancer cells promotes immune evasion. CRISPR knockout, in vitro T-cell co-culture killing assay, in vivo syngeneic tumor growth assay, pharmacologic inhibition (swainsonine) Clinical cancer research High 32723834
2024 The MAN2A1-FER fusion protein ectopically phosphorylates the extracellular domains of PDGFRA, MET, AXL, and N-cadherin, activating their kinase activities and downstream signaling cascades. A monoclonal antibody targeting the ectopic phosphotyrosine epitope in the extracellular domain of PDGFRA deactivated growth signaling and induced cell death selectively in MAN2A1-FER-positive cancer cells, with no effect on MAN2A1-FER-knockout cells. Phosphorylation analysis, monoclonal antibody development and specificity assay, cell growth assay, xenograft mouse model with antibody-drug conjugate Hepatology communications Medium 39082961
2017 Cas9-mediated insertion of a suicide gene (HSV1-tk) at the genomic breakpoint of the MAN2A1-FER fusion caused breakpoint-dependent EGFP-tk expression and ganciclovir-mediated apoptosis selectively in cancer cells carrying the fusion, demonstrating that the MAN2A1-FER genomic rearrangement breakpoint is transcriptionally active and targetable. CRISPR-Cas9 nickase, homology-directed repair, adenoviral delivery, ganciclovir treatment, xenograft mouse model Nature biotechnology Medium 28459452
2021 Knockout of MAN2A1 and MAN2A2 together in HEK293 cells (M2D-KO) abolished production of complex-type N-glycans, resulting in proteins carrying predominantly hybrid-type N-glycans, establishing that MAN2A1 and MAN2A2 together are responsible for complex N-glycan maturation in this cell line. Gene knockout (MAN2A1 and MAN2A2 double KO), glycan structural analysis, recombinant protein expression Journal of biochemistry Medium 33878161
2015 Swainsonine treatment of BRL-3A cells significantly reduced alpha-mannosidase activity and decreased MAN2A1 mRNA and protein expression in a time- and dose-dependent manner, establishing that swainsonine acts as both an inhibitor of MAN2A1 enzymatic activity and a suppressor of MAN2A1 expression. Enzyme substrate activity assay, qPCR, Western blot Toxicon Medium 25797317
2014 Reduced expression of MAN2A1 (man2a1 gene) during TGFβ-induced epithelial-mesenchymal transition in HCV29 bladder epithelial cells contributed to decreased proportions of bi-, tri-, and tetra-antennary complex N-glycans and increased hybrid-type N-glycans, as confirmed by mass spectrometry, lectin microarray, and RT-PCR. GlycoV4 microarray, mass spectrometry, lectin microarray, RT-PCR Molecules (Basel, Switzerland) Medium 25470275

Source papers

Stage 0 corpus · 54 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2006 Modulation of therapeutic antibody effector functions by glycosylation engineering: influence of Golgi enzyme localization domain and co-expression of heterologous beta1, 4-N-acetylglucosaminyltransferase III and Golgi alpha-mannosidase II. Biotechnology and bioengineering 232 16435400
2009 Elevated Golgi pH impairs terminal N-glycosylation by inducing mislocalization of Golgi glycosyltransferases. Journal of cellular physiology 113 19277980
2017 Targeting genomic rearrangements in tumor cells through Cas9-mediated insertion of a suicide gene. Nature biotechnology 100 28459452
1990 Incomplete synthesis of N-glycans in congenital dyserythropoietic anemia type II caused by a defect in the gene encoding alpha-mannosidase II. Proceedings of the National Academy of Sciences of the United States of America 90 2217175
2011 Conserved oligomeric Golgi complex specifically regulates the maintenance of Golgi glycosylation machinery. Glycobiology 89 21421995
2010 Spatial regulation of Golgi phosphatidylinositol-4-phosphate is required for enzyme localization and glycosylation fidelity. Traffic (Copenhagen, Denmark) 60 20573065
2008 Association of polymorphisms in complement component C3 gene with susceptibility to systemic lupus erythematosus. Rheumatology (Oxford, England) 57 18174230
2020 Glycobiology and schizophrenia: a biological hypothesis emerging from genomic research. Molecular psychiatry 55 32377000
2020 Vesicular and uncoated Rab1-dependent cargo carriers facilitate ER to Golgi transport. Journal of cell science 52 32616562
2017 MAN2A1-FER Fusion Gene Is Expressed by Human Liver and Other Tumor Types and Has Oncogenic Activity in Mice. Gastroenterology 51 28245430
2020 Inhibition of MAN2A1 Enhances the Immune Response to Anti-PD-L1 in Human Tumors. Clinical cancer research : an official journal of the American Association for Cancer Research 47 32723834
1997 Immunocytochemical analysis of Uukuniemi virus budding compartments: role of the intermediate compartment and the Golgi stack in virus maturation. Journal of virology 47 8995638
2014 Alteration of N-glycans and expression of their related glycogenes in the epithelial-mesenchymal transition of HCV29 bladder epithelial cells. Molecules (Basel, Switzerland) 37 25470275
2008 Expression of rat beta(1,4)-N-acetylglucosaminyltransferase III in Nicotiana tabacum remodels the plant-specific N-glycosylation. Plant biotechnology journal 36 18778316
2009 A human embryonic kidney 293T cell line mutated at the Golgi alpha-mannosidase II locus. The Journal of biological chemistry 28 19465480
2019 Detection of fusion transcripts in the serum samples of patients with hepatocellular carcinoma. Oncotarget 27 31164957
2016 Study of Ethanol-Induced Golgi Disorganization Reveals the Potential Mechanism of Alcohol-Impaired N-Glycosylation. Alcoholism, clinical and experimental research 19 27748959
2002 Possible implication of Golgi-nucleating function for the centrosome. Biochemical and biophysical research communications 19 11855815
2021 Complex N-Glycans Are Important for Normal Fruit Ripening and Seed Development in Tomato. Frontiers in plant science 17 33767719
2017 Development of a pre-glycoengineered CHO-K1 host cell line for the expression of antibodies with enhanced Fc mediated effector function. mAbs 17 29173063
2014 Effect of swainsonine in Oxytropis kansuensis on Golgi α-mannosidase II expression in the brain tissues of Sprague-Dawley rats. Journal of agricultural and food chemistry 16 24741992
2021 Detection of fusion gene transcripts in the blood samples of prostate cancer patients. Scientific reports 14 34417538
2024 Alteration of the metabolite interconversion enzyme in sperm and Sertoli cell of non-obstructive azoospermia: a microarray data and in-silico analysis. Scientific reports 12 39472682
2018 Eleutheroside B1 mediates its anti-influenza activity through POLR2A and N-glycosylation. International journal of molecular medicine 12 30226535
1982 Metabolism and pharmacokinetics of dibromodulcitol (DBD, NSC-104800) in man--II. pharmacokinetics of DBD. European journal of cancer & clinical oncology 12 6891658
2020 Possible immune regulation mechanisms for the progression of chronic thromboembolic pulmonary hypertension. Thrombosis research 11 33316641
2024 Serum Fusion Transcripts to Assess the Risk of Hepatocellular Carcinoma and the Impact of Cancer Treatment through Machine Learning. The American journal of pathology 10 38537933
2022 The Competitive Endogenous RNA (ceRNA) Regulation in Porcine Alveolar Macrophages (3D4/21) Infected by Swine Influenza Virus (H1N1 and H3N2). International journal of molecular sciences 10 35163797
2018 Intestinal Epithelial Cell-specific Deletion of α-Mannosidase II Ameliorates Experimental Colitis. Cell structure and function 10 29343654
2024 Acceptor-Reactivity-Controlled Stereoconvergent Synthesis and Immunological Activity of a Unique Pentasaccharide from the Cell Wall Polysaccharide of Cutibacterium acnes C7. Angewandte Chemie (International ed. in English) 9 38651620
2022 Imputed expression of schizophrenia-associated genes and cognitive measures in patients with schizophrenia. Molecular genetics & genomic medicine 9 35488718
2019 Functional Annotation of Genetic Loci Associated With Sepsis Prioritizes Immune and Endothelial Cell Pathways. Frontiers in immunology 9 31475010
2015 The effects of swainsonine on the activity and expression of α-mannosidase in BRL-3A cells. Toxicon : official journal of the International Society on Toxinology 9 25797317
2021 Cell engineering for the production of hybrid-type N-glycans in HEK293 cells. Journal of biochemistry 8 33878161
2019 Mannose glycosylation is an integral step for NIS localization and function in human breast cancer cells. Journal of cell science 7 31455607
2019 BML-265 and Tyrphostin AG1478 Disperse the Golgi Apparatus and Abolish Protein Transport in Human Cells. Frontiers in cell and developmental biology 7 31681765
2003 Development of a comprehensive comparative radiation hybrid map of bovine chromosome 7 (BTA7) versus human chromosomes 1 (HSA1), 5 (HSA5) and 19 (HSA19). Cytogenetic and genome research 6 14970674
2021 N-Glycosylation Network Construction and Analysis to Modify Glycans on the Spike (S) Glycoprotein of SARS-CoV-2. Frontiers in bioinformatics 5 36303733
2004 The synthesis of a series of modified mannotrisaccharides as probes of the enzymes involved in the early stages of mammalian complex N-glycan formation. Carbohydrate research 5 15476709
2025 MAN2A2-related glycosylation defects in autism and cognitive delay. Scientific reports 4 40628855
2024 Immuno-targeting the ectopic phosphorylation sites of PDGFRA generated by MAN2A1-FER fusion in HCC. Hepatology communications 4 39082961
2024 Employing Multi-Omics Analyses to Understand Changes during Kidney Development in Perinatal Interleukin-6 Animal Model. Cells 4 39404429
2023 Optimizing effector functions of monoclonal antibodies via tailored N-glycan engineering using a dual landing pad CHO targeted integration platform. Scientific reports 4 37731040
2023 High-fat diet alters N-glycosylation of PTPRJ in murine liver. The Journal of nutritional biochemistry 4 37875230
2022 Homozygous truncating variant in MAN2A2 causes a novel congenital disorder of glycosylation with neurological involvement. Journal of medical genetics 4 36357165
2015 Association of chromosome 5q21.3 polymorphisms with the exploratory eye movement dysfunction in schizophrenia. Scientific reports 4 26242244
2025 Comparisons of miRNA profiles of exosomes derived from human iPSCs, ADSCs, and BMSCs and effects on chondrocyte function. Bone & joint research 2 40823751
2025 Integrated Multi-Level Investigation of Friend Leukemia Integration 1 Transcription Factor as a Novel Immune-Inflammatory Biomarker in Rheumatoid Arthritis: Bridging Bioinformatics, Clinical Cohorts, and Mechanistic Validation. Journal of inflammation research 1 40059948
2024 Whole-exome sequencing reveals novel genomic signatures and potential therapeutic targets during the progression of rectal neuroendocrine neoplasm. Cell death & disease 1 39548061
2026 Impact of afucosylation strategy on antibody function: a comparative study of glycoengineered anti-CD20 antibodies Obinutuzumab and Obinutuzumab beta. mAbs 0 41972453
2025 ManNAc attenuates Man5 glycoform abundance through GNE-mediated metabolic channeling of UDP-GlcNAc to N-glycosylation modifications via CMP-Neu5Ac biosynthesis. mAbs 0 40963238
2025 Plasma Protein Biomarkers to Detect Early Gastric Preneoplasia and Cancer: A Prospective Study. International journal of molecular sciences 0 41155404
2025 Green alga Ulva lactuca polysaccharide inhibits hepatocellular carcinoma through miRNA-mediated regulation and metabolite modulation. International journal of biological macromolecules 0 41297821
2023 High accumulation of the Man5GlcNAc2 structure by combining N-acetylglucosaminyltransferase I gene suppression and mannosidase I gene overexpression in Nicotiana tabacum SR1. Journal of bioscience and bioengineering 0 37311682

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