Affinage

LGALS4

Galectin-4 · UniProt P56470

Round 2 corrected
Length
323 aa
Mass
35.9 kDa
Annotated
2026-04-28
130 papers in source corpus 14 papers cited in narrative 14 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

Galectin-4 (LGALS4) is a tandem-repeat β-galactoside-binding lectin expressed predominantly in gastrointestinal epithelium that organizes apical membrane trafficking and modulates cell signaling and immune cell function. It clusters lipid rafts by binding sulfatides and complex N-glycans in detergent-resistant membranes, thereby directing post-Golgi apical delivery of glycoproteins in polarized enterocytes; its depletion causes intracellular accumulation of apical cargo without affecting basolateral markers, and it mediates basolateral-to-apical transcytosis of the transferrin receptor through recognition of a specific N-glycan (PMID:15883199, PMID:19192249, PMID:25179596). Beyond trafficking, galectin-4 binds sulfated glycosphingolipids, cholesterol 3-sulfate (via Arg45), and CEA on colon cell surfaces to promote cell adhesion, and induces phosphatidylserine exposure on activated neutrophils in a carbohydrate-dependent manner (PMID:15546874, PMID:17545668, PMID:16940423). In colorectal cancer cells, galectin-4 suppresses proliferation and migration by downregulating Wnt signaling and the IL-6/NF-κB/STAT3 axis, with its loss enhancing xenograft tumor growth (PMID:21064109, PMID:23378274).

Mechanistic history

Synthesis pass · year-by-year structured walk · 12 steps
  1. 1997 Medium

    Cloning of human galectin-4 established it as a two-CRD tandem-repeat lectin with expression restricted to the alimentary tract, defining the tissue context for all subsequent functional studies.

    Evidence cDNA cloning, sequencing, and Northern blot analysis of human tissues

    PMID:9310382

    Open questions at the time
    • No functional data beyond expression pattern
    • Post-translational modifications uncharacterized
  2. 1997 Medium

    Subcellular localization studies revealed that galectin-4 concentrates at sites of cell-substrate contact and lamellipodia, and immobilized galectin-4 directly promotes epithelial cell adhesion, providing the first evidence for an adhesion-related function.

    Evidence Immunofluorescence and cell adhesion assays in T84 colon carcinoma cells

    PMID:9162064

    Open questions at the time
    • Cell-surface receptors mediating adhesion not identified
    • In vivo relevance untested
  3. 2002 High

    Quantitative binding studies demonstrated that galectin-4 preferentially recognizes 3-O-sulfated glycans over unsulfated or sialylated core structures, shifting the field from generic β-galactoside binding to sulfation-dependent specificity.

    Evidence Surface plasmon resonance and competitive inhibition assays with defined saccharide ligands

    PMID:11971864

    Open questions at the time
    • Which CRD (N- or C-terminal) mediates sulfated glycan preference was unknown
    • In vivo ligands in intestinal epithelium not confirmed
  4. 2004 High

    Identification of sulfated glycosphingolipids and CEA as galectin-4 ligands on colon cell surfaces, with co-localization in lipid microdomains, connected galectin-4's carbohydrate specificity to a concrete cell-surface organizing function.

    Evidence Co-immunoprecipitation, SPR binding, confocal microscopy, and adhesion assays in colon carcinoma cells

    PMID:15546874

    Open questions at the time
    • Whether galectin-4–CEA interaction has signaling consequences was untested
    • Functional role of lipid microdomain organization not yet established
  5. 2005 High

    RNAi knockdown demonstrated that galectin-4 is required for apical delivery of membrane glycoproteins via lipid raft clustering, establishing it as a sorting lectin in polarized epithelial trafficking.

    Evidence Retroviral shRNA knockdown, DRM fractionation, immunofluorescence, and mass spectrometry in HT-29 5M12 enterocytic cells

    PMID:15883199

    Open questions at the time
    • How galectin-4 itself is sorted to the apical pathway was unclear
    • Whether galectin-4 acts alone or with co-factors in raft clustering was unknown
  6. 2007 High

    Discovery that galectin-4 binds cholesterol 3-sulfate—a non-β-galactoside lipid—via Arg45, revealed an unprecedented ligand-recognition mode for galectins and expanded the range of membrane lipids galectin-4 can organize.

    Evidence Site-directed mutagenesis (R45A), gel filtration, cross-linking, and detergent-insoluble fraction analysis in porcine tissues

    PMID:17545668

    Open questions at the time
    • Structural basis of cholesterol 3-sulfate recognition not resolved at atomic level
    • Physiological concentration of cholesterol 3-sulfate in intestinal membranes not measured
  7. 2009 High

    Mechanistic dissection showed that galectin-4 recruits apical glycoproteins bearing complex N-glycans into DRMs post-Golgi, and that galectin-4 itself follows an apical endocytic-recycling route, placing it in a self-reinforcing apical sorting circuit.

    Evidence shRNA knockdown, DRM fractionation, N-glycan mass spectrometry, and endocytic recycling assays in HT-29 cells

    PMID:19192249

    Open questions at the time
    • Mechanism of galectin-4 externalization (non-classical secretion) unresolved
    • Whether other galectins compensate in vivo remains unknown
  8. 2010 Medium

    Gain- and loss-of-function experiments in colorectal cancer cells linked galectin-4 to Wnt pathway suppression, cell-cycle arrest, and reduced migration, establishing a tumor-suppressive role beyond its trafficking function.

    Evidence Forced expression and shRNA knockdown with co-IP of Wnt pathway proteins, proliferation/migration assays, and apoptosis sensitization in CRC cell lines

    PMID:21064109

    Open questions at the time
    • Direct molecular target within the Wnt pathway not identified
    • Mechanism connecting lectin activity to Wnt suppression unclear
  9. 2011 Medium

    Galectin-4 was shown to modulate innate immunity by binding monocytes and macrophages, inducing monocyte apoptosis and cytokine release, and suppressing T-cell activation indirectly, extending its biology beyond epithelial trafficking.

    Evidence FACS binding, Transwell migration, phagocytosis, cytokine, and T-cell co-culture assays with primary human monocytes/macrophages

    PMID:21724180

    Open questions at the time
    • Monocyte surface receptor for galectin-4 not identified
    • In vivo relevance in gut immune homeostasis untested
  10. 2013 Medium

    Silencing galectin-4 activated the IL-6/NF-κB/STAT3 signaling axis and enhanced xenograft tumor growth, confirming galectin-4 as a suppressor of pro-inflammatory oncogenic signaling in colorectal cancer.

    Evidence shRNA knockdown, Western blot and qPCR array for signaling, and xenograft model in HT-29 cells

    PMID:23378274

    Open questions at the time
    • Whether galectin-4 directly inhibits NF-κB or acts upstream via membrane organization not distinguished
    • Single cell line for in vivo studies
  11. 2014 High

    Galectin-4 was identified as a mediator of basolateral-to-apical transcytosis of the transferrin receptor through recognition of a specific N-glycan (N727), with its loss diverting TfR to lysosomal degradation, revealing a glycan-dependent cargo sorting decision.

    Evidence shRNA knockdown and N-glycosylation site mutagenesis with pulse-chase trafficking in RPE, kidney, and MDCK cells

    PMID:25179596

    Open questions at the time
    • Whether other cargo glycoproteins use the same N-glycan-dependent transcytosis mechanism is unknown
    • Structural basis of galectin-4 selectivity for N727-linked glycan not resolved
  12. 2015 High

    Crystal structures of the C-terminal CRD with multiple ligands provided atomic-level understanding of how galectin-4 accommodates sulfated, fucosylated, and blood group saccharides, revealing that sulfate recognition does not involve a direct amino acid–sulfate contact.

    Evidence X-ray crystallography of galectin-4C with lactose, lactose-3'-sulfate, 2'-fucosyllactose, lacto-N-tetraose, and lacto-N-neotetraose

    PMID:26077389

    Open questions at the time
    • N-terminal CRD structures in complex with sulfated ligands not reported
    • No structure of full-length tandem-repeat protein available

Open questions

Synthesis pass · forward-looking unresolved questions
  • Key unresolved questions include the in vivo role of galectin-4 in intestinal epithelial polarity (no knockout mouse phenotype reported), the mechanism of its non-classical secretion, and whether its tumor-suppressive effects operate through lectin-mediated membrane organization or independent intracellular signaling.
  • No galectin-4 knockout animal model phenotype reported
  • Non-classical secretion mechanism unknown
  • Relationship between trafficking and tumor-suppressive functions unresolved

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0008289 lipid binding 5 GO:0098631 cell adhesion mediator activity 2
Localization
GO:0005886 plasma membrane 3 GO:0005576 extracellular region 2 GO:0031410 cytoplasmic vesicle 2 GO:0005829 cytosol 1
Pathway
R-HSA-5653656 Vesicle-mediated transport 3 R-HSA-9609507 Protein localization 3 R-HSA-162582 Signal Transduction 2 R-HSA-168256 Immune System 2
Partners
CEACAM5TFR1

Evidence

Reading pass · 14 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1997 Human galectin-4 was cloned and identified as a 323-amino-acid protein with two carbohydrate recognition domains, orthologous to rat and porcine galectin-4 (~76–79% amino acid identity). Its mRNA expression was found to be restricted to the small intestine, colon, and rectum. cDNA cloning, sequencing, and Northern blot tissue distribution analysis European Journal of Biochemistry Medium 9310382
1997 Galectin-4 and galectin-3 display strikingly different subcellular localizations in human colon adenocarcinoma T84 cells: galectin-4 is predominantly cytosolic and concentrated at the basal membrane in confluent cells and at the leading edge of lamellipodia in subconfluent cells, consistent with a role in cell adhesion. Immobilized recombinant galectin-4 stimulated T84 cell adhesion. Immunofluorescence microscopy, cell fractionation, cell adhesion assay with immobilized recombinant protein The Journal of Biological Chemistry Medium 9162064
2002 Recombinant human galectin-4 shows high-affinity binding (Kd ~3.4 × 10⁻⁶ M) to SO₃⁻→3Galβ1→3GalNAc pyranoside (3-O-sulfated core 1), two orders of magnitude tighter than to core 1 alone, while having weak affinity to lactose, type 1, and type 2 oligosaccharides. 3'-Sialylated core 1 had very weak affinity, demonstrating that sulfation rather than sialylation at the 3' position is critical for high-affinity recognition. Surface plasmon resonance, galectin-Sepharose column chromatography, competitive inhibition assay Glycobiology High 11971864
2004 Galectin-4 binds specifically to sulfated glycosphingolipids (SB1a, SM3, SM4s, SB2, SM2a, GM1) carrying 3-O-sulfated Gal residues but not to sialylated glycosphingolipids. On the surface of human colon adenocarcinoma cells (CCK-81 and LS174T), galectin-4 co-localizes with SB1a, GM1, and carcinoembryonic antigen (CEA) in cell-surface patches distinct from caveolin. CEA co-immunoprecipitated with galectin-4 from LS174T cell lysates (Kd ~2 × 10⁻⁸ M). Immobilized galectin-4 promoted cell adhesion through SB1a. ELISA, surface plasmon resonance, confocal immunocytochemistry, co-immunoprecipitation, cell adhesion assay The Journal of Biological Chemistry High 15546874
2005 Galectin-4 is a major component of detergent-resistant membranes (DRMs/lipid rafts) in polarized HT-29 5M12 cells and is present in post-Golgi carrier vesicles. RNAi-mediated depletion of galectin-4 causes intracellular accumulation of apical membrane markers without affecting basolateral markers, and alters the DRM-association of apical proteins. Sulfatides with long-chain hydroxylated fatty acids were identified as high-affinity ligands for galectin-4, implicating galectin-4–sulfatide interaction in lipid raft clustering for apical delivery. Retrovirus-mediated RNAi knockdown, detergent-resistant membrane fractionation, immunofluorescence, ligand identification by mass spectrometry and binding assays The Journal of Cell Biology High 15883199
2006 Human galectin-4 induces phosphatidylserine (PS) exposure on the outer leaflet of activated (but not resting) human neutrophils and several leukocyte cell lines in a carbohydrate-dependent manner, without causing apoptosis. Galectin-4 binds well to activated T lymphocytes but does not induce PS exposure or apoptosis in T cells, demonstrating cell-type specificity. Flow cytometry (annexin V staining for PS exposure), cell viability assays, carbohydrate inhibition experiments Blood Medium 16940423
2007 Galectin-4 binds to cholesterol 3-sulfate, a ligand lacking a β-galactoside moiety — a unique property within the galectin family. Site-directed mutagenesis (R45A) abolishes this binding, identifying Arg45 as essential for cholesterol 3-sulfate recognition. Galectin-4 exists partially as dimers, and this multivalency enhances avidity for cholesterol 3-sulfate. Cholesterol 3-sulfate co-fractionates with galectin-4 in detergent-insoluble fractions of porcine esophagus and intestine. Site-directed mutagenesis, gel filtration, chemical cross-linking, binding assays, detergent-insoluble fraction analysis The Journal of Biological Chemistry High 17545668
2009 Galectin-4 drives raft-dependent apical trafficking of glycoproteins in enterocyte-like HT-29 cells by recruiting apical glycoproteins into detergent-resistant membranes (DRMs). In galectin-4-knockdown cells, apical glycoproteins exit the Golgi but accumulate intracellularly. DRM-associated glycoproteins are identified as galectin-4 ligands enriched in complex N-glycans. Galectin-4, synthesized on free cytoplasmic ribosomes, is externalized apically and follows an apical endocytic-recycling pathway required for the apical biosynthetic pathway. shRNA-mediated knockdown, DRM fractionation, mass spectrometry glycan analysis, immunofluorescence, endocytic recycling assays Traffic High 19192249
2010 Forced expression of galectin-4 in galectin-4-negative colorectal cancer cells induces cell cycle arrest, reduces cell migration and motility, and sensitizes cells to camptothecin-induced apoptosis. Galectin-4 knockdown increases proliferation, migration, and motility. Galectin-4 was found to associate with Wnt signaling proteins and its expression led to downregulation of Wnt signaling target genes. Forced expression and shRNA knockdown, cell proliferation assays, migration/motility assays, cell cycle analysis, apoptosis assays, co-immunoprecipitation with Wnt signaling proteins, gene expression analysis International Journal of Cancer Medium 21064109
2011 Galectin-4 binds to human monocytes and macrophages, inhibits Salmonella-induced MHC II upregulation, induces cytokine release, inhibits monocyte migration, and potently induces monocyte (but not macrophage) apoptosis. Supernatants from macrophages co-cultured with galectin-4 significantly restrict T-cell activation, linking galectin-4-mediated innate immune regulation to adaptive immune suppression. Flow cytometry (FACS binding, MHCII, apoptosis), Transwell migration assay, phagocytosis assay, cytokine measurement, T-cell co-culture Cellular Immunology Medium 21724180
2013 shRNA-mediated silencing of galectin-4 in HT-29 colorectal cancer cells increases cell proliferation and activates NF-κB and STAT3 signaling with IL-6 upregulation, upregulates NF-κB target genes, and enhances in vivo xenograft tumor growth, establishing galectin-4 as a suppressor of the IL-6/NF-κB/STAT3 signaling axis. shRNA knockdown, Western blotting, qPCR array, immunofluorescence, xenograft tumor assay Cellular Oncology Medium 23378274
2014 Galectin-4 induces secretion of pro-angiogenic and pro-metastatic cytokines (G-CSF, IL-6, MCP-1, GROα) from blood vascular endothelial cells in vitro and in mice, and this galectin-induced cytokine secretion enhances endothelial cell surface adhesion molecule expression, increases cancer-endothelial adhesion, and promotes endothelial tubule formation. Cytokine array, in vitro endothelial cell treatment assays, in vivo mouse model, multiple regression analysis of patient serum British Journal of Cancer Medium 24384681
2014 Galectin-4 mediates basolateral-to-apical transcytosis of the transferrin receptor (TfR) in epithelial cells lacking the basolateral sorting adaptor AP-1B (RPE and kidney proximal tubule cell lines, AP-1B-knockdown MDCK cells). Mutation of the N-glycan linked to N727 of TfR or knockdown of galectin-4 inhibits TfR transcytosis to apical recycling endosomes and the apical plasma membrane, instead promoting TfR lysosomal targeting and degradation. shRNA knockdown, N-glycosylation site mutagenesis, immunofluorescence, subcellular fractionation, pulse-chase trafficking assays Journal of Cell Science High 25179596
2015 The X-ray crystal structure of the human galectin-4 C-terminal carbohydrate recognition domain (galectin-4C) was solved in complex with lactose, lactose-3'-sulfate, 2'-fucosyllactose, lacto-N-tetraose, and lacto-N-neotetraose. The structures reveal that galectin-4C recognizes sulfated saccharides without a specific amino acid contact with the sulfate group, and that the non-reducing-end galactose of lacto-N-tetraose and lacto-N-neotetraose adopt different binding orientations. Recognition of A and B blood group antigens is enabled by structural features of the extended binding site, while the 2'-fucose of blood group antigens does not interact directly with the protein. X-ray crystallography, molecular dynamics simulations The FEBS Journal High 26077389

Source papers

Stage 0 corpus · 130 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2005 A human protein-protein interaction network: a resource for annotating the proteome. Cell 1704 16169070
2002 Generation and initial analysis of more than 15,000 full-length human and mouse cDNA sequences. Proceedings of the National Academy of Sciences of the United States of America 1479 12477932
1994 Galectins. Structure and function of a large family of animal lectins. The Journal of biological chemistry 1222 8063692
2020 A reference map of the human binary protein interactome. Nature 849 32296183
1994 Stimulation of GAL4 derivative binding to nucleosomal DNA by the yeast SWI/SNF complex. Science (New York, N.Y.) 726 8016655
1985 Specific DNA binding of GAL4, a positive regulatory protein of yeast. Cell 628 3886158
1988 Negative effect of the transcriptional activator GAL4. Nature 602 3412449
2006 Hsp90 cochaperone Aha1 downregulation rescues misfolding of CFTR in cystic fibrosis. Cell 517 17110338
2008 Genetic dissection of neural circuits by Tol2 transposon-mediated Gal4 gene and enhancer trapping in zebrafish. Proceedings of the National Academy of Sciences of the United States of America 450 18202183
2004 The status, quality, and expansion of the NIH full-length cDNA project: the Mammalian Gene Collection (MGC). Genome research 438 15489334
1996 Generation and analysis of 280,000 human expressed sequence tags. Genome research 376 8889549
2007 Transactivation from Gal4-VP16 transgenic insertions for tissue-specific cell labeling and ablation in zebrafish. Developmental biology 329 17335798
2009 G-TRACE: rapid Gal4-based cell lineage analysis in Drosophila. Nature methods 280 19633663
1989 An amino-terminal fragment of GAL4 binds DNA as a dimer. Journal of molecular biology 280 2511324
1988 GAL4 derivatives function alone and synergistically with mammalian activators in vitro. Cell 259 3044607
1991 Facilitated binding of GAL4 and heat shock factor to nucleosomal templates: differential function of DNA-binding domains. Genes & development 245 2065977
2018 A gene-specific T2A-GAL4 library for Drosophila. eLife 243 29565247
1982 Isolation of the yeast regulatory gene GAL4 and analysis of its dosage effects on the galactose/melibiose regulon. Proceedings of the National Academy of Sciences of the United States of America 240 6294669
1984 Primary structure of the Saccharomyces cerevisiae GAL4 gene. Molecular and cellular biology 234 6366516
2005 Galectin-4 and sulfatides in apical membrane trafficking in enterocyte-like cells. The Journal of cell biology 213 15883199
1992 GAL4 fusion vectors for expression in yeast or mammalian cells. Gene 213 1511877
2014 Extracellular matrix signatures of human primary metastatic colon cancers and their metastases to liver. BMC cancer 203 25037231
1988 Cooperative DNA binding of the yeast transcriptional activator GAL4. Proceedings of the National Academy of Sciences of the United States of America 190 3124106
1991 Regulated expression of the GAL4 activator gene in yeast provides a sensitive genetic switch for glucose repression. Proceedings of the National Academy of Sciences of the United States of America 182 1924319
2008 Gal4/UAS transgenic tools and their application to zebrafish. Zebrafish 163 18554173
2018 Genetic Reagents for Making Split-GAL4 Lines in Drosophila. Genetics 156 29535151
1995 GAL4 interacts with TATA-binding protein and coactivators. Molecular and cellular biology 151 7739564
2005 The F box protein Dsg1/Mdm30 is a transcriptional coactivator that stimulates Gal4 turnover and cotranscriptional mRNA processing. Cell 144 15797387
1982 Isolation and preliminary characterization of the GAL4 gene, a positive regulator of transcription in yeast. Proceedings of the National Academy of Sciences of the United States of America 143 6294656
1987 Genetic evidence that zinc is an essential co-factor in the DNA binding domain of GAL4 protein. Nature 142 3299106
1990 GAL11P: a yeast mutation that potentiates the effect of weak GAL4-derived activators. Cell 137 2124519
1990 A potent GAL4 derivative activates transcription at a distance in vitro. Science (New York, N.Y.) 137 2405489
2010 Characterization of midgut stem cell- and enteroblast-specific Gal4 lines in drosophila. Genesis (New York, N.Y. : 2000) 134 20681020
1987 Interaction of GAL4 and GAL80 gene regulatory proteins in vitro. Molecular and cellular biology 134 3316976
2006 Human galectin-1, -2, and -4 induce surface exposure of phosphatidylserine in activated human neutrophils but not in activated T cells. Blood 127 16940423
1992 Nondissociation of GAL4 and GAL80 in vivo after galactose induction. Science (New York, N.Y.) 125 1598579
2009 Galectin-4-regulated delivery of glycoproteins to the brush border membrane of enterocyte-like cells. Traffic (Copenhagen, Denmark) 124 19192249
1989 Mutations in RNA polymerase II enhance or suppress mutations in GAL4. Proceedings of the National Academy of Sciences of the United States of America 121 2495535
1987 Characterization of a positive regulatory gene, LAC9, that controls induction of the lactose-galactose regulon of Kluyveromyces lactis: structural and functional relationships to GAL4 of Saccharomyces cerevisiae. Molecular and cellular biology 120 3550430
2004 Galectin-4 binds to sulfated glycosphingolipids and carcinoembryonic antigen in patches on the cell surface of human colon adenocarcinoma cells. The Journal of biological chemistry 106 15546874
2008 The GAL4 system : a versatile system for the expression of genes. Methods in molecular biology (Clifton, N.J.) 105 18641942
1993 ADA3: a gene, identified by resistance to GAL4-VP16, with properties similar to and different from those of ADA2. Molecular and cellular biology 102 8413201
2015 A 3D Searchable Database of Transgenic Zebrafish Gal4 and Cre Lines for Functional Neuroanatomy Studies. Frontiers in neural circuits 100 26635538
2010 Galectin-4 functions as a tumor suppressor of human colorectal cancer. International journal of cancer 95 21064109
2002 High-affinity binding of recombinant human galectin-4 to SO(3)(-)-->3Galbeta1-->3GalNAc pyranoside. Glycobiology 95 11971864
2003 GAL4 causes developmental defects and apoptosis when expressed in the developing eye of Drosophila melanogaster. Genetics and molecular research : GMR 90 12917801
1997 Strikingly different localization of galectin-3 and galectin-4 in human colon adenocarcinoma T84 cells. Galectin-4 is localized at sites of cell adhesion. The Journal of biological chemistry 87 9162064
2001 Efficient and heritable functional knock-out of an adult phenotype in Drosophila using a GAL4-driven hairpin RNA incorporating a heterologous spacer. Nucleic acids research 82 11410678
1990 Analysis of the GAL3 signal transduction pathway activating GAL4 protein-dependent transcription in Saccharomyces cerevisiae. Genetics 81 2199310
2023 A searchable image resource of Drosophila GAL4 driver expression patterns with single neuron resolution. eLife 75 36820523
2007 Expression of Gal4-dependent transgenes in cells of the mononuclear phagocyte system labeled with enhanced cyan fluorescent protein using Csf1r-Gal4VP16/UAS-ECFP double-transgenic mice. Journal of leukocyte biology 75 17971498
2019 The midbody interactome reveals unexpected roles for PP1 phosphatases in cytokinesis. Nature communications 74 31586073
1986 Functional domains of the yeast regulatory protein GAL4. Proceedings of the National Academy of Sciences of the United States of America 74 2944111
2016 Tissue- and cell-specific localization of galectins, β-galactose-binding animal lectins, and their potential functions in health and disease. Anatomical science international 73 27590897
2012 Depleting gene activities in early Drosophila embryos with the "maternal-Gal4-shRNA" system. Genetics 72 23105012
1997 Cloning and expression of the mRNA of human galectin-4, an S-type lectin down-regulated in colorectal cancer. European journal of biochemistry 71 9310382
2014 Circulating galectins -2, -4 and -8 in cancer patients make important contributions to the increased circulation of several cytokines and chemokines that promote angiogenesis and metastasis. British journal of cancer 70 24384681
2008 Structural basis for dimerization in DNA recognition by Gal4. Structure (London, England : 1993) 68 18611375
2006 Proteolytic turnover of the Gal4 transcription factor is not required for function in vivo. Nature 68 16929306
2009 The Tol2-mediated Gal4-UAS method for gene and enhancer trapping in zebrafish. Methods (San Diego, Calif.) 67 19835787
2016 cGAL, a temperature-robust GAL4-UAS system for Caenorhabditis elegans. Nature methods 65 27992408
2015 Assembly, organization and regulation of cell-surface receptors by lectin-glycan complexes. The Biochemical journal 64 26173257
2004 Characterization of Drosophila fruitless-gal4 transgenes reveals expression in male-specific fruitless neurons and innervation of male reproductive structures. The Journal of comparative neurology 60 15211467
2014 Galectin-4 serves as a prognostic biomarker for the early recurrence / metastasis of hepatocellular carcinoma. Cancer science 58 25230111
2014 Hexameric GFP and mCherry reporters for the Drosophila GAL4, Q, and LexA transcription systems. Genetics 57 24451596
2014 Establishment of Gal4 transgenic zebrafish lines for analysis of development of cerebellar neural circuitry. Developmental biology 56 25300581
2013 Abrogation of galectin-4 expression promotes tumorigenesis in colorectal cancer. Cellular oncology (Dordrecht, Netherlands) 55 23378274
2020 The Drosophila Split Gal4 System for Neural Circuit Mapping. Frontiers in neural circuits 54 33240047
2000 A GAL4-like protein is involved in the switch between biotrophic and necrotrophic phases of the infection process of Colletotrichum lindemuthianum on common bean. The Plant cell 54 11006333
2018 Development of a temperature-responsive yeast cell factory using engineered Gal4 as a protein switch. Biotechnology and bioengineering 50 29315481
2007 Recognition mechanism of galectin-4 for cholesterol 3-sulfate. The Journal of biological chemistry 50 17545668
1993 Wild type GAL4 binds cooperatively to the GAL1-10 UASG in vitro. The Journal of biological chemistry 50 8486650
2020 Analysis of Gal4 Expression Patterns in Adult Drosophila Females. G3 (Bethesda, Md.) 48 32917721
2016 Gal4 Driver Transgenic Zebrafish: Powerful Tools to Study Developmental Biology, Organogenesis, and Neuroscience. Advances in genetics 47 27503354
2013 Galectin-4, a novel predictor for lymph node metastasis in lung adenocarcinoma. PloS one 47 24339976
2011 Galectins distinctively regulate central monocyte and macrophage function. Cellular immunology 47 21724180
2021 Mediator subunit Med15 dictates the conserved "fuzzy" binding mechanism of yeast transcription activators Gal4 and Gcn4. Nature communications 44 33850123
2009 Adaptation of GAL4 activators for GAL4 enhancer trapping in zebrafish. Developmental dynamics : an official publication of the American Association of Anatomists 44 19191223
2015 Structural characterization of human galectin-4 C-terminal domain: elucidating the molecular basis for recognition of glycosphingolipids, sulfated saccharides and blood group antigens. The FEBS journal 43 26077389
2002 GAL4 directs nucleosome sliding induced by NURF. The EMBO journal 42 11889046
2001 The Gal4 activation domain binds Sug2 protein, a proteasome component, in vivo and in vitro. The Journal of biological chemistry 42 11418596
2006 Quantitative analysis of the GAL4/UAS system in Drosophila oogenesis. Genesis (New York, N.Y. : 2000) 41 16425298
1993 Promoter elements determining weak expression of the GAL4 regulatory gene of Saccharomyces cerevisiae. Molecular and cellular biology 41 8393142
2007 Neuronal death in Drosophila triggered by GAL4 accumulation. The European journal of neuroscience 40 17313569
2001 Recruitment of the transcriptional machinery through GAL11P: structure and interactions of the GAL4 dimerization domain. Genes & development 37 11316794
1997 Gene transfer using a novel fusion protein, GAL4/invasin. Human gene therapy 37 9215742
2017 Facilitating Neuron-Specific Genetic Manipulations in Drosophila melanogaster Using a Split GAL4 Repressor. Genetics 36 28363977
2015 The commonly used eye-specific sev-GAL4 and GMR-GAL4 drivers in Drosophila melanogaster are expressed in tissues other than eyes also. Journal of genetics 36 26440079
2012 Simultaneous determination of serum galectin-3 and -4 levels detects metastases in colorectal cancer patients. Cellular oncology (Dordrecht, Netherlands) 34 23117840
2017 Drosophila enhancer-Gal4 lines show ectopic expression during development. Royal Society open science 33 28405401
2016 A galectin-specific signature in the gut delineates Crohn's disease and ulcerative colitis from other human inflammatory intestinal disorders. BioFactors (Oxford, England) 33 26891020
1991 The yeast GAL4 protein transactivates the polyomavirus origin of DNA replication in mouse cells. Journal of virology 33 1645781
2023 Using single-cell RNA sequencing to generate predictive cell-type-specific split-GAL4 reagents throughout development. Proceedings of the National Academy of Sciences of the United States of America 32 37523539
2014 Galectin-4-mediated transcytosis of transferrin receptor. Journal of cell science 32 25179596
2019 The roles of galectin-3 and galectin-4 in the idiopatic Parkinson disease and its progression. Clinical neurology and neurosurgery 30 31147178
2019 A Catalog of GAL4 Drivers for Labeling and Manipulating Circadian Clock Neurons in Drosophila melanogaster. Journal of biological rhythms 30 31856635
2019 GAL4 Drivers Specific for Type Ib and Type Is Motor Neurons in Drosophila. G3 (Bethesda, Md.) 27 30530644
2009 The expanding reach of the GAL4/UAS system into the behavioral neurobiology of Drosophila. BMB reports 27 19944010
2023 A split GAL4 RUBY assay for visual in planta detection of protein-protein interactions. The Plant journal : for cell and molecular biology 26 37323061
2020 Colorectal cancer screening: Assessment of CEACAM6, LGALS4, TSPAN8 and COL1A2 as blood markers in faecal immunochemical test negative subjects. Journal of advanced research 26 32257432
2017 Regulating the UAS/GAL4 system in adult Drosophila with Tet-off GAL80 transgenes. PeerJ 25 29259847
2013 Targeted expression in zebrafish primordial germ cells by Cre/loxP and Gal4/UAS systems. Marine biotechnology (New York, N.Y.) 25 23535913
2016 Tol2-mediated transgenesis, gene trapping, enhancer trapping, and Gal4-UAS system. Methods in cell biology 24 27443919
2005 Development and evaluation of a Gal4-mediated LUC/GFP/GUS enhancer trap system in Arabidopsis. BMC plant biology 24 15941484
2000 Technical advance: the DNA-binding activity of gal4 is inhibited by methylation of the gal4 binding site in plant chromatin. The Plant journal : for cell and molecular biology 24 10929109
2007 Design and isolation of temperature-sensitive mutants of Gal4 in yeast and Drosophila. Journal of molecular biology 23 17553522
1998 Alterations in the GAL4 DNA-binding domain can affect transcriptional activation independent of DNA binding. The Journal of biological chemistry 23 9593720
2003 Cardiac expression of Gal4 causes cardiomyopathy in a dose-dependent manner. Journal of muscle research and cell motility 22 14609031
2001 JAB1/CSN5 interacts with the GAL4 DNA binding domain: a note of caution about two-hybrid interactions. Biochimie 22 11728635
2000 A GAL4-HP1 fusion protein targeted near heterochromatin promotes gene silencing. Chromosoma 22 11151674
2025 A split-GAL4 driver line resource for Drosophila neuron types. eLife 21 39854223
2023 split-intein Gal4 provides intersectional genetic labeling that is repressible by Gal80. Proceedings of the National Academy of Sciences of the United States of America 21 37276389
2020 Molecular and cytological analysis of widely-used Gal4 driver lines for Drosophila neurobiology. BMC genetics 21 33092520
2016 Molecular Characterization of Arabidopsis GAL4/UAS Enhancer Trap Lines Identifies Novel Cell-Type-Specific Promoters. Plant physiology 21 27208300
2000 Biochemical characterization of the TATA-binding protein-Gal4 activation domain complex. The Journal of biological chemistry 19 10931832
2015 High-resolution analysis of central nervous system expression patterns in zebrafish Gal4 enhancer-trap lines. Developmental dynamics : an official publication of the American Association of Anatomists 18 25694140
1997 Refined solution structure of the DNA-binding domain of GAL4 and use of 3J(113Cd,1H) in structure determination. Journal of biomolecular NMR 17 9460244
1994 Directing transcription of an RNA polymerase III gene via GAL4 sites. Proceedings of the National Academy of Sciences of the United States of America 17 7991561
2023 TGM2, HMGA2, FXYD3, and LGALS4 genes as biomarkers in acquired oxaliplatin resistance of human colorectal cancer: A systems biology approach. PloS one 16 37535601
2015 Transcriptional Activity and DNA Methylation Dynamics of the Gal4/UAS System in Zebrafish. Marine biotechnology (New York, N.Y.) 16 25997914
1993 GAL4-I kappa B alpha and GAL4-I kappa B gamma activate transcription by different mechanisms. Nucleic acids research 16 8502557
2020 Identification of Split-GAL4 Drivers and Enhancers That Allow Regional Cell Type Manipulations of the Drosophila melanogaster Intestine. Genetics 15 32988987
2013 Gene trapping using gal4 in zebrafish. Journal of visualized experiments : JoVE 15 24121167
2017 The 9aaTAD Is Exclusive Activation Domain in Gal4. PloS one 14 28056036
2013 Self-association of the Gal4 inhibitor protein Gal80 is impaired by Gal3: evidence for a new mechanism in the GAL gene switch. Molecular and cellular biology 14 23858060
2001 GCN5 dependence of chromatin remodeling and transcriptional activation by the GAL4 and VP16 activation domains in budding yeast. Molecular and cellular biology 14 11416135
2017 Genetic and Epigenetic Strategies Potentiate Gal4 Activation to Enhance Fitness in Recently Diverged Yeast Species. Current biology : CB 13 29153325
2014 Inactivation of a GAL4-like transcription factor improves cell fitness and product yield in glycoengineered Pichia pastoris strains. Applied and environmental microbiology 13 25344235
2012 Mediator acts upstream of the transcriptional activator Gal4. PLoS biology 13 22479149
1995 Genetic analysis of growth inhibition by GAL4-L kappa B-alpha in Saccharomyces cerevisiae. Cell growth & differentiation : the molecular biology journal of the American Association for Cancer Research 13 7547500