Affinage

HMGN2

Non-histone chromosomal protein HMG-17 · UniProt P05204

Length
90 aa
Mass
9.4 kDa
Annotated
2026-04-28
100 papers in source corpus 32 papers cited in narrative 32 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

HMGN2 is a nucleosome-binding chromatin architectural protein that modulates higher-order chromatin structure to regulate transcription, replication, and DNA repair. Its 30-amino-acid nucleosome-binding domain contacts two sites per nucleosome core near the dyad axis and entry/exit regions, requiring histone tails for binding and antagonizing linker histone H1 occupancy; incorporation during chromatin assembly unfolds chromatin and enhances RNA polymerase II transcription initiation and DNA replication from nucleosomal templates (PMID:7649479, PMID:9545265, PMID:8107104, PMID:1453455, PMID:28035005). HMGN2 preferentially associates with acetylated H3 and H2A.Z-containing nucleosomes, maintains active histone marks (H3K27ac, H3K9ac, H3K4me3) at target loci, suppresses H3K27 methylation, and its chromatin binding is negatively regulated by acetylation at K2 (written by PCAF, erased by HDAC6), SUMOylation at K17/K35 (by PIAS1), and succinylation at K30 (PMID:10207070, PMID:41325801, PMID:24872413, PMID:34458839, PMID:27358110). Beyond its chromatin role, HMGN2 directly interacts with homeodomain transcription factors (PITX2, Lef-1, Dlx2) to inhibit their DNA binding, modulates NF-κB and Nrf2 signaling in innate immune responses, and its alpha-helical domain possesses intrinsic antimicrobial activity against Gram-negative bacteria (PMID:18045789, PMID:35872015, PMID:21518253, PMID:16115376, PMID:31596045).

Mechanistic history

Synthesis pass · year-by-year structured walk · 19 steps
  1. 1984 Medium

    Early work established that HMGN2 alters DNA topology by facilitating topoisomerase-mediated catenation, indicating a role in modulating DNA structure beyond simple binding.

    Evidence In vitro topoisomerase catenation assay with purified HMG-17

    PMID:6326673

    Open questions at the time
    • Single lab, single method
    • Physiological relevance of catenation-promoting activity unclear
    • No chromatin context tested
  2. 1987 High

    Immunofractionation of oligonucleosomes revealed that HMGN2 is selectively enriched downstream of transcription start sites in active genes, establishing its association with transcriptionally active chromatin domains.

    Evidence Monoclonal antibody immunoisolation of oligonucleosomes from chicken liver and oviduct with gene-specific probes

    PMID:3665881

    Open questions at the time
    • Causal relationship between HMGN2 presence and transcription not tested
    • Mechanism of preferential deposition at active genes unknown
  3. 1992 High

    Mapping of the nucleosome-binding domain to a 30-amino-acid peptide that autonomously recognizes nucleosome cores and requires histone tails defined the minimal structural module for HMGN2–chromatin interaction.

    Evidence Mobility shift, thermal denaturation, DNase I footprinting, and trypsin digestion with synthetic peptides and recombinant protein

    PMID:1453455

    Open questions at the time
    • Specific histone tail residues required not identified
    • No structure of the domain–nucleosome complex
  4. 1993 High

    Cell-free chromatin assembly experiments demonstrated that HMGN2 must be incorporated during replication-coupled assembly to enhance transcription, revealing that it acts as a constitutive chromatin component rather than a post-hoc modifier.

    Evidence Xenopus egg extract chromatin assembly with transcription assays on 5S RNA gene and satellite I templates

    PMID:8404854

    Open questions at the time
    • Mechanism of co-assembly incorporation unknown
    • Whether replication-independent assembly also suffices not tested
  5. 1994 High

    Hydroxyl radical footprinting resolved the precise binding geometry: two HMGN2 molecules bind per nucleosome near the dyad and ~25 bp from core DNA ends, overlapping linker histone sites, explaining how HMGN2 and H1 compete for the same nucleosome.

    Evidence Hydroxyl radical footprinting of nucleosome cores and H1/H5-depleted chromatosomes

    PMID:8107104

    Open questions at the time
    • No atomic-resolution structure
    • How two HMGN2 molecules cooperatively bind is not resolved
  6. 1995 High

    Reconstituted chromatin transcription assays quantified a 7–40-fold stimulation of RNA polymerase II transcription initiation, establishing HMGN2 as a bona fide chromatin-specific transcriptional coactivator.

    Evidence In vitro transcription from regularly spaced nucleosomal arrays with GAL4-VP16 activator

    PMID:7649479

    Open questions at the time
    • Whether elongation is also affected was not resolved
    • Generality across diverse promoters not tested
  7. 1997 Medium

    Discovery that HMGN2-containing nucleosomes cluster in contiguous runs of ~6 and are mutually exclusive with HMGN1 revealed that HMGN variants define distinct chromatin domains rather than being randomly distributed.

    Evidence Immunofractionation and confocal immunofluorescence microscopy

    PMID:9417927

    Open questions at the time
    • Single lab
    • Mechanism of domain segregation unknown
    • Functional consequence of clustering not tested
  8. 1998 High

    Three concurrent studies resolved HMGN2 dynamics and function: it colocalizes with active pol II sites, is released from chromatin during mitosis and re-imported via importin-α in telophase, and stimulates SV40 replication efficiency only from chromatin assembled with the protein.

    Evidence Immunofluorescence through cell cycle, nuclear import assays, peptide microinjection, SV40 minichromosome replication

    PMID:9545265 PMID:9843505 PMID:9852141

    Open questions at the time
    • Whether mitotic release serves a regulatory function or is passive is unknown
    • Replication stimulation mechanism at the molecular level unresolved
  9. 1999 High

    Identification of PCAF as the acetyltransferase that acetylates K2, with acetylation reducing nucleosome affinity, established the first post-translational switch controlling HMGN2–chromatin interaction; reciprocally, nucleosome-bound HMGN2 inhibits PCAF acetylation of H3.

    Evidence In vitro acetyltransferase assay, mass spectrometry, equilibrium dialysis

    PMID:10207070

    Open questions at the time
    • In vivo dynamics of K2 acetylation not characterized
    • Identity of additional acetyltransferases not excluded
  10. 2005 Medium

    Domain mapping revealed that the alpha-helical region (residues 18–48, overlapping the nucleosome-binding domain) has direct antimicrobial activity against Gram-negative bacteria and Candida, indicating an extranuclear function.

    Evidence In vitro MIC/MBC assays with recombinant fragments and synthetic peptides

    PMID:16115376

    Open questions at the time
    • Single lab
    • In vivo antimicrobial relevance not demonstrated
    • Mechanism of membrane disruption not characterized
  11. 2007 High

    Discovery that HMGN2 directly inhibits PITX2 DNA binding, with β-catenin converting the complex into a transcriptional activator, and that homozygous HMGN2 knockout mice are embryonic lethal, established HMGN2 as a molecular switch for homeodomain transcription factor activity essential for development.

    Evidence Co-IP, EMSA, reporter assays, confocal microscopy, homozygous knockout mouse

    PMID:18045789

    Open questions at the time
    • Stage and cause of embryonic lethality not defined
    • Whether all homeodomain interactions share the same mechanism unclear
  12. 2009 High

    HMGN2 knockout DT40 cells showed impaired removal of UV-induced lesions from chromatin without affecting core NER catalysis, demonstrating that HMGN2 facilitates the global genome repair subpathway through chromatin accessibility rather than repair enzyme function.

    Evidence DT40 gene knockout, UV sensitivity, host cell reactivation, DNA lesion kinetics

    PMID:19843163

    Open questions at the time
    • Mechanism of chromatin opening for NER factors not defined
    • Mammalian confirmation not provided
  13. 2011 Medium

    Two studies connected HMGN2 to innate immune transcriptional regulation: it promotes LPS-induced β-defensin-2 expression by enhancing NF-κB p65 nuclear retention and acetylation, and binds nuclear prolactin receptor in a phosphorylation-dependent manner to enable Stat5a-driven transcription.

    Evidence siRNA knockdown with ChIP, reporter assays, HAT activity measurement; Co-IP and ChIP for Stat5a promoter binding

    PMID:21518253 PMID:21816901

    Open questions at the time
    • Single labs for each study
    • Whether HMGN2–p65 interaction is direct or chromatin-mediated unclear
    • In vivo relevance of PRLr interaction not established
  14. 2014 High

    Identification of PIAS1-mediated SUMOylation at K17 and K35 within the nucleosome-binding domain, reversed by SENP1, provided a second PTM switch that reduces nucleosome affinity, paralleling acetylation at K2.

    Evidence In vitro SUMOylation with site mutagenesis, reconstituted nucleosome binding assays

    PMID:24872413

    Open questions at the time
    • In vivo SUMOylation dynamics and downstream transcriptional consequences not characterized
    • Interplay between SUMOylation and acetylation not tested
  15. 2016 High

    The acetyltransferase/deacetylase cycle at K2 was completed by identifying HDAC6 as the eraser; deacetylation promotes Stat5a transcription and breast cancer growth, linking HMGN2 PTM status to oncogenic signaling. Concurrently, epistasis experiments showed HMGN2 antagonizes linker histone H1 to enable STAT5 chromatin access.

    Evidence HDAC6 inhibitor treatment in vitro/in vivo with Stat5a target gene readouts; double knockdown of HMGN2 and H1 with ChIP

    PMID:27358110 PMID:28035005

    Open questions at the time
    • Whether HDAC6-HMGN2 axis operates in non-breast contexts unknown
    • Structural basis for H1–HMGN2 competition at specific loci not resolved
  16. 2019 High

    Loss of HMGN2 in embryonal carcinoma cells globally reduced H3K9ac and disrupted active marks at pluripotency loci (Nanog, Oct4), causing spontaneous neuronal differentiation and establishing HMGN2 as a guardian of the active chromatin landscape required for pluripotency.

    Evidence HMGN2 knockout cell lines, ChIP-seq for multiple histone marks, differentiation marker analysis

    PMID:31831052

    Open questions at the time
    • Whether effect is direct binding at pluripotency loci or indirect global chromatin change unresolved
    • In vivo embryonic stem cell confirmation lacking
  17. 2021 High

    Site-specific succinylation at K30 in the nucleosome-binding domain was shown to decrease nucleosome binding and promote DNA unwrapping at entry/exit sites, adding a third class of PTM that regulates HMGN2–nucleosome interaction.

    Evidence Succinyl lysine analogue incorporation with mononucleosome binding and DNA accessibility assays

    PMID:34458839

    Open questions at the time
    • Physiological writer/eraser for K30 succinylation not identified
    • In vivo prevalence of K30 succinylation unknown
  18. 2022 High

    HMGN2 was shown to interact with and inhibit Lef-1 DNA binding in addition to PITX2, Dlx2, and FoxJ1, with Hmgn2 ablation in mice increasing amelogenin expression; miR-23a/b were identified as post-transcriptional regulators of Hmgn2, broadening the homeodomain switch model and revealing upstream control of HMGN2 levels.

    Evidence BiFC, pull-down, Co-IP, EMSA, ChIP, mouse knockout, luciferase reporters

    PMID:35872015

    Open questions at the time
    • Whether miR-23a/b regulation is tissue-specific not determined
    • Structural basis for homeodomain recognition by HMGN2 unknown
  19. 2025 High

    Two studies extended HMGN2 biology: it preferentially binds acetylated-H3 and H2A.Z nucleosomes, limits p300-mediated H3K27 acetylation, and suppresses H3K27me2/me3 (with HMGN1); separately, PIAS1-mediated SUMOylation of HMGN2 enhances its interaction with PAX5 to drive macrophage M1 polarization and NF-κB signaling in atherosclerosis.

    Evidence Reconstituted nucleosome binding with modified histones, p300 acetylation assay, epiproteomic MS in CRISPR KO mESCs; Co-IP with PIAS1 knockdown in ApoE−/− mouse model

    PMID:40834970 PMID:41325801

    Open questions at the time
    • How HMGN2 binding to acetylated nucleosomes mechanistically suppresses H3K27me3 not resolved
    • Whether SUMOylation-dependent PAX5 interaction occurs genome-wide unknown
    • Relative contributions of HMGN1 vs HMGN2 to H3K27me3 suppression not separated

Open questions

Synthesis pass · forward-looking unresolved questions
  • No high-resolution structure of HMGN2 bound to a nucleosome exists, the mechanism by which HMGN2 is selectively deposited into active chromatin domains during replication is unknown, and how multiple PTM switches (acetylation, SUMOylation, succinylation) are coordinated in vivo to tune chromatin architecture remains unresolved.
  • No atomic structure of HMGN2–nucleosome complex
  • Mechanism of replication-coupled chromatin incorporation unknown
  • Combinatorial PTM regulation in vivo not addressed
  • Cause of embryonic lethality in knockout mice not defined

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0098772 molecular function regulator activity 6 GO:0140110 transcription regulator activity 6 GO:0042393 histone binding 5
Localization
GO:0005694 chromosome 5 GO:0000228 nuclear chromosome 3 GO:0005634 nucleus 3
Pathway
R-HSA-74160 Gene expression (Transcription) 7 R-HSA-4839726 Chromatin organization 6 R-HSA-168256 Immune System 5 R-HSA-69306 DNA Replication 2 R-HSA-73894 DNA Repair 1
Partners
CTNNB1HDAC6LEF1PAX5PCAFPIAS1PITX2PRLR

Evidence

Reading pass · 32 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1999 PCAF acetyltransferase specifically acetylates HMGN2 (HMG-17) at lysine 2, and this acetylation reduces the affinity of HMGN2 for nucleosome cores; conversely, nucleosome-bound HMGN2 inhibits PCAF-mediated acetylation of histone H3. In vitro acetyltransferase assay, mass spectrometry sequence analysis, equilibrium dialysis, competition studies with deletion mutants Molecular and cellular biology High 10207070
1995 HMGN2 (HMG-17) acts as a chromatin-specific transcriptional coactivator that increases the efficiency of RNA polymerase II transcription initiation (7- to 40-fold stimulation) only from nucleosomal chromatin templates assembled with the protein, not from protein-free DNA or post-assembly added protein. In vitro chromatin assembly and transcription assay with regularly spaced nucleosomal arrays, GAL4-VP16 activator, structural analysis Genes & development High 7649479
1993 HMGN2 (HMG-17) is incorporated into nascent chromatin during replication in a cell-free Xenopus egg system, stabilizes nucleosomal core structure, improves nucleosomal periodicity, and significantly increases transcriptional potential only when incorporated during (not after) chromatin assembly. Cell-free Xenopus egg extract chromatin assembly, transcription assays with 5S RNA gene and satellite I chromatin, kinetic studies The EMBO journal High 8404854
1998 HMGN2 (HMG-17) stimulates replication efficiency of chromatin templates (SV40 minichromosomes) only when incorporated during chromatin assembly, not post-assembly, and only from chromatin templates, not protein-free DNA; the effect is associated with induction of extended chromatin structure. In vitro SV40 replication system, minichromosome assembly in Xenopus egg extracts, structural analysis The Journal of biological chemistry High 9545265
1994 Hydroxyl radical footprinting mapped HMGN2 (HMG-17) binding sites on nucleosome cores: two molecules bind per nucleosome, protecting DNA ~25 bp from the end of the core and near the nucleosomal dyad axis, bridging two adjacent DNA strands on the surface, with binding sites overlapping those of linker histones near the dyad. Hydroxyl radical footprinting of nucleosome cores and H1/H5-depleted chromatosomes Journal of molecular biology High 8107104
1992 The nucleosome-binding domain of HMGN2 (HMG-17) — a 30 amino acid peptide — functions as an independent module: it specifically shifts nucleosome core mobility, elevates thermal denaturation of cores, protects the same DNase I sites as intact protein, and requires histone tails for binding. Mobility shift assays, thermal denaturation, DNase I digestion, trypsin digestion of histone tails with synthetic peptides and recombinant protein Journal of molecular biology High 1453455
1998 HMGN2 (HMG-17) is released from chromatin during mitosis (metaphase/anaphase), then re-imported into the nucleus in late telophase via active nuclear import requiring energy and importin alpha; the protein contains an intrinsic bipartite nuclear localization signal. Immunofluorescence colocalization through cell cycle, reconstituted nuclei and permeabilized cell import assays, energy depletion experiments The Journal of cell biology High 9852141
1998 HMGN2 (HMG-17) colocalizes with sites of active RNA polymerase II transcription; a peptide corresponding to the nucleosomal binding domain of HMG-14/17 displaces HMGN2 from chromatin and arrests pol II transcription; upon transcriptional inhibition, HMGN2 redistributes from chromatin to interchromatin granule clusters (SC35-positive). Immunofluorescence in tissue culture cells, peptide microinjection into permeabilized cells, alpha-amanitin and actinomycin D treatment The EMBO journal High 9843505
2007 HMGN2 (HMG-17) physically interacts with the PITX2 homeodomain protein to inhibit its DNA-binding activity; beta-catenin forms a ternary PITX2/HMGN2/beta-catenin complex that switches HMGN2 from a transcriptional repressor to an activator; HMG-17 homozygous knockout mice show early embryonic lethality. Co-immunoprecipitation, EMSA, reporter assays, pull-down, confocal microscopy, homozygous knockout mouse model Nucleic acids research High 18045789
2011 HMGN2 inducibly binds a novel transactivation domain in nuclear prolactin receptor (PRLr) in a phosphorylation-dependent manner; the HMGN2-PRLr interaction enables Stat5a-responsive promoter binding and transcriptional activation, promoting anchorage-independent growth. Co-immunoprecipitation, ChIP assays, reporter assays, soft agar anchorage-independent growth assay Molecular endocrinology High 21816901
2016 HDAC6 deacetylates HMGN2 at lysine K2; deacetylation of HMGN2 promotes Stat5a-mediated transcription and breast cancer growth; HDAC6 inhibition increases HMGN2 acetylation and reduces Stat5a signaling in vitro and in vivo. HDAC6 inhibitor treatment in vitro and in vivo, acetylation Western blot, Stat5a target gene expression, breast cancer growth assays Molecular cancer research High 27358110
2016 HMGN2 promotes STAT5 accessibility at prolactin-responsive promoter DNA by facilitating dissociation of linker histone H1; H1 knockdown rescues decreases in PRL-induced transcription caused by HMGN2 knockdown, demonstrating that HMGN2 acts antagonistically to H1 to enable STAT5 chromatin recruitment. siRNA knockdown of HMGN2 and H1, ChIP for STAT5 and H1, gene expression analysis, cell proliferation assays The Journal of biological chemistry High 28035005
2014 HMGN2 is SUMOylated at lysines K17 and K35 (within the nucleosome-binding domain) by SUMO E3 ligase PIAS1; SENP1 deSUMOylates HMGN2; SUMOylation decreases HMGN2 binding affinity to nucleosome core particles. In vitro SUMOylation assay, identification of SUMOylated residues by mutagenesis, SUMO1-conjugated HMGN2 reconstituted in E. coli, nucleosome binding assays The Journal of biological chemistry High 24872413
2009 HMGN2 is a component of the global genome repair subpathway of nucleotide excision repair (NER): DT40 cells lacking HMGN2 are hypersensitive to UV irradiation, show increased G2-M checkpoint arrest and apoptosis, and have slower removal of UV-induced DNA lesions from native chromatin, though nucleotide excision repair per se (measured by host cell reactivation) remains intact. Gene knockout in DT40 cells, UV sensitivity assays, apoptosis and cell cycle analysis, host cell reactivation assay, DNA lesion removal kinetics The FEBS journal High 19843163
2019 Loss of HMGN2 in pluripotent embryonal carcinoma cells leads to global reduction in H3K9 acetylation and disrupts H3K4me3, H3K9ac, H3K27ac, and H3K122ac at Nanog and Oct4 loci, causing loss of pluripotency markers and increased spontaneous neuronal differentiation. HMGN2 knockout cell lines, ChIP-seq for histone marks, gene expression analysis, differentiation marker assays Epigenetics & chromatin High 31831052
2021 Lysine succinylation at K30 within the HMGN2 nucleosome-binding domain (installed via a succinyl lysine analogue) significantly decreases HMGN2 binding to mononucleosomes and promotes nucleosomal DNA unwrapping at entry/exit regions, increasing DNA accessibility. Site-specific incorporation of succinyl lysine analogue, mononucleosome binding assays, nucleosomal DNA accessibility assays RSC chemical biology High 34458839
2013 Binding of HMGN2 to reconstituted nucleosomes increases the winding angle of nucleosomal DNA (measured by circular dichroism), with the extent of structural change differing from that of HMGN1, suggesting distinct abilities to facilitate nucleosome remodeling. CD spectroscopy of nucleosomes reconstituted from unmodified recombinant histones and synthetic positioning DNA, with purified HMGN1 and HMGN2 FEBS open bio Medium 23772392
2011 HMGN2 promotes LPS-induced beta-defensin-2 expression in A549 cells by prolonging nuclear retention of NF-κB p65, enhancing p65 acetylation, increasing histone acetyltransferase activity, and promoting p65-Ser536 phosphorylation; ChIP shows HMGN2 and p65 synergistically bind the HBD-2 promoter. siRNA knockdown, ChIP, Western blot for p65 nuclear retention and phosphorylation, HAT activity assay, reporter assay The FEBS journal Medium 21518253
2022 HMGN2 interacts with transcription factor Lef-1 through its HMG-box domain and with Dlx2, FoxJ1, and Pitx2; HMGN2 binding to Lef-1 inhibits Lef-1 DNA-binding activity; HMGN2 associates with H4K5ac and H3K4me2 marks at the Dlx2 promoter; miR-23a/b directly target Hmgn2 to post-transcriptionally suppress its expression; Hmgn2 ablation in mice increases amelogenin expression due to increased transcriptional activity of Pitx2, Dlx2, Lef-1, and FoxJ1. Bimolecular fluorescence complementation, pull-down, co-immunoprecipitation, EMSA, ChIP, mouse knockout, in situ hybridization, luciferase reporter assays The Journal of biological chemistry High 35872015
2025 HMGN1 and HMGN2 preferentially bind nucleosomes with acetylated H3 tail residues and H2A.Z-containing nucleosomes; HMGN1/HMGN2 binding to nucleosomes reduces p300-mediated acetylation of H3K18, H3K23, and H3K27; Hmgn1/Hmgn2 double-knockout mESCs show increased H3K27me2/me3 and altered expression of ~1000 genes. Nucleosome binding assays, in vitro p300 acetylation assays, epiproteomic mass spectrometry, RNA-seq in CRISPR-engineered knockout mESCs The Journal of biological chemistry High 41325801
1987 HMGN2 (HMG17) is present only downstream of transcription start sites in active chromatin of vitellogenin II, lysozyme, and ovalbumin genes, as demonstrated by immunofractionation of oligonucleosomes with monoclonal antibodies and analysis by two independent experimental approaches. Monoclonal antibody immunoisolation of oligonucleosomes, gene-specific DNA probe analysis of active chromatin from chicken liver and oviduct The EMBO journal High 3665881
1984 HMG17 (HMGN2) greatly facilitates catenation of double-stranded DNA rings by DNA topoisomerases types I and II, promoting formation of large catenated networks even at low DNA concentrations where catenanes are otherwise absent. In vitro topoisomerase catenation assay, gel electrophoresis, electron microscopy, restriction enzyme cleavage Archives of biochemistry and biophysics Medium 6326673
1997 HMGN2 (HMG-17) nucleosomes are organized in clusters of ~6 contiguous HMG-17-containing nucleosomes in cellular chromatin, and nucleosomes containing HMG-17 are devoid of HMG-14 (the two segregate into distinct nuclear domains), as shown by immunofractionation and confocal immunofluorescence. Confocal immunofluorescence microscopy, immunofractionation with affinity-purified antibodies, quantitative analysis Journal of molecular biology Medium 9417927
2002 Butyrate-induced hyperacetylation of HMGN2 alters its binding properties to chromatin in HT29 colon adenocarcinoma cells: acetylated HMGN2 is solubilized from chromatin, as demonstrated by FRAP of HMG-N2-EGFP fusion proteins and solubilization of endogenous acetylated HMGN2 in permeabilized cells. Butyrate treatment, transfected EGFP fusion protein dynamics, solubilization of acetylated HMGN2 from permeabilized cells, Northern and Western blot International journal of cancer Medium 11807779
2024 HMGN2 binds to histones and promotes stability of H3K27ac at the CDC20 promoter region, enhancing CDC20 transcriptional activity and accelerating glioma cell proliferation; HMGN2 knockout reduces CDC20 expression and cell cycle progression. ChIP for H3K27ac at CDC20 promoter, HMGN2 knockdown/knockout, gene expression and cell proliferation assays Genes & diseases Medium 40092489
2025 SUMOylated HMGN2 (via PIAS1) enhances interaction with transcription factor PAX5, inhibiting PAX5 activity and driving macrophage M1 polarization; PIAS1 knockdown reduces HMGN2 SUMOylation, restoring PAX5-mediated M2 polarization and suppressing NF-κB inflammatory signaling in atherosclerosis. Co-immunoprecipitation, PIAS1 knockdown, foam macrophage model, ApoE-/- mouse model, ELISA, Western blot, immunofluorescence Experimental cell research Medium 40834970
2005 HMGN2 exhibits antimicrobial activity against Gram-negative bacteria and Candida; the alpha-helical domain (residues 18-48) is essential and sufficient for antimicrobial activity, while N-terminal and C-terminal fragments are inactive. In vitro antimicrobial assays (MIC, MBC) with recombinant full-length and domain fragments, synthetic peptides Acta pharmacologica Sinica Medium 16115376
2017 HMGN2 facilitates Nrf2 nuclear translocation upon pyocyanin stimulation in A549 cells, elevating antioxidant gene expression and reducing ROS; HMGN2 also regulates actin cytoskeleton rearrangement to attenuate Pseudomonas aeruginosa internalization via ROS elimination. HMGN2 knockdown/overexpression, ROS measurement, Nrf2 nuclear fractionation, actin staining, bacterial invasion assays Free radical biology & medicine Medium 28408162
2019 HMGN2 deficiency in macrophages promotes M1 polarization during NTM infection by enhancing activation of NF-κB and MAPK signaling, leading to increased iNOS, IFNγ, TNF-α, IL-1β, IL-6 expression and NO production, which affects NTM survival. siRNA knockdown, macrophage polarization marker analysis, NF-κB/MAPK signaling assays, intracellular bacterial survival assay Journal of cellular and molecular medicine Medium 31596045
2012 HMGN2 acts as a positive regulator of LPS-induced mouse beta-defensin-3 and -4 expression in vivo; HMGN2 shRNA interference reduces mBD-3 and mBD-4 but not mBD-1 expression in maternal, embryonic, and neonatal mouse tissues upon LPS stimulation. shRNA knockdown in vivo in pregnant ICR mice, real-time PCR for defensin and HMGN2 expression across developmental timepoints and tissues Inflammation Medium 21594618
2016 HMGN2 knockdown in A549 cells increases α5β1 integrin expression on cell membranes, talin expression, FAK and Src phosphorylation, and actin polymerization, resulting in increased Klebsiella pneumoniae internalization. siRNA knockdown, cDNA microarray, flow cytometry for integrin surface expression, phospho-Western blot, bacterial invasion assay International journal of molecular medicine Medium 27460641
2025 HMGN2 knockout in RAW264.7 macrophages enhances bactericidal and phagocytic activity by transcriptionally promoting CD14 expression via increased H3K4me3, H3K9ac, and H3K27ac at the CD14 gene promoter, activating CD14-mediated MAPK signaling and NO production. CRISPR-Cas9 knockout, ChIP for histone marks at CD14 promoter, bacterial killing assays, MAPK signaling analysis Frontiers in immunology Medium 41246296

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2002 A fragment of the HMGN2 protein homes to the nuclei of tumor cells and tumor endothelial cells in vivo. Proceedings of the National Academy of Sciences of the United States of America 205 12032302
1982 Autoantibodies to nucleosomal proteins: antibodies to HMG-17 in autoimmune diseases. Science (New York, N.Y.) 84 6460317
1999 Specific acetylation of chromosomal protein HMG-17 by PCAF alters its interaction with nucleosomes. Molecular and cellular biology 80 10207070
1987 Chromatin from transcribed genes contains HMG17 only downstream from the starting point of transcription. The EMBO journal 73 3665881
1995 HMG17 is a chromatin-specific transcriptional coactivator that increases the efficiency of transcription initiation. Genes & development 69 7649479
1992 Nucleosome core binding region of chromosomal protein HMG-17 acts as an independent functional domain. Journal of molecular biology 66 1453455
1986 Chromosomal protein HMG-17. Complete human cDNA sequence and evidence for a multigene family. The Journal of biological chemistry 65 3754870
1994 The footprint of chromosomal proteins HMG-14 and HMG-17 on chromatin subunits. Journal of molecular biology 60 8107104
1993 Deposition of chromosomal protein HMG-17 during replication affects the nucleosomal ladder and transcriptional potential of nascent chromatin. The EMBO journal 54 8404854
1998 Chromosomal proteins HMG-14 and HMG-17 are released from mitotic chromosomes and imported into the nucleus by active transport. The Journal of cell biology 52 9852141
1998 Dynamic relocation of chromosomal protein HMG-17 in the nucleus is dependent on transcriptional activity. The EMBO journal 51 9843505
1998 Stimulation of replication efficiency of a chromatin template by chromosomal protein HMG-17. The Journal of biological chemistry 50 9545265
1986 Isolation of oligonucleosomes from active chromatin using HMG17-specific monoclonal antibodies. Nucleic acids research 49 3703677
1997 Clusters of nucleosomes containing chromosomal protein HMG-17 in chromatin. Journal of molecular biology 48 9417927
2011 Expression of HMGB1 and HMGN2 in gingival tissues, GCF and PICF of periodontitis patients and peri-implantitis. Archives of oral biology 46 21570059
1986 Immunofractionation of DNA sequences associated with HMG-17 in chromatin. Experimental cell research 46 3743668
1983 Affinity of HMG17 for a mononucleosome is not influenced by the presence of ubiquitin-H2A semihistone but strongly depends on DNA fragment size. Nucleic acids research 45 6298725
1997 H1 and HMG17 extracted from calf thymus nuclei are efficient DNA carriers in gene transfer. Gene therapy 40 9231075
2007 Chromatin-associated HMG-17 is a major regulator of homeodomain transcription factor activity modulated by Wnt/beta-catenin signaling. Nucleic acids research 39 18045789
1979 The complete amino-acid sequence of a trout-testis non-histone protein, H6, localized in a subset of nucleosomes and its similarity to calf-thymus non-histone proteins HMG-14 and HMG-17. European journal of biochemistry 39 456349
1981 Immunochemical detection of chromosomal protein HMG-17 in chromatin subunits. Biochemistry 37 6452161
1987 Retropseudogenes for human chromosomal protein HMG-17. Journal of molecular biology 36 3441004
2016 HDAC6 Deacetylates HMGN2 to Regulate Stat5a Activity and Breast Cancer Growth. Molecular cancer research : MCR 35 27358110
2005 HMGN2: a novel antimicrobial effector molecule of human mononuclear leukocytes? Journal of leukocyte biology 35 16204630
1991 Recombinant human chromosomal proteins HMG-14 and HMG-17. Nucleic acids research 31 2057367
1989 Human non-histone chromosomal protein HMG-17: identification, characterization, chromosome localization and RFLPs of a functional gene from the large multigene family. Nucleic acids research 30 2565024
2016 Histone H1 and Chromosomal Protein HMGN2 Regulate Prolactin-induced STAT5 Transcription Factor Recruitment and Function in Breast Cancer Cells. The Journal of biological chemistry 29 28035005
1981 Nonhistone chromatin proteins HMG-14 and HMG-17 bind preferentially to single-stranded DNA. Nucleic acids research 29 7279673
1992 Autoantibodies to the chromosomal protein HMG-17 in juvenile rheumatoid arthritis. Arthritis and rheumatism 28 1567496
1980 The isolation, characterization and partial sequences of the chicken erythrocyte non-histone chromosomal proteins HMG14 and HMG17. Comparison with the homologous calf thymus proteins. The Biochemical journal 28 7396821
1993 Identification and genetic mapping of the murine gene and 20 related sequences encoding chromosomal protein HMG-17. Mammalian genome : official journal of the International Mammalian Genome Society 26 8094303
2019 Maintenance of active chromatin states by HMGN2 is required for stem cell identity in a pluripotent stem cell model. Epigenetics & chromatin 25 31831052
1986 Chromosomal proteins HMG-14 and HMG-17. Distinct multigene families coding for similar types of transcripts. The Journal of biological chemistry 25 3782108
1984 HMG17 protein facilitates the DNA catenation reaction catalyzed by DNA topoisomerases. Archives of biochemistry and biophysics 25 6326673
2002 Modulation of HMG-N2 binding to chromatin by butyrate-induced acetylation in human colon adenocarcinoma cells. International journal of cancer 24 11807779
1987 Cell cycle regulated synthesis of an abundant transcript for human chromosomal protein HMG-17. Nucleic acids research 24 3575100
1988 Single copy gene for the chicken non-histone chromosomal protein HMG-17. The Journal of biological chemistry 23 2831214
2011 HMGN2 inducibly binds a novel transactivation domain in nuclear PRLr to coordinate Stat5a-mediated transcription. Molecular endocrinology (Baltimore, Md.) 21 21816901
2001 HMG-17 is an early marker of inductive interactions in the developing mouse kidney. Differentiation; research in biological diversity 20 11683498
1998 HMG-17, a chromosomal non-histone protein, shows developmental regulation during organogenesis. The International journal of developmental biology 20 9727833
1977 Physicochemical studies of non-histone protein HMG17 with DNA. Biochimica et biophysica acta 20 911836
2019 HMGN2 regulates non-tuberculous mycobacteria survival via modulation of M1 macrophage polarization. Journal of cellular and molecular medicine 19 31596045
2009 Inhibitory effect of HMGN2 protein on human hepatitis B virus expression and replication in the HepG2.2.15 cell line. Antiviral research 19 19150374
2017 Nuclear protein HMGN2 attenuates pyocyanin-induced oxidative stress via Nrf2 signaling and inhibits Pseudomonas aeruginosa internalization in A549 cells. Free radical biology & medicine 18 28408162
2014 HMGN2, a new anti-tumor effector molecule of CD8⁺ T cells. Molecular cancer 18 25060707
2018 Effect of HMGN2 on proliferation and apoptosis of MCF-7 breast cancer cells. Oncology letters 17 30655878
2009 The nucleosome-binding protein HMGN2 modulates global genome repair. The FEBS journal 17 19843163
1988 Chicken chromosomal protein HMG-14 and HMG-17 cDNA clones: isolation, characterization and sequence comparison. Gene 15 3384337
2014 High mobility group nucleosomal binding domain 2 (HMGN2) SUMOylation by the SUMO E3 ligase PIAS1 decreases the binding affinity to nucleosome core particles. The Journal of biological chemistry 14 24872413
2013 Nucleosome-binding protein HMGN2 exhibits antitumor activity in oral squamous cell carcinoma. Oncology letters 14 24348831
2014 Nucleosome-binding protein HMGN2 exhibits antitumor activity in human SaO2 and U2-OS osteosarcoma cell lines. Oncology reports 13 25530340
1990 Mapping the human gene coding for chromosomal protein HMG-17. Human genetics 13 2394451
2012 The chromosomal protein HMGN2 mediates the LPS-induced expression of β-defensins in mice. Inflammation 12 21594618
2011 High-mobility group protein N2 (HMGN2) inhibited the internalization of Klebsiella pneumoniae into cultured bladder epithelial cells. Acta biochimica et biophysica Sinica 12 21778192
1997 Neither HMG-14a nor HMG-17 gene function is required for growth of chicken DT40 cells or maintenance of DNaseI-hypersensitive sites. Nucleic acids research 12 9016555
1993 Autoantibodies to HMG-17 nucleosomal protein in autoimmune rheumatic diseases. Correlation with systemic lupus erythematosus clinical activity and with antibodies to double-stranded DNA. Arthritis and rheumatism 12 8318042
2011 The chromosomal protein HMGN2 mediates lipopolysaccharide-induced expression of β-defensins in A549 cells. The FEBS journal 11 21518253
1995 The chicken HMG-17 gene is dispensable for cell growth in vitro. Molecular and cellular biology 11 7565703
1991 Differentiation-dependent alteration in the chromatin structure of chromosomal protein HMG-17 gene during erythropoiesis. Journal of molecular biology 11 1988681
1982 Effect of high mobility group nonhistone proteins HMG-20 (ubiquitin) and HMG-17 on histone deacetylase activity assayed in vitro. Nucleic acids research 11 6280157
2018 Non-histone nuclear protein HMGN2 differently regulates the urothelium barrier function by altering expression of antimicrobial peptides and tight junction protein genes in UPEC J96-infected bladder epithelial cell monolayer. Acta biochimica Polonica 10 29549670
2013 Nucleosome structural changes induced by binding of non-histone chromosomal proteins HMGN1 and HMGN2. FEBS open bio 10 23772392
1999 Deletion of HMG17 in uterine leiomyomas with ring chromosome 1. Cancer genetics and cytogenetics 10 9973936
2022 HMGN2 represses gene transcription via interaction with transcription factors Lef-1 and Pitx2 during amelogenesis. The Journal of biological chemistry 9 35872015
2018 HMGN2: An Antitumor Effector Molecule of γδT Cells. Journal of immunotherapy (Hagerstown, Md. : 1997) 9 29401165
2011 Expression of HMGB1 and HMGN2 in gingival tissues, GCF and PICF of periodontitis patients and peri-implantitis. Brazilian journal of microbiology : [publication of the Brazilian Society for Microbiology] 9 24031744
1994 Autoantibodies to HMG-17 nucleosomal protein in patients with scleroderma. Journal of autoimmunity 9 8037838
2021 Prolactin Drives a Dynamic STAT5A/HDAC6/HMGN2 Cis-Regulatory Landscape Exploitable in ER+ Breast Cancer. Endocrinology 8 33589921
2016 Knockdown of HMGN2 increases the internalization of Klebsiella pneumoniae by respiratory epithelial cells through the regulation of α5β1 integrin expression. International journal of molecular medicine 8 27460641
2005 Alpha-helical domain is essential for antimicrobial activity of high mobility group nucleosomal binding domain 2 (HMGN2). Acta pharmacologica Sinica 8 16115376
1990 Expression of chromosomal proteins HMG-14 and HMG-17 in transformed human cells. Cancer research 8 2317791
2020 Human HMGN1 and HMGN2 are not required for transcription-coupled DNA repair. Scientific reports 7 32152397
1993 Isolation by a new method and sequence analysis of chromosomal HMG-17 protein from porcine thymus. Archives of biochemistry and biophysics 7 8512325
2022 Recombinant jurkat cells (HMGN2-T cells) secrete cytokines and inhibit the growth of tumor cells. Journal of molecular histology 6 35861945
2021 Lysine succinylation on non-histone chromosomal protein HMG-17 (HMGN2) regulates nucleosomal DNA accessibility by disrupting the HMGN2-nucleosome association. RSC chemical biology 6 34458839
2014 HMGN2 protein inhibits the growth of infected T24 cells in vitro. Journal of cancer research and therapeutics 6 25022381
2017 Evidence For Hmgn2 Involvement in Mouse Embryo Implantation and Decidualization. Cellular physiology and biochemistry : international journal of experimental cellular physiology, biochemistry, and pharmacology 5 29216626
1984 Conformation of the HMG17-nucleosome complex. Biochimica et biophysica acta 5 6477923
2020 Exogenous HMGN2 inhibits the migration and invasion of osteosarcoma cell lines. Translational cancer research 4 35117527
2004 Content of the HMG-17 chromosomal protein in porcine tissues. Protein and peptide letters 4 15579118
1994 Sera from JRA patients contain antibodies against a defined epitope in chromosomal protein HMG-17. Autoimmunity 4 7517709
1990 Protein HMG-17 is hyper-expressed in rat glucagonoma. Single-step isolation and sequencing. European journal of biochemistry 4 2169420
2024 HMGN2 accelerates the proliferation and cell cycle progression of glioblastoma by regulating CDC20 expression. Genes & diseases 3 40092489
2023 HMGN2 and Histone H1.2: potential targets of a novel probiotic mixture for seasonal allergic rhinitis. Frontiers in microbiology 3 37869664
2012 Ectopic expression of Hmgn2 antagonizes mouse erythroid differentiation in vitro. Cell biology international 3 21988615
2002 Rabbit anti-HMG-17 antibodies recognize similar epitopes on the HMG-17 molecule as lupus autoantibodies. Relation with histone H1 defined epitopes. Journal of peptide science : an official publication of the European Peptide Society 3 12523645
2005 [E. coli-based production of recombinant HMG-17 and its antibacterial domain]. Sheng wu yi xue gong cheng xue za zhi = Journal of biomedical engineering = Shengwu yixue gongchengxue zazhi 1 16156270
1996 Chromosomal proteins HMG-14 and HMG-17 are synthesized throughout the S-phase in Burkitt's lymphoma. Biochemical and biophysical research communications 1 8670211
1990 Expression of human chromosomal proteins HMG-14 and HMG-17 in Saccharomyces cerevisiae. Experimental cell research 1 2226652
2026 Antitumor effects of STING agonists on nervous system tumors via tumor-intrinsic STING-STAT1-mediated HMGN2 expression. Cancer biology & medicine 0 41700809
2025 Inhibition of HMGN2 SUMOylation ameliorates atherosclerosis by activating PAX5 expression to induce macrophage M2 polarization. Experimental cell research 0 40834970
2025 Deficiency of HMGN2 enhances antibacterial activity of macrophages by promoting H3 histone modification-mediated CD14/iNOS expression. Frontiers in immunology 0 41246296
2025 HMGN1 and HMGN2 are recruited to acetylated and histone variant H2A.Z-containing nucleosomes to regulate chromatin state and transcription. The Journal of biological chemistry 0 41325801
2024 An evolutionarily distinct Hmgn2 variant influences shape recognition in Medaka Fish. Communications biology 0 39179658
2016 [The Role of HMGN2 in the Development of Periodontitis Dental Plaque]. Sichuan da xue xue bao. Yi xue ban = Journal of Sichuan University. Medical science edition 0 28591948
2011 [Inhibitory effect of recombinant HMGN2 protein on human hepatitis B viral]. Sichuan da xue xue bao. Yi xue ban = Journal of Sichuan University. Medical science edition 0 21866626
2010 [Isolation and purification of antimicrobial polypeptide HMGN2 from human lymph node and analysis of its distribution]. Sheng wu yi xue gong cheng xue za zhi = Journal of biomedical engineering = Shengwu yixue gongchengxue zazhi 0 20842856
2005 [Production of HMGN2 polyclonal antibody by immunization with recombinant GST-HMGN2 fusion protein and its application to analysis of HMGN2 distribution in human monocytes]. Sichuan da xue xue bao. Yi xue ban = Journal of Sichuan University. Medical science edition 0 16078558
2005 [Application of HMGN2-tag constructs to analysis of HMGN2 distribution in HeLa cells]. Sheng wu yi xue gong cheng xue za zhi = Journal of biomedical engineering = Shengwu yixue gongchengxue zazhi 0 16294743
2000 Regional fine mapping of HMG17 to chromosomal band 1p35. Cancer genetics and cytogenetics 0 10640150