Affinage

Showing POU3F2 — BRN2 is a alias.

POU3F2

POU domain, class 3, transcription factor 2 · UniProt P20265

Length
443 aa
Mass
46.9 kDa
Annotated
2026-06-10
95 papers in source corpus 48 papers cited in narrative 48 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 10/10 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

POU3F2 (BRN2) is a class III POU-domain transcription factor that acts as a master regulator of neural lineage specification and neuronal differentiation, and is co-opted in cancer as a driver of phenotype switching and invasion (PMID:8274283, PMID:25991548, PMID:29990501). In the nervous system it is required at a penultimate step of hypothalamic neurosecretory neuron differentiation, enabling activation of neuropeptide genes including vasopressin and oxytocin (PMID:8543156, PMID:8543155), and it sits downstream of SIM1 to control oxytocin expression (PMID:26833329). In cortical and stem-cell neurogenesis it governs progenitor proliferative capacity and the switch from direct to indirect neurogenesis, suppressing the Notch effector Hes5 while promoting Tbr2/Tbr1 and acting through direct targets including Zic1, NTF3, SOX2, SOX13 and ADNP (PMID:22892427, PMID:25991548, PMID:29549646, PMID:35245119, PMID:40498903, PMID:39271675). In Schwann cells it functions redundantly with Oct-6 to drive the promyelinating-to-myelinating transition by directly activating a Krox20 enhancer in synergy with Sox10 (PMID:12782656, PMID:16311519). Mechanistically, BRN2 binds DNA through its POU domain, can homodimerize on palindromic MORE-type elements, and its activity is tuned by phosphorylation within the POU domain (T361/S362), which switches its transcriptional output between proliferative and migratory programs (PMID:14741405, PMID:22290434). In melanoma, BRN2 integrates BRAF/MAPK, Wnt/β-catenin, PI3K-Pax3, and CDKN2A-E2F1 inputs and occupies thousands of genomic loci, where it directly represses MITF and CDH13 and activates an invasive program partly through an NFIB-EZH2 axis and KIT-ligand signaling (PMID:15024080, PMID:15024079, PMID:22988297, PMID:29990501, PMID:18829533, PMID:23069940, PMID:28119061). The BRN2/PAX3 balance forms a rheostat setting MITF plasticity during MAPK inhibition, and BRN2 also induces PTEN to restrain PI3K signaling, giving it a context-dependent tumor-suppressive role (PMID:30277012, PMID:34140478). Beyond transcription, BRN2 performs a non-transcriptional function at sites of DNA damage, binding PARP1 and Ku70/Ku80, promoting Ku80 recruitment and error-prone non-homologous end-joining while suppressing apoptosis (PMID:30804224). BRN2 is also required for neuroendocrine differentiation programs in prostate and small-cell lung cancers downstream of androgen receptor suppression (PMID:27784708, PMID:23530560). Loss-of-function POU3F2 mutations are found in individuals with autism spectrum disorder (PMID:40498903).

Mechanistic history

Synthesis pass · year-by-year structured walk · 20 steps
  1. 1995 High

    Established BRN2 as a genetically required factor for terminal differentiation of hypothalamic neurosecretory neurons, defining its developmental role in vivo.

    Evidence Germline gene knockout in mice with histology and neuropeptide gene expression analysis

    PMID:8543155 PMID:8543156

    Open questions at the time
    • Direct transcriptional targets at the neuropeptide loci not defined
    • Did not establish whether the requirement is cell-autonomous
  2. 1993 Medium

    Showed BRN2 is required to establish neural cell lineages rather than merely refine them, since its loss diverts embryonal carcinoma cells to non-neural fates.

    Evidence Antisense RNA inhibition in P19 cells with lineage marker analysis

    PMID:8274283

    Open questions at the time
    • No direct target genes identified
    • Antisense specificity not orthogonally confirmed
  3. 2003 High

    Defined BRN2 as functionally redundant with Oct-6 in driving Schwann cell myelination, resolving how POU-III factors cooperate in peripheral glia.

    Evidence Transgenic overexpression rescue in Oct-6 knockout and compound knockout in mice

    PMID:12782656

    Open questions at the time
    • Did not identify the direct enhancer targets driving the transition (addressed later)
  4. 2006 High

    Placed BRN2 directly upstream of Krox20 in the myelination hierarchy by identifying a bound enhancer element and Sox10 synergy.

    Evidence Reporter assays, transgenic enhancer analysis and TF binding studies

    PMID:16311519

    Open questions at the time
    • Stoichiometry of BRN2/Oct-6/Sox10 occupancy not resolved
  5. 2000 Medium

    Defined BRN2's biochemical interaction repertoire, showing homodimerization via homeodomain/linker and low-affinity contacts with general and melanocyte transcription factors.

    Evidence GAL4-VP16 two-hybrid, GST-pulldown and EMSA with melanoma nuclear extracts

    PMID:11029584

    Open questions at the time
    • Functional consequence of TBP/p300/Sox10/Pax3 contacts not established
    • In vitro affinities not validated in cells
  6. 2004 High

    Identified BRN2 as a downstream effector of both BRAF/MAPK and Wnt/β-catenin signaling driving melanoma proliferation, integrating two oncogenic inputs at a single transcription factor.

    Evidence Promoter reporter assays, siRNA knockdown and transgenic mice with proliferation assays

    PMID:15024079 PMID:15024080

    Open questions at the time
    • Downstream proliferative targets of BRN2 not enumerated in these studies
  7. 2008 Medium

    Revealed BRN2 as a regulator of melanoma heterogeneity by directly repressing MITF, with the two factors marking mutually exclusive tumor subpopulations.

    Evidence Promoter reporter assays, siRNA knockdown and biopsy immunohistochemistry

    PMID:18829533

    Open questions at the time
    • Mechanism of switching between BRN2-high and MITF-high states not defined
  8. 2010 Medium

    Mapped BRN2's genome-wide occupancy in melanoma (~2108 loci), establishing it as a broad regulator including direct control of MITF and KIT-ligand signaling.

    Evidence ChIP-chip occupancy profiling with luciferase reporter validation

    PMID:20337985

    Open questions at the time
    • Functional consequence validated only for Kitl
    • ChIP-chip lower resolution than later ChIP-seq
  9. 2012 High

    Showed BRN2 transcriptional output is switched by POU-domain phosphorylation at T361/S362, explaining how one factor toggles between proliferation and migration programs.

    Evidence Point mutagenesis with transgenic mouse and in vitro phenotypic assays

    PMID:22290434

    Open questions at the time
    • Kinase responsible for T361/S362 phosphorylation not identified
    • Structural basis of the phospho-switch unresolved
  10. 2012 Medium

    Connected upstream PI3K-Pax3 signaling and downstream CDH13 repression to BRN2, extending its melanoma invasion circuitry.

    Evidence Promoter binding/EMSA, PI3K inhibition, siRNA knockdown and invasion assays

    PMID:22988297 PMID:23069940

    Open questions at the time
    • Cross-talk between PI3K-Pax3 and MAPK inputs not integrated
  11. 2013 Medium

    Established BRN2 as a high-level driver of neuroendocrine differentiation in small-cell lung cancer, upstream of ASCL1/ND1 and directly activating TTF1.

    Evidence siRNA knockdown, ectopic expression, ChIP and cell-cycle assays in SCLC cells

    PMID:23358112 PMID:23530560

    Open questions at the time
    • Direct vs indirect regulation of ASCL1/ND1 not fully separated
  12. 2015 High

    Used integrated ChIP-seq/RNA-seq to define a direct BRN2-Zic1 axis required for neuronal fate specification in differentiating ESCs.

    Evidence shRNA knockdown with RNA-seq, ChIP-seq and differentiation assays

    PMID:25991548

    Open questions at the time
    • Zic1 may be one of several effectors; full direct target set not parsed
  13. 2016 Medium

    Embedded BRN2 in a β-catenin/Brn2/Tbr2 cascade controlling basal progenitor expansion and linked its deregulation to behavioral abnormalities rescuable by Wnt activation.

    Evidence Dvl mutant mouse model with pharmacological Wnt activation and SIM1-epistasis zebrafish models

    PMID:26830142 PMID:26833329

    Open questions at the time
    • Direct vs indirect position of BRN2 within the β-catenin/Tbr2 cascade not fully resolved
  14. 2017 Medium

    Identified the BRN2-NFIB-EZH2-MITF axis as a mechanism by which BRN2 epigenetically suppresses MITF to promote invasive, low-proliferative melanoma states.

    Evidence Overexpression/siRNA knockdown with migration/invasion and expression assays

    PMID:28119061

    Open questions at the time
    • Direct vs indirect induction of NFIB not distinguished
  15. 2018 High

    Placed BRN2 downstream of androgen-receptor suppression and CDKN2A-E2F1, establishing it as a driver of neuroendocrine prostate differentiation and melanoma metastasis initiation.

    Evidence ChIP of AR/E2F1 at BRN2 promoter, CRISPR-engineered primary cells, xenograft and in vivo rescue

    PMID:27784708 PMID:29990501 PMID:30277012

    Open questions at the time
    • How BRN2 selects invasion vs differentiation programs across tissues not defined
  16. 2019 High

    Uncovered a non-transcriptional role for BRN2 at DNA damage sites, where it binds PARP1 and Ku70/80 to promote error-prone NHEJ and suppress apoptosis.

    Evidence Reciprocal Co-IP, live-cell damage recruitment, NHEJ/HR repair assays and apoptosis assays

    PMID:30804224

    Open questions at the time
    • Whether DNA-binding domain or POU domain mediates the repair function unclear
    • Link between transcriptional and repair functions not integrated
  17. 2021 High

    Revealed BRN2's behavior on mitotic chromatin, binding condensed chromosomes non-sequence-specifically to enable early transcriptional reactivation of targets like Nestin during mitotic exit.

    Evidence Live-cell imaging, ChIP-seq, single-molecule RNA-FISH and mitotic dominant-negative

    PMID:34168041

    Open questions at the time
    • How non-specific mitotic binding transitions to specific G1 occupancy unresolved
  18. 2021 Medium

    Demonstrated a context-dependent tumor-suppressive function whereby BRN2 directly induces PTEN to restrain PI3K signaling, opposing its pro-invasive activity.

    Evidence Conditional knockout mouse models with promoter binding and PI3K signaling analysis

    PMID:34140478

    Open questions at the time
    • Determinants of pro- versus anti-tumor BRN2 output not defined
  19. 2024 High

    Extended BRN2's neurogenic function to primate and cross-species cortical development, controlling radial glia expansion and the direct-to-indirect neurogenesis switch via SOX2/STAT3/HOPX and NOTCH interplay.

    Evidence Biallelic CRISPR knockout in cynomolgus monkeys and conditional knockouts in mice/ferrets with single-cell transcriptomics

    PMID:35245119 PMID:39271675

    Open questions at the time
    • Primate-specific regulatory mechanisms not fully dissected at the cis level
  20. 2025 Medium

    Linked POU3F2 loss-of-function to autism via reduced Wnt signaling and premature radial glia specification, identifying SOX13 and ADNP as direct mediating targets.

    Evidence CRISPR disruption in human NPCs with RNA-seq, ChIP-seq and Wnt reporter assays

    PMID:40498903

    Open questions at the time
    • Genotype-phenotype correlation in patients not established
    • Single-lab human NPC model

Open questions

Synthesis pass · forward-looking unresolved questions
  • How BRN2 integrates its sequence-specific transcriptional, mitotic-bookmarking, and DNA-repair activities into a unified control of cell fate and survival remains unresolved.
  • No structural model linking POU-domain DNA binding to PARP1/Ku association
  • Kinases and signals that select among BRN2's divergent outputs are unmapped
  • Whether DNA-repair function operates outside melanoma is untested

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140110 transcription regulator activity 6 GO:0003677 DNA binding 4
Localization
GO:0005634 nucleus 3 GO:0000228 nuclear chromosome 1
Pathway
R-HSA-1266738 Developmental Biology 5 R-HSA-1643685 Disease 5 R-HSA-74160 Gene expression (Transcription) 4 R-HSA-162582 Signal Transduction 3 R-HSA-73894 DNA Repair 2
Partners
ARNT2NFYBPARP1PAX3PQBP1SOX10XRCC5XRCC6

Evidence

Reading pass · 48 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1995 Deletion of the Brn-2 genomic locus in mice results in failure of differentiation of paraventricular and supraoptic nucleus neurons, inability to activate neuropeptide-encoding genes (including vasopressin and oxytocin), and failure of correct axonal projections, demonstrating that Brn-2 is required at a penultimate step in hypothalamic neurosecretory neuron differentiation. Gene knockout (homologous recombination in ES cells), histology, gene expression analysis Genes & development High 8543155 8543156
1993 Antisense RNA-mediated inhibition of Brn-2 induction in P19 embryonal carcinoma cells blocks differentiation into neurons and astrocytes, causing cells to instead differentiate into non-neural cell types (smooth and skeletal muscle), demonstrating that Brn-2 is required for establishing neural cell lineages. Antisense RNA inhibition in P19 cells, morphological and lineage marker analysis Neuron Medium 8274283
2003 Brn-2 is expressed in Schwann cells with a developmental profile similar to Oct-6; overexpression of Brn-2 under the Oct-6 Schwann cell enhancer partially rescues the developmental delay in Oct-6-deficient Schwann cells; compound disruption of both Brn-2 and Oct-6 produces a much more severe myelination phenotype, demonstrating that Brn-2 function largely overlaps with Oct-6 in driving the promyelinating-to-myelinating transition. Transgenic overexpression rescue in Oct-6 knockout, compound knockout, developmental expression profiling Genes & development High 12782656
2006 Oct6 and Brn2 directly activate Krox20 expression in Schwann cells by binding to a defined Krox20 enhancer element during the promyelin-to-myelin transition; Sox10 synergizes with these POU proteins on this enhancer, placing Brn2 directly upstream of Krox20 in the myelination transcriptional hierarchy. Cell culture reporter assays, transgenic enhancer analysis, transcription factor binding studies EMBO reports High 16311519
2004 Brn-2 expression in melanoma is strongly upregulated downstream of Ras and MAPK signaling; the Brn-2 promoter is stimulated by kinase-activating BRAF mutants; siRNA-mediated depletion of BRAF reduces endogenous Brn-2 expression; and siRNA depletion of Brn-2 in BRAF-activated melanoma cells decreases proliferation, placing Brn-2 as a downstream effector of BRAF signaling in melanoma proliferation. Promoter reporter assays, siRNA knockdown, proliferation assays Molecular and cellular biology High 15024080
2004 Brn-2 expression is directly controlled by the Wnt/beta-catenin signaling pathway in melanoma cell lines and transgenic mice; siRNA inhibition of Brn-2 in beta-catenin-overexpressing melanoma cells results in decreased proliferation, identifying Brn-2 as a convergence point for BRAF and Wnt/beta-catenin signaling in melanoma proliferation. Promoter reporter assays, siRNA knockdown, transgenic mice, proliferation assays Molecular and cellular biology High 15024079
2008 Brn-2 directly represses Mitf expression in melanoma cells, and Brn-2 and Mitf mark distinct non-overlapping subpopulations within melanoma biopsies, demonstrating Brn-2 as a regulator of tumor heterogeneity through direct MITF repression. Promoter reporter assays, siRNA knockdown, immunohistochemistry of biopsies Cancer research Medium 18829533
1999 PQBP-1, a novel polyglutamine tract-binding protein, binds directly to the polyglutamine tract of Brn-2 and inhibits Brn-2-mediated transcriptional activation in cells. Protein identification by binding, co-transfection transcriptional assays, nuclear localization studies Human molecular genetics Medium 10332029
2000 Brn-2 homodimerizes in vivo; dimerization requires both the homeodomain and linker regions of the POU domain. Brn-2 also shows low-affinity interactions with TBP, the transcriptional coactivator p300, and with the melanocyte transcription factors Sox-10 and Pax-3. GAL4-VP16 two-hybrid assay, GST-pulldown, EMSA with melanoma nuclear extracts European journal of biochemistry Medium 11029584
2012 Brn-2 directly represses CDH13 (T-cadherin) promoter activity in melanoma cells; BRN2 binds to a specific element at -219 bp of the CDH13 promoter (5'-CATGCAAAA-3'); ectopic BRN2 suppresses CDH13 promoter activity, and BRN2 knockdown restores T-cadherin expression. Reporter gene assays, EMSA, siRNA knockdown, ectopic expression Laboratory investigation Medium 23069940
2012 PI3K signaling positively regulates Brn-2 expression in melanoma via Pax3; Pax3 directly binds and activates the Brn-2 promoter; PI3K inhibition reduces both Pax3 and Brn-2 expression and decreases melanoma invasiveness, placing Brn-2 downstream of a PI3K-Pax3 axis. PI3K inhibitor treatment, promoter binding assays, siRNA knockdown, invasion assays Molecular and cellular biology Medium 22988297
2012 Phosphorylation of BRN2 at T361 and S362 within the POU domain differentially regulates its transcriptional targets: phosphorylated BRN2 (BRN2TS) induces proliferation and represses migration and activates PAX3 transcription, while non-phosphorylatable BRN2 (BRN2AA) represses both proliferation and migration, and represses PAX3. Both forms repress MITF-M. Point mutagenesis, transgenic mouse models, in vitro cell assays, gene expression analysis Molecular and cellular biology High 22290434
2010 POU3F2/BRN2 occupies approximately 2108 genomic loci in melanoma cells including the MITF promoter and the Kit ligand (Kitl) proximal promoter; BRN2 regulates Kitl expression via a cluster of four closely spaced binding sites, indicating BRN2 can drive autocrine KIT ligand signaling. ChIP-chip genome-wide occupancy profiling, luciferase reporter assays Pigment cell & melanoma research Medium 20337985
2016 Androgen receptor (AR) directly suppresses BRN2 transcription in prostate cancer; loss of AR activity leads to elevated BRN2 expression; BRN2 is required for neuroendocrine prostate cancer (NEPC) and regulates the NEPC marker SOX2, placing BRN2 downstream of AR suppression as a driver of neuroendocrine differentiation. AR pathway inhibition models, ChIP assays demonstrating direct AR binding to BRN2 promoter, siRNA knockdown, in vitro and xenograft functional assays Cancer discovery High 27784708
2019 BRN2 associates with DNA damage response proteins PARP1 and Ku70/Ku80 (not transcriptional cofactors); BRN2 is rapidly recruited to DNA damage sites in a PARP1-dependent manner; BRN2 facilitates Ku80 recruitment and promotes non-homologous end joining (NHEJ) at the expense of homologous recombination; BRN2 also suppresses an apoptosis-associated gene expression program, protecting against UVB-, chemotherapy-, and vemurafenib-induced apoptosis. Co-immunoprecipitation, live-cell imaging of DNA damage recruitment, NHEJ/HR repair assays, gene expression analysis, apoptosis assays Genes & development High 30804224
2018 Loss of CDKN2A leads to increased BRN2 expression via E2F1; E2F1 directly regulates BRN2 transcription; BRN2 inhibition rescues metastatic dissemination caused by CDKN2A loss in human melanoma lines in mice, placing BRN2 downstream of CDKN2A/p16-E2F1 in melanoma invasion initiation. CRISPR-Cas9 engineering of primary human melanocytes, xenograft models, E2F1 promoter binding assays Cancer cell High 29990501
2011 BRN2 activates NOTCH pathway signaling in melanoma cells, while MITF represses it; siRNA-mediated depletion of BRN2 decreased NOTCH pathway member expression, and depletion of MITF increased it; loss of both BRN2 and MITF decreased melanosphere-forming capability, cell adhesion, and invasion. siRNA knockdown, gene expression analysis, melanosphere formation assay, invasion assay Oncogene Medium 21358674
2013 BRN2 is required for expression of ASCL1, ND1, and neuroendocrine markers (NCAM1, SYP, CHGA) in small cell lung cancer cells; BRN2 knockdown in SCLC cells causes significant growth retardation with decreased S-to-G2 phase population; forced BRN2 expression in non-SCLC cells induces expression of these neuroendocrine markers, placing BRN2 as a higher-level regulator than ASCL1 and ND1. siRNA knockdown, ectopic expression, cell cycle analysis, growth assays Pathology international Medium 23530560
2013 BRN2 directly activates TTF1 expression in SCLC cells by binding to the TTF1 isoform 2 promoter; BRN2 binding to the TTF1 promoter was confirmed by chromatin immunoprecipitation; BRN2 knockdown significantly reduced TTF1 expression. Promoter reporter assays, ChIP, siRNA knockdown Laboratory investigation Medium 23358112
2005 Brn-2 directly interacts with Jab1 (Jun-activation-domain-binding protein 1); interaction was identified by yeast two-hybrid and confirmed by surface plasmon resonance biosensor. Yeast two-hybrid screen, surface plasmon resonance biosensor Neuroscience letters Low 15911144
2001 Brn-2 activates the neuronal promoter of the aromatic L-amino acid decarboxylase (AADC) gene; NF-Y binding is required for Brn-2-mediated transactivation; Brn-2 directly interacts with NF-Y via its POU domain (GST-pulldown), and NF-Y and Brn-2 bind cooperatively to the AADC neuronal promoter. Reporter gene assays, dominant-negative NF-Y, GST-pulldown, EMSA Brain research. Molecular brain research Medium 11311976
2004 Brn-2 POU domain dimerizes cooperatively on a palindromic MORE+2-type element (TB motif) in the AADC neuronal promoter; site-directed mutagenesis of the POU-specific domain hydrophobic pocket and C-terminal POU homeodomain abolished dimerization on TB; both TB and ONF sites contribute to Brn-2-mediated promoter activation. EMSA with POU domain mutants, site-directed mutagenesis, reporter assays in COS-7 cells Brain research. Molecular brain research Medium 14741405
2017 BRN2 drives melanoma cell migration and invasion through induction of NFIB; NFIB in turn upregulates EZH2, which epigenetically suppresses MITF expression; this BRN2-NFIB-EZH2-MITF axis promotes an invasive, less proliferative melanoma phenotype. Overexpression and siRNA knockdown experiments, migration/invasion assays, gene expression analysis EBioMedicine Medium 28119061
2012 POU-III transcription factors Pou3f2 (Brn2) and Pou3f3 (Brn1) are expressed in ventricular zone progenitors and function to promote upper-layer cortical neuron fate; using an Engrailed dominant-negative repressor, sustained neurogenesis after the deep-to-upper-layer transition requires Pou3f activity; single-gene overexpression of any Pou3f in early neural progenitors is sufficient to specify precocious Satb2+ upper-layer neuron birth. Pou3fs suppress Notch effector Hes5 and promote Tbr2 and Tbr1 expression. Dominant-negative repressor, single-gene overexpression, marker analysis, in utero electroporation Cerebral cortex Medium 22892427
2015 Brn2 is essential for the neuronal differentiation program in retinoic acid-treated embryonic stem cells; integrated RNA-seq (after Brn2 silencing) and ChIP-seq identified Zic1 as a direct Brn2 target gene; shRNA silencing of Zic1 blocked neural precursor specification, defining a hierarchical Brn2-Zic1 axis specifying neuronal fate. shRNA knockdown, RNA-seq, ChIP-seq, differentiation assays Journal of cell science High 25991548
2016 A prenatal β-catenin/Brn2/Tbr2 transcriptional cascade governs basal neural progenitor cell expansion; in Dvl1/Dvl3 mutant mice with social and repetitive behavioral abnormalities, this cascade is deregulated; pharmacological Wnt activation rescues the transcriptional cascade and prevents adult behavioral deficits. Genetic mouse model (Dvl mutants), pharmacological Wnt activation, transcriptional cascade analysis Molecular psychiatry Medium 26830142
2016 POU3F2 lies downstream of SIM1 and controls oxytocin expression in the hypothalamic neuroendocrine preoptic area, as demonstrated using morpholino and mutant zebrafish models. Morpholino knockdown and mutant zebrafish models, oxytocin expression analysis American journal of human genetics Medium 26833329
2018 POU3F2 directly regulates NTF3 promoter activity; POU3F2 binds to the NTF3 promoter as demonstrated by ChIP-seq and mutation/deletion of the POU3F2 binding site in the NTF3 promoter decreased luciferase reporter activity; POU3F2 knockdown downregulates NTF3 expression; exogenous NTF3 rescues neuronal differentiation in POU3F2-knockdown cells. ChIP-seq, promoter luciferase reporter assays with binding site mutation, siRNA knockdown, rescue experiments Molecular neurobiology High 29549646
2018 POU3F2 regulates TRIM8 expression by binding to a SNP-containing promoter element (rs5011218); luciferase reporter and EMSA showed that POU3F2 binds this element and that the SCZ-associated SNP affects binding efficiency; POU3F2 or TRIM8 knockdown in neural progenitor cells promotes NPC proliferation, inhibits neuronal differentiation, and impairs excitatory synaptic transmission. Luciferase reporter assays, EMSA, RNA-seq after knockdown, functional NPC assays, electrophysiology Molecular psychiatry Medium 32929213
2021 Brn2 associates with condensed chromatin throughout cell division in neural stem cells (live-cell imaging); ChIP-seq shows Brn2 mitotic chromosome binding is not sequence-specific (relies mostly on electrostatic forces); competing with Brn2 binding during mitotic exit using a mitotic-specific dominant-negative reduces transcription of its target gene Nestin, demonstrating Brn2's role in early transcriptional reactivation during mitosis-to-G1 transition. Live-cell imaging, ChIP-seq, single-molecule RNA-FISH, mitotic dominant-negative approach Genes & development High 34168041
2010 Mutant huntingtin N-terminal fragment (from R6/2 HD mice) sequesters Brn-2, reducing its DNA binding; mutant huntingtin also reduces Brn-2 transcription; both mechanisms reduce Brn-2 function, leading to reduced hypothalamic neuropeptide expression. Brn-1, a related protein, is not sequestered. Comprehensive analysis of transcription factor DNA binding, protein interaction studies, R6/2 model mice Human molecular genetics Medium 20185558
2020 BRN2 directly binds PARP1 and Ku70/Ku80 and is rapidly recruited to DNA damage sites in a PARP1-dependent manner; BRN2 facilitates Ku80 recruitment and promotes non-homologous end-joining (NHEJ) at the expense of homologous recombination; this is associated with high somatic mutation burden in melanoma. Co-immunoprecipitation, live-cell imaging, NHEJ/HR repair pathway assays Genes & development High 30804224
2020 BRN2 expression increases anoikis resistance in melanoma cells; induction of BRN2 increases c-MET protein levels and STAT3 phosphorylation; treatment with c-MET inhibitors decreases viability of BRN2-expressing cells under non-adherent conditions. Doxycycline-inducible BRN2 expression, genome-wide profiling, non-adherent viability assays, c-MET inhibitor treatment Oncogenesis Medium 32632141
2021 BRN2 directly induces PTEN expression and represses PI3K signaling; BRN2 haplo-insufficiency in a BrafV600E/PtenF/+ context promotes melanoma initiation and metastasis; MITF (a BRN2 target) represses PTEN transcription, suggesting a tumor suppressive role for BRN2 via PTEN induction. Conditional knockout mouse models, promoter binding assays, PI3K signaling analysis Nature communications Medium 34140478
2018 BRAF regulates MITF expression through a BRN2/PAX3 rheostat: PAX3 activates MITF transcription while BRN2 represses it; the balance between BRN2 and PAX3 determines MITF expression plasticity, providing robustness during MAPK inhibitor treatment. Reporter assays, knockdown/overexpression, quantification of transcription factor levels Pigment cell & melanoma research Medium 30277012
2020 Cdk5 phosphorylates Tet3 at S1310 and S1379 in its catalytic domain; phosphorylated Tet3 generates higher 5hmC levels at the BRN2 promoter (compared to phospho-mutant Tet3), correlating with higher BRN2 expression and more efficient neuronal differentiation of ESCs; differential H2A.Z occupancy at BRN2 promoter also participates in this regulation. In vitro dioxygenase assay with phospho-mutant Tet3, 5hmC mapping at BRN2 promoter, ESC differentiation assays, Tet triple-KO rescue experiments Nucleic acids research Medium 31807777
2018 POU3F2 regulates a coexpression network module in brain; knockdown and overexpression experiments in neural cells validate POU3F2 as a regulator of hsa-miR-320e and target mRNAs in this module; POU3F2 knockdown or overexpression in SH-SY5Y cells and human neural progenitor cells modulates gene expression in schizophrenia/bipolar-associated gene networks. siRNA knockdown and overexpression in neural cell lines and human neural progenitor cells, RNA-seq Science translational medicine Medium 30545964
2024 BRN2 in cynomolgus monkey biallelic knockout (lethal before midgestation) decreases radial glia cell expansion, induces precocious differentiation, and alters neurogenesis trajectory in the telencephalon; BRN2 directly regulates SOX2 and STAT3 and maintains HOPX expression; BRN2 controls specification and differentiation of ganglionic eminences, revealing primate-specific mechanistic functions. Biallelic knockout in cynomolgus monkeys (CRISPR), single-cell transcriptome, histology, target gene expression analysis Science advances High 35245119
2024 BRN1 and BRN2 have an evolutionarily conserved function in neocortical progenitors to control proliferative capacity and the switch from direct to indirect neurogenesis; BRN1/2 act in concert with NOTCH and primary microcephaly genes; this is validated in mice, ferrets, and transcriptomic data from genetically modified macaques. Conditional knockout in mice and ferrets, transcriptomics in genetically modified macaques, functional epistasis Nature communications High 39271675
2025 POU3F2 mutation in human neural progenitor cells reduces canonical Wnt signalling and decreases proliferation, causing premature radial glia specification; SOX13 and ADNP are direct transcriptional targets of POU3F2 that mediate its effects on Wnt signaling; loss-of-function mutations in POU3F2 are found in individuals with autism spectrum disorder. CRISPR-mediated POU3F2 disruption in human NPCs, RNA-seq, ChIP-seq, Wnt pathway reporter assays Brain Medium 40498903
2023 POU3F2 promotes radioresistance in triple-negative breast cancer by interacting with ARNT2 and enhancing Akt pathway activation; Co-IP demonstrated POU3F2-ARNT2 interaction; POU3F2 knockdown decreased radioresistance and reduced proliferation and invasion. Co-immunoprecipitation, siRNA knockdown, clonogenic survival assays, xenograft model Breast cancer research and treatment Medium 36797433
2024 RFX4 directly interacts with the promoters of POU3F2 and NEUROD1, acting as an upstream regulator; ectopic expression of RFX4 drives human embryonic stem cells toward a neuronal fate; CRISPR-Cas9 knockout of RFX4 reduces POU3F2 expression and impairs neuronal differentiation. Multi-omics (ATAC-seq, ChIP-seq, Hi-C, RNA-seq), RFX4 knock-in and CRISPR-Cas9 knockout, neuronal differentiation assays Cellular and molecular life sciences Medium 38386071
2021 Integrin α3β1 promotes BRN2 expression in triple-negative breast cancer cells via Akt signaling; RNAi suppression of α3β1 reduces Brn-2 mRNA, protein, and BRN2 promoter activity; Akt inhibitor reduces both Brn-2 expression and invasion; exogenous Brn-2 partially restores invasion in α3β1-suppressed cells. RNAi knockdown, promoter reporter assays, pharmacological Akt inhibition, invasion assays in vitro and in vivo Cancers Medium 33513758
2017 Deletion of the three homopolymeric amino acid repeats (polyG, polyQ, polyP) from the Pou3f2 transactivation domain (Pou3f2Δ/Δ mice) causes cognitive impairment and reduced adult hippocampal neurogenesis, with fewer newborn neurons in the dentate gyrus, demonstrating a functional role for these mammalian-specific repeats in cognitive function and neurogenesis. Knock-in mouse model, behavioral testing (object recognition/location), immunohistochemistry for doublecortin and BrdU/NeuN co-labeling Genes, brain, and behavior Medium 28782255
2014 Replacement of mammalian Pou3f2 with the Xenopus ortholog lacking homopolymeric amino acid repeats (nonmammalized mice) causes deficits in maternal pup retrieval behavior and decreases in dopamine and serotonin synthesis rate-limiting enzymes; knock-in mice lacking the mammalian-specific polyAA repeats show similar results, demonstrating that these mammalian-specific sequences are required for normal monoamine levels and maternal behavior. Knock-in mice with Xenopus ortholog replacement, knock-in mice with polyAA deletions, behavioral testing, immunohistochemical quantification of monoamine synthesis enzymes Genome biology and evolution Medium 24709564
1999 Brn-2 is required as an intermediary for retinoic acid-induced CRF gene transcription in human neuroblastoma BE(2)-M17 cells; antisense Brn-2 aborts RA-mediated CRF induction, but Brn-2 overexpression alone is not sufficient for CRF expression without RA. Antisense RNA inhibition, ectopic overexpression, reporter gene assays Molecular endocrinology Low 10446900
2008 BRN2 knockdown does not alter nestin expression in melanoma cells (negative result); SOX9 and SOX10, but not BRN2, are required for nestin expression in melanoma cells, as SOX9/SOX10 knockdown markedly decreased nestin levels. siRNA knockdown, Western blot and mRNA analysis The Journal of investigative dermatology Medium 18923447
2003 Adenoviral overexpression of Brn-2 in the rat paraventricular nucleus had no effect on vasopressin hnRNA levels, and did not upregulate CRF mRNA in vivo, despite Brn-2 transactivating the proximal CRF promoter in vitro; this indicates Brn-2 is not rate-limiting for VP or CRF expression in adult hypothalamus. Adenoviral-mediated in vivo overexpression in rat PVN, hnRNA and mRNA analysis Molecular and cellular endocrinology Medium 12644309

Source papers

Stage 0 corpus · 95 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2016 The Master Neural Transcription Factor BRN2 Is an Androgen Receptor-Suppressed Driver of Neuroendocrine Differentiation in Prostate Cancer. Cancer discovery 355 27784708
1995 Development and survival of the endocrine hypothalamus and posterior pituitary gland requires the neuronal POU domain factor Brn-2. Genes & development 247 8543156
2003 The POU proteins Brn-2 and Oct-6 share important functions in Schwann cell development. Genes & development 240 12782656
1995 The POU domain transcription factor Brn-2 is required for the determination of specific neuronal lineages in the hypothalamus of the mouse. Genes & development 230 8543155
2009 Intravital imaging reveals transient changes in pigment production and Brn2 expression during metastatic melanoma dissemination. Cancer research 168 19826052
2012 POU-III transcription factors (Brn1, Brn2, and Oct6) influence neurogenesis, molecular identity, and migratory destination of upper-layer cells of the cerebral cortex. Cerebral cortex (New York, N.Y. : 1991) 162 22892427
1999 PQBP-1, a novel polyglutamine tract-binding protein, inhibits transcription activation by Brn-2 and affects cell survival. Human molecular genetics 160 10332029
2008 Brn-2 represses microphthalmia-associated transcription factor expression and marks a distinct subpopulation of microphthalmia-associated transcription factor-negative melanoma cells. Cancer research 159 18829533
2011 Melanoma cell invasiveness is regulated by miR-211 suppression of the BRN2 transcription factor. Pigment cell & melanoma research 150 21435193
2006 Control of myelination in Schwann cells: a Krox20 cis-regulatory element integrates Oct6, Brn2 and Sox10 activities. EMBO reports 144 16311519
2018 Bi-allelic Loss of CDKN2A Initiates Melanoma Invasion via BRN2 Activation. Cancer cell 105 29990501
2004 The Brn-2 transcription factor links activated BRAF to melanoma proliferation. Molecular and cellular biology 97 15024080
2004 Brn-2 expression controls melanoma proliferation and is directly regulated by beta-catenin. Molecular and cellular biology 95 15024079
2003 Human melanoblasts in culture: expression of BRN2 and synergistic regulation by fibroblast growth factor-2, stem cell factor, and endothelin-3. The Journal of investigative dermatology 83 14708619
2018 The transcription factor POU3F2 regulates a gene coexpression network in brain tissue from patients with psychiatric disorders. Science translational medicine 79 30545964
1993 A CNS-specific POU transcription factor, Brn-2, is required for establishing mammalian neural cell lineages. Neuron 78 8274283
2011 Inverse expression states of the BRN2 and MITF transcription factors in melanoma spheres and tumour xenografts regulate the NOTCH pathway. Oncogene 76 21358674
2017 NFIB Mediates BRN2 Driven Melanoma Cell Migration and Invasion Through Regulation of EZH2 and MITF. EBioMedicine 72 28119061
2019 BRN4 Is a Novel Driver of Neuroendocrine Differentiation in Castration-Resistant Prostate Cancer and Is Selectively Released in Extracellular Vesicles with BRN2. Clinical cancer research : an official journal of the American Association for Cancer Research 67 31371344
1995 The brn-2 gene regulates the melanocytic phenotype and tumorigenic potential of human melanoma cells. Oncogene 67 7651733
2016 Prenatal β-catenin/Brn2/Tbr2 transcriptional cascade regulates adult social and stereotypic behaviors. Molecular psychiatry 65 26830142
2008 POU domain transcription factors: BRN2 as a regulator of melanocytic growth and tumourigenesis. Pigment cell & melanoma research 54 18983536
2017 MITF and BRN2 contribute to metastatic growth after dissemination of melanoma. Scientific reports 48 28883623
2010 Genome-wide analysis of POU3F2/BRN2 promoter occupancy in human melanoma cells reveals Kitl as a novel regulated target gene. Pigment cell & melanoma research 48 20337985
2018 BRN2, a POUerful driver of melanoma phenotype switching and metastasis. Pigment cell & melanoma research 46 29781575
2013 POU domain transcription factor BRN2 is crucial for expression of ASCL1, ND1 and neuroendocrine marker molecules and cell growth in small cell lung cancer. Pathology international 46 23530560
2012 A phosphatidylinositol 3-kinase-Pax3 axis regulates Brn-2 expression in melanoma. Molecular and cellular biology 42 22988297
2010 Mutant huntingtin fragment selectively suppresses Brn-2 POU domain transcription factor to mediate hypothalamic cell dysfunction. Human molecular genetics 41 20185558
2008 SOX9 and SOX10 but not BRN2 are required for nestin expression in human melanoma cells. The Journal of investigative dermatology 40 18923447
2000 Domains of Brn-2 that mediate homodimerization and interaction with general and melanocytic transcription factors. European journal of biochemistry 40 11029584
2016 Small 6q16.1 Deletions Encompassing POU3F2 Cause Susceptibility to Obesity and Variable Developmental Delay with Intellectual Disability. American journal of human genetics 37 26833329
1997 Predominant expression of Brn-2 in the postmitotic neurons of the developing mouse neocortex. Brain research 36 9106466
1996 Preferential differentiation of P19 mouse embryonal carcinoma cells into smooth muscle cells. Use of retinoic acid and antisense against the central nervous system-specific POU transcription factor Brn-2. Circulation research 36 8593698
2018 NTF3 Is a Novel Target Gene of the Transcription Factor POU3F2 and Is Required for Neuronal Differentiation. Molecular neurobiology 35 29549646
2018 A PAX3/BRN2 rheostat controls the dynamics of BRAF mediated MITF regulation in MITFhigh /AXLlow melanoma. Pigment cell & melanoma research 34 30277012
2002 Expression of the POU-domain transcription factors SCIP/Oct-6 and Brn-2 is associated with Schwann cell but not oligodendrocyte remyelination of the CNS. Molecular and cellular neurosciences 33 12213447
2019 BRN2 suppresses apoptosis, reprograms DNA damage repair, and is associated with a high somatic mutation burden in melanoma. Genes & development 32 30804224
2018 Regulation of tNOX expression through the ROS-p53-POU3F2 axis contributes to cellular responses against oxaliplatin in human colon cancer cells. Journal of experimental & clinical cancer research : CR 29 30029680
2020 LncRNA BCYRN1/miR-490-3p/POU3F2, served as a ceRNA network, is connected with worse survival rate of hepatocellular carcinoma patients and promotes tumor cell growth and metastasis. Cancer cell international 27 31920461
2017 POU3F2 participates in cognitive function and adult hippocampal neurogenesis via mammalian-characteristic amino acid repeats. Genes, brain, and behavior 27 28782255
2013 Neural lineage-specific homeoprotein BRN2 is directly involved in TTF1 expression in small-cell lung cancer. Laboratory investigation; a journal of technical methods and pathology 27 23358112
2021 Integrin α3β1 Promotes Invasive and Metastatic Properties of Breast Cancer Cells through Induction of the Brn-2 Transcription Factor. Cancers 26 33513758
2020 BRN2 expression increases anoikis resistance in melanoma. Oncogenesis 26 32632141
2020 Transcription factor POU3F2 regulates TRIM8 expression contributing to cellular functions implicated in schizophrenia. Molecular psychiatry 26 32929213
2018 Brn2 Alone Is Sufficient to Convert Astrocytes into Neural Progenitors and Neurons. Stem cells and development 25 29635978
2015 A Brn2-Zic1 axis specifies the neuronal fate of retinoic-acid-treated embryonic stem cells. Journal of cell science 25 25991548
1995 Isolation of the human genomic brain-2/N-Oct 3 gene (POUF3) and assignment to chromosome 6q16. Genomics 25 7601453
2020 MicroRNA-107 is a novel tumor suppressor targeting POU3F2 in melanoma. Biological research 23 32169117
2012 BRN2 is a transcriptional repressor of CDH13 (T-cadherin) in melanoma cells. Laboratory investigation; a journal of technical methods and pathology 23 23069940
2005 The neuronal POU transcription factor Brn-2 interacts with Jab1, a gene involved in the onset of neurodegenerative diseases. Neuroscience letters 23 15911144
1999 A requirement for the POU transcription factor, Brn-2, in corticotropin-releasing hormone expression in a neuronal cell line. Molecular endocrinology (Baltimore, Md.) 23 10446900
2012 Phosphorylation of BRN2 modulates its interaction with the Pax3 promoter to control melanocyte migration and proliferation. Molecular and cellular biology 22 22290434
2006 Exclusion of mutations in the PRNP, JPH3, TBP, ATN1, CREBBP, POU3F2 and FTL genes as a cause of disease in Portuguese patients with a Huntington-like phenotype. Journal of human genetics 22 16858508
2021 Hierarchical reactivation of transcription during mitosis-to-G1 transition by Brn2 and Ascl1 in neural stem cells. Genes & development 21 34168041
2020 A Novel miR-146a-POU3F2/SMARCA5 Pathway Regulates Stemness and Therapeutic Response in Glioblastoma. Molecular cancer research : MCR 21 32973101
2008 Histologic and epidemiologic correlates of P-MAPK, Brn-2, pRb, p53, and p16 immunostaining in cutaneous melanomas. Melanoma research 20 18781132
2024 RFX4 is an intrinsic factor for neuronal differentiation through induction of proneural genes POU3F2 and NEUROD1. Cellular and molecular life sciences : CMLS 19 38386071
2016 Capsaicin Inhibited Aggressive Phenotypes through Downregulation of Tumor-Associated NADH Oxidase (tNOX) by POU Domain Transcription Factor POU3F2. Molecules (Basel, Switzerland) 19 27271588
2020 Phosphorylation of Tet3 by cdk5 is critical for robust activation of BRN2 during neuronal differentiation. Nucleic acids research 18 31807777
2006 Rb induces a proliferative arrest and curtails Brn-2 expression in retinoblastoma cells. Molecular cancer 17 17163992
2014 Mammalian-specific sequences in pou3f2 contribute to maternal behavior. Genome biology and evolution 16 24709564
2021 Circ_0031242 Silencing Mitigates the Progression and Drug Resistance in DDP-Resistant Hepatoma Cells by the miR-924/POU3F2 Axis. Cancer management and research 15 33531841
2001 NF-Y binding is required for transactivation of neuronal aromatic L-amino acid decarboxylase gene promoter by the POU-domain protein Brn-2. Brain research. Molecular brain research 15 11311976
2024 Conserved transcriptional regulation by BRN1 and BRN2 in neocortical progenitors drives mammalian neural specification and neocortical expansion. Nature communications 14 39271675
2021 BRN2 is a non-canonical melanoma tumor-suppressor. Nature communications 13 34140478
2018 A De Novo Missense Variant in POU3F2 Identified in a Child with Global Developmental Delay. Neuropediatrics 13 30199896
2010 Brn2 is a transcription factor regulating keratinocyte differentiation with a possible role in the pathogenesis of lichen planus. PloS one 13 20967260
2023 Overexpression of POU3F2 promotes radioresistance in triple-negative breast cancer via Akt pathway activation. Breast cancer research and treatment 11 36797433
2022 BRN2 as a key gene drives the early primate telencephalon development. Science advances 11 35245119
2023 Integrin α3 Mediates Stemness and Invasion of Glioblastoma by Regulating POU3F2. Current protein & peptide science 10 36843258
2025 The promoting effect of the POU3F2/METTL16/PFKM cascade on glycolysis and tumorigenesis of hepatocellular carcinoma. Annals of hepatology 8 39756795
2023 Monoallelic intragenic POU3F2 variants lead to neurodevelopmental delay and hyperphagic obesity, confirming the gene's candidacy in 6q16.1 deletions. American journal of human genetics 8 37207645
2019 Peptide R18H from BRN2 Transcription Factor POU Domain Displays Antitumor Activity In Vitro and In Vivo and Induces Apoptosis in B16F10-Nex2 Cells. Anti-cancer agents in medicinal chemistry 7 30417795
2017 Effects of Brn2 overexpression on eccrine sweat gland development in the mouse paw. Biochemical and biophysical research communications 7 28648603
2023 CREB1 Facilitates GABAergic Neural Differentiation of Human Mesenchymal Stem Cells through BRN2 for Pain Alleviation and Locomotion Recovery after Spinal Cord Injury. Cells 6 38201271
2015 Partial Dedifferentiation of Murine Radial Glia-Type Neural Stem Cells by Brn2 and c-Myc Yields Early Neuroepithelial Progenitors. Journal of molecular biology 6 26555748
2022 CircPOLR2A Promotes Proliferation and Impedes Apoptosis of Glioblastoma Multiforme Cells by Up-regulating POU3F2 to Facilitate SOX9 Transcription. Neuroscience 5 35398178
2020 BRN2 and MITF together impact AXL expression in melanoma. Experimental dermatology 5 33119145
2004 Cooperative dimerization of the POU domain protein Brn-2 on a new motif activates the neuronal promoter of the human aromatic L-amino acid decarboxylase gene. Brain research. Molecular brain research 5 14741405
2003 Adenoviral-mediated over-expression of Brn2 in the rat paraventricular nucleus: no effect on vasopressin or corticotrophin releasing factor RNA levels. Molecular and cellular endocrinology 5 12644309
1998 Transcriptional regulatory region of Brn-2 required for its expression in developing olfactory epithelial cells. Brain research. Developmental brain research 5 9706393
2022 Deficient maternal behavior in multiparous Pou3f2⊿ mice is associated with an impaired exploratory activity. Behavioural brain research 4 35306097
2017 Immunohistochemical study of the neural development transcription factors (TTF1, ASCL1 and BRN2) in neuroendocrine prostate tumours. Actas urologicas espanolas 4 28285791
2017 Neural differentiation of fibroblasts induced by intracellular co-delivery of Ascl1, Brn2 and FoxA1 via a non-viral vector of cationic polysaccharide. Biomedical materials (Bristol, England) 4 28855426
2016 In silico studies of the interaction between BRN2 protein and MORE DNA. Journal of molecular modeling 4 27568376
2025 POU3F2 regulates canonical Wnt signalling via SOX13 and ADNP to expand the neural progenitor population. Brain : a journal of neurology 3 40498903
2021 Reciprocal Regulation of BRN2 and NOTCH1/2 Signaling Synergistically Drives Melanoma Cell Migration and Invasion. The Journal of investigative dermatology 3 34958806
2024 Neuroendocrine prostate cancer drivers SOX2 and BRN2 confer differential responses to imipridones ONC201, ONC206, and ONC212 in prostate cancer cell lines. American journal of translational research 2 39822524
2026 Resistance to neoadjuvant talazoparib in triple-negative breast cancer by BRN2-induced ATR/STAT3 pathways or SHLD2 subclone expansion. Proceedings of the National Academy of Sciences of the United States of America 1 41961819
2024 Peptide L13S Derived from the BRN2 POU Domain Reduces Metastasis In Vivo and Inhibits Cell Migration and Invasion In Vitro. Anticancer research 1 38160013
2026 Reprogramming of astrocytes into motoneuron-like cells by a Ascl1-Myt1l-Pou3f2-Isl1 cocktail. Experimental cell research 0 41763511
2026 Transposable element-mediated evolutionary expansion of Sox2- and Brn2-binding regulatory modules for mammalian neural-cell differentiation. Genome biology 0 41952195
2025 Peptides derived from the POU domain of BRN2 show antitumor activity against murine melanoma model cells in vitro and in vivo. Cancer chemotherapy and pharmacology 0 40601014
2025 A BRN2:MYC transcriptional axis regulates interconversion between therapy-resistant and tumorigenic phenotypes in melanoma. Cell reports 0 41405996
2024 Neuroendocrine Prostate Cancer Drivers SOX2 and BRN2 Confer Differential Responses to Imipridones ONC201, ONC206, and ONC212 in Prostate Cancer Cell Lines. bioRxiv : the preprint server for biology 0 39257809

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