Affinage

RAX

Retinal homeobox protein Rx · UniProt Q9Y2V3

Round 2 corrected
Length
346 aa
Mass
36.7 kDa
Annotated
2026-04-28
130 papers in source corpus 34 papers cited in narrative 34 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

RAX encodes a paired-type homeodomain transcription factor that functions as a master regulator of vertebrate eye development, hypothalamic patterning, and photoreceptor differentiation; it also shares its gene symbol with a distinct dsRNA-binding protein (PACT/RAX) that activates PKR. As a retinal homeobox gene, RAX is essential for optic cup formation and retinal progenitor cell specification — its targeted deletion causes anophthalmia — and acts genetically upstream of Pax6 while being regulated by Otx2 and Sox2 via the CNS1 enhancer (PMID:9177348, PMID:11105055, PMID:18385377). RAX directly activates the Otx2 enhancer EELPOT in late-stage retinal progenitors and cooperates physically with CRX to transactivate rhodopsin and cone opsin promoters, thereby controlling photoreceptor gene expression, maturation, and survival (PMID:22090505, PMID:25986607). Mutations in the human RAX homeodomain cause anophthalmia and sclerocornea (PMID:14662654), and beyond the eye, RAX functions as a selector gene for ventromedial and arcuate hypothalamic neuronal identities and for α2-tanycyte differentiation (PMID:23283339, PMID:23939786, PMID:27212025).

Mechanistic history

Synthesis pass · year-by-year structured walk · 16 steps
  1. 1997 High

    Identification of Rx/RAX as a retinal-specific homeodomain gene essential for eye formation resolved a long-standing question of what transcription factors initiate vertebrate eye development, as targeted null deletion produced complete anophthalmia and gain-of-function induced ectopic retinal tissue.

    Evidence cDNA cloning, in situ hybridization, gene knockout in mice (anophthalmia), and Rx mRNA injection in Xenopus (ectopic retina)

    PMID:9096350 PMID:9177348

    Open questions at the time
    • Direct DNA-binding targets of Rx/RAX were not identified
    • Upstream regulators of Rx expression were unknown
    • Mechanism by which Rx promotes proliferation was uncharacterized
  2. 1998 High

    Independent discovery of PACT/RAX as the first cellular protein activator of PKR established that PKR can be activated without dsRNA, revealing a new stress-signaling paradigm — though this is a distinct protein from the retinal homeobox RAX.

    Evidence In vitro PKR activation reconstitution, co-immunoprecipitation, yeast growth assays

    PMID:10336432 PMID:9687506

    Open questions at the time
    • The phosphorylation event required for RAX/PACT-mediated PKR activation was not yet mapped
    • Physiological stress signals upstream of RAX/PACT phosphorylation were incompletely defined
  3. 2000 High

    Genetic epistasis placed Rx upstream of Pax6 in retinal progenitor specification and showed Rx also promotes Müller glial fate via Hes1/Notch1, broadening its role from progenitor establishment to cell-fate diversification within the retina.

    Evidence Rx-null vs Pax6-null epistasis in mouse embryos; retroviral transduction and reporter assays for glial markers

    PMID:10839357 PMID:11105055

    Open questions at the time
    • Whether Rx directly binds Hes1/Notch1 regulatory regions was not demonstrated by ChIP
    • The mechanism of Rx-mediated glial vs neuronal fate choice remained unclear
  4. 2001 High

    The eyeless (ey1) mouse mutation was mapped to a Met10Leu change in Rx affecting an alternative translation initiation site, revealing that leaky scanning-based translational regulation of Rx protein levels is physiologically critical and that even partial Rx reduction causes anophthalmia.

    Evidence Genetic mapping, sequencing, in vitro translation assay in ZRDCT strain

    PMID:11668677

    Open questions at the time
    • Quantitative relationship between Rx protein level and eye phenotype severity was not established
    • Other tissues affected by reduced Rx were not systematically examined
  5. 2003 High

    Discovery of human RAX homeodomain mutations (Q147X, R192Q) causing anophthalmia and sclerocornea established RAX as a Mendelian disease gene for human eye malformations.

    Evidence Patient sequencing and in vitro biochemical characterization of mutant proteins

    PMID:14662654

    Open questions at the time
    • Genotype-phenotype correlations across a larger patient cohort were lacking
    • The precise biochemical defect of R192Q (DNA-binding vs protein interaction) was not resolved
  6. 2004 High

    Mapping Ser18 as the critical phosphorylation site for PACT/RAX-mediated PKR activation defined a sequential activation mechanism (RAX–PKR association → Ser18 phosphorylation → PKR kinase activation → eIF2α phosphorylation), and showed that non-phosphorylatable S18A acts as a dominant negative blocking stress-induced apoptosis.

    Evidence Site-directed mutagenesis, stable cell lines, eIF2α phosphorylation and apoptosis assays

    PMID:15299031

    Open questions at the time
    • The kinase responsible for Ser18 phosphorylation was not identified
    • Structural basis of how Ser18 phosphorylation converts RAX from an inactive to an active PKR cofactor was unknown
  7. 2006 High

    Multiple studies converged to show PACT/RAX is required for PKR activation across diverse stresses (cytokines, serum deprivation, ethanol, viral infection) and participates in RISC assembly as a Dicer-associated dsRNA-binding protein, greatly expanding its functional scope beyond a simple PKR cofactor.

    Evidence siRNA knockdown, dominant-negative S18A, co-IP with Dicer/Ago2/TRBP, miRNA accumulation assays, viral infection assays

    PMID:16424907 PMID:16574643 PMID:16861340

    Open questions at the time
    • Whether PKR-activating and RISC-assembly functions of PACT/RAX are mutually exclusive or concurrent was not determined
    • The stoichiometry of PACT within the RISC-loading complex was not established
  8. 2008 High

    ChIP demonstration that endogenous Otx2 and Sox2 bind the conserved CNS1 enhancer to synergistically activate Rax transcription placed Rax within a defined upstream regulatory hierarchy (Otx2/Sox2 → Rax → Pax6) for eye field specification.

    Evidence ChIP with endogenous antibodies, reporter assays, co-immunoprecipitation in Xenopus

    PMID:18385377

    Open questions at the time
    • Whether additional upstream factors besides Otx2 and Sox2 regulate Rax through CNS1 was not explored
    • Chromatin accessibility dynamics at the Rax locus during eye field specification were not assessed
  9. 2011 High

    Two advances clarified RAX's role in photoreceptor fate: RAX directly activates the Otx2 enhancer EELPOT in terminal progenitor divisions (competing with Notch-HES repression), and RAX is required for retinal regeneration in Xenopus, extending its function beyond initial development.

    Evidence Enhancer mapping, conditional Rax KO in mouse retina, shRNA knockdown and rescue in Xenopus regeneration

    PMID:21334323 PMID:22090505

    Open questions at the time
    • Whether RAX binds EELPOT directly or through a cofactor complex was not resolved by ChIP
    • The regenerative role of RAX in mammalian retina was not tested
  10. 2012 High

    PACT/RAX was shown to interact with the SUMO E2 ligase Ubc9 to promote p53 sumoylation at K386, causing p53 stabilization and G1 arrest — a PKR-dependent but mechanistically distinct function linking PACT/RAX to cell cycle control.

    Evidence Co-immunoprecipitation, sumoylation assays, cell cycle analysis, p53 K386R mutant, SENP1 co-expression

    PMID:22214662

    Open questions at the time
    • Whether p53 sumoylation by the RAX-Ubc9 axis occurs under physiological stress was not demonstrated
    • The relative contribution of this pathway vs direct eIF2α phosphorylation to RAX-mediated growth arrest was not quantified
  11. 2013 High

    Conditional ablation experiments established RAX as a selector gene for mediobasal hypothalamic cell types (VMH/ARC neurons), demonstrating that RAX's developmental role extends well beyond the eye to include hypothalamic neuronal identity specification.

    Evidence Conditional KO with Shh::Cre and Six3::Cre drivers, genetic fate mapping, immunostaining in mouse

    PMID:23283339

    Open questions at the time
    • Direct transcriptional targets of RAX in hypothalamic progenitors were not identified
    • Whether RAX interacts with hypothalamic-specific cofactors distinct from retinal cofactors was unknown
  12. 2014 High

    RAX haploinsufficiency revealed a selective requirement for RAX in α2-tanycyte differentiation and CSF-hypothalamus barrier function, while in Xenopus, Rax-null presumptive retina was shown to adopt diencephalic/telencephalic identity, redefining Rax as a suppressor of alternative forebrain fates.

    Evidence Rax haploinsufficient mice with Evans Blue diffusion assays; Xenopus rax mutant RNA-seq with hesx1/fezf2 epistasis

    PMID:23939786 PMID:25224223

    Open questions at the time
    • Whether tanycyte loss contributes to metabolic phenotypes in Rax-deficient animals was not tested
    • Genome-wide direct RAX targets in tanycytes were not mapped
  13. 2015 High

    Conditional inactivation of Rax in postnatal photoreceptors demonstrated that RAX cooperates with CRX through direct physical interaction to transactivate rhodopsin and cone opsin promoters, and that loss causes cone cell death — establishing RAX as required not just for photoreceptor specification but for ongoing photoreceptor maintenance and survival.

    Evidence Tamoxifen-inducible conditional KO, co-immunoprecipitation, luciferase reporter assays

    PMID:25986607

    Open questions at the time
    • The structural basis of the RAX-CRX interaction was not determined
    • Whether RAX has CRX-independent transcriptional targets in mature photoreceptors was not addressed
  14. 2015 High

    A surprising tissue-specific reversal was discovered: in anterior pituitary, RAX/PACT acts as a negative regulator of PKR, and pituitary hypoplasia in Rax-null mice is rescued by PKR deletion or kinase-dead PKR knockin, demonstrating context-dependent modulation of PKR activity.

    Evidence Double-knockout and knockin mouse genetics, pituitary cell RAX knockdown

    PMID:26414443

    Open questions at the time
    • The molecular basis for opposite PKR regulation in pituitary vs hematopoietic cells was not explained
    • Whether post-translational modifications of RAX differ between tissues was not examined
  15. 2016 High

    Timed conditional Rax inactivation before E8.5 demonstrated that RAX is required for dorsoventral patterning of the hypothalamic neuroepithelium and controls Shh expression in the dorsomedial hypothalamus, linking RAX to a core morphogen pathway.

    Evidence Tamoxifen-timed conditional KO, in situ hybridization for neuropeptides and Shh

    PMID:27212025

    Open questions at the time
    • Whether RAX directly binds the Shh locus in hypothalamic progenitors was not tested
    • The interaction between RAX-dependent and BMP/Wnt-dependent hypothalamic patterning signals was unexplored
  16. 2021 High

    Rax+ tanycytes were shown to be quiescent stem-like cells that re-enter the cell cycle upon injury for self-renewal, with oncogenic BRAF activation sufficient to transform them into craniopharyngioma-like tumors, establishing Rax+ tanycytes as a cell of origin for hypothalamic tumors.

    Evidence Rax-CreERT2 lineage tracing, neural injury model, Igf1r conditional KO, BrafV600E conditional activation

    PMID:33863883

    Open questions at the time
    • Whether RAX transcriptional activity is required for tanycyte self-renewal or only marks these cells was not distinguished
    • The tumor-suppressive vs permissive role of endogenous RAX in craniopharyngioma initiation was not tested

Open questions

Synthesis pass · forward-looking unresolved questions
  • Major open questions include: (1) the genome-wide direct binding targets of RAX in hypothalamic vs retinal progenitors, (2) the structural basis of the RAX-CRX cooperative interaction, (3) the molecular explanation for tissue-specific opposing effects of PACT/RAX on PKR activity, and (4) whether RAX transcription factor and PACT/RAX PKR-activator functions intersect in any shared cell type.
  • No genome-wide ChIP-seq for RAX in primary tissues has been reported
  • No crystal structure of RAX homeodomain-DNA complex exists
  • Tissue-specific post-translational regulation of PACT/RAX has not been systematically characterized

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140110 transcription regulator activity 6 GO:0003677 DNA binding 5 GO:0098772 molecular function regulator activity 5 GO:0003723 RNA binding 3
Localization
GO:0005634 nucleus 4
Pathway
R-HSA-1266738 Developmental Biology 6 R-HSA-162582 Signal Transduction 5 R-HSA-5357801 Programmed Cell Death 4 R-HSA-112316 Neuronal System 3 R-HSA-392499 Metabolism of proteins 3 R-HSA-74160 Gene expression (Transcription) 3 R-HSA-8953854 Metabolism of RNA 1
Partners
AGO2CRXDICER1OTX2PAX6PKRTRBPUBC9
Complex memberships
RISC-loading complex (Dicer/TRBP/PACT)

Evidence

Reading pass · 34 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1997 The Rx homeobox gene (retinal and anterior neural fold homeobox) encodes a paired-type homeodomain protein with an octapeptide and a novel C-terminal OAR domain. It is expressed in the anterior neural fold and developing retina of mouse embryos, and its targeted null deletion results in complete failure of optic cup formation and anophthalmia, establishing Rx as essential for vertebrate eye formation and retinal progenitor cell establishment. cDNA cloning, in situ hybridization, and targeted gene knockout in mice with phenotypic analysis Nature / Proceedings of the National Academy of Sciences High 9096350 9177348
1997 Misexpression of Rx RNA in Xenopus embryos induces ectopic retinal tissue formation and hyperproliferation of neuroretinal cells, demonstrating that Rx gain-of-function promotes retinal progenitor cell proliferation. Injection of synthetic Rx RNA into Xenopus embryos with morphological and histological analysis Nature High 9177348
1999 RAX (mouse PKR-associated protein X) was identified as a cellular activator of the double-stranded RNA-dependent protein kinase PKR. RAX directly activates PKR in vitro in the absence of dsRNA. Following stress stimuli (IL-3 deprivation, arsenite, thapsigargin, H2O2), RAX is rapidly phosphorylated, associates with PKR, and activates PKR, linking transmembrane stress signals to inhibition of protein synthesis via eIF2α phosphorylation. cDNA library screening, in vitro PKR activation assay, co-immunoprecipitation, overexpression in IL-3-dependent cells The Journal of Biological Chemistry High 10336432
2000 rax (retinal homeobox) promotes formation of Müller glia from postnatal retinal progenitor cells. Retroviral transduction of rax into retinal progenitors induces expression of glial markers. rax cotransfection with reporter constructs containing Hes1 or notch1 regulatory regions leads to upregulation of reporter transcription, placing rax upstream of Hes1 and Notch1 in a regulatory hierarchy controlling glial versus neuronal fate. Retroviral transduction, reporter gene assays (cotransfection), immunostaining for glial markers Neuron High 10839357
2000 Rx function, but not Pax6 function, is required cell-autonomously for formation of retinal progenitor cells. In Rx-null embryos, Otx2, Six3, and Pax6 show normal activation in anterior neural plate but fail to upregulate in the optic vesicle primordium. Conversely, in Pax6-null (Small eye) embryos, Rx shows normal expression in optic vesicle/retinal progenitors, establishing Rx as genetically upstream of Pax6 in retinal progenitor specification. Genetic epistasis analysis using Rx-null and Pax6-null mouse embryos with in situ hybridization Genesis High 11105055
2001 Ceramide activates PKR through the cellular PKR activator RAX. Ceramide promotes RAX phosphorylation and eIF2α phosphorylation, and RAX overexpression potentiates ceramide-induced cell killing and inhibition of protein synthesis. The serine/threonine kinase inhibitor 2-aminopurine blocks both ceramide-induced RAX/eIF2α phosphorylation and ceramide killing, identifying a novel ceramide–RAX–PKR axis regulating protein synthesis and apoptosis. Overexpression of RAX, ceramide treatment, protein synthesis assays, pharmacological inhibition with 2-aminopurine The Journal of Biological Chemistry Medium 11148216
2001 The eyeless mouse mutation ey1 (ZRDCT strain) is caused by a Met10→Leu mutation in the Rx/rax homeobox gene affecting a conserved AUG codon that serves as an alternative translation initiation site, reducing Rx protein abundance. This hypomorphic allele causes anophthalmia and hypothalamic abnormalities without neonatal lethality, in contrast to the complete null which is lethal, demonstrating that the leaky scanning mechanism for Rx translation is physiologically important. Genetic mapping (F2 genome scan), sequencing, in vitro translation assay Genesis High 11668677
2003 Human RAX gene mutations cause anophthalmia and sclerocornea. A compound heterozygote patient carries a truncating allele (Q147X) and a missense mutation (R192Q), both within the DNA-binding homeodomain. Biochemical characterization in vitro demonstrated altered properties of these mutant proteins, confirming that the homeodomain is critical for RAX protein function in human eye development. Patient sequencing, in vitro biochemical characterization of mutant proteins Human Molecular Genetics High 14662654
2004 RAX serine 18 phosphorylation is required for PKR activation. The non-phosphorylatable RAX(S18A) mutant retains the ability to bind dsRNA and associate with PKR but fails to activate PKR following stress. Stable expression of RAX(S18A) acts as a dominant negative, preventing eIF2α phosphorylation, delaying translation inhibition, and blocking rapid apoptosis upon IL-3 removal. This defines a sequential mechanism: RAX association with PKR → RAX phosphorylation at Ser18 → PKR activation. Site-directed mutagenesis, stable cell line expression, eIF2α phosphorylation assays, apoptosis assays The Journal of Biological Chemistry High 15299031
2004 Ectopic expression of Rx/rax transcription factor in mouse embryonic stem cells specifies retinal cell fate. CCE-RX/E cells (ES cells expressing Rx) migrate into retinal explants and differentiate into ganglion and horizontal cell types with appropriate electrophysiological properties, demonstrating that Rx is sufficient to direct ES cells toward retinal neuron identities. Retroviral transduction of ES cells, retinal explant coculture, electrophysiology, immunostaining Molecular and Cellular Biology Medium 15121868
2005 Mouse ES cells can be directed to differentiate into Rx+/Pax6+ neural retinal precursors under defined conditions (SFEB culture with Dkk1/LeftyA/activin/serum), demonstrating that Rx expression marks functional neural retinal precursors competent to generate photoreceptors (rhodopsin+, recoverin+) that integrate selectively into the outer nuclear layer. Directed differentiation of ES cells, immunostaining, organotypic coculture, retinal integration assays Proceedings of the National Academy of Sciences Medium 16076961
2006 RAX, the PKR activator, is required to activate PKR in response to a broad range of stresses including serum deprivation, inflammatory cytokines (IFNγ/TNFα), chemotherapy (mitomycin C), and viral infection. siRNA-mediated knockdown of RAX by 80% prevents IFNγ/TNFα-induced PKR activation, eIF2α phosphorylation, IκB degradation, IRF-1 expression, and STAT1 phosphorylation. Expression of RAX(S18A) dominant-negative prevents PKR activation and blocks Fanconi anemia group C-null MEF hypersensitivity to cytokines. Reduced RAX also facilitates productive VSV infection and promotes anchorage-independent growth. siRNA knockdown, dominant-negative overexpression, PKR activation assays, viral infection assays, colony formation assay Blood High 16861340
2006 RAX-PKR interaction modulates the effect of ethanol on protein synthesis and neuronal survival. Ethanol increases PKR and eIF2α phosphorylation in developing cerebellum. Overexpression of wild-type RAX dramatically enhances ethanol-induced PKR/eIF2α phosphorylation, translational inhibition, and cell death, while S18A RAX acts as a dominant negative, blocking these effects. Ethanol promotes PKR-RAX physical association in a Ser18-dependent manner. Overexpression/dominant-negative expression, co-immunoprecipitation, eIF2α phosphorylation assays, PKR null fibroblasts The Journal of Biological Chemistry High 16574643
2006 The Rx-like gene (Rx-L/Qrx) in Xenopus is expressed specifically in the developing photoreceptor layer and ciliary marginal zone. Rx-L binds PCE-1 (photoreceptor conserved element-1) oligonucleotides and functions as a transcriptional activator stronger than Rx. Morpholino-mediated knockdown of Rx-L reduces rhodopsin and red cone opsin expression and shortens rod and cone outer segments, placing Rx-L as a regulator of photoreceptor-specific gene expression. EMSA (electrophoretic mobility shift assay), morpholino knockdown, in situ hybridization, promoter assays Investigative Ophthalmology & Visual Science Medium 17003412
2006 Nucleosome regulator Xhmgb3 functions downstream of Xenopus rax/Rx1 to promote cell proliferation in the developing eye and brain. ChIP assays showed Xhmgb3 is recruited to the c-myc promoter to enhance c-myc transcription. Morpholino knockdown of Xhmgb3 reduces eye and brain size, while overexpression increases them. This defines a rax→Xhmgb3→c-myc cascade controlling retinal progenitor proliferation. ChIP assay, morpholino knockdown, overexpression, BrdU proliferation assays Developmental Biology Medium 16445903
2008 Otx2 and Sox2 proteins are direct upstream regulators of Xenopus Rax expression. Endogenous Otx2 and Sox2 proteins bind to the conserved noncoding sequence CNS1 (~2 kb upstream of the Rax promoter), which is required for potent transcriptional activity. Otx2 and Sox2 synergistically activate Rax transcription via CNS1, and the proteins physically interact with each other; Sox2 missense mutations found in ocular disorders disrupt this interaction. ChIP with endogenous proteins, reporter assays, co-immunoprecipitation, mutagenesis of CNS1 Proceedings of the National Academy of Sciences High 18385377
2008 RAX (PKR activator) is required for Drosophila nervous system development and for mouse embryogenesis. dRax mutant flies display severe defects in commissural axon structure of the CNS, reduced locomotion, and lethality. Deletion of the entire mouse Rax gene (encoding the PKR activator) results in preimplantation lethality, as no homozygous null embryos are recovered at E3.5, E7, or E14, indicating that this RAX is essential at the blastocyst stage. Transposon insertion mutant analysis in Drosophila, mouse Rax gene knockout by targeted deletion, embryonic chimera analysis Mechanisms of Development Medium 18634873
2009 Rx function is required cell-autonomously in the mammalian retina and posterior pituitary. In embryonic chimeras of wild-type and Rx-/- cells, Rx-deficient cells cannot participate in formation of the neuroretina, retinal pigment epithelium, or distal optic stalk. Rx-/- cells also cannot form posterior pituitary. Furthermore, Rx-/- and wild-type cells segregate before morphogenesis, suggesting Rx is required for cell sorting to form distinct retinal/pituitary fields prior to morphogenesis. Embryonic chimera generation (wild-type and Rx-/- cells), histological analysis, cell fate tracking PLoS One High 19229337
2011 RAX homeoprotein interacts with the conserved Otx2 enhancer EELPOT (~500 bp) to transactivate Otx2 expression mainly in the final cell cycle of retinal progenitors. Conditional inactivation of Rax in vivo results in downregulation of Otx2 expression in the retina. NOTCH-HES signaling negatively regulates EELPOT to suppress Otx2, thus competing with RAX-mediated activation in photoreceptor fate determination. Enhancer identification, reporter assays, conditional Rax knockout mice, in situ hybridization The Journal of Neuroscience High 22090505
2011 The Retinal Homeobox (Rx) gene is necessary for retinal regeneration in Xenopus. After partial retinal resection, wound-repopulating retinal progenitor cells express Rx. shRNA-mediated silencing of Rx in tadpoles impairs retinal regeneration, including defects in wound repopulation and RPE repair. Exogenous Rx rescues these regeneration defects, demonstrating that Rx function is required during both retinal development and regeneration. Partial retinal resection, shRNA knockdown in vivo, rescue with exogenous Rx, marker gene analysis Developmental Biology High 21334323
2011 miR-29b regulates neuronal apoptosis through the SP1/RAX/PKR cascade. Ethanol suppresses miR-29b in developing cerebellar granule neurons, leading to RAX upregulation, PKR activation, and apoptosis. Overexpression of miR-29b protects neurons from ethanol-induced apoptosis through this cascade. miRNA overexpression, ethanol treatment of cerebellar neurons, apoptosis assays, Western blotting The Journal of Biological Chemistry Medium 24554719
2012 RAX/PACT interacts with the SUMO E2 ligase Ubc9 to stimulate p53-Ubc9 association and reversible p53 sumoylation on lysine 386. RAX/PACT expression promotes p53 stability, increases p53 target gene expression, and causes G1 arrest when co-expressed with p53. This G1 arrest requires RAX/PACT-dependent PKR activity and p53 sumoylation at K386 (sumoylation-deficient p53(K386R) or co-expression with SENP1 desumoylase prevents arrest). Co-immunoprecipitation, sumoylation assays, cell cycle analysis, dominant-negative PKR, p53 mutants Cell Cycle High 22214662
2012 Combinations of transcription factors CRX, RX, and NEUROD can generate photosensitive photoreceptor cells from human iris cells. CRX+RX induces blue opsin and green/red opsin but not rhodopsin; CRX+NEUROD induces rhodopsin and blue opsin. Adding NEUROD to CRX+RX promotes photoreceptor maturation. Patch-clamp recordings confirm that generated cells respond to light with inward currents. Transcription factor overexpression, qRT-PCR, immunostaining, patch-clamp electrophysiology PLoS One Medium 22558175
2013 Rax is a selector gene for mediobasal hypothalamic cell types. Rax is expressed in ARC and VMH progenitors, and genetic fate mapping shows Rax+ lineages give rise to VMH neurons. Conditional ablation of Rax using Shh::Cre causes a fate switch from VMH neuronal phenotype to hypothalamic but non-VMH identity. Broader elimination using Six3::Cre causes severe loss of both VMH and ARC cellular phenotypes. Conditional knockout with Shh::Cre and Six3::Cre drivers, genetic fate mapping, immunostaining The Journal of Neuroscience High 23283339
2014 Rax is required for hypothalamic tanycyte differentiation and the cerebrospinal fluid-hypothalamus barrier. Rax haploinsufficient mice show thinning of the third ventricular wall, reduction of tanycyte and ependymal markers, ectopic ependymal cells in the α2 tanycytic zone, and reduced diffusion of Evans Blue tracer from the ventricle to hypothalamic parenchyma, demonstrating that Rax selectively controls α2 tanycyte differentiation and barrier function. Rax haploinsufficient mouse analysis, immunostaining, Evans Blue diffusion assay, quantitative marker analysis The Journal of Comparative Neurology High 23939786
2014 Xenopus rax mutant tissue fated to form retina instead adopts diencephalic and telencephalic character, indicating that a key role of rax is to prevent alternative forebrain identities in the presumptive retina. RNA-seq analysis identified hesx1 and fezf2 as genes upregulated in rax mutant presumptive retinal tissue; morpholino knockdown of hesx1 and fezf2 partially rescues the retinal-to-forebrain identity transformation in the rax mutant. Xenopus tropicalis rax nonsense mutant analysis, RNA-seq, morpholino knockdown, in situ hybridization Developmental Biology High 25224223
2014 Human dermal fibroblasts can be directly differentiated into photoreceptor cells by the transcription factor combination CRX, RAX, OTX2, and NEUROD. Transduction induces photoreceptor-specific genes (recoverin, blue opsin, PDE6C), upregulates NRL and NR2E3 endogenously, and generates cells that respond to light in patch-clamp recordings. Global gene expression analysis confirms photoreceptor-related functional gene upregulation. Transcription factor transduction, microarray gene expression profiling, patch-clamp electrophysiology Genes to Cells Medium 24456169
2015 Rax homeoprotein cooperates with Crx to regulate photoreceptor gene expression and photoreceptor cell maturation and survival. Conditional inactivation of Rax in postnatal photoreceptors (Rax iCKO mice) causes decreased rod and cone photoreceptor gene expression and specific cone cell loss. Luciferase assays show Rax and Crx cooperatively transactivate Rhodopsin and cone opsin promoters with an optimum Rax expression level. Co-immunoprecipitation in cultured cells confirms physical Rax-Crx interaction. Conditional knockout (tamoxifen-inducible), luciferase reporter assays, co-immunoprecipitation, immunostaining Molecular and Cellular Biology High 25986607
2015 RAX functions as a negative regulator of PKR in anterior pituitary tissue. Mice deficient in RAX (rax-/-) show pituitary hypoplasia, reproductive defects, and craniofacial abnormalities not observed in PKR-deficient mice. Generating rax-/- mice lacking PKR or carrying kinase-inactive PKR(K271R) rescues the developmental defects, demonstrating that the rax-/- phenotype is PKR kinase-activity dependent. In vitro reduction of RAX in anterior pituitary cells increases PKR activity and p21WAF1/CIP1 induction. Double-knockout mouse genetics, kinase-inactive PKR knockin, eIF2α S51A knockin, in vitro RAX knockdown in pituitary cells The FEBS Journal High 26414443
2015 Ethanol-induced neurotoxicity in the developing cerebellum is mediated through RAX/PKR association. N-PKR-/- mice (lacking the RAX-binding domain of PKR) show reduced ethanol-induced brain/body mass reduction, less cerebellar neuronal loss, and attenuated IL-1β secretion compared to wild-type mice following ethanol exposure, confirming that RAX-PKR physical interaction is required for PKR-regulated ethanol neurotoxicity in vivo. N-PKR-/- knockin mice, ethanol exposure model, cerebellar neuronal counting, IL-1β ELISA Cerebellum High 25592072
2016 Rax is essential for early patterning of the mammalian hypothalamus. Conditional inactivation of Rax prior to E8.5 in mice causes general underdevelopment of the hypothalamic neuroepithelium, impaired neurogenesis, loss of dorsomedial hypothalamic markers (POMC, Somatostatin), and invasion of ventromedial/infundibular gene expression domains into dorsal hypothalamic territory, demonstrating Rax is required for dorsoventral patterning. Loss of Rax also causes downregulation of Shh in the dorsomedial hypothalamus, phenocopying Shh conditional knockout. Conditional knockout (tamoxifen timing), immunostaining, in situ hybridization for neuropeptides and patterning genes Developmental Biology High 27212025
1998 PACT (human ortholog of mouse RAX) was identified as the first cellular protein activator of PKR. PACT heterodimerizes with PKR and activates it in vitro in the absence of dsRNA. In mammalian cells, PACT overexpression causes PKR activation; in yeast, PACT co-expression enhances the anti-growth effect of PKR. In vitro PKR activation assay, co-immunoprecipitation/heterodimerization, yeast growth assay, mammalian overexpression The EMBO journal High 9687506
2006 PACT (human RAX ortholog) is a component of the RNA-induced silencing complex (RISC). PACT associates with an ~500 kDa complex containing Dicer, hAgo2, and TRBP. The interaction with Dicer involves the third dsRNA-binding domain (dsRBD) of PACT and the N-terminal helicase region of Dicer. PACT depletion strongly reduces mature miRNA accumulation in vivo and moderately reduces siRNA-induced RNAi efficiency. Co-immunoprecipitation, domain mapping, siRNA depletion, miRNA Northern blotting The EMBO Journal High 16424907
2021 Rax+ tanycytes in the median eminence are largely quiescent but rapidly enter the cell cycle upon neural injury for self-renewal and regeneration. IGF1 receptor (Igf1r) signaling in tanycytes is required for tissue repair under injury conditions. Oncogenic BRAF activation (BrafV600E) is sufficient to transform Rax+ tanycytes into actively dividing tumor cells that develop into papillary craniopharyngioma-like tumors. Rax-CreERT2 lineage tracing, neural injury model, Igf1r conditional knockout, BrafV600E conditional activation, BrdU/EdU proliferation assays Nature Communications High 33863883

Source papers

Stage 0 corpus · 130 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2006 Global, in vivo, and site-specific phosphorylation dynamics in signaling networks. Cell 2861 17081983
2005 A human protein-protein interaction network: a resource for annotating the proteome. Cell 1704 16169070
2002 Generation and initial analysis of more than 15,000 full-length human and mouse cDNA sequences. Proceedings of the National Academy of Sciences of the United States of America 1479 12477932
2016 ATPase-Modulated Stress Granules Contain a Diverse Proteome and Substructure. Cell 1233 26777405
2015 The BioPlex Network: A Systematic Exploration of the Human Interactome. Cell 1118 26186194
2017 Architecture of the human interactome defines protein communities and disease networks. Nature 1085 28514442
2015 A human interactome in three quantitative dimensions organized by stoichiometries and abundances. Cell 1015 26496610
2014 A proteome-scale map of the human interactome network. Cell 977 25416956
2012 The mRNA-bound proteome and its global occupancy profile on protein-coding transcripts. Molecular cell 973 22681889
2020 A reference map of the human binary protein interactome. Nature 849 32296183
2003 Complete sequencing and characterization of 21,243 full-length human cDNAs. Nature genetics 754 14702039
2007 Large-scale mapping of human protein-protein interactions by mass spectrometry. Molecular systems biology 733 17353931
2001 TIRAP: an adapter molecule in the Toll signaling pathway. Nature immunology 729 11526399
2021 Dual proteome-scale networks reveal cell-specific remodeling of the human interactome. Cell 705 33961781
2011 Phylogenetic-based propagation of functional annotations within the Gene Ontology consortium. Briefings in bioinformatics 656 21873635
2018 High-Density Proximity Mapping Reveals the Subcellular Organization of mRNA-Associated Granules and Bodies. Molecular cell 580 29395067
1997 The Rx homeobox gene is essential for vertebrate eye development. Nature 574 9177348
2020 Comparative host-coronavirus protein interaction networks reveal pan-viral disease mechanisms. Science (New York, N.Y.) 564 33060197
2017 Anticancer sulfonamides target splicing by inducing RBM39 degradation via recruitment to DCAF15. Science (New York, N.Y.) 533 28302793
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