{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,3]],"date-time":"2026-07-03T00:02:31Z","timestamp":1783036951295,"version":"3.54.6"},"reference-count":90,"publisher":"The Company of Biologists","issue":"2","license":[{"start":{"date-parts":[[2026,1,23]],"date-time":"2026-01-23T00:00:00Z","timestamp":1769126400000},"content-version":"vor","delay-in-days":8,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100007137","name":"Helse Nord RHF","doi-asserted-by":"crossref","award":["HNF1546-20"],"award-info":[{"award-number":["HNF1546-20"]}],"id":[{"id":"10.13039\/501100007137","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100007465","name":"UiT The Arctic University of Norway","doi-asserted-by":"crossref","id":[{"id":"10.13039\/100007465","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,1,15]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>Paraspeckles are stress-induced nuclear RNA\u2013protein condensates that assemble on the long non-coding RNA NEAT1. Their increased formation under certain cellular circumstances has gained growing interest due to their association with serious human diseases, such as neurodegenerative disorders and cancer. The biological functions of paraspeckles still appear obscure, but increasing evidence suggests that they contribute to regulation of gene expression by recruiting specific proteins and RNA molecules. Here, we have characterized and compared two stress-enriched interactomes of the essential paraspeckle protein NONO in both wild-type and paraspeckle-deficient NEAT1 knockout cells. We identified Hsp70 as part of stress-enriched NONO complexes in wild-type but not in NEAT1-depleted cells. We show that proteotoxic stress-induced paraspeckle formation and NEAT1 expression are strictly dependent on Hsp70 chaperone activity. Our data demonstrate that both NONO and Hsp70 transiently translocate to the nucleolus during heat shock and that paraspeckle formation during recovery follows Hsp70-dependent relocation of NONO from the nucleolus to the nucleoplasm. Taken together, we demonstrate an important role of Hsp70 in paraspeckle assembly and identify a possible link between the nuclear protein quality control system and paraspeckles.<\/jats:p>","DOI":"10.1242\/jcs.264115","type":"journal-article","created":{"date-parts":[[2025,12,19]],"date-time":"2025-12-19T09:54:29Z","timestamp":1766138069000},"source":"Crossref","is-referenced-by-count":1,"title":["Stress-specific NONO interactomes reveal a key role of Hsp70 chaperone activity in regulation of paraspeckle formation"],"prefix":"10.1242","volume":"139","author":[{"given":"Isaac","family":"Odonkor","sequence":"first","affiliation":[{"name":"UiT the Arctic University of Norway 1 Department of Medical Biology, Faculty of Health Sciences , , 9037 Troms\u00f8 ,","place":["Norway"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Birendra Kumar","family":"Shrestha","sequence":"additional","affiliation":[{"name":"UiT the Arctic University of Norway 1 Department of Medical Biology, Faculty of Health Sciences , , 9037 Troms\u00f8 ,","place":["Norway"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Stephanie Rose","family":"Nielsen","sequence":"additional","affiliation":[{"name":"UiT the Arctic University of Norway 1 Department of Medical Biology, Faculty of Health Sciences , , 9037 Troms\u00f8 ,","place":["Norway"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Athanasios","family":"Kournoutis","sequence":"additional","affiliation":[{"name":"UiT the Arctic University of Norway 1 Department of Medical Biology, Faculty of Health Sciences , , 9037 Troms\u00f8 ,","place":["Norway"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ida Emilie","family":"Bj\u00f8rlo","sequence":"additional","affiliation":[{"name":"UiT the Arctic University of Norway 1 Department of Medical Biology, Faculty of Health Sciences , , 9037 Troms\u00f8 ,","place":["Norway"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Saikat","family":"Das Sajib","sequence":"additional","affiliation":[{"name":"UiT the Arctic University of Norway 1 Department of Medical Biology, Faculty of Health Sciences , , 9037 Troms\u00f8 ,","place":["Norway"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Annica","family":"Hedberg","sequence":"additional","affiliation":[{"name":"UiT the Arctic University of Norway 1 Department of Medical Biology, Faculty of Health Sciences , , 9037 Troms\u00f8 ,","place":["Norway"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Toril Anne","family":"Gr\u00f8nset","sequence":"additional","affiliation":[{"name":"UiT the Arctic University of Norway 1 Department of Medical Biology, Faculty of Health Sciences , , 9037 Troms\u00f8 ,","place":["Norway"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kenneth Bowitz","family":"Larsen","sequence":"additional","affiliation":[{"name":"UiT the Arctic University of Norway 1 Department of Medical Biology, Faculty of Health Sciences , , 9037 Troms\u00f8 ,","place":["Norway"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jack-Ansgar","family":"Bruun","sequence":"additional","affiliation":[{"name":"UiT the Arctic University of Norway 1 Department of Medical Biology, Faculty of Health Sciences , , 9037 Troms\u00f8 ,","place":["Norway"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Erik","family":"Knutsen","sequence":"additional","affiliation":[{"name":"UiT the Arctic University of Norway 1 Department of Medical Biology, Faculty of Health Sciences , , 9037 Troms\u00f8 ,","place":["Norway"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Seyed Mohammad","family":"Lellahi","sequence":"additional","affiliation":[{"name":"UiT the Arctic University of Norway 1 Department of Medical Biology, Faculty of Health Sciences , , 9037 Troms\u00f8 ,","place":["Norway"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maria","family":"Perander","sequence":"additional","affiliation":[{"name":"UiT the Arctic University of Norway 1 Department of Medical Biology, Faculty of Health Sciences , , 9037 Troms\u00f8 ,","place":["Norway"]},{"name":"Centre for Clinical Research and Education, University Hospital of North Norway 2 , 9038 Troms\u00f8 ,","place":["Norway"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","published-online":{"date-parts":[[2026,1,23]]},"reference":[{"key":"2026012305022997800_JCS264115C1","doi-asserted-by":"publisher","first-page":"861","DOI":"10.1038\/nm.4135","article-title":"p53 induces formation of NEAT1 lncRNA-containing paraspeckles that modulate replication stress response and chemosensitivity","volume":"22","author":"Adriaens","year":"2016","journal-title":"Nat. Med."},{"key":"2026012305022997800_JCS264115C2","doi-asserted-by":"publisher","first-page":"1681","DOI":"10.1261\/rna.071456.119","article-title":"The long noncoding RNA NEAT1_1 is seemingly dispensable for normal tissue homeostasis and cancer cell growth","volume":"25","author":"Adriaens","year":"2019","journal-title":"RNA"},{"key":"2026012305022997800_JCS264115C3","doi-asserted-by":"publisher","first-page":"115173","DOI":"10.1016\/j.celrep.2024.115173","article-title":"C9ORF72 poly-PR induces TDP-43 nuclear condensation via NEAT1 and is modulated by HSP70 activity","volume":"44","author":"Agnihotri","year":"2025","journal-title":"Cell Rep."},{"key":"2026012305022997800_JCS264115C4","doi-asserted-by":"publisher","first-page":"3271","DOI":"10.1111\/febs.17390","article-title":"Regulation of physiological and pathological condensates by molecular chaperones","volume":"292","author":"Akaree","year":"2025","journal-title":"FEBS J."},{"key":"2026012305022997800_JCS264115C5","doi-asserted-by":"publisher","first-page":"3557","DOI":"10.1242\/jcs.00671","article-title":"Formation of nuclear stress granules involves HSF2 and coincides with the nucleolar localization of Hsp70","volume":"116","author":"Alastalo","year":"2003","journal-title":"J. Cell Sci."},{"key":"2026012305022997800_JCS264115C6","doi-asserted-by":"publisher","first-page":"196","DOI":"10.1038\/s41580-020-00326-6","article-title":"Biomolecular condensates at the nexus of cellular stress, protein aggregation disease and ageing","volume":"22","author":"Alberti","year":"2021","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"2026012305022997800_JCS264115C7","doi-asserted-by":"publisher","first-page":"849","DOI":"10.1016\/j.tcb.2019.08.002","article-title":"The nucleolus as a proteostasis regulator","volume":"29","author":"Amer-Sarsour","year":"2019","journal-title":"Trends Cell Biol."},{"key":"2026012305022997800_JCS264115C8","doi-asserted-by":"publisher","first-page":"4127","DOI":"10.1083\/jcb.201904098","article-title":"Stress granules regulate stress-induced paraspeckle assembly","volume":"218","author":"An","year":"2019","journal-title":"J. Cell Biol."},{"key":"2026012305022997800_JCS264115C9","doi-asserted-by":"publisher","first-page":"119058","DOI":"10.1016\/j.bbamcr.2021.119058","article-title":"Connecting the \u201cdots\u201d: RNP granule network in health and disease","volume":"1868","author":"An","year":"2021","journal-title":"Biochim. Biophys. Acta Mol. Cell Res."},{"key":"2026012305022997800_JCS264115C10","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/S0960-9822(01)00650-9","article-title":"Directed proteomic analysis of the human nucleolus","volume":"12","author":"Andersen","year":"2002","journal-title":"Curr. Biol."},{"key":"2026012305022997800_JCS264115C11","doi-asserted-by":"publisher","first-page":"e45205","DOI":"10.7554\/eLife.45205","article-title":"Protein quality control in the nucleolus safeguards recovery of epigenetic regulators after heat shock","volume":"8","author":"Azkanaz","year":"2019","journal-title":"eLife"},{"key":"2026012305022997800_JCS264115C12","doi-asserted-by":"publisher","first-page":"285","DOI":"10.1038\/nrm.2017.7","article-title":"Biomolecular condensates: organizers of cellular biochemistry","volume":"18","author":"Banani","year":"2017","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"2026012305022997800_JCS264115C13","doi-asserted-by":"publisher","first-page":"10","DOI":"10.3389\/fmolb.2015.00010","article-title":"The nucleotide exchange factors of Hsp70 molecular chaperones","volume":"2","author":"Bracher","year":"2015","journal-title":"Front. Mol. Biosci."},{"key":"2026012305022997800_JCS264115C14","doi-asserted-by":"publisher","first-page":"880","DOI":"10.1038\/nbt.4201","article-title":"Efficient proximity labeling in living cells and organisms with TurboID","volume":"36","author":"Branon","year":"2018","journal-title":"Nat. Biotechnol."},{"key":"2026012305022997800_JCS264115C15","doi-asserted-by":"publisher","first-page":"467","DOI":"10.1016\/j.molcel.2009.06.027","article-title":"Altered nuclear retention of mRNAs containing inverted repeats in human embryonic stem cells: functional role of a nuclear noncoding RNA","volume":"35","author":"Chen","year":"2009","journal-title":"Mol. Cell"},{"key":"2026012305022997800_JCS264115C16","doi-asserted-by":"publisher","first-page":"3971","DOI":"10.1038\/s41596-020-0399-0","article-title":"Proximity labeling in mammalian cells with TurboID and split-TurboID","volume":"15","author":"Cho","year":"2020","journal-title":"Nat. Protoc."},{"key":"2026012305022997800_JCS264115C17","doi-asserted-by":"publisher","first-page":"4482","DOI":"10.1038\/onc.2014.378","article-title":"Tumor hypoxia induces nuclear paraspeckle formation through HIF-2alpha dependent transcriptional activation of NEAT1 leading to cancer cell survival","volume":"34","author":"Choudhry","year":"2015","journal-title":"Oncogene"},{"key":"2026012305022997800_JCS264115C18","doi-asserted-by":"publisher","first-page":"717","DOI":"10.1016\/j.molcel.2009.01.026","article-title":"An architectural role for a nuclear noncoding RNA: NEAT1 RNA is essential for the structure of paraspeckles","volume":"33","author":"Clemson","year":"2009","journal-title":"Mol. Cell"},{"key":"2026012305022997800_JCS264115C19","doi-asserted-by":"publisher","first-page":"289","DOI":"10.1016\/bs.apcsb.2019.08.002","article-title":"Liquid-liquid phase transitions and amyloid aggregation in proteins related to cancer and neurodegenerative diseases","volume":"118","author":"de Oliveira","year":"2019","journal-title":"Adv. Protein Chem. Struct. Biol."},{"key":"2026012305022997800_JCS264115C20","doi-asserted-by":"publisher","first-page":"4085","DOI":"10.1093\/nar\/21.17.4085","article-title":"Purification and cDNA cloning of HeLa cell p54nrb, a nuclear protein with two RNA recognition motifs and extensive homology to human splicing factor PSF and Drosophila NONA\/BJ6","volume":"21","author":"Dong","year":"1993","journal-title":"Nucleic Acids Res."},{"key":"2026012305022997800_JCS264115C21","doi-asserted-by":"publisher","first-page":"2024.07.03.601239","DOI":"10.1101\/2024.07.03.601239","article-title":"Spatial proteomic mapping of human nuclear bodies reveals new functional insights into RNA regulation","author":"Dyakov","year":"2024","journal-title":"bioRxiv"},{"key":"2026012305022997800_JCS264115C22","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1016\/S0960-9822(01)00632-7","article-title":"Paraspeckles: a novel nuclear domain","volume":"12","author":"Fox","year":"2002","journal-title":"Curr. Biol."},{"key":"2026012305022997800_JCS264115C23","doi-asserted-by":"publisher","first-page":"5304","DOI":"10.1091\/mbc.e05-06-0587","article-title":"P54nrb forms a heterodimer with PSP1 that localizes to paraspeckles in an RNA-dependent manner","volume":"16","author":"Fox","year":"2005","journal-title":"Mol. Biol. Cell"},{"key":"2026012305022997800_JCS264115C24","doi-asserted-by":"publisher","first-page":"124","DOI":"10.1016\/j.tibs.2017.12.001","article-title":"Paraspeckles: where long noncoding RNA meets phase separation","volume":"43","author":"Fox","year":"2018","journal-title":"Trends Biochem. Sci."},{"key":"2026012305022997800_JCS264115C25","doi-asserted-by":"publisher","first-page":"7128","DOI":"10.1074\/jbc.TM118.001190","article-title":"Prion-like low-complexity sequences: key regulators of protein solubility and phase behavior","volume":"294","author":"Franzmann","year":"2019","journal-title":"J. Biol. Chem."},{"key":"2026012305022997800_JCS264115C26","doi-asserted-by":"publisher","first-page":"342","DOI":"10.1126\/science.aaw9157","article-title":"The nucleolus functions as a phase-separated protein quality control compartment","volume":"365","author":"Frottin","year":"2019","journal-title":"Science"},{"key":"2026012305022997800_JCS264115C27","doi-asserted-by":"publisher","first-page":"796","DOI":"10.1016\/j.molcel.2016.07.021","article-title":"A surveillance function of the HSPB8-BAG3-HSP70 chaperone complex ensures stress granule integrity and dynamism","volume":"63","author":"Ganassi","year":"2016","journal-title":"Mol. Cell"},{"key":"2026012305022997800_JCS264115C28","doi-asserted-by":"publisher","first-page":"4","DOI":"10.1038\/nrm.2017.73","article-title":"Regulation of heat shock transcription factors and their roles in physiology and disease","volume":"19","author":"Gomez-Pastor","year":"2018","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"2026012305022997800_JCS264115C29","doi-asserted-by":"publisher","first-page":"1024","DOI":"10.1038\/s41422-021-00526-5","article-title":"Hsp70 chaperones TDP-43 in dynamic, liquid-like phase and prevents it from amyloid aggregation","volume":"31","author":"Gu","year":"2021","journal-title":"Cell Res."},{"key":"2026012305022997800_JCS264115C30","doi-asserted-by":"publisher","first-page":"1593","DOI":"10.1016\/j.chembiol.2024.08.009","article-title":"RNA and condensates: disease implications and therapeutic opportunities","volume":"31","author":"Han","year":"2024","journal-title":"Cell Chem. Biol."},{"key":"2026012305022997800_JCS264115C31","doi-asserted-by":"publisher","first-page":"529","DOI":"10.1083\/jcb.201504117","article-title":"Prion-like domains in RNA binding proteins are essential for building subnuclear paraspeckles","volume":"210","author":"Hennig","year":"2015","journal-title":"J. Cell Biol."},{"key":"2026012305022997800_JCS264115C32","doi-asserted-by":"publisher","first-page":"jcs238402","DOI":"10.1242\/jcs.238402","article-title":"New insights into transcriptional reprogramming during cellular stress","volume":"132","author":"Himanen","year":"2019","journal-title":"J. Cell Sci."},{"key":"2026012305022997800_JCS264115C33","doi-asserted-by":"publisher","first-page":"169","DOI":"10.1091\/mbc.e13-09-0558","article-title":"NEAT1 long noncoding RNA regulates transcription via protein sequestration within subnuclear bodies","volume":"25","author":"Hirose","year":"2014","journal-title":"Mol. Biol. Cell"},{"key":"2026012305022997800_JCS264115C34","doi-asserted-by":"publisher","first-page":"288","DOI":"10.1038\/s41580-022-00558-8","article-title":"A guide to membraneless organelles and their various roles in gene regulation","volume":"24","author":"Hirose","year":"2023","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"2026012305022997800_JCS264115C35","doi-asserted-by":"publisher","first-page":"6593","DOI":"10.1074\/jbc.RA117.001451","article-title":"Crystal structure of a SFPQ\/PSPC1 heterodimer provides insights into preferential heterodimerization of human DBHS family proteins","volume":"293","author":"Huang","year":"2018","journal-title":"J. Biol. Chem."},{"key":"2026012305022997800_JCS264115C36","doi-asserted-by":"publisher","first-page":"1902","DOI":"10.1016\/j.cell.2018.11.027","article-title":"CARM1 and paraspeckles regulate pre-implantation mouse embryo development","volume":"175","author":"Hupalowska","year":"2018","journal-title":"Cell"},{"key":"2026012305022997800_JCS264115C37","doi-asserted-by":"publisher","first-page":"39","DOI":"10.1146\/annurev-cellbio-100913-013325","article-title":"Liquid-liquid phase separation in biology","volume":"30","author":"Hyman","year":"2014","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"2026012305022997800_JCS264115C38","doi-asserted-by":"publisher","first-page":"393","DOI":"10.1016\/j.molcel.2014.01.009","article-title":"Long noncoding RNA NEAT1-dependent SFPQ relocation from promoter region to paraspeckle mediates IL8 expression upon immune stimuli","volume":"53","author":"Imamura","year":"2014","journal-title":"Mol. Cell"},{"key":"2026012305022997800_JCS264115C39","doi-asserted-by":"publisher","first-page":"a034066","DOI":"10.1101\/cshperspect.a034066","article-title":"Tailoring of proteostasis networks with heat shock factors","volume":"11","author":"Joutsen","year":"2019","journal-title":"Cold Spring Harb. Perspect Biol."},{"key":"2026012305022997800_JCS264115C40","doi-asserted-by":"publisher","first-page":"105","DOI":"10.1007\/s12192-008-0068-7","article-title":"Guidelines for the nomenclature of the human heat shock proteins","volume":"14","author":"Kampinga","year":"2009","journal-title":"Cell Stress Chaperones"},{"key":"2026012305022997800_JCS264115C41","doi-asserted-by":"publisher","first-page":"7","DOI":"10.1007\/s12192-018-0948-4","article-title":"Function, evolution, and structure of J-domain proteins","volume":"24","author":"Kampinga","year":"2019","journal-title":"Cell Stress Chaperones"},{"key":"2026012305022997800_JCS264115C42","doi-asserted-by":"publisher","first-page":"fcaf261","DOI":"10.1093\/braincomms\/fcaf261","article-title":"Downregulation of NEAT1 due to loss of TDP-43 function exacerbates motor neuron degeneration in amyotrophic lateral sclerosis","volume":"7","author":"Kawakami","year":"2025","journal-title":"Brain Commun."},{"key":"2026012305022997800_JCS264115C43","doi-asserted-by":"publisher","first-page":"3989","DOI":"10.1093\/nar\/gkw271","article-title":"The DBHS proteins SFPQ, NONO and PSPC1: a multipurpose molecular scaffold","volume":"44","author":"Knott","year":"2016","journal-title":"Nucleic Acids Res."},{"key":"2026012305022997800_JCS264115C44","doi-asserted-by":"publisher","first-page":"578","DOI":"10.1016\/j.cell.2012.02.058","article-title":"Hikeshi, a nuclear import carrier for Hsp70s, protects cells from heat shock-induced nuclear damage","volume":"149","author":"Kose","year":"2012","journal-title":"Cell"},{"key":"2026012305022997800_JCS264115C45","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1038\/s41580-020-0272-6","article-title":"The nucleolus as a multiphase liquid condensate","volume":"22","author":"Lafontaine","year":"2021","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"2026012305022997800_JCS264115C46","doi-asserted-by":"publisher","first-page":"18965","DOI":"10.1074\/jbc.RA118.004473","article-title":"The long noncoding RNA NEAT1 and nuclear paraspeckles are up-regulated by the transcription factor HSF1 in the heat shock response","volume":"293","author":"Lellahi","year":"2018","journal-title":"J. Biol. Chem."},{"key":"2026012305022997800_JCS264115C47","doi-asserted-by":"publisher","first-page":"872","DOI":"10.1261\/rna.059477.116","article-title":"Functional dissection of NEAT1 using genome editing reveals substantial localization of the NEAT1_1 isoform outside paraspeckles","volume":"23","author":"Li","year":"2017","journal-title":"RNA"},{"key":"2026012305022997800_JCS264115C48","doi-asserted-by":"publisher","first-page":"104356","DOI":"10.1016\/j.isci.2022.104356","article-title":"Hsp70 exhibits a liquid-liquid phase separation ability and chaperones condensed FUS against amyloid aggregation","volume":"25","author":"Li","year":"2022","journal-title":"iScience"},{"key":"2026012305022997800_JCS264115C49","doi-asserted-by":"publisher","first-page":"1378","DOI":"10.1038\/s41556-022-00988-8","article-title":"Heat-shock chaperone HSPB1 regulates cytoplasmic TDP-43 phase separation and liquid-to-gel transition","volume":"24","author":"Lu","year":"2022","journal-title":"Nat. Cell Biol."},{"key":"2026012305022997800_JCS264115C50","doi-asserted-by":"publisher","first-page":"215","DOI":"10.1038\/s41580-020-00303-z","article-title":"A framework for understanding the functions of biomolecular condensates across scales","volume":"22","author":"Lyon","year":"2021","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"2026012305022997800_JCS264115C51","doi-asserted-by":"publisher","first-page":"1669","DOI":"10.15252\/embj.201695957","article-title":"An aberrant phase transition of stress granules triggered by misfolded protein and prevented by chaperone function","volume":"36","author":"Mateju","year":"2017","journal-title":"EMBO J."},{"key":"2026012305022997800_JCS264115C52","doi-asserted-by":"publisher","first-page":"670","DOI":"10.1007\/s00018-004-4464-6","article-title":"Hsp70 chaperones: cellular functions and molecular mechanism","volume":"62","author":"Mayer","year":"2005","journal-title":"Cell. Mol. Life Sci."},{"key":"2026012305022997800_JCS264115C53","doi-asserted-by":"publisher","first-page":"2603","DOI":"10.1091\/mbc.e06-12-1079","article-title":"Inhibition of the ubiquitin-proteasome system induces stress granule formation","volume":"18","author":"Mazroui","year":"2007","journal-title":"Mol. Biol. Cell"},{"key":"2026012305022997800_JCS264115C54","doi-asserted-by":"publisher","first-page":"e2000245","DOI":"10.1002\/bies.202000245","article-title":"Paraspeckle nuclear condensates: global sensors of cell stress?","volume":"43","author":"Mccluggage","year":"2021","journal-title":"BioEssays"},{"key":"2026012305022997800_JCS264115C55","doi-asserted-by":"publisher","first-page":"951","DOI":"10.1016\/j.molcel.2019.03.041","article-title":"Cross-regulation between TDP-43 and paraspeckles promotes pluripotency-differentiation transition","volume":"74","author":"Modic","year":"2019","journal-title":"Mol. Cell"},{"key":"2026012305022997800_JCS264115C56","doi-asserted-by":"publisher","first-page":"214","DOI":"10.1016\/j.molcel.2018.01.004","article-title":"Cellular handling of protein aggregates by disaggregation machines","volume":"69","author":"Mogk","year":"2018","journal-title":"Mol. Cell"},{"key":"2026012305022997800_JCS264115C57","doi-asserted-by":"publisher","first-page":"M111.009241","DOI":"10.1074\/mcp.M111.009241","article-title":"Quantitative proteomics and dynamic imaging of the nucleolus reveal distinct responses to UV and ionizing radiation","volume":"10","author":"Moore","year":"2011","journal-title":"Mol. Cell. Proteomics"},{"key":"2026012305022997800_JCS264115C58","doi-asserted-by":"publisher","first-page":"4020","DOI":"10.1038\/emboj.2012.251","article-title":"Alternative 3'-end processing of long noncoding RNA initiates construction of nuclear paraspeckles","volume":"31","author":"Naganuma","year":"2012","journal-title":"EMBO J."},{"key":"2026012305022997800_JCS264115C59","doi-asserted-by":"publisher","first-page":"31","DOI":"10.1083\/jcb.201011110","article-title":"Paraspeckles are subpopulation-specific nuclear bodies that are not essential in mice","volume":"193","author":"Nakagawa","year":"2011","journal-title":"J. Cell Biol."},{"key":"2026012305022997800_JCS264115C60","doi-asserted-by":"publisher","first-page":"4618","DOI":"10.1242\/dev.110544","article-title":"The lncRNA Neat1 is required for corpus luteum formation and the establishment of pregnancy in a subpopulation of mice","volume":"141","author":"Nakagawa","year":"2014","journal-title":"Development"},{"key":"2026012305022997800_JCS264115C61","doi-asserted-by":"publisher","first-page":"12038","DOI":"10.1073\/pnas.201112398","article-title":"Dynamic changes in the localization of thermally unfolded nuclear proteins associated with chaperone-dependent protection","volume":"98","author":"Nollen","year":"2001","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2026012305022997800_JCS264115C62","doi-asserted-by":"publisher","first-page":"3095","DOI":"10.1002\/j.1460-2075.1984.tb02264.x","article-title":"Hsp70 accelerates the recovery of nucleolar morphology after heat shock","volume":"3","author":"Pelham","year":"1984","journal-title":"EMBO J."},{"key":"2026012305022997800_JCS264115C63","doi-asserted-by":"publisher","first-page":"D543","DOI":"10.1093\/nar\/gkae1011","article-title":"The PRIDE database at 20 years: 2025 update","volume":"53","author":"Perez-Riverol","year":"2025","journal-title":"Nucleic Acids Res."},{"key":"2026012305022997800_JCS264115C64","doi-asserted-by":"publisher","first-page":"354","DOI":"10.1101\/gad.350518.123","article-title":"RNA granules: functional compartments or incidental condensates?","volume":"37","author":"Putnam","year":"2023","journal-title":"Genes Dev."},{"key":"2026012305022997800_JCS264115C65","doi-asserted-by":"publisher","first-page":"5775","DOI":"10.1016\/j.cell.2021.10.014","article-title":"RNA promotes the formation of spatial compartments in the nucleus","volume":"184","author":"Quinodoz","year":"2021","journal-title":"Cell"},{"key":"2026012305022997800_JCS264115C66","doi-asserted-by":"publisher","first-page":"2281","DOI":"10.1038\/nprot.2013.143","article-title":"Genome engineering using the CRISPR-Cas9 system","volume":"8","author":"Ran","year":"2013","journal-title":"Nat. Protoc."},{"key":"2026012305022997800_JCS264115C67","doi-asserted-by":"publisher","first-page":"183","DOI":"10.1038\/s41580-020-0264-6","article-title":"RNA contributions to the form and function of biomolecular condensates","volume":"22","author":"Roden","year":"2021","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"2026012305022997800_JCS264115C68","doi-asserted-by":"publisher","first-page":"665","DOI":"10.1038\/s41580-019-0133-3","article-title":"The Hsp70 chaperone network","volume":"20","author":"Rosenzweig","year":"2019","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"2026012305022997800_JCS264115C69","doi-asserted-by":"publisher","first-page":"961","DOI":"10.1016\/j.tibs.2020.06.007","article-title":"Biomolecular Condensates in the Nucleus","volume":"45","author":"Sabari","year":"2020","journal-title":"Trends Biochem. Sci."},{"key":"2026012305022997800_JCS264115C70","doi-asserted-by":"publisher","first-page":"1231","DOI":"10.1083\/jcb.201612111","article-title":"Shaping proteostasis at the cellular, tissue, and organismal level","volume":"216","author":"Sala","year":"2017","journal-title":"J. Cell Biol."},{"key":"2026012305022997800_JCS264115C71","doi-asserted-by":"publisher","first-page":"2525","DOI":"10.1073\/pnas.0807899106","article-title":"MENepsilon\/beta noncoding RNAs are essential for structural integrity of nuclear paraspeckles","volume":"106","author":"Sasaki","year":"2009","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2026012305022997800_JCS264115C72","doi-asserted-by":"publisher","first-page":"52","DOI":"10.1261\/rna.078997.121","article-title":"RNA in formation and regulation of transcriptional condensates","volume":"28","author":"Sharp","year":"2022","journal-title":"RNA"},{"key":"2026012305022997800_JCS264115C73","doi-asserted-by":"publisher","first-page":"2395","DOI":"10.1091\/mbc.e04-11-0992","article-title":"Dynamic sorting of nuclear components into distinct nucleolar caps during transcriptional inhibition","volume":"16","author":"Shav-Tal","year":"2005","journal-title":"Mol. Biol. Cell"},{"key":"2026012305022997800_JCS264115C74","doi-asserted-by":"publisher","first-page":"eaaf4382","DOI":"10.1126\/science.aaf4382","article-title":"Liquid phase condensation in cell physiology and disease","volume":"357","author":"Shin","year":"2017","journal-title":"Science"},{"key":"2026012305022997800_JCS264115C75","doi-asserted-by":"publisher","first-page":"1844","DOI":"10.1261\/rna.047332.114","article-title":"The long noncoding RNA Neat1 is required for mammary gland development and lactation","volume":"20","author":"Standaert","year":"2014","journal-title":"RNA"},{"key":"2026012305022997800_JCS264115C76","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1101\/gr.087775.108","article-title":"MEN epsilon\/beta nuclear-retained non-coding RNAs are up-regulated upon muscle differentiation and are essential components of paraspeckles","volume":"19","author":"Sunwoo","year":"2009","journal-title":"Genome Res."},{"key":"2026012305022997800_JCS264115C77","doi-asserted-by":"publisher","first-page":"1011","DOI":"10.1261\/rna.079972.124","article-title":"The essential role of architectural noncoding RNA Neat1 in cold-induced beige adipocyte differentiation in mice","volume":"30","author":"Toya","year":"2024","journal-title":"RNA"},{"key":"2026012305022997800_JCS264115C78","doi-asserted-by":"publisher","first-page":"914873","DOI":"10.3389\/fmolb.2022.914873","article-title":"In vivo proximity labeling of nuclear and nucleolar proteins by a stably expressed, DNA damage-responsive NONO-APEX2 fusion protein","volume":"9","author":"Trifault","year":"2022","journal-title":"Front. Mol. Biosci."},{"key":"2026012305022997800_JCS264115C79","doi-asserted-by":"publisher","first-page":"3050","DOI":"10.1093\/nar\/gkae022","article-title":"Nucleolar detention of NONO shields DNA double-strand breaks from aberrant transcripts","volume":"52","author":"Trifault","year":"2024","journal-title":"Nucleic Acids Res."},{"key":"2026012305022997800_JCS264115C80","doi-asserted-by":"publisher","first-page":"655","DOI":"10.1016\/0092-8674(84)90345-3","article-title":"hsp70: nuclear concentration during environmental stress and cytoplasmic storage during recovery","volume":"36","author":"Velazquez","year":"1984","journal-title":"Cell"},{"key":"2026012305022997800_JCS264115C81","doi-asserted-by":"publisher","first-page":"1145","DOI":"10.1038\/s41556-018-0204-2","article-title":"Genome-wide screening of NEAT1 regulators reveals cross-regulation between paraspeckles and mitochondria","volume":"20","author":"Wang","year":"2018","journal-title":"Nat. Cell Biol."},{"key":"2026012305022997800_JCS264115C82","doi-asserted-by":"publisher","first-page":"443","DOI":"10.1016\/j.molcel.2020.06.019","article-title":"Stress induces dynamic, cytotoxicity-antagonizing TDP-43 nuclear bodies via paraspeckle LncRNA NEAT1-mediated liquid-liquid phase separation","volume":"79","author":"Wang","year":"2020","journal-title":"Mol. Cell"},{"key":"2026012305022997800_JCS264115C83","doi-asserted-by":"publisher","first-page":"4501","DOI":"10.1016\/S0021-9258(17)43075-4","article-title":"Nuclear and nucleolar localization of the 72,000-dalton heat shock protein in heat-shocked mammalian cells","volume":"259","author":"Welch","year":"1984","journal-title":"J. Biol. Chem."},{"key":"2026012305022997800_JCS264115C84","doi-asserted-by":"publisher","first-page":"102735","DOI":"10.1016\/j.sbi.2023.102735","article-title":"Protein disaggregation machineries in the human cytosol","volume":"83","author":"Wentink","year":"2023","journal-title":"Curr. Opin. Struct. Biol."},{"key":"2026012305022997800_JCS264115C85","doi-asserted-by":"publisher","first-page":"1","DOI":"10.2741\/s420","article-title":"The building process of the functional paraspeckle with long non-coding RNAs","volume":"7","author":"Yamazaki","year":"2015","journal-title":"Front. Biosci. (Elite Ed)"},{"key":"2026012305022997800_JCS264115C86","doi-asserted-by":"publisher","first-page":"1038","DOI":"10.1016\/j.molcel.2018.05.019","article-title":"Functional domains of NEAT1 architectural lncRNA induce paraspeckle assembly through phase separation","volume":"70","author":"Yamazaki","year":"2018","journal-title":"Mol. Cell"},{"key":"2026012305022997800_JCS264115C87","doi-asserted-by":"publisher","first-page":"2738","DOI":"10.1016\/j.molcel.2022.05.010","article-title":"Condensates induced by transcription inhibition localize active chromatin to nucleoli","volume":"82","author":"Yasuhara","year":"2022","journal-title":"Mol. Cell"},{"key":"2026012305022997800_JCS264115C88","doi-asserted-by":"publisher","first-page":"eabb4309","DOI":"10.1126\/science.abb4309","article-title":"HSP70 chaperones RNA-free TDP-43 into anisotropic intranuclear liquid spherical shells","volume":"371","author":"Yu","year":"2021","journal-title":"Science"},{"key":"2026012305022997800_JCS264115C89","doi-asserted-by":"publisher","first-page":"4991","DOI":"10.1242\/jcs.01373","article-title":"Hsp70 dynamics in vivo: effect of heat shock and protein aggregation","volume":"117","author":"Zeng","year":"2004","journal-title":"J. Cell Sci."},{"key":"2026012305022997800_JCS264115C90","doi-asserted-by":"publisher","first-page":"1523","DOI":"10.1038\/s41467-019-09234-6","article-title":"Metascape provides a biologist-oriented resource for the analysis of systems-level datasets","volume":"10","author":"Zhou","year":"2019","journal-title":"Nat. Commun."}],"container-title":["Journal of Cell Science"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/journals.biologists.com\/jcs\/article-pdf\/doi\/10.1242\/jcs.264115\/3677816\/jcs264115.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/jcs\/article-pdf\/doi\/10.1242\/jcs.264115\/3681807\/jcs264115.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/jcs\/article-pdf\/doi\/10.1242\/jcs.264115\/3681807\/jcs264115.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,1,23]],"date-time":"2026-01-23T10:02:38Z","timestamp":1769162558000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.biologists.com\/jcs\/article\/139\/2\/jcs264115\/370448\/Stress-specific-NONO-interactomes-reveal-a-key"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,1,15]]},"references-count":90,"journal-issue":{"issue":"2","published-print":{"date-parts":[[2026,1,15]]}},"URL":"https:\/\/doi.org\/10.1242\/jcs.264115","relation":{},"ISSN":["0021-9533","1477-9137"],"issn-type":[{"value":"0021-9533","type":"print"},{"value":"1477-9137","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,1,15]]},"article-number":"jcs264115"}}