{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T15:48:00Z","timestamp":1792079280528,"version":"4.4.0"},"reference-count":105,"publisher":"Elsevier BV","issue":"20","license":[{"start":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T00:00:00Z","timestamp":1790812800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T00:00:00Z","timestamp":1790812800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2026,9,12]],"date-time":"2026-09-12T00:00:00Z","timestamp":1789171200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100021800","name":"Hubrecht Institute","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100021800","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000781","name":"European Research Council","doi-asserted-by":"publisher","award":["TF-Dynamics- 949708"],"award-info":[{"award-number":["TF-Dynamics- 949708"]}],"id":[{"id":"10.13039\/501100000781","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["cell.com","elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Molecular Cell"],"published-print":{"date-parts":[[2026,10]]},"DOI":"10.1016\/j.molcel.2026.09.011","type":"journal-article","created":{"date-parts":[[2026,9,29]],"date-time":"2026-09-29T15:13:39Z","timestamp":1790694819000},"page":"4234-4249.e9","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":0,"title":["Transcription factor clusters enhance gene expression in vivo without modulating bursting dynamics"],"prefix":"10.1016","volume":"86","author":[{"given":"Emilia","family":"Cadar","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anthony","family":"Birnie","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Richard","family":"Marsman","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rianne","family":"Grond","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7335-7252","authenticated-orcid":false,"given":"Jacques P.","family":"Bothma","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.molcel.2026.09.011_bib1","doi-asserted-by":"crossref","DOI":"10.1126\/science.aar2555","article-title":"Imaging dynamic and selective low-complexity domain interactions that control gene transcription","volume":"361","author":"Chong","year":"2018","journal-title":"Science"},{"key":"10.1016\/j.molcel.2026.09.011_bib2","doi-asserted-by":"crossref","first-page":"331","DOI":"10.1016\/j.cell.2023.12.005","article-title":"Direct observation of a condensate effect on super-enhancer controlled gene bursting","volume":"187","author":"Du","year":"2024","journal-title":"Cell"},{"key":"10.1016\/j.molcel.2026.09.011_bib3","doi-asserted-by":"crossref","first-page":"664","DOI":"10.1126\/science.1239053","article-title":"Real-time dynamics of RNA polymerase II clustering in live human cells","volume":"341","author":"Cisse","year":"2013","journal-title":"Science"},{"key":"10.1016\/j.molcel.2026.09.011_bib4","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.40497","article-title":"Dynamic multifactor hubs interact transiently with sites of active transcription in Drosophila embryos","volume":"7","author":"Mir","year":"2018","journal-title":"eLife"},{"key":"10.1016\/j.molcel.2026.09.011_bib5","doi-asserted-by":"crossref","first-page":"412","DOI":"10.1126\/science.aar4199","article-title":"Mediator and RNA polymerase II clusters associate in transcription-dependent condensates","volume":"361","author":"Cho","year":"2018","journal-title":"Science"},{"key":"10.1016\/j.molcel.2026.09.011_bib6","doi-asserted-by":"crossref","first-page":"1842","DOI":"10.1016\/j.cell.2018.10.042","article-title":"Transcription Factors Activate Genes through the Phase-Separation Capacity of Their Activation Domains","volume":"175","author":"Boija","year":"2018","journal-title":"Cell"},{"key":"10.1016\/j.molcel.2026.09.011_bib7","doi-asserted-by":"crossref","DOI":"10.1126\/sciadv.adp3251","article-title":"Transcription factor clusters as information transfer agents","volume":"11","author":"Munshi","year":"2025","journal-title":"Sci. Adv."},{"key":"10.1016\/j.molcel.2026.09.011_bib8","doi-asserted-by":"crossref","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":"10.1016\/j.molcel.2026.09.011_bib9","doi-asserted-by":"crossref","first-page":"3970","DOI":"10.1016\/j.molcel.2022.09.021","article-title":"\u201cStructure\u201d-function relationships in eukaryotic transcription factors: The role of intrinsically disordered regions in gene regulation","volume":"82","author":"Ferrie","year":"2022","journal-title":"Mol. Cell"},{"key":"10.1016\/j.molcel.2026.09.011_bib10","doi-asserted-by":"crossref","DOI":"10.1038\/s42003-024-05892-5","article-title":"Transcriptional condensates: a blessing or a curse for gene regulation?","volume":"7","author":"Stortz","year":"2024","journal-title":"Commun. Biol."},{"key":"10.1016\/j.molcel.2026.09.011_bib11","doi-asserted-by":"crossref","first-page":"1579","DOI":"10.1016\/j.molcel.2021.03.046","article-title":"Is transcriptional regulation just going through a phase?","volume":"81","author":"Narlikar","year":"2021","journal-title":"Mol. Cell"},{"key":"10.1016\/j.molcel.2026.09.011_bib12","doi-asserted-by":"crossref","DOI":"10.1016\/j.ceb.2024.102421","article-title":"Spandrels of the cell nucleus","volume":"90","author":"Solovei","year":"2024","journal-title":"Curr. Opin. Cell Biol."},{"key":"10.1016\/j.molcel.2026.09.011_bib13","doi-asserted-by":"crossref","first-page":"2201","DOI":"10.1016\/j.molcel.2022.05.018","article-title":"A conceptual framework for understanding phase separation and addressing open questions and challenges","volume":"82","author":"Mittag","year":"2022","journal-title":"Mol. Cell"},{"key":"10.1016\/j.molcel.2026.09.011_bib14","doi-asserted-by":"crossref","first-page":"5449","DOI":"10.1093\/nar\/gkad227","article-title":"Transcription factor clusters enable target search but do not contribute to target gene activation","volume":"51","author":"Meeussen","year":"2023","journal-title":"Nucleic Acids Res."},{"key":"10.1016\/j.molcel.2026.09.011_bib15","doi-asserted-by":"crossref","first-page":"20072","DOI":"10.1073\/pnas.0806476105","article-title":"How the Dorsal gradient works: Insights from postgenome technologies","volume":"105","author":"Hong","year":"2008","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.molcel.2026.09.011_bib16","doi-asserted-by":"crossref","first-page":"6680","DOI":"10.1038\/sj.onc.1209954","article-title":"Introduction to NF-\u03baB: players, pathways, perspectives","volume":"25","author":"Gilmore","year":"2006","journal-title":"Oncogene"},{"key":"10.1016\/j.molcel.2026.09.011_bib17","doi-asserted-by":"crossref","first-page":"725","DOI":"10.1038\/nri910","article-title":"NF-\u03baB regulation in the immune system","volume":"2","author":"Li","year":"2002","journal-title":"Nat. Rev. Immunol."},{"key":"10.1016\/j.molcel.2026.09.011_bib18","doi-asserted-by":"crossref","first-page":"1387","DOI":"10.1016\/j.cub.2019.03.019","article-title":"The Drosophila Pioneer Factor Zelda Modulates the Nuclear Microenvironment of a Dorsal Target Enhancer to Potentiate Transcriptional Output","volume":"29","author":"Yamada","year":"2019","journal-title":"Curr. Biol."},{"key":"10.1016\/j.molcel.2026.09.011_bib19","doi-asserted-by":"crossref","first-page":"619","DOI":"10.1101\/gad.6.4.619","article-title":"The functional domains of the Drosophila morphogen dorsal: evidence from the analysis of mutants","volume":"6","author":"Isoda","year":"1992","journal-title":"Genes Dev."},{"key":"10.1016\/j.molcel.2026.09.011_bib20","doi-asserted-by":"crossref","first-page":"841","DOI":"10.1016\/S0969-2126(99)80107-5","article-title":"Structure of the specificity domain of the Dorsal homologue Gambif1 bound to DNA","volume":"7","author":"Cramer","year":"1999","journal-title":"Structure"},{"key":"10.1016\/j.molcel.2026.09.011_bib21","doi-asserted-by":"crossref","first-page":"1869","DOI":"10.1242\/dev.128.10.1869","article-title":"Activation and repression by the C-terminal domain of Dorsal","volume":"128","author":"Flores-Saaib","year":"2001","journal-title":"Development"},{"key":"10.1016\/j.molcel.2026.09.011_bib22","doi-asserted-by":"crossref","first-page":"323","DOI":"10.1002\/dvg.20468","article-title":"GFP-p65 knock-in mice as a tool to study NF-\u03baB dynamics in vivo","volume":"47","author":"De Lorenzi","year":"2009","journal-title":"Genesis"},{"key":"10.1016\/j.molcel.2026.09.011_bib23","doi-asserted-by":"crossref","first-page":"542","DOI":"10.1016\/j.gde.2012.08.005","article-title":"Temporal dynamics, spatial range, and transcriptional interpretation of the Dorsal morphogen gradient","volume":"22","author":"Rushlow","year":"2012","journal-title":"Curr. Opin. Genet. Dev."},{"key":"10.1016\/j.molcel.2026.09.011_bib24","doi-asserted-by":"crossref","first-page":"i","DOI":"10.1038\/nmeth.f.388","article-title":"The Airyscan detector from ZEISS: confocal imaging with improved signal-to-noise ratio and super-resolution","volume":"12","author":"Huff","year":"2015","journal-title":"Nat. Methods"},{"key":"10.1016\/j.molcel.2026.09.011_bib25","doi-asserted-by":"crossref","first-page":"213","DOI":"10.1038\/s41580-024-00789-x","article-title":"Transcription regulation by biomolecular condensates","volume":"26","author":"Pei","year":"2025","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"10.1016\/j.molcel.2026.09.011_bib26","doi-asserted-by":"crossref","first-page":"687","DOI":"10.1016\/S0092-8674(02)01087-5","article-title":"Whole-Genome Analysis of Dorsal-Ventral Patterning in the Drosophila Embryo","volume":"111","author":"Stathopoulos","year":"2002","journal-title":"Cell"},{"key":"10.1016\/j.molcel.2026.09.011_bib27","doi-asserted-by":"crossref","first-page":"2135","DOI":"10.1016\/j.cub.2013.08.053","article-title":"Live imaging of bicoid-dependent transcription in Drosophila embryos","volume":"23","author":"Lucas","year":"2013","journal-title":"Curr. Biol."},{"key":"10.1016\/j.molcel.2026.09.011_bib28","doi-asserted-by":"crossref","first-page":"2140","DOI":"10.1016\/j.cub.2013.08.054","article-title":"Quantitative imaging of transcription in living Drosophila embryos links polymerase activity to patterning","volume":"23","author":"Garcia","year":"2013","journal-title":"Curr. Biol."},{"key":"10.1016\/j.molcel.2026.09.011_bib29","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1038\/nrm757","article-title":"The snail superfamily of zinc-finger transcription factors","volume":"3","author":"Nieto","year":"2002","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"10.1016\/j.molcel.2026.09.011_bib30","doi-asserted-by":"crossref","first-page":"395","DOI":"10.1038\/330395a0","article-title":"The Drosophila developmental gene snail encodes a protein with nucleic acid binding fingers","volume":"330","author":"Boulay","year":"1987","journal-title":"Nature"},{"key":"10.1016\/j.molcel.2026.09.011_bib31","doi-asserted-by":"crossref","first-page":"983","DOI":"10.1242\/dev.111.4.983","article-title":"The snail gene required for mesoderm formation in Drosophila is expressed dynamically in derivatives of all three germ layers","volume":"111","author":"Alberga","year":"1991","journal-title":"Development"},{"key":"10.1016\/j.molcel.2026.09.011_bib32","doi-asserted-by":"crossref","first-page":"84","DOI":"10.1038\/35000034","article-title":"The transcription factor snail is a repressor of E-cadherin gene expression in epithelial tumour cells","volume":"2","author":"Batlle","year":"2000","journal-title":"Nat. Cell Biol."},{"key":"10.1016\/j.molcel.2026.09.011_bib33","doi-asserted-by":"crossref","first-page":"963","DOI":"10.2174\/15680096113136660102","article-title":"The Role of Snail in EMT and Tumorigenesis","volume":"13","author":"Wang","year":"2013","journal-title":"Curr. Cancer Drug Targets"},{"key":"10.1016\/j.molcel.2026.09.011_bib34","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.07956","article-title":"Enhancer additivity and non-additivity are determined by enhancer strength in the Drosophila embryo","volume":"4","author":"Bothma","year":"2015","journal-title":"eLife"},{"key":"10.1016\/j.molcel.2026.09.011_bib35","doi-asserted-by":"crossref","first-page":"335","DOI":"10.1038\/341335a0","article-title":"The products of the Drosophila gap genes hunchback and Kr\u00fcppel bind to the hunchback promoters","volume":"341","author":"Treisman","year":"1989","journal-title":"Nature"},{"key":"10.1016\/j.molcel.2026.09.011_bib36","doi-asserted-by":"crossref","first-page":"13570","DOI":"10.1073\/pnas.1109873108","article-title":"Multiple enhancers ensure precision of gap gene-expression patterns in the Drosophila embryo","volume":"108","author":"Perry","year":"2011","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.molcel.2026.09.011_bib37","doi-asserted-by":"crossref","first-page":"1568","DOI":"10.1101\/gad.5.9.1568","article-title":"twist and snail as positive and negative regulators during Drosophila mesoderm development","volume":"5","author":"Leptin","year":"1991","journal-title":"Genes Dev."},{"key":"10.1016\/j.molcel.2026.09.011_bib38","doi-asserted-by":"crossref","first-page":"3173","DOI":"10.1038\/onc.2010.92","article-title":"TWISTing an embryonic transcription factor into an oncoprotein","volume":"29","author":"Ansieau","year":"2010","journal-title":"Oncogene"},{"key":"10.1016\/j.molcel.2026.09.011_bib39","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pgen.1000857","article-title":"Molecular Evolution and Functional Characterization of Drosophila Insulin-Like Peptides","volume":"6","author":"Gr\u00f6nke","year":"2010","journal-title":"PLoS Genet."},{"key":"10.1016\/j.molcel.2026.09.011_bib40","doi-asserted-by":"crossref","first-page":"10327","DOI":"10.1073\/pnas.93.19.10327","article-title":"Identification of novel genes in Drosophila reveals the complex regulation of early gene activity in the mesoderm","volume":"93","author":"Casal","year":"1996","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.molcel.2026.09.011_bib41","doi-asserted-by":"crossref","first-page":"1518","DOI":"10.1101\/gad.6.8.1518","article-title":"dorsal-twist interactions establish snail expression in the presumptive mesoderm of the Drosophila embryo","volume":"6","author":"Ip","year":"1992","journal-title":"Genes Dev."},{"key":"10.1016\/j.molcel.2026.09.011_bib42","doi-asserted-by":"crossref","DOI":"10.1126\/science.1160631","article-title":"Shadow Enhancers as a Source of Evolutionary Novelty","volume":"321","author":"Hong","year":"2008","journal-title":"Science"},{"key":"10.1016\/j.molcel.2026.09.011_bib43","doi-asserted-by":"crossref","DOI":"10.1242\/dev.074377","article-title":"Complex interactions between cis-regulatory modules in native conformation are critical for Drosophila snail expression","volume":"138","author":"Dunipace","year":"2011","journal-title":"Development"},{"key":"10.1016\/j.molcel.2026.09.011_bib44","doi-asserted-by":"crossref","first-page":"1562","DOI":"10.1016\/j.cub.2010.07.043","article-title":"Shadow Enhancers Foster Robustness of Drosophila Gastrulation","volume":"20","author":"Perry","year":"2010","journal-title":"Curr. Biol."},{"key":"10.1016\/j.molcel.2026.09.011_bib45","doi-asserted-by":"crossref","first-page":"3683","DOI":"10.1242\/dev.124.19.3683","article-title":"Differential regulation of gastrulation and neuroectodermal gene expression by Snail in the Drosophila embryo","volume":"124","author":"Hemavathy","year":"1997","journal-title":"Development"},{"key":"10.1016\/j.molcel.2026.09.011_bib46","doi-asserted-by":"crossref","DOI":"10.1093\/genetics\/iyaf058","article-title":"The proximal enhancer of the snail gene mediates negative autoregulatory feedback in Drosophila melanogaster","volume":"230","author":"Dunipace","year":"2025","journal-title":"Genetics"},{"key":"10.1016\/j.molcel.2026.09.011_bib47","doi-asserted-by":"crossref","first-page":"1898","DOI":"10.1016\/j.devcel.2023.07.007","article-title":"Chromatin accessibility in the Drosophila embryo is determined by transcription factor pioneering and enhancer activation","volume":"58","author":"Brennan","year":"2023","journal-title":"Dev. Cell"},{"key":"10.1016\/j.molcel.2026.09.011_bib48","doi-asserted-by":"crossref","first-page":"395","DOI":"10.1038\/nbt.3121","article-title":"ChIP-nexus enables improved detection of in vivo transcription factor binding footprints","volume":"33","author":"He","year":"2015","journal-title":"Nat. Biotechnol."},{"key":"10.1016\/j.molcel.2026.09.011_bib49","doi-asserted-by":"crossref","first-page":"1837","DOI":"10.1002\/j.1460-2075.1992.tb05235.x","article-title":"The same dorsal binding site mediates both activation and repression in a context-dependent manner","volume":"11","author":"Pan","year":"1992","journal-title":"EMBO J."},{"key":"10.1016\/j.molcel.2026.09.011_bib50","doi-asserted-by":"crossref","first-page":"D174","DOI":"10.1093\/nar\/gkad1059","article-title":"JASPAR 2024: 20th anniversary of the open-access database of transcription factor binding profiles","volume":"52","author":"Rauluseviciute","year":"2024","journal-title":"Nucleic Acids Res."},{"key":"10.1016\/j.molcel.2026.09.011_bib51","doi-asserted-by":"crossref","first-page":"347","DOI":"10.1093\/genetics\/108.2.347","article-title":"Mutations and chromosomal rearrangements affecting the expression of snail, a gene involved in embryonic patterning in Drosophila melanogaster","volume":"108","author":"Grau","year":"1984","journal-title":"Genetics"},{"key":"10.1016\/j.molcel.2026.09.011_bib52","doi-asserted-by":"crossref","first-page":"615","DOI":"10.1093\/genetics\/105.3.615","article-title":"Maternal-Zygotic Gene Interactions during Formation of the Dorsoventral Pattern in Drosophila Embryos","volume":"105","author":"Simpson","year":"1983","journal-title":"Genetics"},{"key":"10.1016\/j.molcel.2026.09.011_bib53","doi-asserted-by":"crossref","first-page":"267","DOI":"10.1007\/BF00848156","article-title":"Mutations affecting the pattern of the larval cuticle inDrosophila melanogaster: I. Zygotic loci on the second chromosome","volume":"193","author":"N\u00fcsslein-Volhard","year":"1984","journal-title":"Wilhelm Rouxs Arch. Dev. Biol."},{"key":"10.1016\/j.molcel.2026.09.011_bib54","doi-asserted-by":"crossref","first-page":"80","DOI":"10.1006\/dbio.2000.9890","article-title":"The Alternative Migratory Pathways of the Drosophila Tracheal Cells Are Associated with Distinct Subsets of Mesodermal Cells","volume":"227","author":"Franch-Marro","year":"2000","journal-title":"Dev. Biol."},{"key":"10.1016\/j.molcel.2026.09.011_bib55","doi-asserted-by":"crossref","DOI":"10.1038\/ncomms10624","article-title":"Optogenetic control of nuclear protein export","volume":"7","author":"Niopek","year":"2016","journal-title":"Nat. Commun."},{"key":"10.1016\/j.molcel.2026.09.011_bib56","doi-asserted-by":"crossref","DOI":"10.1038\/s41467-024-53539-0","article-title":"Optogenetic dissection of transcriptional repression in a multicellular organism","volume":"15","author":"Zhao","year":"2024","journal-title":"Nat. Commun."},{"key":"10.1016\/j.molcel.2026.09.011_bib57","doi-asserted-by":"crossref","first-page":"2348","DOI":"10.1016\/j.devcel.2021.07.011","article-title":"Extremely rapid and reversible optogenetic perturbation of nuclear proteins in living embryos","volume":"56","author":"K\u00f6gler","year":"2021","journal-title":"Dev. Cell"},{"key":"10.1016\/j.molcel.2026.09.011_bib58","doi-asserted-by":"crossref","DOI":"10.1242\/dev.204706","article-title":"Optogenetic manipulation of nuclear Dorsal reveals temporal requirements and consequences for transcription","volume":"152","author":"Pimmett","year":"2025","journal-title":"Development"},{"key":"10.1016\/j.molcel.2026.09.011_bib59","doi-asserted-by":"crossref","DOI":"10.1016\/j.celrep.2022.110543","article-title":"Optogenetic control of the Bicoid morphogen reveals fast and slow modes of gap gene regulation","volume":"38","author":"Singh","year":"2022","journal-title":"Cell Rep."},{"key":"10.1016\/j.molcel.2026.09.011_bib60","doi-asserted-by":"crossref","first-page":"1810","DOI":"10.1016\/j.cell.2018.03.069","article-title":"LlamaTags: A Versatile Tool to Image Transcription Factor Dynamics in Live Embryos","volume":"173","author":"Bothma","year":"2018","journal-title":"Cell"},{"key":"10.1016\/j.molcel.2026.09.011_bib61","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1038\/nsmb.1727","article-title":"Modulation of protein properties in living cells using nanobodies","volume":"17","author":"Kirchhofer","year":"2010","journal-title":"Nat. Struct. Mol. Biol."},{"key":"10.1016\/j.molcel.2026.09.011_bib62","doi-asserted-by":"crossref","first-page":"1253","DOI":"10.1038\/nmeth.3170","article-title":"A robust pipeline for rapid production of versatile nanobody repertoires","volume":"11","author":"Fridy","year":"2014","journal-title":"Nat. Methods"},{"key":"10.1016\/j.molcel.2026.09.011_bib63","doi-asserted-by":"crossref","DOI":"10.3390\/cells8010048","article-title":"Nanobody Detection of Standard Fluorescent Proteins Enables Multi-Target DNA-PAINT with High Resolution and Minimal Displacement Errors","volume":"8","author":"Sograte-Idrissi","year":"2019","journal-title":"Cells"},{"key":"10.1016\/j.molcel.2026.09.011_bib64","article-title":"Four-color single-molecule imaging with engineered tags resolves the molecular architecture of signaling complexes in the plasma membrane","volume":"2","author":"Sotolongo Bell\u00f3n","year":"2022","journal-title":"Cell Rep. Methods"},{"key":"10.1016\/j.molcel.2026.09.011_bib65","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1016\/j.bbrc.2021.05.089","article-title":"Structural insights into two distinct nanobodies recognizing the same epitope of green fluorescent protein","volume":"565","author":"Zhong","year":"2021","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"10.1016\/j.molcel.2026.09.011_bib66","doi-asserted-by":"crossref","first-page":"411","DOI":"10.1016\/S0092-8674(00)81286-6","article-title":"marelle acts downstream of the Drosophila HOP\/JAK kinase and encodes a protein similar to the mammalian STATs","volume":"84","author":"Hou","year":"1996","journal-title":"Cell"},{"key":"10.1016\/j.molcel.2026.09.011_bib67","doi-asserted-by":"crossref","first-page":"2598","DOI":"10.1038\/sj.onc.1203482","article-title":"The roles of the Drosophila JAK\/STAT pathway","volume":"19","author":"Zeidler","year":"2000","journal-title":"Oncogene"},{"key":"10.1016\/j.molcel.2026.09.011_bib68","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1146\/annurev-biochem-011520-105250","article-title":"Transcription in Living Cells: Molecular Mechanisms of Bursting","volume":"89","author":"Rodriguez","year":"2020","journal-title":"Annu. Rev. Biochem."},{"key":"10.1016\/j.molcel.2026.09.011_bib69","doi-asserted-by":"crossref","first-page":"1695","DOI":"10.1042\/BST20231286","article-title":"Transcriptional bursting: from fundamentals to novel insights","volume":"52","author":"Hebenstreit","year":"2024","journal-title":"Biochem. Soc. Trans."},{"key":"10.1016\/j.molcel.2026.09.011_bib70","doi-asserted-by":"crossref","first-page":"1959","DOI":"10.1038\/s41594-025-01615-4","article-title":"A conserved coupling of transcriptional ON and OFF periods underlies bursting dynamics","volume":"32","author":"Chen","year":"2025","journal-title":"Nat. Struct. Mol. Biol."},{"key":"10.1016\/j.molcel.2026.09.011_bib71","doi-asserted-by":"crossref","first-page":"836","DOI":"10.1073\/pnas.1912500117","article-title":"Multimodal transcriptional control of pattern formation in embryonic development","volume":"117","author":"Lammers","year":"2020","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.molcel.2026.09.011_bib72","doi-asserted-by":"crossref","first-page":"835","DOI":"10.1016\/j.cell.2018.09.056","article-title":"Diverse Spatial Expression Patterns Emerge from Unified Kinetics of Transcriptional Bursting","volume":"175","author":"Zoller","year":"2018","journal-title":"Cell"},{"key":"10.1016\/j.molcel.2026.09.011_bib73","doi-asserted-by":"crossref","first-page":"1387","DOI":"10.1016\/j.cub.2017.03.069","article-title":"Rapid Rates of Pol II Elongation in the Drosophila Embryo","volume":"27","author":"Fukaya","year":"2017","journal-title":"Curr. Biol."},{"key":"10.1016\/j.molcel.2026.09.011_bib74","doi-asserted-by":"crossref","DOI":"10.1016\/j.celrep.2023.113225","article-title":"Regulation of the dynamic RNA Pol II elongation rate in Drosophila embryos","volume":"42","author":"Keller","year":"2023","journal-title":"Cell Rep."},{"key":"10.1016\/j.molcel.2026.09.011_bib75","doi-asserted-by":"crossref","first-page":"3449","DOI":"10.1093\/hmg\/ddi373","article-title":"Cancer development induced by graded expression of Snail in mice","volume":"14","author":"P\u00e9rez-Mancera","year":"2005","journal-title":"Hum. Mol. Genet."},{"key":"10.1016\/j.molcel.2026.09.011_bib76","doi-asserted-by":"crossref","first-page":"197","DOI":"10.1016\/j.ccr.2005.07.009","article-title":"The transcriptional repressor Snail promotes mammary tumor recurrence","volume":"8","author":"Moody","year":"2005","journal-title":"Cancer Cell"},{"key":"10.1016\/j.molcel.2026.09.011_bib77","doi-asserted-by":"crossref","first-page":"1567","DOI":"10.1038\/s41594-024-01322-6","article-title":"MYC phase separation selectively modulates the transcriptome","volume":"31","author":"Yang","year":"2024","journal-title":"Nat. Struct. Mol. Biol."},{"key":"10.1016\/j.molcel.2026.09.011_bib78","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1016\/j.cell.2017.02.007","article-title":"A Phase Separation Model for Transcriptional Control","volume":"169","author":"Hnisz","year":"2017","journal-title":"Cell"},{"key":"10.1016\/j.molcel.2026.09.011_bib79","doi-asserted-by":"crossref","first-page":"2718","DOI":"10.1016\/j.molcel.2025.06.008","article-title":"Reconciling competing models on the roles of condensates and soluble complexes in transcription factor function","volume":"85","author":"Bremer","year":"2025","journal-title":"Mol. Cell"},{"key":"10.1016\/j.molcel.2026.09.011_bib80","doi-asserted-by":"crossref","first-page":"1553","DOI":"10.1101\/gad.14.13.1553","article-title":"CBP\/p300 in cell growth, transformation, and development","volume":"14","author":"Goodman","year":"2000","journal-title":"Genes Dev."},{"key":"10.1016\/j.molcel.2026.09.011_bib81","doi-asserted-by":"crossref","first-page":"1145","DOI":"10.1016\/j.bcp.2004.03.045","article-title":"CBP and p300: HATs for different occasions","volume":"68","author":"Kalkhoven","year":"2004","journal-title":"Biochem. Pharmacol."},{"key":"10.1016\/j.molcel.2026.09.011_bib82","doi-asserted-by":"crossref","first-page":"5543","DOI":"10.1007\/s00018-021-03878-8","article-title":"CDK9 keeps RNA polymerase II on track","volume":"78","author":"Egloff","year":"2021","journal-title":"Cell. Mol. Life Sci."},{"key":"10.1016\/j.molcel.2026.09.011_bib83","doi-asserted-by":"crossref","first-page":"1375","DOI":"10.1128\/MCB.01365-08","article-title":"Brd4 Coactivates Transcriptional Activation of NF-\u03baB via Specific Binding to Acetylated RelA","volume":"29","author":"Huang","year":"2009","journal-title":"Mol. Cell. Biol."},{"key":"10.1016\/j.molcel.2026.09.011_bib84","doi-asserted-by":"crossref","first-page":"393","DOI":"10.1016\/j.molcel.2022.12.006","article-title":"P-TEFb: The master regulator of transcription elongation","volume":"83","author":"Fujinaga","year":"2023","journal-title":"Mol. Cell"},{"key":"10.1016\/j.molcel.2026.09.011_bib85","article-title":"Novel Fluorescent and Photoconvertible Fusions Reveal Dorsal Activator Dynamics","author":"Turner","year":"2025","journal-title":"bioRxiv"},{"key":"10.1016\/j.molcel.2026.09.011_bib86","doi-asserted-by":"crossref","first-page":"160","DOI":"10.1016\/j.tig.2023.11.003","article-title":"Time will tell: comparing timescales to gain insight into transcriptional bursting","volume":"40","author":"Meeussen","year":"2024","journal-title":"Trends Genet."},{"key":"10.1016\/j.molcel.2026.09.011_bib87","doi-asserted-by":"crossref","first-page":"707","DOI":"10.1016\/j.cell.2007.01.015","article-title":"The role of chromatin during transcription","volume":"128","author":"Li","year":"2007","journal-title":"Cell"},{"key":"10.1016\/j.molcel.2026.09.011_bib88","doi-asserted-by":"crossref","first-page":"92","DOI":"10.1126\/science.ade5308","article-title":"Chromatin expansion microscopy reveals nanoscale organization of transcription and chromatin","volume":"381","author":"Pownall","year":"2023","journal-title":"Science"},{"key":"10.1016\/j.molcel.2026.09.011_bib89","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1038\/s41576-022-00526-0","article-title":"The spatial organization of transcriptional control","volume":"24","author":"Hafner","year":"2023","journal-title":"Nat. Rev. Genet."},{"key":"10.1016\/j.molcel.2026.09.011_bib90","first-page":"89","article-title":"Fluorescent protein and peptide tags alter condensate formation and dynamics in vivo and in vitro","volume":"27","author":"D\u00f6rner","year":"2025","journal-title":"EMBO Rep."},{"key":"10.1016\/j.molcel.2026.09.011_bib91","article-title":"Transcription factor hubs exhibit gene-specific properties that tune expression","author":"Fallacaro","year":"2025","journal-title":"bioRxiv"},{"key":"10.1016\/j.molcel.2026.09.011_bib92","article-title":"Global maps of transcription factor properties reveal threshold-based formation of DNA-bound and mobile clusters","volume":"12","author":"Dima","year":"2025","journal-title":"Sci. Adv."},{"key":"10.1016\/j.molcel.2026.09.011_bib93","doi-asserted-by":"crossref","first-page":"544","DOI":"10.1016\/j.devcel.2011.12.007","article-title":"Dorsal-Ventral Gene Expression in the Drosophila Embryo Reflects the Dynamics and Precision of the Dorsal Nuclear Gradient","volume":"22","author":"Reeves","year":"2012","journal-title":"Dev. Cell"},{"key":"10.1016\/j.molcel.2026.09.011_bib94","doi-asserted-by":"crossref","DOI":"10.1242\/dev.200940","article-title":"Shadow enhancers modulate distinct transcriptional parameters that differentially effect downstream patterning events","volume":"149","author":"Whitney","year":"2022","journal-title":"Development"},{"key":"10.1016\/j.molcel.2026.09.011_bib95","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0017762","article-title":"Robust RT-qPCR Data Normalization: Validation and Selection of Internal Reference Genes during Post-Experimental Data Analysis","volume":"6","author":"Ling","year":"2011","journal-title":"PLOS One"},{"key":"10.1016\/j.molcel.2026.09.011_bib96","doi-asserted-by":"crossref","first-page":"2839","DOI":"10.1016\/j.cub.2023.04.005","article-title":"Precisely timed regulation of enhancer activity defines the binary expression pattern of Fushi tarazu in the Drosophila embryo","volume":"33","author":"Birnie","year":"2023","journal-title":"Curr. Biol."},{"key":"10.1016\/j.molcel.2026.09.011_bib97","doi-asserted-by":"crossref","first-page":"676","DOI":"10.1038\/nmeth.2019","article-title":"Fiji: An open-source platform for biological-image analysis","volume":"9","author":"Schindelin","year":"2012","journal-title":"Nat. Methods"},{"key":"10.1016\/j.molcel.2026.09.011_bib98","doi-asserted-by":"crossref","first-page":"1881","DOI":"10.1101\/gad.5.10.1881","article-title":"The dorsal morphogen gradient regulates the mesoderm determinant twist in early Drosophila embryos","volume":"5","author":"Jiang","year":"1991","journal-title":"Genes Dev."},{"key":"10.1016\/j.molcel.2026.09.011_bib99","doi-asserted-by":"crossref","first-page":"439","DOI":"10.1016\/0092-8674(91)90651-E","article-title":"The dorsal morphogen is a sequence-specific DNA-binding protein that interacts with a long-range repression element in drosophila","volume":"64","author":"Ip","year":"1991","journal-title":"Cell"},{"key":"10.1016\/j.molcel.2026.09.011_bib100","doi-asserted-by":"crossref","first-page":"741","DOI":"10.1016\/0092-8674(93)90402-C","article-title":"Binding affinities and cooperative interactions with bHLH activators delimit threshold responses to the dorsal gradient morphogen","volume":"72","author":"Jiang","year":"1993","journal-title":"Cell"},{"key":"10.1016\/j.molcel.2026.09.011_bib101","doi-asserted-by":"crossref","first-page":"1747","DOI":"10.1093\/bioinformatics\/bti173","article-title":"Drosophila DNase I footprint database: a systematic genome annotation of transcription factor binding sites in the fruitfly, Drosophila melanogaster","volume":"21","author":"Bergman","year":"2005","journal-title":"Bioinformatics"},{"key":"10.1016\/j.molcel.2026.09.011_bib102","doi-asserted-by":"crossref","first-page":"2018","DOI":"10.1101\/gr.132811.111","article-title":"Uncovering cis -regulatory sequence requirements for context-specific transcription factor binding","volume":"22","author":"Y\u00e1\u00f1ez-Cuna","year":"2012","journal-title":"Genome Res."},{"key":"10.1016\/j.molcel.2026.09.011_bib103","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1146\/annurev-cellbio-100617-062719","article-title":"Low-Affinity Binding Sites and the Transcription Factor Specificity Paradox in Eukaryotes","volume":"35","author":"Kribelbauer","year":"2019","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"10.1016\/j.molcel.2026.09.011_bib104","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.85997","article-title":"Modulation of protein-DNA binding reveals mechanisms of spatiotemporal gene control in early Drosophila embryos","volume":"12","author":"Syed","year":"2023","journal-title":"eLife"},{"key":"10.1016\/j.molcel.2026.09.011_bib105","series-title":"Computer Vision: Models, Learning, and Inference","author":"Prince","year":"2012"}],"container-title":["Molecular Cell"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S1097276526006544?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S1097276526006544?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T15:23:14Z","timestamp":1792077794000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S1097276526006544"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10]]},"references-count":105,"journal-issue":{"issue":"20","published-print":{"date-parts":[[2026,10]]}},"alternative-id":["S1097276526006544"],"URL":"https:\/\/doi.org\/10.1016\/j.molcel.2026.09.011","relation":{},"ISSN":["1097-2765"],"issn-type":[{"value":"1097-2765","type":"print"}],"subject":[],"published":{"date-parts":[[2026,10]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Transcription factor clusters enhance gene expression in vivo without modulating bursting dynamics","name":"articletitle","label":"Article Title"},{"value":"Molecular Cell","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.molcel.2026.09.011","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2026 The Author(s). Published by Elsevier Inc.","name":"copyright","label":"Copyright"}]}}