{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T10:46:30Z","timestamp":1782902790808,"version":"3.54.5"},"reference-count":100,"publisher":"The Company of Biologists","issue":"13","license":[{"start":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T00:00:00Z","timestamp":1782864000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"name":"Lohengrin Foundation"},{"name":"Eunice Kennedy Shriver National Institute of Child Health and Human Development","award":["T32HD055164"],"award-info":[{"award-number":["T32HD055164"]}]},{"DOI":"10.13039\/100000065","name":"National Institute of Neurological Disorders and Stroke","doi-asserted-by":"crossref","award":["R01NS116365"],"award-info":[{"award-number":["R01NS116365"]}],"id":[{"id":"10.13039\/100000065","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100007234","name":"University of Chicago","doi-asserted-by":"crossref","id":[{"id":"10.13039\/100007234","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,7,1]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>Motor neuron (MN) diversity is essential for generating animal movement, yet the molecular mechanisms specifying MN subtypes remain poorly understood. We investigate how Hox genes and their PBX co-factors establish cholinergic MN identity along the anterior-posterior axis of the Caenorhabditis elegans ventral nerve cord. In anterior MNs, the Hox genes ceh-13 (Lab\/Hox1) and lin-39 (Scr\/Dfd\/Hox4-5) collaborate with the co-factor ceh-20 (Exd\/Pbx1-4) and terminal selector unc-3 (Collier\/Ebf1-4) to activate terminal identity genes. In posterior MNs, the Hox gene mab-5 (Antp\/Hox6-8) represses terminal identity genes by antagonizing unc-3 in a ceh-20-dependent manner. Both mab-5 and ceh-20 are required not only during early development but also in later life stages to maintain posterior MN identity. In lumbar MNs, egl-5 (Abd-A\/Abd-B\/Hox9-13) collaborates with unc-3 to activate lumbar-specific identity genes independently of ceh-20. We also find that ceh-20 is required for Hox gene expression in ventral nerve cord MNs, supporting a model in which Hox-positive autoregulation depends on PBX activity. Together, these findings provide a conceptual framework for understanding how spatial patterning information is integrated with terminal selectors to generate and maintain neuronal subtype diversity.<\/jats:p>","DOI":"10.1242\/dev.205511","type":"journal-article","created":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T10:08:27Z","timestamp":1782900507000},"source":"Crossref","is-referenced-by-count":0,"title":["PBX-dependent and -independent Hox programs establish and maintain motor neuron terminal identity"],"prefix":"10.1242","volume":"153","author":[{"given":"Manasa","family":"Prahlad","sequence":"first","affiliation":[{"name":"University of Chicago 1 Department of Neurobiology , , Chicago, IL 60637 ,","place":["USA"]},{"name":"Center for Motor Neuron Disease, University of Chicago 2 , Chicago, IL 60637 ,","place":["USA"]},{"name":"Neuroscience Institute, University of Chicago 3 , Chicago, IL 60637 ,","place":["USA"]},{"name":"Committee on Genetics, Genomics, and Systems Biology 4 , Chicago, IL 60637 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Weidong","family":"Feng","sequence":"additional","affiliation":[{"name":"University of Chicago 1 Department of Neurobiology , , Chicago, IL 60637 ,","place":["USA"]},{"name":"Center for Motor Neuron Disease, University of Chicago 2 , Chicago, IL 60637 ,","place":["USA"]},{"name":"Neuroscience Institute, University of Chicago 3 , Chicago, IL 60637 ,","place":["USA"]},{"name":"Committee on Development, Regeneration, and Stem Cell Biology 5 , Chicago, IL 60637,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ian Q.","family":"Weigle","sequence":"additional","affiliation":[{"name":"University of Chicago 1 Department of Neurobiology , , Chicago, IL 60637 ,","place":["USA"]},{"name":"Center for Motor Neuron Disease, University of Chicago 2 , Chicago, IL 60637 ,","place":["USA"]},{"name":"Neuroscience Institute, University of Chicago 3 , Chicago, IL 60637 ,","place":["USA"]},{"name":"Committee on Neurobiology 6 , Chicago, IL 60637 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Oyunsuvd","family":"Bat-Erdene","sequence":"additional","affiliation":[{"name":"University of Chicago 1 Department of Neurobiology , , Chicago, IL 60637 ,","place":["USA"]},{"name":"Center for Motor Neuron Disease, University of Chicago 2 , Chicago, IL 60637 ,","place":["USA"]},{"name":"Neuroscience Institute, University of Chicago 3 , Chicago, IL 60637 ,","place":["USA"]},{"name":"Committee on Neurobiology 6 , Chicago, IL 60637 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0000-6095-2052","authenticated-orcid":false,"given":"Yihan","family":"Chen","sequence":"additional","affiliation":[{"name":"University of Chicago 1 Department of Neurobiology , , Chicago, IL 60637 ,","place":["USA"]},{"name":"Center for Motor Neuron Disease, University of Chicago 2 , Chicago, IL 60637 ,","place":["USA"]},{"name":"Neuroscience Institute, University of Chicago 3 , Chicago, IL 60637 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Filipe","family":"Marques","sequence":"additional","affiliation":[{"name":"University of Chicago 1 Department of Neurobiology , , Chicago, IL 60637 ,","place":["USA"]},{"name":"Center for Motor Neuron Disease, University of Chicago 2 , Chicago, IL 60637 ,","place":["USA"]},{"name":"Neuroscience Institute, University of Chicago 3 , Chicago, IL 60637 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1363-9271","authenticated-orcid":false,"given":"Paschalis","family":"Kratsios","sequence":"additional","affiliation":[{"name":"University of Chicago 1 Department of Neurobiology , , Chicago, IL 60637 ,","place":["USA"]},{"name":"Center for Motor Neuron Disease, University of Chicago 2 , Chicago, IL 60637 ,","place":["USA"]},{"name":"Neuroscience Institute, University of Chicago 3 , Chicago, IL 60637 ,","place":["USA"]},{"name":"Committee on Genetics, Genomics, and Systems Biology 4 , Chicago, IL 60637 ,","place":["USA"]},{"name":"Committee on Development, Regeneration, and Stem Cell Biology 5 , Chicago, IL 60637,","place":["USA"]},{"name":"Committee on Neurobiology 6 , Chicago, IL 60637 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","published-online":{"date-parts":[[2026,7,1]]},"reference":[{"key":"2026070106063340300_DEV205511C1","doi-asserted-by":"publisher","first-page":"3071","DOI":"10.1534\/g3.120.401515","article-title":"cnd-1\/NeuroD1 functions with the Homeobox gene ceh-5\/Vax2 and Hox gene ceh-13\/labial to specify aspects of RME and DD neuron fate in Caenorhabditis elegans","volume":"10","author":"Aquino-Nunez","year":"2020","journal-title":"G3 (Bethesda)"},{"key":"2026070106063340300_DEV205511C2","doi-asserted-by":"publisher","first-page":"751","DOI":"10.1016\/j.tins.2015.10.005","article-title":"Homeotic transformations of neuronal cell identities","volume":"38","author":"Arlotta","year":"2015","journal-title":"Trends Neurosci."},{"key":"2026070106063340300_DEV205511C3","doi-asserted-by":"publisher","first-page":"2027","DOI":"10.1242\/dev.090902","article-title":"Dual role for Hox genes and Hox co-factors in conferring leg motoneuron survival and identity in Drosophila","volume":"140","author":"Baek","year":"2013","journal-title":"Development"},{"key":"2026070106063340300_DEV205511C4","doi-asserted-by":"publisher","first-page":"209","DOI":"10.1016\/S0896-6273(02)01181-9","article-title":"A pulse of the Drosophila Hox protein Abdominal-A schedules the end of neural proliferation via neuroblast apoptosis","volume":"37","author":"Bello","year":"2003","journal-title":"Neuron"},{"key":"2026070106063340300_DEV205511C5","doi-asserted-by":"publisher","first-page":"71","DOI":"10.1093\/genetics\/77.1.71","article-title":"The genetics of Caenorhabditis elegans","volume":"77","author":"Brenner","year":"1974","journal-title":"Genetics"},{"key":"2026070106063340300_DEV205511C6","doi-asserted-by":"publisher","first-page":"1537","DOI":"10.1242\/dev.126.7.1537","article-title":"Anterior organization of the Caenorhabditis elegans embryo by the labial-like Hox gene ceh-13","volume":"126","author":"Brunschwig","year":"1999","journal-title":"Development"},{"key":"2026070106063340300_DEV205511C7","doi-asserted-by":"publisher","first-page":"e70766","DOI":"10.7554\/eLife.70766","article-title":"Control of spinal motor neuron terminal differentiation through sustained Hoxc8 gene activity","volume":"11","author":"Catela","year":"2022","journal-title":"eLife"},{"key":"2026070106063340300_DEV205511C8","doi-asserted-by":"publisher","first-page":"663","DOI":"10.1101\/gad.9.6.663","article-title":"Pbx proteins display hexapeptide-dependent cooperative DNA binding with a subset of Hox proteins","volume":"9","author":"Chang","year":"1995","journal-title":"Genes Dev."},{"key":"2026070106063340300_DEV205511C9","doi-asserted-by":"publisher","first-page":"1734","DOI":"10.1128\/MCB.16.4.1734","article-title":"Pbx modulation of Hox homeodomain amino-terminal arms establishes different DNA-binding specificities across the Hox locus","volume":"16","author":"Chang","year":"1996","journal-title":"Mol. Cell. Biol."},{"key":"2026070106063340300_DEV205511C10","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1016\/0092-8674(93)90293-Y","article-title":"Control of cell fates in the central body region of C. elegans by the homeobox gene lin-39","volume":"74","author":"Clark","year":"1993","journal-title":"Cell"},{"key":"2026070106063340300_DEV205511C11","doi-asserted-by":"publisher","first-page":"200","DOI":"10.1016\/S0012-1606(02)00018-0","article-title":"Autonomous and nonautonomous functions for Hox\/Pbx in branchiomotor neuron development","volume":"253","author":"Cooper","year":"2003","journal-title":"Dev. Biol."},{"key":"2026070106063340300_DEV205511C12","doi-asserted-by":"publisher","first-page":"e1011355","DOI":"10.1371\/journal.pgen.1011355","article-title":"Ambivalent partnership of the Drosophila posterior class Hox protein Abdominal-B with Extradenticle and Homothorax","volume":"21","author":"Curt","year":"2025","journal-title":"PLoS Genet."},{"key":"2026070106063340300_DEV205511C13","doi-asserted-by":"publisher","first-page":"5664","DOI":"10.1038\/s41598-019-42096-y","article-title":"The human HOXA9 protein uses paralog-specific residues of the homeodomain to interact with TALE-class cofactors","volume":"9","author":"Dard","year":"2019","journal-title":"Sci. Rep."},{"key":"2026070106063340300_DEV205511C14","doi-asserted-by":"publisher","first-page":"ar130","DOI":"10.1091\/mbc.E24-08-0347","article-title":"The Hox Gene, abdominal-A, controls the size and timely mitotic entry of neural stem cells during CNS patterning in Drosophila","volume":"36","author":"Das","year":"2025","journal-title":"Mol. Biol. Cell"},{"key":"2026070106063340300_DEV205511C15","doi-asserted-by":"publisher","first-page":"304","DOI":"10.1016\/j.cell.2008.06.019","article-title":"Hox repertoires for motor neuron diversity and connectivity gated by a single accessory factor, FoxP1","volume":"134","author":"Dasen","year":"2008","journal-title":"Cell"},{"key":"2026070106063340300_DEV205511C16","doi-asserted-by":"publisher","first-page":"113049","DOI":"10.1016\/j.celrep.2023.113049","article-title":"Determinants of motor neuron functional subtypes important for locomotor speed","volume":"42","author":"D'Elia","year":"2023","journal-title":"Cell Rep."},{"key":"2026070106063340300_DEV205511C18","doi-asserted-by":"publisher","first-page":"1391","DOI":"10.1101\/gad.217224.113","article-title":"A combinatorial regulatory signature controls terminal differentiation of the dopaminergic nervous system in C. elegans","volume":"27","author":"Doitsidou","year":"2013","journal-title":"Genes Dev."},{"key":"2026070106063340300_DEV205511C19","doi-asserted-by":"publisher","first-page":"7666","DOI":"10.1073\/pnas.90.16.7666","article-title":"Local alterations of Krox-20 and Hox gene expression in the hindbrain suggest lack of rhombomeres 4 and 5 in homozygote null Hoxa-1 (Hox-1.6) mutant embryos","volume":"90","author":"Doll\u00e9","year":"1993","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2026070106063340300_DEV205511C20","doi-asserted-by":"publisher","first-page":"6225","DOI":"10.1523\/JNEUROSCI.19-15-06225.1999","article-title":"Coordinated transcriptional regulation of the unc-25 glutamic acid decarboxylase and the unc-47 GABA vesicular transporter by the Caenorhabditis elegans UNC-30 homeodomain protein","volume":"19","author":"Eastman","year":"1999","journal-title":"J. Neurosci."},{"key":"2026070106063340300_DEV205511C21","doi-asserted-by":"publisher","first-page":"170","DOI":"10.1016\/j.gene.2007.03.002","article-title":"Efficient and cell specific knock-down of gene function in targeted C. elegans neurons","volume":"395","author":"Esposito","year":"2007","journal-title":"Gene"},{"key":"2026070106063340300_DEV205511C22","doi-asserted-by":"publisher","first-page":"e50065","DOI":"10.7554\/eLife.50065","article-title":"A terminal selector prevents a Hox transcriptional switch to safeguard motor neuron identity throughout life","volume":"9","author":"Feng","year":"2020","journal-title":"eLife"},{"key":"2026070106063340300_DEV205511C23","doi-asserted-by":"publisher","first-page":"6097","DOI":"10.1038\/s41467-022-33781-0","article-title":"Maintenance of neurotransmitter identity by Hox proteins through a homeostatic mechanism","volume":"13","author":"Feng","year":"2022","journal-title":"Nat. Commun."},{"key":"2026070106063340300_DEV205511C24","doi-asserted-by":"publisher","first-page":"850","DOI":"10.1016\/j.celrep.2015.12.077","article-title":"Hox function is required for the development and maintenance of the Drosophila feeding motor unit","volume":"14","author":"Friedrich","year":"2016","journal-title":"Cell Rep."},{"key":"2026070106063340300_DEV205511C25","doi-asserted-by":"publisher","first-page":"3115","DOI":"10.1242\/dev.129.13.3115","article-title":"Hox repression of a target gene: extradenticle-independent, additive action through multiple monomer binding sites","volume":"129","author":"Galant","year":"2002","journal-title":"Development"},{"key":"2026070106063340300_DEV205511C26","doi-asserted-by":"publisher","first-page":"5191","DOI":"10.1242\/dev.00730","article-title":"Hox3 genes coordinate mechanisms of genetic suppression and activation in the generation of branchial and somatic motoneurons","volume":"130","author":"Gaufo","year":"2003","journal-title":"Development"},{"key":"2026070106063340300_DEV205511C27","doi-asserted-by":"publisher","first-page":"1775","DOI":"10.1126\/science.1196914","article-title":"Integrative analysis of the Caenorhabditis elegans genome by the modENCODE project","volume":"330","author":"Gerstein","year":"2010","journal-title":"Science"},{"key":"2026070106063340300_DEV205511C28","doi-asserted-by":"publisher","first-page":"eadg0506","DOI":"10.1126\/sciadv.adg0506","article-title":"Whole-body gene expression atlas of an adult metazoan","volume":"9","author":"Ghaddar","year":"2023","journal-title":"Sci. Adv."},{"key":"2026070106063340300_DEV205511C29","doi-asserted-by":"publisher","first-page":"859","DOI":"10.1038\/nrn2254","article-title":"Patterning and axon guidance of cranial motor neurons","volume":"8","author":"Guthrie","year":"2007","journal-title":"Nat. Rev. Neurosci."},{"key":"2026070106063340300_DEV205511C30","doi-asserted-by":"publisher","first-page":"1005","DOI":"10.1016\/j.neuron.2016.07.043","article-title":"Parallel Pbx-dependent pathways govern the coalescence and fate of motor columns","volume":"91","author":"Hanley","year":"2016","journal-title":"Neuron"},{"key":"2026070106063340300_DEV205511C31","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1242\/dev.143875","article-title":"The Drosophila Hox gene Ultrabithorax acts in both muscles and motoneurons to orchestrate formation of specific neuromuscular connections","volume":"144","author":"Hessinger","year":"2017","journal-title":"Development"},{"key":"2026070106063340300_DEV205511C32","doi-asserted-by":"publisher","first-page":"iyac085","DOI":"10.1093\/genetics\/iyac085","article-title":"Defining characteristics and conservation of poorly annotated genes in Caenorhabditis elegans using WormCat 2.0","volume":"221","author":"Higgins","year":"2022","journal-title":"Genetics"},{"key":"2026070106063340300_DEV205511C33","doi-asserted-by":"publisher","first-page":"455","DOI":"10.1016\/bs.ctdb.2015.12.007","article-title":"Terminal selectors of neuronal identity","volume":"116","author":"Hobert","year":"2016","journal-title":"Curr. Top. Dev. Biol."},{"key":"2026070106063340300_DEV205511C34","doi-asserted-by":"publisher","first-page":"97","DOI":"10.1016\/j.conb.2018.12.006","article-title":"Neuronal identity control by terminal selectors in worms, flies, and chordates","volume":"56","author":"Hobert","year":"2019","journal-title":"Curr. Opin. Neurobiol."},{"key":"2026070106063340300_DEV205511C35","doi-asserted-by":"publisher","first-page":"e1001351","DOI":"10.1371\/journal.pbio.1001351","article-title":"Hox proteins display a common and ancestral ability to diversify their interaction mode with the PBC class cofactors","volume":"10","author":"Hudry","year":"2012","journal-title":"PLoS Biol."},{"key":"2026070106063340300_DEV205511C36","doi-asserted-by":"publisher","first-page":"387","DOI":"10.1242\/dev.117.1.387","article-title":"Negative autoregulation by Ultrabithorax controls the level and pattern of its expression","volume":"117","author":"Irvine","year":"1993","journal-title":"Development"},{"key":"2026070106063340300_DEV205511C37","doi-asserted-by":"publisher","first-page":"780","DOI":"10.1038\/372780a0","article-title":"Control of type-D GABAergic neuron differentiation by C. elegans UNC-30 homeodomain protein","volume":"372","author":"Jin","year":"1994","journal-title":"Nature"},{"key":"2026070106063340300_DEV205511C38","doi-asserted-by":"publisher","first-page":"530","DOI":"10.1016\/j.cell.2007.09.024","article-title":"Functional specificity of a Hox protein mediated by the recognition of minor groove structure","volume":"131","author":"Joshi","year":"2007","journal-title":"Cell"},{"key":"2026070106063340300_DEV205511C39","doi-asserted-by":"publisher","first-page":"781","DOI":"10.1016\/j.neuron.2010.08.008","article-title":"Global control of motor neuron topography mediated by the repressive actions of a single hox gene","volume":"67","author":"Jung","year":"2010","journal-title":"Neuron"},{"key":"2026070106063340300_DEV205511C40","doi-asserted-by":"publisher","first-page":"2751","DOI":"10.1242\/dev.126.12.2751","article-title":"Specification of distinct motor neuron identities by the singular activities of individual Hox genes","volume":"126","author":"Jungbluth","year":"1999","journal-title":"Development"},{"key":"2026070106063340300_DEV205511C41","doi-asserted-by":"publisher","first-page":"477","DOI":"10.1016\/0092-8674(86)90668-9","article-title":"A gene involved in the development of the posterior body region of C. elegans","volume":"46","author":"Kenyon","year":"1986","journal-title":"Cell"},{"key":"2026070106063340300_DEV205511C42","doi-asserted-by":"publisher","first-page":"205","DOI":"10.1038\/nn.2989","article-title":"Coordinated regulation of cholinergic motor neuron traits through a conserved terminal selector gene","volume":"15","author":"Kratsios","year":"2011","journal-title":"Nat. Neurosci."},{"key":"2026070106063340300_DEV205511C43","doi-asserted-by":"publisher","first-page":"1282","DOI":"10.1016\/j.cub.2015.03.028","article-title":"Transcriptional coordination of synaptogenesis and neurotransmitter signaling","volume":"25","author":"Kratsios","year":"2015","journal-title":"Curr. Biol."},{"key":"2026070106063340300_DEV205511C44","doi-asserted-by":"publisher","first-page":"e25751","DOI":"10.7554\/eLife.25751","article-title":"An intersectional gene regulatory strategy defines subclass diversity of C. elegans motor neurons","volume":"6","author":"Kratsios","year":"2017","journal-title":"eLife"},{"key":"2026070106063340300_DEV205511C45","doi-asserted-by":"publisher","first-page":"191","DOI":"10.1016\/0092-8674(94)90290-9","article-title":"Hox genes in vertebrate development","volume":"78","author":"Krumlauf","year":"1994","journal-title":"Cell"},{"key":"2026070106063340300_DEV205511C46","doi-asserted-by":"publisher","first-page":"1006","DOI":"10.1242\/bio.20121966","article-title":"The labial gene is required to terminate proliferation of identified neuroblasts in postembryonic development of the Drosophila brain","volume":"1","author":"Kuert","year":"2012","journal-title":"Biol. Open"},{"key":"2026070106063340300_DEV205511C47","doi-asserted-by":"publisher","first-page":"102","DOI":"10.1016\/j.ydbio.2014.03.021","article-title":"Neuroblast lineage identification and lineage-specific Hox gene action during postembryonic development of the subesophageal ganglion in the Drosophila central brain","volume":"390","author":"Kuert","year":"2014","journal-title":"Dev. Biol."},{"key":"2026070106063340300_DEV205511C48","doi-asserted-by":"publisher","first-page":"477","DOI":"10.1016\/0092-8674(88)90034-7","article-title":"Autoregulation of a Drosophila homeotic selector gene","volume":"55","author":"Kuziora","year":"1988","journal-title":"Cell"},{"key":"2026070106063340300_DEV205511C49","doi-asserted-by":"publisher","first-page":"175","DOI":"10.1016\/S0736-5748(00)00090-3","article-title":"The acquisition of motoneuron subtype identity and motor circuit formation","volume":"19","author":"Landmesser","year":"2001","journal-title":"Int. J. Dev. Neurosci."},{"key":"2026070106063340300_DEV205511C101","doi-asserted-by":"publisher","first-page":"2060","DOI":"10.1101\/gad.1103303","article-title":"Structure of HoxA9 and Pbx1 bound to DNA: Hox hexapeptide and DNA recognition anterior to posterior","volume":"17","author":"LaRonde-LeBlanc","year":"2003","journal-title":"Genes Dev."},{"key":"2026070106063340300_DEV205511C50","doi-asserted-by":"publisher","first-page":"10.17912\/micropub.biology.000453","DOI":"10.17912\/micropub.biology.000453","article-title":"Transgenic reporter analysis of ChIP-Seq-defined enhancers identifies novel target genes for the terminal selector UNC-3\/Collier\/Ebf","volume":"2021","author":"Li","year":"2021","journal-title":"MicroPubl. Biol."},{"key":"2026070106063340300_DEV205511C51","doi-asserted-by":"publisher","first-page":"e59464","DOI":"10.7554\/eLife.59464","article-title":"Establishment and maintenance of motor neuron identity via temporal modularity in terminal selector function","volume":"9","author":"Li","year":"2020","journal-title":"eLife"},{"key":"2026070106063340300_DEV205511C52","doi-asserted-by":"publisher","first-page":"5179","DOI":"10.1242\/dev.127.23.5179","article-title":"Overlapping roles of two Hox genes and the exd ortholog ceh-20 in diversification of the C. elegans postembryonic mesoderm","volume":"127","author":"Liu","year":"2000","journal-title":"Development"},{"key":"2026070106063340300_DEV205511C53","doi-asserted-by":"publisher","first-page":"997","DOI":"10.1016\/S0896-6273(01)00544-X","article-title":"Assigning the positional identity of spinal motor neurons: rostrocaudal patterning of Hox-c expression by FGFs, Gdf11, and retinoids","volume":"32","author":"Liu","year":"2001","journal-title":"Neuron"},{"key":"2026070106063340300_DEV205511C54","doi-asserted-by":"publisher","first-page":"893","DOI":"10.1038\/s41592-021-01216-1","article-title":"A 4D single-cell protein atlas of transcription factors delineates spatiotemporal patterning during embryogenesis","volume":"18","author":"Ma","year":"2021","journal-title":"Nat. Methods"},{"key":"2026070106063340300_DEV205511C55","doi-asserted-by":"publisher","first-page":"423","DOI":"10.1016\/S0959-437X(98)80113-5","article-title":"Hox proteins meet more partners","volume":"8","author":"Mann","year":"1998","journal-title":"Curr. Opin. Genet. Dev."},{"key":"2026070106063340300_DEV205511C56","doi-asserted-by":"publisher","first-page":"63","DOI":"10.1016\/S0070-2153(09)88003-4","article-title":"Hox specificity unique roles for cofactors and collaborators","volume":"88","author":"Mann","year":"2009","journal-title":"Curr. Top. Dev. Biol."},{"key":"2026070106063340300_DEV205511C57","doi-asserted-by":"publisher","first-page":"334","DOI":"10.1016\/j.tig.2016.03.004","article-title":"To be specific or not: the critical relationship between Hox and TALE proteins","volume":"32","author":"Merabet","year":"2016","journal-title":"Trends Genet."},{"key":"2026070106063340300_DEV205511C58","doi-asserted-by":"publisher","first-page":"193","DOI":"10.1016\/j.ydbio.2005.10.032","article-title":"Hox cofactors in vertebrate development","volume":"291","author":"Moens","year":"2006","journal-title":"Dev. Biol."},{"key":"2026070106063340300_DEV205511C59","doi-asserted-by":"publisher","first-page":"245","DOI":"10.1101\/gr.114587.110","article-title":"Diverse transcription factor binding features revealed by genome-wide ChIP-seq in C. elegans","volume":"21","author":"Niu","year":"2011","journal-title":"Genome Res."},{"key":"2026070106063340300_DEV205511C60","doi-asserted-by":"publisher","first-page":"1991","DOI":"10.1242\/dev.125.11.1991","article-title":"Expression of the murine Hoxa4 gene requires both autoregulation and a conserved retinoic acid response element","volume":"125","author":"Packer","year":"1998","journal-title":"Development"},{"key":"2026070106063340300_DEV205511C61","doi-asserted-by":"publisher","first-page":"eaax1971","DOI":"10.1126\/science.aax1971","article-title":"A lineage-resolved molecular atlas of C. elegans embryogenesis at single-cell resolution","volume":"365","author":"Packer","year":"2019","journal-title":"Science"},{"key":"2026070106063340300_DEV205511C62","doi-asserted-by":"publisher","first-page":"dev168179","DOI":"10.1242\/dev.168179","article-title":"Control of Hox transcription factor concentration and cell-to-cell variability by an auto-regulatory switch","volume":"146","author":"Papadopoulos","year":"2019","journal-title":"Development"},{"key":"2026070106063340300_DEV205511C63","doi-asserted-by":"publisher","first-page":"714","DOI":"10.1038\/17833","article-title":"Structure of a DNA-bound Ultrabithorax-Extradenticle homeodomain complex","volume":"397","author":"Passner","year":"1999","journal-title":"Nature"},{"key":"2026070106063340300_DEV205511C64","doi-asserted-by":"publisher","first-page":"677","DOI":"10.1093\/genetics\/156.2.677","article-title":"Regulation by homeoproteins: a comparison of deformed-responsive elements","volume":"156","author":"Pederson","year":"2000","journal-title":"Genetics"},{"key":"2026070106063340300_DEV205511C65","doi-asserted-by":"publisher","first-page":"1209","DOI":"10.1101\/gad.4.7.1209","article-title":"Mutations in the Drosophila gene extradenticle affect the way specific homeo domain proteins regulate segmental identity","volume":"4","author":"Peifer","year":"1990","journal-title":"Genes Dev."},{"key":"2026070106063340300_DEV205511C66","doi-asserted-by":"publisher","first-page":"12","DOI":"10.1016\/j.neuron.2013.09.020","article-title":"Hox genes: choreographers in neural development, architects of circuit organization","volume":"80","author":"Philippidou","year":"2013","journal-title":"Neuron"},{"key":"2026070106063340300_DEV205511C67","doi-asserted-by":"publisher","first-page":"1636","DOI":"10.1038\/nn.3242","article-title":"Sustained Hox5 gene activity is required for respiratory motor neuron development","volume":"15","author":"Philippidou","year":"2012","journal-title":"Nat. Neurosci."},{"key":"2026070106063340300_DEV205511C68","doi-asserted-by":"publisher","first-page":"2032","DOI":"10.1093\/emboj\/16.8.2032","article-title":"A model for extradenticle function as a switch that changes HOX proteins from repressors to activators","volume":"16","author":"Pinsonneault","year":"1997","journal-title":"EMBO J."},{"key":"2026070106063340300_DEV205511C69","doi-asserted-by":"publisher","first-page":"587","DOI":"10.1016\/S0092-8674(00)80662-5","article-title":"Structure of a HoxB1-Pbx1 heterodimer bound to DNA: role of the hexapeptide and a fourth homeodomain helix in complex formation","volume":"96","author":"Piper","year":"1999","journal-title":"Cell"},{"key":"2026070106063340300_DEV205511C70","doi-asserted-by":"publisher","first-page":"1031","DOI":"10.1016\/S0092-8674(05)80008-X","article-title":"Segmental expression of Hoxb-1 is controlled by a highly conserved autoregulatory loop dependent upon exd\/pbx","volume":"81","author":"P\u00f6pperl","year":"1995","journal-title":"Cell"},{"key":"2026070106063340300_DEV205511C71","doi-asserted-by":"publisher","first-page":"255","DOI":"10.1016\/S1097-2765(00)00027-7","article-title":"lazarus is a novel pbx gene that globally mediates hox gene function in zebrafish","volume":"6","author":"P\u00f6pperl","year":"2000","journal-title":"Mol. Cell"},{"key":"2026070106063340300_DEV205511C72","doi-asserted-by":"publisher","first-page":"207","DOI":"10.1016\/j.ydbio.2008.08.029","article-title":"unc-3-dependent repression of specific motor neuron fates in Caenorhabditis elegans","volume":"323","author":"Prasad","year":"2008","journal-title":"Dev. Biol."},{"key":"2026070106063340300_DEV205511C73","doi-asserted-by":"publisher","first-page":"W160","DOI":"10.1093\/nar\/gkw257","article-title":"deepTools2: a next generation web server for deep-sequencing data analysis","volume":"44","author":"Ramirez","year":"2016","journal-title":"Nucleic Acids Res."},{"key":"2026070106063340300_DEV205511C74","doi-asserted-by":"publisher","first-page":"3561","DOI":"10.1002\/j.1460-2075.1994.tb06663.x","article-title":"Coordinate regulation of downstream genes by extradenticle and the homeotic selector proteins","volume":"13","author":"Rauskolb","year":"1994","journal-title":"EMBO J."},{"key":"2026070106063340300_DEV205511C75","doi-asserted-by":"publisher","first-page":"e1003252","DOI":"10.1371\/journal.pgen.1003252","article-title":"Antagonism versus cooperativity with TALE cofactors at the base of the functional diversification of Hox protein function","volume":"9","author":"Rivas","year":"2013","journal-title":"PLoS Genet."},{"key":"2026070106063340300_DEV205511C76","doi-asserted-by":"publisher","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":"2026070106063340300_DEV205511C77","doi-asserted-by":"publisher","first-page":"103133","DOI":"10.1016\/j.xpro.2024.103133","article-title":"Protocol for auxin-inducible protein degradation in C. elegans using different auxins and TIR1-expressing strains","volume":"5","author":"Sharma","year":"2024","journal-title":"STAR Protoc."},{"key":"2026070106063340300_DEV205511C78","doi-asserted-by":"publisher","first-page":"898","DOI":"10.1093\/nar\/24.5.898","article-title":"Hox homeodomain proteins exhibit selective complex stabilities with Pbx and DNA","volume":"24","author":"Shen","year":"1996","journal-title":"Nucleic Acids Res."},{"key":"2026070106063340300_DEV205511C79","doi-asserted-by":"publisher","first-page":"284","DOI":"10.1186\/1471-2164-15-284","article-title":"ngs.plot: quick mining and visualization of next-generation sequencing data by integrating genomic databases","volume":"15","author":"Shen","year":"2014","journal-title":"BMC Genomics"},{"key":"2026070106063340300_DEV205511C80","doi-asserted-by":"publisher","first-page":"25875","DOI":"10.1074\/jbc.C110.150482","article-title":"Structural determination of functional domains in early B-cell factor (EBF) family of transcription factors reveals similarities to Rel DNA-binding proteins and a novel dimerization motif","volume":"285","author":"Siponen","year":"2010","journal-title":"J. Biol. Chem."},{"key":"2026070106063340300_DEV205511C81","doi-asserted-by":"publisher","first-page":"1270","DOI":"10.1016\/j.cell.2011.10.053","article-title":"Cofactor binding evokes latent differences in DNA binding specificity between Hox proteins","volume":"147","author":"Slattery","year":"2011","journal-title":"Cell"},{"key":"2026070106063340300_DEV205511C82","doi-asserted-by":"publisher","first-page":"113857","DOI":"10.1016\/j.celrep.2024.113857","article-title":"A molecular atlas of adult C. elegans motor neurons reveals ancient diversity delineated by conserved transcription factor codes","volume":"43","author":"Smith","year":"2024","journal-title":"Cell Rep."},{"key":"2026070106063340300_DEV205511C83","doi-asserted-by":"publisher","first-page":"733","DOI":"10.1016\/j.neuron.2015.07.031","article-title":"Regulatory logic of pan-neuronal gene expression in C. elegans","volume":"87","author":"Stefanakis","year":"2015","journal-title":"Neuron"},{"key":"2026070106063340300_DEV205511C84","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1348\/000711005X48266","article-title":"K-means clustering: a half-century synthesis","volume":"59","author":"Steinley","year":"2006","journal-title":"Br. J. Math. Stat. Psychol."},{"key":"2026070106063340300_DEV205511C85","doi-asserted-by":"publisher","first-page":"25","DOI":"10.1016\/S0092-8674(00)80603-0","article-title":"MIG-13 positions migrating cells along the anteroposterior body axis of C. elegans","volume":"98","author":"Sym","year":"1999","journal-title":"Cell"},{"key":"2026070106063340300_DEV205511C86","doi-asserted-by":"publisher","first-page":"661","DOI":"10.1016\/j.ydbio.2006.09.049","article-title":"Transcriptional control of Notch signaling by a HOX and a PBX\/EXD protein during vulval development in C. elegans","volume":"302","author":"Takacs-Vellai","year":"2007","journal-title":"Dev. Biol."},{"key":"2026070106063340300_DEV205511C87","doi-asserted-by":"publisher","first-page":"78","DOI":"10.1186\/1471-213X-10-78","article-title":"The C. elegans Hox gene ceh-13 regulates cell migration and fusion in a non-colinear way. Implications for the early evolution of Hox clusters","volume":"10","author":"Tihanyi","year":"2010","journal-title":"BMC Dev. Biol."},{"key":"2026070106063340300_DEV205511C88","doi-asserted-by":"publisher","first-page":"617","DOI":"10.1016\/0092-8674(94)90526-6","article-title":"Extradenticle raises the DNA binding specificity of homeotic selector gene products","volume":"78","author":"van Dijk","year":"1994","journal-title":"Cell"},{"key":"2026070106063340300_DEV205511C89","doi-asserted-by":"publisher","first-page":"e10255","DOI":"10.15252\/msb.202110255","article-title":"Single-cell RNA sequencing of motoneurons identifies regulators of synaptic wiring in Drosophila embryos","volume":"18","author":"Velten","year":"2022","journal-title":"Mol. Syst. Biol."},{"key":"2026070106063340300_DEV205511C90","doi-asserted-by":"publisher","first-page":"125","DOI":"10.1016\/S0074-7742(05)69005-8","article-title":"The motor circuit","volume":"69","author":"von Stetina","year":"2006","journal-title":"Int. Rev. Neurobiol."},{"key":"2026070106063340300_DEV205511C91","doi-asserted-by":"publisher","first-page":"550","DOI":"10.1016\/j.ydbio.2006.05.008","article-title":"Identification of cis-regulatory elements from the C. elegans Hox gene lin-39 required for embryonic expression and for regulation by the transcription factors LIN-1, LIN-31 and LIN-39","volume":"297","author":"Wagmaister","year":"2006","journal-title":"Dev. Biol."},{"key":"2026070106063340300_DEV205511C92","doi-asserted-by":"publisher","first-page":"1413","DOI":"10.1242\/dev.01569","article-title":"The roles of two C. elegans HOX co-factor orthologs in cell migration and vulva development","volume":"132","author":"Yang","year":"2005","journal-title":"Development"},{"key":"2026070106063340300_DEV205511C93","doi-asserted-by":"publisher","first-page":"287","DOI":"10.1006\/dbio.2002.0849","article-title":"Auto\/cross-regulation of Hoxb3 expression in posterior hindbrain and spinal cord","volume":"252","author":"Yau","year":"2002","journal-title":"Dev. Biol."},{"key":"2026070106063340300_DEV205511C94","doi-asserted-by":"publisher","first-page":"272","DOI":"10.1016\/j.cell.2020.12.012","article-title":"NeuroPAL: a multicolor atlas for whole-brain neuronal identification in C. elegans","volume":"184","author":"Yemini","year":"2021","journal-title":"Cell"},{"key":"2026070106063340300_DEV205511C95","doi-asserted-by":"publisher","first-page":"2382","DOI":"10.1093\/bioinformatics\/btv145","article-title":"ChIPseeker: an R\/Bioconductor package for ChIP peak annotation, comparison and visualization","volume":"31","author":"Yu","year":"2015","journal-title":"Bioinformatics"},{"key":"2026070106063340300_DEV205511C96","doi-asserted-by":"publisher","first-page":"R137","DOI":"10.1186\/gb-2008-9-9-r137","article-title":"Model-based analysis of ChIP-Seq (MACS)","volume":"9","author":"Zhang","year":"2008","journal-title":"Genome Biol."},{"key":"2026070106063340300_DEV205511C97","doi-asserted-by":"publisher","first-page":"4374","DOI":"10.1242\/dev.129635","article-title":"The auxin-inducible degradation (AID) system enables versatile conditional protein depletion in C. elegans","volume":"142","author":"Zhang","year":"2015","journal-title":"Development"},{"key":"2026070106063340300_DEV205511C98","doi-asserted-by":"publisher","first-page":"514","DOI":"10.1016\/j.neuron.2015.09.049","article-title":"Hox genes promote neuronal subtype diversification through posterior induction in Caenorhabditis elegans","volume":"88","author":"Zheng","year":"2015","journal-title":"Neuron"},{"key":"2026070106063340300_DEV205511C99","doi-asserted-by":"publisher","first-page":"1343","DOI":"10.1016\/j.celrep.2015.10.044","article-title":"Hox proteins act as transcriptional guarantors to ensure terminal differentiation","volume":"13","author":"Zheng","year":"2015","journal-title":"Cell Rep."},{"key":"2026070106063340300_DEV205511C100","doi-asserted-by":"publisher","first-page":"e1010092","DOI":"10.1371\/journal.pgen.1010092","article-title":"Nervous system-wide analysis of Hox regulation of terminal neuronal fate specification in Caenorhabditis elegans","volume":"18","author":"Zheng","year":"2022","journal-title":"PLoS Genet."}],"container-title":["Development"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/journals.biologists.com\/dev\/article-pdf\/doi\/10.1242\/dev.205511\/3706749\/dev205511.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/dev\/article-pdf\/doi\/10.1242\/dev.205511\/3706749\/dev205511.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T10:08:30Z","timestamp":1782900510000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.biologists.com\/dev\/article\/153\/13\/dev205511\/372120\/PBX-dependent-and-independent-Hox-programs"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,1]]},"references-count":100,"journal-issue":{"issue":"13","published-print":{"date-parts":[[2026,7,1]]}},"URL":"https:\/\/doi.org\/10.1242\/dev.205511","relation":{},"ISSN":["0950-1991","1477-9129"],"issn-type":[{"value":"0950-1991","type":"print"},{"value":"1477-9129","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,1]]},"article-number":"dev205511"}}