{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,16]],"date-time":"2026-07-16T16:56:05Z","timestamp":1784220965706,"version":"3.55.0"},"reference-count":60,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,1,14]],"date-time":"2026-01-14T00:00:00Z","timestamp":1768348800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Neurosci."],"abstract":"<jats:p>\n                    This study characterizes the requirements of the\n                    <jats:italic>kayak<\/jats:italic>\n                    (\n                    <jats:italic>kay<\/jats:italic>\n                    ) gene in\n                    <jats:italic>Drosophila melanogaster<\/jats:italic>\n                    adult eye biology by examining mutant phenotypes in photoreceptor development, external eye morphology, corneal and bristle ultrastructure, and visually guided behaviors, such as phototaxis, in\n                    <jats:italic>kay<\/jats:italic>\n                    strong loss-of-function homozygous mutant clones. Despite previous studies on\n                    <jats:italic>kay<\/jats:italic>\n                    , there is a dearth of phenotypic characterization of the morphological and behavioral consequences of\n                    <jats:italic>kay<\/jats:italic>\n                    loss-of-function alleles in the adult eye. We find that\n                    <jats:italic>kay<\/jats:italic>\n                    is expressed in developing ommatidia in eye discs. The\n                    <jats:italic>kay<\/jats:italic>\n                    mutant ommatidia are misaligned, lack photoreceptors, have malformed corneal surfaces, and have misshaped, misplaced, and fewer mechanosensory bristles. Corneal nipples, while present in mutant corneas on the corneal surface, are disorganized and malformed. With an average of 30% of the eye territory mutant, flies have a significantly lower response in a behavioral phototaxis assay. Altogether,\n                    <jats:italic>kay<\/jats:italic>\n                    function is required for multiple cell types in the adult retina, and this stands in stark contrast with other jun kinase genes, like the fly homologs of\n                    <jats:italic>jun kinase<\/jats:italic>\n                    and\n                    <jats:italic>jun<\/jats:italic>\n                    , genes not required for adult eye morphogenesis. This is consistent with Kayak functions that are independent of heterodimerizing with Jun proteins or requiring activation of the jun kinase pathway.\n                  <\/jats:p>","DOI":"10.3389\/fnins.2025.1703753","type":"journal-article","created":{"date-parts":[[2026,1,14]],"date-time":"2026-01-14T22:41:07Z","timestamp":1768430467000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":1,"title":["The fos homolog kayak is required for adult eye formation and function in Drosophila"],"prefix":"10.3389","volume":"19","author":[{"given":"Manuel","family":"Z\u00faniga-Garc\u00eda","sequence":"first","affiliation":[{"name":"Laboratorio D-04, Departamento de Neurobiolog\u00eda del Desarrollo y Neurofisiolog\u00eda, Instituto de Neurobiolog\u00eda, Universidad Nacional Aut\u00f3noma de M\u00e9xico","place":["Quer\u00e9taro, Mexico"]},{"name":"Posgrado en Ciencias Biol\u00f3gicas, Unidad de Posgrado, Ciudad Universitaria, Universidad Nacional Aut\u00f3noma de M\u00e9xico","place":["Coyoac\u00e1n, Mexico"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Juan Rafael","family":"Riesgo-Escovar","sequence":"additional","affiliation":[{"name":"Laboratorio D-04, Departamento de Neurobiolog\u00eda del Desarrollo y Neurofisiolog\u00eda, Instituto de Neurobiolog\u00eda, Universidad Nacional Aut\u00f3noma de M\u00e9xico","place":["Quer\u00e9taro, Mexico"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,1,14]]},"reference":[{"key":"ref1","doi-asserted-by":"publisher","first-page":"73","DOI":"10.1016\/j.mod.2018.05.006","article-title":"Fos metamorphoses: lessons from mutants in model organisms","volume":"154","author":"Alfonso-Gonzalez","year":"2018","journal-title":"Mech. Dev."},{"key":"ref2","doi-asserted-by":"publisher","first-page":"252","DOI":"10.1016\/j.ymeth.2014.04.007","article-title":"Eye development","volume":"68","author":"Baker","year":"2014","journal-title":"Methods"},{"key":"ref3","volume-title":"The development of","author":"Bate","year":"1993"},{"key":"ref4","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1387\/ijdb.11902676","article-title":"Maternal-effect loci involved in Drosophila oogenesis and embryogenesis: P element-induced mutations on the third chromosome","volume":"46","author":"Bellotto","year":"2002","journal-title":"Int. J. Dev. Biol."},{"key":"ref5","doi-asserted-by":"publisher","first-page":"e3001494","DOI":"10.1371\/journal.pbio.3001494","article-title":"Fos regulates macrophage infiltration against surrounding tissue resistance by a cortical actin-based mechanism in Drosophila","volume":"20","author":"Belyaeva","year":"2022","journal-title":"PLoS Biol."},{"key":"ref6","doi-asserted-by":"publisher","first-page":"1112","DOI":"10.1073\/pnas.58.3.1112","article-title":"Behavioral mutants of Drosophila isolated by countercurrent distribution","volume":"58","author":"Benzer","year":"1967","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref7","doi-asserted-by":"publisher","first-page":"1","DOI":"10.18637\/jss.v031.i10","article-title":"CircStat: a MATLAB toolbox for circular statistics","volume":"31","author":"Berens","year":"2009","journal-title":"J. Stat. Softw."},{"key":"ref8","doi-asserted-by":"publisher","first-page":"10750","DOI":"10.1073\/pnas.1505748112","article-title":"Diverse set of turing nanopatterns coat corneae across insect lineages","volume":"112","author":"Blagodatski","year":"2015","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref9","doi-asserted-by":"publisher","first-page":"7205","DOI":"10.1038\/s41467-023-43037-0","article-title":"A single cell genomics atlas of the Drosophila larval eye reveals distinct photoreceptor developmental timelines","volume":"14","author":"Bozllepogu Raja","year":"2023","journal-title":"Nat. Commun."},{"key":"ref10","doi-asserted-by":"publisher","first-page":"401","DOI":"10.1242\/dev.118.2.401","article-title":"Targeted gene expression as a means of altering cell fates and generating dominant phenotypes","volume":"118","author":"Brand","year":"1993","journal-title":"Development"},{"key":"ref11","doi-asserted-by":"publisher","first-page":"1241","DOI":"10.1016\/j.bbalip.2014.05.007","article-title":"C-Fos: an AP-1 transcription factor with an additional cytoplasmic, non-genomic lipid synthesis activation capacity","volume":"1841","author":"Caputto","year":"2014","journal-title":"Biochim. Biophys. Acta"},{"key":"ref12","doi-asserted-by":"publisher","first-page":"153","DOI":"10.1023\/A:1007508020173","article-title":"Immediate early gene expression within the visual system: light and circadian regulation in the retina and the suprachiasmatic nucleus","volume":"25","author":"Caputto","year":"2000","journal-title":"Neurochem. Res."},{"key":"ref13","doi-asserted-by":"publisher","first-page":"463","DOI":"10.1016\/0092-8674(95)90497-2","article-title":"Phyllopod functions in the fate determination of a subset of photoreceptors in Drosophila","volume":"80","author":"Chang","year":"1995","journal-title":"Cell"},{"key":"ref14","doi-asserted-by":"publisher","first-page":"259","DOI":"10.1016\/0092-8674(84)90219-8","article-title":"Viral and cellular fos proteins: a comparative analysis","volume":"36","author":"Curran","year":"1984","journal-title":"Cell"},{"key":"ref15","doi-asserted-by":"publisher","first-page":"47","DOI":"10.1016\/S0925-4773(01)00418-X","article-title":"Top-DER- and Dpp-dependent requirements for the Drosophila fos\/kayak gene in follicular epithelium morphogenesis","volume":"106","author":"Dequier","year":"2001","journal-title":"Mech. Dev."},{"key":"ref16","doi-asserted-by":"publisher","first-page":"453","DOI":"10.1016\/0092-8674(95)90496-4","article-title":"Control of Drosophila photoreceptor cell fates by phyllopod, a novel nuclear protein acting downstream of the Raf kinase","volume":"80","author":"Dickson","year":"1995","journal-title":"Cell"},{"key":"ref17","doi-asserted-by":"publisher","first-page":"698","DOI":"10.1126\/science.151.3711.698","article-title":"Virus induction of osteosarcomas in mice","volume":"151","author":"Finkel","year":"1966","journal-title":"Science"},{"key":"ref18","doi-asserted-by":"publisher","first-page":"128","DOI":"10.1016\/j.ydbio.2010.12.040","article-title":"Two-color in vivo imaging of photoreceptor apoptosis and development in Drosophila","volume":"351","author":"Gambis","year":"2011","journal-title":"Dev. Biol."},{"key":"ref19","doi-asserted-by":"publisher","first-page":"511","DOI":"10.1111\/j.1748-1716.1966.tb03232.x","article-title":"Ultrastructural ontogenesis of cornea and corneal nipples in the compound eye of insects","volume":"66","author":"Gemne","year":"1966","journal-title":"Acta Physiol. Scand."},{"key":"ref20","doi-asserted-by":"publisher","first-page":"3236","DOI":"10.1093\/plcell\/koad157","article-title":"The pyrenoid: the eukaryotic CO2-concentrating organelle","volume":"35","author":"He","year":"2023","journal-title":"Plant Cell"},{"key":"ref21","doi-asserted-by":"publisher","first-page":"pdb prot5199","DOI":"10.1101\/pdb.prot5199","article-title":"Recovery of DNA sequences flanking P-element insertions in Drosophila: inverse PCR and plasmid rescue","volume":"2009","author":"Huang","year":"2009","journal-title":"Cold Spring Harb. Protoc."},{"key":"ref22","doi-asserted-by":"publisher","first-page":"76","DOI":"10.1016\/j.gene.2008.05.001","article-title":"The gene structure of the Drosophila melanogaster proto-oncogene, kayak, and its nested gene, fos-intronic gene","volume":"420","author":"Hudson","year":"2008","journal-title":"Gene"},{"key":"ref23","doi-asserted-by":"publisher","first-page":"8293","DOI":"10.1128\/MCB.02455-05","article-title":"Control of G2\/M transition by Drosophila Fos","volume":"26","author":"Hyun","year":"2006","journal-title":"Mol. Cell. Biol."},{"key":"ref24","volume-title":"Statistical analysis and modelling of spatial point patterns","author":"Illian","year":"2008"},{"key":"ref25","doi-asserted-by":"publisher","first-page":"577","DOI":"10.1016\/0092-8674(92)90592-Z","article-title":"Pleiotropic effects of a null mutation in the c-fos proto-oncogene","volume":"71","author":"Johnson","year":"1992","journal-title":"Cell"},{"key":"ref26","doi-asserted-by":"publisher","first-page":"283","DOI":"10.1007\/BF00848157","article-title":"Mutations affecting the pattern of the larval cuticle in Drosophila melanogaster: II. Zygotic loci on the third chromosome","volume":"193","author":"Jurgens","year":"1984","journal-title":"Wilhelm Roux Arch. Dev. Biol."},{"key":"ref27","doi-asserted-by":"publisher","first-page":"1040","DOI":"10.1002\/1873-3468.13388","article-title":"Why the c-Fos\/c-Jun complex is extremely conserved: an in vitro evolution exploration by combining cDNA display and proximity ligation","volume":"593","author":"Kang","year":"2019","journal-title":"FEBS Lett."},{"key":"ref28","doi-asserted-by":"publisher","first-page":"e1006281","DOI":"10.1371\/journal.pgen.1006281","article-title":"C-Fos repression by piwi regulates Drosophila ovarian germline formation and tissue morphogenesis","volume":"12","author":"Klein","year":"2016","journal-title":"PLoS Genet."},{"key":"ref29","doi-asserted-by":"publisher","first-page":"287","DOI":"10.1016\/S0197-0186(98)00023-0","article-title":"C-Fos as a transcription factor: a stressful (re)view from a functional map","volume":"33","author":"Kovacs","year":"1998","journal-title":"Neurochem. Int."},{"key":"ref30","doi-asserted-by":"publisher","first-page":"136","DOI":"10.1002\/dvdy.23707","article-title":"Building an ommatidium one cell at a time","volume":"241","author":"Kumar","year":"2012","journal-title":"Dev. Dyn."},{"key":"ref31","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1002\/dvdy.24585","article-title":"The fly eye: through the looking glass","volume":"247","author":"Kumar","year":"2018","journal-title":"Dev. Dyn."},{"key":"ref32","doi-asserted-by":"publisher","first-page":"741","DOI":"10.1523\/JNEUROSCI.01-07-00741.1981","article-title":"Anatomy and physiology of experimentally produced striped tecta","volume":"1","author":"Law","year":"1981","journal-title":"J. Neurosci."},{"key":"ref33","doi-asserted-by":"publisher","first-page":"26","DOI":"10.1002\/gene.20022","article-title":"Drosophila eye disc margin is a center for organizing long-range planar polarity","volume":"39","author":"Lim","year":"2004","journal-title":"Genesis"},{"key":"ref34","doi-asserted-by":"publisher","first-page":"380","DOI":"10.1038\/sj.emboj.7601484","article-title":"Foxo and Fos regulate the decision between cell death and survival in response to UV irradiation","volume":"26","author":"Luo","year":"2007","journal-title":"EMBO J."},{"key":"ref35","doi-asserted-by":"publisher","first-page":"374","DOI":"10.1016\/j.ydbio.2017.06.023","article-title":"A population of G2-arrested cells are selected as sensory organ precursors for the interommatidial bristles of the Drosophila eye","volume":"430","author":"Meserve","year":"2017","journal-title":"Dev. Biol."},{"key":"ref36","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1016\/0092-8674(84)90073-4","article-title":"C-fos protein can induce cellular transformation: a novel mechanism of activation of a cellular oncogene","volume":"36","author":"Miller","year":"1984","journal-title":"Cell"},{"key":"ref37","doi-asserted-by":"publisher","first-page":"78","DOI":"10.3389\/fnmol.2015.00078","article-title":"Role of immediate-early genes in synaptic plasticity and neuronal ensembles underlying the memory trace","volume":"8","author":"Minatohara","year":"2015","journal-title":"Front. Mol. Neurosci."},{"key":"ref38","doi-asserted-by":"publisher","first-page":"3966","DOI":"10.1016\/j.cub.2024.07.060","article-title":"The JNK and hippo pathways control epithelial integrity and prevent tumor initiation by regulating an overlapping transcriptome","volume":"34","author":"Mitchell","year":"2024","journal-title":"Curr. Biol."},{"key":"ref39","doi-asserted-by":"publisher","first-page":"171","DOI":"10.1016\/j.mod.2016.12.003","article-title":"Piragua encodes a zinc finger protein required for development in Drosophila","volume":"144","author":"Nazario-Yepiz","year":"2017","journal-title":"Mech. Dev."},{"key":"ref40","doi-asserted-by":"publisher","first-page":"iyad211","DOI":"10.1093\/genetics\/iyad211","article-title":"FlyBase: updates to the Drosophila genes and genomes database","volume":"227","author":"Ozturk-Colak","year":"2024","journal-title":"Genetics"},{"key":"ref41","doi-asserted-by":"publisher","first-page":"822","DOI":"10.1101\/gad.4.5.822","article-title":"The Drosophila Fos-related AP-1 protein is a developmentally regulated transcription factor","volume":"4","author":"Perkins","year":"1990","journal-title":"Genes Dev."},{"key":"ref42","doi-asserted-by":"publisher","first-page":"e20230015","DOI":"10.1590\/1980-5764-dn-2023-0015","article-title":"Exercise induction at expression immediate early gene (c-Fos, ARC, EGR-1) in the hippocampus: a systematic review","volume":"18","author":"Rahmi","year":"2024","journal-title":"Dement. Neuropsychol."},{"key":"ref43","doi-asserted-by":"publisher","first-page":"471","DOI":"10.1101\/gad.2.12b.1687","article-title":"Common DNA binding site for Fos protein complexes and transcription factor AP-1","volume":"52","author":"Rauscher","year":"1988","journal-title":"Cell"},{"key":"ref44","doi-asserted-by":"publisher","first-page":"3353","DOI":"10.1242\/dev.124.17.3353","article-title":"D-Fos, a target gene of decapentaplegic signalling with a critical role during Drosophila endoderm induction","volume":"124","author":"Riese","year":"1997","journal-title":"Development"},{"key":"ref45","doi-asserted-by":"publisher","first-page":"669","DOI":"10.1126\/science.278.5338.669","article-title":"Common and distinct roles of DFos and DJun during Drosophila development","volume":"278","author":"Riesgo-Escovar","year":"","journal-title":"Science"},{"key":"ref46","doi-asserted-by":"publisher","first-page":"1717","DOI":"10.1101\/gad.11.13.1717","article-title":"Drosophila Jun kinase regulates expression of decapentaplegic via the ETS-domain protein Aop and the AP-1 transcription factor DJun during dorsal closure","volume":"11","author":"Riesgo-Escovar","year":"","journal-title":"Genes Dev."},{"key":"ref47","doi-asserted-by":"publisher","first-page":"2759","DOI":"10.1101\/gad.10.21.2759","article-title":"The Drosophila Jun-N-terminal kinase is required for cell morphogenesis but not for DJun-dependent cell fate specification in the eye","volume":"10","author":"Riesgo-Escovar","year":"1996","journal-title":"Genes Dev."},{"key":"ref48","volume-title":"Molecular cloning: a laboratory manual","author":"Sambrook","year":"1989"},{"key":"ref49","doi-asserted-by":"publisher","first-page":"477","DOI":"10.1016\/0896-6273(90)90106-P","article-title":"The regulation and function of c-fos and other immediate early genes in the nervous system","volume":"4","author":"Sheng","year":"1990","journal-title":"Neuron"},{"key":"ref50","doi-asserted-by":"publisher","first-page":"200356","DOI":"10.1098\/rsob.200356","article-title":"How do the Fat-Dachsous and core planar polarity pathways act together and independently to coordinate polarized cell behaviours?","volume":"11","author":"Strutt","year":"2021","journal-title":"Open Biol."},{"key":"ref51","doi-asserted-by":"publisher","first-page":"71","DOI":"10.1016\/S0925-4773(00)00455-X","article-title":"An autoregulatory function of Dfos during Drosophila endoderm induction","volume":"98","author":"Szuts","year":"2000","journal-title":"Mech. Dev."},{"key":"ref52","doi-asserted-by":"publisher","first-page":"355","DOI":"10.1006\/meth.1998.0591","article-title":"Use of FLP\/FRT system to study Drosophila development","volume":"14","author":"Theodosiou","year":"1998","journal-title":"Methods"},{"key":"ref53","doi-asserted-by":"publisher","first-page":"iyac041","DOI":"10.1093\/genetics\/iyac041","article-title":"The early history of the eye-antennal disc of Drosophila melanogaster","volume":"221","author":"Weasner","year":"2022","journal-title":"Genetics"},{"key":"ref54","doi-asserted-by":"publisher","first-page":"110","DOI":"10.1016\/j.ydbio.2008.01.016","article-title":"Combinatorial signaling by the frizzled\/PCP and Egfr pathways during planar cell polarity establishment in the Drosophila eye","volume":"316","author":"Weber","year":"2008","journal-title":"Dev. Biol."},{"key":"ref55","doi-asserted-by":"publisher","first-page":"7106","DOI":"10.1073\/pnas.0600869103","article-title":"Acute suppression, but not chronic genetic deficiency, of c-fos gene expression impairs long-term memory in aversive taste learning","volume":"103","author":"Yasoshima","year":"2006","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref56","doi-asserted-by":"publisher","first-page":"3947","DOI":"10.1242\/dev.126.17.3947","article-title":"Thorax closure in Drosophila: involvement of Fos and the JNK pathway","volume":"126","author":"Zeitlinger","year":"1999","journal-title":"Development"},{"key":"ref57","doi-asserted-by":"publisher","first-page":"7393","DOI":"10.1093\/emboj\/16.24.7393","article-title":"Defective dorsal closure and loss of epidermal decapentaplegic expression in Drosophila fos mutants","volume":"16","author":"Zeitlinger","year":"1997","journal-title":"EMBO J."},{"key":"ref58","doi-asserted-by":"publisher","first-page":"91","DOI":"10.1002\/syn.10115","article-title":"Hippocampal expression of c-fos is not essential for spatial learning","volume":"46","author":"Zhang","year":"2002","journal-title":"Synapse"},{"key":"ref59","doi-asserted-by":"publisher","first-page":"W45","DOI":"10.1093\/nar\/gkae358","article-title":"ChIP-atlas 3.0: a data-mining suite to explore chromosome architecture together with large-scale regulome data","volume":"52","author":"Zou","year":"2024","journal-title":"Nucleic Acids Res."},{"key":"ref60","doi-asserted-by":"publisher","first-page":"203997","DOI":"10.1016\/j.cdev.2025.203997","article-title":"Fos genes in mainly invertebrate model systems: a review of commonalities and some diversities","volume":"181","author":"Zuniga-Garcia","year":"2025","journal-title":"Cells Dev."}],"container-title":["Frontiers in Neuroscience"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fnins.2025.1703753\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,1,14]],"date-time":"2026-01-14T22:41:09Z","timestamp":1768430469000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fnins.2025.1703753\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,1,14]]},"references-count":60,"alternative-id":["10.3389\/fnins.2025.1703753"],"URL":"https:\/\/doi.org\/10.3389\/fnins.2025.1703753","relation":{},"ISSN":["1662-453X"],"issn-type":[{"value":"1662-453X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,1,14]]},"article-number":"1703753"}}