{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,7]],"date-time":"2026-08-07T23:44:26Z","timestamp":1786146266941,"version":"build-2736575974"},"reference-count":82,"publisher":"Elsevier BV","issue":"7","license":[{"start":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T00:00:00Z","timestamp":1782864000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T00:00:00Z","timestamp":1782864000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2026,5,26]],"date-time":"2026-05-26T00:00:00Z","timestamp":1779753600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000049","name":"National Institute on Aging","doi-asserted-by":"publisher","award":["AG000214"],"award-info":[{"award-number":["AG000214"]}],"id":[{"id":"10.13039\/100000049","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000026","name":"Institute of Aging","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100000026","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["10-AG-0423\/CR002894"],"award-info":[{"award-number":["10-AG-0423\/CR002894"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["03-AG-0325"],"award-info":[{"award-number":["03-AG-0325"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["15-AG-0063"],"award-info":[{"award-number":["15-AG-0063"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000062","name":"National Institute of Diabetes and Digestive and Kidney Diseases","doi-asserted-by":"publisher","award":["2UC4DK098085"],"award-info":[{"award-number":["2UC4DK098085"]}],"id":[{"id":"10.13039\/100000062","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["jbc.org","elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Journal of Biological Chemistry"],"published-print":{"date-parts":[[2026,7]]},"DOI":"10.1016\/j.jbc.2026.113209","type":"journal-article","created":{"date-parts":[[2026,5,29]],"date-time":"2026-05-29T06:43:49Z","timestamp":1780037029000},"page":"113209","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":0,"title":["Cell type-specific expression and subcellular localization of the human insulin upstream open reading frame (INSU) protein in pancreatic \u03b2-cells"],"prefix":"10.1016","volume":"302","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-8477-6452","authenticated-orcid":false,"given":"Qing-Rong","family":"Liu","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6614-606X","authenticated-orcid":false,"given":"Min","family":"Zhu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lisa M.","family":"Hartnell","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jane","family":"Tian","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qin","family":"Yao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaoming","family":"Zhong","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chee W.","family":"Chia","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Paritosh","family":"Ghosh","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"M\u00e1ire E.","family":"Doyle","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jennifer F.","family":"O\u2019Connell","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Josephine M.","family":"Egan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.jbc.2026.113209_bib1","doi-asserted-by":"crossref","first-page":"535","DOI":"10.1038\/331535a0","article-title":"Growth-controlling molluscan neurons produce the precursor of an insulin-related peptide","volume":"331","author":"Smit","year":"1988","journal-title":"Nature"},{"key":"10.1016\/j.jbc.2026.113209_bib2","doi-asserted-by":"crossref","first-page":"264","DOI":"10.1038\/s41559-022-01925-6","article-title":"De novo genes with an lncRNA origin encode unique human brain developmental functionality","volume":"7","author":"An","year":"2023","journal-title":"Nat. Ecol. Evol."},{"key":"10.1016\/j.jbc.2026.113209_bib3","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1000734","article-title":"A human-specific de novo protein-coding gene associated with human brain functions","volume":"6","author":"Li","year":"2010","journal-title":"Plos Comput. Biol."},{"key":"10.1016\/j.jbc.2026.113209_bib4","doi-asserted-by":"crossref","first-page":"115","DOI":"10.1016\/j.ygeno.2016.09.003","article-title":"Insights in human epigenomic dynamics through comparative primate analysis","volume":"108","author":"Bell","year":"2016","journal-title":"Genomics"},{"key":"10.1016\/j.jbc.2026.113209_bib5","doi-asserted-by":"crossref","first-page":"8817","DOI":"10.1073\/pnas.1802561115","article-title":"Human exonization through differential nucleosome occupancy","volume":"115","author":"Li","year":"2018","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"10.1016\/j.jbc.2026.113209_bib6","doi-asserted-by":"crossref","first-page":"2453","DOI":"10.1093\/molbev\/msx212","article-title":"Isoform evolution in Primates through independent combination of alternative RNA processing events","volume":"34","author":"Zhang","year":"2017","journal-title":"Mol. Biol. Evol."},{"key":"10.1016\/j.jbc.2026.113209_bib7","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pgen.1002942","article-title":"Hominoid-specific de novo protein-coding genes originating from long non-coding RNAs","volume":"8","author":"Xie","year":"2012","journal-title":"Plos Genet."},{"key":"10.1016\/j.jbc.2026.113209_bib8","doi-asserted-by":"crossref","first-page":"232","DOI":"10.1016\/j.compbiolchem.2010.08.001","article-title":"Computational identification and characterization of primate-specific microRNAs in human genome","volume":"34","author":"Lin","year":"2010","journal-title":"Comput. Biol. Chem."},{"key":"10.1016\/j.jbc.2026.113209_bib9","article-title":"A Human-Specific De Novo Gene Promotes Cortical Expansion and Folding","volume":"10","author":"Qi","year":"2023","journal-title":"Adv. Sci. (Weinh)"},{"key":"10.1016\/j.jbc.2026.113209_bib10","doi-asserted-by":"crossref","first-page":"287","DOI":"10.1016\/j.cell.2022.12.017","article-title":"Resurrection of endogenous retroviruses during aging reinforces senescence","volume":"186","author":"Liu","year":"2023","journal-title":"Cell"},{"key":"10.1016\/j.jbc.2026.113209_bib11","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pbio.1001179","article-title":"Accelerated recruitment of new brain development genes into the human genome","volume":"9","author":"Zhang","year":"2011","journal-title":"Plos Biol."},{"key":"10.1016\/j.jbc.2026.113209_bib12","doi-asserted-by":"crossref","DOI":"10.3390\/cells13221903","article-title":"An insulin upstream open reading frame (INSU) is present in skeletal muscle satellite cells: changes with Age","volume":"13","author":"Liu","year":"2024","journal-title":"Cells"},{"key":"10.1016\/j.jbc.2026.113209_bib13","doi-asserted-by":"crossref","first-page":"424","DOI":"10.1093\/molbev\/msg050","article-title":"Estimation of divergence times for major lineages of primate species","volume":"20","author":"Glazko","year":"2003","journal-title":"Mol. Biol. Evol."},{"key":"10.1016\/j.jbc.2026.113209_bib14","doi-asserted-by":"crossref","first-page":"2947","DOI":"10.2337\/db21-0198","article-title":"Novel human insulin isoforms and Calpha-Peptide product in islets of Langerhans and choroid plexus","volume":"70","author":"Liu","year":"2021","journal-title":"Diabetes"},{"key":"10.1016\/j.jbc.2026.113209_bib15","doi-asserted-by":"crossref","first-page":"770","DOI":"10.2337\/diabetes.51.3.770","article-title":"Unprocessed proinsulin promotes cell survival during neurulation in the chick embryo","volume":"51","author":"Hernandez-Sanchez","year":"2002","journal-title":"Diabetes"},{"key":"10.1016\/j.jbc.2026.113209_bib16","doi-asserted-by":"crossref","first-page":"5","DOI":"10.3389\/fnevo.2012.00005","article-title":"A bird's eye view of Human Language evolution","volume":"4","author":"Berwick","year":"2012","journal-title":"Front Evol. Neurosci."},{"key":"10.1016\/j.jbc.2026.113209_bib17","doi-asserted-by":"crossref","first-page":"5582","DOI":"10.1093\/emboj\/cdg515","article-title":"Upstream AUGs in embryonic proinsulin mRNA control its low translation level","volume":"22","author":"Hernandez-Sanchez","year":"2003","journal-title":"EMBO J."},{"key":"10.1016\/j.jbc.2026.113209_bib18","doi-asserted-by":"crossref","first-page":"128","DOI":"10.1016\/j.tig.2015.01.001","article-title":"Alternative mRNA transcription, processing, and translation: insights from RNA sequencing","volume":"31","author":"de Klerk","year":"2015","journal-title":"Trends Genet."},{"key":"10.1016\/j.jbc.2026.113209_bib19","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1007\/978-3-030-19966-1_5","article-title":"Translational regulation by upstream open reading frames and human diseases","volume":"1157","author":"Silva","year":"2019","journal-title":"Adv. Exp. Med. Biol."},{"key":"10.1016\/j.jbc.2026.113209_bib20","doi-asserted-by":"crossref","first-page":"1076","DOI":"10.1038\/s41467-021-21394-y","article-title":"Determinants of genome-wide distribution and evolution of uORFs in eukaryotes","volume":"12","author":"Zhang","year":"2021","journal-title":"Nat. Commun."},{"key":"10.1016\/j.jbc.2026.113209_bib21","doi-asserted-by":"crossref","first-page":"609","DOI":"10.1101\/gr.230938.117","article-title":"Recognition of the polycistronic nature of human genes is critical to understanding the genotype-phenotype relationship","volume":"28","author":"Brunet","year":"2018","journal-title":"Genome Res."},{"key":"10.1016\/j.jbc.2026.113209_bib22","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pgen.1003529","article-title":"Gene expression regulation by upstream open reading frames and human disease","volume":"9","author":"Barbosa","year":"2013","journal-title":"Plos Genet."},{"key":"10.1016\/j.jbc.2026.113209_bib23","doi-asserted-by":"crossref","first-page":"494","DOI":"10.1038\/nature21036","article-title":"Translation from unconventional 5' start sites drives tumour initiation","volume":"541","author":"Sendoel","year":"2017","journal-title":"Nature"},{"key":"10.1016\/j.jbc.2026.113209_bib24","doi-asserted-by":"crossref","first-page":"4091","DOI":"10.1093\/nar\/8.18.4091","article-title":"Analysis of the regions flanking the human insulin gene and sequence of an Alu family member","volume":"8","author":"Bell","year":"1980","journal-title":"Nucleic Acids Res."},{"key":"10.1016\/j.jbc.2026.113209_bib25","doi-asserted-by":"crossref","first-page":"8125","DOI":"10.1093\/nar\/15.20.8125","article-title":"An analysis of 5'-noncoding sequences from 699 vertebrate messenger RNAs","volume":"15","author":"Kozak","year":"1987","journal-title":"Nucleic Acids Res."},{"key":"10.1016\/j.jbc.2026.113209_bib26","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1005170","article-title":"PreTIS: a tool to predict non-canonical 5' UTR translational initiation sites in human and mouse","volume":"12","author":"Reuter","year":"2016","journal-title":"Plos Comput. Biol."},{"key":"10.1016\/j.jbc.2026.113209_bib27","doi-asserted-by":"crossref","first-page":"e91","DOI":"10.1093\/nar\/gkw104","article-title":"SRAMP: prediction of mammalian N6-methyladenosine (m6A) sites based on sequence-derived features","volume":"44","author":"Zhou","year":"2016","journal-title":"Nucleic Acids Res."},{"key":"10.1016\/j.jbc.2026.113209_bib28","doi-asserted-by":"crossref","first-page":"200","DOI":"10.1007\/s10930-019-09838-3","article-title":"A brief history of protein sorting prediction","volume":"38","author":"Nielsen","year":"2019","journal-title":"Protein J."},{"key":"10.1016\/j.jbc.2026.113209_bib29","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0136360","article-title":"Identification of Human N-Myristoylated proteins from Human complementary DNA resources by cell-free and cellular metabolic labeling analyses","volume":"10","author":"Takamitsu","year":"2015","journal-title":"PLoS One"},{"key":"10.1016\/j.jbc.2026.113209_bib30","doi-asserted-by":"crossref","first-page":"841","DOI":"10.1038\/446841a","article-title":"Chimps lead evolutionary race","volume":"446","author":"Hopkin","year":"2007","journal-title":"Nature"},{"key":"10.1016\/j.jbc.2026.113209_bib31","doi-asserted-by":"crossref","first-page":"3709","DOI":"10.1093\/bioinformatics\/btv449","article-title":"SNPGenie: estimating evolutionary parameters to detect natural selection using pooled next-generation sequencing data","volume":"31","author":"Nelson","year":"2015","journal-title":"Bioinformatics"},{"key":"10.1016\/j.jbc.2026.113209_bib32","doi-asserted-by":"crossref","first-page":"68","DOI":"10.1038\/nature15393","article-title":"A global reference for human genetic variation","volume":"526","author":"Genomes Project","year":"2015","journal-title":"Nature"},{"key":"10.1016\/j.jbc.2026.113209_bib33","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1038\/nature12228","article-title":"Great ape genetic diversity and population history","volume":"499","author":"Prado-Martinez","year":"2013","journal-title":"Nature"},{"key":"10.1016\/j.jbc.2026.113209_bib34","doi-asserted-by":"crossref","first-page":"1143","DOI":"10.1007\/s00592-015-0805-1","article-title":"Insulin gene VNTR polymorphisms -2221MspI and -23HphI are associated with type 1 diabetes and latent autoimmune diabetes in adults: a meta-analysis","volume":"52","author":"Zhang","year":"2015","journal-title":"Acta Diabetol."},{"key":"10.1016\/j.jbc.2026.113209_bib35","doi-asserted-by":"crossref","first-page":"54","DOI":"10.3390\/jcm7030054","article-title":"The cells of the islets of Langerhans","volume":"7","author":"Da Silva Xavier","year":"2018","journal-title":"J. Clin. Med."},{"key":"10.1016\/j.jbc.2026.113209_bib36","doi-asserted-by":"crossref","first-page":"4469","DOI":"10.1091\/mbc.e08-05-0499","article-title":"Translationally repressed mRNA transiently cycles through stress granules during stress","volume":"19","author":"Mollet","year":"2008","journal-title":"Mol. Biol. Cell"},{"key":"10.1016\/j.jbc.2026.113209_bib37","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1016\/j.redox.2016.11.004","article-title":"Inhibition of autophagy with bafilomycin and chloroquine decreases mitochondrial quality and bioenergetic function in primary neurons","volume":"11","author":"Redmann","year":"2017","journal-title":"Redox Biol."},{"issue":"Suppl 1","key":"10.1016\/j.jbc.2026.113209_bib38","doi-asserted-by":"crossref","first-page":"1719","DOI":"10.2337\/db20-1719-P","article-title":"Novel mass spectrometry based selected reaction monitoring proteomics for analysis of low abundant insulin levels in cerebrospinal fluid [abstract]","volume":"69","author":"Liu","year":"2020","journal-title":"Diabetes"},{"key":"10.1016\/j.jbc.2026.113209_bib39","doi-asserted-by":"crossref","first-page":"247","DOI":"10.1055\/s-2007-1012285","article-title":"Prolonged plasma half-life of insulin in patients with a genetic defect of high affinity binding sites","volume":"18","author":"Dreyer","year":"1986","journal-title":"Horm. Metab. Res."},{"key":"10.1016\/j.jbc.2026.113209_bib40","doi-asserted-by":"crossref","first-page":"501","DOI":"10.1038\/nm.4289","article-title":"Autoimmunity against a defective ribosomal insulin gene product in type 1 diabetes","volume":"23","author":"Kracht","year":"2017","journal-title":"Nat. Med."},{"key":"10.1016\/j.jbc.2026.113209_bib41","doi-asserted-by":"crossref","first-page":"884","DOI":"10.1007\/s00125-023-05882-y","article-title":"Presence of immunogenic alternatively spliced insulin gene product in human pancreatic delta cells","volume":"66","author":"van Tienhoven","year":"2023","journal-title":"Diabetologia"},{"key":"10.1016\/j.jbc.2026.113209_bib42","doi-asserted-by":"crossref","first-page":"511","DOI":"10.1093\/bioinformatics\/btv639","article-title":"Gapped sequence alignment using artificial neural networks: application to the MHC class I system","volume":"32","author":"Andreatta","year":"2016","journal-title":"Bioinformatics"},{"key":"10.1016\/j.jbc.2026.113209_bib43","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1016\/j.coi.2007.11.005","article-title":"Insulin as an autoantigen in NOD\/human diabetes","volume":"20","author":"Zhang","year":"2008","journal-title":"Curr. Opin. Immunol."},{"key":"10.1016\/j.jbc.2026.113209_bib44","doi-asserted-by":"crossref","first-page":"241","DOI":"10.1038\/308241a0","article-title":"Point mutations close to the AUG initiator codon affect the efficiency of translation of rat preproinsulin in vivo","volume":"308","author":"Kozak","year":"1984","journal-title":"Nature"},{"key":"10.1016\/j.jbc.2026.113209_bib45","doi-asserted-by":"crossref","first-page":"504","DOI":"10.1016\/j.molcel.2017.10.002","article-title":"m(6)A facilitates eIF4F-Independent mRNA translation","volume":"68","author":"Coots","year":"2017","journal-title":"Mol. Cell"},{"key":"10.1016\/j.jbc.2026.113209_bib46","doi-asserted-by":"crossref","first-page":"585","DOI":"10.1016\/j.gendis.2020.06.011","article-title":"Reversible N6-methyladenosine of RNA: the regulatory mechanisms on gene expression and implications in physiology and pathology","volume":"7","author":"Fang","year":"2020","journal-title":"Genes Dis."},{"key":"10.1016\/j.jbc.2026.113209_bib47","doi-asserted-by":"crossref","first-page":"237","DOI":"10.2337\/db23-0360","article-title":"METTL3-mediated m6A methylation controls pancreatic bipotent progenitor fate and islet Formation","volume":"73","author":"Sun","year":"2024","journal-title":"Diabetes"},{"key":"10.1016\/j.jbc.2026.113209_bib48","doi-asserted-by":"crossref","first-page":"321","DOI":"10.1007\/s00125-009-1562-x","article-title":"The long lifespan and low turnover of human islet beta cells estimated by mathematical modelling of lipofuscin accumulation","volume":"53","author":"Cnop","year":"2010","journal-title":"Diabetologia"},{"key":"10.1016\/j.jbc.2026.113209_bib49","doi-asserted-by":"crossref","first-page":"5988","DOI":"10.1074\/jbc.M108375200","article-title":"Regulated translation termination at the upstream open reading frame in s-adenosylmethionine decarboxylase mRNA","volume":"277","author":"Raney","year":"2002","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.jbc.2026.113209_bib50","doi-asserted-by":"crossref","first-page":"90","DOI":"10.1186\/s13059-015-0651-z","article-title":"Oxygen and glucose deprivation induces widespread alterations in mRNA translation within 20 minutes","volume":"16","author":"Andreev","year":"2015","journal-title":"Genome Biol."},{"key":"10.1016\/j.jbc.2026.113209_bib51","doi-asserted-by":"crossref","first-page":"1024","DOI":"10.1016\/j.molmet.2017.06.001","article-title":"Endoplasmic reticulum stress and eIF2alpha phosphorylation: the Achilles heel of pancreatic beta cells","volume":"6","author":"Cnop","year":"2017","journal-title":"Mol. Metab."},{"key":"10.1016\/j.jbc.2026.113209_bib52","doi-asserted-by":"crossref","first-page":"706","DOI":"10.15252\/embj.201592759","article-title":"Upstream ORFs are prevalent translational repressors in vertebrates","volume":"35","author":"Johnstone","year":"2016","journal-title":"EMBO J."},{"key":"10.1016\/j.jbc.2026.113209_bib53","doi-asserted-by":"crossref","first-page":"1165","DOI":"10.1016\/S1097-2765(01)00265-9","article-title":"Translational control is required for the unfolded protein response and in vivo glucose homeostasis","volume":"7","author":"Scheuner","year":"2001","journal-title":"Mol. Cell"},{"key":"10.1016\/j.jbc.2026.113209_bib54","doi-asserted-by":"crossref","first-page":"3201","DOI":"10.2337\/db06-0788","article-title":"Comparative analysis of insulin gene promoters: implications for diabetes research","volume":"55","author":"Hay","year":"2006","journal-title":"Diabetes"},{"key":"10.1016\/j.jbc.2026.113209_bib55","doi-asserted-by":"crossref","first-page":"979","DOI":"10.1083\/jcb.201106098","article-title":"Biogenesis of a novel compartment for autophagosome-mediated unconventional protein secretion","volume":"195","author":"Bruns","year":"2011","journal-title":"J. Cell Biol."},{"key":"10.1016\/j.jbc.2026.113209_bib56","doi-asserted-by":"crossref","DOI":"10.1016\/j.lfs.2024.122460","article-title":"Pancreatic beta cell derived extracellular vesicles containing surface preproinsulin are involved in glucose stimulated insulin secretion","volume":"340","author":"Ghosh","year":"2024","journal-title":"Life Sci."},{"key":"10.1016\/j.jbc.2026.113209_bib57","doi-asserted-by":"crossref","DOI":"10.1242\/jcs.260741","article-title":"The role of crinophagy in quality control of the regulated secretory pathway","volume":"136","author":"Szenci","year":"2023","journal-title":"J. Cell Sci."},{"key":"10.1016\/j.jbc.2026.113209_bib58","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1007\/s00418-023-02256-8","article-title":"Insulin granule morphology and crinosome formation in mice lacking the pancreatic beta cell-specific phogrin (PTPRN2) gene","volume":"161","author":"Yasui","year":"2024","journal-title":"Histochem. Cell Biol."},{"key":"10.1016\/j.jbc.2026.113209_bib59","doi-asserted-by":"crossref","DOI":"10.1083\/jcb.202110164","article-title":"VAMP4 regulates insulin levels by targeting secretory granules to lysosomes","volume":"221","author":"Li","year":"2022","journal-title":"J. Cell Biol."},{"key":"10.1016\/j.jbc.2026.113209_bib60","article-title":"Anti-Inflammatory and pro-autophagy effects of the cannabinoid receptor CB2R: possibility of modulation in type 1 diabetes","volume":"12","author":"Liu","year":"2021","journal-title":"Front Pharmacol."},{"key":"10.1016\/j.jbc.2026.113209_bib61","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1086\/381662","article-title":"Meat-adaptive genes and the evolution of slower aging in humans","volume":"79","author":"Finch","year":"2004","journal-title":"Q. Rev. Biol."},{"key":"10.1016\/j.jbc.2026.113209_bib62","doi-asserted-by":"crossref","first-page":"56","DOI":"10.1016\/j.domaniend.2018.04.006","article-title":"Tissue-specific expression and circulating concentrations of nesfatin-1 in domestic animals","volume":"65","author":"Morton","year":"2018","journal-title":"Domest. Anim. Endocrinol."},{"key":"10.1016\/j.jbc.2026.113209_bib63","doi-asserted-by":"crossref","first-page":"3886S","DOI":"10.1093\/jn\/133.11.3886S","article-title":"The critical role played by animal source foods in human (Homo) evolution","volume":"133","author":"Milton","year":"2003","journal-title":"J. Nutr."},{"key":"10.1016\/j.jbc.2026.113209_bib64","doi-asserted-by":"crossref","first-page":"952","DOI":"10.1038\/s41586-025-09399-9","article-title":"The evolution of hominin bipedalism in two steps","volume":"645","author":"Senevirathne","year":"2025","journal-title":"Nature"},{"key":"10.1016\/j.jbc.2026.113209_bib65","doi-asserted-by":"crossref","first-page":"391","DOI":"10.1186\/s12864-019-5775-1","article-title":"Translation of upstream open reading frames in a model of neuronal differentiation","volume":"20","author":"Rodriguez","year":"2019","journal-title":"BMC Genomics"},{"key":"10.1016\/j.jbc.2026.113209_bib66","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1073\/pnas.1713611115","article-title":"Two types of aggression in human evolution","volume":"115","author":"Wrangham","year":"2018","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"10.1016\/j.jbc.2026.113209_bib67","doi-asserted-by":"crossref","first-page":"95","DOI":"10.2174\/1874609809666160211124947","article-title":"The origins of aging: evidence that aging is an adaptive phenotype","volume":"9","author":"Singer","year":"2016","journal-title":"Curr. Aging Sci."},{"key":"10.1016\/j.jbc.2026.113209_bib68","doi-asserted-by":"crossref","first-page":"867","DOI":"10.1038\/nrg3322","article-title":"Adaptive evolution: evaluating empirical support for theoretical predictions","volume":"13","author":"Olson-Manning","year":"2012","journal-title":"Nat. Rev. Genet."},{"key":"10.1016\/j.jbc.2026.113209_bib69","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1016\/j.arr.2016.02.002","article-title":"Apolipoprotein E (APOE) epsilon4 and episodic memory decline in Alzheimer's disease: a review","volume":"27","author":"El Haj","year":"2016","journal-title":"Ageing Res. Rev."},{"key":"10.1016\/j.jbc.2026.113209_bib70","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0047760","article-title":"The apolipoprotein E (APOE) gene appears functionally monomorphic in chimpanzees (Pan troglodytes)","volume":"7","author":"McIntosh","year":"2012","journal-title":"PLoS One"},{"key":"10.1016\/j.jbc.2026.113209_bib71","article-title":"Quantification of preproinsulin and insulin isoforms derived from upstream open reading frames in human clinical samples by selected reaction monitoring assay","volume":"71","author":"LIU","year":"2022","journal-title":"Diabetes"},{"key":"10.1016\/j.jbc.2026.113209_bib72","doi-asserted-by":"crossref","first-page":"170","DOI":"10.2337\/db16-0754","article-title":"Insulin resistance is associated with reduced mitochondrial oxidative capacity measured by 31P-Magnetic resonance spectroscopy in participants without diabetes from the Baltimore longitudinal Study of aging","volume":"66","author":"Fabbri","year":"2017","journal-title":"Diabetes"},{"key":"10.1016\/j.jbc.2026.113209_bib73","doi-asserted-by":"crossref","first-page":"635","DOI":"10.1038\/s43587-022-00243-7","article-title":"Longitudinal phenotypic aging metrics in the Baltimore longitudinal Study of Aging","volume":"2","author":"Kuo","year":"2022","journal-title":"Nat. Aging"},{"key":"10.1016\/j.jbc.2026.113209_bib74","doi-asserted-by":"crossref","first-page":"741","DOI":"10.2174\/1567205016666190913155950","article-title":"A pilot Study of exenatide actions in Alzheimer's Disease","volume":"16","author":"Mullins","year":"2019","journal-title":"Curr. Alzheimer Res."},{"key":"10.1016\/j.jbc.2026.113209_bib75","doi-asserted-by":"crossref","first-page":"2101","DOI":"10.1101\/gr.948003","article-title":"Global haplotype diversity in the human insulin gene region","volume":"13","author":"Stead","year":"2003","journal-title":"Genome Res."},{"key":"10.1016\/j.jbc.2026.113209_bib76","doi-asserted-by":"crossref","first-page":"28","DOI":"10.1186\/s13195-019-0475-2","article-title":"Neurexin 3 transmembrane and soluble isoform expression and splicing haplotype are associated with neuron inflammasome and Alzheimer's disease","volume":"11","author":"Hishimoto","year":"2019","journal-title":"Alzheimers Res. Ther."},{"key":"10.1016\/j.jbc.2026.113209_bib77","doi-asserted-by":"crossref","first-page":"W521","DOI":"10.1093\/nar\/gkae241","article-title":"The EMBL-EBI Job Dispatcher sequence analysis tools framework in 2024","volume":"52","author":"Madeira","year":"2024","journal-title":"Nucleic Acids Res."},{"key":"10.1016\/j.jbc.2026.113209_bib78","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1038\/35052055","article-title":"Rab proteins as membrane organizers","volume":"2","author":"Zerial","year":"2001","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"10.1016\/j.jbc.2026.113209_bib79","doi-asserted-by":"crossref","first-page":"8032","DOI":"10.1073\/pnas.1921619117","article-title":"Host membrane lipids are trafficked to membranes of intravacuolar bacterium Ehrlichia chaffeensis","volume":"117","author":"Lin","year":"2020","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"10.1016\/j.jbc.2026.113209_bib80","doi-asserted-by":"crossref","DOI":"10.1002\/pmic.201600237","article-title":"A novel, multiplexed targeted mass spectrometry assay for quantification of complement factor H (CFH) variants and CFH-related proteins 1-5 in human plasma","volume":"17","author":"Zhang","year":"2017","journal-title":"Proteomics"},{"key":"10.1016\/j.jbc.2026.113209_bib81","doi-asserted-by":"crossref","first-page":"697","DOI":"10.1126\/science.157.3789.697","article-title":"Insulin biosynthesis: evidence for a precursor","volume":"157","author":"Steiner","year":"1967","journal-title":"Science"},{"key":"10.1016\/j.jbc.2026.113209_bib82","first-page":"597","article-title":"Differential expression of vacuolar-type H+-ATPase between normal human pancreatic islet B-cells and insulinoma cells","volume":"11","author":"Ohta","year":"1997","journal-title":"Int. J. Oncol."}],"container-title":["Journal of Biological Chemistry"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0021925826020818?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0021925826020818?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,8,7]],"date-time":"2026-08-07T23:30:21Z","timestamp":1786145421000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0021925826020818"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7]]},"references-count":82,"journal-issue":{"issue":"7","published-print":{"date-parts":[[2026,7]]}},"alternative-id":["S0021925826020818"],"URL":"https:\/\/doi.org\/10.1016\/j.jbc.2026.113209","relation":{},"ISSN":["0021-9258"],"issn-type":[{"value":"0021-9258","type":"print"}],"subject":[],"published":{"date-parts":[[2026,7]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Cell type-specific expression and subcellular localization of the human insulin upstream open reading frame (INSU) protein in pancreatic \u03b2-cells","name":"articletitle","label":"Article Title"},{"value":"Journal of Biological Chemistry","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.jbc.2026.113209","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"Published by Elsevier Inc on behalf of American Society for Biochemistry and Molecular Biology.","name":"copyright","label":"Copyright"}],"article-number":"113209"}}