{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,21]],"date-time":"2026-10-21T14:37:56Z","timestamp":1792593476320,"version":"4.5.0"},"reference-count":91,"publisher":"Elsevier BV","issue":"11","license":[{"start":{"date-parts":[[2026,11,1]],"date-time":"2026-11-01T00:00:00Z","timestamp":1793491200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,11,1]],"date-time":"2026-11-01T00:00:00Z","timestamp":1793491200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2026,9,12]],"date-time":"2026-09-12T00:00:00Z","timestamp":1789171200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100012914","name":"Vidyasirimedhi Institute of Science and Technology","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100012914","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004192","name":"National Science and Technology Development Agency","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100004192","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":[[2026,11]]},"DOI":"10.1016\/j.jbc.2026.113568","type":"journal-article","created":{"date-parts":[[2026,9,18]],"date-time":"2026-09-18T08:50:19Z","timestamp":1789721419000},"page":"113568","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":0,"title":["Biochemical, bioluminescence and bioreporter properties of functionally improved firefly luciferases"],"prefix":"10.1016","volume":"302","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-8839-5814","authenticated-orcid":false,"given":"Chadaporn","family":"Kantiwiriyawanitch","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Vinutsada","family":"Pongsupasa","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Pratchaya","family":"Watthaisong","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jittima","family":"Phonbuppha","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6565-998X","authenticated-orcid":false,"given":"Aisaraphon","family":"Phintha","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Daisuke","family":"Sasaki","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Thanyaporn","family":"Wongnate","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ruchanok","family":"Tinikul","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3367-3127","authenticated-orcid":false,"given":"Kazuki","family":"Niwa","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7422-730X","authenticated-orcid":false,"given":"Yoshihiro","family":"Nakajima","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yoshihiro","family":"Ohmiya","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ubolsree","family":"Leartsakulpanich","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8533-1604","authenticated-orcid":false,"given":"Pimchai","family":"Chaiyen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.jbc.2026.113568_bib1","doi-asserted-by":"crossref","first-page":"5668","DOI":"10.1039\/D0CS01492C","article-title":"Applications of bioluminescence in biotechnology and beyond","volume":"50","author":"Syed","year":"2021","journal-title":"Chem. Soc. Rev."},{"key":"10.1016\/j.jbc.2026.113568_bib2","doi-asserted-by":"crossref","first-page":"180","DOI":"10.3390\/s110100180","article-title":"In vivo bioluminescent imaging (BLI): noninvasive visualization and interrogation of biological processes in living animals","volume":"11","author":"Close","year":"2011","journal-title":"Sensors (Basel)"},{"key":"10.1016\/j.jbc.2026.113568_bib3","doi-asserted-by":"crossref","first-page":"453","DOI":"10.1177\/1087057112465184","article-title":"Affordable luciferase reporter assay for cell-based high-throughput screening","volume":"18","author":"Siebring-van Olst","year":"2013","journal-title":"J. Biomol. Screen"},{"key":"10.1016\/j.jbc.2026.113568_bib4","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1007\/978-1-4939-3673-1_14","article-title":"A luciferase reporter gene system for high-throughput screening of \u03b3-Globin gene activators","volume":"1439","author":"Xie","year":"2016","journal-title":"Methods Mol. Biol."},{"key":"10.1016\/j.jbc.2026.113568_bib5","doi-asserted-by":"crossref","first-page":"2351","DOI":"10.1021\/jacs.6b11737","article-title":"Orthogonal luciferase-luciferin pairs for bioluminescence imaging","volume":"139","author":"Jones","year":"2017","journal-title":"J. Am. Chem. Soc."},{"key":"10.1016\/j.jbc.2026.113568_bib6","doi-asserted-by":"crossref","first-page":"640","DOI":"10.1016\/j.tibtech.2017.03.012","article-title":"In vivo molecular bioluminescence imaging","volume":"35","author":"Mezzanotte","year":"2017","journal-title":"New Tools Appl. Trends Biotechnol."},{"key":"10.1016\/j.jbc.2026.113568_bib7","doi-asserted-by":"crossref","DOI":"10.1016\/j.ejmech.2020.113111","article-title":"Recent achievements of bioluminescence imaging based on firefly luciferin-luciferase system","volume":"211","author":"Li","year":"2021","journal-title":"Eur. J. Med. Chem."},{"key":"10.1016\/j.jbc.2026.113568_bib8","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1016\/j.tibtech.2004.03.011","article-title":"Biotechnological applications of bioluminescence and chemiluminescence","volume":"22","author":"Roda","year":"2004","journal-title":"Trends Biotechnol."},{"key":"10.1016\/j.jbc.2026.113568_bib9","doi-asserted-by":"crossref","first-page":"646","DOI":"10.1016\/j.chembiol.2010.05.012","article-title":"Illuminating insights into firefly luciferase and other bioluminescent reporters used in chemical biology","volume":"17","author":"Thorne","year":"2010","journal-title":"Chem. Biol."},{"key":"10.1016\/j.jbc.2026.113568_bib10","doi-asserted-by":"crossref","first-page":"835","DOI":"10.1517\/17460441.2010.506213","article-title":"Bioluminescence assays: multicolor luciferase assay, secreted luciferase assay and imaging luciferase assay","volume":"5","author":"Nakajima","year":"2010","journal-title":"Expert Opin. Drug Discov."},{"key":"10.1016\/j.jbc.2026.113568_bib11","first-page":"127","article-title":"Bioluminescent assays for high-throughput screening ASSAY and drug","volume":"5","author":"Fan","year":"2007","journal-title":"Dev. Tech."},{"key":"10.1016\/j.jbc.2026.113568_bib12","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0142736","article-title":"Establishment of green fluorescent protein and firefly luciferase expressing mouse primary macrophages for in vivo bioluminescence imaging","volume":"10","author":"Pajarinen","year":"2015","journal-title":"PLoS One"},{"key":"10.1016\/j.jbc.2026.113568_bib13","doi-asserted-by":"crossref","first-page":"2707","DOI":"10.1021\/acschembio.1c00549","article-title":"Brightening up biology: advances in luciferase systems for in vivo imaging","volume":"16","author":"Liu","year":"2021","journal-title":"ACS Chem. Biol."},{"key":"10.1016\/j.jbc.2026.113568_bib14","doi-asserted-by":"crossref","first-page":"796","DOI":"10.1007\/s00259-021-05439-4","article-title":"Firefly luciferase offers superior performance to AkaLuc for tracking the fate of administered cell therapies","volume":"49","author":"Amadeo","year":"2022","journal-title":"Eur. J. Nucl. Med. Mol. Imaging"},{"key":"10.1016\/j.jbc.2026.113568_bib15","doi-asserted-by":"crossref","first-page":"750","DOI":"10.1109\/JPROC.2005.844261","article-title":"Using in vivo bioluminescence imaging to shed light on","volume":"93","author":"Thorne","year":"2005","journal-title":"Cancer Biol. Proc. IEEE"},{"key":"10.1016\/j.jbc.2026.113568_bib16","doi-asserted-by":"crossref","first-page":"67","DOI":"10.1039\/D5CB00287G","article-title":"Red-shifted d-luciferin analogues and their bioluminescence characteristics","volume":"7","author":"Watthaisong","year":"2026","journal-title":"RSC Chem. Biol."},{"key":"10.1016\/j.jbc.2026.113568_bib17","doi-asserted-by":"crossref","first-page":"7592","DOI":"10.1021\/jacs.5b03820","article-title":"Experimental support for a single electron-transfer oxidation mechanism in firefly bioluminescence","volume":"137","author":"Branchini","year":"2015","journal-title":"J. Am. Chem. Soc."},{"key":"10.1016\/j.jbc.2026.113568_bib18","doi-asserted-by":"crossref","first-page":"1085","DOI":"10.1039\/b809935a","article-title":"Kinetics of inhibition of firefly luciferase by oxyluciferin and dehydroluciferyl-adenylate","volume":"7","author":"Ribeiro","year":"2008","journal-title":"Photochem. Photobiol. Sci."},{"key":"10.1016\/j.jbc.2026.113568_bib19","doi-asserted-by":"crossref","first-page":"1039","DOI":"10.1039\/c0pp00379d","article-title":"Kinetics of inhibition of firefly luciferase by dehydroluciferyl-coenzyme A, dehydroluciferin and L-luciferin","volume":"10","author":"da Silva","year":"2011","journal-title":"Photochem. Photobiol. Sci."},{"key":"10.1016\/j.jbc.2026.113568_bib20","doi-asserted-by":"crossref","first-page":"387","DOI":"10.1007\/s00018-009-0170-8","article-title":"Firefly luciferase: an adenylate-forming enzyme for multicatalytic functions","volume":"67","author":"Inouye","year":"2010","journal-title":"Cell Mol. Life Sci."},{"key":"10.1016\/j.jbc.2026.113568_bib21","doi-asserted-by":"crossref","first-page":"110","DOI":"10.1002\/cbic.200300735","article-title":"Identification of luciferyl adenylate and luciferyl coenzyme a synthesized by firefly luciferase","volume":"5","author":"Fraga","year":"2004","journal-title":"Chembiochem"},{"key":"10.1016\/j.jbc.2026.113568_bib22","doi-asserted-by":"crossref","first-page":"146","DOI":"10.1039\/b719181b","article-title":"Firefly luminescence: a historical perspective and recent developments photochem","volume":"7","author":"Fraga","year":"2008","journal-title":"Photobiol. Sci."},{"key":"10.1016\/j.jbc.2026.113568_bib23","doi-asserted-by":"crossref","first-page":"5206","DOI":"10.1111\/j.1742-4658.2005.04895.x","article-title":"Coenzyme A affects firefly luciferase luminescence because it acts as a substrate and not as an allosteric effector","volume":"272","author":"Fraga","year":"2005","journal-title":"FEBS J."},{"key":"10.1016\/j.jbc.2026.113568_bib24","doi-asserted-by":"crossref","first-page":"1528","DOI":"10.1016\/S0021-9258(18)49367-2","article-title":"The synthesis and function of luciferyl-adenylate and oxyluciferyl-adenylate","volume":"233","author":"Rhodes","year":"1958","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.jbc.2026.113568_bib25","doi-asserted-by":"crossref","first-page":"445","DOI":"10.1006\/bbrc.1997.7161","article-title":"Synthesis of dehydroluciferin by firefly luciferase: effect of dehydroluciferin, coenzyme A and nucleoside triphosphates on the luminescent reaction","volume":"237","author":"Fontes","year":"1997","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"10.1016\/j.jbc.2026.113568_bib26","doi-asserted-by":"crossref","first-page":"2161","DOI":"10.1007\/s00216-008-2117-6","article-title":"An optimized luciferase bioluminescent assay for coenzyme A","volume":"391","author":"Marques","year":"2008","journal-title":"Anal. Bioanal. Chem."},{"key":"10.1016\/j.jbc.2026.113568_bib27","doi-asserted-by":"crossref","first-page":"929","DOI":"10.1002\/cbic.200500443","article-title":"Firefly luciferase produces hydrogen peroxide as a coproduct in dehydroluciferyl adenylate formation","volume":"7","author":"Fraga","year":"2016","journal-title":"Chembiochem"},{"key":"10.1016\/j.jbc.2026.113568_bib28","doi-asserted-by":"crossref","first-page":"2156","DOI":"10.1002\/elps.201500031","article-title":"Stability of energy metabolites\u2014An often overlooked issue in metabolomics studies: a review","volume":"36","author":"Gil","year":"2015","journal-title":"Electrophoresis"},{"key":"10.1016\/j.jbc.2026.113568_bib29","doi-asserted-by":"crossref","first-page":"324","DOI":"10.1162\/1535350042973553","article-title":"Thermostability of firefly luciferases affects efficiency of detection by in vivo bioluminescence","volume":"3","author":"Baggett","year":"2004","journal-title":"Mol. Imaging"},{"key":"10.1016\/j.jbc.2026.113568_bib30","doi-asserted-by":"crossref","first-page":"115","DOI":"10.1016\/S0003-2670(01)01496-9","article-title":"Development of a thermostable firefly luciferase","volume":"457","author":"Tisi","year":"2002","journal-title":"Anal. Chim. Acta"},{"key":"10.1016\/j.jbc.2026.113568_bib31","doi-asserted-by":"crossref","first-page":"2628","DOI":"10.1021\/acsomega.7b02068","article-title":"Firefly luciferase mutant with enhanced activity and thermostability","volume":"3","author":"Pozzo","year":"2018","journal-title":"ACS Omega"},{"key":"10.1016\/j.jbc.2026.113568_bib32","doi-asserted-by":"crossref","first-page":"305","DOI":"10.1042\/BJ20051847","article-title":"Mutagenesis of solvent-exposed amino acids in Photinus pyralis luciferase improves thermostability and pH-tolerance","volume":"397","author":"Law","year":"2006","journal-title":"Biochem. J."},{"key":"10.1016\/j.jbc.2026.113568_bib33","first-page":"119","article-title":"Development of a pH-tolerant Thermostable Photinus Pyralis Luciferase for Brighter in Vivo Imaging Bioluminescence-recent Advances in Oceanic Measurements and Laboratory Applications","author":"Jathoul","year":"2012","journal-title":"IntechOpen"},{"key":"10.1016\/j.jbc.2026.113568_bib34","doi-asserted-by":"crossref","first-page":"203","DOI":"10.2144\/05392ST02","article-title":"Comparison of the responsiveness of the pGL3 and pGL4 luciferase reporter vectors to ster hormones","volume":"39","author":"Dougherty","year":"2005","journal-title":"Biotechniques"},{"key":"10.1016\/j.jbc.2026.113568_bib35","doi-asserted-by":"crossref","first-page":"127","DOI":"10.4161\/epi.1.3.3327","article-title":"Functional analysis of promoter CPG-Methylation using a CpG-Free luciferase reporter vector","volume":"1","author":"Klug","year":"2006","journal-title":"Epigenetics"},{"key":"10.1016\/j.jbc.2026.113568_bib36","doi-asserted-by":"crossref","first-page":"104639","DOI":"10.1016\/j.jbc.2023.104639","article-title":"High sensitivity and low-cost flavin luciferase (FLUXVc)-based reporter gene for mammalian cell expression","volume":"299","author":"Phonbuppha","year":"2023","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.jbc.2026.113568_bib37","doi-asserted-by":"crossref","first-page":"534","DOI":"10.1038\/s41929-022-00800-8","article-title":"Mechanism-guided tunnel engineering to increase the efficiency of a flavin-dependent halogenase","volume":"5","author":"Prakinee","year":"2022","journal-title":"Nat. Catal."},{"key":"10.1016\/j.jbc.2026.113568_bib38","first-page":"2614","article-title":"Rational and Mechanistic Approaches for Improving Biocatalyst Performance Chem Catalysis","volume":"2","author":"Phintha","year":"2022","journal-title":"Chem Catal"},{"key":"10.1016\/j.jbc.2026.113568_bib39","doi-asserted-by":"crossref","DOI":"10.1039\/D4CB00102H","article-title":"Enhancement of tryptophan 2-monooxygenase thermostability by semi-rational enzyme engineering: a strategic design to minimize experimental investigation","author":"Kongjaroon","year":"2024","journal-title":"RSC Chem. Biol."},{"key":"10.1016\/j.jbc.2026.113568_bib40","doi-asserted-by":"crossref","first-page":"1481","DOI":"10.1002\/cbic.201900737","article-title":"Creating flavin reductase variants with thermostable and solvent-tolerant properties by rational-design engineering","volume":"21","author":"Maenpuen","year":"2020","journal-title":"ChemBioChem"},{"key":"10.1016\/j.jbc.2026.113568_bib41","series-title":"Rational-Design Engineering to Improve Enzyme Thermostability in Enzyme Engineering: Methods and Protocols","first-page":"159","author":"Pongsupasa","year":"2022"},{"key":"10.1016\/j.jbc.2026.113568_bib42","doi-asserted-by":"crossref","first-page":"3020","DOI":"10.1002\/cbic.201900413","article-title":"Identification of a hotspot residue for improving the thermostability of a flavin-dependent monooxygenase","volume":"20","author":"Pongpamorn","year":"2019","journal-title":"ChemBioChem"},{"key":"10.1016\/j.jbc.2026.113568_bib43","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1080\/07388551.2019.1682963","article-title":"Recent advances in the improvement of enzyme thermostability by structure modification","volume":"40","author":"Xu","year":"2020","journal-title":"Crit. Rev. Biotechnol."},{"key":"10.1016\/j.jbc.2026.113568_bib44","article-title":"Luciferin synthesis and pesticide detection by luminescence enzymatic cascades","volume":"61","author":"Watthaisong","year":"2022","journal-title":"Angew. Chem. Int. Ed."},{"key":"10.1016\/j.jbc.2026.113568_bib45","doi-asserted-by":"crossref","first-page":"13388","DOI":"10.1002\/ange.201904923","article-title":"A chemo-enzymatic cascade for the smart detection of nitro-and halogenated phenols","volume":"131","author":"Watthaisong","year":"2019","journal-title":"Angew. Chem."},{"key":"10.1016\/j.jbc.2026.113568_bib46","doi-asserted-by":"crossref","first-page":"549","DOI":"10.1016\/j.molmed.2004.09.003","article-title":"Nrf2\u2013Keap1 defines a physiologically important stress response mechanism","volume":"10","author":"Motohashi","year":"2004","journal-title":"Trends Mol. Med."},{"key":"10.1016\/j.jbc.2026.113568_bib47","doi-asserted-by":"crossref","first-page":"7130","DOI":"10.1128\/MCB.24.16.7130-7139.2004","article-title":"Oxidative stress sensor Keap1 functions as an adaptor for Cul3-based E3 ligase to regulate proteasomal degradation of Nrf2","volume":"24","author":"Kobayashi","year":"2004","journal-title":"Mol. Cell. Biol."},{"key":"10.1016\/j.jbc.2026.113568_bib48","doi-asserted-by":"crossref","first-page":"1304","DOI":"10.1016\/j.freeradbiomed.2009.07.035","article-title":"Nrf2: INrf2 (Keap1) signaling in oxidative stress","volume":"47","author":"Kaspar","year":"2009","journal-title":"Free Radic. Biol. Med."},{"key":"10.1016\/j.jbc.2026.113568_bib49","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1004556","article-title":"FireProt: energy- and evolution-based computational design of thermostable multiple-point mutants","volume":"11","author":"Bednar","year":"2015","journal-title":"PLoS Comput. Biol."},{"key":"10.1016\/j.jbc.2026.113568_bib50","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1021\/ci700237g","article-title":"Thermostability of salt bridges versus hydrophobic interactions in proteins probed by statistical potentials","volume":"48","author":"Folch","year":"2008","journal-title":"J. Chem. Inf. Model."},{"key":"10.1016\/j.jbc.2026.113568_bib51","doi-asserted-by":"crossref","first-page":"220","DOI":"10.1016\/j.cbpa.2007.01.685","article-title":"Stability of biocatalysts","volume":"11","author":"Polizzi","year":"2007","journal-title":"Curr. Opin. Chem. Biol."},{"key":"10.1016\/j.jbc.2026.113568_bib52","doi-asserted-by":"crossref","first-page":"1325","DOI":"10.1023\/A:1025771421906","article-title":"Physical stability of proteins in aqueous solution: mechanism and driving forces in nonnative protein aggregation","volume":"20","author":"Chi","year":"2003","journal-title":"Pharm. Res."},{"key":"10.1016\/j.jbc.2026.113568_bib53","article-title":"Immobilization of firefly bioluminescent System: development and application of reagents","volume":"13","author":"Esimbekova","year":"2022","journal-title":"Biosensors (Basel)"},{"key":"10.1016\/j.jbc.2026.113568_bib54","doi-asserted-by":"crossref","first-page":"3427","DOI":"10.1002\/anie.200462934","article-title":"Synthesis of Luciferyl coenzyme A: a bioluminescent substrate for firefly luciferase in the presence of AMP","volume":"44","author":"Fraga","year":"2005","journal-title":"Angew. Chem. Int. Ed."},{"key":"10.1016\/j.jbc.2026.113568_bib55","doi-asserted-by":"crossref","first-page":"180","DOI":"10.1016\/0167-4838(95)00150-S","article-title":"Does the sulfhydryl or the adenine moiety of CoA enhance firefly luciferase activity?","volume":"1252","author":"Ford","year":"1995","journal-title":"Biochim. Biophys. Acta"},{"key":"10.1016\/j.jbc.2026.113568_bib56","doi-asserted-by":"crossref","DOI":"10.3389\/fchem.2022.1018124","article-title":"Use of whole-cell bioreporters to assess bioavailability of contaminants in aquatic systems","volume":"10","author":"Zhu","year":"2022","journal-title":"Front. Chem."},{"key":"10.1016\/j.jbc.2026.113568_bib57","doi-asserted-by":"crossref","first-page":"3234","DOI":"10.1038\/s41598-020-60182-4","article-title":"A universal reporter cell line for bioactivity evaluation of engineered cytokine products","volume":"10","author":"Mock","year":"2020","journal-title":"Sci. Rep."},{"key":"10.1016\/j.jbc.2026.113568_bib58","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1186\/s40643-025-00932-2","article-title":"Advancements in the application of reporter gene cell lines in bioactivity evaluation of biological products","volume":"12","author":"Yi","year":"2025","journal-title":"Bioresour. Bioproc."},{"key":"10.1016\/j.jbc.2026.113568_bib59","doi-asserted-by":"crossref","DOI":"10.3389\/fphys.2016.00300","article-title":"A guide to transient expression of membrane proteins in hek-293 cells for functional characterization","volume":"7","author":"Ooi","year":"2016","journal-title":"Front. Physiol."},{"key":"10.1016\/j.jbc.2026.113568_bib60","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1016\/j.jbiotec.2022.01.004","article-title":"Statistical experimental designs to optimize the transient transfection of HEK 293T cells and determine a transfer criterion from adherent cells to larger-scale cell suspension cultures","volume":"346","author":"Dekevic","year":"2022","journal-title":"J. Biotechnol."},{"key":"10.1016\/j.jbc.2026.113568_bib61","doi-asserted-by":"crossref","DOI":"10.7717\/peerj.11165","article-title":"Transfection types, methods and strategies: a technical review","volume":"9","author":"Chong","year":"2021","journal-title":"PeerJ"},{"key":"10.1016\/j.jbc.2026.113568_bib62","series-title":"Protocols in in Vitro Hepatocyte Research","first-page":"77","article-title":"Culture and functional characterization of human hepatoma HepG2 cells","author":"Donato","year":"2015"},{"key":"10.1016\/j.jbc.2026.113568_bib63","first-page":"372","article-title":"Optimizing a lipocomplex-based gene transfer method into hela cell line","volume":"15","author":"Asgharian","year":"2014","journal-title":"Cell J."},{"key":"10.1016\/j.jbc.2026.113568_bib64","doi-asserted-by":"crossref","DOI":"10.1038\/mtna.2016.8","article-title":"Lipid-based transfection reagents exhibit cryo-induced increase in transfection efficiency","volume":"5","author":"Sork","year":"2016","journal-title":"Mol. Ther. Nucleic Acids"},{"key":"10.1016\/j.jbc.2026.113568_bib65","doi-asserted-by":"crossref","DOI":"10.3389\/fbioe.2021.796991","article-title":"HEK293 cell line as a platform to produce recombinant proteins and viral vectors","volume":"9","author":"Tan","year":"2021","journal-title":"Front. Bioeng. Biotechnol."},{"key":"10.1016\/j.jbc.2026.113568_bib66","doi-asserted-by":"crossref","DOI":"10.1155\/2019\/8592348","article-title":"Potential applications of NRF2 inhibitors in cancer therapy","volume":"2019","author":"Panieri","year":"2019","journal-title":"Oxid Med. Cell Longev."},{"key":"10.1016\/j.jbc.2026.113568_bib67","doi-asserted-by":"crossref","DOI":"10.1038\/cddis.2016.389","article-title":"Therapeutic advantage of Pro-electrophilic drugs to activate the Nrf2\/ARE pathway in Alzheimer\u2019s disease models","volume":"7","author":"Lipton","year":"2016","journal-title":"Cell Death Dis."},{"key":"10.1016\/j.jbc.2026.113568_bib68","doi-asserted-by":"crossref","first-page":"1116","DOI":"10.1111\/j.1471-4159.2007.05039.x","article-title":"Carnosic acid, a catechol-type electrophilic compound, protects neurons both in vitro and in vivo through activation of the Keap1\/Nrf2 pathway via S-alkylation of targeted cysteines on Keap1","volume":"104","author":"Satoh","year":"2008","journal-title":"J. Neurochem."},{"key":"10.1016\/j.jbc.2026.113568_bib69","doi-asserted-by":"crossref","first-page":"103","DOI":"10.1016\/j.expneurol.2014.11.008","article-title":"Nrf2-ARE activator carnosic acid decreases mitochondrial dysfunction, oxidative damage and neuronal cytoskeletal degradation following traumatic brain injury in mice","volume":"264","author":"Miller","year":"2015","journal-title":"Exp. Neurol."},{"key":"10.1016\/j.jbc.2026.113568_bib70","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.freeradbiomed.2012.12.011","article-title":"Administration of the Nrf2\u2013ARE activators sulforaphane and carnosic acid attenuates 4-hydroxy-2-nonenal-induced mitochondrial dysfunction ex vivo","volume":"57","author":"Miller","year":"2013","journal-title":"Free Radic. Biol. Med."},{"key":"10.1016\/j.jbc.2026.113568_bib71","doi-asserted-by":"crossref","first-page":"1848","DOI":"10.1021\/cb3002478","article-title":"Engineered luciferase reporter from a deep Sea shrimp utilizing a novel imidazopyrazinone substrate","volume":"7","author":"Hall","year":"2012","journal-title":"ACS Chem. Biol."},{"key":"10.1016\/j.jbc.2026.113568_bib72","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1149\/05029.0001ecst","article-title":"Evaluation of bioluminescence activity of firefly luciferin nucleotide","volume":"50","author":"Iwano","year":"2013","journal-title":"Derivatives ECS Trans."},{"key":"10.1016\/j.jbc.2026.113568_bib73","doi-asserted-by":"crossref","first-page":"1046","DOI":"10.1111\/j.1751-1097.2010.00777.x","article-title":"Quantum yields and kinetics of the firefly bioluminescence reaction of beetle luciferases","volume":"86","author":"Niwa","year":"2010","journal-title":"Photochem. Photobiol."},{"key":"10.1016\/j.jbc.2026.113568_bib74","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1246\/cl.2010.291","article-title":"Quantum yield measurements of firefly bioluminescence reactions using a commercial luminometer","volume":"39","author":"Niwa","year":"2010","journal-title":"Chem. Lett."},{"key":"10.1016\/j.jbc.2026.113568_bib75","doi-asserted-by":"crossref","first-page":"44","DOI":"10.1038\/nphoton.2007.251","article-title":"Firefly bioluminescence quantum yield and colour change by pH-sensitive green emission","volume":"2","author":"Ando","year":"2008","journal-title":"Nat. Photon."},{"key":"10.1016\/j.jbc.2026.113568_bib76","doi-asserted-by":"crossref","first-page":"372","DOI":"10.1038\/nature04542","article-title":"Structural basis for the spectral difference in luciferase bioluminescence","volume":"440","author":"Nakatsu","year":"2006","journal-title":"Nature"},{"key":"10.1016\/j.jbc.2026.113568_bib77","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0010011","article-title":"Enhanced beetle luciferase for high-resolution bioluminescence imaging","volume":"5","author":"Nakajima","year":"2010","journal-title":"PLoS One"},{"key":"10.1016\/j.jbc.2026.113568_bib78","doi-asserted-by":"crossref","first-page":"373","DOI":"10.1186\/s13104-024-07038-6","article-title":"Nrf2 induction potency of plant-derived compounds determined using an antioxidant response element luciferase reporter and conventional NAD(P)H-quinone acceptor oxidoreductase 1 activity assay","volume":"17","author":"Tamaru","year":"2024","journal-title":"BMC Res. Notes"},{"key":"10.1016\/j.jbc.2026.113568_bib79","doi-asserted-by":"crossref","first-page":"W393","DOI":"10.1093\/nar\/gkx285","article-title":"FireProt: web server for automated design of thermostable proteins","volume":"45","author":"Musil","year":"2017","journal-title":"Nucleic Acids Res."},{"key":"10.1016\/j.jbc.2026.113568_bib80","doi-asserted-by":"crossref","DOI":"10.1093\/bib\/bbad425","article-title":"FireProt 2.0: web-based platform for the fully automated design of thermostable proteins","volume":"25","author":"Musil","year":"2023","journal-title":"Brief. Bioinform."},{"key":"10.1016\/j.jbc.2026.113568_bib81","series-title":"Rational and Semirational Protein Design in Protein Engineering: Methods and Protocols","first-page":"15","author":"Korendovych","year":"2018"},{"key":"10.1016\/j.jbc.2026.113568_bib82","doi-asserted-by":"crossref","first-page":"70","DOI":"10.1016\/j.enzmictec.2013.02.012","article-title":"Library construction and evaluation for site saturation mutagenesis","volume":"53","author":"Sullivan","year":"2013","journal-title":"Enzyme Microb. Technol."},{"key":"10.1016\/j.jbc.2026.113568_bib83","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1006\/abio.1996.0319","article-title":"SYPRO Orange and SYPRO red protein Gel stains: One-Step fluorescent staining of denaturing gels for detection of nanogram levels of protein","volume":"239","author":"Steinberg","year":"1996","journal-title":"Anal. Biochem."},{"key":"10.1016\/j.jbc.2026.113568_bib84","doi-asserted-by":"crossref","first-page":"28.9.1","DOI":"10.1002\/0471140864.ps2809s79","article-title":"Analysis of protein stability and ligand interactions by thermal shift assay","volume":"79","author":"Huynh","year":"2015","journal-title":"Curr. Protoc. Protein Sci."},{"key":"10.1016\/j.jbc.2026.113568_bib85","doi-asserted-by":"crossref","first-page":"366","DOI":"10.1038\/s41598-021-04223-6","article-title":"The N-terminal tail of the hydrophobin SC16 is not required for rodlet formation","volume":"12","author":"Vergunst","year":"2022","journal-title":"Sci. Rep."},{"key":"10.1016\/j.jbc.2026.113568_bib86","doi-asserted-by":"crossref","first-page":"6493","DOI":"10.1021\/bi300934s","article-title":"Crystal structure of firefly luciferase in a second catalytic conformation supports a domain alternation mechanism","volume":"51","author":"Sundlov","year":"2012","journal-title":"Biochemistry"},{"key":"10.1016\/j.jbc.2026.113568_bib87","doi-asserted-by":"crossref","first-page":"1859","DOI":"10.1002\/jcc.20945","article-title":"CHARMM-GUI: a web-based graphical user interface for CHARMM","volume":"29","author":"Jo","year":"2008","journal-title":"J. Comput. Chem."},{"key":"10.1016\/j.jbc.2026.113568_bib88","doi-asserted-by":"crossref","DOI":"10.1016\/j.jmb.2023.167995","article-title":"CHARMM-GUI PDB manipulator: various PDB structural modifications for biomolecular modeling and simulation","volume":"435","author":"Park","year":"2023","journal-title":"J. Mol. Biol."},{"key":"10.1016\/j.jbc.2026.113568_bib89","doi-asserted-by":"crossref","first-page":"129","DOI":"10.1002\/pro.4200","article-title":"PDB-101: Educational resources supporting molecular explorations through biology and medicine","volume":"31","author":"Zardecki","year":"2022","journal-title":"Protein Sci."},{"key":"10.1016\/j.jbc.2026.113568_bib90","doi-asserted-by":"crossref","first-page":"609","DOI":"10.1002\/prot.10465","article-title":"Improved protein-ligand docking using GOLD","volume":"52","author":"Verdonk","year":"2003","journal-title":"Proteins"},{"key":"10.1016\/j.jbc.2026.113568_bib91","doi-asserted-by":"crossref","first-page":"727","DOI":"10.1006\/jmbi.1996.0897","article-title":"Development and validation of a genetic algorithm for flexible docking","volume":"267","author":"Jones","year":"1997","journal-title":"J. Mol. Biol."}],"container-title":["Journal of Biological Chemistry"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0021925826024403?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0021925826024403?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,10,21]],"date-time":"2026-10-21T13:38:18Z","timestamp":1792589898000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0021925826024403"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,11]]},"references-count":91,"journal-issue":{"issue":"11","published-print":{"date-parts":[[2026,11]]}},"alternative-id":["S0021925826024403"],"URL":"https:\/\/doi.org\/10.1016\/j.jbc.2026.113568","relation":{},"ISSN":["0021-9258"],"issn-type":[{"value":"0021-9258","type":"print"}],"subject":[],"published":{"date-parts":[[2026,11]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Biochemical, bioluminescence and bioreporter properties of functionally improved firefly luciferases","name":"articletitle","label":"Article Title"},{"value":"Journal of Biological Chemistry","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.jbc.2026.113568","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2026 The Authors. Published by Elsevier Inc on behalf of American Society for Biochemistry and Molecular Biology.","name":"copyright","label":"Copyright"}],"article-number":"113568"}}