{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,3]],"date-time":"2026-09-03T12:12:17Z","timestamp":1788437537349,"version":"build-2803163510"},"reference-count":60,"publisher":"American Chemical Society (ACS)","issue":"34","license":[{"start":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T00:00:00Z","timestamp":1787011200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-029"},{"start":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T00:00:00Z","timestamp":1787011200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-037"},{"start":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T00:00:00Z","timestamp":1787011200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-045"}],"funder":[{"DOI":"10.13039\/100031919","name":"Southern Marine Science and Engineering Guangdong Laboratory","doi-asserted-by":"publisher","award":["SML2024SP024"],"award-info":[{"award-number":["SML2024SP024"]}],"id":[{"id":"10.13039\/100031919","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100014219","name":"National Science Fund for Distinguished Young Scholars","doi-asserted-by":"publisher","award":["52425001"],"award-info":[{"award-number":["52425001"]}],"id":[{"id":"10.13039\/501100014219","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/na","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["U23A2049"],"award-info":[{"award-number":["U23A2049"]}],"id":[{"id":"10.13039\/na","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,9,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Biogenic hydrogen sulfide produced in sewer systems causes odor nuisance and concrete corrosion, necessitating effective biocidal control. Free nitrous acid (FNA) has emerged as a promising biocide, but its unclear mechanisms complicate dosage optimization and risk assessment. Here, using Desulfovibrio vulgaris as a model lysogenic bacterium, we demonstrate that low-dose FNA (0.2\u20134.0 mg N\/L) induces bacterial inactivation via prophage activation-associated lysis in addition to chemical oxidation. Reactive nitrogen species (RNS) scavenging tests revealed that RNS-mediated oxidative stress was closely associated with prophage activation. Activated phages further infected new hosts, reducing the viability of freshly cultured D. vulgaris cells by 25.7% and increasing total phage production 15.2-fold. The phenomenon was further validated in real sewage biofilms, where increased phage production and decreased bacterial viability were observed at a low FNA dose of 0.2 mg N\/L, below the dose required for cell destruction by chemical oxidation. Furthermore, metagenomic analysis of 896 sewage samples worldwide revealed that 71.9% of recovered sewage-derived microbial genomes harbor prophages, indicating a widespread genomic basis for prophage activation-mediated bacterial inactivation. Overall, this study expands current understanding of the biocidal mechanisms of FNA and contributes to the development of environmentally sustainable biocidal strategies.<\/jats:p>","DOI":"10.1021\/acs.est.6c01580","type":"journal-article","created":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T08:02:13Z","timestamp":1787040133000},"page":"24165-24176","source":"Crossref","is-referenced-by-count":0,"title":["Prophage\nActivation as an Overlooked Mechanism Underlying\nthe Biocidal Effect of Free Nitrous Acid in Sewers"],"prefix":"10.1021","volume":"60","author":[{"given":"Yuming","family":"Zhen","sequence":"first","affiliation":[{"name":"Sun Yat-sen University , , ,","place":["Guangzhou, China, 510275"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Juntao","family":"Xia","sequence":"additional","affiliation":[{"name":"Sun Yat-sen University , , ,","place":["Guangzhou, China, 510275"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yan-Ying","family":"Qiu","sequence":"additional","affiliation":[{"name":"Sun Yat-sen University , , ,","place":["Guangzhou, China, 510275"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jiahua","family":"Guo","sequence":"additional","affiliation":[{"name":"Sun Yat-sen University , , ,","place":["Guangzhou, China, 510275"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8198-7454","authenticated-orcid":true,"given":"Feng","family":"Jiang","sequence":"additional","affiliation":[{"name":"Sun Yat-sen University , , ,","place":["Guangzhou, China, 510275"]},{"name":"Sun Yat-sen University , , ,","place":["Guangzhou, China, 510275"]},{"name":"Southern Marine Science and Engineering Guangdong Laboratory (Zhuhai) , ,","place":["Zhuhai, China, 519000"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,8,18]]},"reference":[{"issue":"16","key":"2026090307124513300_cit1","doi-asserted-by":"publisher","first-page":"11625","DOI":"10.1021\/acs.est.2c01156","article-title":"Understanding the Effect\nof Free Nitrous Acid on Biofilms","volume":"56","author":"Chislett","year":"2022","journal-title":"Environ. Sci.\nTechnol."},{"issue":"12","key":"2026090307124513300_cit2","doi-asserted-by":"publisher","first-page":"3735","DOI":"10.1016\/j.watres.2011.04.026","article-title":"The strong biocidal effect of free\nnitrous acid on anaerobic sewer biofilms","volume":"45","author":"Jiang","year":"2011","journal-title":"Water\nRes."},{"issue":"13","key":"2026090307124513300_cit3","doi-asserted-by":"publisher","first-page":"4331","DOI":"10.1016\/j.watres.2013.05.024","article-title":"Dosing free nitrous acid for sulfide\ncontrol in sewers: results of field trials in Australia","volume":"47","author":"Jiang","year":"2013","journal-title":"Water Res."},{"issue":"16","key":"2026090307124513300_cit4","doi-asserted-by":"publisher","first-page":"10301","DOI":"10.1021\/acs.est.0c01453","article-title":"Structural Changes\nin Cell-Wall and Cell-Membrane Organic Materials Following Exposure\nto Free Nitrous Acid","volume":"54","author":"Chislett","year":"2020","journal-title":"Environ. Sci. Technol."},{"issue":"20","key":"2026090307124513300_cit5","doi-asserted-by":"publisher","first-page":"11897","DOI":"10.1021\/es402933b","article-title":"Free nitrous\nacid (FNA)-based pretreatment enhances methane production from waste\nactivated sludge","volume":"47","author":"Wang","year":"2013","journal-title":"Environ. Sci. Technol."},{"key":"2026090307124513300_cit6","doi-asserted-by":"publisher","first-page":"115382","DOI":"10.1016\/j.watres.2019.115382","article-title":"Improving wastewater management using free nitrous\nacid (FNA)","volume":"171","author":"Duan","year":"2020","journal-title":"Water Res."},{"key":"2026090307124513300_cit7","doi-asserted-by":"publisher","first-page":"118401","DOI":"10.1016\/j.watres.2022.118401","article-title":"Reactive nitrogen\nspecies from free nitrous acid (FNA) cause cell lysis","volume":"217","author":"Chislett","year":"2022","journal-title":"Water Res."},{"issue":"10","key":"2026090307124513300_cit8","doi-asserted-by":"publisher","first-page":"5305","DOI":"10.1021\/acs.est.6b00288","article-title":"Determining\nMultiple Responses of\nPseudomonas aeruginosa PAO1 to an Antimicrobial Agent, Free Nitrous\nAcid","volume":"50","author":"Gao","year":"2016","journal-title":"Environ. Sci. Technol."},{"issue":"17","key":"2026090307124513300_cit9","doi-asserted-by":"publisher","first-page":"12542","DOI":"10.1021\/acs.est.2c01136","article-title":"Reactive Nitrogen\nSpecies Mediated Inactivation of Pathogenic Microorganisms during\nUVA Photolysis of Nitrite at Surface Water Levels","volume":"56","author":"Wang","year":"2022","journal-title":"Environ. Sci. Technol."},{"issue":"51","key":"2026090307124513300_cit10","doi-asserted-by":"publisher","first-page":"22480","DOI":"10.1021\/acs.est.4c03086","article-title":"Formation and Fate of Reactive Nitrogen\nduring Biological Nitrogen Removal from Water: Important Yet Often\nIgnored Chemical Aspects of the Nitrogen Cycle","volume":"58","author":"Su","year":"2024","journal-title":"Environ. Sci. Technol."},{"key":"2026090307124513300_cit11","doi-asserted-by":"publisher","first-page":"116553","DOI":"10.1016\/j.watres.2020.116553","article-title":"Structural changes in model compounds\nof sludge extracellular polymeric substances caused by exposure to\nfree nitrous acid","volume":"188","author":"Chislett","year":"2021","journal-title":"Water Res."},{"issue":"5","key":"2026090307124513300_cit12","doi-asserted-by":"publisher","first-page":"2305","DOI":"10.1007\/s00253-014-6211-8","article-title":"The concentration-determined\nand\npopulation-specific antimicrobial effects of free nitrous acid on\nPseudomonas aeruginosa PAO1","volume":"99","author":"Gao","year":"2015","journal-title":"Appl. Microbiol.\nBiotechnol."},{"issue":"4","key":"2026090307124513300_cit13","doi-asserted-by":"publisher","first-page":"1937","DOI":"10.1021\/acs.est.8b06148","article-title":"Overcoming Nitrite Oxidizing Bacteria\nAdaptation through Alternating Sludge Treatment with Free Nitrous\nAcid and Free Ammonia","volume":"53","author":"Duan","year":"2019","journal-title":"Environ. Sci. Technol."},{"key":"2026090307124513300_cit14","doi-asserted-by":"publisher","first-page":"119183","DOI":"10.1016\/j.watres.2022.119183","article-title":"Phage-host interactions: The neglected\npart of biological wastewater treatment","volume":"226","author":"Tang","year":"2022","journal-title":"Water\nRes."},{"issue":"10","key":"2026090307124513300_cit15","doi-asserted-by":"publisher","first-page":"641","DOI":"10.1038\/nrmicro3527","article-title":"A new perspective\non lysogeny: prophages as active\nregulatory switches of bacteria","volume":"13","author":"Feiner","year":"2015","journal-title":"Nat. Rev. Microbiol."},{"issue":"7","key":"2026090307124513300_cit16","doi-asserted-by":"publisher","first-page":"1511","DOI":"10.1038\/ismej.2017.16","article-title":"Lysogeny in nature:\nmechanisms, impact and ecology of temperate phages","volume":"11","author":"Howard-Varona","year":"2017","journal-title":"ISME J."},{"key":"2026090307124513300_cit17","doi-asserted-by":"publisher","first-page":"264","DOI":"10.1016\/j.watres.2019.01.003","article-title":"Coupled\nvirus - bacteria interactions and ecosystem function in an engineered\nmicrobial system","volume":"152","author":"Brown","year":"2019","journal-title":"Water Res."},{"key":"2026090307124513300_cit18","doi-asserted-by":"publisher","first-page":"122200","DOI":"10.1016\/j.watres.2024.122200","article-title":"Prophage induction by non-antibiotic compounds promotes\ntransformation\nof released antibiotic resistance genes from cell lysis","volume":"263","author":"Lu","year":"2024","journal-title":"Water Res."},{"key":"2026090307124513300_cit19","doi-asserted-by":"publisher","first-page":"124010","DOI":"10.1016\/j.watres.2025.124010","article-title":"Prophage induction\nand quorum sensing enhance biofilm\nstability and resistance under ammonia-oxidizing bacteria-mediated\noxidative stress","volume":"284","author":"Ju","year":"2025","journal-title":"Water Res."},{"issue":"12","key":"2026090307124513300_cit20","doi-asserted-by":"publisher","first-page":"8920","DOI":"10.1021\/acs.est.2c01516","article-title":"Bacterial Inactivation\nand Biofilm Disruption through Indigenous Prophage Activation Using\nLow-Intensity Cold Atmospheric Plasma","volume":"56","author":"Gu","year":"2022","journal-title":"Environ.\nSci. Technol."},{"issue":"18","key":"2026090307124513300_cit21","doi-asserted-by":"publisher","first-page":"5563","DOI":"10.1128\/AEM.01655-16","article-title":"Antimicrobial\nEffects of Free Nitrous\nAcid on Desulfovibrio vulgaris: Implications for Sulfide-Induced Corrosion\nof Concrete","volume":"82","author":"Gao","year":"2016","journal-title":"Appl. Environ. Microbiol."},{"issue":"3","key":"2026090307124513300_cit22","doi-asserted-by":"publisher","first-page":"415","DOI":"10.1021\/acsestengg.0c00163","article-title":"In Situ Exploration\nof the Sulfidogenic Process at the Water-Sediment Interface in Sewers:\nMechanism and Implications","volume":"1","author":"Zuo","year":"2021","journal-title":"ACS ES&T Eng."},{"issue":"3","key":"2026090307124513300_cit23","doi-asserted-by":"publisher","first-page":"1270","DOI":"10.1021\/acs.est.7b05121","article-title":"Co-Variation\nbetween Distribution of Microbial Communities and Biological Metabolization\nof Organics in Urban Sewer Systems","volume":"52","author":"Jin","year":"2018","journal-title":"Environ.\nSci. Technol."},{"issue":"32","key":"2026090307124513300_cit24","doi-asserted-by":"publisher","first-page":"17160","DOI":"10.1021\/acs.est.4c13397","article-title":"Urban Sewer\nSystem Is an Incubator of Sewage-Derived\nMercury-Methylating Microorganisms: A New Public Health Challenge","volume":"59","author":"Xia","year":"2025","journal-title":"Environ. Sci. Technol."},{"key":"2026090307124513300_cit25","doi-asserted-by":"publisher","first-page":"123248","DOI":"10.1016\/j.watres.2025.123248","article-title":"Integrated application of nanoscale\nzero-valent iron for sulfide and methane control in sewers and improved\nwastewater treatment","volume":"276","author":"Cen","year":"2025","journal-title":"Water Res."},{"issue":"15","key":"2026090307124513300_cit26","doi-asserted-by":"publisher","first-page":"6196","DOI":"10.1021\/acs.est.2c08443","article-title":"Prophage\nInduction Causes Geobacter Electroactive Biofilm Decay","volume":"57","author":"Liu","year":"2023","journal-title":"Environ. Sci. Technol."},{"issue":"7","key":"2026090307124513300_cit27","doi-asserted-by":"publisher","first-page":"1104","DOI":"10.1038\/s41396-023-01425-w","article-title":"Lysogenic bacteriophages encoding\narsenic resistance determinants promote bacterial community adaptation\nto arsenic toxicity","volume":"17","author":"Tang","year":"2023","journal-title":"ISME J."},{"issue":"8013","key":"2026090307124513300_cit28","doi-asserted-by":"publisher","first-page":"886","DOI":"10.1038\/s41586-024-07396-y","article-title":"Ligand cross-feeding\nresolves bacterial vitamin B(12) auxotrophies","volume":"629","author":"Wienhausen","year":"2024","journal-title":"Nature"},{"issue":"17","key":"2026090307124513300_cit29","doi-asserted-by":"publisher","first-page":"i884","DOI":"10.1093\/bioinformatics\/bty560","article-title":"fastp: an ultra-fast\nall-in-one FASTQ preprocessor","volume":"34","author":"Chen","year":"2018","journal-title":"Bioinformatics"},{"issue":"10","key":"2026090307124513300_cit30","doi-asserted-by":"publisher","first-page":"1674","DOI":"10.1093\/bioinformatics\/btv033","article-title":"MEGAHIT:\nan ultra-fast single-node solution for large and complex metagenomics\nassembly via succinct de Bruijn graph","volume":"31","author":"Li","year":"2015","journal-title":"Bioinformatics"},{"issue":"7","key":"2026090307124513300_cit31","doi-asserted-by":"publisher","first-page":"836","DOI":"10.1038\/s41564-018-0171-1","article-title":"Recovery\nof genomes from metagenomes\nvia a dereplication, aggregation and scoring strategy","volume":"3","author":"Sieber","year":"2018","journal-title":"Nat. Microbiol."},{"issue":"1","key":"2026090307124513300_cit32","doi-asserted-by":"publisher","first-page":"158","DOI":"10.1186\/s40168-018-0541-1","article-title":"MetaWRAP-a flexible\npipeline for\ngenome-resolved metagenomic data analysis","volume":"6","author":"Uritskiy","year":"2018","journal-title":"Microbiome"},{"issue":"7","key":"2026090307124513300_cit33","doi-asserted-by":"publisher","first-page":"1043","DOI":"10.1101\/gr.186072.114","article-title":"CheckM: assessing the quality of\nmicrobial genomes\nrecovered from isolates, single cells, and metagenomes","volume":"25","author":"Parks","year":"2015","journal-title":"Genome Res."},{"issue":"6","key":"2026090307124513300_cit34","doi-asserted-by":"publisher","first-page":"1925","DOI":"10.1093\/bioinformatics\/btz848","article-title":"GTDB-Tk: a toolkit\nto classify genomes with the Genome Taxonomy Database","volume":"36","author":"Chaumeil","year":"2020","journal-title":"Bioinformatics"},{"issue":"1","key":"2026090307124513300_cit35","doi-asserted-by":"publisher","first-page":"2500","DOI":"10.1038\/s41467-020-16366-7","article-title":"Precise phylogenetic analysis of\nmicrobial isolates\nand genomes from metagenomes using PhyloPhlAn 3.0","volume":"11","author":"Asnicar","year":"2020","journal-title":"Nat. Commun."},{"issue":"1","key":"2026090307124513300_cit36","doi-asserted-by":"publisher","first-page":"37","DOI":"10.1186\/s40168-020-00990-y","article-title":"VirSorter2:\na multi-classifier, expert-guided\napproach to detect diverse DNA and RNA viruses","volume":"9","author":"Guo","year":"2021","journal-title":"Microbiome"},{"issue":"1","key":"2026090307124513300_cit37","doi-asserted-by":"publisher","first-page":"64","DOI":"10.1007\/s40484-019-0187-4","article-title":"Identifying viruses\nfrom metagenomic data using deep learning","volume":"8","author":"Ren","year":"2020","journal-title":"Quant.\nBiol."},{"issue":"23","key":"2026090307124513300_cit38","doi-asserted-by":"publisher","first-page":"3150","DOI":"10.1093\/bioinformatics\/bts565","article-title":"CD-HIT: accelerated\nfor clustering the next-generation sequencing data","volume":"28","author":"Fu","year":"2012","journal-title":"Bioinformatics"},{"issue":"5","key":"2026090307124513300_cit39","doi-asserted-by":"publisher","first-page":"578","DOI":"10.1038\/s41587-020-00774-7","article-title":"CheckV assesses the quality and completeness\nof metagenome-assembled viral genomes","volume":"39","author":"Nayfach","year":"2021","journal-title":"Nat.\nBiotechnol."},{"issue":"14","key":"2026090307124513300_cit40","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gkac341","article-title":"vRhyme enables\nbinning of viral genomes from metagenomes","volume":"50","author":"Kieft","year":"2022","journal-title":"Nucleic\nAcids Res."},{"issue":"7","key":"2026090307124513300_cit41","doi-asserted-by":"publisher","first-page":"1009","DOI":"10.1093\/bioinformatics\/btr039","article-title":"Easyfig:\na genome comparison visualizer","volume":"27","author":"Sullivan","year":"2011","journal-title":"Bioinformatics"},{"key":"2026090307124513300_cit42","doi-asserted-by":"publisher","DOI":"10.7717\/peerj.3788","article-title":"MetaCRAST: reference-guided\nextraction of CRISPR spacers from unassembled\nmetagenomes","volume":"5","author":"Moller","year":"2017","journal-title":"PeerJ"},{"issue":"1","key":"2026090307124513300_cit43","doi-asserted-by":"publisher","DOI":"10.1002\/cpps.71","article-title":"Sequence Similarity Searching","volume":"95","author":"Hu","year":"2019","journal-title":"Current protocols in protein science"},{"key":"2026090307124513300_cit44","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/978-1-4939-9173-0_1","article-title":"tRNAscan-SE: Searching for tRNA Genes in Genomic Sequences","volume":"1962","author":"Chan","year":"2019","journal-title":"Methods Mol. Biol."},{"issue":"1","key":"2026090307124513300_cit45","doi-asserted-by":"publisher","first-page":"90","DOI":"10.1186\/s40168-020-00867-0","article-title":"VIBRANT: automated\nrecovery, annotation\nand curation of microbial viruses, and evaluation of viral community\nfunction from genomic sequences","volume":"8","author":"Kieft","year":"2020","journal-title":"Microbiome"},{"issue":"4","key":"2026090307124513300_cit46","doi-asserted-by":"publisher","first-page":"1560","DOI":"10.1093\/bib\/bbx121","article-title":"PHAST, PHASTER and\nPHASTEST: Tools for finding prophage in bacterial genomes","volume":"20","author":"Arndt","year":"2019","journal-title":"Brief. Bioinform."},{"issue":"1","key":"2026090307124513300_cit47","doi-asserted-by":"publisher","first-page":"9262","DOI":"10.1038\/s41467-024-53479-9","article-title":"Global metagenomic survey identifies\nsewage-derived hgcAB(+) microorganisms as key contributors to riverine\nmethylmercury production","volume":"15","author":"Xia","year":"2024","journal-title":"Nat. Commun."},{"issue":"15","key":"2026090307124513300_cit48","doi-asserted-by":"publisher","first-page":"4550","DOI":"10.1016\/j.watres.2010.04.040","article-title":"Various physico-chemical\nstress factors\ncause prophage induction in Nitrosospira multiformis 25196--an ammonia\noxidizing bacteria","volume":"44","author":"Choi","year":"2010","journal-title":"Water Res."},{"issue":"2","key":"2026090307124513300_cit49","doi-asserted-by":"publisher","first-page":"277","DOI":"10.1046\/j.1365-2958.2003.03580.x","article-title":"Prophages\nand bacterial genomics: what have we learned so far?","volume":"49","author":"Casjens","year":"2003","journal-title":"Mol. Microbiol."},{"issue":"6","key":"2026090307124513300_cit50","doi-asserted-by":"publisher","first-page":"845","DOI":"10.1038\/nprot.2015.053","article-title":"The Phyre2\nweb portal for protein modeling, prediction and analysis","volume":"10","author":"Kelley","year":"2015","journal-title":"Nat. Protoc."},{"issue":"5","key":"2026090307124513300_cit51","doi-asserted-by":"publisher","first-page":"503","DOI":"10.1080\/10409238.2017.1337705","article-title":"Lytic transglycosylases:\nconcinnity in concision of the bacterial cell wall","volume":"52","author":"Dik","year":"2017","journal-title":"Crit. Rev. Biochem. Mol. Biol."},{"issue":"2","key":"2026090307124513300_cit52","doi-asserted-by":"publisher","first-page":"201","DOI":"10.1111\/j.1574-6968.2006.00523.x","article-title":"Peptidoglycan\nlytic activity of the Pseudomonas aeruginosa phage phiKZ gp144 lytic\ntransglycosylase","volume":"266","author":"Paradis-Bleau","year":"2007","journal-title":"FEMS Microbiol. Lett."},{"issue":"3","key":"2026090307124513300_cit53","doi-asserted-by":"publisher","DOI":"10.1128\/aem.01716-22","article-title":"Exposure of\nShewanella oneidensis MR-1 to Sublethal Doses of Ionizing Radiation\nTriggers Short-Term SOS Activation and Longer-Term Prophage Activation","volume":"89","author":"Sweet","year":"2023","journal-title":"Appl. Environ. Microbiol."},{"issue":"1","key":"2026090307124513300_cit54","doi-asserted-by":"publisher","first-page":"6013","DOI":"10.1038\/s41467-023-41699-4","article-title":"A systematic analysis of marine lysogens and proviruses","volume":"14","author":"Yi","year":"2023","journal-title":"Nat. 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