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The rice rhizosphere Pseudomonas strain PA1201 contains two reiterated gene clusters, phz1 and phz2, for phenazine-1-carboxylic acid (PCA) biosynthesis; PCA is further converted into PCN by this strain using a functional phzH-encoding glutamine aminotransferase. However, PCN levels in PA1201 constitute approximately one-fifth of PCA levels and the optimal temperature for PCN synthesis is 28 \u00b0C. In this study, the phzH open reading frame (ORF) and promoter region were investigated and reannotated. phzH promoter PphzH was found to be a weak promoter, and PhzH levels were not sufficient to convert all of the native PCA into PCN. Following RNA Seq and promoter-lacZ fusion analyses, a strong quorum sensing (QS)- and thermo-regulated promoter PrhlI was identified and characterized. The activity of PphzH is approximately 1% of PrhlI in PA1201. After three rounds of promoter editing and swapping by PrhlI, a new PCN-overproducing strain UP46 was generated. The optimal fermentation temperature for PCN biosynthesis in UP46 was increased from 28 to 37 \u00b0C and the PCN fermentation titer increased 179.5-fold, reaching 14.1 g\/L, the highest ever reported.<\/jats:p>","DOI":"10.1021\/acssynbio.0c00161","type":"journal-article","created":{"date-parts":[[2020,6,25]],"date-time":"2020-06-25T15:02:50Z","timestamp":1593097370000},"page":"1802-1812","source":"Crossref","is-referenced-by-count":27,"title":["Identification of a Strong Quorum Sensing- and Thermo-Regulated\nPromoter for the Biosynthesis of a New Metabolite Pesticide Phenazine-1-carboxamide\nin\n                    <i>Pseudomonas<\/i>\n                    strain PA1201"],"prefix":"10.1021","volume":"9","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1785-7479","authenticated-orcid":true,"given":"Zi-Jing","family":"Jin","sequence":"first","affiliation":[{"name":"Shanghai Jiao Tong University , , , ,","place":["Shanghai, China, 200240"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lian","family":"Zhou","sequence":"additional","affiliation":[{"name":"Shanghai Jiao Tong University , , , ,","place":["Shanghai, China, 200240"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shuang","family":"Sun","sequence":"additional","affiliation":[{"name":"Shandong Normal University , , , ,","place":["Ji\u2019nan, China, 250014"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ying","family":"Cui","sequence":"additional","affiliation":[{"name":"Shanghai Jiao Tong University , , , ,","place":["Shanghai, China, 200240"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kai","family":"Song","sequence":"additional","affiliation":[{"name":"Shanghai Jiao Tong University , , , ,","place":["Shanghai, China, 200240"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4894-0555","authenticated-orcid":true,"given":"Xuehong","family":"Zhang","sequence":"additional","affiliation":[{"name":"Shanghai Jiao Tong University , , , ,","place":["Shanghai, China, 200240"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7349-924X","authenticated-orcid":true,"given":"Ya-Wen","family":"He","sequence":"additional","affiliation":[{"name":"Shanghai Jiao Tong University , , , ,","place":["Shanghai, China, 200240"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2020,6,25]]},"reference":[{"key":"2026081714221579600_cit1","first-page":"1","article-title":"Blasticidin\nS, a new antibiotic","volume":"111","author":"Takeuchi","year":"1958","journal-title":"J. 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Biochem.\nPhysiol."},{"key":"2026081714221579600_cit4","doi-asserted-by":"publisher","first-page":"251","DOI":"10.1016\/j.copbio.2017.08.016","article-title":"Learn from microbial\nintelligence for avermectins overproduction","volume":"48","author":"Gao","year":"2017","journal-title":"Curr. Opin. Biotechnol."},{"key":"2026081714221579600_cit5","doi-asserted-by":"publisher","first-page":"1700769","DOI":"10.1002\/biot.201700769","article-title":"Strategies for enhancing\nthe yield of the potent insecticide spinosad in actinomycetes","volume":"14","author":"Tao","year":"2019","journal-title":"Biotechnol. J."},{"key":"2026081714221579600_cit6","doi-asserted-by":"publisher","first-page":"599","DOI":"10.1016\/j.molmed.2004.10.002","article-title":"The role of pyocyanin\nin Pseudomonas aeruginosa infection","volume":"10","author":"Lau","year":"2004","journal-title":"Trends Mol. 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Biotechnol."},{"key":"2026081714221579600_cit9","doi-asserted-by":"crossref","first-page":"177","DOI":"10.1007\/978-3-642-40573-0_9","article-title":"Genomic features and regulation of phenazine\nbiosynthesis in the\nrhizosphere strain Pseudomonas aeruginosa M18","volume-title":"Microbial Phenazines","author":"Xu","year":"2013"},{"key":"2026081714221579600_cit10","doi-asserted-by":"publisher","first-page":"503","DOI":"10.1046\/j.1469-8137.2003.00686.x","article-title":"Phenazines and their role in biocontrol\nby Pseudomonas bacteria","volume":"157","author":"Chin-A-Woeng","year":"2003","journal-title":"New\nPhytol."},{"key":"2026081714221579600_cit11","doi-asserted-by":"publisher","first-page":"977","DOI":"10.1038\/ismej.2009.33","article-title":"Phenazine\nantibiotics produced by fluorescent pseudomonads contribute to natural\nsoil suppressiveness to Fusarium wilt","volume":"3","author":"Mazurier","year":"2009","journal-title":"ISME J."},{"key":"2026081714221579600_cit12","doi-asserted-by":"publisher","first-page":"675","DOI":"10.1111\/j.1462-2920.2012.02846.x","article-title":"Recent insights into the diversity,\nfrequency and ecological roles of phenazines in fluorescent Pseudomonas spp","volume":"15","author":"Mavrodi","year":"2013","journal-title":"Environ. 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Eng."},{"key":"2026081714221579600_cit22","doi-asserted-by":"publisher","first-page":"1408","DOI":"10.1039\/b817075b","article-title":"Genomics of secondary metabolite\nproduction by Pseudomonas spp","volume":"26","author":"Gross","year":"2009","journal-title":"Nat. Prod. Rep."},{"key":"2026081714221579600_cit23","doi-asserted-by":"publisher","first-page":"1006","DOI":"10.1094\/MPMI.2001.14.8.1006","article-title":"Introduction of the phzH Gene of Pseudomonas chlororaphis PCL1391 Extends the Range of Biocontrol\nAbility of Phenazine-1-Carboxylic Acid-Producing Pseudomonas spp. Strains","volume":"14","author":"Chin-A-Woeng","year":"2001","journal-title":"Mol. Plant-Microbe Interact."},{"key":"2026081714221579600_cit24","doi-asserted-by":"publisher","first-page":"703","DOI":"10.1111\/j.1365-2672.2009.04466.x","article-title":"Purification, crystal structure and\nantimicrobial activity of phenazine-1-carboxamide produced by a growth-promoting\nbiocontrol bacterium, Pseudomonas aeruginosa MML2212","volume":"108","author":"Shanmugaiah","year":"2010","journal-title":"J. Appl. Microbiol."},{"key":"2026081714221579600_cit25","doi-asserted-by":"publisher","first-page":"3429","DOI":"10.1038\/s41467-018-05683-7","article-title":"Wheat microbiome bacteria\ncan reduce\nvirulence of a plant pathogenic fungus by altering histone acetylation","volume":"9","author":"Chen","year":"2018","journal-title":"Nat. Commun."},{"key":"2026081714221579600_cit26","doi-asserted-by":"publisher","first-page":"32","DOI":"10.1016\/j.pestbp.2017.10.006","article-title":"Effects and\ninhibition mechanism of phenazine-1-carboxamide on the mycelial morphology\nand ultrastructure of Rhizoctonia solani","volume":"147","author":"Xiang","year":"2018","journal-title":"Pestic. Biochem. Physiol."},{"key":"2026081714221579600_cit27","doi-asserted-by":"publisher","first-page":"774","DOI":"10.1111\/1758-2229.12310","article-title":"Interplay\nbetween orfamides, sessilins and phenazines in the control of Rhizoctonia\ndiseases by Pseudomonas sp. CMR12a","volume":"7","author":"Olorunleke","year":"2015","journal-title":"Environ. Microbiol. Rep."},{"key":"2026081714221579600_cit28","doi-asserted-by":"publisher","first-page":"896","DOI":"10.1111\/1758-2229.12454","article-title":"Role of phenazines\nand cyclic lipopeptides produced by Pseudomonas,\nsp. CMR12a in induced systemic resistance on rice and bean","volume":"8","author":"Ma","year":"2016","journal-title":"Environ. Microbiol. Rep."},{"key":"2026081714221579600_cit29","doi-asserted-by":"publisher","first-page":"438","DOI":"10.1186\/1471-2164-12-438","article-title":"Genomic analysis and\ntemperature-dependent transcriptome profiles\nof the rhizosphere originating strain Pseudomonas aeruginosa M18","volume":"12","author":"Wu","year":"2011","journal-title":"BMC Genomics"},{"key":"2026081714221579600_cit30","doi-asserted-by":"publisher","first-page":"574","DOI":"10.1023\/A:1026047301457","article-title":"The Production\nof antifungal metabolites\nby Pseudomonas chlororaphis grown on different nutrient\nsources","volume":"72","author":"Shtark","year":"2003","journal-title":"Microbiology"},{"key":"2026081714221579600_cit31","doi-asserted-by":"publisher","first-page":"610","DOI":"10.1111\/j.1365-2958.2005.04961.x","article-title":"Genome scale analysis of diffusible signal factor regulon\nin Xanthomonas campestris pv. campestris: identification of novel cell-cell communication-dependent genes\nand functions","volume":"59","author":"He","year":"2006","journal-title":"Mol. Microbiol."},{"key":"2026081714221579600_cit32","doi-asserted-by":"publisher","first-page":"44","DOI":"10.1016\/j.plasmid.2011.11.001","article-title":"Construction of a novel sacB-based system for marker-free\ngene deletion in Corynebacterium\nglutamicum","volume":"67","author":"Tan","year":"2012","journal-title":"Plasmid"},{"key":"2026081714221579600_cit33","first-page":"2080","volume-title":"Molecular Cloning: A Laboratory Manual","author":"Green","year":"2012","edition":"4"},{"key":"2026081714221579600_cit34","doi-asserted-by":"publisher","first-page":"948","DOI":"10.2144\/00295bm04","article-title":"Integration-proficient Pseudomonas aeruginosa vectors for isolation of single-copy\nchromosomal lacZ and lux gene fusions","volume":"29","author":"Becher","year":"2000","journal-title":"BioTechniques"},{"key":"2026081714221579600_cit35","doi-asserted-by":"publisher","first-page":"1007","DOI":"10.1007\/s00253-012-4466-5","article-title":"Comparative analysis of temperature-dependent transcriptome\nof Pseudomonas aeruginosa strains from rhizosphere\nand human habitats","volume":"96","author":"Wu","year":"2012","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"2026081714221579600_cit36","doi-asserted-by":"publisher","first-page":"101","DOI":"10.1111\/j.1574-6968.2002.tb11251.x","article-title":"Role of\nthe Pseudomonas aeruginosa las and rhl quorum-sensing systems in rhlI regulation","volume":"212","author":"Kievit","year":"2002","journal-title":"FEMS Microbiol. Lett."},{"key":"2026081714221579600_cit37","doi-asserted-by":"publisher","first-page":"318","DOI":"10.1186\/1471-2164-15-318","article-title":"Pseudomonas\naeruginosa clinical and environmental isolates constitute\na single population with high phenotypic diversity","volume":"15","author":"Grosso-Becerra","year":"2014","journal-title":"BMC Genomics"},{"key":"2026081714221579600_cit38","doi-asserted-by":"publisher","first-page":"6198","DOI":"10.1038\/s41598-019-42722-9","article-title":"Phenazine-1-carboxamide\nfunctionalized mesoporous silica nanoparticles as antimicrobial coatings\non silicone urethral catheters","volume":"9","author":"Kanugala","year":"2019","journal-title":"Sci. Rep."},{"key":"2026081714221579600_cit39","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1080\/14786419.2018.1451997","article-title":"Synthesis\nand bioactivities of diamide\nderivatives containing a phenazine-1-carboxamide scaffold","volume":"33","author":"Zhu","year":"2019","journal-title":"Nat. Prod. Res."},{"key":"2026081714221579600_cit40","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1007\/978-3-642-40573-0_5","article-title":"Fermentative Production of Bacterial Phenazines","volume-title":"Microbial\nPhenazines","author":"Chincholkar","year":"2013"},{"key":"2026081714221579600_cit41","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1016\/j.mex.2014.07.001","article-title":"Standardized\nchemical synthesis of Pseudomonas aeruginosa pyocyanin","volume":"1","author":"Cheluvappa","year":"2014","journal-title":"MethodsX."},{"key":"2026081714221579600_cit42","doi-asserted-by":"publisher","first-page":"130","DOI":"10.1186\/s13068-018-1123-y","article-title":"Engineering\nand systems-level analysis of Pseudomonas chlororaphis for production of phenazine-1-carboxamide using glycerol as the\ncost-effective carbon source","volume":"11","author":"Yao","year":"2018","journal-title":"Biotechnol. Biofuels"},{"key":"2026081714221579600_cit43","doi-asserted-by":"publisher","first-page":"754","DOI":"10.1016\/j.jmb.2011.01.018","article-title":"Mechanism\nof bacterial transcription\ninitiation: RNA polymerase-promoter binding, isomerization to initiation-competent\nopen complexes, and initiation of RNA synthesis","volume":"412","author":"Saecker","year":"2011","journal-title":"J. Mol. Biol."},{"key":"2026081714221579600_cit44","doi-asserted-by":"publisher","first-page":"19862","DOI":"10.1038\/srep19862","article-title":"Characterization\nof the GntR family regulator HpaR1 of the crucifer black rot pathogen Xanthomonas campestris pathovar campestris","volume":"28","author":"Su","year":"2016","journal-title":"Sci. Rep."},{"key":"2026081714221579600_cit45","doi-asserted-by":"publisher","first-page":"57","DOI":"10.1038\/nrmicro787","article-title":"The regulation of bacterial transcription initiation","volume":"2","author":"Browning","year":"2004","journal-title":"Nat. Rev. Microbiol."},{"key":"2026081714221579600_cit46","doi-asserted-by":"publisher","first-page":"331","DOI":"10.1093\/femsre\/fuv022","article-title":"Small RNAs, 5\u2032 UTR elements\nand RNA-binding proteins in intracellular bacteria: impact on metabolism\nand virulence","volume":"39","author":"Oliva","year":"2015","journal-title":"Fems. Microbiol. Rev."},{"key":"2026081714221579600_cit47","doi-asserted-by":"publisher","first-page":"182","DOI":"10.1016\/j.mib.2009.01.005","article-title":"Quorum sensing\nand environmental adaptation in Pseudomonas aeruginosa: a tale of regulatory networks and\nmultifunctional signal molecules","volume":"12","author":"Williams","year":"2009","journal-title":"Curr. Opin.\nMicrobiol."},{"key":"2026081714221579600_cit48","doi-asserted-by":"publisher","first-page":"6568","DOI":"10.1128\/AEM.01148-09","article-title":"Temperature-dependent expression of phzM and its\nregulatory genes lasI and ptsP in\nrhizosphere isolate Pseudomonas sp. strain M18","volume":"75","author":"Huang","year":"2009","journal-title":"Appl. Environ. Microbiol."},{"key":"2026081714221579600_cit49","doi-asserted-by":"publisher","first-page":"e02253","DOI":"10.1128\/mBio.02253-19","article-title":"A rhlI 5\u2032UTR-derived\nsRNA regulates RhlR-dependent\nquorum sensing in Pseudomonas aeruginosa","volume":"10","author":"Thomason","year":"2019","journal-title":"mBio"}],"container-title":["ACS Synthetic Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/pubs.acs.org\/doi\/pdf\/10.1021\/acssynbio.0c00161","content-type":"application\/pdf","content-version":"vor","intended-application":"unspecified"},{"URL":"https:\/\/pubs.acs.org\/asbcd6\/article-pdf\/9\/7\/1802\/5929976\/sb0c00161.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/pubs.acs.org\/asbcd6\/article-pdf\/9\/7\/1802\/5929976\/sb0c00161.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,17]],"date-time":"2026-08-17T18:47:47Z","timestamp":1786992467000},"score":1,"resource":{"primary":{"URL":"https:\/\/pubs.acs.org\/asbcd6\/article\/9\/7\/1802\/592019\/Identification-of-a-Strong-Quorum-Sensing-and"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,6,25]]},"references-count":49,"journal-issue":{"issue":"7","published-online":{"date-parts":[[2020,6,25]]},"published-print":{"date-parts":[[2020,7,17]]}},"URL":"https:\/\/doi.org\/10.1021\/acssynbio.0c00161","relation":{},"ISSN":["2161-5063"],"issn-type":[{"value":"2161-5063","type":"electronic"}],"subject":[],"published":{"date-parts":[[2020,6,25]]}}}