{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,6]],"date-time":"2026-03-06T06:55:32Z","timestamp":1772780132649,"version":"3.50.1"},"reference-count":103,"publisher":"MDPI AG","issue":"2","license":[{"start":{"date-parts":[[2022,2,1]],"date-time":"2022-02-01T00:00:00Z","timestamp":1643673600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001871","name":"Funda\u00e7\u00e3o para a Ci\u00eancia e Tecnologia","doi-asserted-by":"publisher","award":["PTDC\/ASP-PLA\/28266\/2017"],"award-info":[{"award-number":["PTDC\/ASP-PLA\/28266\/2017"]}],"id":[{"id":"10.13039\/501100001871","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001871","name":"Funda\u00e7\u00e3o para a Ci\u00eancia e Tecnologia","doi-asserted-by":"publisher","award":["PTDC\/ASP-PLA\/28263\/2017"],"award-info":[{"award-number":["PTDC\/ASP-PLA\/28263\/2017"]}],"id":[{"id":"10.13039\/501100001871","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001871","name":"Funda\u00e7\u00e3o para a Ci\u00eancia e Tecnologia","doi-asserted-by":"publisher","award":["SFRH\/BD\/145321\/2019"],"award-info":[{"award-number":["SFRH\/BD\/145321\/2019"]}],"id":[{"id":"10.13039\/501100001871","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001871","name":"Funda\u00e7\u00e3o para a Ci\u00eancia e Tecnologia","doi-asserted-by":"publisher","award":["UIDB\/05183\/2020"],"award-info":[{"award-number":["UIDB\/05183\/2020"]}],"id":[{"id":"10.13039\/501100001871","id-type":"DOI","asserted-by":"publisher"}]},{"name":"European Union through the European Regional Development Fund","award":["ALT20-03-0145-FEDER-028266"],"award-info":[{"award-number":["ALT20-03-0145-FEDER-028266"]}]},{"name":"European Union through the European Regional Development Fund","award":["ALT20-03-0145-FEDER-028263"],"award-info":[{"award-number":["ALT20-03-0145-FEDER-028263"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Biology"],"abstract":"<jats:p>Tomato, one of the most cultivated and economically important vegetable crops throughout the world, is affected by a panoply of different pathogens that reduce yield and affect product quality. The study of tomato\u2013pathogen system arises as an ideal system for better understanding the molecular mechanisms underlying disease resistance, offering an opportunity of improving yield and quality of the products. Among several genes already identified in tomato response to pathogens, we highlight those encoding the transcription factors (TFs). TFs act as transcriptional activators or repressors of gene expression and are involved in large-scale biological phenomena. They are key regulators of central components of plant innate immune system and basal defense in diverse biological processes, including defense responses to pathogens. Here, we present an overview of recent studies of tomato TFs regarding defense responses to biotic stresses. Hence, we focus on different families of TFs, selected for their abundance, importance, and availability of functionally well-characterized members in response to pathogen attack. Tomato TFs\u2019 roles and possibilities related to their use for engineering pathogen resistance in tomato are presented. With this review, we intend to provide new insights into the regulation of tomato defense mechanisms against invading pathogens in view of plant breeding.<\/jats:p>","DOI":"10.3390\/biology11020235","type":"journal-article","created":{"date-parts":[[2022,2,1]],"date-time":"2022-02-01T22:16:18Z","timestamp":1643753778000},"page":"235","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":47,"title":["Defense Strategies: The Role of Transcription Factors in Tomato\u2013Pathogen Interaction"],"prefix":"10.3390","volume":"11","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-9262-2880","authenticated-orcid":false,"given":"Maria Doroteia","family":"Campos","sequence":"first","affiliation":[{"name":"MED\u2014Mediterranean Institute for Agriculture, Environment and Development, Instituto de Investiga\u00e7\u00e3o e Forma\u00e7\u00e3o Avan\u00e7ada, Universidade de \u00c9vora, P\u00f3lo da Mitra, Ap. 94, 7006-554 \u00c9vora, Portugal"}]},{"given":"Maria do Ros\u00e1rio","family":"F\u00e9lix","sequence":"additional","affiliation":[{"name":"MED\u2014Mediterranean Institute for Agriculture, Environment and Development & Departamento de Fitotecnia, Escola de Ci\u00eancias e Tecnologia, Universidade de \u00c9vora, P\u00f3lo da Mitra, Ap. 94, 7006-554 \u00c9vora, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0094-6782","authenticated-orcid":false,"given":"Mariana","family":"Patanita","sequence":"additional","affiliation":[{"name":"MED\u2014Mediterranean Institute for Agriculture, Environment and Development, Instituto de Investiga\u00e7\u00e3o e Forma\u00e7\u00e3o Avan\u00e7ada, Universidade de \u00c9vora, P\u00f3lo da Mitra, Ap. 94, 7006-554 \u00c9vora, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5769-1963","authenticated-orcid":false,"given":"Patrick","family":"Materatski","sequence":"additional","affiliation":[{"name":"MED\u2014Mediterranean Institute for Agriculture, Environment and Development, Instituto de Investiga\u00e7\u00e3o e Forma\u00e7\u00e3o Avan\u00e7ada, Universidade de \u00c9vora, P\u00f3lo da Mitra, Ap. 94, 7006-554 \u00c9vora, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1189-9803","authenticated-orcid":false,"given":"Andr\u00e9","family":"Albuquerque","sequence":"additional","affiliation":[{"name":"MED\u2014Mediterranean Institute for Agriculture, Environment and Development, Instituto de Investiga\u00e7\u00e3o e Forma\u00e7\u00e3o Avan\u00e7ada, Universidade de \u00c9vora, P\u00f3lo da Mitra, Ap. 94, 7006-554 \u00c9vora, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1895-2523","authenticated-orcid":false,"given":"Joana A.","family":"Ribeiro","sequence":"additional","affiliation":[{"name":"MED\u2014Mediterranean Institute for Agriculture, Environment and Development, Instituto de Investiga\u00e7\u00e3o e Forma\u00e7\u00e3o Avan\u00e7ada, Universidade de \u00c9vora, P\u00f3lo da Mitra, Ap. 94, 7006-554 \u00c9vora, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6915-1793","authenticated-orcid":false,"given":"Carla","family":"Varanda","sequence":"additional","affiliation":[{"name":"MED\u2014Mediterranean Institute for Agriculture, Environment and Development, Instituto de Investiga\u00e7\u00e3o e Forma\u00e7\u00e3o Avan\u00e7ada, Universidade de \u00c9vora, P\u00f3lo da Mitra, Ap. 94, 7006-554 \u00c9vora, Portugal"}]}],"member":"1968","published-online":{"date-parts":[[2022,2,1]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","unstructured":"Campos, M.D., Patanita, M., Varanda, C., Materatski, P., and F\u00e9lix, M.R. (2021). Plant-Pathogen Interaction. Biology, 10.","DOI":"10.3390\/biology10050444"},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Burdon, J.J., and Zhan, J. (2020). Climate change and disease in plant communities. PLoS Biol., 18.","DOI":"10.1371\/journal.pbio.3000949"},{"key":"ref_3","doi-asserted-by":"crossref","unstructured":"Campos, M.D., Patanita, M., Campos, C., Materatski, P., Varanda, C.M.R., Brito, I., and F\u00e9lix, M.R. (2019). Detection and quantification of Fusarium spp. (F. oxysporum, F. verticillioides, F. graminearum) and Magnaporthiopsis maydis in maize using real-time PCR targeting the ITS region. Agronomy, 9.","DOI":"10.3390\/agronomy9020045"},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Varanda, C.M.R., Materatski, P., Landum, M., Campos, M.D., and F\u00e9lix, M.R. (2019). Fungal communities associated with peacock and cercospora leaf spots in olive. Plants, 8.","DOI":"10.3390\/plants8060169"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"694","DOI":"10.3389\/fpls.2019.00694","article-title":"Establishment of a sensitive qPCR methodology for detection of the olive-infecting viruses in portuguese and tunisian orchards","volume":"10","author":"Campos","year":"2019","journal-title":"Front. Plant Sci."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1016\/j.pbi.2014.04.004","article-title":"Transcriptional control of plant defence responses","volume":"20","author":"Buscaill","year":"2014","journal-title":"Curr. Opin. Plant Biol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"539","DOI":"10.1038\/nrg2812","article-title":"Plant immunity: Towards an integrated view of plant\u2013pathogen interactions","volume":"11","author":"Dodds","year":"2010","journal-title":"Nat. Rev. Genet."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"328","DOI":"10.1094\/MPMI-04-10-0098","article-title":"Biocontrol treatments confer protection against Verticillium dahliae infection of potato by inducing antimicrobial metabolites","volume":"24","author":"Adam","year":"2011","journal-title":"Mol. Plant-Microbe Interact."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"797","DOI":"10.1038\/nrg3748","article-title":"Microbial genome-enabled insights into plant-microorganism interactions","volume":"15","author":"Guttman","year":"2014","journal-title":"Nat. Rev. Genet."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"335","DOI":"10.2174\/1389203717666160619185308","article-title":"Transcription factors involved in plant resistance to pathogens","volume":"18","author":"Amorim","year":"2016","journal-title":"Curr. Protein Pept. Sci."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"803","DOI":"10.1016\/j.cell.2006.02.008","article-title":"Host-microbe interactions: Shaping the evolution of the plant immune response","volume":"124","author":"Chisholm","year":"2006","journal-title":"Cell"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"323","DOI":"10.1038\/nature05286","article-title":"The plant immune system","volume":"444","author":"Jones","year":"2006","journal-title":"Nature"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1111\/j.1439-0434.2006.01073.x","article-title":"Signal transduction and transcriptional regulation of plant defence responses","volume":"154","author":"Jalali","year":"2006","journal-title":"J. Phytopathol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"397","DOI":"10.1016\/S0065-2296(09)51010-5","article-title":"Chapter 10 Transcriptional regulation of plant defense responses","volume":"51","author":"Gatz","year":"2009","journal-title":"Adv. Bot. Res."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"6383","DOI":"10.1038\/s41598-017-04792-5","article-title":"Pathogen recognition in compatible plant-microbe interactions","volume":"7","author":"Rezzonico","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"110","DOI":"10.1016\/j.gde.2017.02.007","article-title":"Transcription factor\u2013DNA binding: Beyond binding site motifs","volume":"43","author":"Inukai","year":"2017","journal-title":"Curr. Opin. Genet. Dev."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"260","DOI":"10.1093\/bfgp\/elv011","article-title":"Functional studies of transcription factors involved in plant defenses in the genomics era","volume":"14","author":"Seo","year":"2015","journal-title":"Brief. Funct. Genomics"},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Javed, T., Shabbir, R., Ali, A., Afzal, I., Zaheer, U., and Gao, S.J. (2020). Transcription factors in plant stress responses: Challenges and potential for sugarcane improvement. Plants, 9.","DOI":"10.3390\/plants9040491"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1038\/s41438-021-00607-x","article-title":"High throughput sequencing unravels tomato-pathogen interactions towards a sustainable plant breeding","volume":"8","author":"Campos","year":"2021","journal-title":"Hortic. Res."},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Zhao, M., Ji, H.M., Gao, Y., Cao, X.X., Mao, H.Y., Ouyang, S.Q., and Liu, P. (2018). An integrated analysis of mRNA and srna transcriptional profiles in tomato root: Insights on tomato wilt disease. PLoS ONE, 13.","DOI":"10.1371\/journal.pone.0206765"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"881","DOI":"10.1007\/s11240-014-0664-4","article-title":"Tomato (Solanum lycopersicum L.) in the service of biotechnology","volume":"120","author":"Gerszberg","year":"2015","journal-title":"Plant Cell. Tissue Organ Cult."},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Panno, S., Davino, S., Caruso, A.G., Bertacca, S., Crnogorac, A., and Mandi, A. (2021). A review of the most common and economically important diseases that undermine the cultivation of tomato crop in the mediterranean basin. Agronomy, 11.","DOI":"10.3390\/agronomy11112188"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1080\/07352689.2021.1941605","article-title":"Genomics and marker-assisted improvement of vegetable crops","volume":"40","author":"Simko","year":"2021","journal-title":"CRC. Crit. Rev. Plant Sci."},{"key":"ref_24","doi-asserted-by":"crossref","unstructured":"Salava, H., Thula, S., Mohan, V., Kumar, R., and Maghuly, F. (2021). Application of genome editing in tomato breeding: Mechanisms, advances, and prospects. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22020682"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1007\/s13205-017-0896-1","article-title":"Disease management of tomato through PGPB: Current trends and future perspective","volume":"7","author":"Singh","year":"2017","journal-title":"3 Biotech"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"482","DOI":"10.1038\/s41598-017-00578-x","article-title":"Rapid generation of a transgene-free powdery mildew resistant tomato by genome deletion","volume":"7","author":"Nekrasov","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"239","DOI":"10.1007\/s13205-017-0870-y","article-title":"Genetic engineering strategies for biotic and abiotic stress tolerance and quality enhancement in horticultural crops: A comprehensive review","volume":"7","author":"Parmar","year":"2017","journal-title":"3 Biotech"},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Capriotti, L., Baraldi, E., Mezzetti, B., Limera, C., and Sabbadini, S. (2020). Biotechnological approaches: Gene overexpression, gene silencing, and genome editing to control fungal and oomycete diseases in grapevine. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21165701"},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Louwaars, N., and Jochemsen, H. (2021). An ethical and societal analysis for biotechnological methods in plant breeding. Agronomy, 11.","DOI":"10.3390\/agronomy11061183"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"2727","DOI":"10.1007\/s00122-021-03874-3","article-title":"Advances in application of genome editing in tomato and recent development of genome editing technology","volume":"134","author":"Xia","year":"2021","journal-title":"Theor. Appl. Genet."},{"key":"ref_31","doi-asserted-by":"crossref","unstructured":"Pathak, K., and Gogoi, B. (2016). RNA interference (RNAi): Application in crop improvement: A review. Agric. Rev., 37.","DOI":"10.18805\/ag.v37i3.3540"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/978-1-62703-278-0_1","article-title":"Virus-induced gene silencing (VIGS) in plants: An overview of target species and the virus-derived vector systems","volume":"975","author":"Lange","year":"2013","journal-title":"Methods Mol. Biol."},{"key":"ref_33","doi-asserted-by":"crossref","unstructured":"Varanda, C.M., F\u00e9lix, M.R., Campos, M.D., Patanita, M., and Materatski, P. (2021). Plant viruses: From targets to tools for CRISPR. Viruses, 13.","DOI":"10.3390\/v13010141"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1038\/s41438-019-0159-x","article-title":"CRISPR technology is revolutionizing the improvement of tomato and other fruit crops","volume":"6","author":"Wang","year":"2019","journal-title":"Hortic. Res."},{"key":"ref_35","unstructured":"Chaudhary, R., and Atamian, H.S. (2017). Resistance-gene-mediated defense responses against biotic stresses in the crop model plant tomato. J. Plant Pathol. Microbiol., 8."},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Huibers, R.P., Loonen, A.E.H.M., Gao, D., Van den Ackerveken, G., Visser, R.G.F., and Bai, Y. (2013). Powdery mildew resistance in tomato by impairment of SlPMR4 and SlDMR1. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0067467"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"247","DOI":"10.1016\/j.tplants.2010.02.006","article-title":"WRKY transcription factors","volume":"15","author":"Rushton","year":"2010","journal-title":"Trends Plant Sci."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1186\/s42483-019-0022-x","article-title":"WRKY transcription factors: Evolution, binding, and action","volume":"1","author":"Chen","year":"2019","journal-title":"Phytopathol. Res."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Baillo, E.H., Kimotho, R.N., Zhang, Z., and Xu, P. (2019). Transcription factors associated with abiotic and biotic stress tolerance and their potential for crops improvement. Genes, 10.","DOI":"10.3390\/genes10100771"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"801","DOI":"10.3389\/fpls.2018.00801","article-title":"The role of tomato WRKY genes in plant responses to combined abiotic and biotic stresses","volume":"9","author":"Bai","year":"2018","journal-title":"Front. Plant Sci."},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"(2005). Xie, Zhen, Zhong-Lin Zhang, Xiaolu Zou, Jie Huang, Paul Ruas, Daniel Thompson, and Q. J.S. Annotations and functional analyses of the rice WRKY gene superfamily reveal positive and negative regulators of abscisic acid signaling in., 137, 176\u2013189.","DOI":"10.1104\/pp.104.054312"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1023\/A:1020780022549","article-title":"Expression profiles of the Arabidopsis WRKY gene superfamily during plant defense response","volume":"51","author":"Dong","year":"2003","journal-title":"Plant Mol. Biol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"23101","DOI":"10.1038\/srep23101","article-title":"Genomic identification of WRKY transcription factors in carrot (Daucus carota) and analysis of evolution and homologous groups for plants","volume":"6","author":"Li","year":"2016","journal-title":"Sci. Rep."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"e27700","DOI":"10.4161\/psb.27700","article-title":"Wrky transcription factors jack of many trades in plants","volume":"9","author":"Bakshi","year":"2014","journal-title":"Plant Signal. Behav."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"2105","DOI":"10.1126\/science.290.5499.2105","article-title":"Arabidopsis transcription factors: Genome-wide comparative analysis among eukaryotes","volume":"290","author":"Riechmann","year":"2000","journal-title":"Science"},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Zhang, H., Kang, H., Su, C., Qi, Y., Liu, X., and Pu, J. (2018). Genome-wide identification and expression profile analysis of the NAC transcription factor family during abiotic and biotic stress in woodland strawberry. PLoS ONE, 13.","DOI":"10.1371\/journal.pone.0197892"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"217","DOI":"10.1007\/s12155-009-9047-9","article-title":"A bioinformatic analysis of NAC genes for plant cell wall development in relation to lignocellulosic bioenergy production","volume":"2","author":"Shen","year":"2009","journal-title":"Bioenergy Res."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"547","DOI":"10.1007\/s00438-008-0386-6","article-title":"Systematic sequence analysis and identification of tissue-specific or stress-responsive genes of NAC transcription factor family in rice","volume":"280","author":"Fang","year":"2008","journal-title":"Mol. Genet. Genom."},{"key":"ref_49","doi-asserted-by":"crossref","unstructured":"Jin, J.F., Wang, Z.Q., He, Q.Y., Wang, J.Y., Li, P.F., Xu, J.M., Zheng, S.J., Fan, W., and Yang, J.L. (2020). Genome-wide identification and expression analysis of the NAC transcription factor family in tomato (Solanum lycopersicum) during aluminum stress. BMC Genomics, 21.","DOI":"10.1186\/s12864-020-6689-7"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"1900","DOI":"10.3389\/fpls.2018.01900","article-title":"NAC Family Transcription factors in tobacco and their potential role in regulating leaf senescence","volume":"9","author":"Li","year":"2018","journal-title":"Front. Plant Sci."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1038\/s41438-018-0036-z","article-title":"Comprehensive analysis of NAC transcription factors and their expression during fruit spine development in cucumber (Cucumis sativus L.)","volume":"5","author":"Liu","year":"2018","journal-title":"Hortic. Res."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1186\/s42483-018-0008-0","article-title":"NAC transcription factors in plant immunity","volume":"1","author":"Yuan","year":"2019","journal-title":"Phytopathol. Res."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"354","DOI":"10.1111\/pbi.12776","article-title":"NAC transcription factor JUNGBRUNNEN1 enhances drought tolerance in tomato","volume":"16","author":"Thirumalaikumar","year":"2018","journal-title":"Plant Biotechnol. J."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"248","DOI":"10.3389\/fmicb.2013.00248","article-title":"Roles of NAC transcription factors in the regulation of biotic and abiotic stress responses in plants","volume":"4","author":"Nuruzzaman","year":"2013","journal-title":"Front. Microbiol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"639","DOI":"10.1111\/nph.12291","article-title":"APETALA2\/Ethylene Responsive Factor (AP2\/ERF) transcription factors: Mediators of stress responses and developmental programs","volume":"199","author":"Licausi","year":"2013","journal-title":"New Phytol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"426","DOI":"10.1007\/s13205-018-1454-1","article-title":"DREB2 (dehydration-responsive element-binding protein 2) type transcription factor in sorghum (Sorghum bicolor): Genome-wide identification, characterization and expression profiles under cadmium and salt stresses","volume":"8","author":"Akbudak","year":"2018","journal-title":"3 Biotech"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"483","DOI":"10.1007\/s11103-020-01052-5","article-title":"Structural insights into Arabidopsis ethylene response factor 96 with an extended N-terminal binding to GCC box","volume":"104","author":"Chen","year":"2020","journal-title":"Plant Mol. Biol."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"465","DOI":"10.1016\/j.pbi.2004.04.007","article-title":"Regulation of disease resistance pathways by AP2\/ERF transcription factors","volume":"7","author":"Gutterson","year":"2004","journal-title":"Curr. Opin. Plant Biol."},{"key":"ref_59","doi-asserted-by":"crossref","unstructured":"Chopra, R., Burow, G., Hayes, C., Emendack, Y., Xin, Z., and Burke, J. (2015). Transcriptome profiling and validation of gene based single nucleotide polymorphisms (SNPs) in sorghum genotypes with contrasting responses to cold stress. BMC Genom., 16.","DOI":"10.1186\/s12864-015-2268-8"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"95","DOI":"10.1017\/S0021859610000742","article-title":"Transgenic expression of sorghum DREB2 in rice improves tolerance and yield under water limitation","volume":"149","author":"Bihani","year":"2011","journal-title":"J. Agric. Sci."},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"Guo, B., Wei, Y., Xu, R., Lin, S., Luan, H., Lv, C., Zhang, X., Song, X., and Xu, R. (2016). Genome-wide analysis of APETALA2\/ethylene-responsive factor (AP2\/ERF) gene family in barley (Hordeum vulgare L.). PLoS ONE, 11.","DOI":"10.1371\/journal.pone.0161322"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"6","DOI":"10.1186\/s40529-016-0159-1","article-title":"Multiple regulatory roles of AP2\/ERF transcription factor in angiosperm","volume":"58","author":"Gu","year":"2017","journal-title":"Bot. Stud."},{"key":"ref_63","doi-asserted-by":"crossref","unstructured":"Liu, R., Song, J., Liu, S., Chen, C., Zhang, S., Wang, J., Xiao, Y., Cao, B., Lei, J., and Zhu, Z. (2021). Genome-wide identification of the capsicum bHLH transcription factor family: Discovery of a candidate regulator involved in the regulation of species-specific bioactive metabolites. BMC Plant Biol., 21.","DOI":"10.1186\/s12870-021-03004-7"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"510","DOI":"10.1016\/j.pld.2020.10.004","article-title":"Genome-wide identification and expression analysis of NtbHLH gene family in tobacco (Nicotiana tabacum L.) and the role of NtbHLH86 in drought adaptation","volume":"43","author":"Bai","year":"2021","journal-title":"Plant Divers."},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"Wang, R., Zhao, P., Kong, N., Lu, R., Pei, Y., Huang, C., Ma, H., and Chen, Q. (2018). Genome-wide identification and characterization of the potato bHLH transcription factor family. Genes, 9.","DOI":"10.3390\/genes9010054"},{"key":"ref_66","doi-asserted-by":"crossref","unstructured":"Sun, H., Fan, H.J., and Ling, H.Q. (2015). Genome-wide identification and characterization of the bHLH gene family in tomato. BMC Genom., 16.","DOI":"10.1186\/s12864-014-1209-2"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"1749","DOI":"10.1105\/tpc.013839","article-title":"The Arabidopsis basic\/helix-loop-helix transcription factor family","volume":"15","author":"Huq","year":"2003","journal-title":"Plant Cell"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"501","DOI":"10.1007\/PL00006494","article-title":"Positional dependence, cliques, and predictive motifs in the bHLH protein domain","volume":"48","author":"Atchley","year":"1999","journal-title":"J. Mol. Evol."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"828","DOI":"10.1105\/tpc.113.121111","article-title":"The bHLH transcription factor HBI1 mediates the trade-off between growth and pathogen-associated molecular pattern-triggered immunity in Arabidopsis","volume":"26","author":"Fan","year":"2014","journal-title":"Plant Cell"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"740","DOI":"10.1093\/pcp\/pcs033","article-title":"The bHLH Rac immunity1 (RAI1) is activated by OsRac1 via OsMAPK3 and OsMAPK6 in rice immunity","volume":"53","author":"Kim","year":"2012","journal-title":"Plant Cell Physiol."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"1614","DOI":"10.1094\/MPMI-05-19-0122-R","article-title":"Tomato bHLH132 transcription factor controls growth and defense and is activated by Xanthomonas euvesicatoria effector XopD during pathogenesis","volume":"32","author":"Kim","year":"2019","journal-title":"Mol. Plant-Microbe Interact."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"106","DOI":"10.1016\/S1360-1385(01)02223-3","article-title":"bZIP transcription factors in Arabidopsis","volume":"7","author":"Jakoby","year":"2002","journal-title":"Trends Plant Sci."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"463","DOI":"10.1093\/dnares\/dss026","article-title":"Genome-wide analysis of bZIP-encoding genes in maize","volume":"19","author":"Wei","year":"2012","journal-title":"DNA Res."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"1","DOI":"10.3389\/fpls.2020.00139","article-title":"Characterization of the bZIP transcription factor family in pepper (Capsicum annuum L.): CabZIP25 positively modulates the salt Tolerance","volume":"11","author":"Gai","year":"2020","journal-title":"Front. Plant Sci."},{"key":"ref_75","doi-asserted-by":"crossref","unstructured":"Li, D., Fu, F., Zhang, H., and Song, F. (2015). Genome-wide systematic characterization of the bZIP transcriptional factor family in tomato (Solanum lycopersicum L.). BMC Genom., 16.","DOI":"10.1186\/s12864-015-1990-6"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"336","DOI":"10.1104\/pp.106.095299","article-title":"Genetic interactions of TGA transcription factors in the regulation of pathogenesis-related genes and disease resistance in Arabidopsis","volume":"144","author":"Kesarwani","year":"2007","journal-title":"Plant Physiol."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"825","DOI":"10.1094\/MPMI-10-14-0313-R","article-title":"Expression and functional roles of the pepper pathogen-induced bZIP transcription factor CabZIP2 in enhanced disease resistance to bacterial pathogen infection","volume":"28","author":"Lim","year":"2015","journal-title":"Mol. Plant-Microbe Interact."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"635","DOI":"10.1038\/nature11119","article-title":"The tomato genome sequence provides insights into fleshy fruit evolution","volume":"485","author":"Sato","year":"2012","journal-title":"Nature"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1016\/j.plantsci.2014.08.001","article-title":"Tomato WRKY transcriptional factor SlDRW1 is required for disease resistance against Botrytis cinerea and tolerance to oxidative stress","volume":"227","author":"Liu","year":"2014","journal-title":"Plant Sci."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"67","DOI":"10.1007\/s11240-015-0815-2","article-title":"SpWRKY1 mediates resistance to Phytophthora infestans and tolerance to salt and drought stress by modulating reactive oxygen species homeostasis and expression of defense-related genes in tomato","volume":"123","author":"Li","year":"2015","journal-title":"Plant Cell. Tissue Organ Cult."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"787","DOI":"10.1007\/s00122-017-3035-9","article-title":"Transcriptome signatures of tomato leaf induced by Phytophthora infestans and functional identification of transcription factor SpWRKY3","volume":"131","author":"Cui","year":"2018","journal-title":"Theor. Appl. Genet."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"495","DOI":"10.1007\/s00438-012-0696-6","article-title":"Genome-wide analysis of WRKY transcription factors in Solanum lycopersicum","volume":"287","author":"Huang","year":"2012","journal-title":"Mol. Genet. Genom."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"52","DOI":"10.1007\/s12374-014-0407-4","article-title":"sheng Over-expression of SlWRKY39 leads to enhanced resistance to multiple stress factors in tomato","volume":"58","author":"Sun","year":"2015","journal-title":"J. Plant Biol."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"229","DOI":"10.1111\/j.1365-313X.2010.04232.x","article-title":"WRKY72-type transcription factors contribute to basal immunity in tomato and Arabidopsis as well as gene-for-gene resistance mediated by the tomato R gene Mi-1","volume":"63","author":"Bhattarai","year":"2010","journal-title":"Plant J."},{"key":"ref_85","doi-asserted-by":"crossref","unstructured":"Chinnapandi, B., Bucki, P., and Braun Miyara, S. (2017). SlWRKY45, nematode-responsive tomato WRKY gene, enhances susceptibility to the root knot nematode; M. javanica infection. Plant Signal. Behav., 12.","DOI":"10.1080\/15592324.2017.1356530"},{"key":"ref_86","doi-asserted-by":"crossref","unstructured":"Huang, Y., Li, M.Y., Wu, P., Xu, Z.S., Que, F., Wang, F., and Xiong, A.S. (2016). Members of WRKY Group III transcription factors are important in TYLCV defense signaling pathway in tomato (Solanum lycopersicum). BMC Genom., 17.","DOI":"10.1186\/s12864-016-3123-2"},{"key":"ref_87","doi-asserted-by":"crossref","unstructured":"Wang, J., Zheng, C., Shao, X., Hu, Z., Li, J., Wang, P., Wang, A., Yu, J., and Shi, K. (2020). Transcriptomic and genetic approaches reveal an essential role of the NAC transcription factor SlNAP1 in the growth and defense response of tomato. Hortic. Res., 7.","DOI":"10.1038\/s41438-020-00442-6"},{"key":"ref_88","doi-asserted-by":"crossref","unstructured":"Liu, B., Ouyang, Z., Zhang, Y., Li, X., Hong, Y., Huang, L., Liu, S., Zhang, H., Li, D., and Song, F. (2014). Tomato NAC transcription factor SlSRN1 positively regulates defense response against biotic stress but negatively regulates abiotic stress response. PLoS ONE, 9.","DOI":"10.1371\/journal.pone.0102067"},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"1214","DOI":"10.1111\/nph.12096","article-title":"SlNAC1, a stress-related transcription factor, is fine-tuned on both the transcriptional and the post-translational level","volume":"197","author":"Huang","year":"2013","journal-title":"New Phytol."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"311","DOI":"10.1105\/tpc.104.027235","article-title":"A NAC domain protein interacts with tomato leaf curl virus replication accessory protein and enhances viral replication","volume":"17","author":"Selth","year":"2005","journal-title":"Plant Cell"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1016\/j.sajb.2016.07.001","article-title":"Identification of transcription factors in tomato, potentially related to early blight resistance at invasion in host tissue using, microarray expression profiling","volume":"106","author":"Upadhyay","year":"2016","journal-title":"S. Afr. J. Bot."},{"key":"ref_92","doi-asserted-by":"crossref","unstructured":"Yang, H., Sun, Y., Wang, H., Zhao, T., Xu, X., Jiang, J., and Li, J. (2021). Genome-wide identification and functional analysis of the ERF2 gene family in response to disease resistance against Stemphylium lycopersici in tomato. BMC Plant Biol., 21.","DOI":"10.1186\/s12870-021-02848-3"},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1007\/s11103-013-0117-1","article-title":"The transcription factor SlSHINE3 modulates defense responses in tomato plants","volume":"84","author":"Buxdorf","year":"2014","journal-title":"Plant Mol. Biol."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"5529","DOI":"10.1021\/acs.jafc.9b08069","article-title":"CRISPR\/Cas9-mediated SlMYC2 mutagenesis adverse to tomato plant growth and MeJA-induced fruit resistance to Botrytis cinerea","volume":"68","author":"Shu","year":"2020","journal-title":"J. Agric. Food Chem."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"2191","DOI":"10.1111\/j.1365-3040.2010.02220.x","article-title":"The transcription factor SlAREB1 confers drought, salt stress tolerance and regulates biotic and abiotic stress-related genes in tomato","volume":"33","author":"Orellana","year":"2010","journal-title":"Plant Cell Environ."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1016\/j.plgene.2017.11.002","article-title":"Genome wide expression analysis of WRKY genes in tomato (Solanum lycopersicum) under drought stress","volume":"13","author":"Karkute","year":"2018","journal-title":"Plant Gene"},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"299","DOI":"10.1007\/s00425-011-1509-6","article-title":"SlWRKY70 is required for Mi-1-mediated resistance to aphids and nematodes in tomato","volume":"235","author":"Atamian","year":"2012","journal-title":"Planta"},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1038\/s41438-018-0111-5","article-title":"A NAC transcription factor, NOR-like1, is a new positive regulator of tomato fruit ripening","volume":"5","author":"Gao","year":"2018","journal-title":"Hortic. Res."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1093\/pcp\/pct162","article-title":"A new tomato NAC (NAM ATAF1\/2\/CUC2) transcription factor, SlNAC4, functions as a positive regulator of fruit ripening and carotenoid accumulation","volume":"55","author":"Zhu","year":"2014","journal-title":"Plant Cell Physiol."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"3167","DOI":"10.1105\/tpc.114.128272","article-title":"Closely related NAC transcription factors of tomato differentially regulate stomatal closure and reopening during pathogen attack","volume":"26","author":"Du","year":"2014","journal-title":"Plant Cell"},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"plantgenome2015-09","DOI":"10.3835\/plantgenome2015.09.0082","article-title":"AP2\/ERF transcription actors involved in response to tomato yellow leaf curly virus in tomato","volume":"9","author":"Huang","year":"2016","journal-title":"Plant Genome"},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"817","DOI":"10.1105\/tpc.000794","article-title":"Tomato transcription factors Pti4, Pti5, and Pti6 activate defense responses when expressed in Arabidopsis","volume":"14","author":"Gu","year":"2002","journal-title":"Plant Cell"},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"686","DOI":"10.1093\/mp\/sss128","article-title":"MYC2: The master in action","volume":"6","author":"Kazan","year":"2013","journal-title":"Mol. 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