vavilovВавиловский журнал генетики и селекцииVavilov Journal of Genetics and Breeding2500-3259Institute of Cytology and Genetics of Siberian Branch of the RAS10.18699/VJ15.105vavilov-469Research ArticleФизиологическая генетика растений. ОБЗОРPhysiological genetics of plants. REVIEWМеханизмы регуляции передачи этиленового сигнала у растенийRegulatory mechanisms tuning ethylen signaling in plantsЗемлянскаяЕ. В.ZemlyanskayaE. V.ОмельянчукН. А.OmelyanchukN. A.ЕрмаковА. А.ErmakovA. A.МироноваВ. В.MironovaV. V.Федеральное государственное бюджетное научное учреждение «Федеральный исследовательский центр Институт цитологии и генетики Сибирского отделения Российской академии наук», Новосибирск, Россия Федеральное государственное автономное образовательное учреждение высшего образования «Новосибирский национальный исследовательский государственный университет», Новосибирск, РоссияРоссияInstitute of Cytology and Genetics SB RAS, Novosibirsk, Russia Novosibirsk National Research State University, Novosibirsk, RussiaRussian FederationФедеральное государственное бюджетное научное учреждение «Федеральный исследовательский центр Институт цитологии и генетики Сибирского отделения Российской академии наук», Новосибирск, РоссияРоссияInstitute of Cytology and Genetics SB RAS, Novosibirsk, RussiaRussian Federation201615082016203386395Copyright © Землянская Е.В., Омельянчук Н.А., Ермаков А.А., Миронова В.В., 20162016Землянская Е.В., Омельянчук Н.А., Ермаков А.А., Миронова В.В.Zemlyanskaya E.V., Omelyanchuk N.A., Ermakov A.A., Mironova V.V.This work is licensed under a Creative Commons Attribution 4.0 License.https://vavilov.elpub.ru/jour/article/view/469

Фитогормон этилен регулирует широкий спектр физиологических процессов на разных этапах онтогенеза растений и ответов на воздействие различных стрессовых факторов. Среди прочих под контролем этого фитогормона находятся такие практически значимые характеристики сельскохозяйственных культур как скорость созревания плодов и устойчивость растений к неблагоприятным условиям. Вследствие этого понимание молекулярных механизмов, лежащих в основе действия этилена, на сегодняшний день является одним из основных вопросов биологии растений, как с точки зрения фундаментальных исследований, так и для решения практических задач. Биосинтез этилена из аминокислоты метионина и основные этапы пути передачи его сигнала в клетке от мембранных рецепторов до эффекторных генов изучены достаточно детально, и результаты этих исследований представлены в виде многочисленных обзоров. Гораздо меньше известно о генетической регуляции этих двух процессов, хотя именно благодаря этой регуляции обеспечивается быстрая и адекватная реакция растения на различные внутренние и внешние стимулы, а также разнообразие физиологических ответов растения на действие этилена. В настоящем обзоре обобщены данные о механизмах регуляции биосинтеза этилена и передачи его сигнала. Описываются ключевые факторы транскрипционной и посттрансляционой регуляции, контролирующие экспрессию и стабильность ключевых компонентов путей биосинтеза и передачи сигнала этилена, а также множественные обратные связи, дополняющие линейную модель сигнального пути. Особое внимание уделяется роли взаимодействия этилена с сигнальными путями других фитогормонов. Разные механизмы их взаимодействия проиллюстрированы на примере синергии или антагонизма этилена с ауксином, жасмонатами, цитокнинами и брассиностероидами. Кроме того, обсуждаются возможные молекулярные основы разнообразия физиологических ответов на этилен.

Plant hormone ethylene regulates a wide range of physiological processes during plant development and coordinates different stress responses. Among others ethylene controls such practically significant characteristics of agricultural crops as fruit ripening rates and plant tolerance to stress conditions. That is why understanding molecular mechanisms underlying ethylene action is one of the basic questions in plant biology that is addressed in the context of both fundamental research and application in agriculture. Ethylene biosynthesis from methionine amino acid and the main points of its signaling pathway from membrane receptors to effector genes are studied in details and widely reviewed. Far less is known about genetic regulation of these two processes although it is the one that ensures accurate plant reaction to different endogenous and exogenous signals and causes the multiplicity of different physiological responses to ethylene. This review summarizes data about regulatory mechanisms of ethylene biosynthesis and signaling. It reports the key transcriptional and post-translational regulatory factors which control expression and stability of the main components of ethylene biosynthesis and signaling pathways, and describes multiple feed-backs supplementing the linear model of ethylene signaling. Particular attention is given to the role of hormonal crosstalk in the process. Different mechanisms of hormonal interaction are illustrated by synergy or antagonism of ethylene and auxin, jasmonates, cytokinins, brassinosteroids. Possible molecular basics of multiplicity of different physiological responses to ethylene is also discussed.

этиленфитогормоныморфогенезпуть передачи сигналарегуляция транскрипциипосттрансляционная регуляцияethyleneplant hormonemorphogenesissignaling pathwaytranscriptional regulationposttranslational regulationРФФИ, ИЦиГ СО РАНReferencesКудрякова Н.В., Бурханова Э.А., Яковлева Л.А., Ракитин В.Ю., Смит А.Р., Холл М.А., Кулаева О.Н. Этилен и цитокинины в регуляции старения срезанных листьев мутанта eti5 Arabidopsis thaliana и исходного дикого типа. Физиология растений. 2001;48(5):723-727.Abel S., Nguyen M.D., Chow W., Theologis A. ACS4, a primary indoleacetic acid- responsive gene encoding 1-aminocyclopropane-1-carboxylate synthase in Arabidopsis thaliana: structural characterization, expression in Escherichia coli, and expression characteristics in response to auxin [corrected]. J. Biol. Chem. 1995;270(32): 19093-19099.Черных О.А., Левицкий В.Г., Омельянчук Н.А., Миронова В.В. Компьютерный анализ и функциональная аннотация сайтов связывания транскрипционных факторов AP2/ERF в геноме Arabidopsis thaliana L. Вавиловский журнал генетики и селекции. 2014;18(4/2):887-897.Abeles F.B., Morgan P.W., Saltveit M.E. Ethylene in plant biology. San Diego: Acad. Press, 1992.Abel S., Nguyen M.D., Chow W., Theologis A. ACS4, a primary indoleacetic acid-responsive gene encoding 1-aminocyclopropane-1-carboxylate synthase in Arabidopsis thaliana: structural characterization, expression in Escherichia coli, and expression characteristics in response to auxin [corrected]. J. Biol. Chem. 1995;270(32):19093-19099.Alexander L., Grierson D. Ethylene biosynthesis and action in tomato: a model for climacteric fruit ripening. J. Exp. Bot. 2002;53(377): 2039-2055.Abeles F.B., Morgan P.W., Saltveit M.E. Ethylene in plant biology. San Diego: Acad. Press, 1992.Alonso J.M., Stepanova A.N., Solano R., Wisman E., Ferrari S., Ausubel F.M., Ecker J.R. Five components of the ethylene-response pathway identified in a screen for weak ethylene-insensitive mutants in Arabidopsis. Proc. Natl. Acad. Sci. USA. 2003;100(5):2992-2997.Alexander L., Grierson D. Ethylene biosynthesis and action in tomato: a model for climacteric fruit ripening. J. Exp. Bot. 2002;53(377):2039-2055.An F., Zhang X., Zhu Z., Ji Y., He W., Jiang Z., Li M., Guo H. Coordinated regulation of apical hook development by gibberellins and ethylene in etiolated Arabidopsis seedlings. Cell Res. 2012;22(5): 915-927. DOI 10.1038/cr.2012.29Alonso J.M., Stepanova A.N., Solano R., Wisman E., Ferrari S., Ausubel F.M., Ecker J.R. Five components of the ethylene-response pathway identified in a screen for weak ethylene-insensitive mutants in Arabidopsis. Proc. Natl. Acad. Sci. USA. 2003;100(5):2992-2997.An F., Zhao Q., Ji Y., Li W., Jiang Z., Yu X., Zhang C., Han Y., He W., Liu Y., Zhang S., Ecker J.R., Guo H. Ethylene-induced stabilization of ETHYLENE-INSENSITIVE3 and EIN3-LIKE1 is mediated by proteasomal degradation of EIN3 binding F-box 1 and 2 that requires EIN2 in Arabidopsis. Plant Cell. 2010;22(7):2384-2401. DOI 10.1105/tpc.110.076588An F., Zhang X., Zhu Z., Ji Y., He W., Jiang Z., Li M., Guo H. Coordinated regulation of apical hook development by gibberellins and ethylene in etiolated Arabidopsis seedlings. Cell Res. 2012;22(5):915-927. DOI: 10.1038/cr.2012.29Cary A.J., Liu W., Howell S.H. Cytokinin action is coupled to ethylene in its effects on the inhibition of root and hypocotyl elongation in Arabidopsis thaliana seedlings. Plant Physiol. 1995;107(4):1075- 1082. DOI 10.1104/pp.107.4.1075An F., Zhao Q., Ji Y., Li W., Jiang Z., Yu X., Zhang C., Han Y., He W., Liu Y., Zhang S., Ecker J.R., Guo H. Ethylene-induced stabilization of ETHYLENE-INSENSITIVE3 and EIN3-LIKE1 is mediated by proteasomal degradation of EIN3 binding F-box 1 and 2 that requires EIN2 in Arabidopsis. Plant Cell. 2010;22(7):2384-2401. DOI: 10.1105/tpc.110.076588Chae H.S., Faure F., Kieber J.J. The eto1, eto2 and eto3 mutations and cytokinin treatment increase ethylene biosynthesis in Arabidopsis by increasing the stability of the ACS protein. Plant Cell. 2003;15(2):545-559. DOI 10.1105/tpc.006882Chang K.N., Zhong S., Weirauch M.T., Hon G., Pelizzola M., Li H., Huang S.S.C., Schmitz R.J., Urich M.A., Kuo D., Nery J.R., Qiao H., Yang A., Jamali A., Chen H., Ideker T., Ren B., Bar-Joseph Z., Hughes T.R., Ecker J.R. Temporal transcriptional response to ethylene gas drives growth hormone cross-regulation in Arabidopsis. eLife. 2013;2:e00675. DOI: 10.7554/eLife.00675Chang K.N., Zhong S., Weirauch M.T., Hon G., Pelizzola M., Li H., Huang S.S.C., Schmitz R.J., Urich M.A., Kuo D., Nery J.R., Qiao H., Yang A., Jamali A., Chen H., Ideker T., Ren B., Bar-Joseph Z., Hughes T.R., Ecker J.R. Temporal transcriptional response to ethylene gas drives growth hormone cross-regulation in Arabidopsis. eLife. 2013;2:e00675. DOI 10.7554/eLife.00675Cary A.J., Liu W., Howell S.H. Cytokinin action is coupled to ethylene in its effects on the inhibition of root and hypocotyl elongation in Arabidopsis thaliana seedlings. Plant Physiol. 1995;107(4):1075-1082. DOI:10.1104/pp.107.4.1075Chen Y.F., Shakeel S.N., Bowers J., Zhao X.C., Etheridge N., Schaller G.E. Ligand- induced degradation of the ethylene receptor ETR2. J. Biol. Chem. 2007;282(34):24752-24758.Chae H.S., Faure F., Kieber J.J. The eto1, eto2 and eto3 mutations and cytokinin treatment increase ethylene biosynthesis in Arabidopsis by increasing the stability of the ACS protein. Plant Cell. 2003;15(2):545-559. DOI: 10.1105/tpc.006882Chernykh O.A., Levitsky V.G., Omelyanchuk N.A., Mironova V.V. Computational analysis and functional annotation of AP2/ERF translocation factor binding sites in Arabidopsis thaliana L. genome. Vavilovskii Zhurnal Genetiki i Selektsii = Vavilov Journal of Genetics and Breeding. 2014;18(4/2):887-897.Chen Y.F., Shakeel S.N., Bowers J., Zhao X.C., Etheridge N., Schaller G.E. Ligand-induced degradation of the ethylene receptor ETR2. J. Biol. Chem. 2007;282(34):24752-24758.Chilley P.M., Casson S.A., Tarkowski P., Hawkins N., Wang K.L., Hussey P.J., Beale M., Ecker J.R., Sandberg G.K., Lindsey K. The POLARIS peptide of Arabidopsis regulates auxin transport and root growth via effects on ethylene signaling. Plant Cell. 2006;18(11): 3058-3072.Chilley P.M., Casson S.A., Tarkowski P., Hawkins N., Wang K.L., Hussey P.J., Beale M., Ecker J.R., Sandberg G.K., Lindsey K. The POLARIS peptide of Arabidopsis regulates auxin transport and root growth via effects on ethylene signaling. Plant Cell. 2006;18(11):3058-3072.Cho Y.H., Yoo S.D. Novel connections and gaps in ethylene signaling from the ER membrane to the nucleus. Front. Plant Sci. 2015;5:733. DOI 10.3389/fpls.2014.00733Cho Y.H., Yoo S.D. Novel connections and gaps in ethylene signaling from the ER membrane to the nucleus. Front. Plant Sci. 2015;5:733. DOI: 10.3389/fpls.2014.00733De Paepe A., Vuylsteke M., Van Hummelen P., Zabeau M., Van Der Straeten D. Transcriptional profiling by cDNA-AFLP and microarray analysis reveals novel insights into the early response to ethylene in Arabidopsis. Plant J. 2004;39(4):537-559. DOI 10.1111/j.1365-313X.2004.02156.xDe Paepe A., Vuylsteke M., Van Hummelen P., Zabeau M., Van Der Straeten D. Transcriptional profiling by cDNA-AFLP and microarray analysis reveals novel insights into the early response to ethylene in Arabidopsis. Plant J. 2004;39(4):537-559. DOI: 10.1111/j.1365-313X.2004.02156.xDíaz J., Álvarez-Buylla E.R. A model of the ethylene signaling pathway and its gene response in Arabidopsis thaliana: Pathway crosstalk and noise-filtering properties. Chaos. 2006;16(2):023112. DOI 10.1063/1.2189974Díaz J., Álvarez-Buylla E.R. A model of the ethylene signaling pathway and its gene response in Arabidopsis thaliana: Pathway crosstalk and noise-filtering properties. Chaos. 2006;16(2):023112. DOI: 10.1063/1.2189974Dong C.H., Rivarola M., Resnick J.S., Maggin B.D., Chang C. Subcellular co- localization of Arabidopsis RTE1 and ETR1 supports a regulatory role for RTE1 in ETR1 ethylene signaling. Plant J. 2008; 53(2):275-286.Dong C.H., Rivarola M., Resnick J.S., Maggin B.D., Chang C. Subcellular co-localization of Arabidopsis RTE1 and ETR1 supports a regulatory role for RTE1 in ETR1 ethylene signaling. Plant J. 2008;53(2):275-286.Dugardeyn J., Vandenbussche F., Van Der Straeten D. To grow or not to grow: what can we learn on ethylene–gibberellin cross-talk by in silico gene expression analysis? J. Exp. Bot. 2008;59(1):1-16. DOI 10.1093/jxb/erm349Dugardeyn J., Vandenbussche F., Van Der Straeten D. To grow or not to grow: what can we learn on ethylene–gibberellin cross-talk by in silico gene expression analysis? J. Exp. Bot. 2008;59(1):1-16. DOI: 10.1093/jxb/erm349Ecker J.R. The ethylene signal transduction pathway in plants. Science. 1995;268:667-675.Ecker J.R. The ethylene signal transduction pathway in plants. Science. 1995;268:667-675.Fujimoto S.Y., Ohta M., Usui A., Shinshi H., Ohme-Takagi M. Arabidopsis ethylene- responsive element binding factors act as transcriptional activators or repressors of GCC box-mediated gene expression. Plant Cell. 2000;12(3):393-404.Fujimoto S.Y., Ohta M., Usui A., Shinshi H., Ohme-Takagi M. Arabidopsis ethylene-responsive element binding factors act as transcriptional activators or repressors of GCC box-mediated gene expression. Plant Cell. 2000;12(3):393-404.Gallie D.R., Geisler-Lee J., Chen J., Jolley B. Tissue-specific expression of the ethylene biosynthetic machinery regulates root growth in maize. Plant. Mol. Biol. 2009;69(1-2):195-211. DOI 10.1007/s11103-008-9418-1Gallie D.R., Geisler-Lee J., Chen J., Jolley B. Tissue-specific expression of the ethylene biosynthetic machinery regulates root growth in maize. Plant. Mol. Biol. 2009;69(1-2):195-211. DOI: 10.1007/s11103-008-9418-1Gazzarrini S., McCourt P. Cross-talk in plant hormone signalling: What Arabidopsis mutants are telling us. Ann. Bot. 2003;91(6):605-612.DOI 10.1093/aob/mcg064Gazzarrini S., McCourt P. Cross-talk in plant hormone signalling: What Arabidopsis mutants are telling us. Ann. Bot. 2003;91(6):605-612. DOI: 10.1093/aob/mcg064Giovannoni J.J. Fruit ripening mutants yield insights into ripening control. Curr. Opin. Plant Biol. 2007;10(3):283-289. DOI 10.1016/j.pbi.2007.04.008Giovannoni J.J. Fruit ripening mutants yield insights into ripening control. Curr. Opin. Plant Biol. 2007;10(3):283-289. DOI: 10.1016/j.pbi.2007.04.008Han L., Li G.J., Yang KY., Mao G., Wang R., Liu Y., Zhang S. Mitogenactivated protein kinase 3 and 6 regulate Botrytis cinerea-induced ethylene production in Arabidopsis. Plant J. 2010;64(1):114-127. DOI 10.1111/j.1365-313X.2010.04318.xHan L., Li G.J., Yang KY., Mao G., Wang R., Liu Y., Zhang S. Mitogenactivated protein kinase 3 and 6 regulate Botrytis cinerea-induced ethylene production in Arabidopsis. Plant J. 2010;64(1):114-127. DOI: 10.1111/j.1365-313X.2010.04318.xHansen M., Chae H.S., Kieber J.J. Regulation of ACS protein stability by cytokinin and brassinosteroid. Plant J. 2009;57(4):606-614. DOI 10.1111/j.1365- 313X.2008.03711.xHansen M., Chae H.S., Kieber J.J. Regulation of ACS protein stability by cytokinin and brassinosteroid. Plant J. 2009;57(4):606-614. DOI:10.1111/j.1365-313X.2008.03711.xIkeda Y., Men S., Fischer U., Stepanova A.N., Alonso J.M., Ljung K., Grebe M. Local auxin biosynthesis modulates gradient-directed planar polarity in Arabidopsis. Nat. Cell Biol. 2009;11(6):731-738. DOI 10.1038/ncb1879Ikeda Y., Men S., Fischer U., Stepanova A.N., Alonso J.M., Ljung K., Grebe M. Local auxin biosynthesis modulates gradient-directed planar polarity in Arabidopsis. Nat. Cell Biol. 2009;11(6):731-738. DOI: 10.1038/ncb1879Itkin M., Seybold H., Breitel D., Rogachev I., Meir S., Aharoni A. TOMATO AGAMOUS- LIKE 1 is a component of the fruit ripening regulatory network. Plant J. 2009;60(6):1081-1095. DOI 10.1111/j.1365-313X.2009.04064.xItkin M., Seybold H., Breitel D., Rogachev I., Meir S., Aharoni A. TOMATO AGAMOUS-LIKE 1 is a component of the fruit ripening regulatory network. Plant J. 2009;60(6):1081-1095. DOI: 10.1111/j.1365-313X.2009.04064.xIto Y., Kitagawa M., Ihashi N., Yabe K., Kimbara J., Yasuda J., Ito H., Inakuma T., Hiroi S., Kasumi T. DNA-binding specificity, transcriptional activation potential, and the rin mutation effect for the tomato fruit-ripening regulator RIN. Plant J. 2008;55(2):212-223. DOI 10.1111/j.1365-313X.2008.03491.xIto Y., Kitagawa M., Ihashi N., Yabe K., Kimbara J., Yasuda J., Ito H., Inakuma T., Hiroi S., Kasumi T. DNA-binding specificity, transcriptional activation potential, and the rin mutation effect for the tomato fruit-ripening regulator RIN. Plant J. 2008;55(2):212-223. DOI: 10.1111/j.1365-313X.2008.03491.xJoo S., Seo Y.S., Kim S.M., Hong D.K., Park K.Y., Kim W.T. Brassinosteroid induction of AtACS4 encoding an auxin-responsive 1-aminocyclopropane-1-carboxylate synthase 4 in Arabidopsis seedlings. Physiol. Plant. 2006;126(4):592-604. DOI 10.1111/j.1399-3054.2005.00602.xJoo S., Seo Y.S., Kim S.M., Hong D.K., Park K.Y., Kim W.T. Brassinosteroid induction of AtACS4 encoding an auxin-responsive 1-aminocyclopropane-1-carboxylate synthase 4 in Arabidopsis seedlings. Physiol. Plant. 2006;126(4):592-604. DOI: 10.1111/j.1399-3054.2005.00602.xJu C., Chang C. Advances in ethylene signalling: protein complexes at the endoplasmic reticulum membrane. AoB Plants. 2012:pls031. DOI 10.1093/aobpla/pls031Ju C., Chang C. Advances in ethylene signalling: protein complexes at the endoplasmic reticulum membrane. AoB Plants. 2012:pls031. DOI:10.1093/aobpla/pls031Ju C., Yoon G.M., Shemansky J.M., Lin D.Y., Ying Z.I., Chang J., Garrett W.M., Kessenbrock M., Groth G., Tucker M.L., Cooper B., Kieber J.J., Chang C. CTR1 phosphorylates the central regulator EIN2 to control ethylene hormone signaling from the ER membrane to the nucleus in Arabidopsis. Proc. Natl. Acad. Sci. USA. 2012; 109(47):19486-19491. DOI 10.1073/pnas.1214848109Ju C., Yoon G.M., Shemansky J.M., Lin D.Y., Ying Z.I., Chang J., Garrett W.M., Kessenbrock M., Groth G., Tucker M.L., Cooper B., Kieber J.J., Chang C. CTR1 phosphorylates the central regulator EIN2 to control ethylene hormone signaling from the ER membrane to the nucleus in Arabidopsis. Proc. Natl. Acad. Sci. USA. 2012;109(47):19486-19491. DOI: 10.1073/pnas.1214848109Karlova R., Chapman N., David K., Angenent G.C., Seymour G.B., de Maagd R.A. Transcriptional control of fleshy fruit development and ripening. J. Exp. Bot. 2014;65(16):4527-4541. DOI 10.1093/jxb/eru316Karlova R., Chapman N., David K., Angenent G.C., Seymour G.B., de Maagd R.A. Transcriptional control of fleshy fruit development and ripening. J. Exp. Bot. 2014;65(16):4527-4541. DOI:10.1093/jxb/eru316Kendrick M.D., Chang C. Ethylene signaling: new levels of complexity and regulation. Curr. Opin. Plant Biol. 2008;11(5):479-485. DOI 10.1016/j.pbi.2008.06.011Kendrick M.D., Chang C. Ethylene signaling: new levels of complexity and regulation. Curr. Opin. Plant Biol. 2008;11(5):479-485. DOI: 10.1016/j.pbi.2008.06.011Kevany B.M., Tieman D.M., Taylor M.G., Cin V.D., Klee H.J. Ethylene receptor degradation controls the timing of ripening in tomato fruit. Plant J. 2007;51(3):458-467. DOI 10.1111/j.1365-313X.2007.03170.xKevany B.M., Tieman D.M., Taylor M.G., Cin V.D., Klee H.J. Ethylene receptor degradation controls the timing of ripening in tomato fruit. Plant J. 2007;51(3):458-467. DOI: 10.1111/j.1365-313X.2007.03170.xKonishi M., Yanagisawa S. Ethylene signaling in Arabidopsis involves feedback regulation via the elaborate control of EBF2 expression by EIN3. Plant J. 2008;55(5):821-831. DOI 10.1111/j.1365-313X.2008.03551.xKosugi S., Ohashi Y. Cloning and DNA-binding properties of a tobacco Ethylene-Insensitive3 (EIN3) homolog. Nucleic Acids Res. 2000;28(4):960-967.Kosugi S., Ohashi Y. Cloning and DNA-binding properties of a tobacco Ethylene- Insensitive3 (EIN3) homolog. Nucleic Acids Res. 2000; 28(4):960-967.Konishi M., Yanagisawa S. Ethylene signaling in Arabidopsis involves feedback regulation via the elaborate control of EBF2 expression by EIN3. Plant J. 2008;55(5):821-831. DOI: 10.1111/j.1365-313X.2008.03551.xKudryakova N.V., Burkhanova E.A., Yakovleva L.A., Rakitin V.Yu., Smith A.R., Hall M.A., Kulaeva O.N. Ethylene and cytokinin in the control of senescence in detached leaves of Arabidopsis thaliana eti- 5 mutant and wild-type plants. Fiziologiya rasteniy = Plant Physiology (Moscow). 2001;48(5):723-727.Lacey R.F., Binder B.M. How plants sense ethylene gas — The ethylene receptors. J. Inorg. Biochem. 2014;133:58-62. DOI: 10.1016/j.jinorgbioLacey R.F., Binder B.M. How plants sense ethylene gas – The ethylene receptors. J. Inorg. Biochem. 2014;133:58-62. DOI 10.1016/j.jinorgbioLewis D.R., Negi S., Sukumar P., Muday G.K. Ethylene inhibits lateral root development, increases IAA transport and expression of PIN3 and PIN7 auxin efflux carriers. Development. 2011;138(16):3485-3495. DOI:10.1242/dev.065102Lewis D.R., Negi S., Sukumar P., Muday G.K. Ethylene inhibits lateral root development, increases IAA transport and expression of PIN3 and PIN7 auxin efflux carriers. Development. 2011;138(16):3485-3495. DOI 10.1242/dev.065102Li G., Meng X., Wang R., Mao G., Han L., Liu Y., Zhang S. Dual-level regulation of ACC synthase activity by MPK3/MPK6 cascade and its downstream WRKY transcription factor during ethylene induction in Arabidopsis. PLoS Genetics. 2012;8(6):e1002767. DOI: 10.1371/journal.pgen.1002767Li G., Meng X., Wang R., Mao G., Han L., Liu Y., Zhang S. Duallevel regulation of ACC synthase activity by MPK3/MPK6 cascade and its downstream WRKY transcription factor during ethylene induction in Arabidopsis. PLoS Genetics. 2012;8(6):e1002767. DOI 10.1371/journal.pgen.1002767Liu Q., Wen C.K. Cooperative ethylene receptor signaling. Plant Signal. Behav. 2012a;7(8):1009-1013. DOI: 10.4161/psb.20937Li W., Ma M., Feng Y., Li H., Wang Y., Ma Y., Li M., An F., Guo H. EIN2-directed translational regulation of ethylene signaling in Arabidopsis. Cell. 2015;163(3):670-683. DOI 10.1016/j.cell.2015.09.037Liu Q., Wen C.K. Arabidopsis ETR1 and ERS1 Differentially Repress the Ethylene Response in Combination with Other Ethylene Receptor genes. Plant Physiol. 2012b;158(3):1193-1207. DOI: 10.1104/pp.111.187757Liu Q., Wen C.K. Cooperative ethylene receptor signaling. Plant Signal. Behav. 2012a;7(8):1009-1013. DOI 10.4161/psb.20937Lorenzo O., Piqueras R., Sanchez-Serrano J.J., Solano R. ETHYLENE RESPONSE FACTOR1 integrates signals from ethylene and jasmonate pathways in plant defense. Plant Cell. 2003;15(1):165-178. DOI: 10.1105/tpc.007468Liu Q., Wen C.K. Arabidopsis ETR1 and ERS1 differentially repress the ethylene response in combination with other ethylene receptor genes. Plant Physiol. 2012b;158(3):1193-1207. DOI 10.1104/pp.111.187757Ludwikówa A., Cieśla A., Kasprowicz-Maluśki A., Mituła F., Tajdel M., Gałgański Ł., Ziółkowski P.A., Kubiak P., Małecka A., Piechalak A., Szabat M., Górska A., Dąbrowski M., Ibragimow I., Sadowski J. Arabidopsis protein phosphatase 2C ABI1 interacts with type I ACC synthases and is involved in the regulation of ozone-induced ethylene biosynthesis. Mol. Plant. 2014;7(6):960–976. DOI: 10.1093/mp/ssu025Lorenzo O., Piqueras R., Sanchez-Serrano J.J., Solano R. ETHYLENE RESPONSE FACTOR1 integrates signals from ethylene and jasmonate pathways in plant defense. Plant Cell. 2003;15(1):165-178. DOI 10.1105/tpc.007468Lyzenga W.J., Stone S.L. Regulation of ethylene biosynthesis through protein degradation. Plant Signal. Behav. 2012;7(11):1438-1442. DOI: 10.4161/psb.21930Ludwikówa A., Cieśla A., Kasprowicz-Maluśki A., Mituła F., Tajdel M., Gałgański Ł., Ziółkowski P.A., Kubiak P., Małecka A., Piechalak A., Szabat M., Górska A., Dąbrowski M., Ibragimow I., Sadowski J. Arabidopsis protein phosphatase 2C ABI1 interacts with type I ACC synthases and is involved in the regulation of ozone- induced ethylene biosynthesis. Mol. Plant. 2014;7(6):960-976. DOI 10.1093/mp/ssu025McKeon T., Yang S.F. Biosynthesis and metabolism of ethylene. In: Plant hormones and their role in plant growth and development. P.J. Davides (ed.); Dordrecht: Martinus Nijhoff Publishers, 1987:94-112.Lyzenga W.J., Stone S.L. Regulation of ethylene biosynthesis through protein degradation. Plant Signal. Behav. 2012;7(11):1438-1442. DOI 10.4161/psb.21930McManus M.T The plant hormone ethylene. Annual Plant Reviews. Oxford: Wiley-Blackwell, 2012(44).McKeon T., Yang S.F. Biosynthesis and metabolism of ethylene. Plant Hormones and Their Role in Plant Growth and Development. Ed. P.J. Davides. Dordrecht: Martinus Nijhoff Publ., 1987.Merchante C., Alonso J.M., Stepanova A.N. Ethylene signaling: simple ligand, complex regulation. Curr. Opin. Plant Biol. 2013;16(5):554-560. DOI: 10.1016/j.pbi.2013.08.001McManus M.T. The plant hormone ethylene. Annual Plant Reviews. Oxford: Wiley-Blackwell, 2012;44.Muday G.K., Rahman A., Binder B.M. Auxin and ethylene: collaborators or competitors? Trends Plant Sci. 2012;17(4):181-195. DOI: 10.1016/j.tplants.2012.02.001Merchante C., Alonso J.M., Stepanova A.N. Ethylene signaling: simple ligand, complex regulation. Curr. Opin. Plant Biol. 2013;16(5):554- 560. DOI 10.1016/j.pbi.2013.08.001Murr D.P., Yang S.F. Conversion of 5’-methylthioadenosine to methionine by apple tissue. Phytochemistry. 1975;14:1291-1292. DOI: 10.1016/S0031- 9422(00)98613-8.Muday G.K., Rahman A., Binder B.M. Auxin and ethylene: collaborators or competitors? Trends Plant Sci. 2012;17(4):181-195. DOI 10.1016/j.tplants.2012.02.001Ohme-Takagi M., Shinshi H. Ethylene-lnducible DNA binding proteins that interact with an ethylene-responsive element. Plant Cell. 1995;7(2):173-182.Murr D.P., Yang S.F. Conversion of 5’-methylthioadenosine to methionine by apple tissue. Phytochemistry. 1975;14:1291-1292. DOI 10.1016/S0031-9422(00)98613-8Pirrello J., Prasad B.C., Zhang W., Chen K., Mila I., Zouine M., Latché A., Pech J.C., Ohme-Takagi M., Regad F., Bouzayen M. Functional analysis and binding affinity of tomato ethylene response factors provide insight on the molecular bases of plant differential responses to ethylene. BMC Plant Biol. 2012;12:190. DOI: 10.1186/1471-2229-12-190Ohme-Takagi M., Shinshi H. Ethylene-lnducible DNA binding proteins that interact with an ethylene-responsive element. Plant Cell. 1995;7(2):173-182.Pre M., Atallah M., Champion A., De Vos M., Pieterse C.M., Memelink J. The AP2/ERF domain transcription factor ORA59 integrates jasmonic acid and ethylene signals in plant defense. Plant Physiol. 2008;147(3):1347-1357. DOI: 10.1104/pp.108.117523Pirrello J., Prasad B.C., Zhang W., Chen K., Mila I., Zouine M., Latché A., Pech J.C., Ohme-Takagi M., Regad F., Bouzayen M. Functional analysis and binding affinity of tomato ethylene response factors provide insight on the molecular bases of plant differential responses to ethylene. BMC Plant Biol. 2012;12:190. DOI 10.1186/1471- 2229-12-190Qiao H., Shen Z., Huang S.C., Schmitz R.J., Urich M.A., Briggs S.P., Ecker J.R. Processing and subcellular trafficking of ER-tethered EIN2 control response to ethylene gas. Science. 2012;338(6105):390-393. DOI: 10.1126/science.1225974Pre M., Atallah M., Champion A., De Vos M., Pieterse C.M., Memelink J. The AP2/ERF domain transcription factor ORA59 integrates jasmonic acid and ethylene signals in plant defense. Plant Physiol. 2008;147(3):1347-1357. DOI 10.1104/pp.108.117523Riechmann J. L., Heard J., Martin G., Reuber L., Jiang C.-Z., Keddie J., Adam L., Pineda O., Ratcliffe O. J., Samaha R. R., Creelman R., Pilgrim M., Broun P., Zhang J. Z., Ghandehari D., Sherman B. K., Yu G.-L. Arabidopsis transcription factors: genome-wide comparative analysis among eukaryotes. Science. 2000;290(5499):2105-2110.Qiao H., Shen Z., Huang S.C., Schmitz R.J., Urich M.A., Briggs S.P., Ecker J.R. Processing and subcellular trafficking of ER-tethered EIN2 control response to ethylene gas. Science. 2012;338(6105):390-393. DOI 10.1126/science.1225974Rudus I., Sasiak M., Kepczynski J. Regulation of ethylene biosynthesis at the level of 1-aminocyclopropane-1-carboxylate oxidase (ACO) gene. Acta Physiol. Plant. 2013;35(2):295-307. DOI: 10.1007/s11738-012-1096-6Riechmann J.L., Heard J., Martin G., Reuber L., Jiang C.-Z., Keddie J., Adam L., Pineda O., Ratcliffe O.J., Samaha R.R., Creelman R., Pilgrim M., Broun P., Zhang J.Z., Ghandehari D., Sherman B.K., Yu G.- L. Arabidopsis transcription factors: genome-wide comparative analysis among eukaryotes. Science. 2000;290(5499):2105- 2110.Ruzicka K., Ljung K., Vanneste S., Podhorska R., Beeckman T., Friml J., Benkova E. Ethylene regulates root growth through effects on auxin biosynthesis and transport-dependent auxin distribution. Plant Cell. 2007;19(7):2197-2212. DOI: 10.1105/tpc.107.052126Rudus I., Sasiak M., Kepczynski J. Regulation of ethylene biosynthesis at the level of 1-aminocyclopropane-1-carboxylate oxidase (ACO) gene. Acta Physiol. Plant. 2013;35(2):295-307. DOI 10.1007/s11738-012-1096-6Rzewuski G., Suter M. Ethylene biosynthesis and signaling in rice. Plant Sci. 2008;175:32-42. DOI: 10.1016/j.plantsci.2008.01.012Ruzicka K., Ljung K., Vanneste S., Podhorska R., Beeckman T., Friml J., Benkova E. Ethylene regulates root growth through effects on auxin biosynthesis and transport- dependent auxin distribution. Plant Cell. 2007;19(7):2197-2212. DOI 10.1105/tpc.107.052126Shakeel S., Gao Z., Amir M., Chen Y.F., Rai M.I., Haq N.U., Schaller G.E. Ethylene regulates levels of ethylene-receptor/CTR1 signaling complexes in Arabidopsis thaliana. J. Biol. Chem. 2015;290(19):12415-12424. DOI: 10.1074/jbc.M115.652503Rzewuski G., Suter M. Ethylene biosynthesis and signaling in rice. Plant Sci. 2008;175:32-42. DOI 10.1016/j.plantsci.2008.01.012Shin K., Lee S., Song W.Y., Lee R.A., Lee I., Ha K., Koo J.C., Park S.K., Nam H.G., Lee Y., Soh M.S. Genetic Identification of ACCRESISTANT2 reveals involvement of LYSINE HISTIDINE TRANSPORTER1 in the uptake of 1-aminocyclopropane-1-carboxylic acid in Arabidopsis thaliana. Plant Cell Physiol. 2015;56(3):572-582. DOI: 10.1093/pcp/pcu201Shakeel S., Gao Z., Amir M., Chen Y.F., Rai M.I., Haq N.U., Schaller G.E. Ethylene regulates levels of ethylene-receptor/CTR1 signaling complexes in Arabidopsis thaliana. J. Biol. Chem. 2015; 290(19):12415-12424. DOI 10.1074/jbc.M115.652503Skottke K.R., Yoon G.M., Kieber J.J., DeLong A. Protein phosphatase 2A controls ethylene biosynthesis by differentially regulating the turnover of ACC synthase isoforms. PLoS Genet. 2011;7(4):e1001370. DOI: 10.1371/journal.pgen.1001370Shin K., Lee S., Song W.Y., Lee R.A., Lee I., Ha K., Koo J.C., Park S.K., Nam H.G., Lee Y., Soh M.S. Genetic Identification of ACC-RESISTANT2 reveals involvement of LYSINE HISTIDINE TRANSPORTER1 in the uptake of 1-aminocyclopropane-1-carboxylic acid in Arabidopsis thaliana. Plant Cell Physiol. 2015;56(3):572-582.DOI 10.1093/pcp/pcu201Solano R., Stepanova A., Chao Q., Ecker J.R. Nuclear events in ethylene signaling: a transcriptional cascade mediated by ETHYLENEINSENSITIVE3 and ETHYLENE-RESPONSE-FACTOR1. Genes Dev. 1998;12(23):3703-3714.Skottke K.R., Yoon G.M., Kieber J.J., DeLong A. Protein phosphatase 2A controls ethylene biosynthesis by differentially regulating the turnover of ACC synthase isoforms. PLoS Genet. 2011;7(4):e1001370. DOI 10.1371/journal.pgen.1001370Stepanova A.N., Alonso J.M. Ethylene signaling and response: where different regulatory modules meet. Curr. Opin. Plant Biol. 2009;12(5):548-555. DOI: 10.1016/j.pbi.2009.07.009Solano R., Stepanova A., Chao Q., Ecker J.R. Nuclear events in ethylene signaling: a transcriptional cascade mediated by ETHYLENEINSENSITIVE3 and ETHYLENE-RESPONSE- FACTOR1. Genes Dev. 1998;12(23):3703-3714.Stepanova A.N., Hoyt J.M., Hamilton A.A., Alonso J.M. A link between ethylene and auxin uncovered by the characterization of two root-specific ethylene-insensitive mutants in Arabidopsis. Plant Cell. 2005;17(8):2230-2242. DOI: 10.1105/tpc.105.033365Stepanova A.N., Alonso J.M. Ethylene signaling and response: where different regulatory modules meet. Curr. Opin. Plant Biol. 2009; 12(5):548-555. DOI 10.1016/j.pbi.2009.07.009Stepanova A.N., Yun J., Likhacheva A.V., Alonso J.M. Multilevel interactions between ethylene and auxin in Arabidopsis roots. Plant Cell. 2007;19(7):2169-2185. DOI: 10.1105/tpc.107.052068Stepanova A.N., Hoyt J.M., Hamilton A.A., Alonso J.M. A link between ethylene and auxin uncovered by the characterization of two root-specific ethylene-insensitive mutants in Arabidopsis. Plant Cell. 2005;17(8):2230-2242. DOI 10.1105/tpc.105.033365Swarup R., Perry P., Hagenbeek D., Van Der Straeten D., Beemster G.T.S., Sandberg G., Bhalerao R., Ljung K., Bennett M.J. Ethylene upregulates auxin biosynthesis in Arabidopsis seedlings to enhance inhibition of root cell elongation. Plant Cell. 2007;19(7):2186-2196. DOI: 10.1105/tpc.107.052100Stepanova A.N., Yun J., Likhacheva A.V., Alonso J.M. Multilevel interactions between ethylene and auxin in Arabidopsis roots. Plant Cell. 2007;19(7):2169-2185. DOI 10.1105/tpc.107.052068Takatsuka H., Umeda M. Hormonal control of cell division and elongation along differentiation trajectories in roots. J. Exp. Bot. 2014;65(10):2633-2643. DOI: 10.1093/jxb/ert485Swarup R., Perry P., Hagenbeek D., Van Der Straeten D., Beemster G.T.S., Sandberg G., Bhalerao R., Ljung K., Bennett M.J. Ethylene upregulates auxin biosynthesis in Arabidopsis seedlings to enhance inhibition of root cell elongation. Plant Cell. 2007;19(7): 2186-2196. DOI 10.1105/tpc.107.052100Tsuchisaka A., Theologis A. Unique and overlapping expression patterns among the Arabidopsis 1-amino-cyclopropane-1-carboxylate synthase gene family members. Plant Physiol. 2004;136(2):2982-3000. DOI:10.1104/pp.104.049999Takatsuka H., Umeda M. Hormonal control of cell division and elongation along differentiation trajectories in roots. J. Exp. Bot. 2014;65(10):2633-2643. DOI 10.1093/jxb/ert485Tsuchisaka A., Yu G., Jin H., Alonso J.M., Ecker J.R., Zhang X., Gao S., Theologis A. A combinatorial interplay among the 1-aminocyclopropane-1-carboxylate isoforms regulates ethylene biosynthesis in Arabidopsis thaliana. Genetics. 2009;183(3):979-1003. DOI: 10.1534/genetics.109.107102Tsuchisaka A., Theologis A. Unique and overlapping expression patterns among the Arabidopsis 1-amino-cyclopropane-1-carboxylate synthase gene family members. Plant Physiol. 2004;136(2):2982-3000. DOI 10.1104/pp.104.049999Van de Poel B., Bulens I., Hertog M.L., Nicolai B.M., Geeraerd A.H. A transcriptomics-based kinetic model for ethylene biosynthesis in tomato (Solanum lycopersicum) fruit: development, validation and exploration of novel regulatory mechanisms. New Phytol. 2014;202(3):952-963. DOI: 10.1111/nph.12685Tsuchisaka A., Yu G., Jin H., Alonso J.M., Ecker J.R., Zhang X., Gao S., Theologis A. A combinatorial interplay among the 1-aminocyclopropane- 1-carboxylate isoforms regulates ethylene biosynthesis in Arabidopsis thaliana. Genetics. 2009;183(3):979- 1003. DOI 10.1534/genetics.109.107102Van de Poel B., Bulens I., Markoula A., Hertog M.L.A.T.M., Dreesen R., Wirtz M., Vandoninck S., Oppermann Y., Keulemans J., Hell R., Waelkens E., De Proft M.P., Sauter M., Nicolai B.M., Geeraerd A.H. Targeted systems biology profiling of tomato fruit reveals coordination of the Yang Cycle and a distinct regulation of ethylene biosynthesis during postclimacteric ripening. Plant Physiol. 2012;160(3):1498-1514. DOI: 10.1104/pp.112.206086Van de Poel B., Bulens I., Hertog M.L., Nicolai B.M., Geeraerd A.H. A transcriptomics-based kinetic model for ethylene biosynthesis in tomato (Solanum lycopersicum) fruit: development, validation and exploration of novel regulatory mechanisms. New Phytol. 2014; 202(3):952-963. DOI 10.1111/nph.12685Van de Poel B., Van Der Straeten D. 1-aminocyclopropane-1-carboxylic acid (ACC) in plants: more than just the precursor of ethylene! Front. Plant Sci. 2014;5:640. DOI: 10.3389/fpls.2014.00640Van de Poel B., Bulens I., Markoula A., Hertog M.L.A.T.M., Dreesen R., Wirtz M., Vandoninck S., Oppermann Y., Keulemans J., Hell R., Waelkens E., De Proft M.P., Sauter M., Nicolai B.M., Geeraerd A.H. Targeted systems biology profiling of tomato fruit reveals coordination of the Yang Cycle and a distinct regulation of ethylene biosynthesis during postclimacteric ripening. Plant Physiol. 2012; 160(3):1498-1514. DOI 10.1104/pp.112.206086Voß U., Bishopp A., Farcot E., Bennett M.J. Modelling hormonal response and development. Trends Plant Sci. 2014;19(5):311-319. DOI: 10.1016/j.tplants.2014.02.004Van de Poel B., Van Der Straeten D. 1-aminocyclopropane-1-carboxylic acid (ACC) in plants: more than just the precursor of ethylene! Front. Plant Sci. 2014;5:640. DOI 10.3389/fpls.2014.00640Vogel J.P., Woeste K.E., Theologis A., Kieber J.J. Recessive and dominant mutations in the ethylene biosynthetic gene ACS5 of Arabidopsis confer cytokinin insensitivity and ethylene overproduction, respectively. Proc. Natl. Acad. Sci. USA. 1998;95(8):4766-4771.Voß U., Bishopp A., Farcot E., Bennett M.J. Modelling hormonal response and development. Trends Plant Sci. 2014;19(5):311-319. DOI 10.1016/j.tplants.2014.02.004Vrebalov J., Ruezinsky D., Padmanabhan V., White R., Medrano D., Drake R., Schuch W., Giovannoni J. A MADS-box gene necessary for fruit ripening at the Tomato Ripening-Inhibitor (Rin) locus. Science. 2002;296:343-346. DOI: 10.1126/science.1068181Vogel J.P., Woeste K.E., Theologis A., Kieber J.J. Recessive and dominant mutations in the ethylene biosynthetic gene ACS5 of Arabidopsis confer cytokinin insensitivity and ethylene overproduction, respectively. Proc. Natl. Acad. Sci. USA. 1998;95(8):4766-4771.Wang K.L.-C., Yoshida H., Lurin C., Ecker J.R. Regulation of ethylene gas biosynthesis by the Arabidopsis ETO1 protein. Nature. 2004;428(6986):945-950.Vrebalov J., Ruezinsky D., Padmanabhan V., White R., Medrano D., Drake R., Schuch W., Giovannoni J. A MADS-box gene necessary for fruit ripening at the tomato ripening-inhibitor (rin) locus. Science. 2002;296:343-346. DOI 10.1126/science.1068181Xiong L., Xiao D., Xu X., Guo Z., Wang N.N. The non-catalytic Nterminal domain of ACS7 is involved in the post-translational regulation of this gene in Arabidopsis. J. Exp. Bot. 2014;65(15):4397-4408. DOI: 10.1093/jxb/eru211Wang K.L.-C., Yoshida H., Lurin C., Ecker J.R. Regulation of ethylene gas biosynthesis by the Arabidopsis ETO1 protein. Nature. 2004; 428(6986):945-950.Yamagami T., Tsuchisaka A., Yamada K., Haddon W.F., Harden L.A., Theologis A. Biochemical diversity among the 1-amino-cyclopropane-1-carboxylate synthase isozymes encoded by the Arabidopsis gene family. J. Biol. Chem. 2003;278(49):49102-49112.Xiong L., Xiao D., Xu X., Guo Z., Wang N.N. The non-catalytic Nterminal domain of ACS7 is involved in the post-translational regulation of this gene in Arabidopsis. J. Exp. Bot. 2014;65(15):4397-4408. DOI 10.1093/jxb/eru211Yang S.F., Hoffman N.E. Ethylene biosynthesis and its regulation in higher-plants. Annu. Rev. Plant Physiol. Mol. Biol. 1984;35:155–189. DOI: 10.1146/annurev.pp.35.060184.001103Yamagami T., Tsuchisaka A., Yamada K., Haddon W.F., Harden L.A., Theologis A. Biochemical diversity among the 1-amino-cyclopropane- 1-carboxylate synthase isozymes encoded by the Arabidopsis gene family. J. Biol. Chem. 2003;278(49):49102- 49112.Zarei A., Korbes A.P., Younessi P., Montiel G., Champion A., Memelink J. Two GCC boxes and AP2/ERF-domain transcription factor ORA59 in jasmonate/ethylene-mediated activation of the PDF1.2 promoter in Arabidopsis. Plant Mol. Biol. 2011;75(4-5):321-331. DOI: 10.1007/s11103-010-9728-yYang S.F., Hoffman N.E. Ethylene biosynthesis and its regulation in higher-plants. Annu. Rev. Plant Physiol. Mol. Biol. 1984;35:155- 189. DOI 10.1146/annurev.pp.35.060184.001103Zhang J., Yu J., Wen C.K. An alternate route of ethylene receptor signaling. Front. Plant Sci. 2014a;5:648. DOI: 10.3389/fpls.2014.00648Zarei A., Korbes A.P., Younessi P., Montiel G., Champion A., Memelink J. Two GCC boxes and AP2/ERF-domain transcription factor ORA59 in jasmonate/ethylene-mediated activation of the PDF1.2 promoter in Arabidopsis. Plant Mol. Biol. 2011;75(4-5):321- 331. DOI 10.1007/s11103-010-9728-yZhang X., Zhu Z., An F., Hao D., Li P., Song J., Yi C., Guo H. Jasmonate-activated MYC2 represses ETHYLENE INSENSITIVE3 activity to antagonize ethylene-promoted apical hook formation in Arabidopsis. Plant Cell. 2014b;26(3):1105-1117. DOI: 10.1105/tpc.113.122002Zhang J., Yu J., Wen C.K. An alternate route of ethylene receptor signaling. Front. Plant Sci. 2014a;5:648. DOI 10.3389/fpls.2014.00648Zhao Q., Guo H.W. Paradigms and paradox in the ethylene signaling pathway and interaction network. Mol. Plant. 2011;4(4):626-634. DOI: 10.1093/mp/ssr042Zhang X., Zhu Z., An F., Hao D., Li P., Song J., Yi C., Guo H. Jasmonate-activated MYC2 represses ETHYLENE INSENSITIVE3 activity to antagonize ethylene-promoted apical hook formation in Arabidopsis. Plant Cell. 2014b;26(3):1105-1117. DOI 10.1105/tpc.113.122002Zhao R., Xie H., Lv S., Zheng Y., Yu M., Shen L., Sheng J. LeMAPK4 participated in cold-induced ethylene production in tomato fruit. J. Sci. Food Agric. 2013;93(5):1003-1009. DOI: 10.1002/jsfa.5790Zhao Q., Guo H.W. Paradigms and paradox in the ethylene signaling pathway and interaction network. Mol. Plant. 2011;4(4):626-634. DOI 10.1093/mp/ssr042Zhu Z., An F., Feng Y., Li P., Xue L., Mu A., Jiang Z., Kim J.M., To T.K., Li W., Zhang X., Yu Q., Dong Z., Chen W.Q., Seki M., Zhou J.M., Guo H. Derepression of ethylene-stabilized transcription factors (EIN3/EIL1) mediates jasmonate and ethylene signaling synergy in Arabidopsis. Proc. Natl. Acad. Sci. USA. 2011;108(30):12539-12544. DOI: 10.1073/pnas.1103959108Zhao R., Xie H., Lv S., Zheng Y., Yu M., Shen L., Sheng J. LeMAPK4 participated in cold-induced ethylene production in tomato fruit. J. Sci. Food Agric. 2013;93(5):1003-1009. DOI 10.1002/jsfa.5790Zhu Z., Lee B. Friends or Foes: New insights in jasmonate and ethylene co-actions. Plant Cell Physiol. 2015;56(3):414-420.Zhu Z., An F., Feng Y., Li P., Xue L., Mu A., Jiang Z., Kim J.M., To T.K., Li W., Zhang X., Yu Q., Dong Z., Chen W.Q., Seki M., Zhou J.M., Guo H. Derepression of ethylene-stabilized transcription factors (EIN3/EIL1) mediates jasmonate and ethylene signaling synergy in Arabidopsis. Proc. Natl. Acad. Sci. USA. 2011;108(30):12539-12544. DOI 10.1073/pnas.1103959108Zhu Z., Lee B. Friends or Foes: New insights in jasmonate and ethylene co-actions. Plant Cell Physiol. 2015;56(3):414-420.Zhu Z., Lee B. Friends or Foes: New insights in jasmonate and ethylene co-actions. Plant Cell Physiol. 2015;56(3):414-420.

The authors declare that there are no conflicts of interest present.