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Functional characterization of tomato Sl-IAA3 and Sl-hls genes. Role ...

Functional characterization of tomato Sl-IAA3 and Sl-hls genes. Role ...

ChapitreI: Bibliographic

ChapitreI: Bibliographic review receptors causes inactivation of a Raf-like kinase CTR1 and the consequent derepression of EIN2, a protein of unknown biochemical function that is essential for the ethylene response. Downstream of EIN2, a family of transcription factors composed of EIN3 and EIN3-like proteins triggers a transcriptional cascade that results in the activation/repression of hundreds of target genes (Alonso and Stepanova., 2004; Guo and Ecker., 2004) (Figure 22). The first step in the cascade initiated by EIN3/EILs involves other transcription factors, such as the AP2/EREBP family members ERF1 (Solano et al., 1998) and EDF1-4 (Alonso et al., 2003). These different EIN3 targets probably represent branching points in the ethylene response that can be turned on or off not only by ethylene but also by other factors and therefore could be used to provide specificity in response to ethylene. In the case of ethylene and auxin, several common target genes have (Bishop et al., 2006) Figure 22. Ethylene signalling: At low concentrations of the ligand, ethylene receptors (ETR1, ERS1, ETR2, EIN4, and ERS2) are active and can therefore stimulate the negative regulator CTR1, which in turn shuts down ethylene signalling by allowing EIN3 degradation. Ethylene binding inactivates the receptors and therefore they are unable to stimulate CTR1-mediated inhibition. As a result, EIN2 is active and prevents EIN3 degradation, which leads to EIN3 accumulation and activation of ethylene responsive gene transcription. 32

ChapitreI: Bibliographic review also been identified (Zhong and Burns., 2003). However it’s still unclear if whether or not they participate in common biological processes and related regulation. roots VII.6.a Multilevel interaction between auxin and ethylene in the Despite the accumulated knowledge on each individual hormone signaling and response pathway, very little is known about the mechanisms that govern the interactions between ethylene and auxin. The ethylene-mediated regulation of auxin biosynthesis through the activation of WEI2/ASA1 and WEI7/ASB1, a anthranilase synthase subunits that catalyse the fist step in tryptophane biosynthesis (Figure 2) (Stepanova et al., 2005) as well as the reciprocal effect of auxin on ethylene biosynthesis through the activation of several ACC synthases, represent two elegant examples of the molecular mechanisms controlling the ethylene–auxin crosstalk. Recently, for better understand the molecular mechanisms behind the ethylene–auxin interactions in the roots, a comprehensive study relying on physiological, cellular, genetic and genomic approaches was performed by Stepanova et al., 2007. Quantification of the morphological effects of ethylene and auxin in a variety of mutant backgrounds indicates that auxin biosynthesis, transport, signalling and response are required for the ethylene-induced growth inhibition in roots. This analysis suggests a simple mechanistic model for the interaction between these two hormones in roots, according to which ethylene and auxin can reciprocally regulate each other’s biosynthesis, influence each other’s response pathways and/or act independently on the same target genes. This model not only implies existence of several levels of interaction but also provides a likely explanation for the strong ethylene response defects observed in auxin mutants (Figure 23) VII.6.b Interaction between auxin and ethylene in the hook The induction of apical hook formation in Arabidopsis represents one of the best described examples of auxin-ethylene cross-talk in plants (Lehman et al., 33

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