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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 climacteric fruits do not increase respiration at ripening and often have no requirement for ethylene to complete maturation. So, climacteric fruits such as tomato are distinguished from non-climacteric by their increased ethylene biosynthesis rates during ripening. This is one of the main reasons that the majority of biochemical research has concentrated on this hormone. Initial molecular studies focused on the isolation of ethylene-regulated genes which include those encoding the ethylene biosynthesis enzymes (Sadenosylmethionine, SAM-synthase, 1-aminocyclopropane carboxylic acid, ACC-synthase and ACC oxidase) (reviewed by Redgwell and Fischer, 2002 Giovanini et al., 2001). It was later demonstrated, using the reverse genetic approach, that delaying ethylene production constituted a successful strategy to extend the shelf-life of fruits (Grierson, 1992). In 2002, Klee group proposed a model for ethylene perception and metabolism. As the receptor also acts as a negative regulator of downstream responses, in the absence of ethylene, receptors actively suppress expression of ethylene responsive genes. Recently, several further ethylene inducible genes have been identified in tomato, including mitochondrial translation elongation factors (Benichou et al., 2003) and CTR-1 (Leclercq et al., 2002). It seems likely, given the development of micro-array resources for tomato that significant advances will be made in our understanding of ethylene signal transduction and its role in ripening fruit. Examples of this have already been started by using tomato cDNA microarray containing 12000 unique elements encoding 9000 genes covering a range of metabolic and developmental processes (http://bti.cornell.edu/CGEP/CGEP.html) (Fei et al., 2004; Baxter et al., 2005). More recently, a joined effort between four countries (France, Spain, Italy and USA) led to the construction of an oligo-based new generation of tomato chips. A complete list of 70 mer long oligonucleotides was first derived from 27000 individual clones composing the unigene collection of tomato ESTs. Thereafter, 12000 clones were selected for printing in the first version of oligo DNA-chips named EU-TOM1. A Laboratory Information Management Systems (LIMS) for microarrays production and traceability including the set up of a dedicated tomato microarrays database providing access to the gene ID of the EU-TOM1 clones has been installed by the French partner 46

ChapitreI: Bibliographic review (http://bioinfo.genopole-toulouse.prd.fr/eusol/base/). The printing of the second set of tomato oligo-based DNA chips (EU-TOM2) has been scheduled within the EU-SOL program for 2007 and will include the remaining tomato unigenes estimated at 15000 clones. VIII.7.a Revealing the secrets of ripening mutant Tomato has proved to be an excellent model system for the analysis of fruit ripening and development, in part due to the availability of well characterized ripening mutants. These include pleiotropic ripening mutations, such as Colorless nonripening (Cnr), ripening-inhibitor (rin), Never-ripe (Nr) and Green-ripe (Gr) (Figure 33). WT Nr Gr Cnr rin (Giovannoni., 2007) Figure 33. Normal and mutant tomato fruit. Normal tomato cultivar Ailsa Craig ripe fruit ten days post breaker and equivalent age fruit homozygous for the Never-ripe (Nr/Nr), Green-ripe (Gr/Gr), Colorless non-ripening (Cnr/Cnr) and ripening-inhibitor (rin/rin) mutations. Recent cloning of this tomato ripening genes, that were previously known only through mutation, have created new inroads into understanding of the primary ripening control mechanisms, including transcription factors such as those encoded by the RIPENING-INHIBITOR (RIN) MADS-box and COLOURLESS NON-RIPENING (CNR) SPB-box genes, which are necessary for the progression of virtually all ripening processes. These discoveries have also facilitated the elucidation of downstream signal transduction components that impact the hormonal and environmental stimuli that coordinate and modulate ripening phenotypes (Figure 34). Ripening mutant 47

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