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All parts of the bodies of multicellular organisms have to be able to respond to wounding and, in animals, this process involves cells that play specific roles in the healing process. Some of these cells actually migrate into wounded tissues from the surrounding tissues. In contrast, cell migration does not occur in plants and it is not known whether discrete cell types in the leaf have specialised roles in wound perception and in the production of wound hormones such as jasmonates. Jasmonates accumulate rapidly in the vicinity of wounds (in a timescale of tens of seconds) but little is known about where jasmonate synthesis takes place and how this is regulated. In which cell types are jasmonates made and which cells need to be wounded to initiate their synthesis? Do the abilities to generate jasmonate-synthesis activating signals and to make JA-Ile reside in all cells within the plant body, or do these events take place in discrete or even in exclusive cell types? This proposal focusses on these spatial aspects in the intiation of jasmonate synthesis using jasmonoyl-isoleucine (JA-Ile) as a marker. JA-Ile quantitation will be coupled to laser-targetted wounding to localise the areas in Arabidopsis seedlings that must be wounded in order to intitate JA-Ile accumulation 30 min after wounding. JAZ10-based reporters will be used as additional readouts for these experiments. Making use of a knowledge of JA-Ile accumulation in seedlings and the importance of parastichies in leaf-to-leaf wound signalling in rosettes we will employ promoter-specific rescue of the jasmonate biosynthesis gene ALLENE OXIDE SYNTHASE (AOS) in an aos mutant background. Based on results from this we will produce sector mosaic plants by selectively ablating AOS expression domains, again using tissue-specific promoters. Finally, using knowledge gained from an earlier part of the project on seedlings we will initiate a novel genetic screen to look for mutants with defective spatial responses to wounding. The proposed work should reveal to what extent different cell types in plants cooperate to counter the threat posed by wounding and whether these cell types originate from the same meristematic layers.Questions: 1. Which tissue types in 5 d-old seedlings need to be wounded in order to initiate JA-Ile and JA accumulation and JAZ10 expression?2. What is the timeframe of the long-distance signal(s) leading to JA-Ile accumulation in distal organs of 4 d-old seedlings after wounding one cotyledon?3. Which cell types can produce JA-Ile?4. Can mutants with altered spatial JAZ10 expression patterns be identified in wounded seedlings.