Adecatrieonic acid (), are converted into their hydroperoxide types by way of oxygenation by precise lipoxygenases (LOXs).The resulting (S)hydroxyperoxyoctadecadi(tri)enoic acid (HPOT) and (S)hydroperoxyhexadeca(tri)enoic acid (HPHT) in their turn form a sizable assortment of oxylipins, including JA, by way of no less than six alternative pathways .The two JA precursors now comply with two parallel pathways; the octadecanoid pathway from HPOT plus the hexadecanoid pathway from HPHT .The first step is performed by allene oxide synthases (AOS) that catalyzes dehydrations to type the unstable allene oxides ,(S)epoxyoctadecaenoic acid (,EOT) and ,(S)epoxyhexadeca(tri)enoic acid (,EHT).In aqueous media, ,EOT undergoes cyclisation to kind cisoxophytodienoic acid (cisOPDA), a reaction mediated by allene oxide cyclase (AOC).Four stereoisomers of OPDA may well be formed, but only S,SOPDA is actually a precursor for biologically active JA.The carbon homologue dinorOPDA (dnOPDA) is generated in the parallel pathway from ,EHT .OPDA and dnOPDA are then transported into the peroxisomes, via a mechanism nevertheless unresolved.The Arabidopsis ATPbinding cassette (ABC) transporter COMATOSE (CTSPXAPED) has been showed to catalyze the ATPdependent import of fatty acids into peroxisomes as substrates for oxidation .However, other pathways for dnOPDA will have to exist, as knockout mutants lack JAdeficiency symptoms (for instance male sterility).After within the peroxisomes, S,SOPDA is lowered by oxophytodienoate reductase (OPR) to yield oxo(‘(Z)pentenyl)cyclopentaneoctanoic acid (OPC), and dnOPDA is lowered for the corresponding hexanoic acid derivative (OPC) .OPC and OPC are then activated through CoA esterification of the carboxylic moiety assisted by OPC CoA ligase (OPCL) , plus a still unknown ligase for OPC.The hexanoic and octanoic side chains of OPC and OPC are shortened by two or three rounds of oxidation.The oxidation entails three core enzymes; acylCoA oxidase (ACX), a multifunctional protein (MFP, comprising enoylCoA hydratase and hydroxyacylCoA dehydrogenase activities) and ketoacylCoA thiolase (KAT) forming JACoA.The last biosynthetic step could be the release of the JACoA ester from JA, that is catalyzed by an acylthioesterase (ACH), forming the reactive epijasmonic acid that effortlessly epimerize towards the extra steady epijasmonic acid .Upon the subsequent transport towards the cytoplasm, JA is further modified to methylepijasmonate (MeepiJA) through the assistance of a JA methyl transferase, a epijasmonylLisoleucine (epiJALIle) PubMed ID:http://www.ncbi.nlm.nih.gov/pubmed/21601637 catalyzed by a JA amido synthetase, or other derivatives .Int.J.Mol.SciFigure .Biosynthesis of jasmonic acid within the chloroplast and peroxisome.Polyunsaturated fatty acids ( and ) released in the cell, chloroplast andor thylakoid membrane are precursors for the biosynthesis of jasmonic acid (JA).Within the chloroplast, cisoxophytodienoic acid (cisOPDA) and dinorOPDA (dnOPDA) are formed via the octa and hexadecanoid pathways.Following transport into the peroxisome, OPDA (dnOPDA) is SPQ Epigenetic Reader Domain reduced to OPC (OPC) and undergoes three (two) cycles of oxidation that benefits inside the production of epijasmonic acid.The reactions are catalyzed by lipoxygenases (LOX), allene oxide synthase (AOS), allene oxide cyclase (AOC), ATPbinding cassette (ABC) transporter COMATOSE (CTSPXAPED), oxophytodienoate reductase (OPR), OPC CoA ligase (OPCL), acylthioesterase (ACH), ketoacylCoA thiolase , acylCoA oxidase (ACX) and a multifunctional protein (MFP).Enzymes are shown in blue.Arrows show the effectively characterized reactions.