Jasmonates in plant stress responses and development Claus Wasternack Halle/Saale, Germany May 2010 Leibniz Institute of Plant Biochemistry Past: Department of Natural Product Biotechnology, Presence: Molecular Signal Processing

pollen maturation flower development germination pollen maturation flower development fruit / seed development cell elongation leaf senescence root growth pathogens (virus, bacteria fungi) herbivores attraction of predators of parasitoides by release of volatiles osmotic stress/drought mycorrhiza pathogens (virus, bacteria fungi)

0 1 2 3 4 d H2O JM Jasmonate-induced senescence in Hordeum vulgare

Feussner and Wasternack, Ann Rev. Plant Biol. 2002 Localization of enzymes of jasmonate biosynthesis tomato barley barley Feussner and Wasternack, Ann Rev. Plant Biol. 2002

4-CL-like CoA synthetases activate JA precursors in peroxisomes for ß-oxidation of the carboxylic acid side chain after Kienow et al., J. Exp. Bot. 59: 403-419 (2008), Collab. E. Kombrink (-)-JA

w-7-fatty acid desaturase Mutants Gene product Phenotype JA biosynthesis dgl Galactolipase A1 Reduced JA level in leaves dad1 Phospholipase A1 Reduced filament elongation, male sterile, JA-deficient in flowers spr21 w-7-fatty acid desaturase Deficient in a-LeA and JA levels, no wound response dde2-2 AOS Male sterile, delayed anther development, opr3 OPR3 JA-deficient, reduced filament elongation, male sterile acx11 Acyl-CoA oxidase JA-deficient reduced wound response comatose COMATOSE/PXA1, ABC transporter Reduced JA content Zhai et al. 2007 Constitutive JA response cev1 constitutive expression of vegetative storage proteins hy1-101 Heme oxygenase HY1 Increased JA level, stunted growth, phytochrome chromophore deficiency joe2 ? Reduced inhibition of root growth, increased expression of LOX2 Others   ore9, max2 F-box protein Delayed leaf senescence, more axillary branches Reduced sensitivity to JA coi1 F-box protein COI1/ JA receptor Reduced root growth inhibition, male sterile, reduced filament elongation, enhanced sensitivity to necrotrophic pathogens jai11 Tomato homologue of COI1 Female sterile, decreased wound-response jar1/jin4/jai2 JA amino acid conjugate synthase Reduced root growth inhibition by JA jin1/jai1 AtMYC2 (bHLHzip transcription factor) Reduced root growth inhibition axr1 RUB-activating enzyme jai4/sgt1b AtSGT1b Reduced root growth inhibition in the ein3 background  1tomato mutants

Jasmonate biosynthesis in Arabidopsis COOH 1 13-LOX AOS AOCs Arabidopsis a-linolenic acid 13-LOX 13-HPOT COOH HOO 13 AOS COOH O AOC3 AOC4 AOC1 AOC2 COOH O 12 cis-(+)-12-oxo-phytodienoic acid [OPDA] OPR3 b-oxidation COOH 3 7 O COOH O coi1 Gene expression (+)-7-iso-JA (-)-JA Leibniz Institute of Plant Biochemistry Department of Natural Product Biotechnology 7

There is more than JA Wasternack, in H. Hirt “Plant Stress Biology“, 2009

Levels of 12-OH-JA, 12-HSO4-JA, JA and OPDA differ dramatically pmole per g f.w. OPDA JA 12-OH-JA 12-HSO4-JA 12-O-Glc-JA Z. mays dry seeds 254 82 502 992 112 G. max immat. seeds 601 259 71,050 34,655 112,753 V. faba pericarp 9,773 9,286 69 645 3,937 C. pepo seeds 184 20 n.d. 218 1,257 Z. mays d1 leaves 2,528 127 320 117,461 n.d. S. esculentum cv. Lukullus leaves 550 20 51 872 584 n.d.: not detectable; errors in the range of 2-10% Miersch et al., New Phytol., 177: 114-127 (2008)

JA signaling is switched off for a subset of genes by hydroxylation and sulfation

Jasmonate biosynthesis in tomato COOH 1 a-linolenic acid 13-LOX 13-HPOT COOH HOO 13 13-LOX anti-AOC AOS COOH O GUS-staining AOS AOC COOH O 12 Ovule-specific AOC expression, preferential accumulation of jasmonates in the pistil AOC cis-(+)-12-oxo-phytodienoic acid [OPDA] OPR3 Leaf cross section b-oxidation COOH 3 7 O COOH O Gene expression Is there any role of oxylipins in embryo development ? (+)-7-iso-JA (-)-JA jai1 11

Which JA compound is the ligand of the JA receptor ?

5, 344-350 (2009) Collaboration between R. Solano´s group (Madrid), M. Hamberg (Stockholm) and the Halle group

is most active in pulldown experiments yeast two hybrid interactions root growth assays - anthocyanin accumulation Epimerization as a mechanism activating a pre-existing pool of inactive JA compounds ?