Distinct developmental defense activations in barley embryos identified by transcriptome profiling

Proper embryo development is crucial for normal growth and development of barley. Numerous related aspects of this process--for example how the embryo establishes and sustains disease resistance for extended periods during dormancy--remain largely unknown. Here we report the results of microarray analyses of >22,000 genes, which together with measurements of jasmonic acid and salicylic acid during embryo development provide new information on the initiation in the developing barley embryo of at least two distinct types of developmental defense activation (DDA). Early DDA is characterized by the up-regulation of a specific set of genes around 20 days after flowering, including co-regulation of those for encoding 9-lipoxygenase and several oxylipin-generating enzymes, possibly leading to the formation of α-ketols. The same developmental phase includes an up-regulation of several defense genes, and indications of co-regulation of those for enzymes involved in the generation of phenylpropanoid phytoalexins. Late DDA is initiated prior to grain desiccation, around 37 days after flowering, with up-regulation of several genes encoding proteins with roles in antioxidant responses as well as a simultaneous up-regulation of several PR genes is notable. Throughout barley embryo development, there are no indications of an increased biosynthesis of either jasmonic acid or salicylic acid. Collectively, the results help explain how the proposed DDA enables protection of the developing barley embryo and grain for purposes of disease resistance.