Conifers dominate the boreal forests of the Northern Hemisphere, and especially members of the family Pinaceae have great economic and ecological significance. Part of their success is thought to arise from the vast array of secondary metabolites they produce. The products of secondary metabolism are essential for plants to survive in the ever-changing environment. In Scots pine (Pinus sylvestris L.), two groups of secondary metabolites, stilbenes and resin acids, are crucial for decay resistance of heartwood timber and for active defense responses against herbivores and fungal pathogens. Several studies have shown that stilbenes improve decay resistance of pine heartwood. Since there is wide variation in the concentration of stilbenes between individuals and the trait has high heritability, it may be possible to breed heartwood that is more decay-resistant. However, breeding for heartwood properties is slow, since the decay resistance characteristics can be estimated at the earliest from 30-year-old trees. Early selection methods utilizing genetic markers or chemical screening are needed, but we do not yet understand which genes control the biosynthesis of stilbenes and what the genetic differences are between individuals that explain the variation in the capacity to produce stilbenes. Importantly, there is genetic correlation between stress-induced stilbene biosynthesis in seedlings and the heartwood stilbene content in their adult mother trees. Here, we examined the pine transcriptional responses under two conditions that were previously known to activate stilbene biosynthesis: heartwood formation in adult trees and ultraviolet (UV)-C treatment of needles in seedlings. We found that these two conditions had very little in common, except for the activation of stilbene pathway genes. For example, the regulators of the two responses seemed not to be shared. The activation of the stilbene pathway in response to UV-C treatment occurred a few hours after the onset of the treatment and was independent of translation. Stilbene biosynthesis seems to be an early defense response in Scots pine. Heartwood formation, an important developmental process in the senescence of secondary xylem, is poorly understood. Based on transcriptomic analysis, stilbene biosynthesis occurs in situ in the transition zone between the sapwood and heartwood, but resin acids were synthesized primarily in the sapwood. Bifunctional nuclease, an enzyme involved in the process of developmentally programmed cell death (dPCD), is a useful marker for heartwood formation and aided us in defining the timing of the process, from spring to late autumn. Expression of this marker, which is strictly confined to dPCD conditions, further clarified that heartwood formation truly is a process that is initiated by intrinsic programming instead of environmental cues. The transcriptomic data revealed that the expression of the previously characterized pinosylvin O-methyltransferase gene, PMT1, was not induced under stilbene-forming conditions. A new PMT-encoding gene, PMT2, was identified by coexpression analysis. The gene showed an inducible expression pattern very similar to that of the stilbene synthase gene under all conditions studied. PMT2 furthermore methylated pinosylvin with high specificity, in contrast to PMT1, which accepted several substrates.
|Myöntöpäivämäärä||15 joulukuuta 2017|
|Tila||Julkaistu - 15 joulukuuta 2017|
|OKM-julkaisutyyppi||G5 Tohtorinväitöskirja (artikkeli)|
LisätietojaYhteenveto-osa 50 s. ja kolme eripainosta
- 414 Maatalouden bioteknologia
- 4112 Metsätiede