'Defence lignin' and hydroxycinnamyl alcohol dehydrogenase activities in wounded Eucalyptus gunnii

To learn more about lignin formation in response to wounding in trees, we adopted two complementary approaches: (1) microscopic and histochemical studies of the wound response in 3.5-month-old Eucalyptus gunnii plantlets and (2) biochemical investigations of hydroxycinnamyl alcohol dehydrogenase activities in wounded 6-year-old, field-grown E. gunnii trees. The first approach revealed that a barrier zone was formed in response to wounding in both ground tissues (cortex barrier and pith reaction zone) and vascular tissues. The barrier zone was barely detectable after 24 h but well-developed 7 days after wounding. Microscopic analyses indicated that the barrier zone was formed by the reinforcement of cell walls with 'lignin-like material' in both ground tissues and vascular tissue, and that, in addition, the lumen of certain xylem cells (vessels and fibres) were blocked by the deposition of polymeric phenolic material. Histochemical characterization revealed that the lignin-like material ('defence lignin') deposited in ground tissue cell walls and xylem cell blockages was poor in syringyl (S-type) lignin units and therefore differed from the usual mixed guaiacyl-syringyl (G-S) lignin unit composition of E. gunnii developmental lignin. In contrast, S-type lignin appeared to be deposited in the cell walls of immature developing secondary xylem cells at a stage when the cell walls of comparable cells from unwounded control stems contained lignin poor in syringyl units. The second approach indicated that two different types of cinnamyl alcohol dehydrogenase activity are induced, and apparently regulated differentially, in response to wounding in E. gunnii trees. Coniferyl alcohol dehydrogenase activity was induced immediately and continued to increase throughout the first 15 days of the 17-day experimental period, while sinapyl alcohol dehydrogenase activity was first detected at 8 days after wounding and continued to increase throughout the experimental period. The biological roles of the two alcohol dehydrogenase activities are discussed in relation to the formation of defence lignin versus developmental lignin in trees.