Antimicrobial defences in the wood of living trees

The wood (xylem) in a living tree is protected from microbial attack by the secondary plant surface (periderm and rhytidome), which provides an effective barrier preventing the entry of most potential pathogens, and by constitutive and induced defence mechanisms in the bark (cortex and phloem). Although a few pathogens are able to penetrate these outer tissues directly, most xylem pathogens gain entry through wounds that expose this tissue and render it more vulnerable to attack. In functional sapwood, microbial colonization might be restricted by active defence mechanisms, or by passive microenvironmental restriction consequent upon the high water content and low availability of O-2 in healthy conductive xylem. These factors are not mutually exclusive: indeed, they may operate in concert in many host-pathogen interactions. Sapwood lesions made by wood-decaying fungi are characteristically bounded by multicellular walls or barrier zones (compartmentalization wall 4 barriers and reaction zones (column boundary layers)), which may function both as inhibitory or degradation-resistant barriers to further pathogen spread, and as seals to maintain xylem function and prevent the drying and aeration that could predispose to further infection. A range of putative antimicrobial defence mechanisms contributing to the effectiveness of such barriers has been identified in sapwood tissues. This includes cell wall alterations, constitutive and induced antimicrobial compounds, necrotic responses of living cells and the deposition of gummy materials, often resinous or polyphenolic, at the host-pathogen interface. Nutritional, environmental and anatomical features of living wood might also contribute to pathogen restriction. Although there might be differences in detail, the defences operating in gymnosperms and angiosperms are generally similar. Defence responses against a variety of pathogen categories also hare much in common. Dynamic studies are crucial in elucidating the roles of the various components of the host-pathogen interaction in the wood of living trees. A model for the protection and defence of xylem tissues in woody angiosperms is suggested, based largely upon results from dynamic studies of host-pathogen interactions in the wood of sycamore (Acer pseudoplatanus L.).