Antimicrobial defences in the wood of living trees

Abstract

summary 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 O2 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 multiccllular walls OT 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 gymnospenns and angiosperms are generally similar. Defence responses against a variety of pathogen categories also have 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.).

Pattern of delignification inAilanthus excelsaRoxb. wood byInonotus hispidus(Bull.: Fr.) Karst.

“…Tyloses are an outgrowth of adjacent ray or axial parenchyma cells into the lumen which block the vessel partially or completely. It is a common feature and reported in several earlier studies (Shain 1967(Shain , 1979Pearce 1996).…”

Controls on Coarse Wood Decay in Temperate Tree Species: Birth of the LOGLIFE Experiment

“…do not. Wood outer bark forms a formidable physical (and chemical) barrier to penetration, which can considerably retard decomposer penetration (Käärik 1974;Pearce 1996). However, this protection only prevails as long as wood moisture content stays constant as drying out of the outer wood and bark tissue will lead to crack formation and entrance ports for decomposers.…”

“…These factors explain the relatively high decomposition resistance of heartwood in comparison to sapwood, although due to the nature of organic substances in different species also heartwood resistance varies substantially (Hillis 1987;Eriksson et al 1990). Heartwood formation is a genetically determined process that occurs in many broad-leaved and coniferous species and indicates the transformation of partly living tissue, which is actively involved in water transport and storage, into dead tissue, which is made impermeable and impregnated with-often toxic-organic substances (Panshin and de Zeeuw 1980;Bamber and Fukazawa 1985;Pearce 1996;Taylor et al 2002;Cornwell et al 2009). …”

Identification of hydrolysable tannins in the reaction zone of Eucalyptus nitens wood by high performance liquid chromatography–electrospray ionisation mass spectrometry

“…Reaction zones are a major component of active antimicrobial defence in the secondary xylem of woody plants (Shain, 1979;Pearce 1996). The reaction zone forms at the interface between living sapwood and fungal infection, and is able to restrict or slow fungal decay.…”