[논문]Penetration of Picea sitchensis root bark by Armillaria mellea, Armillaria ostoyae and Heterobasidion annosum

SOLLA, A.; TOMLINSON, F.; WOODWARD, S.

doi:10.1046/j.1439-0329.2002.00265.x

[해외논문] Penetration of Picea sitchensis root bark by Armillaria mellea, Armillaria ostoyae and Heterobasidion annosum

Forest pathology = Journal de pathologie forestière = Zeitschrift für Forstpathologie, v.32 no.1, 2002년, pp.55 - 70

SOLLA, A. , TOMLINSON, F. , WOODWARD, S.

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Penetration of root bark tissues of Picea sitchensis by Armillaria ostoyae, Armillaria mellea and Heterobasidion annosum was examined in the absence of wounds, in superficial wounds (rhytidome tissues removed to expose the secondary phloem) and in wounds to the depth of the vascular cambium (deep wounding). Both species of Armillaria penetrated bark without prior wounding, but neither species formed rhizomorphs in this treatment. Armillaria ostoyae penetrated to 39 cell layers in depth by 48 days after inoculation of unwounded bark, whereas A. mellea penetrated 25 cell layers in the same time. Armillaria mellea penetrated superficial wounds significantly more rapidly than did A. ostoyae. Both species produced rhizomorphs within wounded host tissues. Inoculation of deep wounds with Armillaria resulted in a greater depth of bark necrosis with A. mellea than with A. ostoyae. In the absence of wounding, H. annosum failed to penetrate root bark tissues, but in both superficial and deep wounds hyphae penetrated beyond the ligno–suberized boundary zone (LSZ) by 12 days after inoculation. Where no inoculations were made, superficial or deep wounding led within 25 days to the restoration of a structurally continuous LSZ, and by day 48 the wound periderm (WP) was fully differentiated. In inoculated wounds, however, formation of the LSZ and WP was delayed or inhibited in most trees, particularly following inoculation with A. ostoyae or A. mellea. Suberization in the LSZ and WP remained diffuse and discontinuous 48 days after inoculation. Moreover, the presence of WP did not prevent further penetration of the tissues by the pathogens. Variations between trees in the depth of pathogen penetration were noted, possibly indicating differing susceptibilities of individual host genotypes. The possible host factors involved in resistance to penetration of root bark tissues by Armillaria and Heterobasidion are discussed.

Abstract

Penetration der Wurzelrinde von Picea sitchensis durch Armillaria mellea, Armillaria ostoyae und Heterobasidion annosumDie Penetration der Wurzelrindengewebe von Picea sitchensis durch A. ostoyae, A. mellea und H. annosum wurde unter folgenden Bedingungen untersucht: ohne Verletzung, mit oberflächlicher Verletzung (sekundäres Phloem exponiert) und tiefe Verletzung (bis zum vasculären Kambium). Beide Armillaria-Arten durchdrangen die unverletzte Rinde, aber sie bildeten keine Rhizomorphen. Nach Inokulation der unverletzten Rinde drang A. ostoyae in 48 Tagen 39 Zellschichten tief ein, A.mellea in derselben Zeit 25 Zellschichten tief. Armillaria mellea durchwuchs das Gewebe nach oberflächlicher Verletzung signifikant schneller als A. ostoyae. Beide Arten bildeten im verletzten Wirtsgewebe Rhizomorphen. Bei Inokulation von tiefen Wunden waren die Rindennekrosen bei A. mellea grösser als bei A. ostoyae. Heterobasidionannosum konnte unverletztes Rindengewebe nicht durchdringen, nach oberflächlicher und tiefer Verletzung durchwuchs das Myzel jedoch die lignifizierte und suberinisierte Abgrenzungszone (LSZ) 12 Tage nach der Inokulation. Bei oberflächlicher oder tiefer Verletzung ohne Inokulation war innerhalb von 25 Tagen eine strukturell zusammenhängende LSZ vorhanden, und nach 48 Tagen war das Wundperiderm (WP) voll ausdifferenziert. Nach der Inokulation war die Bildung der LSZ und des Wundperiderms in den meisten Bäumen jedoch verzögert oder gehemmt, insbesondere nach einer Inokulation mit A. ostoyae oder A. mellea. Die Suberinisierung in der LSZ und dem WP blieb hier 48 Tage nach der Inokulation diffus und unzusammenhängend. Zudem verhinderte ein vorhandenes Wundperiderm das weitere Vordringen der Pathogene nicht. Es gab Unterschiede zwischen den Bäumen in der Tiefe der Ausbreitung der Pathogene, was möglicherweise auf unterschiedliche Anfälligkeit individueller Wirtsgenotypen hinweist. Mögliche Resistenzfaktoren auf Seiten des Wirtes bei der Penetration des Rindengewebes durch Armillaria und Heterobasidion werden diskutiert.

Abstract

Pénétration de l'écorce des racines de Picea sitchensis par Armillaria mellea, Armillaria ostoyae et Heterobasidion annosumLa pénétration d'Armillaria ostoyae, A. mellea et Heterobasidion annosum dans les tissus corticaux racinaires de Picea sitchensis a étéétudiée. Trois cas ont été examinés: sans blessure, avec blessure superficielle (rhythidome enlevé jusqu'au phloème secondaire) et avec blessure allant jusqu'au cambium vascularisé (blessure profonde). Les deux espèces d'Armillaire pénétraient l'écorce sans blessure préalable, mais aucune d'elles n'avait formé de rhizomorphes. Armillaria ostoyae avait traversé jusqu'à 39 couches de cellules 48 jours après l'inoculation sans blessure alors que A. mellea en avait traversé 25. Armillaria mellea pénétrait les blessures superficielles significativement plus rapidement que A. ostoyae. Les deux espèces produisaient des rhizomorphes à l'intérieur des tissus blessés. L'inoculation des blessures profondes avec les Armillaires aboutissait à une nécrose d'écorce, plus profonde avec A. mellea qu'avec A. ostoyae. En l'absence de blessure, H. annosum ne parvenait pas à pénétrer l'écorce, mais sur blessures superficielles ou profondes, les hyphes, en 12 jours, pénétraient au-delà de la zone limite ligno-subérisée (LSZ). En l'absence d'inoculation, chez les blessures superficielles ou profondes il y avait en 25 jours restauration d'une structure LSZ continue, et en 48 jours le périderme de blessure (WP) était complètement différencié. Chez les blessures inoculées, cependant, la formation de LSZ et de WP était retardée ou inhibée chez la plupart des arbres, surtout à les suite de l'inoculation par A. ostoyae et A. mellea. La subérisation des structures LSZ et WP demeurait diffuse et discontinue 48 jours après l'inoculation. De plus, la présence de WP n'empêchait pas les parasites de continuer de pénétrer dans les tissus. Des différences de profondeur de pénétration ont été observées entre les arbres, ce qui peut indiquer des différences de sensibilité entre génotypes. Les facteurs possibles de l'hôte, impliqués dans la résistance à la pénétration d'Armillaria et d'Heterobasidion dans les tissus corticaux racinaires sont discutés.

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