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Esca De La Vigne Et Communauté Fongique Schweizerische Eidgenossenschaft Station de recherche Agroscope Changins-Wädenswil ACW Confédération suisse Directeur: Jean-Philippe Mayor • www.acw.admin.ch Confederazione Svizzera Confederaziun svizra Esca de la vigne et communauté fongique V. HOFSTETTER, L. CASIERI, O. VIRET et K. GINDRO, Station de recherche Agroscope Changins-Wädenswil ACW, CP 1012, 1260 Nyon E-mail: [email protected] @ Tél. (+41) 22 36 34 374. Résumé Les études s’intéressant aux champignons associés au bois de vigne se limitent généralement aux parties nécrotiques de ceps atteints d’esca. Ces champignons sont nombreux et appartiennent à des groupes systé - matiques très divers. Les espèces fongiques considérées comme res - ponsables de l’esca ont également été isolées de jeunes plantes. Ces ré - sultats suggèrent que l’esca pourrait être due à un déséquilibre de la communauté fongique normalement associée à la vigne. Pour étudier cette hypothèse, la communauté fongique a été isolée de ceps apoplec - tiques et de boutures saines de Chasselas. Les isolats fongiques ont été identifiés sur la base de leur morphologie, de leur vitesse de croissance, de leur expression enzymatique et de leur séquence ITS, puis classés phylogénétiquement par analyse combinée de quatre gènes. Les com - munautés fongiques isolées de bois atteints d’esca et sains sont très semblables et toutes deux dominées par les mêmes groupes de champi - gnons, à savoir les Ascomycètes, principalement de la classe des Sorda - riomycetes (ordre des Hypocreales et Xylariales ), mais les espèces qui les composent diffèrent partiellement. Cette analyse a permis de déterminer l’emplacement phylogénétique, jusqu’à présent incertain, de Phaeomo - niella chlamydospora, Oidiodendron truncatum et Trichothecium roseum dans les Ascomycètes. Ces résultats laissent entrevoir des perspectives de diagnostic moléculaire pour le suivi de l’évolution des communautés fongiques du bois de vigne. Introduction groupes de champignons suivant les ré - gions géographiques: Cephalosporium L’esca est la principale maladie du bois spp., Eutypa lata, Phialophora spp., de vigne ( Vitis vinifera L.), responsable Acremonium spp., Botryosphaeria spp., d’importantes pertes économiques dans Phomopsis viticola et Fomitiporia medi - le monde entier. Les symptômes exter- terranea (Armengol et al. , 2001; Crous nes de l’esca apparaissent généralement et al. , 1996; Fourie et al. , 2000; Rum - au cours de l’été sous deux formes dis - bos et Rumbou, 2001). Récemment, tinctes: une forme lente, caractérisée par Gimenez-Jaime et al. (2006) ont dé - une décoloration progressive des feuil- montré que les champignons impliqués les et une forme foudroyante, dite apo - dans la maladie de Petri, responsable plectique, qui tue la plante en quelques du dépérissement de jeunes plants de jours. Certains auteurs (Larignon et Du- vigne, étaient les mêmes que ceux qui bos, 1997) considèrent que l’esca est due engendrent l’esca (s’attaquant généra - à une succession fongique impliquant lement à des ceps plus âgés), mais avec des espèces pionnières, comme Phaeo - une prédominance de différentes es - moniella chlamydospora et Phaeoacre - pèces selon les régions géographiques. monium aleophilum , suivies d’un cor - Graniti et al. (2000) considèrent que la Forme lente (en haut) et apoplectique (en tège d’espèces appartenant à différents maladie de l’esca est causée par des bas) de l’esca de la vigne. Revue suisse Vitic. Arboric. Hortic. Vol. 41 (4): 247-253, 2009 247 groupes de champignons systématique - L’alignement obtenu (214 espèces/4 gènes) plus fréquemment isolées (tabl. 2), il est ment très divers qui agissent de concert a été utilisé pour placer certaines espèces, intéressant de noter que toutes possè - ou successivement. Cependant, les dont la position systématique était incer - dent des activités cellulolytiques, leur études sur l’esca se sont essentielle - taine (incertae sedis) , dans la phylogénie permettant de déstructurer les éléments des Ascomycètes. Les analyses ont été ment intéressées aux champignons iso - effectuées dans le programme PAUP*v.4 cellulosiques du bois (cellulose et hé - lés des nécroses du bois, alors que, (Swofford, 2001). Une analyse de ‘maxi - micellulose). Seulement un tiers de comme toutes les autres plantes terres - mum parsimony’ a été effectuée, compre - celles-ci sont capables de digérer tous tres, la vigne saine héberge de nom - nant 500 recherches heuristiques, avec les éléments du bois, dont la lignine, breux champignons endophytes, ce qui MAXTREE = ‘unlimited’ et ‘branch swap - causant par là des dommages irrépara - signifie qu’ils vivent dans une plante au ping’ = TBR. L’analyse mesurant la robus - bles à la structure et à l’intégrité du tesse des branches est basée sur 500 réplicats moins une partie de leur vie sans causer de ‘bootstrap ’, comprenant chacun 10 ‘heu - bois de vigne. Cette communauté fon - de maladie apparente (Halleen et al. , ristic searches’. Seules les branches de la gique est largement dominée par la li - 2001; Schweigkofler et Prillinger, phylogénie obtenue recevant 70% ou plus gnée des Ascomycota qui comprennent 1999). Ces derniers, très peu étudiés, de support de ‘bootstrap’ sont considérées 19 des 21 espèces de champignons le jouent probablement un rôle dans comme robustes. Des amorces permettant plus fréquemment isolées. Les lignées l’équilibre de la communauté fongique d’amplifier sélectivement les ordres et fa - Basidiomycota et Zygomycota sont associée à la vigne. Certains de ces en - milles dominant la communauté fongique chacune représentées par une seule es - associée à Vitis vinifera ont été développées. dophytes peuvent être des pathogènes pèce. Parmi les 19 Ascomycète les plus latents, aussi bien que des antagonistes fréquemment isolés, 3 espèces ont en - protecteurs. core une position systématique incer - Ce travail présente des résultats prélimi - Résultats et discussion taine: Phaeomoniella chlamydospora, naires sur les communautés fongiques Trichothecium roseum et Oidiodendron isolées de plantes apoplectiques et de Les 339 isolats fongiques obtenus en cul - truncatum . Une analyse phylogénétique jeunes plantes apparemment saines de ture pure représentent 42 espèces diffé - combinant les séquences obtenues pour Chasselas (porte-greffe 3309), toutes rentes de champignons. La figure 2 et 4 gènes indique que P. chlamydospora issues du même vignoble, puis compa - le tableau 2 illustrent la manière dont se place à la base de l’ordre des Pyre - rées afin de déterminer quels groupes ces espèces ont été caractérisées. Parmi nulales (fig. 3B) et n’appartient pas à la de champignons dominent la commu - ces espèces, 21 ont été isolées de plus famille des Herpotrichiellaceae placée nauté fongique présente dans la vigne de 5% des plantes et sont considérées dans l’ordre des Chaetothyriales (Crous avant et après l’apparition des symp - comme hôtes réguliers de la vigne et et Gams, 2000). Trichothecium roseum tômes de l’esca. non comme hôtes occasionnels (tabl.1). se place dans l’ordre des Hypocreales Si l’on considère les douze espèces le et Oidiodendron truncatum dans l’ordre Matériel et méthodes Les champignons présents dans 70 plantes apoplectiques de Chasselas (porte-greffe 3309) âgées de trente ans et dans 24 bar - bues saines produites à partir de greffons du même vignoble (Perroy, Vaud, Suisse) ont été isolés en cultures pures. Pour les ceps apoplectiques, six sections du tronc de 2,5 cm d’épaisseur ont été sciées stérilement à dif - férents niveaux (porte-greffe, point de greffe, greffon, tête de cep, zone supérieure de la tête de cep, branche à fruits; fig.1) puis la - vées à l’eau stérile. Quinze disques ont été prélevés de chaque section de bois, coupés en six morceaux et placés sur un milieu gé - losé (PDA + 12,5 mg/l d’auréomycine). Les plantes de pépinière ont été traitées selon Casieri et al. (2009). L’émergence des champignons, repiqués dès leur apparition sur milieu gélosé, a été suivie durant deux semaines. Les isolats fongiques ont été Fig. 1. Parties des ceps apoplec - identifiés d’après leur morphologie, leur ap - tiques analysées: titude à dégrader la lignine (Glenn et Gold, ቢ porte-greffe; 1983) et la cellulose (Smith, 1971), leur vi - ባ point de greffe; tesse de croissance et leur séquence d’ADN ቤ zone au-dessus du point de ribosomique ( Internal Transcribed Spacers greffe; [ITS]). Pour les espèces d’Ascomycètes ብ tête de cep; isolées à une fréquence ≥ 5% du bois des ቦ zone supérieure de la tête de plantes apoplectiques, quatre gènes ont été cep; amplifiés et séquencés (mitSSU, nucLSU, ቧ branche à fruits ( gauche : lo - RPB2, et tef-1). Les séquences obtenues calisation des parties 1-6 sur ont été combinées avec les séquences cor - le cep, droite : sections trans - respondantes obtenues par le projet «As - versales de 2,5 cm d’épais - sembling the Fungal Tree of Life» (AFTOL) seur correspondantes (1-6). pour 185 espèces de champignons (dispo- nibles sur le site: http://aftol.biology.duke. edu/pub/alignments/download_alignments). 248 Fig. 2. Morphotypes macro- et microscopiques de cultures pures des douze champignons le plus fréquemment isolés (≥17%) de troncs de vigne apoplectique (ordre décroissant de fréquence selon le tableau 1). A Cultures vues de des - sus; B Cultures vues de dessous; C-F Aspect des cultures sous la loupe binoculaire ou le microscope (marge de crois - sance mycélienne, mycélium et spores). Revue suisse Vitic. Arboric. Hortic. Vol. 41 (4): 247-253, 2009 249 Tableau 1. Taxonomie, classification et fréquence d’isolement (%) des
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