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Identification of Acremonium Isolates from Grapevines and Evaluation of Their Antagonism Towards Plasmopara Viticola

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Identification of Acremonium Isolates from Grapevines and Evaluation of Their Antagonism Towards Plasmopara Viticola Identification of Acremonium isolates from grapevines and evaluation of their antagonism towards Plasmopara viticola Sandra Lo Piccolo, Antonio Alfonzo, Selene Giambra, Gaetano Conigliaro, Luis V. Lopez-Llorca & Santella Burruano Annals of Microbiology ISSN 1590-4261 Ann Microbiol DOI 10.1007/s13213-015-1082-5 1 23 Your article is protected by copyright and all rights are held exclusively by Springer- Verlag Berlin Heidelberg and the University of Milan. This e-offprint is for personal use only and shall not be self-archived in electronic repositories. If you wish to self- archive your article, please use the accepted manuscript version for posting on your own website. You may further deposit the accepted manuscript version in any repository, provided it is only made publicly available 12 months after official publication or later and provided acknowledgement is given to the original source of publication and a link is inserted to the published article on Springer's website. The link must be accompanied by the following text: "The final publication is available at link.springer.com”. 1 23 Author's personal copy Ann Microbiol DOI 10.1007/s13213-015-1082-5 ORIGINAL ARTICLE Identification of Acremonium isolates from grapevines and evaluation of their antagonism towards Plasmopara viticola Sandra Lo Piccolo1 & Antonio Alfonzo1 & Selene Giambra1 & Gaetano Conigliaro1 & Luis V. Lopez-Llorca2 & Santella Burruano 1 Received: 8 July 2014 /Accepted: 1 April 2015 # Springer-Verlag Berlin Heidelberg and the University of Milan 2015 Abstract Some endophytic fungal genera in Vitis vinifera, Keywords Fungal endophytes . Phylogeny . RAPD . including Acremonium,havebeenreportedasantagonistsof Inhibition . Sporangia germination . Vitis vinifera Plasmopara viticola.EndophyticAcremonium isolates from an asymptomatic grapevine cultivar Inzolia from Italy were identified by morphological features and multigene phyloge- Introduction nies of ITS, 18S and 28S genes, and their intra-specific geno- mic diversity was analyzed by RAPD analysis. Culture fil- Fungal endophytes, both clavicipitaceous and non- trates (CFs) obtained from Acremonium isolates were tested clavicipitaceous groups, colonize plant tissues without caus- in vitro for their inhibitory activity against the P. viticola ing visible disease symptoms and have a profound impact on sporangia germination. Among 94 isolates, 68 belonged to plant communities (Petrini 1991;Arnoldetal.2003; Schulz the Acremonium persicinum and26totheAcremonium and Boyle 2006; Sanchez Marquez et al. 2007; Hyde and sclerotigenum. RAPD analysis grouped the A. persicinum iso- Soytong 2008; Rodriguez et al. 2009). These microorganisms lates into 15 clusters and defined 31 different strains. The display a remarkable diversity, in fact, more than 100 endo- A. sclerotigenum isolates, instead, were clustered into 22 phytic taxa have been detected in some plant species (Stone groups and represented 25 strains. All A. persicinum CFs et al. 2004). Endophytic fungi can also show host, organ and inhibited sporangia germination of P. viticola,whilenotall tissue specificity (Chapela et al. 1991; Bettucci et al. 1997; those of A. sclerotigenum had inhibitory effect. A different Mostert et al. 2000; Ragazzi et al. 2004; Schulz and Boyle degree of inhibition was observed between strains of the same 2006;Peršoh et al. 2010;Yuanetal.2011). species, while some strains of different species showed iden- Several studies have shown the ability of fungal endo- tical inhibitory effect. No correlation was found between phytes to promote growth of their host plants (Rahman and RAPD groups and inhibitory activity in both Acremonium Saiga 2005). Moreover, they can also protect their hosts from species. herbivores, insects and pathogens (Leuchtmann et al. 2000; Arnold and Lewis 2005;Rubinietal.2005; Schulz and Boyle 2005). Endophytic microorganisms may play a key role in host-pathogen interactions even prior to the triggering of the disease. In particular, some endophytes can induce systemic * Selene Giambra resistance mechanisms in their hosts, as well as the expression [email protected] of defence genes against some pathogens (Gwinn and Gavin 1992;Arnoldetal.2003). In addition, endophytic fungi may produce secondary metabolites active against pathogens, con- 1 Dipartimento di Scienze Agrarie e Forestali, University of Palermo, Viale delle Scienze 4, Palermo 90128, Italy stituting an important potential source of biocontrol (Li et al. 2000; Tan and Zou 2001; Schulz et al. 2002; Kongue Tatong 2 Department of Marine Sciences and Applied Biology Multidisciplinary Institute for Environmental Studies (MIES) et al. 2014). BRamón Margalef^, University of Alicante, Aptdo. 99, Alican- Several authors have investigated the diversity and ecolog- te 03080, Spain ical role of endophytic fungal communities in Vitis vinifera L. Author's personal copy Ann Microbiol (von Tiedemann et al. 1988; Mostert et al. 2000; Casieri et al. DNA extraction and PCR amplification 2009; González and Tello 2010). Among these communities, the genera Acremonium, Alternaria, Epicoccum and Single-conidial cultures were also used for DNA extraction. Fusarium have been shown to be effective as biocontrol Genomic DNA was extracted from these pure cultures using a agents of Plasmopara viticola (Berk. & Curtis) Berl. & De standard CTAB-based protocol (O’Donnell et al. 1998). Toni (Falk et al. 1996; Kortekamp 1997; Bakshi et al. 2001; The internal transcribed spacer (ITS) regions, ITS1 and Assante et al. 2005; Musetti et al. 2006;Burruanoetal.2008). ITS2, and the 5.8S gene of the ribosomal DNA (rDNA) oper- In this regard, Acremonium sp. strain A20, isolated from a leaf on, were amplified using primers ITS1F (fungal specific: 5’- of asymptomatic grapevine cultivar Inzolia and identified CTTGGTCATTTAGAGGAAGTAA-3’) (Gardes and Bruns wrongly as Acremonium byssoides W. Gams & G. Lim, has 1993) and ITS4 (universal: 5’-TCCTCCGCTTATTGATAT been reported as hyperparasite of P.viticola gamic and agamic GC-3’)(Whiteetal.1990). The amplification reaction was structures (Burruano et al. 2008) and producer of secondary performed in a total reaction volume of 40 μlcontaining50– metabolites, named acremines (A-N), that inhibit sporangia 100 ng of DNA template, 2 mM of MgCl2, 0.2 mM of each germination (Assante et al. 2005;Arnoneetal.2008, 2009). dNTP, 0.3 μM of each primer, 0.5 U of Dream Taq The aim of the present study was to identify a collection of (Fermentas, Milan, Italy) and 1× Dream Taq buffer endophytic Acremonium isolates from grapevine cv. Inzolia, (Fermentas). The amplification reaction was carried out in to analyze the genetic variability among these isolates, and to T1 Thermocycler (Biometra, Göttingen, Germany) under the evaluate their antagonistic activity towards P. viticola following conditions: initial denaturation at 95 °C for 2 min, sporangia. 35 cycles of denaturation at 95 °C for 30 s, annealing at 55 °C for 20 s, extension at 72 °C for 1 min and final extension at 72 °C for 5 min. To amplify the 18S rRNA gene (18S) and D1/D2 domains Materials and methods of the 28S of the nuclear rRNA gene (28S), the primer pairs NS1 (5’-GTAGTCATATGCTTGTCTC-3’)/NS2 (5’-GGCT Fungal isolates GCTGGCACCAGACTTGC-3’) and NL1 (5’-GCATATCA ATAAGCGGAGGAAAAG-3’)/NL4 (5’-GGTCCGTGTT A total of 94 endophytic fungal isolates belonging to the TCAAGACGG-3’)(Whiteetal.1990;O’Donnell 1993)were Acremonium genus, collected in 2008 from used, respectively. The reaction volume (50 μl) contained asymptomatic grapevine cv. Inzolia located in southern Italy 50 ng of DNA template, 2 mM of MgCl2,0.25mMof (province of Palermo, Sicily), were used in this study dNTPs, 0.2 μM of each primer, 1.5 U of Dream Taq (Burruano et al. 2008). Particularly, 58 isolates were from (Fermentas) and 1× Dream Taq buffer (Fermentas). The am- leaves, 20 from buds, 11 from shoots, four from petioles and plification programs consisted of 94 °C for 8 min; 35 cycles of one from seed (Table 1). All fungi were maintained on 2 % 94 °C for 40 s, 48 °C for 40 s, 72 °C for 45 s; 72 °C for 5 min Malt Extract Agar (MEA; Oxoid, Milan, Italy) at 28 °C±1 °C. for18rRNAgene,andof94°Cfor3min;35cyclesof94°C for 1 min, 50 °C for 30 s, 72 °C for 1 min; 72 °C for 10 min for the 28S rRNA gene. PCR products were analyzed by electro- Fungal identification and genetic variability phoresis on 1.5 % (w/v) agarose gel in 1× TBE (89 mM Tris, 89 mM boric acid, 2 mM EDTA, pH 8.3) buffer. Gels were Morphological characterization stained with SYBR Safe DNA gel stain (Invitrogen, Milan, Italy), visualized by UV transilluminator and acquired by a Single-conidial cultures of each isolate were obtained in 2 % Gel Doc 1000 Video Gel Documentation System (BioRad, MEA plates incubated at 28 °C±1 °C, alternating light and Richmond, V USA). Standard molecular markers were 1 kb darkness (12 h of each), for 7–14 days up to 1 month. Plus DNA Ladder (Invitrogen) and GeneRuler 100 bp Plus Identification at the species level of isolates was performed DNA Ladder (Fermentas). on the basis of their macroscopic (morphology, colour and growth rate of colony) and microscopic (size, col- Sequencing and phylogenetic analysis our, shape of conidia and phialides) features in culture, by comparison of above criteria with those given by PCR products were sequenced in both directions with the Gams (1971) and Domsch et al. (1980, 2007). same primers used for amplification reported in the previous Microscopic characteristics were examined using a light paragraph. Nucleotide sequences were compared to GenBank microscope (Axioskop, Zeiss, Germany), coupled to an sequences through BLASTn searches (Altschul et al. 1997). AxioCam MRc5 (Zeiss) digital camera. Images were Sequences were read and edited by using the Sequencher soft- captured using the software AxioVision 4.6 (Zeiss). ware (Version 4.7, Gene Codes Corporation, Ann Arbor, MI), Author's personal copy Ann Microbiol Table 1 Isolation tissue and nucleotide sequences deposited in GenBank of endophytic Acremonium isolates from grapevine cv. Inzolia and their inhibitory activity against the Plasmopara viticola sporangia germination Strain code Species Tissue GenBank Accession no.
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