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Characterization of Monilinia Linhartiana Isolates

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Characterization of Monilinia Linhartiana Isolates Journal of Plant Pathology (2017), 99 (3), 583-591 Edizioni ETS Pisa, 2017 583 CHARACTERIZATION OF MONILINIA LINHARTIANA ISOLATES A. Lantos1, M. Petróczy1, R. Oláh2 and L. Palkovics1 1Faculty of Horticultural Science, Department of Plant Pathology, Szent István University, 44 Ménesi Road, Budapest H-1118, Hungary 2Faculty of Horticultural Science, Department of Entomology, Szent István University, 44 Ménesi Road, Budapest H-1118, Hungary SUMMARY as reported from the Mediterranean region (Altinyay, 1972; Moral et al., 2008; Nevado et al., 2011). Monilinia linhartiana is a significant fungal pathogen of M. linhartiana (Prill. and Delacr.) N.F. Buchw. (ana- quince causing leaf blotch, shoot blight and mummifica- morph: Monilia cydoniae Schellenb.) was first described tion of developing young fruits. In our study, five isolates by the Hungarian botanist and mycologist György Lin- originating from infected quince shoots were character- hart, and was named after him (Wormald, 1926). In Hun- ized. Morphological characteristics were determined be- gary, the disease has been observed since the 1930s (Glits, fore performing molecular assays. Three genomic regions 2000), although the pathogen has rarely appeared in recent were sequenced and analysed: the ITS region, a polymor- decades. Nowadays the total area of quince orchards is phic region and the β-tubulin gene. The ITS region (ITS1- steadily increasing in Hungary, thus the pathogen showed 5.8 rDNA-ITS2) sequences of different Monilinia species up again causing infection more frequently. In Turkey, the showed high similarity, thus M. linhartiana isolates could fungus was studied in detail in the 1970s (Altinyay, 1972) not be safely separated from M. aucupariae isolates. By con- and it is still present nowadays (Ünlü and Boyraz, 2010), verse, based on the 28S rDNA sequences, M. linhartiana but no further studies have been conducted. Recent stud- isolates could be distinguished from the closely related ies were only carried out in Spain by Moral et al. (2011) species, offering prospects for the unambiguous identifi- who examined the pathogen by molecular methods, and cation of Monilinia species. The 28S rDNA and β-tubulin reconsidered the existing plant protection strategies. genes of M. linhartiana were sequenced for the first time. M. linhartiana colonises leaves, shoots and young fruits of quince (Naidenov, 1915), overwintering in pseudoscle- Keywords: beta-tubulin, disjunctors, ITS, Monilinia lin- rotial or mycelial form in twigs (Altinyay, 1972). Two in- hartiana, quince, 28S rDNA. fection cycles can be distinguished during the vegetation period. Ascospores as primary inoculum infect the young leaves and the mycelium spreads along the shoots. The secondary inoculum develops on the surface of primary infected leaves, and infects the flowers or young fruits. In early spring, apothecia with ascospores develop on stromatized fruit mummies (pseudosclerotia). The life INTRODUCTION cycle of M. linhartiana is closely related to the host plant phenology (Moral et al., 2011), as the first apothecia ap- Quinces (Cydonia oblonga) have been grown for centu- pear at the bud break stage of quince. Ascospores infect ries in Europe, where summers are sufficiently hot for the the young leaves, causing foliar necrosis along the primary fruit to fully ripen, Turkey being the world’s leading pro- veins starting from the base of the leaf. Wormald (1926) ducer (Topcu et al., 2015). Shoot blight and fruit rot caused observed that the infection spreads rapidly; the fungus by Monilinia spp. are significant diseases of quince, which penetrates through the petioles into the axis of the shoots requires chemical treatments every year. Brown rot symp- and blights them. Conidial coating appears on the necro- toms caused by Monilinia fructigena or Monilinia fructicola, tised tissues (Sumstine, 1913). Conidia induce the second- occasionally by Monilinia laxa or Monilinia polystroma, ap- ary infection and attack the young fruits. The infected pear on mature fruits (Chalkley, 2015). Monilinia linhar- fruits shrivel on the shoots or fall to the ground, but co- tiana causes leaf blotch, shoot blight and mummification nidia never develop on their surfaces (Batra, 1991). These of young fruits. However, the disease caused by M. lin- infected fruits turn into pseudosclerotial structures. hartiana is one of the most destructive diseases of quince M. linhartiana belongs to the section Disjunctoriae of worldwide (Batra, 1991). Yield losses could reach 90-95%, the genus Monilinia. Conidial chains contain small modi- fied conidia called disjunctors, which can be found be- tween macroconidia (Honey, 1936). The species of the sec- Corresponding author: L. Palkovics Fax: + 3613057324 tion are monotrophic, inhabiting a single host, or are oli- E-mail: [email protected] gotrophic for they attack several species of one host genus 584 Characterization of M. linhartiana isolates Journal of Plant Pathology (2017), 99 (3), 583-591 (Batra, 1991). In the section Junctoriae, the macroconidia sucrose, 5.0 g peptone, 15.0 g agar powder, 10.0 g malt are joined directly, like the conidial chains of M. laxa or extract/litre) media. Mycelial plugs (5 mm in diameter) M. fructicola. These pathogens are polytrophics, infecting from single-spore cultures were transferred to the center several genera of a family. The identification and classifi- of the agar plates, and incubated in the dark at 24°C. cation of the species is feasible based on host specificity, Colour, structure, growth rate, pigmentation, conidia morphology of stromata and conidial chains, shape and production and growth rate (mm/24 h) of the isolates size of the conidia, and cultural characteristics on cultiva- were determined. tion media (Batra, 1991). However, closely related species cannot be differentiated in all cases by morphology and Pathogenicity tests. The pathogenicity of M. linhartiana cultural characteristics (van Leeuwen and van Kesteren, isolates was tested on young leaves, young fruits (approx. 1998), whereas reliable identification can be obtained by 3-4 cm in diameter) and fruits of Cydonia oblonga. Before PCR-based methods (De Cal and Melgarejo, 1999). inoculation, the surface of leaves and fruits was disinfected Molecular studies of the internal transcribed spacer with 70% ethanol for 1 min, then washed in sterile water. (ITS) region are generally applied for the identification of The petioles of the leaves and the fruit peels were pricked Ascomycota fungi, and are also appropriate for phylogenet- by a sterile needle, then mycelial plugs (3 mm in diam- ic and evolutionary studies (Capote et al., 2012). Carbone eter) were placed into the punctures. For control, sterile and Kohn (1993) examined the ITS region of Sclerotiniace- PDA plugs were used. Five pieces of each organ (shoots ae, separating the sections Junctoriae and Disjunctoriae of and fruits) per isolate were inoculated. On each shoot two the genus Monilinia. Several brown rot species have been leaves were inoculated, while in the case of young and ma- characterized by the ITS1-5.8S rDNA-ITS2 region (Holst- ture fruits only one mycelial plug was inserted per fruit. Jensen et al., 1997; Fulton et al., 1999; Ioos and Frey, 2000). Inoculated materials were placed into large sterile glass Holst-Jensen et al. (1997) found cospeciation between host vessels on glass beads and incubated under natural light and pathogen. Species of the section Junctoriae infect rosa- at 22-25°C for 3 weeks in 95 ± 2 RH. ceous hosts with stone or pome fruits. Four evolutionary lineages were found in the section Disjunctoriae: one group Molecular investigations. DNA was extracted using the of species on dry stone fruits of rosaceous hosts, one group cetyl-trimethyl-ammonium-bromide (CTAB) method (Ma- of species on capsular fruits of ericaceous hosts, one group niatis et al., 1989), followed by phenol-chloroform extrac- of species on sweet berry fruits of ericaceous hosts, and tion and isoamyl alcohol (24:1) precipitation carried out one parasite species of Crataegus sp. Moral et al. (2011) according to a protocol described by Gell et al. (2007b). studied the ITS region of M. linhartiana, and the phyloge- Three genomic regions were examined by PCR: the ITS netic analyses demonstrated that the pathogen was closely region and the 28S rDNA gene (primers ITS5 and NL4), related to Monilinia mali and Monilinia amelanchieris and the β-tubulin gene (primers BT-forw and BT-rev), and a other Monilinia species infecting dry stone fruits of rosa- polymorphic region (primers UniMon-F and UniMon-R) ceous hosts. (Table 2). The BT primer pair was designed using the data of McKay et al. (1998). Each PCR reaction mixture con- tained 100 ng of fungal DNA, 1 mM of both primers, and MATERIALS AND METHODS Dream Taq Green PCR Master Mix 2× (Thermo Scien- tific, Lithuania) diluted to a final volume of 50 μl. During Sample collection and isolation. Infected shoots show- PCR reactions, the initial denaturation step (94°C, 3 min) ing typical symptoms were collected in different locations was followed by 35 cycles as shown in Table 2. Final elon- across Hungary in 2013 and 2014, and in Turkey in 2015 gation was 10 min at 72°C. PCR products were separated from orchards and gardens (Table 1). Conidia from infect- in 1% GelRed (Biotium Inc. USA) stained agarose gel, ed leaves were transferred onto Petri dishes containing and purified using the High Pure PCR Product Purifi- potato dextrose agar (PDA, BioLab Zrt., Hungary), and cation Kit (Roche, Germany) according to the manufac- individual conidia were separated and placed onto culture turer’s protocol. Final DNA concentrations were verified medium under an inverted microscope. Petri dishes were by spectrophotometry (NanoDrop 2000). Fragments were incubated in the dark at 24°C for 10 days. sequenced in an ABI Prism automatic sequencer (Base- Clear B.V., Leiden, The Netherlands) in three replicates. Morphological and cultural characteristics. The Nucleotide sequence identities and protein sequence simi- symptoms visible on the infected parts and the appear- larities were determined by BLAST analyses (Altschul et ance of conidial chains were observed, and the length al., 1999).
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