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Pathogen Profile View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Academica-e MOLECULAR PLANT PATHOLOGY (2012) 13(9), 998–1009 DOI: 10.1111/j.1364-3703.2012.00816.x Pathogen profile Pseudomonas savastanoi pv. savastanoi: Some like it knot Cayo Ramos1, Isabel M. Matas1, Leire Bardaji2, Isabel M. Aragón1, Jesús Murillo2* 1 Instituto de Hortofruticultura Subtropical y Mediterránea “La Mayora”, Universidad de Málaga-Consejo Superior de Investigaciones Científicas (IHSM-UMA-CSIC), Área de Genética, Facultad de Ciencias, Campus Teatinos s/n, E-29010 Málaga, Spain 2 Departamento de Producción Agraria, ETS Ingenieros Agrónomos, Universidad Pública de Navarra, 31006 Pamplona, Spain * Corresponding author: Jesús Murillo; [email protected] Key words: Pseudomonas syringae pv. savastanoi, olive knot, tumour, genome sequencing, plant disease, control, pathogenicity, virulence, effector. SUMMARY molecular biology that has elevated the foliar pathogens of the P. syringae complex to Pseudomonas savastanoi pv. savastanoi is the supermodels. A series of studies in the last years causal agent of olive (Olea europaea) knot have made significant advances in the biology, disease and an unorthodox member of the P. ecology and genetics of P. savastanoi pv. syringae complex, causing aerial tumours instead savastanoi, paving the way for the molecular of the foliar necroses and cankers characteristic dissection of its interaction with other non- of most members of this complex. Olive knot is pathogenic bacteria and their woody hosts. The present wherever olive is grown; although losses selection of a genetically pliable model strain was are difficult to assess, it is assumed that olive knot soon followed by the development of rapid is one of the most important diseases of the olive methods for virulence assessment with crop. The last century has witnessed a good deal micropropagated olive plants and the analysis of of scientific articles describing the biology, cellular interactions with the plant host. The epidemiology and control of this pathogen. generation of a draft genome of strain NCPPB However, most P. savastanoi pv. savastanoi 3335 and the closed sequence of its three native strains are highly recalcitrant to genetic plasmids has allowed for functional and manipulation, which has effectively left the comparative genomic analyses for the pathogen out of the scientific progress in identification of its pathogenicity gene Pseudomonas savastanoi pv. savastanoi complement. This includes 34 putative type III knots) on the stems and branches of the host effector genes and genomic regions, shared with plant and, occasionally, on leaves and fruits. other pathogens of woody hosts, that encode Epidemiology: The pathogen can survive and metabolic pathways associated with the multiply on aerial plant surfaces, as well as in degradation of lignin-derived compounds. Now, knots, from where it can be dispersed by rain, the time is right to explore the molecular basis of wind, insects and human activities, entering the the P. savastanoi pv. savastanoi-olive interaction plant through wounds. Populations are very and to get insights into why some pathovars like it unevenly distributed in the plant, and suffer necrotic and why some like it knot. drastic fluctuations throughout the year, with maximum numbers of bacteria occurring during Synonyms: Pseudomonas syringae pv. rainy and warm months. Populations of P. savastanoi savastanoi pv. savastanoi are normally Taxonomy: Kingdom Bacteria; Phylum associated to non-pathogenic bacteria, both Proteobacteria; Class Gammaproteobacteria; epiphytically and endophytically, and were Family Pseudomonadaceae; Genus demonstrated to form mutualistic consortia with Pseudomonas; included in genomospecies 2 Erwinia toletana and Pantoea agglomerans that together with at least P. amygdali, P. ficuserectae, could result in increased bacterial populations and P. meliae and 16 other pathovars from the P. disease symptoms. syringae complex (aesculi, ciccaronei, Disease control: Based on preventive measures, dendropanacis, eriobotryae, glycinea, hibisci, mostly sanitary and cultural practices. Integrated mellea, mori, myricae, phaseolicola, photiniae, control programs benefit from regular applications sesami, tabaci, ulmi, and certain strains of of copper formulations, which should be lachrymans and morsprunorum); when a formal maintained at least a few years for maximum proposal is made for the unification of these benefit. Olive cultivars vary in their susceptibility bacteria, the species name P. amygdali would to olive knot, but there are no known cultivars with take priority over P. savastanoi. full resistance to the pathogen. Microbiological properties: Gram-negative rods, Useful websites: http://www.pseudomonas- 0.4-0.8 by 1.0-3.0 µm, aerobic. Motile by one to syringae.org/; http://genome.ppws.vt.edu/cgi- four polar flagella, rather slow growing, optimal bin/MLST/home.pl; ASAP access to the P. temperatures for growth of 25–30 °C, oxidase savastanoi pv. savastanoi NCPPB 3335 genome negative, arginine dihydrolase negative, elicits the sequence hypersensitive response on tobacco, most https://asap.ahabs.wisc.edu/asap/logon.php. isolates are fluorescent and levan negative although some isolates are non-fluorescent and levan positive. Host range: P. savastanoi pv. savastanoi causes INTRODUCTION tumours in cultivated and wild olive and ash (Fraxinus excelsior). Although strains from olive Pseudomonas syringae is an economically were reported to infect oleander (Nerium important pathogen and one of the most relevant oleander), this is generally not the case; however, models for the study of plant-microbe interactions strains of P. savastanoi pv. nerii can infect olive. (e.g., Mansfield, 2009, Mansfield et al., 2012). Pathovars fraxini and nerii differentiate from pv. The species is currently a taxonomic conundrum savastanoi mostly in their host range, and were and has been pulled together with P. amygdali, P. not formally recognized until 1996. Literature avellanae, P. cannabina, P. caricapapayae, P. previous to about 1996 generally name strains of ficuserectae, P. meliae, P. savastanoi, P. tremae the three pathovars as P. syringae subsp. and P. viridiflava into a group designated as the savastanoi or P. savastanoi subsp. savastanoi, P. syringae complex, which could correspond to contributing to confusion about host range and at least nine different species (Gardan et al., biological properties. 1999, Young, 2010, Parkinson et al., 2011). Disease symptoms: Symptoms of infected trees Pathovars of the P. syringae complex include hyperplastic growths (tumorous galls or generally exploit the plant apoplast as a parasitic C. Ramos et al. niche and cause foliar necrosis in diverse plant the P. syringae complex, encompassing at least hosts, with a minority of strains causing other 60 pathovars and several other Pseudomonas types of symptoms, such as vascular diseases on species (Bull et al., 2010, Young, 2010). A study woody plants (Agrios, 2005). A remarkable limited to a few taxa, formally classified pathovars exception are a few pathovars producing aerial glycinea, phaseolicola and savastanoi into the tumours in woody plants, including P. savastanoi new species P. savastanoi (Gardan et al., 1992), pv. savastanoi. P. savastanoi pv. savastanoi is to which pathovars fraxini, nerii and retacarpa the causal agent of olive (Olea europaea) knot were later added (Bull et al., 2010). DNA-DNA disease, whose symptoms include hyperplastic hybridization distributed P. syringae into at least growths (tumorous galls or knots) on the stems nine separate genomospecies (Gardan et al., and branches of the host plant and, occasionally, 1999, Young, 2010). P. savastanoi pv. savastanoi on leaves and fruits (Fig. 1). Olive knot is present was included in genomospecies 2, together with 16 worldwide, wherever olive is grown, and it is other P. savastanoi-P. syringae pathovars (see considered one of the most important diseases of Taxonomy, above) and the species P. amygdali, P. olive (CMI, 1987, Young, 2004, Quesada et al., ficuserectae and P. meliae; when genomospecies 2 2010a). Diverse research groups worldwide have is formally named, however, the species should be made substantial contributions towards designated Pseudomonas amygdali and not P. understanding the biology, epidemiology and savastanoi (Gardan et al., 1999). Multilocus control of this pathogen; however, most strains of sequence analyses show that P. savastanoi pv. P. savastanoi pv. savastanoi are highly recalcitrant savastanoi NCPPB 3335 is evolutionarily closer to to genetic manipulation (Pérez-Martínez et al., P. syringae pathovars aesculi 2250 and NCPPB 2007), which has significantly slowed down their 3681, tabaci ATCC 11528 and phaseolicola 1448A molecular analysis. (genomospecies 2) than to P. syringae pv. tomato The growing availability of microbial genomes DC3000 (genomospecies 3) or P. syringae pv. has spurred a new research era in the field of syringae B728a (genomospecies 1) (Fig. S1) plant-microbe interactions, leading to the (Sarkar & Guttman, 2004, Parkinson et al., 2011). identification of potentially comprehensive These studies support the genomospecies 2 repertoires of putative virulence genes and the grouping and indicate that it might encompass at emergence of unified models of interaction least 9 further pathovars (broussonetiae, between prototypical pathogens and plant hosts castaneae, cerasicola, cunninghamiae, (Lindeberg et al., 2008, Mansfield,
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