Invasive Species Compendium

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Pear blister canker viroid

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Datasheet

Pear blister canker viroid

Summary

  • Last modified
  • 23 January 2019
  • Datasheet Type(s)
  • Documented Species
  • Pest
  • Preferred Scientific Name
  • Pear blister canker viroid
  • Taxonomic Tree
  • Domain: Virus
  •   Unknown: Viroids
  •     Family: Pospiviroidae
  •       Genus: Apscaviroid
  •         Species: Pear blister canker viroid
  • Summary of Invasiveness
  • Pear blister canker disease, first reported in France in the 1960’s, is caused by pear blister canker viroid (PBCVd). Diseases can occur in several cultivars of pear (Pyrus communis), its major host, but most p...

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    Compendia
    CAB International
    Wallingford
    Oxfordshire
    OX10 8DE
    UK
    compend@cabi.org
  • Distribution map More information

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Identity

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Preferred Scientific Name

  • Pear blister canker viroid

International Common Names

  • English: Pear blister canker apscaviroid

Summary of Invasiveness

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Pear blister canker disease, first reported in France in the 1960’s, is caused by pear blister canker viroid (PBCVd). Diseases can occur in several cultivars of pear (Pyrus communis), its major host, but most pear cultivars are tolerant and do not produce symptoms of infection. PBCVd was initially reported in pear and quince, followed by wild pear and nashi and can experimentally infect species in other genera (Chaenomeles, Cydonia, Sorbus, Malus). Transmission through mechanical inoculation from pruning tools and grafting to infective propagative materials is a potential pathway for spread; no animal vector is known and it is not known to be seed transmitted. PBCVd has been reported in several European countries, Malta, Tunisia, Turkey, Australia, Japan, China, and the Americas; the geographic distribution may be underestimated because of symptomless infections in certain hosts. PBCVd was placed on the A1 list in Canada in 1995, listed as a quarantine pest in the United States in 1989, is regarded as quarantine pathogen for Australia, and is listed in a certification scheme to produce clean Pyrus and Cydonia sp. planting material in an OEPP/EPPO Bulletin in 1999 (OEPP/EPPO, 1999).

Taxonomic Tree

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  • Domain: Virus
  •     Unknown: Viroids
  •         Family: Pospiviroidae
  •             Genus: Apscaviroid
  •                 Species: Pear blister canker viroid

Notes on Taxonomy and Nomenclature

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Viroids are small covalently-Di closed, circular single-stranded, highly base-paired RNA molecules that range in size from 249 to 434 nucleotides, do not encode peptides or proteins, and use host proteins for replication, movement, and processing of replication intermediates, which distinguishes them from plant viruses (Diener, 1971, 1987). Pear blister canker disease was first reported in pear (Pyrus communis) in France (Desvignes, 1970). Bark symptoms associated with the disease in the pear indicator host cv. "A20" was shown to be caused by a viroid, and the name pear blister canker viroid (PBCVd) was coined (Flores et al., 1991; Ambrós et al., 1995a; Flores et al., 2011). The nucleotide sequence revealed that PBCVd is a member of the family Pospiviroidae, genus Apscaviroid (Hernández et al., 1992a,b; Di Serio et al., 2014). The reference isolate is 315 nt in length and the most stable predicted secondary structure was branched (Hernández et al., 1992a); additional variants range in size from 312-316 nt in length. PBCVd is most closely related phylogenetically to grapevine yellow speckle viroid (GYSVd) (Di Serio et al., 2014).

Description

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The sequence of the pear blister canker viroid reference isolate (EMBL accession number S46812) is 315 nt in length and the most stable predicted secondary structure is branched (Hernández et al., 1992a); additional variants range in size from 312-316 nt in length (Ambrós et al., 1995b; Loreti et al., 1997; Barone et al., 2004; Joyce et al., 2006; Lolic et al., 2007; Attard et al., 2007; Yesilcollou et al., 2010).

Distribution

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Pear blister canker disease was first reported in pear (Pyrus communis) in France (Desvignes, 1970). Bark symptoms associated with the disease in the pear indicator host cv. "A20" were shown to be caused by pear blister canker viroid (PBCVd) (Flores et al., 1991; Ambrós et al., 1995a; Flores et al., 2011). An improved bioassay on new pear indicator hosts revealed the PBCVd was present in 10% of 152 old French pear cultivars (Desvignes et al., 1999). With the development of molecular technologies for rapid detection of PBCVd, it has since been reported in several European countries, including France, Italy, Spain, Greece, Boznia-Herzogovinia and Albania, and also in Malta, North Africa (Tunisia), Turkey, Australia, China, Japan and the Americas (Argentina and Canada) (see Distribution Table for details and references).

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

ChinaPresentIntroduced2004 Not invasive Shamloul et al., 2004; CABI/EPPO, 2013
JapanPresentIntroduced Not invasive Sano et al., 1997; CABI/EPPO, 2013Detected in European pears cultivated in two prefectures in Japan
TurkeyPresent Not invasive Yesilcollou et al., 2010; CABI/EPPO, 2013Pear germplasm collection only

Africa

TunisiaPresent Not invasive Hassen et al., 2004; CABI/EPPO, 2013

North America

CanadaPresent Not invasive Torchetti et al., 2012
-OntarioPresent Not invasive Torchetti et al., 2012
USAPresent only in captivity/cultivationLin et al., 2011Only in research locations/quarantine stations

South America

ArgentinaPresentNome et al., 2011; CABI/EPPO, 2013

Europe

AlbaniaPresentNavarro et al., 2011; CABI/EPPO, 2013
Bosnia-HercegovinaPresent Not invasive Lolic et al., 2007; CABI/EPPO, 2013Present in native and ancient pear cultivars
FrancePresentNative Not invasive Desvignes, 1970; Desvignes et al., 1999; CABI/EPPO, 2013
GreeceWidespreadKaponi et al., 2010; CABI/EPPO, 2013
-CretePresentCABI/EPPO, 2013
ItalyPresent Not invasive Loreti et al., 1997; Barone et al., 2004; CABI/EPPO, 2013
MaltaPresent2006 Not invasive Attard et al., 2007; CABI/EPPO, 2013
SpainRestricted distribution Not invasive Flores et al., 1991; CABI/EPPO, 2013
UKPresentCABI/EPPO, 2013

Oceania

AustraliaPresentJoyce et al., 2006; CABI/EPPO, 2013
-QueenslandPresentConstable et al., 2007
-South AustraliaPresentConstable et al., 2007
-VictoriaPresentConstable et al., 2007

History of Introduction and Spread

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Since being identified in France in the 1960’s, improved diagnostic techniques have led to the viroid being found in other European countries and in parts of North Africa, Asia, Australia and the Americas. Among recent records, it was identified for the first time in Malta in 2006 (Attard et al., 2007), and in Albania and Argentina in 2010 (Navarro et al., 2011; Nome et al., 2011). PBCVd has been detected in imported propagation materials tested for their sanitary status in research locations in the United States (Lin et al., 2011). The geographic distribution may be underestimated because of symptomless infections in certain pears. Quince, which is a symptomless carrier that is used as a pear rootstock, may have contributed to its spread. 

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
Terrestrial – ManagedCultivated / agricultural land Present, no further details Harmful (pest or invasive)
Managed forests, plantations and orchards Present, no further details Harmful (pest or invasive)

Hosts/Species Affected

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The disease caused by PBCVd, pear blister canker disease, can infect several cultivars of pear (Pyrus communis), its major host, but most pear cultivars are tolerant and do not produce symptoms of infection (Desvignes et al., 1999). On the indicator pear host, cv. "A20", sensitive infected trees display pustule bark cankers and can slowly decline and eventually die. PBCVd was initially reported in pear and quince (Cydonia oblonga), followed by wild pear (Pyrus amygdaliformis) (Kyriakopoulou et al., 2001) and nashi (Pyrus serotina) (Joyce et al., 2006). PBCVd can be experimentally transmitted to and infect species in other woody hosts (15 species of Pyrus, 16 commercial pear cultivars, Chaenomeles, Sorbus, and five species of Malus) although they were all symptomless (Desvignes et al., 1999). PBCVd has been reported in wild and cultivated apples in Greece (Kaponi et al., 2010).

Therefore, PBCVd susceptibility is not limited to the genus Pyrus. PBCVd can be mechanically transmitted to cucumber (Cucumis sativis), which displays mild leaf symptoms or no symptoms (Flores et al., 1991).

Growth Stages

Top of page Vegetative growing stage

Symptoms

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Most commercial pear cultivars do not express bark symptoms when infected with PBCVd (Desvignes et al., 1999), however, symptoms on the pear indicator host, cv. “A20” and proposed replacement indicator pear indicator hosts, cv. "Fieudiére" (Fieud 37 and Fieud 110), and some susceptible pear varieties, include petiole and leaf necrosis, bark pustules, bark scaling, bark splitting and tree death (Desvignes, 1970; Loreti et al., 1997; Desvignes et al., 1999). On the pear indicator host cv. "A20", differing severity depends on the PBCVd isolate (Desvignes et al., 1999). Pear and quince (used as a pear rootstock) are generally tolerant (Devisgnes et al., 1999). There are no reported symptoms on fruit.                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                       

List of Symptoms/Signs

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SignLife StagesType
Leaves / necrotic areas
Stems / discoloration of bark

Biology and Ecology

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Genetics

The genome of PBCVd is a single-stranded, covalently-closed circular single-stranded RNA molecule of 312-315 nt in length. It does not contain a protein coat and does not encode proteins.  

Reproductive Biology

Viroids replicate autonomously in plant cells and are dependent on cellular RNA polymerases, ribonucleases (enzymes or ribozymes) and RNA ligases for replication. PBCVd, as a member of the Pospiviroidae, replicates in the nucleus via a rolling circle mechanism that produces plus and minus sense greater-than-unit length RNAs, which are processed to the mature, circular form. Viroids spread from cell-to-cell and systemically in the infected host through the vascular system.

Physiology and Phenology

Most of the natural hosts of PBCVd are woody perennials.

Environmental Requirements

The distribution of PBCVd is dependent on the growing conditions for its woody perennial hosts and tree species (Pyrus sp., Malus sp., Cydonia sp.).

Climate

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ClimateStatusDescriptionRemark
C - Temperate/Mesothermal climate Preferred Average temp. of coldest month > 0°C and < 18°C, mean warmest month > 10°C
D - Continental/Microthermal climate Preferred Continental/Microthermal climate (Average temp. of coldest month < 0°C, mean warmest month > 10°C)

Means of Movement and Dispersal

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Transmission of PBCVd is through mechanical inoculation from pruning tools and grafting/budding to infective propagative materials and is a potential pathway for spread; no animal vector is known and it is not known to be seed transmitted (Flores et al., 2011).

Seedborne Aspects

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No seed transmission has been reported for this pathogen.

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Breeding and propagation Yes Yes
Crop production Yes Yes
Nursery trade Yes Yes
Research Yes Yes

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Germplasm Yes Yes
Plants or parts of plants Yes Yes

Impact: Economic

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Most pear cultivars are tolerant to PBCVd (Desvignes et al., 1999) and 10% of old French cultivars are known to be infected without known significant economic impact. However, pear indicator hosts are severely affected, implying that in the future, newer pear and apple cultivars may be impacted by PBCVd infection.

Risk and Impact Factors

Top of page Invasiveness
  • Benefits from human association (i.e. it is a human commensal)
Impact outcomes
  • Host damage
Impact mechanisms
  • Pest and disease transmission
  • Pathogenic
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Difficult to identify/detect as a commodity contaminant
  • Difficult to identify/detect in the field

Diagnosis

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PBCVd infection is symptomless on many of its woody hosts. PBCVd can be identified by bioassay on pear indicator hosts cv. “A20”, and improved cvs. “Fieudiére” (Fieud 37 and Fieud 110) (Ambros et al., 1995b; Desvignes et al., 1999).

Double polyacrylamide gel electrophoresis (gel electrophoresis of known and unknown viroids based on the physical properties of circular viroid molecules) was used to identify the etiology of pear blister canker disease (Flores et al., 1991). Characterization of the nucleotide sequence of PBCVd (Hernández et al., 1992a) allowed the development of rapid and sensitive dot-blot and tissue-print hybridization assays using radioactive or non-radioactive PBCVd-specific probes (Ambrós et al., 1995b;  Desvignes et al., 1999; Lolić et al., 2007). More recent detection methods for PBCVd include reverse transcription-polymerase chain reaction (RT-PCR) using either pome viroid-specific or PBCVd-specific primer pairs (Ragozzino et al., 2004; Hassen et al., 2006; Lin et al., 2011; Faggioli and Ragozzino, 2017), RT-PCR-ELISA (Shamloul et al., 2002), and quantitative real-time PCR (Malandraki et al., 2015).

Prevention and Control

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As contaminated equipment and infected propagation material are the primary means of transmission, viroid-free propagation material should be used and tools should be disinfected to prevent secondary spread (Flores et al., 2011). Infected materials should be discarded. Sanitary measures include disinfection of tools and greenhouse benches with diluted household bleach (10% solution of household bleach) (Eastwell and Sano, 2009; Sano, 2009; Flores et al., 2011).

Gaps in Knowledge/Research Needs

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To determine the geographic distribution of PBCVd, more surveys are needed using molecular tools in pear growing regions of the world.

References

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Ambrós, S., Desvignes, J. C., Llácer, G., Flores, R., 1995. Pear blister canker viroid: sequence variability and causal role in pear blister canker disease. Journal of General Virology, 76(10), 2625-2629. doi: 10.1099/0022-1317-76-10-2625

Ambrós, S., Llácer, G., Desvignes, J. C., Flores, R., 1995. Peach latent mosaic and pear blister canker viroids: detection by molecular hybridization and relationships with specific maladies affecting peach and pear trees. In: Acta Horticulturae [XVIth International Symposium on fruit tree virus diseases, 27 June - 2 July, 1994, Rome, Italy], (No. 386) [ed. by Barba, M., Hadidi, A.]. 515-521.

Attard, D., Afechtal, M., Agius, M., Matic, S., Gatt, M., Myrta, A., Serio, F. di, 2007. First report of Pear blister canker viroid in Malta. Journal of Plant Pathology, 89(Suppl.3), S71. http://www.agr.unipi.it/sipav/jpp/index.html

CABI/EPPO, 2013. Pear blister canker viroid. [Distribution map]. Distribution Maps of Plant Diseases, No.April. Wallingford, UK: CABI, Map 1142 (Edition 1)

Constable, F. E., Joyce, P. A., Rodoni, B. C., 2007. A survey of key Australian pome fruit growing districts for exotic and endemic pathogens. Australasian Plant Pathology, 36(2), 165-172. http://www.publish.csiro.au/nid/39.htm doi: 10.1071/AP07003

Desvignes, J. C., Cornaggia, D., Grasseau, N., Ambrós, S., Flores, R., 1999. Pear blister canker viroid: host range and improved bioassay with two new pear indicators, Fieud 37 and Fieud 110. Plant Disease, 83(5), 419-422. doi: 10.1094/PDIS.1999.83.5.419

Desvignes, J.C., 1970. Pear virus diseases and their detection, [Les maladies á virus du poirer et leur détection] CTIFL Doc. 26, 1-12

Di Serio, F. , Afechtal, M., Attard, D., Choueiri, E., Gumus, M., Kaymak, S., Lolic, B., Matic, S., Navarro, B., Yesilcollou, S., Myrta, A., 2010. Detection by tissue printing hybridization of Pome fruit viroids in the Mediterranean basin. In: Julius-Kühn-Archiv,(No.427) . Quedlinburg, Germany: Julius Kühn Institut, Bundesforschungsinstitut für Kulturpflanzen. 357-360. http://pub.jki.bund.de/index.php/JKA/issue/archive

Di Serio, F., Flores, R., Verhoeven, J. Th. J., Li, S.-F., Pallás, V., Randles, J. W., Sano, T., Vidalakis, G., Owens, R. A., 2014. Current status of viroid taxonomy. Archives of Virology, 159, 3467-3478.

Di Serio, F., Torchetti, E. M., Flores, R., Sano, T., 2017. Other apscaviroids infecting pome fruit trees. In: Viroid and Satellites, [ed. by Hadidi, A., Flores, R., Randles, J. W., Palukaitis, P.]. London, UK: Academic Press. 229-241.

Diener, T. O., 1971. Potato spindle tuber “virus”. IV. A replicating, low molecular weight RNA. Virology, 45, 411-428.

Diener, T. O., 1987. The Viroids, New York, USA: Plenum Press.344 pp.

Eastwell, K., Sano, T., 2009. Hop stunt disease. In: The Compendium of Hop Diseases, Arthropod Pests and Other Disorders, [ed. by Mahafee, W. F., Pethybridge, S. J., Gent, D. H.]. St. Paul, USA: APS Press. 48-51.

Elleuch, A., Hamdi, I., Bessaies, N., Fakhfakh, H., 2013. Single-strand conformation polymorphism for molecular variability studies of six viroid species. Bioscience, Biotechnology and Biochemistry, 77(1), 182-188. https://www.jstage.jst.go.jp/article/bbb/77/1/77_120714/_article doi: 10.1271/bbb.120714

Faggioli, F., Ragozzino, E., 2002. Detection of pome fruit viroids by RT-PCR using a single primer pair. Journal of Plant Pathology, 84, 125-128.

Flores, R., Ambrós, S., Llácer, G., Hernández, C., 2011. Pear blister canker viroid. In: Virus and Virus-like Diseases of Pome and Stone Fruits, [ed. by Hadidi, A., Barba, M., Candresse, T., Jelkmann, W.]. St. Paul, MN, USA: APS Press. 63-66.

Flores, R., Hernández, C., Llácer, G., Desvignes, J. C., 1991. Identification of a new viroid as the putative causal agent of pear blister canker disease. Journal of General Virology, 72(6), 1199-1204. doi: 10.1099/0022-1317-72-6-1199

Hassen, I. F., Kummert, J., Marbot, S., Fakhfakh, H., Marrakchi, M., Jijakli, M. H., 2004. First report of Pear blister canker viroid, Peach latent mosaic viroid, and Hop stunt viroid infecting fruit trees in Tunisia. Plant Disease, 88(10), 1164. http://www.apsnet.org doi: 10.1094/PDIS.2004.88.10.1164A

Hassen, I. F., Roussel, S., Kummert, J., Fakhfakh, H., Marrakchi, M., Jijakli, M. H., 2006. Development of a rapid RT-PCR test for the detection of peach latent mosaic viroid, pear blister canker viroid, hop stunt viroid and apple scar skin viroid in fruit trees from Tunisia. Journal of Phytopathology, 154(4), 217-223. doi: 10.1111/j.1439-0434.2006.01087.x

Hernández C., Llácer G., Desvignes J. C., Flores R., 1992b. Evidence supporting a viroid etiology for Pear blister canker disease. Acta Horticulturae, 309, 319-324.

Hernández, C., Elena, S. F., Moya, A., Flores, R., 1992. Pear blister canker viroid is a member of the apple scar skin subgroup (apscaviroids) and also has sequence homology with viroids from other subgroups. Journal of General Virology, 73(10), 2503-2507. doi: 10.1099/0022-1317-73-10-2503

Joyce, P. A., Constable, F. E., Crosslin, J., Eastwell, K., Howell, W. E., Rodoni, B. C., 2006. Characterisation of Pear blister canker viroid isolates from Australian pome fruit orchards. Australasian Plant Pathology, 35(4), 465-471. http://www.publish.csiro.au/nid/39.htm doi: 10.1071/AP06050

Kaponi MS, Luigi M, Barba M, Kyriakopoulou PE, 2010. Pospiviroidae viroids in naturally infected stone and pome fruits in Greece. Julius-Kühn-Archiv [Proceedings of the 21st International Conference on Virus and other Graft Transmissible Diseases of Fruit Crops, Neustadt, Germany, 5-10 July, 2009.], No.427:353-356. http://pub.jki.bund.de/index.php/JKA/issue/archive

Kaponi, M., Luigi, M., Kyriakopoulou, P. E., 2012. Mixed infections of pome and stone fruit viroids in cultivated and wild trees in Greece. New Disease Reports, 26, 8. http://www.ndrs.org.uk/article.php?id=026008 doi: 10.5197/j.2044-0588.2012.026.008

Kyriakopoulou, P. E., Giunchedi, L., Hadidi, A., 2001. Peach latent mosaic and pome fruit viroids in naturally infected cultivated pear Pyrus communis and wild pear P. amygdaliformis: implications on possible origin of these viroids in the Mediterranean region. Journal of Plant Pathology, 83(1), 51-62.

Lin, L. M., Li, R. H., Mock, R., Kinard, G., 2011. Development of a polyprobe to detect six viroids of pome and stone fruit trees. Journal of Virological Methods, 171(1), 91-97. http://www.sciencedirect.com/science/journal/01660934 doi: 10.1016/j.jviromet.2010.10.006

Lolic, B., Afechtal, M., Matic, S., Myrta, A., Serio, F. di, 2007. Detection by tissue-printing of pome fruit viroids and charaterization of Pear blister canker viroid in Bosnia and Herzegovina. Journal of Plant Pathology, 89(3), 369-375. http://www.agr.unipi.it/sipav/jpp/index.html

Loreti, S., Faggioli, F., Barba, M., 1997. Identification and characterization of an Italian isolate of pear blister canker viroid. Journal of Phytopathology, 145(11/12), 541-544. doi: 10.1111/j.1439-0434.1997.tb00363.x

Malandraki, I., Varveri, C., Olmos, A., Vassilakos, N., 2015. One-step multiplex quantitative RT-PCR for the simultaneous detection of viroids and phytoplasmas of pome fruit trees. Journal of Virological Methods, 213, 12-17. http://www.sciencedirect.com/science/journal/01660934 doi: 10.1016/j.jviromet.2014.11.010

Malfitano, M., Barone, M., Ragozzino, E., Alioto, D., Flores, R., 2004. Identification and molecular characterization of Pear blister canker viroid isolates in Campania (Southern Italy). Acta Horticulturae, (No.657), 367-371. http://www.actahort.org

Malfitano, M., Barone, M., Ragozzino, E., Alioto, D., Flores, R., 2004. Identification and molecular characterization of Pear blister canker viroid isolates in Campania (Southern Italy). Acta Horticulturae, (No.657), 367-371. http://www.actahort.org

Navarro B, Bacu A, Torchetti M, Kongjika E, Susuri L, Serio Fdi, Myrta A, 2011. First record of Pear blister canker viroid on pear in Albania. Journal of Plant Pathology, 93(4, Supplement):S4.70. http://sipav.org/main/jpp/index.php/jpp/issue/view/118

Nome CF, Difeo LV, Giayetto A, Rossini M, Frayssinet S, Nieto A, 2011. First report of Pear blister canker viroid in pear trees in Argentina. Plant Disease, 95(7):882. http://apsjournals.apsnet.org/loi/pdis

OEPP/EPPO, 1999. Pathogen-tested material of Malus, Pyrus and Cydonia. In: Bulletin O€PP/€PPO Bulletin,(29) . 239-252.

Ragozzino, E., Faggioli, F., Barba, M., 2004. Development of a one tube-one step RT-PCR protocol for the detection of seven viroids in four genera: Apscaviroid, Hostuviroid, Pelamoviroid and Pospiviroid. Journal of Virological Methods, 121(1), 25-29. doi: 10.1016/j.jviromet.2004.05.012

Sano, T., Apple fruit crinkle viroid (AFCVd). In: The Compendium of Hop Diseases, Arthropod Pests and Other Disorders, [ed. by Mahafee, W. F., Pethybridge, S. J., Gent, D. H.]. St. Paul, MN, USA: APS Press. 39.

Sano, T., Li, S. F., Ogata, T., Ochiai, M., Suzuki, C., Ohnuma, S., Shikata, E., 1997. Pear blister canker viroid isolated from European pear in Japan. Annals of the Phytopathological Society of Japan, 63(2), 89-94. doi: 10.3186/jjphytopath.63.89

Shamloul, A. M., Faggioli, F., Keith, J. M., Hadidi, A., 2002. A novel multiplex RT-PCR probe capture hybridization (RT-PCR-ELISA) for simultaneous detection of six viroids in four genera: Apscaviroid, Hostuviroid, Pelamoviroid, and Pospiviroid. Journal of Virological Methods, 105(1), 115-121. doi: 10.1016/S0166-0934(02)00090-3

Shamloul, A. M., Yang, X., Han, L., Hadidi, A., 2004. Characterization of a new variant of Apple scar skin viroid associated with pear fruit crinkle disease. Journal of Plant Pathology, 86(3), 249-256.

Torchetti, E. M., Birch, C., Cooper, M., Masters, C., Arocha, R. Y., Serio F. di, Michelutti, R., 2012. Detection and identification of pear blister canker viroid occurring in pear trees in Canada. Petria, 22(3), 123-458.

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30/09/18 Original text by:

Rosemarie W Hammond, Molecular Plant Pathology Laboratory, USDA-ARS, Beltsville, Maryland, USA

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