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Options Méditerranéennes Série B n.55 Directory of Infectious Diseases of Grapevines and Viroses and Virus-like Diseases of the Grapevine: Bibliographic Report 1998-2004
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Boudon-Padieu 2006 . Martelli.CIHEAM Centre International de Hautes Etudes Agronomiques Méditerranéennes Options méditerranéennes Directeur de la publication: Bertrand Hervieu Directory of Infectious Diseases of Grapevines and Viroses and Virus-like Diseases of the Grapevine: Bibliographic Report 1998-2004 Editors: G.P. E.
Martelli. Italy. 2006 Reproduction partielle ou totale interdite sans l'autorisation d'«Options Méditerranéennes» Reproduction in whole or in part is not permitted without the consent of «Options Méditerranéennes» .it Directory of Infectious Diseases of Grapevines and Viroses and Virus-like Diseases of the Grapevine: Bibliographic Report 1998-2004 Editors: G. Série B : N.This volume of Options Méditerranéennes Series B has been formatted and paged up by the IAM Bari Editing Board La maquette et la mise en page de ce volume de Options Méditerranéennes Series B ont été réalisées à l'Atelier d'Edition de l'IAM Bari Copy number : 500 Printed by IDEAPRINT -Bari.P. 55 ISSN : 1016-1228 ISBN : 2-85352-338-1 © CIHEAM. E. 279 Options Méditerranéennes.September 2006 tel. Boudon-Padieu Bari : CIHEAM (Centre International de Hautes Etudes Agronomiques Méditerranéennes) p. +39 080 542 45 87 e-mail: ideaprint@virgilio.
TABLE OF CONTENTENTS PREFACE INTRODUCTION INFECTIOUS AGENTS OF GRAPEVINES INFECTIOUS DEGENERATION (GRAPEVINE FANLEAF VIRUS) INFECTIOUS DEGENERATION (EUROPEAN AND MEDITERRANEAN NEPOVIRUSES) GRAPEVINE DEGENERATION AND DECLINE (AMERICAN NEPOVIRUSES) GRAPEVINE LEAFROLL RUGOSE WOOD COMPLEX GRAFT INCOMPATIBILITY FLECK COMPLEX MINOR VIRUS DISEASES VIRUS-LIKE DISEASES VIROIDS GRAPEVINE YELLOWS: GENERAL PROPERTIES GRAPEVINE YELLOWS: INDIVIDUAL DISEASES GRAPEVINE YELLOWS: REFERENCES PIERCE'S DISEASE THE VIROSES AND VIRUS-LIKE DISEASES OF THE GRAPEVINE: BIBLIOGRAPHIC REPORT .1998-2004 INTRODUCTION SUBJECT INDEX REFERENCES 7 11 15 17 33 49 59 76 93 99 107 117 127 133 149 173 195 203 205 207 219 5 .
This raises the question of how to offer to those interested in a particular field the means for following with the least difficulty the research work being done in so many laboratories throughout the world and the information stemming from it. As a whole.P.PREFACE Virus. which reflect on the commercial value and productive life of the vineyards. that fosters coperative studies in grapevine virology.P. ICVG has a long-lasting tradition in these endeavours. Improving the sanitary conditions of the industry. to all those interested in viticultural matters. and practitioners. The “Bibliographic Report” is another most precious achievement by R. they reduce graft compatibility of scions and rootstocks. these diseases cause loss of vigour and often a decline of affected stocks. Bovey. although the technology is now available for detection and elimination of most of the pathogens occurring in propagation material. Sincere thanks are espressed to the authors of these contributions. a distinguished virologist of worldwide reputation and one of the father founders of ICVG. Scientific knowledge is nevertheless advancing rapidly and one consequence of this is the increased number of publications in specialized journals and of papers delivered during technical Meetings and Congresses. G. Boudon-Padieu and the latest Bibliographic Report (1998-2004) on “The Viroses and Virus-like Diseases of the Grapevine” by R. virus-like and phytoplasma diseases of grapevines constitute a major limiting factor to the development and well-being of the world viticultural industry. Bovey The “Directory” is a work of great thoroughness and accuracy. The present issue of Options Méditerrnéennes. Boudon-Padieu has a long lasting experience in the study of phytoplasmas. produced under the auspices of ICVG. Italy 7 . students. grouping diseases and setting them in a well thought out order. to which IAM-B was happy to contribute. whereas E. in the belief of rendering a good service to the scientific community and. Notwithstanding these efforts and the promoting activity of the International Council for the Study of Virus and Virus-like Diseases of the Grapevine (ICVG). being a recognized authority in this field. Cosimo Lacirignola Director of IAM Bari. up to date as to the time of publication. or induce subtle debilitating effects which are difficult to percieve. except when virus-free vines are grown for comparison alongside with non-sanitized sister vines. Special care was taken in the organization of the subject matter. is contributing with direct involvement in international research projects and through the activity the Mediterranean Network on Certification of Grapevines. Martelli and E. hosts a new edition of the “Directory of Infectious Diseases of Grapevines” by G. whose publication IAM-B is happy to support. Martelli is the President of ICVG and a widely known virologist who has much contributed to the advancement of the knowledge on grapevine viruses for the last forty years or so. and to the quality and quantity of the crop. it is most unfortunate that dissemination of infectious diseases continues. by and large. As to the authors. whose Secretatiat he has admirably conducted since its very establishment in 1962. Furthemore. technicians. is therefore a goal of utmost importance to which the Mediterranean Agronomic Institute of Bari (IAM-B). taking also into account the future role of the Mediterranean as a free-trade area. One way of achieving this goal is to produce and make available updated accounts of the state-of-theart of grapevine virology and comprehensive bibliographic reports. promoting the publication of the Bibliografic Report 1985-1997 on virus and virus-like diseases of the grapevine. so as to produce a document which can readily be used by scientits.
Martelli University of Bari. France 2006 .International Council for the Study of Virus and Virus-like Diseases of the Grapevine (ICVG) DIRECTORY OF INFECTIOUS DISEASES OF GRAPEVINES G.P. Italy E. Boudon-Padieu INRA Dijon.
Vitis 18.) undoubtedly represents one of the horticultural crops most widely grown in temperate climates. ICVG was established in 1962 by a group of American and European plant pathologists who realized the importance of creating an organization for promoting research on grapevine virology and favouring the exchange of information among students. The importance of the grapevine industry and the magnitude of the problems caused by these pathogens has generated wide interest which. nurserymen. its 14th Conference having taken place in Italy in 2003 (Bovey and Gugerli. The viroses and virus-like diseases of the grapevine. among other things. Gugerli. As most of the vegetatively propagated crops. has fostered intensive research. Bibliographie des viroses de la vigne des origines à 1965. Office International de la Vigne et du Vin. that has been especially active at the international scale from the late 1950's onwards. having a negative impact on the plant vigour and longevity.B. and endangering the survival of affected vines. 29 (Series B. Locorotondo 2003. 10-172. viroids. ICVG has met regularly every 3 to 4 years.400. growers. W. 1999. all efforts should be made to improve its sanitary conditions. The viroses and virus-like diseases of the grapevine. Infected propagating material is largely responsible for the spread of diseases among and within viticultural countries. Thus. Les virus de la vigne. Hewitt W. and phytoplasmas) recorded from the grapevine can be highly detrimental to this crop. The papers up to 2004 are listed and commented upon in six bibliographic reports: Caudwell A. which were compiled under the auspices of the International Council for the Study of Viruses and Virus Diseases of the Grapevine (ICVG).. Hewitt and R. and P.INTRODUCTION The grapevine (Vitis spp. has produced an impressive series of papers which now number about 5. Bovey. A short history of ICVG. 1972. The viroses and virus-like diseases of the grapevine. 205-279. Options Méditerranéenes. 1-2. 3rd part). and R.P. 11 . A bibliographic report 1971-1978. and administrators on the detrimental effects of infectious diseases on the well-being of the industry. A bibliographic report 1998-2004.. viroids.B. in turn. shortening the productive life of vineyards. To this effect. A bibliographic report 1979-1984. recognizes that a number of the 60 or so infectious agents (viruses. 316-376. as well as on the quality and quantity of the yield. Options Méditerranéenes. 2003. 2003). 303-324. and supporting initiatives for the establishment and implementation of clean stock and certification programmes. Bovey R. and G. 1965.. 1979. grapevines are exposed to the attacks of a variety of pests and pathogens among which infectious intracellular agents (viruses. Since its foundation. The increased attention paid to grapevine virological problems and the like. 1986. ICVG has issued the recommendations that follow: The International Council for the Study of Virus and Virus-like Diseases of the Grapevine (ICVG). Bibliographie de 1965-1970.. phloem-and xylemlimited prokaryotes) play a major role. Extended Abstracts 14th Meeting of ICVG.. Bovey R. Martelli. Bovey R. 227-275. attracting the attention of scientists. 3rd part). 76 pp. and a highly valuable agricultural commodity. 2006. The viroses and virus-like diseases of the grapevine. ICVG has been instrumental in fostering basic and applied research in grapevine virology. A bibliographic report 1985-1997. Bovey R. Vitis 25. Bovey. xx (Series B. Paris. From the very beginning. Caudwell A. causing heavy losses. Vitis 11.
rugose wood (GVA. all efforts should be made to improve its sanitary conditions. Certified selections should be tested for specific pathogens. Woodham. high standards are essential to ensure that no viticultural area is compromised by the introduction and spread of diseases. Grapevine (Vitis) virus and virus-like diseases. Locorotondo. Hewitt. R. bois noir. Class 1 should include only grape nursery stock which tests negative for the most damaging diseases/pathogens. Gubler and J. G. including the causal agents of fleck and rupestris stem pitting. Inadequate certification standards have repeatedly resulted in disease problems for growers and nurserymen.D. As efforts are made to harmonize grapevine certification protocols. Virus and Virus-like Diseases of Grapevines. W. Uyemoto will be published in 2005). Pearson R. Certified grapevines are derived from pathogen tested. Plant Virus Slide Series. viroids and phytoplasmas). 29 pp. Set 1 (O. (a revised edition by J. 2. Lausanne. is regarded as incompatible with an accepted sanitary status. recognises over 70 infectious agents affecting grapevine (viruses. (issued and approved in 2003 at the 14th ICVG Meeting. Certification of grapevine nursery stock is a powerful and effective tool to control these agents. 100 slides. Italy) The Proceedings of all the international ICVG Conferences have been published in extenso or. Martelli and A. and permit tracing the certified grapevines to the originally selected and tested plants.K. USA. 93 pp. Gärtel. Class 2 would be a specific pathogen-tested certification system for stock which remains within regulatory regions and is only distributed with disclosure of health status. G. that enables vineyards to economically and sustainably maintain quality and productivity. W. 181 pp.P. leafroll. St. eds) Clemson University. Martelli and A. GVB and GVD). 12 . Portugal ) The International Council for the Study of Virus and Virus-like Diseases of the Grapevine (ICVG). Uyemoto. Dias.The presence of diseases such as infectious degeneration. and other grapevine yellows). as Extended Abstracts. rugose wood. However. The certification process should specify conditions to prevent and detect subsequent infection of nursery plants by regulated pests. A.G. The American Phytopathological Society Press. Martelli. The agents that should be controlled by the Class 1 certification program are those associated with infectious degeneration and grapevine decline (nepoviruses). They represent a most valuable source of information. clonally selected primary sources. Goheen and H. technology should make it possible to exclude additional disease-causing viruses from the certified stock. under the name of Grapevine Viruses. Their elimination from mother vines intended for propagation should therefore be pursued. leafroll disease and associated closteroviruses (grapevine leafroll associated viruses 1. Clemson. G.C.P.W. and the need for preservation of heritage germplasm. Lisbon. as well as on the quality and quantity of the yield. Infected propagation material is largely responsible for the spread of diseases among and within viticultural countries. Vuittenez. Wilcox. Thus. and fleck. 1988. and A. (issued and approved in 1997 at the 12th ICVG Meeting. Grape Section. In order to preserve valuable grape clones and varieties. we propose two sanitary classes. a moratorium will be established for these viruses. Improvement of the sanitary level can be achieved through selection and sanitation. Paul. the virological problems of grapevines have been extensively treated and illustrated in a number of books and major review articles: Uyemoto J. 1980. and phytoplasmas (flavescence dorée. Goheen. Editions Payot.C. It would move freely between regulatory boundaries.P. (a 2nd revised edition edited by W.K. W. many of which can be highly detrimental to this crop. ensure clonal integrity. Until that time.F. valuable grape genetic resources exist which are infected with virus but are essential to the preservation of world viticultural heritage. Compendium of Grape Diseases. 1978. No other stock should move outside regulatory regions. having a negative impact on plant vigour and longevity. Virus-like Diseases and Other Disorders). and 3). which are best performed in the framework of certification programmes encompassing also clonal selection. regional viticultural conditions. Bovey R. lately. In addition. Tolin. In the future.B. Rowhani will be published in 2005 in the series APS Plant Virus Image CD-ROM.K.F.. The regional certification standards for Class 2 stock should be created at a local level based on the rate of endemic infection.C. Minnesota. Barnett and S.A.
CNRS . Khetarpal. 192 pp. 2000. 1ere partie: Effets des viroses sur la culture de la vigne et ses produits. Paris. which was established in the framework of the International Project RAB/88 sponsored by the United States Development Programme and the Food and Agriculture Organization of the United Nations.H.Frison E. the Steering Committee of ICGV decided to update the Directory and to entrust this task to G. 945-971. and B. Bulletin de l'OIV 69. and G. Boudon-Padieu. Martelli G.). American Phytopathological Society Press. and G. INRA Editions. C. Martelli Professor of Plant Virology. Martelli. Bovey and G. and to Dr. Rome. 1997. Hadidi. Protocols for detection of viruses and virus-like diseases: Les Colloques.P. FAO Publication Division. Director of the Mediterrenean Agronomic Institute of Bari for supporting the publication of this work. Rezaian and R. 225 pp. Paris. Chairman of the Board of Directors and Secretary General. Rome /International Board for Plant Genetic Resources. Hamze and Mr. R.S. bactéries et phytoplasmes de la Vigne. N. Editions Féret.P. Martelli.R.K. 1999. Ridé. B. Koganezawa. (ed). Walter B. Taylor. Virus certification of grapevines.P.A. Bari. Notwithstanding this wealth of published information a "Directory of Major Virus and Virus-like Diseases of Grapevines" was compiled in 1992 by R. Sélection sanitaire de la vigne: sélection sanitaire et sélection pomologique.. Giovanni P. Walter B. The authors hope that this endeavour may serve as a useful guideline and working tool for both experienced researchers and those who are now approaching the intriguing but intricate field of grapevine virology. Influence des viroses et qualité. University of Bari.). Rome. sélection pomologique. and R. At the 14th ICVG Meeting. France 13 . (ed. Scott. FAO/IBPGR Technical Guidelines for the Safe Movement of Grapevine Germplasm. a body now estinguished.P. eds.P. M.137 pp.A.. In: Plant Virus Disease Control (A. E. M.P. Influence des viroses et qualité. Walter B. respectively. 263 pp. Italy. Food and Agriculture Organization of the United Nations. 1998. Martelli G. Detection and Diagnosis of Graft-transmissible Diseases of Grapevines. H. of the Centre International des Hautes Etudes Agronomiques Méditerranéennes. Hervieu. St. Sélection sanitaire de la vigne: sélection sanitaire et sélection pomologique. Bulletin de l'OIV 70. 2ere partie: Sélection sanitaire. Martelli and published under the auspices of the Mediterranean Fruit Crop Improvement Council (MFCIC). Maladies à Virus. Martelli and E. Ikin. 261-276. Collingwood. 1993. Boudon-Padieu and M. They express their deep gratitude to Prof. Sanitary selection of the grapevine. n° 86.Université de Bourgogne Dijon. 1991. Walter. Elisabeth Boudon-Padieu Directeur de Recherches Head Biologie et Ecologie des Phytoplasmes Plante Microbe Environnement INRA . CSIRO Publishing. Lacirignola. 1997. Graft-transmissible Diseases of Grapevines. 1996. Paul. Bordeaux. 54 pp. Walter B. 5-23. Krake L.
INFECTIOUS AGENTS OF GRAPEVINES More than 70 infectious agents among viruses (58). phytoplasmas (8). The viral scenario of Vitis: viruses and their taxonomic affiliation(a) FAMILY GENUS SPECIES A. viroids (5). Viruses belonging to genera included into families BROMOVIRIDAE Alfamovirus Cucumovirus Ilarvirus Alfalfa mosaic virus (AMV) Cucumber mosaic virus (CMV) Grapevine line pattern virus (GLPV) Grapevine angular mosaic virus (GAMoV) BUNYAVIRIDAE CLOSTEROVIRIDAE Tospovirus Closterovirus Ampelovirus Tomato spotted wilt virus (TSWV) Grapevine leafroll-associated virus 2 (GLRaV-2) Grapevine leafroll-associated virus 1 (GLRaV-1) Grapevine leafroll-associated virus 3 (GLRaV-3) Grapevine leafroll-associated virus 4 (GLRaV-4) Grapevine leafroll-associated virus 5 (GLRaV-5) Grapevine leafroll-associated virus 6 (GLRaV-6) Grapevine leafroll-associated virus 7 (GLRaV-7) Grapevine leafroll-associated virus 8 (GLRaV-8) Grapevine leafroll-associated virus 9 (GLRaV-9) Two new putative ampeloviruses are being characterized in France and the USA COMOVIRIDAE Fabavirus Nepovirus Broadbean wilt virus (BBWV) Artichoke italian latent virus (AILV) Arabis mosaic virus (ArMV) Blueberry leaf mottle virus (BBLMV) Cherry leafroll virus (CLRV) Grapevine Bulgarian latent virus (GBLV) Grapevine Anatolian ringspot virus (GARSV) Grapevine deformation virus (GDefV) Grapevine Grapevine fanleaf virus (GFLV) Grapevine Tunisian ringspot virus (GTRV) Peach rosette mosaic virus (PRMV) Raspberry ringspot virus (RpRV) Tobacco ringspot virus (TRSV) Tomato ringspot virus (ToRSV) Tomato blackring virus (TBRV) FLEXIVIRIDAE Foveavirus Grapevine rupestris stem pitting-associated virus (GRSPaV) chrome mosaic virus (GCMV) 15 . and insecttransmitted xylematic bacteria (1) have been recorded form grapevines. This represents the highest number of intracelluar pathogens ever found in a single crop.
. London. Maniloff.M. The names of tentative species are written in Roman characters.Potexvirus (GINV) Potato virus X (PVX) Trichovirus Vitivirus Grapevine berry inner necrosis virus Grapevine virus A (GVA) Grapevine virus B (GVB) Grapevine virus C (GVC) Grapevine virus D (GVD) Carnation mottle virus (CarMV) Tobacco necrosis virus D (TNV-D) Grapevine Algerian latent virus (GALV) Petunia asteroid mosaic virus (PAMV) Grapevine asteroid mosaic-associated virus (GAMaV) Grapevine redglobe virus (GRGV) Grapevine fleck virus (GFkV) Grapevine rupestris vein feathering virrus (GRVFV) TOMBUSVIRIDAE Carmovirus Necrovirus Tombusvirus Marafivirus Maculavirus TYMOVIRIDAE A new putative marafivirus is being characterized in the USA POTYVIRIDAE Potyvirus(?) Unidentified potyvirus-like virus isolated in Japan from a Russian cultivar B. C. Mayo. Virus Taxonomy. (a) 16 . Viruses belonging to unassigned genera Idaeovirus Sadwavirus Sobemovirus Tobamovirus C.A. and Ball. pp.A. Taxonomically unassigned viruses Unnamed filamentous Grapevine Ajinashika virus (GAgV) Grapevine stunt virus (GSV) Grapevine labile rod-shaped virus (GLRSV) Raspberry bushy dwarf virus (RBDV) Strawberry latent ringspot virus (SLRSV) Sowbane mosaic virus (SoMV) Tobacco mosaic virus (TMV) Tomato mosaic virus (ToMV) Scientific names of definitive virus species are written in italics. L. J. (eds). 2005. Desselberger.. The updated taxonomy of all classified grapevine viruses can be found in: Fauquet. Elsevier/Academic Press. 1258.. U. 8Ith Report of the International Committee on Taxonomy of Viruses. M.
INFECTIOUS DEGENERATION .
low proportion of graft take. reduced rooting ability of propagation material. however. but bunches are small and few. Leaves are variously and severely malformed. Chromatic alterations of the leaves vary from a few scattered yellow spots. This disorder was first reported from California as a syndrome elicited by a specific strain of GFLV. and the yield may be much reduced. are diagnostic of grapevines infected by GFLV. the yellowing fades rapidly and the canopy develops a normal green color. The consensus is that fanleaf degeneration may have existed in the Mediterranean Basin and the Near East since the earliest time of grape cultivation. that give the leaf the appearance of an open fan. causing degenerative diseases whose symptoms are similar to. The characterizing symptoms of "Vein banding". 19 . Trabeculae. phloem. puckered. These structures are readily visible by light microscopy in lignified shoots. Often infected grapevines occur in patches. FANLEAF 1. deep lobes.). dégénérescence infectieuse (Fr.e. urticado (Port. and decreased resistance to adverse climatic factors. especially in the basal internodes. a disorder induced by Grapevine fanleaf virus (GFLV). fasciations. arricciamento. and grapevine leaves with typical symptoms were found in herbaria established before the introduction of American rootstock hybrids. Gelbmosaik (Germ.INFECTIOUS DEGENERATION (GRAPEVINE FANLEAF VIRUS) Several nepoviruses infect grapevines in Europe and the Mediterranean area. chlorotic mottling may accompany foliar deformations. records of this disease date back some 150 years. showing progressive decline. GFLV and several of the other grapevine-infecting European nepoviruses have distorting and chromogenic strains and may occur in mixed infections. parenchyma. showing abnormal branching. Main symptoms: Two distinct syndromes caused by different strains of the causal agent characterize this disease. Description Fanleaf is the oldest known and one of the most important and widespread virus disease of the grapevine. and acute denticulations. mosaico giallo.and zigzag growth. Tolerant cultivars produce fairly good crops whereas sensitive cultivars are severely affected. In the European literature. are small-sized and set poorly. With increased ambient temperatures during summer. Yellow mosaic is induced by chromogenic virus strains. and inflorescences). low yields and low fruit quality. and berries ripen irregularly. Infectious malformations are induced by virus strains causing distortions. tendrils. which exhibit widely open petiolar sinuses and abnormally gathered primary veins. More recently.). Bunches are smaller and fewer in number. The foliage develops bright chrome yellow discolorations early in the spring that may affect all vegetative parts (leaves. degenerazione infettiva (Ital. consist of chrome yellow flecks first localized along the main veins of mature leaves and progressing into the interveinal areas which appear in mid to late summer in a limited number of leaves. The foliage and shoots show little if any malformation. Reisigkrankheit (partly). vein banding symptoms were shown to be caused by a co-infection by Grapevine yellow speckle viroid and GFLV. double nodes. panachure. short internodes. asymmetrical. or endocellular cordons. another disease sometimes occurring in vineyards affected by infectious degeneration. Fruit set is poor. Now the disease is known to occur worldwide. roncet. and xylem cells. bunches are straggly. Occasionally.). Main synonyms: court-noué. may show open petiolar sinuses. The name of this virus comes from the peculiar malformation of infected leaves. i. shortened productive life.). shoots. Symptomatic leaves show little malformation. Foliar symptoms develop early in the spring and persist throughout the vegetative season becoming less distinct in summer. radial bars crossing the lumen of epidermal. Their economic impact varies with the tolerance of the cultivar to the viruses. or indistinguishable from those of fanleaf. Shoots are also malformed. sometimes appearing as rings or lines to extensive mottling of the veins and/or interveinal areas to total yellowing .
but is not a specific test. cheap. Indexing on herbaceous hosts by mechanical inoculation requires climatized greenhouses and is less reliable than ELISA. Some V. amaranticolor. These inclusions derive from membrane proliferation. and very sensitive method. Cytopathology: GFLV elicits the formation of intracellular cytopathic structures known as vesiculatevacuolate inclusion bodies which are often apposed to the nucleus. 20 .Agent: Grapevine fanleaf virus (GFLV) is a nepovirus with polyhedral particles of about 30 nm in diameter. Transmission: At a site. The best antigen sources for serological diagnosis are leaves collected in spring or cortical shavings from mature dormant canes. For transformation. However. serologically rather uniform and occurring as a family of minor molecular variants. Molecular assays using radioactive or digoxigenin-labelled probes. but it is still regarded as necessary for confirming freedom from virus infection. A satellite RNA 1114 nt in size is associated with some virus isolates. RT-PCR and immunocapture RT-PCR are becoming increasingly popular. RT-PCR is estimated to be four to sixfold more sensitive than ELISA. C. but is not reliable. index is determined by the viral coat protein. However. Control: Use of virus-tested scionwood and rootstock material in the framework of clean stock or certification programmes. which are desirable as they cause no harm to human health. but show few symptoms. Indexing on Vitis indicators by grafting takes a lot of time and field or greenhouse space. However. Natural GFLV infections have been detected in weeds in Hungary and Iran. Resistance to X. vuittenezi has been suspected but not proven.g. where they occur in a radial orientation. The virus occurs in the pollen of infected grapevines and herbaceous hosts. and is transmitted through seeds of C. Varietal susceptibility: Almost all known Vitis vinifera L. GFLV was the first grapevine virus to be recovered by mechanical inoculation and to be thoroughly characterized physicochemically and molecularly. both required for infectivity. Virus elimination is readily achieved from vegetating shoot tips by heat treatment (38-40 °C for as little as four weeks). Nematode populations transmit local virus isolates with a higher efficiency than those from other geographical areas. rupestris x M. index feeding. and redistribution and are thought to be sites of viral polyprotein processing and RNA replication. Detection: ELISA using polyclonal antisera and monoclonal antibodies is a quick. mannose and xylose. rotundifolia hybrid rootstocks (e. but resists infection when the virus is transmitted by the nematode. rotundifolia can be infected by GFLV when graft inoculated. a number of selectable marker genes toxic to non engineered vines are used. the use of fumigants against nematode vectors gives only a temporary but economically valuable control of the disease. Positive sense single-stranded RNA genome. Transmission by Xiphinema italiae has not been consistently documented. the endosperm of grapevine seeds. There are conflicting reports on seed transmission in grapevines. reorganization. amaranticolor. In the laboratory. Detection of trabeculae can give information on the health of American rootstocks. Muscadinia (Vitis) rotundifolia and Vitis munsoniana are highly resistant to X. in a persistent manner by the longidorid nematode Xiphinema index feeding on the roots of grapevines and retaining the virus for several months. M. Gomphrena globosa). and transmission by X. GFLV can be transmitted by mechanical inoculation from infected grapevine tissues to various herbaceous hosts (e. encapsidated in different particles.g. O36-16) show interesting levels of field resistance to GFLV. Work is under way in different laboratories to create GFLV-resistant rootstocks or cultivar through traditional breeding methods or genetic transformation technology. index (10 individuals) by ELISA and in single nematodes by RT-PCR and immunosorbent electron microscopy. Virus particles are often within tubular structures that accumulate in bundles in the cytoplasm or nucleus. GFLV has been detected in small groups of viruliferous X. use of fumigants is more and more questioned for environmental reasons and is being progressively banned. or by somatic embryogenesis. and soybean. vinifera. are toxic to many plants but not to V. vinifera x M. by in vitro meristem tip culture. although some like Vitis labrusca can be infected. American rootstocks are also susceptible and are generally very sensitive. with variable levels of sensitivity. rotundifolia hybrids is thought to be controlled by a single dominant gene. index in V. Observation of symptoms in the field is useful as a first step in selection. Endocelluar cordons or “trabeculae” are abnormal straight cylindrical spool-like o ribbon-like structures of pectocellulosic nature that cross the cell lumen in different tissues and are especially oustanding in vascular bundles. Chenopodium quinoa. varieties are susceptible. Specific transmission by X. consisting of two functional molecules 7342 nt (RNA-1) and 3774 nt (RNA-2) in size. quinoa. tolerance to infection is widespread in European grapes and a high resistance level of the "host plant resistance" type was found in two accessions from Afghanistan and Iran. C. This resistance is controlled by two unlinked recessive genes. Dissemination over medium and long distances is through infected vegetatively propagated scionwood and rootstocks. In contaminated soils.
. Description of grapevine degeneration in Frontignan (France) Rathay: Description of fanleaf disease from Austria (Zwiewipflereben) Ruggeri : Description of fanleaf disease from Italy (Roncet) Cholin: Description of fanleaf disease from Germany (Reisigkrankheit) Baccarini: First suggestion that fanleaf may be due to a virus. bactéries. France. Pantanelli: Fanleaf caused by contamination through the roots possibly due to heat-labile toxic substances Petri: Disinfection of contaminated soil at 120 oC or filtration of liquid leached from contaminated soil through porcelain filter prevents infection through the roots of grapevine. see the book by Galet.: Hypothesis that court-noué (fanleaf) is caused by a virus transmitted by phylloxera. the ICVG Bibliographic Reports (a) have recorded more than 1000 papers dealing with fanleaf. viroses et phanérogames). Heating whole plants in a thermostatic chamber at 37 °C for several weeks provides a temporary elimination of symptoms on the new growth but no lasting cure. With soil containing phylloxera. research and hypotheses on fanleaf. Tome 1: Les maladies dues à des végétaux (champignons. Report on first experiments on heat treatment of grapevine in order to eliminate fanleaf. as well as on controversies about transmission by phylloxera. No conclusive results were obtained. Arnaud: Court-noué is considered as a soil-borne virus disease. With individual phylloxera (radicicolous or gallicolous) fed on infected vines. 871 pp. Les maladies et les parasites de la vigne. (b) Galet P. Hypothesis about a possible role of phylloxera as a vector. (a) See references in the Introduction. 1977. Bovey: Review on grapevine virus and virus-like diseases. 1929 1931 1937 1937 1946 1950a. Hypothesis that fanleaf is a fungal disease. 1954 1958 Hewitt: Review on grapevine virus and virus-like diseases found in California. Healthy rooted cuttings or seedling of Rupestris du Lot were contaminated: 1. Imprimerie du "Paysan du Midi". With roots of fanleaf-infected grapevines with phylloxera feeding on them. 2. Branas et al. For a comprehensive review on early observations.: Experiments on the capacity of phylloxera to transmit fanleaf. 1977(b) 1865 1882 1895 1896 1902 1906 1910 1912 1912 1917 1918 Cazalis-Allut. Petri: Grapevine "arricciamento" (fanleaf) has a viral origin Arnaud and Arnaud: Hypothesis of a viral origin for grapevine court-noué (fanleaf). No direct proof of transmission by this aphid. Branas et al. 3. but only circumstantial evidence. Historical review From the late 1800 to 1997.2.b Hewitt: Fanleaf and yellow mosaic recorded from California. Montpellier. Schiff-Giorgini: Graft-transmission of fanleaf disease Pantanelli: Fanleaf disease can be trasmitted through the soil Pantanelli : Fanleaf disease has a patchy distribution in the field Petri: Association of trabeculae with fanleaf. 21 .
Potted plants to be cured are grown at 38°C for several weeks and shoot extremities are cut and rooted under mist in a greenhouse. Gerola et al. 1965 1967 1968 1968 1969 1969 1970 22 . Serological relationship with ArMV reported. whereas insecticide treatment has no effect. Hewitt et al. Description of the chipbudding method for indexing.1958 1958 1960 Vuittenez: Fumigation of fanleaf-contaminated soil with nematicides prevents infection of healthy grapevines replanted immediately. Goheen and Hewitt: Description of vein banding as a GFLV-induced disease Dias: Host range and properties of fanleaf and yellow mosaic viruses. Vuittenez: Mechanical transmission of fanleaf virus to Chenopodium quinoa and C. Boubals and Dalmasso: Experiments on soil disinfection against X. Vuittenez: New observations on the effects of soil fumigants on fanleaf in contaminated soils. Transmission of fanleaf virus by X. Graniti and Russo: A light microscope and cytochemical study of endocellular cordons. Brückbauer and Rüdel: The virus (or viruses) of Reisigkrankheit (GFLV and/or other nepoviruses) are seed transmitted in some herbaceous indicator plants.: Description of the Davis method of heat therapy of grapevines. latex test and barium sulfate test. including fanleaf virus: bentonite flocculation test.: Detection of GFLV particles in thin-sectioned grapevine root tissues. Yield was increased by 400 % in comparison with that of untreated controls. Discussion on the possible role of weeds in the epidemiology of the disease. index. yellow mosaic and ArMV 1960a 1960b 1961 1961 1962 1962 1963 1963 1963a. Dichloropropane-dichloropropene (DD) at 1000 l/ha gave satisfactory results.b Martelli and Hewitt: Comparative studies showed that Californian and Italian GFLV isolates are the same. Control of the vector by soil fumigation. Dias and Harrison: Relationships between the viruses causing fanleaf.: Investigations on grapevine virus diseases in California.: Fanleaf virus is transmitted by the nematode Xiphinema index Cadman et al.: Transmission of GFLV by Xiphinema italiae in Israel. Gifford and Hewitt: Use of heat therapy and in vitro shoot tip culture to eliminate fanleaf virus from infected grapevines. and no reinfestation by X. Bercks and Querfurth: Use of latex-test for detecting GFLV and other nepoviruses in grapevine tissue extracts in Germany. Bercks: Research on the use of three serological methods for detecting plant viruses. Cohn et al. index. Hewitt et al. Das and Raski: Studies on the relationships of GFLV with its vector X. Goheen et al. Reproduction of fanleaf symptoms in mechanically inoculated grapevine seedlings is confirmed Galzy: Heat treatment of grapevine plantlets grown aseptically in vitro. amaranticolor is confirmed. Reproduction of fanleaf symptoms in mechanically inoculated grapevine seedlings 1964 1964 1965 Taylor and Hewitt: Description and characterization of Australian isolates of GFLV. index occurred during the 6 year period of observation. index in France.: Transmission of fanleaf virus from grapevine to herbaceous hosts by mechanical inoculation and preliminary characterization of the virus.
Bass and Vuittenez: Thermotherapy was improved by growing shoot apices of heat treated vines aseptically on nutritive media or by grafting them on aseptic grape seedlings in vitro. index. rupestris St. Martelli and Piro: Evidence from a herbarium of dried specimens collected between 1880 and 1886 that fanleaf and yellow mosaic occurred in field-grown grapevine in Sicily in the second half of the 19th century Quacquarelli et al.3 dichloropropene or methyl bromide. Mur et al. but ELISA is more sensitive. Vuittenez: Review paper on grapevine fanleaf. this lining is shed and ingested in the intestine. St. yellow mosaic and vein banding) are transmitted in the same way by X. quinoa. the plant is placed in a heat cabinet for treatment Hévin et al. Raski et al. Walter et al. Uyemoto et al. George for detecting GFLV. Dias: Review paper on grapevine yellow mosaic. Both methods give satisfactory and similar results. Raski et al.: GFLV particles observed in the lumen of the oesophagus of X.: Detailed physico-chemical characterization of GFLV. Alfaro and Goheen: The different strains of fanleaf virus (fanleaf sensu stricto. rupestris is more accurate than mechanical transmission to C.: Control of fanleaf by soil fumigation with 1. During the moult of the nematode. Vineyards replanted in contaminated but treated soils remained healthy for at least 5 years. especially with low titre antisera.3-dichloropropene or methyl bromide. 1970 1970 1970 1971 1971 1972 1972 1973 1973 1973a 1973b 1974 1974 1975 1976 1976 1977 1979 1979 23 .: Comparison of indexing by mechanical inoculation to Chenopodium quinoa and by graft-transmission to V. Taylor: Review paper on grapevine vein banding. Querfurth and Paul: Protein A-coated latex-linked antiserum (PALLAS) method for detecting GFLV and other viruses. Van Velsen and Niejalke: Green budding for indexing grapevine with the indicator cvs. Indexing by budding on V. Goheen and Luhn: New method of heat therapy. A dormant bud of the variety to be cured is grafted onto a healthy potted rootstock. The acquisition time threshold is less than 5 minutes. The sensitivity of the latex test is increased. Hévin et al.: Use of green grafting as a quick and secure method for graft-indexing. index.: GFLV and marbrure (fleck) are not transmitted through the seeds of grapevine. Bercks: Serological detection of grapevine viruses in West Germany. PALLAS is quicker and cheaper than ELISA. Both tests are more sensitive than mechanical inoculation to C. After bud take. Raski and Schmitt: Progress in the control of the fanleaf-nematode complex by soil disinfection with 1. Mission or LN 33. anterior oesophagus and oesophageal bulb of their nematode vectors. quinoa.: Description of GFLV in the CMI/AAB Descriptions of Plant Viruses Taylor and Robertson: GFLV and ArMV are retained as a monolayer of particles adsorbed onto the cuticle lining the lumina of odontophore.: Heat therapy of grape plantlets grown in vitro causes changes in some characteristics of the variety. George.: Comparison between PALLAS latex test and ELISA for detecting GFLV in France.1970 1970 Hewitt et al.
index larvae were present in the X. Efficiency of both methods is compared. vuittenezi population used for the experiments could not be entirely ruled out. Walker et al. in California. These are promising sources of germplasm for obtaining resistant rootstocks. index and fanleaf in grapevines. Monette: Heat therapy of GFLV and ArMV--infected grapevines with alternating temperatures. index. Hafez et al.3-D) or methyl bromide applied 75-90 cm deep with 90 cm spacing for 1. Bouquet: Serological detection of GFLV in its vector X. Bouquet: Muscadinia rotundifolia becomes infected by GFLV when the virus is transmitted by grafting but resists infection when transmission is by X. especially in wood shavings of dormant canes during winter.: In vitro translation of genomic RNAs of GFLV yields two large polyproteins (220 Kd and 125 Kd) which are subsequently processed by proteolytic cleavage to form mature structural and non structural proteins.: Comparison of polyclonal and monoclonal antibodies for detecting fanleaf virus with ELISA in various grapevine tissues. index by ISEM Bovey et al.: Detection of fanleaf virus and other sap-transmissible viruses in grapevine tissues by ISEM. although it is not resistant to the virus itself. A Middle Eastern V. a very common species in vineyards of Rheinhessen and Palatinate.: Use of systemic nematicides for the control of X. Even in the cases where the virus was transmitted. index. Forty days of treatment. Lear et al. Bouquet: M. That X. Methyl bromide and 1. Russo et al. Krake and Woodham: Possibility that the agent of yellow speckle is involved together with GFLV in the etiology of vein banding.: Identification of a natural serological variant of GFLV from Tunisia Huss et al. index. vuittenezi.: Identification of several Vitis species and interspecific hybrids resistant to fanleaf virus. The use of methyl bromide requires a continuous cover with polyethylene sheeting for some time after the treatment. Savino et al. Morris-Krsinich et al. as vector of GFLV. rotundifolia is resistant to fanleaf virus transmission by X. 1980 1980 1980 1981 1981 1981 1981 1983a 1983b 1983 1983 1983 1985 1985 1986 1986 24 . Raski et al. vuittenezi might be a vector of GFLV is therefore uncertain.3-D (1400 l/ha) and 50-75 cm deep with 165 cm spacing for methyl bromide (448 kg/ha) gave a good control of X.: GFLV particles are arrayed in long parallel rows in thin-sectioned mesophyll cells of infected grapevines. Transmission trials gave a few positive results. with temperatures of 39 °C for 6 h followed by 22 °C for 18 h made it possible to eliminate both viruses from the developing shoot tips (2 mm) of in vitro cultured plantlets. vinifera accession represent an excellent example of host plant resistance to GFLV. the possibility that a few X.: Soil fumigation with 1. Rüdel: Discussion on the possible role of X.: Study on the effectiveness of soil fumigation for the control of X.: Detection of fanleaf virus in grapevine tissues by ELISA and ISEM in different periods of the year. RNA-2 contains the cistron coding for the viral coat protein. Vuittenez: Review on serological methods of detection and identification of grapevine viruses. Carbon disulfide gave less satisfactory results. index. index feeding.3-dichloropropene failed to eradicate either nematodes or fanleaf virus from the soil but reduced the incidence of the disease to acceptable levels.1979 1980 1980 Kalasian et al. index by ELISA.3-dichloropropene (1.: Experiments with systemic nematicides for controlling X. Brown and Roberts: Detection of fanleaf virus in its vector X. Raski et al.
Etienne et al. The selection of resistant cultivars and rootstocks is considered of primary importance. SLRV. cultivation of non-host plants and organic soil amendments are recommended. Altmayer: Elimination of GFLV. quinoa and transmitted by heterografting to Vialla and Kober 5BB rootstocks. Mild and severe strains were discriminated on the basis of field performance of infected Vitis.: Detection of GFLV in the vector Xiphinema index by ELISA. Walker et al. No eradication was obtained. Raski and Goheen: Comparison of 1. Rüdel: Severe restrictions set on the use of soil fumigants in West Germany for environmental reasons make control of "Reisigkrankheit" very difficult.1987 1987 1987 Huss et al. Treated vines yielded more for over 4 years.: Study of interactions between GFLV and ArMV isolates grown in C. Walker and Meredith: Identification of two accessions of Vitis vinifera resistant to GFLV. Treatments with soil fumigants are no longer permitted in Germany. Previous experience showed that 1.: Identification of a satellite RNA of GFLV. vinifera x V. Rüdel: Review on the most important virus diseases of grapevines in West Germany. but less consistent results were obtained with 1-10 nematodes.: Determination of the nucleotide sequence of the satellite RNA (RNA-3) of GFLV. Positive. rotundifolia showed good resistance to GFLV in California. TBRV and leafroll from infected grapevines by in vitro meristem tip culture. Resistance is controlled by two unliked recessive genes Walter et al.3-dichloropropene or methyl bromide fumigation following one year fallow period can give a satisfactory control of the disease for at least 12-15 years. Mild strains were shown to confer protection towards severe challenging strains in Chenopodium and grapevine.: Improvement in the serological detection of GFLV and ArMV viruses using monoclonal antibodies. Catalano et al. Walter et al.3-dichloropropene and methyl bromide for controlling X. Walter and Etienne: Detection of GFLV in wood shavings of dormant canes.: Possibility of detecting several nepoviruses or serotypes of nepoviruses in grapevine leaves or wood shavings in a single DAS-ELISA test using a mixture of different polyclonal antisera. Walter et al.: Determination of the complete sequence of GFLV RNA-2.: Evidence of a differential efficiency of GFLV transmission by Xiphinema index populations from different geographical origins. Viruliferous nematodes were crushed in standard extraction buffer and tested in batches of 1-50 by means of DAS-ELISA. the latter being especially damaging on the variety Kerner. GFLV. RNA-3 encodes a non structural protein.: Two rootstock selections derived from crossings V. Catalano et al. RpRSV and ArMV are common.: Use of green grafting technique for sensitive and quick GFLV detection under greenhouse conditions. Long term fallow (about 5 years). Lázár et al. ArMV. Serghini et al.: Detection of GFLV in grapevine seeds and seedlings by ELISA. Martelli and Taylor: Review article on nematode-transmitted viruses. Effect on yield and economic importance. Pinck et al. Fuchs et al. 1988 1988 1988 1989 1989 1989 1989 1989 1990 1990 1990 1990 1990 1991 1991 1991 25 . RpRV. index and GFLV.: Production and use of monoclonal antibodies to GFLV. Reliable results were obtained with samples of 20-50 nematodes. and has strong homologies with the satellite RNA associated with ArMV.
Walter et al. Spielmann et al.1991a 1991b Fuchs et al. Esmenjaud et al.: Genomic RNA-1 of GFLV is completely sequenced and its genetic organization determined. Viry et al.: European virologists propose a certification scheme for grapevine Gemmrich et al: Development and use digoxigenin-labelled cDNA probes for molecular detection of GFLV Nolasco and De Sequeira: Design and use of primers for specific amplification of GFLV sequences by IC-PCR Nolasco and De Sequeira: Molecular variability in the genome of GFLV isolates coming from the same vineyard assessed by IC-PCR combined with RFLP and SSCP analysis. which is also resistant to phylloxera.: Co-inoculation of C.: Two Vitis vinifera x Muscadinia rotundifolia rootstock hybrids (O39-16 and O4343) grafted with Cabernet sauvignon showed a high level of tolerance to GFLV.: GFLV satellite RNA detected in 5 of 34 virus isolates from different geographical locations.: Use of modified GFLV coat protein genes for transformation of different Nicotiana species for inducing resistance. Goussard and Wiid: First application of somatic embryogenesis for sanitation of grapevines.: Evidence that transgenic tobacco plants expressing the coat protein of GFLV are protected from GFLV infection. index. index infested soils.: In naturally GFLV-infected vineyards the hypovirulent ArMV A1 isolate induces delayed infection by GFLV Saldarelli et al. Both became infected in the course of a 12-year trial but had no reduced crop yields. 1993 1993 1993 1993 1993 1994 1994 1994 1994 26 . Ritzenthaler et al. GFLV is a quasispecies occurring in the field as a series of minor molecular variants. Satellite RNA appears to have a modulating effect on virus pathogenicity. Staudt: Study of the spread of GFLV in several Vitis species. identification of their replication determinants and evidence of replication in Chenopodium quinoa protoplasts 1993 1993 1993 1993 Martelli et al. Fuchs et al. quinoa with biologically active transcripts of GFLV F-13 satellite RNA and GFLV strains devoid of satellite.: Production of biologically active transcripts from cloned cDNA of genomic RNAs of GFLV. thus qualifying for use in X. Esmenjaud et al. hybrids and breeding stocks after infection of the roots by means of viruliferous X. index.: Production of GFLV satellite RNA transcripts.:Detection of GFLV in single nematodes by RT-PCR. GFLV is eliminated from somatic embryos obtained from tissue cultures grown at 35 °C 1991 1991 1992 1992 1992a. Walker et al. Staudt and Weischer: Vitis rotundifolia and Vitis munsoniana resist infection by GFLV transmitted by X. O39-16 in particular.: Detection of GFLV in X.: GFLV isolated in Hungary from naturally infected symptomatic plants of Aristolochia clematis and Lagenaria siceraria turbinata. This represents the first substantiated record of a natural GFLV infection in hosts other than Vitis. index by biotin-avidin ELISA Bardonnet et al.b Hans et al. delays symptom expression by 1-2 days and adversely affects virus replication.: Development of cDNA probes to GFLV genomic and satellite RNAs and their use for virus detection directly in grapevine extracts. Horvath et al.
RNase L) genes for inducing resistance to GFLV.: Genetic transformation of Vitis vinifera with the coat protein gene of GFLV for resistance induction. Mauro et al. Gölles et al.: Transformation of Nicotiana species and Vitis rupestris with different virusderived (coat protein. and Asian Vitis species inoculated by green grafting with a GFLV source. Lahogue and Boulard: Search for genes of resistance in grapevines. Spielmann et al. Walker and Jin: V.: Successful transformation of somatic embryos of an European grape cultivar (Russalska 3) with the normal. Belin et al. Pinck: A comprehensive review of the molecular aspects of GFLV genome and its replication strategy.: Attempts to characterize molecularly X. This resistance is controlled by a single dominant gene. rupestris x M. Belin et al.: Identification of the molecular signal accounting for the systemic spread of GFLV in infected hosts. Naraghi-Arani et al.: Identification of membranes derived from the endoplasmic reticulum as sites of GFLV replication.: Variations observed following RT-PCR and RFLP analysis of the coat protein gene of nine GFLV isolates grown in different hosts confirm the quasispecies nature of this virus.: Demostration that the movement protein of GFLV is located on the intracellular tubular structures containing rows of virus particles. index feeding. index populations by PCR-RFLP and sequencing of the ITS region.: The membranous structures appressed to the nucleus of infected cells known as vacuolate-vesiculate inclusion bodies are virus factories as they are the likely site of RNA replication and processing of viral polyproteins. De Luca et al. Krastanova et al. : Identification of RNA2-encoded proteins in the specific transmission of GFLV by X. except for four. 1997 1998 1998 1999 1999 2000 2000 2000 2000 2001 2002 2003 2003 2003 27 . replicase) and exogenous (2. Pfeiffer et al. index. including the two accessions reported as resistant by Walker and Meredith (1990). American. Gaire et al.: Genetic transformation of American roostocks with the coat protein gene of GFLV for resistance induction. Fuchs: Review article on genetic transformation of grapevines for resistance to GFLV and other pathogens. Ritzenthaler et al. rotundifolia hybrids show high resistance to X. The level of resistance obtained looks promising. Pfeiffer et al. Walter and Martelli: Review article on detrimental effects of viruses on grape yields.: Demonstration that a 28 kDa protein coded by GFLV RNA-2 is involved in the replication of this RNA. Of 531 accessions of European. Rowhani et al.1995 1995 1995 1995 1995 1996 1996 Brandt and Himmler: Use of immunocapture RT-PCR for GFLV detection in host tissues. all were susceptible to the virus. Martelli and Walter: Review article on certification of grapevines.: Up to date account of GFLV replication strategy. truncated or nontranslatable coat protein gene of GFLV. 5 oligoadenylate synthase.: Development of a GFLV detection system based on PCR analysis of immobilized virions. Ritzenthaler et al.
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subgroup B. 1978... 1977.F. 3.A. J. Iwanami. and A. San Diego. Many are transmitted by longidorid nematodes (Rüdel. a nematode-borne virus originally classified as a tentative species in the genus Nepovirus. Scholtof.F. Murant (eds). Van Regenmortel et al. Palukaitis.. Bulletin de l'OIV 69. typified by Tomato black ring virus (TBRV). 5. Family Comoviridae. Taylor C. 1955) and are subdivided into subgroups based on physico-chemical properties of member viruses. RT-PCR) are routinely used for their detection in grapevine tissues (primarily cortical scrapings from dormant canes). Martelli and R.. New York. Savino and P. 1996. Eight Report of the International Committee on Taxonomy of Viruses (C. Martelli. Plenum Press. 1981. In: Virus Taxonomy. which are separately encapsidated. Mechanical transmission to herbaceous hosts or indexing on Vitis indicators can also be used. Sixth Report of the International Committee on Taxonomy of Viruses (F. single-stranded RNA genome occurring as two functional species (RNA-1 and RNA-2). Influence des viroses et qualité. Karasev. 1978.. typified by Tomato ringspot virus (ToRSV) (Martelli et al. Istituto Agronomico Mediterraneo. Wellink. 1955. H. 1992. Harrison B. Fauquet. epidemiological. Murant. Vol. causing diseases whose symptoms are similar to. 10281032 Murant A. T. M. Family Comoviridae. has now been assigned to the newly established genus Sadwavirus (Le Gall et al.P.D. a non-taxonomic clustering. T. subgroup C. CAB International. CMI/AAB Descriptions of Plant Viruses No. (E. Seventh Report of the International Committee on Taxonomy of Viruses (M.V. 341-347. Nepoviruses. Taylor and Brown. G. Satellite nucleic acids.e. T. Piazzolla. 35 . 1996. Amsterdam. Ball. G. Milne. 23-39. Taliansky. Polyhedral Virions and Bipartite RNA Genomes.A. family Comoviridae (Goldbach et al. 1ere partie: Effects des viroses sur la culture des vignes et ses produits. Le Gall et al. In: Handbook of Plant Virus Infections: Comparative Diagnosis. Desselberger and L. Quaderno No. Vienna. Le Gall O. These viruses can readily be eliminated by heat therapy or in vitro meristem tip culture... Lehto. Nematode Vectors of Plant Viruses. which were originally included in the Nepovirus group (Harrison and Murant. In: Virus Taxonomy. the Mediterranean and Middle East. Martelli. Rüdel M. Brown. Leibowitz. 2005). ISEM) and molecular assays (hybridization. Kurstak.INFECTIOUS DEGENERATION (EUROPEAN AND MEDITERRANEAN NEPOVIRUSES) Besides GFLV several other nepoviruses can infect grapevine in Europe. i. Sanfaçon. ed). 145-147. Fritsch. 1996. Harrison B.. U. Martelli G..V. ed. eds). 286 pp.J. Several of these viruses have distorting and chromogenic strains and may occur in mixed infections with GFLV. P. 2005. D.A. In: Virus Taxonomy. Gallitelli.J.H.F. 197-238. Walter B. J. C. Their detrimental effects to grapevine culture and products have been summari zed by Walter and Martelli (1996). Jones.P. The Plant Viruses. Mayo M.F. Strawberry latent ringspot virus (SLRSV). (G. K. Murant 1981. physico-chemical. Yoshikawa.A. A.). Bari. and A.. Serological (ELISA. 1992). All have polyhedral particles about 30 nm in diameter and a positive sense. are now classified in the genus Nepovirus.P. Quacquarelli. 1997) and their satellite RNAs (Mayo et al. and molecular characteristics of nepoviruses (Harrison and Murant.D. Murphy et al. V. or indistinguishable from those of fanleaf.G.M. subgroup A typified by Tobacco ringspot virus (TRSV).. A tentative grouping of nepoviruses Phytopathologia Mediterranea 17. Nepovirus Group.E and D. 2000. 2000) are available. K.. eds) Elsevier/Academic Press. eds). M.. References Goldbach R. Nepoviruses. and G. Academic Press. 185. A.P. 2005) Extensive reviews of the biological. 362 pp. Nepoviruses of grapevine and their nematode vectors in the EEC. 945-971. London. Maniloff. Oxon.B. Mayo. 1977) . In: Grapevine Viruses and Certification in EEC Countries: State of the Art. Simons and M. Springer-Verlag. Elsevier/Noth Holland. Sélection clonale de la vigne: sélection sanitaire et sélection pomologique. 1997.A. 807-818. Wetzel and N.
2 x 106 (RNA-1) and 1. consisting of two separately encapsidated molecules with Mol. The genome is a positive sense ssRNA. In other V. This virus has also been found in grapevine in Switzerland. Romania. Its particles are about 30 nm in diameter. ArMV. Rüdel: Transmission of ArMV to grape seedlings by Xiphinema diversicaudatum.: Physico-chemical properties of GFLV. Russo et al. and. SLRSV and TBRV found in grapevines with atypical Reisigkrankheit in West Germany.Genus NEPOVIRUS ARABIS MOSAIC VIRUS (ArMV) 1. Cross-protection between ArMV and GFLV has been reported.4 x 106 and a size of 1104 nt.: Detection of ArMV by ISEM Tanne: Detection of GFLV.: Ultrastructure of ArMV infections in plant tissues Martelli and Lehoczky: Detection of ArMV in grapevine in Hungary. AILV and GCMV. and B). symptoms are of the fanleaf type. and Japan. Iran. Brückbauer and Rüdel: Symptoms of atypical Reisigkrankheit in the vineyard are associated with ArMV in West Germany Bercks et al.: ArMV detected in Japan in grapevines imported from Europe. losses up to 50 % have been recorded. ArMV and TBRV by ELISA in Israel. Coat protein has a single type of subunits with Mr 54 x 103. a typical nepovirus belonging in subgroup A of the genus Nepovirus. always in Germany the severe dieback disease of the cv. Component T is made up of empy protein shells. Hungary.: ArMV. In Germany. It can infect many woody and herbaceous plants and is transmitted to grapevine by the nematode Xiphinema diversicaudatum but not by X. Murant: ArMV description in the CMI/AAB Descriptions of Plant Viruses series. vinifera varieties. Bulgaria. M. and circular about 350 nt in size. respectively. Turkey.: Xiphinema diversicaudatum can transmit ArMV to grapevine.1 x 106 (RNA-2). have a angular outline. USA (California). Quacquarelli et al. Gerola et al. accounting for 27% (component M) and 41% (component B) of the particle weight. Description ArMV. Kerner appears to be caused by ArMV infection. wt of 0. Historical review 1963 1964 1968 1970 1970 1972 1976 1977 1978 1978 1979 1979 1980 1980 1980 1982 Panjan and Saric: ArMV detected in grapevine in Yugoslavia. and sediment as three components (T. Belli et al. Stellmach: Review paper on ArMV in grapevine. whereas components M and Bcontain RNA. TBRV. wt 2. Canada. linear with Mol. Kobayashi et al.: Isolation of ArMV from grapevine in Italy. Two types of RNA-2 molecules have been found which differ slightly in size (3852 and 3711 nt ) but contain a single ORF encoding polypeptides with Mr of 119 and 124 kDa. Vuittenez et al. ArMV occurs often in mixed infections with GFLV in certain areas of France and Italy. Dalmasso et al. 36 . and with other nepoviruses in the Reisigkrankheit complex of western Germany. 2. Transgenic plants expressing the coat protein gene of the virus have been produced. index. the vector of GFLV. The virus supports the replication of two types of satellite RNAs. is serologically related to GFLV. Yugoslavia. Israel.: Interactions between nepoviruses in grapevine and herbaceous hosts.95-2. Jankulova and Kaitasova: ArMV found in grapevine in Bulgaria.
ArMV. The virus cannot be recovered from leaves or buds of the Kerner scions.1982 1984 1985 Brückbauer: Possibility of distinguishing GFLV. Stellmach: Kerner disease is probably caused by ArMV. When a healthy scion is grafted onto an infected rootstock.: Properties of ArMV small satellite RNA.Kerner to ArMV seems to be limited in time.: A hypovirulent ArMV isolate delays GFLV infection in grapevines under field conditions.: ArMV is not seed-transmitted in grapevines Liu et al. RRV. Stellmach and Berres: The susceptibility of cv. Becker et al.: Use of cDNA probes for ArMV detection. MacKenzie et al.: ArMV in Switzerland Lázár et al. Marc-Martin et al. such as RpRSV or GFLV can be found in both rootstock and scion. SLRV and TBRV Belli et al. whereas other nepoviruses.: The presence of ArMV satellite RNA can attenuate symptoms in certain hosts Bertioli et al. Steinkellner et al. Study of histological changes at the graft union level. Hypothesis of a graft incompatibility when the rootstock is infected with ArMV.: Survey for the presence of ArMV in Canadian vineyards Lahogue and Boulard: Search for genes of resistance in grapevines.: Properties of a strain of ArMV isolated from grapevine in Italy. Walter et al.: Evidence that ArMV has two RNA-2 molecules and complete nucleotide sequence of both RNAs Etscheid et al.: Transgenic Nicotiana plants transformed with the coat protien of ArMV produce empty viral shells.: Nucleotide sequence of the ArMV satellite RNA Liu et al.: Transformation of grapevines with the coat protein gene of ArMV. Faber. : Comparison of coat proteins of ArMV and other nepoviruses. whereas the rootstock remains infected.: Cross-protection experiments in Chenopodium quinoa between ArMV and GFLV Gugerli et al. Kaper et al.: Association of ArMV-infected rootstocks with Kerner disease in West Germany. the virus is recovered from the scion only during the first year. Eppler et al. Ipach et al. Yield losses may reach 77% in cv. Molecular assays are as good as ELISA for routine testing. Of 407 accessions of 1986 1987 1988 1989 1989 1989 1989 1990 1990 1990 1991 1991 1992 1992 1993 1993 1994 1995 1995 1995 1996 1996 37 .: ArMV recorded from Romania Huss et al.: Nicotiana plants engineerd with ArMV coat protein gene show different degrees of tolerance to the virus Flak and Gangl: ArMV recorded from Austria Loudes et al.: Nucleotide sequenece of a small circular satelllite RNA Steinkellner et al. Rüdel: In the Palatinate (West Germany) ArMV is transmitted by Xiphinema diversicaudatum and occurs often in mixed infections with GFLV in grapevine.: Detection of ArMV by PCR in herbaceous hosts and grapevines Steinkellner et al.
European, American, and Asian Vitis species inoculated by green grafting with a ArMV source, 42 were apparently resistant. 1998 2000 2003 2004 Akbas and Erdiller: ArMV recorded from Turkey Goelles et al. : Transgenic grapevines expressing ArMV coat protein gene Fuchs: Review on transgenic resistance of grapevines to pathogens Pourrahim et al.: ArMV identified in Iranian grapevines by mechanical transmission to herbaceous hosts and ELISA
3. References Akbas B. and G. Erdiller, 1998. Grapevine virus diseases in Karaman, Konya and Nevsheir provinces. Proceedings VII Congress of the Turkish Phytopathological Society, Ankara 1998, 149-153. Becker A., J. Jäger and B. Altmayer, 1989. Association of arabis mosaic virus-infected rootstocks with the dieback of the Vitis vinifera cv."Kerner" in Germany. Proceedings 9th Meeting of ICVG, Kiryat Anavim, 1987, 57-61. Belli G., A. Fortusini and G. Vegetti, 1982. Il virus del mosaico dell' Arabis isolato da vite in Italia. Rivista di Patologia Vegetale, S IV, 18, 175-177. Belli G., G. Vegetti, S. Cinquanta, C. Soncini, S. Prati and D. Tolentino, 1984. Properties of a strain of arabis mosaic virus isolated from grapevine in Italy. Rivista di Patologia Vegetale, S IV, 20, 56-64. Bercks R., H. Brückbauer, G. Querfurth and M. Rüdel, 1977. Untersuchungen über die Viruskrankheiten der Rebe unter besonderer Berücksichtigung "atypischer Formen" der Reisigkrankheit. Weinberg und Keller 24, 133-180. Bertioli D.J., R.D. Harris, M.L. Edwards, J.I.Cooper and W.S. Hawes, 1991. Transgenic plants and insect cells expressing the coat protein of Arabis mosaic virus produce empty virus-like particles. Journal of General Virology 72, 1801-1809. Brückbauer H. and M. Rüdel, 1976. Untersuchungen über eine "atypische" Form der Reisigkrankheit bei der Rebsorte Silvaner. Weinberg und Keller 23, 53-79. Dalmasso A., M.C. Munck-Cardin and R. Legin, 1972. Résultats préliminaires d'essais de transmission de sérotypes de la mosaïque de l'Arabis trouvés sur vigne, par l'intermédiaire de Xiphinema diversicaudatum. Annales de Phytopathlogie 4, 410 Eppler A., V. Lesan and A. Lázár, 1989 Viruses and virus diseases in some vineyards in Romania. Meded. Facultet Landbouwwetenschappen Rijkuniversiteit Gent 54, 491-497. Etscheid M., M.E. Tousignat and J.M. Kaper, 1995. Small satellite of arabis mosaic virus autolytic processing of in vitro transcripts of (+) and (-) polarity and infectivity of (+) strand transcripts. Journal of General Virology 76, 271-282. Flak W. and H. Gangl, 1994. Grobkartierung des Rebenvirosenbefalls in der Weinbauregion Bungerland mittels ELISA. Mitteilung Klosterneuburg 44, 163-167. Fuchs M., 2003. Transgenic resistance: state of the art. Extended Abstracts 14th Meeting of ICVG, Locorotondo 2003, 221-223. Gerola F.M. M. Bassi and E. Betto, 1964. Shape and localization of arabis mosaic virus in experimentally infected cells of Chenopodium amaranticolor. Phytopathologische Zeitscrift 51, 192-194. Goelles R., R. Moser, H. Puhringer, H. Katinger, M.L. da Camara Machado, A. Minafra, V. Savino, P. Saldarelli, G.P. Martelli and M. Laimer da Camara Machado, 2000. Transgenic grapevines expressing coat protein gene sequences of grapevine fanleaf virus, arabis mosaic virus, grapevine virus A and grapevine virus B. Acta Horticulturae 528, 305-311. Gugerli P., J.J. Brugger and P. Basler, 1990. Les maladies de l'enroulement, du bois strié et de l'ècorce liégeuse de la vigne (grapevine leafroll, rugose wod and corky bark). Revue Suisse de Viticulture, Arboriculture et Horticulture 22, 35-36. Kaper J.M., M.E. Tousignant and M. Steger, 1988. Nucleotide sequence predicts circularity and selfcleavage of 300-nucleotide satellite of arabis mosaic virus. Biochemical and Biophysical Research Communications 154, 318-322. Lahogue F. and G. Boulard, 1996. Recherche de gènes de résistance naturelle à deux viroses de la vigne: le court-noué et l'enroulement. Vitis 35, 43-48. Liu Y.Y., C.U.T. Hellen, J.I. Coper, D.J. Bertioli, D. Coates and G. Bauer, 1990. The nucleotide sequence of a satellite RNA associated with arabis mosaic nepovirus. Journal of General Virology 71, 1259-1263. Liu Y.Y., J.I. Cooper, M.L. Edwards and C.U.T Hellen, 1991. A satellite RNA of arabis mosaic virus and its pathological impact. Annals of Applied Biology 118, 557-587.
Loudes A.M., C. Ritzenthaler, M. Pinck, M.A. Serghini and L.. Pinck, 1995. The 119 kDa and 124 kDa polyprotiens of arabis mosaic nepovirus (isolate S) are encoded by two distinct RNA2 species. Journal of General Virology 76, 899-906. MacKenzie D.J., R.C. Johnson and C. Warner, 1996. Incidence of four important viral pathogens in Canadian vineyards. Plant Disease 80, 955-958. Murant A.F, 1970. Arabis mosaic virus. CMI/AAB Descriptions of Plant Viruses, No. 16 Pourrahim R., A. Ahoonmanesh, Sh. Farzdfar, F.Rakhshandehro and A.R. Golanaraghi. Occurrence of Arabis mosaic virus and Grapevine leafroll-associated virus 3 in Iran. Plant Disease 88, 424. Quacquarelli A., D. Gallitelli, V. Savino, P. Piazzolla and G.P. Martelli, 1979. Some properties of grapevine fanleaf and other nepoviruses infecting the grapevine. Proceedings 6th Meeting of ICVG, Cordoba, 1976. Monografias INIA No. 18 , 41-49. Rüdel M., 1978. Übertragung des Arabis-Mosaik-Virus (AMV) durch den Nematoden Xiphinema diversicaudatum (Micoletzki) Thorne auf Rebensämlinge (Vorläufige Mitteilung). Die WeinWissenschaft 33, 243-247. Rüdel M., 1985. Grapevine damage induced by particular virus-vector combinations. Phytopathologia Mediterranea 24, 183-185. Steinkellner H., G. Himmler, M. Laimer, D. Mattanovich, G. Bisztray and H. Katinger, 1989. Konstruktion von cDNA von Arabis Mosaik Virus und deren Anwendung für Diagnose. Mitteilung Klosterneuburg 39, 242-246. Steinkellner H., G. Himmler, R. Sagl, D. Mattanovich, and H. Katinger, 1992. Amino acid sequence comparison of nepovirus coat proteins. Virus Genes 6, 197-202. Steinkellner H., A. da Camara Machado, M. Laimer-da Camada Machado, M. Gölles and H. Katinger, 1993. Studies on coat protein mediated cross protection of nepoviruses. Extended Abstracts 11th Meeting of ICVG, Montreux 1993, 175. Stellmach G., 1970. Arabis mosaic in Vitis. In: Virus Diseases of Small Fruits and Grapevines- A Handbook (N.W. Frazier, ed). University of California, Division of Agricultural Sciences Berkeley, 233-234. Stellmach G., 1987. Die Kerner Krankheit: Theoretische und praktische Aspekte einer tödlichen Rebenvirose. Die Wein-Wissenschaft 42, 421-427. Stellmach G. and R.-E. Berres, 1986. Begrenzte Infektionsanfälligkeit der Vitis vinifera- Sorte "Kerner" gegenüber dem Arabismosaik-Virus? Zeitschrift für Pflanzenkrankheiten und Pflanzenschutz 93, 356-360. Walter B, P. Bass, P. Cornuet and P.M. Guillaume, 1993. Preliminary results of cross protection experiments against grapevine fanleaf virus (GFLV) in the vineyards. Extended Abstracts 11th Meeting of ICVG, Montreux 1993, 167-168 ARTICHOKE ITALIAN LATENT VIRUS (AILV) 1. Description Artichoke Italian latent virus (AILV), a member of subgroup B of the genus Nepovirus, was isolated in Bulgaria from vines with fanleaf-like symptoms. AILV has isometric particles with angular outline, sedimenting as three components: T (empty shells), M (particles contaning a molecule of RNA-2 with Mol. wt of 1.5 x 106 daltons accounting for 34% of the particle weight) and B (particles containing a molecule of RNA-1 with Mol. wt of 2.4 x 106 daltons, accounting for 41% of the particle weight). Coat protein is made up of a single type of subunits with Mr 54 x 103. AILV is transmitted by the Dorylaimoid nematode Longidorus apulus in vegetable crops but no field transmission to grapevines has been recorded. The virus has limited distribution and economic importance. 2. Historical review 1976 Jankulova et al.: AILV isolated in southern Bulgaria in 1976 from a grapevine with fanleaf-like symptoms. Properties of the virus, cultured in Chenopodium quinoa determined and positive serological reaction with an antiserum to an Italian strain of AILV ascertained. Savino et al.: Comparison of a Bulgarian grapevine isolate of AILV with an Italian isolate from artichoke and two Bulgarian isolates from sowthistle and gladiolus. Martelli et al.: AILV description in the CMI/AAB Descriptions of Plant Viruses series.
3. References Jankulova M., V. Savino, D. Gallitelli, A. Quacquarelli and G.P. Martelli, 1979. Isolation of artichoke Italian latent virus from the grapevine in Bulgaria. Proceedings of the 6th Meeting of ICVG, Cordoba 1976. Monografias INIA 18, 143-148. Martelli G.P., G.L.. Rana and V. Savino, 1977. Artichoke Italian latent virus. CMI/AAB Descriptions of Plant Viruses, No. 176. Savino V., D. Gallitelli, M. Jankulova and G.L. Rana, 1976. A comparison of four isolates of Artichoke Italian latent virus (AILV). Phytopathologia Mediterranea 16, 41-50. CHERRY LEAFROLL VIRUS (CLRV) 1. Description Cherry leafroll virus (CLRV) is a cosmopolitan virus. In Chile it was recovered from vines with fanleaflike symptoms and in Germany from vines with yellow mosaic-like symptoms. Although CLRV is a definitive nepovirus species classified in subgroup C of the genus Nepovirus it differs from most of the other members in the genus being transmitted by pollen rather than nematodes. The vector to grapevine, if any, is unknown. CLRV occurs in nature as multiple strains but is not serologically related to any of the known nepoviruses. Virus particles are isometric, about 30 nm in diameter have angular outline and sediment as three components (T, M, and B). Their coat protein consists of a single type of subunits with Mr of about 54 kDa. The genome is a bipartite, positive sense, single-stranded RNA which has been sequenced only in part. Genomic RNA consists of two separately encapsidated functional molecules with Mol. wt of 2.8 x 106 (RNA-1) , accounting for 46% of the particle weight, and 2.3 x 106 (RNA-2), accounting for 41% of the particle weight. In grapevines CLRV is readily detected by DAS-ELISA. The best woody indicator for the German isolate is reported to be Pinot noir. 2. Historical review 1985 1993 2001 2003 Jones: Description of Cherry leafroll virus in the AAB Descriptions of Plant viruses series. Scott et al.: Partial nucleotide sequence of CLRV RNA-2. Herrera and Madariaga: First record of CLRV from Chile. Field infection is estimated to be 0.2% Ipach et al.: Isolation of CLRV from German vines showing yellow mosaic-like symptoms and reduced crop.
3. References Herrera M.G and V.M. Madariaga, 2001. Presence and incidence of grapeivne viruses in central Chile. Agricultura Tecnica 61, 393-400 Ipach U., L. Kling and D.E. Lesemann, 2003. First record of Cherry leafroll virus on grapevine in Germany. Extended Abstracts 14th Meeting of ICVG, Locorotondo, 2003, 17-18 Scott N.W., J.I. Cooper, and M.L Edwards, 1993. The identification, cloning, and sequence analysis of the coat protein coding region of a birch isolate of cherry leafroll nepovirus. Archives of Virology 131, 209-215. GRAPEVINE ANATOLIAN RINGSPOT VIRUS (GARSV) 1. Description Grapevine Anatolian ringspot Virus (GARSV) was isolated from Turkish grapevines with mild fanleaflike symptoms. The virus belongs in subgroup B of the genus Nepovirus but is not serologically related to any of the known grapevine nepoviruses. Virus particles are isometric c. 30 nm in diameter and sediment as three centrifugal components. RNA-1 has a Mol. wt of 2.2 x 106 Da and RNA-2 a Mol. wt of 1.4 x 106 Da and a size of 4607 nt. Coat protein subunits have a Mr 56 x 103 Da. GARSV can be readily detected by ELISA and PCR using primers designed on the coat protein sequence. The virus has no recognized
GBLV has also been recorded from Hungary and Yugoslavia. Coat protein has a single type of subunits with Mr 54 x 103. Bulletin OEPP/EPPO Bulletin 32. M. M. Historical review 1972 1972 1977 Ferreira and De Sequeira: Description and preliminary characterization of an unidentified virus denoted CM112. Martelli et al. Complete nucleotide sequence of RNA-2 of two Turkish nepoviruses. 2003. GBLV is a typical nepovirus belonging in subgroup C of this genus but its vector is not known. Properties of a previously undescribed nepovirus fron south-east Anatolia.: Complete nucleotide sequence of GARSV RNA-2 3. Sanitary status of grapevine in south eastern and central Anatolia.: Description and thorough characterization of GARSV identified as a new species in the subgroup B of the genus Nepovirus Abou Ghanem-Sabanadzovic et al.1 x 106 (RNA-2). Cigsar I.95-2. isolated in 1970 in Portugal from symptomless vines. Martelli. is not seed borne and was reported only from south-eastern Turkey. 335-340. Concord in New York State. 35-41. consisting of two separately encapsidated molecules with Mol. but different from GBLV. Cigasr. Martelli et al. where it is widespread and infects latently several grapevine varieties growing in widely separared areas. D. Virus particles are about 30 nm in diameter and sediment as three components (T.P. Martelli. The scattered distribution of infected vines in the field suggests that the virus is spread primarily by infected propagative material. Component T is made up of empy protein shells. The virus can be detected directly in grapevine leaf extracts by immunodiffusion in agar plates. De Stradis. N. Kizlar tahtasi showing mild fanleaf-like symptoms Gokalp et al. Virus Genes 30. accounting for 39% (componet B1) and 42% (component B2) of the particle weight. and B2).: A virus serologically related to GBLV isolated from Vitis labrusca cv. A strain of this virus had been found previously in Portugal and described as virus CM112.vector. GRAPEVINE BULGARIAN LATENT VIRUS (GBLV) 1. References Abou Ghanem-Sabanadzovic N. Digiaro and G. 2002. A. Boscia and G. Martelli. The genome is a positive sense ssRNA.2 x 106 (RNA-1) and 1. M. whereas components B1 and B2 contain RNA. Digiaro. Digiaro and G. Uyemoto et al. Journal of Plant Pathology 85. The virus supports the replication of a satellite RNA with Mol.: First isolation by mechanical transmission of an unknown nepovirus from cv.. 1977 1978 41 . S. wt of 2.P. Historical review 2002 2003 2005 Cigsar et al. Sabanadzovic. Description Grapevine Bulgarian latent virus (GBLV) owes it name to the fact that it was found for the first time in Bulgaria in 1971.5 x 106 (less than 1800 nt). 2005.P. B1. 471-475 Gokalp K. I. 2. 2.. The virus occurs as different closely related but serologically distinguishable strains. a virus serologically related to GBLV found in Concord grapes in New York State vineyards is a strain of Blueberry leaf mottle virus (BLMoV) a North American nepovirus species related to. De Mendonça et al.: Description of GBLV. physico-chemical and serological characterization of the virus and assignment to the Nepovirus group (now genus Nepovirus).: Isolation of virus CM112 from in vitro cultures of grapevine tissues. wt 0.: GBLV description in the CMI/AAB Descriptions of Plant Viruses series. Biological. Two isolates of GBLV have been transmitted by mechanical inoculation to seedlings and rooted cuttings of several grapevine cultivars without inducing symptoms. By contrast. Abou Ghanem-Sabanadzovic. The economic importance of the virus is minor.
A.. Savino and O. Abracheva. Monografias INIA 18 . Pereira and V.A. 95-101. Annals of Applied Biology 85.: Detection of GBLV in grapevine leaf extracts by ISEM. Ganeva and M. Abracheva. Annales de Phytopathologie.N. Colmar 1970. Gallitelli. Quacquarelli. and J. De Sequeira. Martelli G. Proceeding 4th Meeting of ICVG. V. 217-222. M. Pocsai: Occurence of GBLV in Hungary. O. Ramsdell D.C. Quacquarelli. Krastanova S. Grapevine Bulgarian latent virus. Colmar. Gallitelli D. M. 51-58. 269-272. A. Numéro hors série. 245-250.P. 113-120. and O. Varennes and De Sequeira: First application of ELISA for the detection of virus CM112. Properties of a distinctive strain of Grapevine Bulgarian latent virus.Proceedings 7th Meeting of ICVG. 468-472. 1992. Savino and A. Pocsai E. Occurence of grapevine Bulgarian latent virus in Hungary.: Detection of virus CM112 in grapevine leaf extracts by ISEM. Gallitelli et al. M. Proceedings 6th Meeting of ICVG.: A comparative study of Bulgarian GBLV isolates and the Portuguese virus CM112 establishes that CM112 is a serologically close but distinguishable strain of GBLV. Dimitrijevic: GBLV found in Yugoslavia De Mendonça et al.P. V. Niagara Falls 1980. Some properties of grapevine Bulgarian latent virus. P. quinoa. 135-141. and R. Niagara Falls 1980. Niagara Falls 1980. De Sequeira.P.A. Annales de Phytopathologie. 42 . Physical and chemical properties of the particles and ribonucleic acid of blueberry leaf mottle virus. 1980. Garcia de Orta Estacao Agronomica 12.. De Sequeira O.. Russo et al. Preliminary studies on an undescribed grapevine virus. The ultrastructure of grapevine Bulgarina latent virus infections in natural and artificial hosts. Yankulova. 1979. 1985.A. 349-354. Proceedings 7th Meeting of ICVG. The Portuguese strain supports the replication of a satellite RNA. Quacquarelli and D.: A comparative study of three GBLV isolates from Bulgaria shows that they are closely related but serologically distiguishable and that can infect seedlings and rooted cuttings of different grapevine cultivars without inducing symptoms. O. 143-145 De Mendonça A. 1980. P.. D. 1972. Gallitelli. 1983. Dimitrijevic B. 1981. 1981. An immunoradiometric assay for the titration of a Portugese strain of grapevine Bulgarian latent virus (GBLV).. Proceeding 4th Meeting of ICVG. 1980.A. D. 1972. De Sequeira and A. A. Jankulova. Simoes. Vasconcelos-Costa.. Phytopathology 71. Martelli G. France. Ferreira. No. Savino. De Sequeira and Vasconcelos-Costa: Use of an immunoradiometric assay for the titration of the Portuguese strain of GBLV. A manually transmissible latent virus of the grapevine from Bulgaria. Savino and A.A. V. Di Franco. Cordoba 1976. Rasteniev'dni Nauki 29. Acta Phytopathologica Academiae Scientiarum Hungaricae 16. Ferreira A.. Ramsdell and Stace-Smith: The New York isolate of GBLV is a strain of BLMoV. De Sequeira. Gallitelli. 186 Martelli G.. 27-32. Numéro hors série. D. Mota.. 1978. A preliminary report.P. CMI/AAB Descriptions of Plant Viruses. Phytopathologia Mediterranea 22. 1980 1980 1980 1980 1981 1981 1982 1983 1985 1992 3. Stace-Smith.1979 Martelli et al. 1977. Martelli G. Proceedings 7th Meeting of ICVG. 1970.: Improvement of ELISA protocol for GBLV detection the whole year round. A. Krastanova et al.. 21-24.Sur l'isolement d'un virus à partir de cultures de tissus de vigne. Russo and V.A. Some properties of the new latent virus from grapevine rootstocks in Yugoslavia. References De Mendonça A. Possibilities for the whole-year ELISA detection of viruses infecting grapevines.: Ultrastructural study of GBLV infections in grapevine and C. Applicability of immunosorbent electron microscopy (ISEM) for the detection and identification of CM112 virus in grapevine. Martelli et al.
index. Croatia and Austria. a size of 7212 nt and accounts for 40% of the particle weight. However. 2.K.8 x 106 . transmitted to herbaceous hosts from Hungarian grapevines with symptoms similar to those of fanleaf and yellow mosaic. index. near Lake Balaton. 1977. called Hungarian chrome mosaic virus. Tobacco pants and the rootstock 110R have been successfully transformed with the viral coat protein for induction of resistance. Mali et al. De Sequeira. Plant Disease Reporter 61. vuittenezi is very frequently found in vineyards with chrome mosaic patches. Lehoczky and Tasnady: A study of the effect of HCMV on yield and sugar content of infected grapevines.: GCMV recorded from Slovakia and report of X. GRAPEVINE CHROME MOSAIC VIRUS (GCMV) 1. The coat protein has a single type of subunits of Mr 52 x 103. vuittenezi transmits GCMV or GFLV. Kenten: GCMV is distantly serologically related to Cacao necrosis virus.: Host range and properties of a spherical virus. The virus appears to be unrelated serologically to GFLV and is not transmitted by X. No evidence that X. index as vector of the virus (unconfirmed results). GCMV is transmitted through grapevine seeds. Although GCMV particles have been detected by immunosorbent electron microscopy in Xiphinema index fed on infected hosts. 949-953. Varennes A. and was originally called Hungarian chrome mosaic virus. wt of 1. and O. wt of 2. sometimes together with X. a size of 4441 nt and accounts for 31% of the particle weight. 1966 1968 1968 1969 1969 1971 1972a 1972b 1972 1975 1977 43 . Agronomia Lusitana 41. Leaves of infected vines are partially or entirely bright yellow or whitish. 269-277. Affected vines lack in vigour and may decline and die. Saric and Hranuelli: GCMV recorded from Croatia. double nodes and short internodes. Description Grapevine chrome mosaic virus (GCMV) was first found in Hungary. Isolation and identification of a strain of grapevine Bulgarian latent virus in Concord grapevines in New York State. Martelli et al. RNA-2 has Mol. Jakó et al: HCMV affects pigment and sugar content of infected grapevine leaves. Virus re-named Grapevine chrome mosaic virus.Uyemoto J. It has been recorded also from Czechoslovakia. Taschenberg and D.A. RNA-1 has Mol. Martelli and Quacquarelli : Physico-chemical characterization of HCMV and comparison with TBRV. Martelli and Sarospataki: X.K. The virus is not serologically related with GFLV. Pozsár et al.F. The virus belongs in the same subgroup of TBRV (subgroup B) to which is distantly related serologically. Detection of CM122 latent grapevine virus by enzyme-linked immunosorbent assay (ELISA). Hummer. Historical review 1966 Martelli et al. symptomless infection may occur.: HCMV adversely affects photosynthetical carbon dioxide fixation. Evaluation of short reaction times and re-use of a-globulin and conjugate. E. pretty much like GFLV..: The isometric virus associated with Hungarian chrome mosaic is serologically distantly related to Tomato black ring virus (TBRV). The genome is bipartite. early reports that this nematode could transmit the virus have not been confirmed. Martelli and Quacquarelli: Description of HCMV in the CMI/AAB Descripitons of Plant Viruses series. Some virus strains induce leaf deformity. a symptom virtually indistiguishable from GFLV-induced yellow mosaic. Martelli: Purification and serology of the virus isolated from Hungarian grapevines with fanleafand yellow mosaic-like symptoms. 1982.6 x 106 .
: Description of a certification scheme for the production of virus-free propagating material in Hungary. O. Le Gall.M.: Detection of GCMV in leaf dips by ISEM. Candresse. 3. Lehoczky: Pinot noir and Jubileum 75 are good indicators for GCMV.: Up-to-date report on virus diseases of grapevines in Hungary and description of the clean stock programme implemented in the country. Le Gall and J. 7809-7819. and G. Dunez. Savino. Lehoczky et al. Bretout et al.: Complete nucleotide sequence of GCMV RNA-1.. Candresse. Dunez. L. index are inconclusive. Delbos.P. Candresse. Genetically engineered resistance against grapevine chrome mosaic virus. A. 1995. Occurrence of grapevine chrome mosaic nepovirus in Austria.: Transformation of roostock 110R with the coat protein gene of GCMV.: GCMV recorded from Austria. Acta Horticulture 235. M. Himmler. Russo et al. Doz et al. V. R. T. 44 . Roberts and Brown: Detection of GCMV in X. 1993.: GCMV and TBRV can recombine. Dimou D. M. 231-238. Further demonstration that the two viruses are related. Dodd and Robinson: GCMV and TBRV are molecularly related.: Development of molecular probes for GCMV detection.: GCMV cross-protects Chenopodium quinoa from the severe apical necrosis induced by a TBRV strain. Le Gall. 89-97. Brault V. Laimer da Camara Machado and V. Nucleic Acid Research 17. Le Gall et al. O. Virus and RNAspecific molecular hydridization probes for two nepoviruses. T.: Tobacco plants genetically engineered with the coat protein gene of GCMV are resistant to infection. Brault et al. Nucleotide sequence and genetic organization of Hungarian grapevine chrome mosaic nepovirus RNA2. Lanneau and J. References Brandt S. M. Plant Molecular Biology 21. 1989. Ravelonandro and J. Le Gall et al. Brault et al. Brault. No assessment of resistance made. Hilbrand. Detection of nepoviruses in ligneous grapevine material using IC/PCR. The virus vector is yet to be identified. Dunez. Brandt and Himmler: Development of a IC-PCR protocol for GCMV detection in cortical scrapings from dormant grapevine canes. D'Onghia. Vitis 34. Bretout C. Journal of Phytopathology 142.: GCMV detected by ELISA in infected field-grown vines. 1994.: Characterization of a GCMV strain and confirmation of its serological relationship with TBRV. Taylor and Brown: Results of GCMV transmission trials with X. 1989. Dimou et al. This finding does not imply vectoring capacity by this nematode. Le Gall et al.. T. 258-262. Brault V. Lázár et al. Lázár et al: Seed transmission of GCMV in grapevine. Kölber et al.. O.1979 1980 1980 1982 1984 1985 1985 1989 1989 1989 1990 1993 1993 1994 1994 1995 1995 1997 2000 Lehoczky et al. 127-128.: Complete nucleotide sequence of GCMV RNA-2. index extracts by ISEM.
J. Phytopathologia Mediterranea 24. 58-72. Bojnansky. 1990. Hungarian chrome mosaic of grapevine and tomato black ring: two similar but unrelated plant viruses. 29-44.P. Kölber M. Studies on the nitrogen metabolism of vines infected with yellow mosaic virus.. 1989.. T.J. Proceedings International Conference on Virus and Vector on Perennial Hosts.. Dunez. Lehoczky J. The effect of fanleaf and chrome mosaic virus diseases on yield and the fruit sugar content of grapevine. Torregrosa .. 1972b. Acta Phytopathologica Academia Scientiarum Hungaricae 3. 1966. 123-141. 1-130. Martelli G. 236-237.P. Pozsár B. Danglot. 1985. Luntz. A Handbook.P. Beczner. Annales de Phytopathologie. 1982. Delbos and J. and S. Sarospataki and J. 1995.. 402-410. Candresse.. Lehoczky and A. 172-173. Vanek and V. University of California.. Brault and J. S. Lehoczky J. T. and D. J. NW Frazier (ed.M. Sarospataki. Numero hors série. 1968. Mali V. Davis 1965. Lehoczky and G. and A... 505.. Journal of General Virology 65. 1969. O. Quacquarelli and J. ArMV) in the seeds and seedlings of grapevine by ELISA. Le Gall O.. 165-173. The change of pigment and sugar content in the chrome mosaic virus infected leaves of grapevine. Annals of Applied Biology 71. Jakó N. Serologische Verwandschaft eines mit dem ungarischen "chrome mosaic" vergesellschafteten Virus mit einem Stamm des "tomato black ring virus". Nucleotide sequence of Hungarian grapevine chrome mosaic nepovirus RNA1. Journal of General Virology 76. G. Annales de Phytopathologie 11. 119-126. Sarospataki. University of California. Agrobacterium-mediated genetic transformation of grapevine somatic embryos and regeneration of transgenic plants expressing the coat protein of grapevine chrome mosaic nepovirus (GCMV).. 1985. J. E. Nucleic Acid Research 17. 1285-1289. J. S. 103. J. Mikulás. 3. Lázár. Dunez.H. Grapevine virus diseases and clean stock program in Hungary. Pol'nohopodarska Veda .P. Berkeley. 1970. 1993. Cardin. Nouvelle évidence d'une parenté sérologique éloignée entre ce virus et celui deas anneaux noirs de la tomate (tBRV). Preliminary report on purification and serology of a virus associated with Hungarian grapevines showing macroscopic symptoms of fanleaf and yellow mosaic. Les Colloques de l'INRA 11. Phytopathologia Mediterranea 24. G. 1984. Prémunition: une competition entre souches faibles et souches sévères pour un voie commune d'expression de symptômes. a serotype ot tomato black ring virus. 567-568. No. Acta Phytopathologica Academia Scientiarum Hungaricae 1. 1972a. 1972. Proceedings International Conference on Virus and Vector on Perennial Hosts. In: Virus Diseases of Small Fruits and Grapevines. Caractérization d'une souche du virus de la mosaïque jaune crome de la vigne (GCMV) isolée en Hongrie de vignes non panachées. GFV-YM. Transfer of the 3' non-translated region of grapevine chrome mosaic virus RNA-1 by recombination to tomato black ring virus RNA-2 in pseudorecombinant isolates. Nematodes of the family Longidoridae (Thorne 1935) Meyl 1960 found in Hungarian vineyards and virus transmission trials with Xiphinema index Thorne et Allen.. 171-170. Kertgazdasag 22.P. Weinberg und Keller 15. Lehoczky and A. Kölber. Bouquet. Jakó N. Nucleotide sequence homologies among RNA species of strains of tomato black ring virus and other nepoviruses. Hajdú. Kölber M. and A. Lázár J. L. Adelaide 2000. Phytopathologia Mediterranea 8. Martelli G. Grapevine chrome mosaic virus.) University of California. A. Lehoczky. Quacquarelli. Lehoczky . Lehoczky and G. Vitis 8. Detection of grapevine chrome mosaic virus in field-grown vines by ELISA. 1971. G. Host range and properties of a virus associated with Hungarian grapevines showing macroscopic symptoms of fanleaf and yellow mosaic. 49-64. Kiserletugyi-Kozlemenyek 64 (1-3). Martelli G. 1731-1740. 1975.. 1969. Lehoczky. Effect of the grape chrome mosaic and grape fanleaf-yellow mosaic virus infection on the photosynthetical carbon dioxide fixation in vine leaves. 2000. Detection of some nepoviruses (GFV. Le Gall O. Martelli G. R. 7795-7807.C. Robinson. 1968.Dodd S.. Division of Agricultural Sciences. A. with Special Reference to Vitis. CMI/AAB Descriptions of Plant Viruses. Lehoczky J. Quacquarelli. Quacquarelli. Le Gall O. Farkas. Extended Abstracts 13th Meeting of ICVG.. S. Montreux 1993.. and G. Sznyegi.P. Quacquarelli. Lehoczky. Martelli G. Horváth. 45 . 1979. with Special Reference to Vitis.J. Plant Science. Szonyegi and M. M. Martelli G.Tasnady. 1966.R. 129-134. Candresse and A. 169-170. Candresse and J. GCMV. 206-210. Sarospataki. Muranyi . The purification and some properties of cocoa necrosis virus. 102. M. 135-140. 1994. V. Y. Transmission by nematodes of some grapevine viruses occurring in Czecholovakia and Hungary. and G.. L. Kenten R. Martelli G. Vuittenez.Ser. Kuszala and A. T. Detection of grapevine chrome mosaic virus in naturally infected vines by indexing.. Kölber and J.P. Division of Agricultural Sciences. Certification scheme for production of virus-free grape propagating material and its results in Hungary. L. Doz B. Davis 1965. Extended Abstracts 11th Meeting of ICVG. 389-401. L. 185-192. 1966. J. Hungarian chrome mosaic. Lázár J. Pacsa and J. Lehoczky J. 1-7. G. J. Dunez. Division of Agricultural Sciences. Devergne. Sarospataki. J..
: Description and thorough characterization of GDefV. and belongs in the subgroup C of the genus Nepovirus. Boscia and G. The genome is bipartite. E.3 x 106 Da and a size of 3753 nt. No vector is known and no information is available on the distribution and economic importance of the virus. 335-340 Cigsar I. Martelli. 5. Martelli and V.P. including all those known to infect grapevine. K.Russo M. Proceedings of the Conference on Excoriosis and Virus Diseases of Grapevine. 2. The virus sediments as three components: T (empty shells). wt of 2 x 106 daltons and apparent size of c. Taylor C. mol. showing leaf and cane deformations Cigsar et al.The virus has no recognized vector. GDefV is readily detected by ELISA and PCR using primers designed on the coat protein sequence. 2. Description Grapevine Tunisian ringspot virus (GTRSV).4 x 106 daltons and apparent size of c. Mostar 1977. Description Grapevine deformation virus (GDefV) was recovered from Turkish grapevines showing distinct fanleaf-like symptoms. 1977. Grapevine deformation virus. Investigations on grapevine viruses in Croatia. Saric A. 1980. Bulletin OEPP/EPPO Bulletin 32.800 nt) and B (particles containing a molecule of RNA-1 with Mol. is not seed-borne and reported only from south-eastern Turkey. 149-151. Virus Genes 30... Martelli. is distantly related serologically to ArMV but not to GFLV.P. S. Oxon.J. 286 pp. G. GRAPEVINE TUNISIAN RINGSPOT VIRUS (GTRSV) 1. Gokalp. M. Sanitary status of grapevine in south eastern and central Anatolia.800 nt. GTRSV is serologically unrelated to any of 19 nepoviruses tested. Hranuelli.). was isolated from a Tunisian grapevine with mild fanleaflike symptoms.. Proceedings of the 7th Meeting of ICVG. 30 nm in diameter and sediment as three components. Savino. RNA-1 has a mol. Martelli. Nematode Vectors of Plant Viruses. Complete nucleotide sequence of RNA-2 of two Turkish nepoviruses. wt of 1. identified as a new species in the subgroup A of the genus Nepovirus.F Brown. wt of 2. Immunosorben electron microscopy for detecting saptransmissible viruses of grapevine. and T. Based on its properties the virus appears to be a new nepovirus serologically unrelated to any of 19 members of the genus and has no known vector. : Complete nucleotide sequence of GDefV RNA-2 3. The scattered distribution of infected vines in the field suggests that the virus is spread primarily by infected propagative material. Niagara Falls 1980. References Abou Ghanem-Sabanadzovic N. A. distantly serologically related with ArMV.. 2005. M (particles contaning a molecule of RNA-2 with Mol. 1997. Digiaro. Particles are isometric c. Historical review 2002 2003 2005 Cigsar et al. 251-257. GRAPEVINE DEFORMATION VIRUS (GDefV) 1. De Stradis. CAB International. 35-41. Digiaro and G.: First isolation by mechanical transmission of an unknown nepovirus from cv. 2002.P. Digiaro and G. Coat protein subunits have a Mr 53 x 103. Abou Ghanem-Sabanadzovic et al. 46 . 2003.6 x 106 Da and RNA-2. Journal of Plant Pathology 85. The virus belongs in the subgroup A of the genus Nepovirus. N. M. and D. Sabanadzovic. 6. 471-475 Cigsar I.: A mechanically transmissible virus was recovered by sap inoculation from Tunisian grapevines showing mild fanleaf-like symptoms. M.. Abou Ghanem-Sabanadzovic. wt of 2. a novel nepovirus from Turkey. Historical review 1991 Ouertani et al. D.P.
Brown. Scottish and English. Mayo. Jolly. CMI/AAB Descriptions of Plant Viruses. Particles are about 30 nm in diameter. M.A. 88-118. Archives of Virology 126. D. M. Reustle.F. Castellano. The viral genome is a bipartite positive sense ssRNA. V. Mögliche Beziehungen zwischen Virus und Symptomausprägung bei der Rebe. The coat protein has a single type of subunits with Mr 54 x 103.6 x 106 (RNA-2).6 x 106 (RNA-1) and 1. These differences strongly suggest that the grapevine-infecting RpRSV may be a different viral species. 16. Die Wein-Wissenschaft 37. Brückbauer: RpRSV can be distinguished from other nepoviruses on the basis of symptoms induced on Vitis idicator plants Blok et al.: Sequencing and molecular characterization of two German isolates of RpRSV from grapevine 2003 3.J. This strain differs considerably from the English type strain of the virus although is serologically closely related to it. 1994.C. W. A Schnabel. The nucleotide sequence of RNA 2 of raspberry ringspot nepovirus.3. McGavin. No. 283-300.P. Raspberry ringspot virus. References Blok V. and sediment as three components (T.. D. 198. The type strain of RpRSV is transmitted by Longidorus macrosoma but the grapevine strain is transmitted by Paralongidorus maximus. Journal of General Virology 73. Savino.M.A. Robinson. consisting of two separately encapsidated molecules with Mol. C. and B). Greco. Edwards and M. Brückbauer H. Extended Abstracts 14th Meeting of ICVG. accounting for 29% (component M) and 43% (component B) of the particle weight.: Biological and physico-chemical characterization of the grapevine strain of RpRSV. 1992. Murant: Description of RpBRSV in the CMI/AAB Plant Virus Description series Stellmach and Querfurth: Study of a strain of RpRSV isolated from cv. Properties of an unusual isolate of raspberry ringspot virus from grapevine in Germany and evidence of its possibile transmission by Paralogidorus maximus. Two strains of different virulence occur in the Palatinate. Murant A. D. Crop losses can be higher than 30 %. G.F. Symptoms are similar to those of fanleaf. Description Raspberry ringspot virus (RpRSV) is a nepovirus belonging in subgroup A of this genus. Locorotondo 2003.. RASPBERRY RINGSPOT VIRUS (RpRSV) 1. Historical review 1970 1978 1978 1982 1992 1994 Vuittenez et al. Boscia. Altmayer.J. The virus has only been found in grapevine in western Germany. 2. Properties of a previously undescribed grapevine nepovirus from Tunisia. A. Jones A. 2189-2194.A. Manoukian. A.. References Ouertani R.L. Molecular characterization of two German Raspberry ringspot virus isolates infecting grapevines and construction of full length infectious clones. The grapevine strain of this virus is serologically very distantly related to the two main serotypes. Heat therapy of infected grapevines. 1978. Wardell J.T. M. Minafra. G. 2003. The virus is transmitted by Paralogidorus maximus Ebel et al.J. Rüdel and B. G. Wetzel. M. RNA-2 is 3928 nt in size and contains a single ORF encoding a polypeptide with Mr of 124 kDa. have angular outline. and differs from the type strain for it often sediments as if it were a single centrifugal component. wt of 2. 107-117. 47 . Martelli and N. 1982. Annals of Applied Biology 124. Krczal and T. Ebel R..: Nucleotide sequence of RpRSV RNA-2 Jones et al. FS4 is a good indicator. 1992. Elbling in West Germany.: Recovery of RpRSV from grapevines of Palatinate.
Tanne: Detection of TBRV by ELISA in Israel. either by agar gel diffusion with extracts of herbaceous hosts previously infected mechanically from grapevine. and G. J. RpRV and TBRV detected serologically in grapevine in West Germany. 1970. RNA-1 is 7356 nt in size and contains a single open reading frame encoding a polypeptide with Mr of 254 kDa. Annales de Phytopathologie 2. rings and lines on the leaves of recently infected plants. wt of 0.. Querfurth. Untersuchungen zur Serologie. is a strain of this virus. then in Yugoslavia. 1978.virus). Meyer et al. about 30 nm in diameter have angular outline and sediment as three components (T. respectively. Virus particles are isometric. Greece.: RRV. Weinberg und Keller 25. Bercks and Querfurth: GFLV. RRV and TBRV are not transmitted by contact of roots or foliage in the vineyard.: Nucleotide sequence of a TBRV satellite RNA. du virus des anneaux noires de la tomate (tomato black ring) et du virus des taches annulaires du framboisier (raspberry ringspot) chez des vignes du Palatinat. riparia 143 A in West Germany. and B).5 x 106 daltons and a size of 1327 nt. The genome is a bipartite positive sense single-stranded RNA occurring as two separately encapsidated functional molecules with Mol. The vector to grapevine is Longidorus attenuatus. Stobbs and Van Schagen: First record of TBRV from Canada. and an increase in graft failure. Bercks: Comparison of three serological tests for detecting several viruses. The virus is a definitive nepovirus species classified in subgroup A of this genus. Joannes Seyve. mottling of the older leaves. 128-136. M. 2. M. 279-327 TOMATO BLACK RING VIRUS (TBRV) 1. TBRV supports the replication of a satellite RNA with Mol. Joannes Seyve virus. known to cause severe damage to the grapevine variety Joannes Seyve in Ontario. latex test and barium sulfate test. Détection et étude selogique du virus latent des taches annulires du frasier (strawberry latent ringspot). Stellmach and Bercks: Further investigations on TBRV in grapevine. and Ontario (Canada). Israel. RNA-2 is 4662 nt in size encoding a polyprotein with Mr of 150 kDa.65 x 106 (RNA-2) accounting for 44% and 37% of the particle weight. Historical review 1963 1965 1966 Stellmach and Bercks: TBRV detected in rootstock Aramon x V. The virus was detected in grapevines in the Niagara Peninsula. SLRV and TBRV found in grapevine in the Palatinate. Vuittenez et al. chlorotic spots. but they can be high. 1967 1970 1970b 1976 1977 1980 1984 1984 1986 48 . Losses are not known precisely.7 x 106 (RNA-1) and 1.Stellmach G. Pathologie und Thermo-Labilität mehrerer Reben-Isolate des Himbeerringflecken-Virus (raspberry-ringspot. ArMV. The latex test is considered as the most sensitive and the least time consuming method. Their coat protein consists of a single type of subunits with Mr of about 57 kDa. its own subgroup. Turkey. Kuszala. Bercks and Stellmach: ArMV. Lehoczky and Burgyan: Occurrence of TBRV in grapevines in Hungary. TBRV produces a reduction in growth and yield. wt of 2. Vuittenez A. Rüdel: Transmission of TBRV to grapevine by Longidorus attenuatus. Description Tomato black ring virus (TBRV) was first found in grapevines in Germany. Stellmach: Review paper on TBRV in grapevine. Rüdel and H Brückbauer. Ontario as the cause of severe damage to cv. or directly in grapevine leaf extracts using bentonite flocculation test. including TBRV : bentonite flocculation test.
Nucleotide sequence of tomato black ring virus RNA-1. Mayo and C. M. Journal of General Virology 65. 1984. respectively. and G. Greif C. Brückbauer. 1993.: SLRSV found in grapevine in Turkey. wt 2. 1984. The virus supports the replication of a satellite RNA 1118 nt in size. Savino et al. and 1.6 x 106 (RNA-2). Hemmer and C.. Bercks et al. Fritsch. Babini and Bertaccini: Electron microscope study SLRSV infections in plant tissues. 1575-1583. Annales de Phytopathologie 2.G. 3-5. Occurrence of tomato black ring virus on grapevine in southern Ontario. J. Fritsch. Canadian Plant Disease Survey 64. Turkiye. Researches on grapevine virus diseases and determination of their incidence in Ankara.: SLRSV recorded from grapevine in Italy.: Nucleotide sequence of SLRSV satellite RNA.W. Their coat protein consists of two types of subunits with Mr 43 x 103 and 27 x 103.: Nucleotide sequence of TBRV RNA-1 Abkas and Erdiller: TBRV recorded from grapevines in Turkey 3. The genome is a bipartite positive sense singlestranded RNA occurring as two separately encapsidated functional molecules with Mol. Virus particles are isometric. 1257-1271 Stobbs L. O. du virus des anneaux noirs de la Tomate (tomato black ring). and J. The virus is transmitted by Xiphinema diversicaudatum.: SLRSV and other nepoviruses isolated from grapevines in Germany Credi et al. Kreiah et al. Fritsch. Journal of Gerneal Virology 67. Erdiller. Vuittenez A. Historical review 1974 1977 1981 1982 1982 1987 1993 1994 2005 Murant: Description of SLRSV in the CMI/AAB Descriptions of Plant vrises series. about 30 nm in diameter have angular outline and sediment as three components (T.. Hemmer and C. Kreiah et al. 2. Symptoms on affected European grapes are of the fanleaf type. 1986.: Nucleotide sequence of SLRSV RNA-2. Genus SADWAVIRUS STRAWBERRY LATENT RINGSPOT VIRUS (SLRSV) 1. 1970. and B). The nucleotide sequence of tomato black ring virus RNA-2. Journal of General Virology 69. O. Van Schagen.6 x 106 (RNA-1) accounting for 38% of the particle weight. M. et du virus des taches annulaires du Framboisier (raspberry ringspot) chez des vignes du Palatinat. M. O. 279-327. Meyer M. 1517-1529.1986 1988 1993 Meyer et al. Journal of Turkish Phytopathology 22. 49 . Rüdel and H.: Nucleotide sequence of TBRV RNA-2 Greif et al. Le Gall et al.A. 55-61. Brückbauer: SLRSV can be distinguished from other nepoviruses on the basis of symptoms induced on Vitis idicator plants. Hemmer. Complete nucleotide sequence of a satellite RNA of tomato black ring virus. 1988. Description Strawberry latent ringspot virus (SLRSV) has been isolated from grapevine in the Palatinate (Germany) and in northern Italy. The virus is a definitive species in the genus Sadwavirus.. Assignement of SLRSV to the new genus Sadwavirus. Kuszala. References Akbas B. Meyer M. Détection et étude sérologique du virus latent des taches annulaires du Fraisier (strawberry latent ringspot). RNA-2 is 3824 nt in size and encodes a single ORF expressing a polypetide with Mr of about 99 kDa. It was also detected in imported vines in Turkey and Portugal.
1993. Murant A. Yoshikawa. Maniloff. 50 .A. Desselberger. T Jones. Sanfaçon. and A. (eds) 799-802 Elsevier/Academic Press. 102-104. Iwanami. 56-63. Bertaccini. References Babini A. Weinberg und Keller 24.I.. 1974. Credi R. 1977. Martelli. Brückbauer. Viral aggregates induced by a distinctive strain of strawberry latent ringspot virus from grapevine. Wetzel and N. 1981. Kreiah S. T. Strawberry latent ringspot virus. Gelli.A. Querfurth and M. 1982. J. Lehto. 1987. Die Wein-Wissenschaft 37. Journal of General Virology 75. Brückbauer H. The nucleotide sequence of a satellite RNA associated with strawberry latent ringspot virus. G. A. H. J. G. 2005. Wellink. Kreiah S..R. Strawberry latent ringspot virus in grapevine.I. Genus Sadwavirus. 2527-2532. T. J. Journal of General Virology. CMI/AAB Descriptions of Plant Viruses No.. Mayo. Eight Report of the International Committee on Taxonomy of Viruses (Fauquet. Strunk. 304-308. Phytopathologische Zeitschrift 104.V. M. H. Yilmaz. D'Onghia and M. and Ball..M.. U. A. Bertaccini and C. Betti. Cooper and G. Turkey. Karasev. 1982. 1994. Mögliche Beziehungen zwischen Virus und Symptomausprägung bei der Rebe. 133-180. L..R.. A distinctive isolate of Strawberry latent ringspot virus from grapevines in Italy. Sequence analysis and location of capsid protein within RNA 2 of strawberry latent ringspot virus. Bercks R. London. Savino V.3.M.. Cooper. Le Gall O.F.. 88-118.. Phytopathologia Mediterranea 20. 1163-1165.P.A. A. Strunk and J. C. In: Virus Taxonomy. L. 126. A.. K. Babini. Rüdel. G. FAO Plant Protection Bulletin 35. Untersuchungen überdie Viruskrankheited der rebe unter besonderer Berüchsichtigung “atypischer Formen” der Reisigkrankheit. 74..
PRMV. which in blueberry is transmitted by pollen. TRSV. rupestris plasma show field resistance to the northern US strain of ToRSV and to TRSV and PRMV.). sedimenting as three components. A number of rootstocks containing V. single-stranded RNA occurring as two separately encapsidated functional molecules with Mol . Partial sequence of the 3' 51 . about 30 nm in diameter with angular outline. Alternative weed hosts that have epidemiological significance are known for ToRSV. berlandieri or V. rivesi transmit ToRSV type strain (decline). PRMV. jaunissement des nervures. and/or ring spots) and distorted leaves. Long distance spread takes place primarily through infected propagating material. No vector is known for BLMoV. Concord it delays bud burst. Infected vines decline slowly over time. californicum transmits ToRSV yellow vein strain. depérissement de la vigne (Fr. The genome is a bipartite. Vitis vinifera.). In warmer climates (Maryland. whereas in V. except for BLMoV which may have been introduced from Europe. ToRSV and BLMoV are also seed transmitted in grapes. Their coat protein consists of a single type of subunits with Mr of about 54 x 103 Da. and poor fruit setting Agent: The above mentioned four distinct nepoviruses. labrusca is also resistant to TRSV.GRAPEVINE DEGENERATION AND DECLINE (AMERICAN NEPOVIRUSES) 1. mottled (oak leaf pattern. exhibiting stunted growth. Blueberry leaf mottle virus (BLMoV) infects latently European grapes. Peach rosette mosaic virus (PRMV) in V. BLMoV is a definitive nepovirus species assigned to subgroup C. Control: Use of virus-free propagating material and resistant rootstocks.e. wt of 2. americanum sensu lato and PRMV by X. elongatus. most V.35 x 106 (RNA-1) and 2. little grape (Eng. americanum sensu stricto.g. labrusca. straggly and shelled clusters. labrusca causes a severe disease characterized by delayed bud burst. and ToRSV separately or in combination. the infecting virus and the climatic conditions.). V. New York State and Ontario) own-rooted European grapes affected by Tomato ringspot virus (ToRSV) and Tobacco ringspot virus (TRSV) decline rapidly.15 x 106 (RNA-2). All these viruses. malformation and mottling of the leaves. deperimento della vite. Adernvergilbung (Germ. Longidorus diadecturus and L. and poor fruit setting. its main host. Bunches are small and straggly (Maryland's grapevine little berry) and leaves may show chrome yellow flecking along the veins (California's yellow vein). riparia. ELISA and molecular tools are useful for testing field-infected material. Varietal susceptibility: There are great variations in the susceptibility of Vitis species and cultivars. interspecific hybrids). grapevine decline. induces fanleaf-like symptoms on leaves and canes. labrusca cv. Description Blueberry leaf mottle virus (BLMoV) is named after the disease induced in highbush blueberry (Vaccinium corymbosum). BLMoV. BLUEBERRY LEAF MOTTLE VIRUS (BLMoV) 1. Virus particles are isometric. TRSV is transmitted by X. V. X. Transmission: These viruses are all transmitted by grafting and mechanical inoculation. Description Main synonyms: Yellow vein. are endemic in North America and thought to be native of the region. California) yield but not vigour is affected. poor fruit setting. ingiallimento nervale (Ital. are involved in the aetiology of North American grapevine degeneration and decline. positive-sense. interestingly. All roostocks and. This type of resistance is hypersensitivity. V.) Main symptoms: Symptomatological responses of grapevines vary according to the species (i. but not conclusive. distortion of canes. vinifera cultivars are reported as immune to the Californian strain of ToRSV Detection: All viruses are transmissible to herbaceous hosts mechanically and to woody indicators by grafting. Nematicidal control of vectors is possible. In cold climates (e. TRSV and PRMV. All other viruses are transmitted by longidorid nematodes: Xiphinema americanum sensu stricto and X.
Vectors are the Dorylaimoid nematodes Xiphinema americanum sensu lato and Longidorus diadecturus. PRMV is soil-borne. RNA1 is 8004 nt in size and contains a single open reading frame encoding a polypeptide with Mr of 240 kDa. The vector is unknown. mostly to vines adjacent to previously infected plants. Healthy grapevines become infected when planted in soils from diseased vineyards. The virus is seed-transmitted in grapevines and C. Physical and chemical properties of the particles and ribonucleic acid of blueberry leaf mottle virus. 52 .C. respectively.W. quinoa. Prolonged fallow is not an effective means of control because viruliferous nematodes remain alive for many years thriving on infected surviving roots and alternative weed hosts. Warkentin.4 x 106 (RNA-1) and 2. Grapevines (Vitis labrusca) are infected in New York State (USA) by a serologically distinct strain of the virus. The virus is transmitted through seeds in grapevines and C. quinoa (90% of infected seedlings). References Bacher J. Occasional. when a vineyard is planted susceptible cultivars may become infected by nematode vectors. sedimenting as three components.. wt of 2. PRMV is seed-borne in both naturally infected dandelion (4% of infected seeldlings) and in artificially infected C. 468-472. Sequence analysis of the 3' temini of RNA 1 and RNA 2 of blueberry leaf mottle virus. smaller and fewer than normal . : Partial nucleotide sequence of BLMoV RNA-1 and RNA-2 1981 1994 3. Stace-Smith. Taschenberg and D. Uyemoto J. Historical review 1977 Uyemoto et al: BLMoV isolated from New York 'Concord' vines showing fanleaf-like symptoms. Infected grapevines show shortened and crooked shoots. Plant Disease Reporter 61. possibily non specific transmission by L. Virus particles are isometric. and with extensive shelling of the berries.F. 145-156 Ramsdell D.. Phytopathology 71. Description Peach rosette mosaic virus (PRMV) is named after the disease induced in peach.2 x 106 (RNA-2) accounting for 44% and 37% of the particle weight.termini of both RNA molecules has been determined. Solanum carolinense (Carolina horse nettle) and Rumex crispus (curly dock). E.Use of resistant roostock hybrids and of certified planting material is recommended. PRMV can also be introduced in a site by infected planting material and be spread by vectors to adjacent vines.F. but not eradicate. 1977. but in highbush blueberry the virus is pollen-borne and suspected to be pollen-transmitted . Concord grapes are apparently virus-free but 9. where the disease occurs in more or less circular patches and spreads slowly. Isolation and identification of a strain of grapevine Bulgarian latent virus in Concord grapevines in New York State.Rasmdel and J. about 28 nm in diameter with angular outline. 1994. quinoa Ramsdell and Stace-Smith: Physico-chemical characterization of BLMoV and evidence that the New York grapevine virus is a strain of BLMoV Bacher et al. which may lead to their death.5% of seedlings from seeds taken from diseased vines proved to be infected. vector populations . 1981. mottled and variously deformed leaves and delayed bud burst. Roguing of infected trees and preplanting autumn fumigation with high rates of fumigant injected at two depths (15-20 cm and 75-90 cm) can effectively reduce. The genome is a bipartite positive sense single-stranded RNA occurring as two separately encapsidated functional molecules with Mol. Clusters are straggly. but identified as a strain of GBLV. D. labrusca cultivars (Concord. As the virus is endemic and seed-borne in some perennial weeds Taraxacum officinale (dandelion). Hancock. Their coat protein consists of a single type of subunits with Mr of about 57 x 103 Da. Crop losses up to 60% and death of susceptible V. The virus is a definitive nepovirus species assigned to subgroup C. especially) and a number of American-French hybrids have been recorded. which induces fanleaf-type symptoms and is distantly related to GBLV.K. elongatus has also been reported. 2.K. and R. PEACH ROSETTE MOSAIC VIRUS (PRMV) 1. Hummer. D. and has no economic importance. Virus Research 33. Vines are stunted and show a progressive decline. 949-953. Pollen grains of cv. one of its crop plant hosts.
Dias and Allen: Physico-chemical characterization of PRMV.A Ebsary. Canadian Journal of Botany 54. References Allen W. Canadian Journal of Plant Pathology 6. Ramsdell et al: High rates of fumigant injected at two depths (15-20 cm and 75-90 cm) during autumn reduce effectively but do not eradicate nematode vector populations in infested soils. Annales de Phytopathologie. 29-32 Allen W.A Ebsary. Rasmdell et al. Ramsdell and Myers: Description of PRMV-induced grapevine degeneration and association of X. J. Numero hors sèrie. Ramsdell: Review article on PRMV. Canadian Journal of Plant Pathology 10. Ramsdell and Gillet: List of grapervine cultivars and roostocks showing differential susceptibility to PRMV. The characterization of a virus responsible for peach rosette mosaic and grape decline in Michigan. 97-103. Transmission of peach rosette mosaic virus by Ontario populations of Longidorus diadecturus and Xiphinema americanum (Nematoda: Longidoridae). 1978 1979 1980 1982 1983 1984 1985 1988 1988 1995 1998 1999 3. S. and D. 16-18. tomato black ring and peach rosette mosaic viruses by an Ontario population of Longidorus elongatus. Ramsdell and Gillet: Description of PRMV in the AAB Descriptions of Plant Viruses.. Transmission of peach rosette mosaic virus to peach grape and cucumber by Longidorus diadecturus obtained from diseased orchards in Ontario. Purification and some characteristics of peach rosette mosaic virus (grape isolate). quinoa and has reproduced in part the field syndrome when inoculated mechanically to Concord grape seedlings. Historical review 1972a 1972b 1974 1976 Dias: Preliminary characterization of the grapevine isolate of PRMV. Canadian Journal of Plant Pathology 4. J. and B. Ramsdell and Myers: Field spread of PRMV is associated with the presence of infected weeds (T.: Xiphinema americanum is an efficient vector of PRMV. 1988. 1228-1239.: Longidorus diadecturus transmits PRMV to grapevines. Dias: Grapevine and peach strains of PRMV can be differentiated serologically. americanum with the disease. Carolinense. Strains of peach rosette mosaic virus differentiated by cross absorption and immunodiffusion tests. 1984. 1972a..: Nucleotide sequence of RNA-1 of the grapevine strain of PRMV. Transmission of raspberry ringspot. Dias H. Dias and Cation: Biological characterization of the grapevine strain of PRMV.S Everleigh. crispus) and transmission through grapevine seeds. Allen and Ebsary: Longidorus attenuatus transmits PRMV non specifically and with low efficiency.R.R.F.. 1976. Ramsdell et al: Use of ELISA for PRMV detection in grapevines. 1-5 Allen W.G Van Schagen and E. 1982.F. officinale.. Numero hors sèrie.2. Dias H. Allen et al.F. Annales de Phytopathologie. Allen et al. 105-106. The virus is seedborne in C. Lammers et al.R.G Van Schagen and B.: Investigation on the susceptibility to PRMV infection of American and European grapevines and hybrid rootstocks. R. 53 . Dias H. 1972b. Cation.
1995. Epidemiology of peach rosette mosaic virus in a Concord grape vineyard. Rose. 1998. Relative susceptibility of American. 1974. sedimenting as three components (T. Bird GW. 1174-1178.7 x 106 (RNA-1) and 1. Peach rosette mosaic virus decline.. Gillet and C.F Allison and D.: Survey of ToRSV and TRSV in Pennsylvanian vineyards. Ramsdell D. No.C.C. Lammers A. 2. Allen. 57-73.C.: A review of viruses infecting grapevines in New York vineyards. Ramsdell D. Phytopathology 64. 1983. Myers. wt of 2.M.C. Plant Disease 79.M. 1985. 1999. 1980. Characterization of the single protein and the two nucleic acids of peach rosette mosaic virus. Andrews. 1747-1754. Coat protein consists of a single type of subunits with Mr of about 57 x 103 Da.C. 1988. and B). and W.). The virus supports the replication of a circular satellite RNA 359 nt in size.C. Gillet. Myers.: Nucleotide sequence of TRSV satellite RNA.. Ramsdell D.W. R. Goheen eds. 364. Superimposed shallow and deep soil fumigation to control Xiphinema americanum and peach rosette mosaic virus reinfection a Concord vineyard.Dias H. Gillet and L. G. accounting for 44% and 28% of B and M particle weight. Ramsdell D.C. 4143. 1978. Virus particles are isometric.M. Description Tobacco ringspot virus (TRSV) is the type species of the genus Nepovirus and the prototype of subgroup A. . In: Compendium of Grape Diseases (R. French hybrids and European grape cultivars to infection by peach rosette mosaic virus. Phytopathologia Mediterrenea 24. Symptoms elicited by TRSV are the same as those of ToRSV in native cultivars. Bird. Susceptibility of American grapevine scion cultivars and French hybrid roostocks and scion cultivars to infection by peach rosette mosaic virus.C.C. Preventive control measures are the use of resistant roostock hybrids and of certified planting material. J.L. Ramsdell D.L.: TRSV agent of a new grapevine disease in New York State..E Morris. respectively. Cloning an sequencing of peach rosette mosaic virus RNA1.. but was recorded from grapevines only in New York State and Pennsylvania.C. Gillet. M. and J. TRSV is soil-borne and is transmitted by Xiphinema americanum sensu stricto. Phytopathology 68. RNA-2 has been sequenced only in part. 154-157. Uyemoto et al. Historical review 1970 1977 1985 1985 1986 1990 Gilmer et al. A comparison between enzyme-linked immunosorbent assay (ELISA) and Chenopodium quinoa for detection of peach rosette mosaic virus in 'Concord' grapevines.W. 625-627.3 x 106 (RNA-2).C Ramsdell.M. J. and J. The genome is a bipartite positive sense single-stranded RNA occurring as two separately encapsidated functional molecules with Mol. TRSV has a relatively wide natural host range. St. TOBACCO RINGSPOT VIRUS (TRSV) 1.M.M. There is no evidence of seed trasmission in the grapevine.C. Buzayan et al.W. 54 . Peach rosette mosaic virus. is endemic in Central and Eastern North America. and R. Gillet and G. Plant Disease 67. American Phytopathological Society Press. 51-52. and R. Peach rosette mosaic virus.R. Virus Research 65. G. Plant Disease Reporter 63. R. about 30 nm in diameter with angular outline. American Vitis species reported to be resistant to ToRSV and TRSV. Paul. Ramsdell D. AAB Descriptions of Plant Viruses. symptomatology and nematodes associated with grapevine 'degeneration' in Michigan.. Pearson and A. but in European grapes responses are similar to those elicited by GFLV. 74-78. Canadian Journal of Botany 58. RNA-1 is 7514 nt in size and contains a single open reading frame encoding a polypeptide with Mr of 225 kDa. Ramsdell D.H. 447-450 Ramsdell D. Stace-Smith: Description of TRSV in the AAB Descriptions of Plant Viruses series. Foster and Morris-Krsinich: In vitro translation of TRSV RNA-1 and TRSV RNA-2 yields major polypeptides with Mr of 225K and 116K. respectively. 1979. Powell et al.
Chemical cleavage of the 5'-linked protein of tobacco ringspot virus genomic RNAs and characterization of the protein-RNA linkage. Forer. ToRSV-induced decline affects European cultivars. Two serological ToRSV variants are known to infect grapevines. labrusca. A new grapevine disease induced by tobacco ringspot virus.B. AAB Descriptions of Plant Viruses. Silva and S. Zallua et al. and B. Incidence of tomato rigspot virus and tobacco ringspot virus in grapevines in Pennsylvania. B. V. Virology 144. Gould. Clusters are straggly. Uyemoto J. Stace-Smith R. Tobacco ringspot virus.L. Preventive control measures are the use of resistant roostock hybrids and of certified planting material. 309. Powell C. Virology 219. Symptomatological responses vary according to the species (V. 1985. 131-136. 1977. ToRSV is soil-borne.A. American Journal of Enology and Viticulture 28. Zalloua P. Nucleotide sequence of satellite tobacco ringspot virus RNA and its relationship to multimeric forms.K. Bruening. 1993. Buzayan and G. 335-349. TOMATO RINGSPOT VIRUS (ToRSV) 1. Phytopathology 60. Plant Disease 74. Bruening. In California ToRSV affects the yield rather than the vine's growth.. and B)..1993 1996 Buckley et al. 1-8. 1970.M. 1985.8 x 106 (RNA-1) and 2. Nucleotide sequence and in vitro expression of the capsid protein gene of tobacco ringspot virus.M.K.L. vinifera. Vectors are the Dorylaimoid nematodes Xiphinema americanum sensu stricto and X. Longenecker and L. Virus Research 30. P.. Gerlach G. 1990. mottled and distorted leaves and short internodes. contrary to the decline strain which is seed-transmitted. 2. Gooding and Hewitt: A mechanically transmissible virus found to be associated with yellow vein.A. Buzayan J. and with extensive shelling of the berries. Virology 151. sedimenting as three components (T. RNA-1 is 8214 nt and RNA-2 is 7273 nt in size. 516-519... 1996. J. about 30 nm in diameter with angular outline. W. Cummins and G. Abawi. and the climatic conditions. smaller and fewer than normal. ToRSV has a relatively wide natural host range and is endemic in North America. Foster R. No. The virus has been occasionaly recorded from grapevines outside of North America. The yellow vein strain of the virus is pollen-borne but is not transmitted through seeds. wt of 2. respectively. Gilmer R. J. interspecific hybrids). Morris-Krsinich. Singh. 619-627. Infected vines have small. rivesi in north American States and Canada and X.: Nucleotide sequence of TRSV RNA-1 3.4 x 106 (RNA-2) accounting for 44% and 41 % of the particle weight. "yellow vein" being the characterizing syndrome of its infections. Description Tomato ringspot virus (ToRSV) is a definitive species in the genus Nepovirus and the prototype of subgroup C. Keese and A. the infecting virus strain. 702-704. Historical review 1954 1956 1962 Hewitt: Report of an "unfruitful vine" condition in California to which a yellow speckling of the leaves is associated.S. The genome is a bipartite positive sense single-stranded RNA occurring as two separately encapsidated functional molecules with Mol. : Partial nucleotide sequence of TRSV RNA-2. Both RNAs contain a single open reading frame encoding polypeptides with Mr of 244 kDa (RNA-1) and 207 kDa (RNA-2). Vines grow vigorously but bear little or no fruit.J.M. Synthesis and processing of the translation products of tobacco ringspot virus in rabbit reticulocyte lysates.M. References Buckley B. Vines are stunted and show a progressive rapid decline.A.S. 186-199. Disease named yellow vein..R. 1986. more severely in colder than in warmer climates.M. 55 . Coat protein consists of a single type of subunits with Mr of about 58 x 103 Da. where it occurs iin the region of the Great Lakes and in the Pacific seaboard from California to British Columbia. J. Hewitt: Successful graft transmission of unfruitful vine condition. Virus particles are isometric. Kelts. J. californicum in California The virus can be introduced in a site by infected planting material and be spread by vectors to adjacent vines. Virus and virus-like diseases affecting grapevines in New York vineyards. especially if self-rooted.L. which often leads to death. Uyemoto and L.R.
Uyemoto: Seed transmission of the decline strain of ToRSV.1963 1966 1968 1972 1972 1975 1977 1977 1977 1978 1980 1982 1982 1984 1985 1985 1986 1987 1987 1988 1989 1989 1990 1991 1992 1993 1995 2001 2004 Gooding: Yellow vein virus identified as a strain of ToRSV. americanum (now X. Stace-Smith: Description of ToRSV in the CMI/AAB Descriptions of Plant Viruses series. Allen et al. californicum). Gonsalves: ToRSV is irregularly distributed in infected vines but can be detected by ELISA. American Vitis species reported to be resistant to ToRSV and TRSV. Yang et al.: Complete nucleotide sequence of ToRSV RNA-1.: A review of viruses infecting grapevines in New York State vineyards.: Complete nucleotide sequence of ToRSV RNA-2.: List of grapevine roostocks and cultivars showing differential susceptibility to ToRSV in Canada. Uyemoto and Gilmer: Spread of ToRSV through the soil of New York State vineyards recorded. Rott et al. Allen and Dias: Physico-chemical characterization of ToRSV. 56 . Podlekis and Corbett: ToRSV is the agent of little grape disease in Maryland. Herrera and Madariaga: ToRSV recorded from grapevine in Chile.: Survey of ToRSV and TRSV in Pennsylvanian vineyards. Uyemoto et al. Martelli: Review of nematode-borne viruses of grapevines and their epidemiology. Teliz et al. Corbett and Podleckis: Ultrastructural study of ToRSV-infected grapevine tissues. Rott et al. Baumgartnerova and Subikova: ToRSV recorded form grapevine in Slovakia. Dias: Record of ToRSV in the Niagara peninsula. Martelli and Taylor: Review of nematode-borne viruses and their vectors. Piazzolla et al. Li et al: ToRSV identified in China in grapevine seedlings grown from seeds imported from France.: Transmission of the yellow vein strain of ToRSV by X.: ToRSV found in grapevines in Taiwan. Cory and Hewitt: The yellow vein strain of ToRSV is not transmitted through seeds. Rowhani et al: Description of sampling strategy for detection of ToRSV.: Confirmation that the grape yellow vein and the the grape decline strains of ToRSV are serological variants of the same virus. Bays and Tolin: ToRSV found in grapevines in Virginia Powell et al. Allen et al. Stace-Smith and Ramsdell: Review of nepoviruses of the Americas. Gilmer and Uyemoto: ToRSV agent of a decline of Baco noir in New York State.: Xiphinema rivesi identified as the main vector of ToRSV in Ontario vineyards. Bitterlin and Gonslaves: ToRSV retained and transmitted by viruliferous Xiphinema rivesi stored for two years at 1-3°C.
Martelli G. Plant Disease Reporter 61. M. Gonsalves D. 31-34. Téliz D. Journal of General Virology 72. Cory L. Bulletin of the California Department of Agriculture 43. Rowhani A. 1987.F. Phytopathology 72..R. 1977. and grape yellow vein viruses by Xiphinema americanum.A. Nematologia Mediterranea 6. Dias and J. Corbett M. Plant Disease Reporter 59. Canadian Journal of Botany 55. J. Susceptibility of grape cultivars and rootstocks to an Ontario isolate of tomato ringspot virus.B. 1954. 1992.A. Plant Disease Reporter 56. 1505-1514. 663 Martelli G. Allen W. CMI/AAB Descriptions of Plant Viruses. Madariaga. and W. Ebsary. Savino.E.. Distribution of viruses and their nematode vectors. Works of the Institute of Experimental Phytopathology and Entomology .R. 861-863. Detection and identification of plant viruses for quarantine in China. Grogan and B. Bitterlin M. Li. 710. Nucleotide sequence of tomato ringspot virus RNA 1.E. 44-50. 1169. Tomato ringspot virus in "Baco noir" grapevines in New York. and W. Phytopathology 58. Zhang and Y. 1990. Identification of tomato rinsgspot virus in leafroll diseased grapevines. 1989. Detection of tomato ringspot virus in grapevines: irregular distribution of virus. Corbett. and E.B. Beijing 2004.F. 702-704. Plant Disease 74. Some virus and virus-like diseases of grapevines.A. Ramsdell.F. 275-277. References Allen W.B. 1985. 131-166.K. 1966.. and S. 1972. and H. 1985. 35-44. Musci. Nucleotide sequence of tomato ringspot virus RNA-2. Gooding G. Niagara Falls 1980. G. Membrane-associated spherical particles in extracts and tissues of virus-infected grapevines. Properties of the single protein and two nucleic acids of tomato ringspot virus. Uyemoto J. Rochon. No. identification.. M. V. Advances in Disease Vector Research 6.M. Y. 133-135. 24-28. Tolin. Proceedings 7th Meeting of IGVG..W. Phytopathologia Mediterranea 24. and D. Purification and serology of a virus associated with the grape yellow vein disease. Van Schagen and B. Hewitt.F. Gonsalves. 1982. Bays D. 2004. Hewitt. Incidence of tomato ringspot virus in grape in Virginia. Tomato ringspot virus. 1984.F. 157-164..G.. 1991. Transmission of tomato ringspot. Lownsberry. 475-480. Stace-Smith R. Phytopathology 56. Piazzolla P.C. Uyemoto. Spatial distribution of Xiphinema rivesi and persistence of tomato ringspot virus and its vector in soil. Phytopathologia Mediterranea 24. J. A. 465-473. 393-400 Hewitt W. J. Grape yellow vein: symptomatology. Canada.V.C. 1-27. Allen W. peach yellow bud mosaic. Li M. Walker and S. Herrera M. Nematode-borne viruses of grapevine. Gooding G.A. N.K.. 408-411. Proceedings 15th International Plant Protection Congress. and B. 2001. 1982. L. L. Tomato ringspot virus associated with little grape disease of Vidal 256 grapevines.R. 98-101. Nepoviruses of the Americas.W. 1989.G and V. Chen.. 1988. Gilmer R. 1993. 658-663.J.3. Stobbs.F. 290 Stace-Smith R. Xiang .V.. Baumgartnerova H.M. 47-64. H. and M. Incidence and geographic distribution of tomato ringspot virus in De Chaunac vineyards in the Niagara peninsula.A. Agricultura Tecnica 61.V. Association of Xiphinema species with soil types ang grapevines infected with tomato ringspot virus in Ontario. A comparison of Grapevine yellow vein virus and a Grapevine isolate of Tomato ringspot virus. Plant Disease 72. 1962. Podleckis. Ivanka pri Dunaji 4. Sampling strategies for the detection of grapevine fanleaf virus and the grapevine strain of tomato ringspot virus. Taylor. Incidence of tomato rigspot virus and tobacco rigspot virus in grapevines in Pennsylvania. Rochon. Castellano and D.V. R.G. Longenecker and L. and C..E.S Whei. Presence and incidence of grapevine viruses in central Chile. Subikova. Journal of General Virology 76. Vitis 31. Rott M. 95-106. Podleckis E. Rott M.B. their epidemiology and control. Some grapevine viruses in pollen and seed. 196-203. 1963.G. Gilchrist. and the association of a mechanically transmissible virus with the disease.L. 13161320 Dias H. Rokni. Van Schagen. Forer. Powell C..F.. 1987. and V. Canadian Journal of Plant Pathology 4. Plant Disease 71. 151-189. 1968. Dias.P. 1995. Phytopathology 53. Tremaine and D'A. A severe outbreak of virus-induced grapevine decline in Cascade grapevines in New York. American Journal of Enology and Viticulture 13. 1977. 57 . Lee and D'A. 1980.. 1978. and J. Current Topics in Vector Research 5. 1028-1037. 1975. M..P..H.K. Phytopathology 79.K.
R. American Journal of Enology and Viticulture 28.S. and R. Spread of tomato ringspot virus in "Baco Noir" grapevines in New York. J.Uyemoto J. 1986. Abawi.K. T. Virus and virus-like diseases affecting grapevines in New York vineyards. 1972. Cummins and G... Uyemoto J. Yang I.K. Plant Disease Reporter 56. 1977. Sap-transmissiible viruses associated with grapevine yellow mottle disease in Taiwan.L.M. 1062-1064. Deng and M. 131-136. 504-510.J. Gilmer.C. 58 . Journal of Agricultural Research China 35. Chen.
the whole leaf surface becomes deep purple. GLRaV-3. especially the phloem. These symptoms progress towards the top of the canes as the season advances. and lowering of sugar content. Description The first descriptions of grapevine leafroll date back to the mid 19th century. GLRaV-4. Sieve elements are obliterated and crusheds. and with many cultivars. Rollkrankheit.e. Hence. Careful observation of field symptoms in infected vines reveals that there are several types of leafroll. but the leaves become chlorotic to yellowish. Leafroll is no less important than fanleaf in economic importance.) Main symptoms: In red-berried cultivars of Vitis vinifera reddish spots develop in the lower leaves in late spring or summer. respectively. GLRaV-2 CP has a Mr of 24 kDa. Other such viruses may exist. A number of other physiological parameters are affected. nine different viruses with filamentous particles. GLRaV-5. potassium depletion in the leaf blade and accumulation in the petioles. graft take and plant vigour. the risk of disseminating the disease is great if untested rootstocks are used. Sizes deduced from the nucleotide sequence of the CP cistron are 22 kDa for GLRaV-2 . Its occurrence in viticultural areas is worldwide. These negative effects are reverted if the disease is eliminated by sanitation treatments. infection of rootstocks is symptomless. the symptoms are similar. 61 . In white-berried cultivars of V. Also the composition and aromatic profile of the musts are modified.). Also plant anatomy is affected. and GLRsV-9 in the newly established genus Ampelovirus. whereas the Mr of all other viruses ranges between 35 and 44 kDa. the same as the size of coat protein (CP) subunits.). usually leaving a narrow green band along the primary and secondary veins. GLRaV-1. Blattrollkrankheit (Germ. accartocciamento. have been found in leafroll-infected vines. called grapevine leafroll-associated viruses (GLRaV). GLRaV-8. These spots enlarge with time and coalesce so that.GRAPEVINE LEAFROLL 1. All GLRaVs belong in the family Closteroviridae. changes in the pattern of peroxidase and polyphenoloxidase isoenzymes. exhibiting open structure and distinct cross banding with a pitch of about 3. i. vinifera. depending on the climate and geographic location. accartocciamento fogliare (Ital. most of the leaf surface becomes reddish. and low in sugar. Leafroll decreases grapevine yield (by 15-20% in average) and affects negatively rooting ability.). The leaf blade becomes thick. in autumn. as estimated by polyacrylamide gel electrophoresis. Agents: To date.). Starch accumulates in degenerated chloroplasts causing increased thickness and brittleness of the leaf blades. GLRaV-6. enrollado (Sp. There are reports of early reddening of grapevine leaves regarded as physiological disorders and referred to as “Rugeau” or “Rossore” in the French and Italian literature. Virus particles are very flexuous filaments about 12 nm wide. thus impairing carbohydrate translocation from foliar parenchymas. differentiation of GLRaVs at the species level was by Roman numerals. except for a variable decrease in vigour. Main synonyms: White Emperor disease (Eng.). Until 1995. In most cases. now is by Arabic numerals: Grapevine leafroll-associated virus 1 (GLRaV-1) Grapevine leafroll-associated virus 2 (GLRaV-2) Grapevine leafroll-associated virus 3 (GLRaV-3) Grapevine leafroll-associated virus 4 (GLRaV-4) Grapevine leafroll-associated virus 5 (GLRaV-5) Grapevine leafroll-associated virus 6 (GLRaV-6) Grapevine leafroll-associated virus 7 (GLRaV-7) Grapevine leafroll-associated virus 8 (GLRaV-8) Grapevine leafroll-associated virus 9 (GLRaV-9) A potyvirus isolated in Israel from leafroll-infected vines is now regarded as an occasional contaminant. brittle and rolls downwards. they are inferior in quantity and quality. Particle length varies from 1400 to 2200 nm according to individual viruses. enroulement (Fr. and is probably the most widespread virus disease of grapevine. Enrolamento de la folha (Port. The fruits often mature late and irregularly. thus suggesting that there can be several causal agents. instead of reddish. enrollamiento de la hoja. reduction of protein content. differing somewhat in aspect and in severity. whereas GLRaV-7 is presently classified as unassigned species to the family. which are differentiated from one another by a progressive number.5 nm. In the most severe cases. GLRaV-2 in the genus Closterovirus.
cloned cDNA probes and riboprobes to GLRaV-1 and GLRaV-3 have been produced from denatured double-stranded RNA (dsRNA) and a number of virus-specific. This has been ascertained experimentally for GLRaV-1.Transmission is semipersistent and does not appear to be vector-specific. singlestranded. leafroll can be detected by its symptoms in the field on red-fruited varieties. ultrastructurally. grafted vines) which is largely responsible for its dissemination over medium and long distances. As to nucleic acid-based assays. roostocks. The genome of GLRaV-2 is 15.35 kDa for both GLRaV-3 and GLRaV-1. The genome of GLRaV-1 is 17. or the hybrid LN 33 is still the most popular method for identifying the disease. Ps. Polyclonal antisera and/or monoclonal antibodies have been raised to each single GLRaV . Indexing on red-fruited cultivars such as 'Cabernet sauvignon'. contains 13 ORFs. GLRaV-5) or from vesiculation of the endoplasmic reticulum (GLRaV-2 and GLRaV-7) Transmission: Leafroll is graft-transmissible and persists in propagative material (budwood. Ps. Symptom expression depends on the variety. only vectors of GLRaV-1. GLRaV-3. The presence of high molecular weight double-stranded RNAs (dsRNA) in phloem tissue extracts can be used as infection marker. broadspectrum. and some of them are used as indicators. and Neopulvinaria innumerabilis. Mealybug vectors of GLRaV-3 are Planococcus ficus. positive sense RNA molecule. or by molecular techniques. American rootstocks are usually symptomless carriers of GLRaVs. The genome of GLRaV-4 and GLRaV-9 are incompletely sequenced but show a structural organization compatible with that of the genus. GLRaV-3. None of the GLRaVs is known to be seedborne. Pl. climate. Ps. Varietal susceptibility and sensitivity: No immune variety or rootstock is known. American Vitis species and rootstocks are the best antigen sources for serological assays. Composite samples should be used to minimize false negative responses that may originate from the unven distribution of GLRaVs in chronically infected vines. GLRaVs differ in various vays (molecularly. biologically. 'Merlot'. GLRaV-5 and GLRaV-9 are phylogenetically close to one another but are serologically unrelated. the type member of the genus. in agreement with the quasispecies nature of viruses. comstocki. has a number of minor biological variants which can be differentiated by the reaction of inoculated Nicotiana species. These reagents are routinely used for ISEM. the only member of the group to be mechanically transmissible. maritimus.528 nt in size. vinifera. viburni and Ps. vinifera varieties show symptoms most clearly because of the reddening of the leaves. Membranous vesicles can derive eitrher from peripheral vesiculation of mitochondria followed by disruption of the organelles (GLRaV-1. GLRaV-5 and GLRaV-9 have been identified. vinfera and cortical shavings from mature dormant canes of V. 29. Ps. Red-berried V. Foliar tissues are not recommended for serological GLRaVs detection in American Vitis species and roostocks. number an types of infecting viruses. affinis. Parthenolecanium corni. Disappearance of dsRNAs from vines submitted to 62 . Pseudococcus longispinus. but it does not discriminate between GLRaVs and was reported to be less sensitive than ELISA. with none of which they are serologically related. longispinus. GLRaV-5 and GLRaV-9 are both transmitted by Ps. So far. GLRaVs show molecular variations which give rise to a population of strains. and epidemiologically) from most of the known closteroviruses. All GLRaVs can be identified by serological and nucleic acid-based techniques. citri. Regardless of whether they belong to the genus Closterovirus or Amplelovirus.647 nt in size and contains 10 major ORFs. Its soft scale insects vectors are Pulvinaria vitis and Neopulvinaria innumerabilis. and has structural organization differing from that of other sequenced closteroviruses. Other GLRaVs may exist in nature as suggested by reports. soil condition and probably. GLRaV-3.5 kDa for GLRaV-4 . 'Cabernet Franc' 'Pinot noir'. GLRaV-2 and GLRaV-3. which is the type species of the novel genus Ampelovirus has a genome 17. contains nine open reading frames (ORF) and has structural organization identical to that of Beet yellow virus (BYV). calceolariae. and degenerate primers have been designed and successfully used for PCR detection of virtually all GLRaVs. GLRaV-1 is transmitted in nature by the pseudococcid mealybugs Heliococcus bohemicus and Phenacoccus aceris and the soft scale insects Pulvinaria vitis.919 nt in size. GLRaVs were also thought to be serologically distinct from one another until a distant serological relationship was found between GLRaV-1 and GLRaV-3 using monoclonal antibodies raised to GLRaV-1. The genome is a monopartite. classical double antibody sandwich ELISA (Chromo-ELISA) or Lumino-ELISA. Natural field spread of leafroll disease has been reported from many countries in Europe and elsewhere. Detection: In many cases. Cytopathology: A characterizing feature of all GLRaV infections is the presence of intracellular inclusions in phloem tissues made up of aggregates of virus particles intermingled with single or clustered mebranous vesicles containing finely stranded material thought to be viral RNA. Leaf tissues or petioles from mature symptomatic leaves of V. Spread at a site is mediated by mealybug and soft scale insect vectors. and some are commercially avaliable. GLRaV-2.
As no apparent spread of the disease was observed.sanitation treatments is regarded as evidence for successful virus elimination. Hypothesis of the viral origin of leafroll.: White Emperor and leafroll are identical diseases. Luhn and Goheen: Leafroll found in the original grapevine stocks imported from Europe into California in 1890.: Leafroll in Czechoslovakia. Lehoczky et al. a grapevine disorder similar to leafroll. Ravaz and Verge: Occurrence in France of “rougeau”. Goheen et al. Belli et al. 2. unless they are hybridized with virus-specific probes. Tanne and Nitzany: Leafroll in Israel Tanne et al. roostocks are suggested as the major souces of leafroll dissemination.: Transmission of a virus to herbaceous plants from a leafroll-infected vine in Israel. Introduction of transgenic resistance to GLRaV-2 and GLRaV-3 is being attempted by engineering different viral genes into rootstocks and European grape cultivars. Fraser: Leafroll in Australia.: Leafroll in Hungary. 1971 1973 1974 1975 63 . Hewitt: Leafroll in California Goheen et al. These particles are probably closteroviruses associated with leafroll. Scheu: Leafroll is widespread in German vineyards. Hoefert and Gifford: Study of the effects of leafroll infection on vine anatomy. Later studies showed that the virus is an occasional contaminant Lider et al. No sources of resistance are known in V. However. vinifera and there is no published information on how to protect healthy stocks from vector-mediated reinfection in the field.: Studies on the effects of leafroll on yield of grapevines in California. Vuittenez: Leafroll in France. a grapevine disorder similar to leafroll Scheu: Demonstration of graft transmission of leafroll from diseased to healthy Vitis vinifera. Mendgen: Presence of filamentous particles in grapevines with symptoms of flavescence dorée in West Germany.: Leafroll in Italy. Control: Production and use of clonally selected and sanitized propagation material is very effective and the only preventive method for leafroll control available. Chamberlain: Leafroll in New Zealand. Blattny et al. Historical review 1906 1924 1935 1936 1946 1954 1958 1958 1958 1960: 1965 1967 1967 1967 1968 1969 1970 1970 Sannino: Occurrence in Italy of “rossore”. dsRNAs cannot be utilized for virus identification. Bovey: Leafroll in Switzerland. Dimitrijevic: Leafroll in Yugoslavia. The incidence if the disease was less than 20% as compared with 80 to 100% in commercial vineyards.: Leafroll virus can be inactivated in vivo by heat therapy. Harmon and Snyder: The "White Emperor" disease is graft-transmissible and is regarded a virus disease.
Von der Brelie and Nienhaus: Light and electron microscope study of cytopathological changes induced by leafroll in grapevines.: Ultrastructural study of leafroll-infected grapevine tissues. Faoro et al. Kliever and Lider: Study of biochemical changes found in grapevine infected with leafroll in California.: Closterovirus-like particles with an estimated length of 1000 nm found in thin sections of phloem tissue and in leaf dip preparations of leafroll-diseased grapevines in Japan. Gugerli et al. Hu and Gonsalves: Monoclonal antibodies produced against GLRaV-3.: Closterovirus-like particles and specific dsRNA found in leafroll-diseased grapevines in New Zealand. Abracheva: Leafroll in Bulgaria. Zee et al.: Aggregates of closterovirus-like particles observed in thin sections of phloem from leafroll-diseased grapevines. Castellano et al.: ELISA testing reveals that GLRaV-3 has an uneven distribution in grapevine tissues. are also present in leafroll-affected grapevines from Italy. Suggestion that a closterovirus may be the agent of the disease. but not in healthy controls. Sasahara et al. Presence of virus-like particles in thin sections of leafroll-diseased vines.: Review on the detrimental effects of viral infection on grapevine physiology. Tanaka: Leafroll in Japan. Mossop et al.1975 1976 1976 1977 1979 Martelli and Piro: Evidence from a herbarium that leafroll occurred in Sicily in the second half of the 19th century. Confirmation that GLRaV-3 and the New York closterovirus isolate cross react serologically. 1981 1981 1982 1982 1983 1984 1984 1984 1985 1986 1986 1987 1987 1988 1988 1989 64 . A third closterovirus type called GLRaV-3. Rosciglione and Gugerli: GLRaV-3 is transmitted by the mealybug Planococcus ficus. Namba et al. Antiserum against a New York isolate also reacted with GLRaV-3 from Europe. Rosciglione and Gugerli: GLRaV-1 and GLRaV-2 with particles of 2200 nm and 1800 nm respectively. Martelli et al.: Closterovirus-like particles purified from leafroll-diseased grapevines in France. Barlass at al.: Extraction and first purification of closterovirus-like particles with maximum particle length of 2200 nm (type I) and 1800 nm (type II) from leafroll-diseased grapevine leaves in Switzerland. A large dsRNA molecule is consistently isolated from leafroll-diseased grapes. Absence of such particles in healthy grapevines. Teliz et al.: First record of successful elimination of leafroll in grapevine by using meristem tip culture in Japan. Corbett et al: Electron microscope observations by negative staining of leaf extracts from leafroll-diseased grapevines in South Africa showed the presence of closterovirus. Production of rabbit and hen antibodies for ELISA and ISEM to GLRaV-1 and GLRaV-3. Hofmann: Symptoms of leafroll in affected clones of Pinot noir and performance in West Germany. Production of polyclonal antisera for use in ELISA. but not in similar praparations from healthy plants. Zimmermann et al.: Elimination of leafroll by in vitro meristm tip culture and apex fragmentation.: Studies on the cytopathology of leafroll-diseased grapevines. previously found in grapevines in Switzerland.like particles. found in grapevines affected by leafroll. Purification and serology of associated closterovirus-like particles.
Auger et al.: Leafroll and associated closteroviruses in Chile Kuhn: Leafroll in Brasil Li et al. Savino et al. but not GLRaV-1. rupestris plasma. symptoms are obtained within 20-70 days. GLRaV-1 and GLRaV-2 were not transmitted. Téliz et al.: Detection of leafroll-associated closterovirus in recently infected grapevines in New York.: Characterization of leafroll-associated closterovirus-like particles from grapevine using also monoclonal antibodies. In Mexico leafroll. stem pitting and corky bark spread rapidly. Gugerli et al. With leafroll. For reliable testing. ELISA is to be applied to cortical scrapings rather than leaf tissues. Transmission of GLRaV-3 by the mealybug Planococcus ficus. whereas meristem tip culture yields up to 100 % sanitation Walter and Zimmermann: Further characterization of closteroviruses associated with leafroll in France.: Evidence of the irregular distribution of GLRaV-3 in American rootstocks. but they are easily detected in inoculated LN33 vines and in V.: Leafroll and associated closteroviruses in China Engelbrecht and Kasdorf: Transmission of GLRaV-3 by Planococcus ficus from grapevine to grapevine in South Africa. 1990 1991 1991 1991 1991 1991 1991 1991 65 . The virus was detected in root tissues. Gugerli: Review of grapevine closteroviruses. Credi and Santucci: GLRaV-1 and GLRaV-3 cannot be detected by direct ELISA in leaves of graft-inoculated American rootstock. ficus fed on infected vines. later in the leaves. Agran et al.: Transmission of GLRaV-3 from grapevine to grapevine by the mealybug Pseudococcus longispinus in Israel. GLRaV-2. b Hu et al. GLRaV-1. Some vines indexing positive for leafroll do not react positively with any ofthe antisera. Identification of GLRaV-5. was detected in P.: Production of monoclonal antibodies to GLRaV-1 and GLRaV-3. 1989 1989 1989 1990 1990 1990a. especially those containing V. Zimmermann et al. Heat therapy requires very long treatments and is only 20-30 % successful.: Immunocytological detection and localization of GLRaV-1 and GLRaV-3 by immunogold labelling in grapevine thin sections. Faoro et al. Pseudococcus longispinus is present on weeds around diseased vineyards.1989 1989 Tanne et al. Gugerli et al.: Comparison of heat therapy and meristem tip culture for eliminating GLRaV-3 from Italian grape varieties. -2 and -3 are common whereas GLRaV-5 is rarely detected. Good results in summer with extracts of basal leaves and in autumn or winter with wood shavings macerated in buffer.: Leafroll in Tunisia Azeri: Leafroll in Turkey Borgo: Serological detection of GLRaV -1 and GLRaV-3 by ELISA in extracts of leaves or wood shavings.: Further characterization of GLRaV-1 and GLRaV-3 by monoclonal antibodies. Boscia et al.: Production and characterization of monoclonal antibodies specific to GLRaV-3. vinifera varieties used as inoculum source. There is evidence that other GLRaVs are involved in leafroll etiology. indicating the presence of other leafroll-associated viruses.: Use of green grafting for detecting virus-like diseases of grapevine. Identification of GLRaV-4 1990 1990 1990 1990 Walter et al.
Leafroll and associated closteroviruses in Yemen. whereas the other assays are more suitable for analyzing samples that gave inconclusive results with ELISA. Namba et al.: Leafroll in Taiwan. Krake: Characterization of leafroll disease based on symptoms shown by field-infected vines and graft-transmission tests. Gugerli and Ramel: Analysis by monolconal antibodies of a Swiss source of cv. Observation of peripherically vesiculated mitochondria. Segura et al. Pop et al. Habili et al. Minafra and Hadidi: Detection of GLRaV-3 in viruliferous mealybugs by PCR. ELISA is recommended for large screening. Flak and Gangl: Leafroll and associated closteroviruses in Austria.: Analysis for the presence of double-stranded RNAs can be used for assessing virus elimination following sanitation treatments.: Purification and physico-chemical characterization of grapevine corky bark associated virus.1% in 1992 Ioannou: Leafroll and natural spread of associated closteroviruses in Cyprus. 1994 1994 1994 1994 1994 1995 1994 1995 1995 1995 1995 Merkuri et al. Boscia et al.: Leafroll and associated closteroviruses in Spain 1991 1991 1991 1991 1991 1991 1992 1993 1993 1993 1993 1993 1993 1994a. Golino et al.: Mechanical transmission of GLRaV-2 and ultrastructural study of infected tissues of Nicotiana benthamiana. Martelli et al.: Leafroll and associated closteroviruses in Romania. Former GLRaV 2 is re-named GLRaV-6. denoted GLRaV 2a and GLRaV 2b.: Leafroll and associated closteroviruses in Albania. Belli et al. 66 . Bondarchuk et al.b Saldarelli et al.1991 Hu et al. later identified as GLRaV-2. Jordan: In a New Zealand commercial vineyard GLRaV-3 incidence increased from 9. Castellano et al.: Leafroll and associated closteroviruses in Ukraine.: Production of radioactive and non-radioactive molecular probes to GLRaV-3 from denatured dsRNA template and their use for virus identification.: Transmission of GLRaV-3 by the soft scale insect Pulvinaria vitis. ISEM and dsRNA analysis.: Leafroll and associated closteroviruses in Moldova. Tzeng et al. Faoro and Carzaniga: Ustrastructural study of GLRaV-1 and GLRV-3 infections. Katis et al: Leafroll and associated closteroviruses in Greece.: Comparison of different assay methods for detecting GLRaVs : ELISA.1% in 1988 to 93. Kassemeyer: Detection of GLRaVs in Germany. Chasselas shows the prsence of two different GLRaV-2. Milkus et al.: Revision of the nomenclature of GLRaVs and use of Arabic numerals in the species names.: Production of antisera to GLRaVs using electrophoretically separated coat protein subunits as antigens. Boehm and Martins: Leafroll in Portugal.: Transmission of GLRaV-3 by Pseudococcus affinis in California. Gozsczynski et al.
Alkowni et al.: Transmission of GLRaV-1 by the soft scale insects Parthenolecanium corni and Neopuvinaria innumerabilis. MacKenzie et al.: Identification of GLRaV-7 and production of a polyclonal antiserum. Ling et al. No vector was identified.: Leafroll and associated closteroviruses in Palestine. American.: Extensive sequencing of GLRaV-3 genome.: Development of a spot-PCR technique for GLRaVs identification. Wilcox et al.: GLRaV-2 and GLRaV-3 genes engineered in grapevine rootstocks to induce resistance.: Serological characterization of GLRaV-6 and production of monoclonal antibodies. Gugerli et al.: Use of degenerate primers for PCR detection of GLRaV-4 and GLRaV-5.1995 1996 1996 1996 1996 1996 Greif et al.: The sequenced genome of GLRaV-2 has the same structural organization of that of Beet yellows virus. Al-Tamimi et al. Choueiri et al.: Association of GLRaV-2 in Italy and France with a graft incompatibility revealed by Kober 5BB. Rowhani and Uyemoto: Comparative trials between indexing and laboratory detection methods show that the latter are more sensitve for GLRaVs detection. Fortusini et al. Routh et al.: Leafroll and associated closteroviruses in Jordan. Petersen and Charles: Pseudococcus calceolariae acts as vector of GLRaV-3 in New Zealand. Gozsczynski et al.: Comprehensive review of the properties of GLRaVs. None of 223 accessions of European. Habili and Nutter: In an Australian commercial vineyard GLRaV-3 incidence increased from 23.: Leafroll and associated closteroviruses in Lebanon.GLRaV-5 induces mitochondrial vesiculation.: GLRaV-3 is transmitted by Planococcus citri in a semipersisten manner. Ling et al.: Elimination of GLRaV-3 by heat therapy improves agronomic performances of a Nebbiolo clone and the quality of the must. Faoro: Comprehensive review of the ultrastructure of GLRaVs infections. Cabaleiro et al. Martelli et al.: GLRaV-3 detected in native American vines in Western New York. and Asian Vitis species inoculated by green grafting with a GLRaV-1 and GLRaV-3 sources were resistant. Zhu et al. Guidoni et al.: Cloning an sequencing of the coat protein gene of GLRaV-3 and its expression in transgenic tobacco. the type specied of the genus Closterovirus. Krastanova et al. Lahogue and Boulard: Search for genes of resistance in grapevines.1% in 1986 to 51. GLRaV-3 appears to be a typical closterovirus.: Identification of two different mechanically transmissibile strains of GLRaV-2. Viruses are irregularly distributed in the vines. 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1998 1998 1998 1998 1998 1998 1998 67 . Haidar et al.: Distribution and incidence of GLRaVs in Canadian viticultural districts.9% in 1996. La Notte et al.
Karasev: Review article on closteroviruses.: Use of degenerate primers for PCR detection of GLRaV-1 and GLRaV-7. maritimus. Golino et al.: Completion of the nucleotide sequence of GLRaV-3 genome and use of the HSP90 and coat protein genes for producing transgenic rootstocks.: Evidence that GLRaV-1 and GLRaV-3 are serologically related based on the cross-reactivity of a monoclonal antibody raised to GLRaV-1.1998 2000 2000 2000 Saldarelli et al. Golino et al. Meng et al. : Partial sequencing of GLRaV-7 genome. transmitted by mealybugs and soft scale insects. Rowhani et al. Gugerli: Detection of GLRaVs by Lumino-ELISA.: Evidence that vines sanitized from leafroll have superior qualitative and quantitative traits. Turturo et al. affinis and Planococcus citri transmit with various efficiency GLRaV-3 in California but not GLRaV-1. Monis: Identification of GLRaV-8 and production of monoclonal antibodies. Mannini and Credi.: Survey of GLRaVs in Mediterranean and Near East countries. Boscia et al. Evidence of the quasispecies nature of the virus. Little et al. Seddas et al. a genus with monopartite RNA species. The type species is GLRaV-3 2000 2000a 2000b 2000 2000 2000 2000 2000 2000 2000 2000 2000 2000 2000 2000 2000 2000 2001 2002 68 .: Revision of the family Closteroviridae. Ps. a chemiluminometric enzyme-linked immuosorbent assay.: Detection in California of a GLRaV-2 strain sharing about 74% sequence homology with GLRaV-2 type strain and reacting weakly serologically with GLRaV-2.: Mealybug species Pseudococcus longispinus. Kim et al. Digiaro et al. GLRaV-2 and GLRaV-4. Martelli et al.: Intriguing association of GLRaV-6 with cv. Ling et al. Ps. a new genus having GLRaV-3 as type species. Sforza et al. Abou Ghanem -Sabanadzovic et al.: Identification of hypervariable regions in GLRaV-1 genome. Cardinal in Italy and Greece. Proposal for the establishment of Vinvirus (Ampelovirus) . Establishment and description of Ampelovirus . Zhou et al. viburni. Demonstration that the membranous vesicles accumulating in the cytoplasm derive from proliferation of the endoplasmic reticulum. Fazeli and Rezaian: Partial sequencing of GLRaV-1 genome. Gonsalves: Review article on the molecular traits of GLRaVs. Castellano et al: Ultrastructural study of GLRaV-2 and GLRV-7 infections.: Leafroll and associated closteroviruses in South Korea. Ps. Production of a new set of monoclonal antibodies.: New monoclonal antibodies to GLRaV-2.: Completion of the nucleotide sequence of GLRaV-2 genome.: Association of an unusual strain of GLRaV-2 with a graft incompatibility condition described from California as young vine decline.: Identification and partial characterization of a new strain of GLRaV-2.: New vectors of GLRaV-1 are the mealybugs Heliococcus bohemicus and Phenacoccus aceris and the soft scale insect Pulvinaria vitis. one of which is especially suited for ELISA testing.
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M. 1238-1243. Ben-Dov and B. Rowhani. 2000. Martelli. Preliminary investigations of genetic variability of Grapeviune leafroll associated virus 3 isolates. Adelaide 2000. and G. A comparison between serological and biological assays in detecting grapevine leafroll-associated viruses.. Sasahara H. 1981. Proceedings 9th Meeting of ICVG. A. Proceedings 9th Meeting of ICVG. Indexing grapes in Japan for viruses.P. 67-69.. Sannino F. 1998. 1997. A. 1989. Le rugeau de la vigne. 38. Segura A.. Progrès Agricole et Viticole 45. 1986. 14. 704-709. Routh G. 2000. Use of degenerate primers for partial sequencing and RT-PCR-based assays of grapevine leafroll-associated viruses. 80-85. Digiaro.. Uyemoto. V. Jelkmann and G. A. Uyemoto. Extended Abstracts 14th Meeting of ICVG.. Cabaleiro. Greif. Tanne. 68-69. P. Kiryat Anavim 1987. Saldarelli. Haidar. 192-196. 2000. Jacquet. Y. C. Chen and D. Guglielmi Montano and G. 162-163.. 799-801. D. Walter. Hu and D. and P.P.. 1973. Transmission of closterolike particles associated with grapevine leafroll by mealybugs (Pseudoccidae) in Israel.. Horticulture 18. Martelli.P.A. Martelli and B. Phytopathologische Zeitschrift 80. Isolation and partial characterization of two new viruses from grapevine. Mein Winzerbuch. 433-436.K. Y. 157-160. Plant Pathology 43. Kiryat Anavim 1987.E. Zhang. 91-96. 1993. 1982a. P.L. Effect of heat therapy and meristem tip culture on the elimination of grapevine leafroll-associated closterovirus type III. Plant Disease 71. Martelli. Establishment of a relationship between grapevine leafroll closteroviruses 1 and 3 by use of monoclonal antibodies. D'Onghia and G. European Journal of Plant Pathology 104. Partial molecular characterization and RT-PCR detection of grapevine leafroll-associated virus 7..L. 71-73. Boscia. 1987. Tanne E. 207211. 1994a. Minafra.. 35-38. Adelaide 2000. Presence of grapevine leafroll in North West Spain.. Zeitschrift für Pflanzenkrankheiten und Pflanzenschutz 89. M. 222-223.. 345-346. Walter. Y.S. Yafeng.S. D.M. Rowhani. 1976. and P. Golino. Adelaide 2000. Non radioactive molecular probes for the detection of three filamentous viruses of the grapevine.. Rowhani A. 176-180. J. 82. G. Gonsalves.M. W. Rosciglione B. 1994. Verge. 1994b. and J. Savino and G. Tazaki. Turturo C. Rowhani A. Phytopathology 88. Saldarelli P. Rosciglione B. T. 1924. 110-113. and F. Martelli. Rivista di Patologia Vegetale 1. 11-17. 1989 Detection of a grapevine leafroll-associated closterovirus in recently infected tissues in New York and spread of the disease in Mexico. Savino V. Gonsalves and F.P. Revue Suisse de Viticulture. Turturo C. Minafra and M. 135-141. Histological and cytological studies on the infectious leafroll disease of the grapevine. Zhang . Extended Abstracts 11th Meeting of ICVG. 222-225 Tanne E. Volos 1990. M. Teliz D. Von der Brelie D. Virus diseses of grapevine in Israel. Seddas A. Sforza R. Transmission of grapevine leafroll virus to herbaceous plants. Nitzany. I. 125-126. 74 . Gugerli 1989. Journal of the Japanese Society for Horticultural Science 50. Zee.P.. 1935. C. 169175. Hummer. D. Plant Pathology 49. D.. Saldarelli and A. with special reference to closteroand vitiviruses of the grapevine. Sela and I. Transmission of grapevine leafroll disease and an associated closterovirus to healthy grapevine by the mealybug Planococcus ficus Signoret. Detection of grapevine leafroll-associated closterovirus III by molecular hybridization. 17-18.Ravaz L. Proceedings 10th Meeting of ICVG. 356358. Greif. Scheu G. 508-517. D. Annals of the Phytopathological Society of Japan 42. and J. The occurrence of grapevine leafroll disease among the main grapevine cultivars and breeding stocks in Taiwan.A . P. Teliz D.. Locorotondo 2003. 1906. Reichnährstand Verlag. 1991.P. Nienhaus. H.P. M. Raccah. Saldarelli. Tanne E.J. M. Extended Abstracts 13th Meeting of ICVG. Regeneration of plantlets by meristem tip culture for virus-free grapevine. Il rossore delle viti.D. New scale insect vectors of grapevine closteroviruses. Rott. Use of degenerate primers in a RT-PCR assay for the identification and analysis of some filamentous viruses. Tzeng. A. E. Berlin.K..C. Scheu G. Arboriculture. Tanaka S. Die Rollkrankheit des Rebstockes. 945-950. Iri. Vitis 33. 8689. Komar and C.. 1974. Cloquemin and B. Plant Disease 81. Gugerli. Tada. Plant Pathology Bulletin 3. Maladies de l'enroulement et du bois strié de la vigne: analyse microscopique et sérologique. Der Deutsche Weinbau 14. Field serological detection of viral antigens associated with grapevine leafroll disease. Phytoparasitica 17. G. Minafra. 156-167.. 1998. 1936. M. and F. Saldarelli P.. K. 2000. Gonzales and C. Vitis 12. Extended Abstracts 13th Meeting of ICVG. Digiaro.K. Saldarelli P. Montreux 1993. Extended Abstracts 13th Meeting of ICVG. A. Harpaz. Tzeng H. V.. Takezawa and M.
. and D. Zhu H. J.V. Further characterization of closterovirus-like particles associated with the grapevine leafroll disease. C. Gonsalves. Walter and O.. Gonsalves. 277-288. R. Potere. D. Z. 1958. 130. 1987. 731-741. Agronomie 8. C. Comptes Rendues de l'Academie d'Agriculture de France 44. Abou-Ghanem. Boscia. 1289-1298. Walter B. Zimmermann. 1427-1434. the closterovirus type member. Vernoy. 2000. Production and characterization of monoclonal antibodies specific for Grapevine leafroll-associated virus 3 and epitope mapping of the coat protein. N.F. Martelli. Pool and R. 1988... 2003. Walter and M.A..S. Castellano. 1991. A. P.. and G. Cytopathology of leafroll diseased grapevines and the purification and serology of associated closteroviruslike particles. 1998. 62-66. Bass. 75 . Phytopathology 77. Wilcox F. Nucleotide sequence and genome organization of Grapevine leafroll-associated virus 2 are similar to Beet yellows virus. 1990. Sommermeyer.. Potere. Saldarelli. O. Journal of General Virology 79. Zee F. Proceedings 10th Meeting of ICVG.W. D. D. 1998 Leafroll virus is common in cultivated American grapevines in Western New York. Turturo. P. Extended Abstracts 14th Meeting of ICVG. Kim. Boscia. Goheen.E. G.P. Zhou Z.. Lee. Collas and G. McFerson and D.Vuittenez A. D. Zimmermann D. 203. K. B. R.R. Purification de particules virales associées à l'enroulement de la vigne et mise au point d'un protocole ELISA permettant leur détection. O. R. 1990.H. 1062. Extended Abstracts 13th Meeting of ICVG. Martin. Jiang and D.E Goszczynski. Walter B. Monoclonal antibodies for detection and characterization of Grapevine leafroll-associated virus 2. Zhou Z. 137-145.Y. Legin. Production and characterization of monoclonal antibodies specific to closterovirus-like particles associated with grapevine leafroll disease. Gonsalves. D.S. Vesselle. Ling. Adelaide 2000. Locorotondo 2003. Transmission par greffage d'une virose type enroulement foliarie commune dans le vignobles de l'Est e du Centre-Est de la France. Zimmermann D. K. B. Plant Disease 82. Van Regenmortel. The use of a greengrafting technique for the detection of virus-like diseases of the grapevine. Goszczynski and M. A. Journal of Phytopathology 130.Y. Journal of Phytopathology 128. Le Gall. Volos 1990. 313-316.
RUGOSE WOOD COMPLEX .
RUGOSE WOOD COMPLEX
The rugose wood complex consists of several diseases (Grapevine rupestris stem pitting, Grapevine kober stem grooving, Grapevine corky bark, Grapevine LN33 stem grooving) that are usually latent in ungrafted Vitis vinifera and American Vitis species and rootstock hybrids, but develop in grafted vines. Woody cylinder alterations resembling rugose wood symptoms are reported in the French literature of the early 1900s as possible physiological disorders. Rugose wood was first identified and described from southern Italy in the early 1960s as a graft-transmissible disease, and was considered to be a local problem until its discovery in Hungary in 1967. Now it is known to occur worldwide. 1. Description Main synonyms: Stem pitting, stem grooving (Eng.); legno riccio (Ital.); bois strié, cannelures du tronc (Fr.); madera rizada (Sp.); lenho rugoso (Port.); corky bark: rough bark (Eng.); suberosi corticale (Ital.); écorce liégeuse (Fr.); Korkrindenkrankheit (Germ.). Symptoms: Affected vines appear less vigorous than normal and may show delayed bud opening in spring. Some decline and die within a few years from planting. Grafted vines often show a swelling above the bud union and a marked difference between the relative diameter of scion and rootstock. With certain cultivars, the bark above the graft union is exceedingly thick and corky, has a spongy texture and a rough appearance, a condition known as "corky rugose wood". The woody cylinder is typically marked by pits and/or grooves which correspond to peg-and ridge-like protrusions on the cambial face of the bark. These alterations may occur on scion, rootstock or both. The severity of wood symptoms vary according to scion/stock combinations. Climatic conditions may have a bearing on symptom expression for under cool and wet climates symptoms are milder or absent. Cases of latent infection in grafted vines are not rare. By contrast, self-rooted European grapes and, sometimes, American roostocks, can show wood alterations. No specific symptoms are seen on the foliage, although certain cultivars show rolling, yellowing or reddening of the leaves similar to those induced by leafroll. Bunches may be fewer and smaller than normal and the crop reduced by 20-30%. The four diseases of the rugose wood complex can be recognized and sorted out by graft transmission to the indicators Vitis rupestris, LN 33 and Kober 5BB: a. Rupestris stem pitting. Distinct basipetal pitting limited to a band extending downwards from the point of inoculation in V. rupestris. LN 33 and Kober 5BB remain symptomless. b. Corky bark. Grooving and pitting of the entire surface of the stem of V. rupestris and LN 33, but no symptoms in Kober 5BB. Severe stunting of LN 33 is accompanied by rolling and reddening of the leaves and by most typical internodal swelling of the canes. c. Kober stem grooving. Marked grooving appear on the stem of Kober 5BB; no symptoms in V. rupestris and LN 33. d. LN 33 stem grooving. Grooves occur on the stem of LN 33, much the same as with corky bark, but no internodal swelling of the shoots nor foliar discolorations are present. V. rupestris and Kober 5BB show no symptoms. Agents: Putative agents of individual diseases of the rugose wood complex are members of the genera Vitivirus or Foveavirus, family Flexiviridae, i.e. viruses with flexuous filamentous particles from about 730 to 800 x 12 nm, with distinct transverse cross banding. Vitiviruses and foveaviruses are phloem-restricted in grapevines, but whereas vitiviruses are mechanically transmissibile to herbaceous hosts, though with difficulty, foveaviruses are not. The genome of all viruses consists of a single species of single-stranded positive sense RNA with Mol. wt 2.6-3.05 x 106 that accounts for c. 5% of the particle weight. Coat protein subunits have a single size and Mr of 22-28 kDa. Rugose wood-associated viruses have a worldwide distribution. Records exist from Europe, the Mediterranean basin, Near and Far East, Australasia, South Africa, and North and South Americas. Grapevine rupestris stem pitting-associated virus (GRSPaV), a definitive member of the genus Foveavirus, is the associated agent of Grapevine rupestris stem pitting disease. Virus particles are about 730 nm in length and are not readily observed with the electron microscope. GRSPaV occurs in nature as
a family of molecular variants. Viral RNA, which has been totally sequenced, has a Mol. wt of about 3.05 x 106 Da and a size of 8726 nt. The viral genome comprises 5 or 6 ORFs encoding, in the order, the replication-associated proteins (244 kDa), movement proteins (triple gene block, 25, 13 and 8 kDa) and the coat protein (28 kDa). The 6th ORF, when present, encodes a 14 kDa proteins with unknown function. GRSPaV seems to be more closely related to potexviruses than carlaviruses both of which have a similar genomic organization. These relationships have evolutionary implications and suggest that GRSPaV may have evolved from an ancient recombiantion event between a carlavirus and a potexvirus, in which ORF 4 and 5 but not the 3' non coding region of the carlavirus were replaced by those of the potexvirus. Grapevine virus A (GVA), the type species of the genus Vitivirus, is the putative agent of Grapevine kober stem grooving. Virus particles are flexuous filaments about 800 nm long. Viral RNA has a Mol. wt of about 2.6 x 106 Da and a size of 7349 nt. The viral genome consists of 5 ORFs encoding, in the order, the replication-associated proteins (195 kDa), a 20 kDa protein with unknown function, the movement protein (31 kDa), the coat protein (22 kDa) and a 10 kDa product which has nucleotide binding properties, is a pathogenicity factor and a gene silencing suppressor. Minor biological and serological variants of the virus are known. Grapevine virus B (GVB) is a vitivirus distantly related serologically to GVA and one of the etiological agents associated with Grapevine corky bark. GVB is also involved in young grapevine decline, a graft incompatibility condition recorded from California. Its totally sequenced RNA has a Mol. wt of about 2.7 x 106 Da, a size of 7599 nt and the same gene sequence and structural organization as GVA. This virus occurs in nature as a family of molecular variants, but biological variants are also known, two groups of which can be differentiated by the reaction of herbaceous hosts. Virus particles coated by both GVA and GVB coat protein occur in cells infected contemporarily by both viruses (phenotypic mixing). Grapevine virus C (GVC) is a little known and poorly characterized virus reported from Canada. Virus particles have a vitivirus morphology and an estimated length of about 725 nm. GVC is serologically distinct from GVA and GVB. Grapevine virus D (GVD), a vitivirus distantly related serologically to GVA and GVB is associated with with corky rugose wood, a field syndrome characterized by the presence of a striking corky condition of affected vines, just above the graft union. Virus particles are flexuous filaments about 825 nm long. The viral genome, which was sequenced only in part, has an estimated size of c. 7600 nt and a 3' terminus structurally comparable to that of GVA and GBV. Cytopathology: Whereas no information is available on the cytopathology of GRSPaV infections, vitivirus-induced cellular modifications have been extensively studied, primarily in herbaceous hosts. Cytopathological features common to all four vitiviruses (GVA, GVB, GVC, and GVD) consist of: (i) virus particle aggregates of various size, forming bundles, whorls, banded bodies, stacked layers that sometimes fill the entire cell lumen; (ii) variously extended wall thickenings originating from deposits of callose-like substances; (iii) proliferation and accumulation of cytoplasmic membranes; (iv) vesiscular evaginations of the tonoplast protruding into the vacuole and containing finely fibrillar material resembling dsRNA. GVA and GVB movement proteins were found to associate with cell walls and plasmodesmata, as detected by gold immunolabelling. Transmission: For many years after its discovery there were no records of natural spread of rugose wood in the field. GVA and GVB are now known to be transmitted from grapevine to grapevine by pseudococcid mealybugs and/or scale insects in a semipersistent manner. GVA vectors are the mealybugs Planococcus citri, Pl. ficus, Pseudococcus longispinus, Ps. affinis, Heliococcus bohemicus, and the scale insect Neopulvinaria innumerabilis, whereas GVB is transmitted by Ps. longispinus, Ps. affinis, and Pl. ficus. GRSPaV has no known vectors, but is suspected to be pollen-borne. There are, however, conflicting reports on its presence within seed and no evidence that it occurs in seedlings from infected vines. None of the putative agents of rugose wood has alternative hosts in nature and, because of the relatively limited range of vector movement, is not disseminated over long distances by natural means. Transport of infected propagative material represents the major means of dispersal. The presence of rugose wood and its causal agents in phylloxera-free countries with a millenial history of ownrooted grapevine cultivation, suggests that the disease originated in the Old World and was distributed worldwide by commercial trading and planting of infected grafted plants. Varietal susceptibility: Most if not all V.vinifera varieties and American rootstocks are susceptible. Although customarily grapevines are infected symptomlessly when ungrafted, rugose wood symptoms
other assays include: single step or nested reverse trascription-polymerase chain reaction (RT-PCR). rupestris. since symptomless infections make sanitary selection not totally reliable. Latent infection can occur also in grafted vines. possibly in relation with the scion/stock combination and climatic conditions. the putative agent of rupestris stem pitting. a virus-like disease. or a combination of the two. Beukman and Goheen: Brief account of the histological modifications of corky bark-affected LN 33. if not otherwise associated with a specific syndrome. to a restricted range of herbaceous hosts (mostly Nicotiana species). though with difficulty. described from California. Thus. Detection: Indexing on indicators (V. Vitiviruses. In general. Faccioli: First histological study of corky bark-affected grapevines. and that it may be a composite disease resulting from the interaction of different viruses among which GFLV. several resistant plant lines were obtained by transformation with the coat protein and the movement protein genes of GVA and GVB. Kober 5BB and LN 33) is the only reliable method for detecting and sorting out the diseases of the complex. 2. Goidanich and Canova: First record of corky bark in Europe. 1965 1965 1967 81 . Martelli et al. Graniti and Martelli: Demonstration of the infectious nature of rugose wood. Goheen et al. Transgene expression was detected in these plants and in transformed grapevine explants.: Graft transmission of rough bark to LN 33.: First record of rugose wood outside of Italy. GVA can be eliminated to a very hight rate (up to 97%) by the procedure used for cryopreservation of grapevine shoot tips. The intensity of wood abnormalities (pitting and grooving) vary. meristem tip culture.were observed in self-rooted cultivars and ungrafted roostock stocks (V. experimental evidence has been obtained of the very close association of GRSPaV. it is plausible to regard 110R as a specific indicator of rupestris stem pitting in addition to V. No natural sources of resistance to any of the rugose wood agents are known but the possibility of using pathogen-derived resistance in Vitis is being explored. are mechanically transmissible. as shown by 110R. immunocapture RT-PCR. Graniti: Detailed description of rugose wood symptoms. rupestris. all sources must be indexed and/or laboratory tested. or spot-RT-PCR using degenerate or virus-specific primers. no strategy has yet been developed for the chemical control of vectors. rupestris and Kober 5BB). Histological study of diseased vines. "stem pitting" and "stem grooving". Historical review Names like "legno riccio". Graniti and Ciccarone: First record of rugose wood from southern Italy. are synonymized with "rugose wood". Suggestion that rugose wood may be a disease of combination requiring the contact of scion and rootstock for the development of symptoms. The best antigen sources for serological diagnosis are cortical shavings from mature dormant canes. Individual viruses can be identified by ELISA or dot immunobinding on nylon membranes using polyclonal antisera and/or monoclonal antibodies when available. Immuno-capture RT-PCR is 1000-fold more sensitive than ELISA for virus detection in grapevines. with vein necrosis. 1954 1961 1962 1963 1963 1964 1965 Hewitt: Rough bark. Hewitt et al.: Corky bark is remarkably heat stable and difficult to eliminate by heat therapy. Recently. However. Suggestion that it may be caused by a virus. Name of the disease changed into corky bark. rugose wood agents can be eliminated with reasonable efficiency by heat therapy. In addition. Control of mealybugs is difficult for they overwinter under the bark of grapevines and possess an unwettable waxy covering. but not foveaviruses. Thus. Control: Use for propagation of virus-free scionwood and rootstocks obtained by sanitary selection combined with sanitation is of paramount importance to avoid introduction of infected vines in the vineyards. Using a Nicotiana benthamiana model system.
1968 1968 Lehoczky et al.: Green grafting useful for corky bark indexing. Boccardo and D'Aquilio: Physicochemical characterization of GSP-AV Abracheva: Survey of over 650 grapevine cultivars and hybrids for rugose wood reaction in Bulgaria.: Observation of rugose wood symptoms in self-rooted vines. Engelbrecht and Nel: Rugose wood and fanleaf are not related. First record of rugose wood symptoms outside of Europe (Israel). is transmitted by the pseudoccid mealybug Pseudococcus longispinus. Mink and Parsons: Use of growth chambers for rapid symptom expression of corky bark in Vitis indicators. Hewitt: Successful graft transmission of Californian rugose wood. Graniti and Martelli: Up-to-date review of rugose wood. Goheen: Evidence that corky bark and leafroll.c: A series of three papers reporting the occurrence and field spread of corky bark in Mexico and evaluating symptoms induced by natural infections of corky bark in formerly virusfree self-rooted or grafted European grape varieties and rootstocks. Goheen and Luhn: Suggestion that corky bark and rugose wood are the same disease. Beukman and Goheen: Up-to-date review of corky bark. based on graft transmission tests.: Rugose wood recorded from Australia. Anonymous: A review of rugose wood in Italy. Teliz et al.b. a. Hewitt and Neja: First record of rugose wood in USA (California). At 38 °C the minimum inactivation period for leafroll is 56 days and for corky bark 98 days. Conti et al. No nepoviruses implicated in their etiology.: Recovery by mechanical inoculation of a closterovirus with particles 800 nm long. Beukman and Gifford: Detailed account of adverse effects of corky bark on the anatomy of Vitis. 1968 1969 1970 1970 1971 1971 1972 1973 1973 1975 1975 1977 1978 1979 1979 1980 1980 1981 1981 1982 1983 82 . Bovey and Brugger: Further evidence that GFLV may not be implicated in the etiology of rugose wood in Switzerland. Lehoczky: Destructive effects of rugose wood registered in Hungary in both self-rooted and grafted European grape varieties. Goheen and Luhn: Heat treatment of dormant buds grafted onto LN 33 is effective against corky bark. Sarooshi et al.Virus provisionally called grapevine stem pitting-associated virus (GSP-AV). Hewitt: Up-to-date review on grapevine virus and virus-like disease worldwide.: First experimental evidence that a filamentous virus (GVA). despite similarities in the s y m p t o m s o n t h e foliage are different diseases. Rugose wood may not require a grafted plant for full symptom expression. Rosciglione et al. Castillo et al. Legin et al.: Heat therapy effective against rugose wood. from a rugose wood-infected vine.
Rugose wood and corky bark are not the same disease. ficus. thesis describing rupestris stem pitting disease in comparison with corky bark. Savino et al. No closterovirus-like particles in samples with rupestris stem pitting. Garau et al. an additional disease of the rugose wood complex.: Experimental confirmation that rugose wood may not express symptoms in grafted indicators.: Transmission of corky bark by the mealybug P. Prudencio: M. Corky bark and Rupestris stem pitting. Suggestion that the disease is not related to closteroviruses associated with GLRaV and corky bark.ficus.: A low molecular weight dsRNA associated with rupestris stem pitting.: First indication of the possible existence of LN 33 stem grooving. Engelbrecht et al.: Application of spot hybridization for the detection of GVA in grapevine sap.: First record of rugose wood from China.: Three types of wood disorders of the stem-grooving type observed in South African grapevines. Li et al. Savino et al. wt of 5. Martelli: Rugose wood recorded in southern Mediterranean and Arab countries. Monette and James: Detection of two biologically distinct but serologically indistinguishable isolates of GVA. Similar dsRNA species were detected.: Two distinct dsRNAs with a mol.Sc. similar to Kober stem grooving.: Evaluation of the effect of rugose wood on cv. Garau et al. GSP-AV re-named Grapevine virus A (GVA). Azzam et al.3 and 4. Savino et al. Engelbrecht and Kasdorf: Natural field spread of corky bark in South Africa associated with the presence of P.4 x 106 associated with rupestris stem pitting in grapevines from California and Canada. First report of Kober stem grooving. ficus. 1985 1985 1985 1985 1985a 1985b 1985 1985 1985 1987 1988 1989 1989 1989 1989 1989 1989 1990 1990 1991 1991 83 . denoted Grapevine virus B (GVB). Kuniyuki and Costa: Rugose wood recorded from Brasil Goheen: First published description of rupestris stem pitting.: Heracleum latent virus and GVA are distantly serologically related. Corbett and Wiid: Closterovirus-like particles found in extracts from vines affected by corky bark and rugose wood in South Africa.: Experimental confirmation of the complex nature of rugose wood based on the differential reaction of woody indicators. Gallitelli et al. Castrovilli and Gallitelli: Physicochemical comparison of two Italian isolates of GVA. Tanne et al. Monette et al.1984 Milne et al. Murant et al.: Evidence that GSP-AV can occur in grapevines together with another similar but serologically unrelated virus with short closterovirus-like particles. Italia propagated on six different rootstocks.: Assessment of crop losses induced by rugose wood to two different European grape varieties. Rosciglione and Castellano: Demonstration that GVA is transmitted also by Planococcus citri and P. but not consistently in grapevines from New York. The first two disorders appear to be spreading in the vineyards.
: Presence of two distinct serotypes of GVA.: Contemporary occurrence of Rupestris stem pitting and Kober stem grooving in symptomless scions of cv.: Rugose wood recorded from Ukraine Boscia et al. Monette and James: A closterovirus with short particles (725 nm) isolated from a corky barkaffected vine induces necrotic local lesions and systemic symptoms in Nicotiana benthamiana.: Purification and properties of GVB.1991 1991 Gugerli et al. Boscia et al.: Production of monoclonal antibodies to GVA and their use for ELISA detection of the virus in infected vines. Virus later identified as Grapevine leafroll-associated virus 2 Tanne and Meir: A dsRNA with a molecular weight higher than 14 Kd identified in extracts from corky bark-affected vines. Saldarelli et al. Martelli et al. Suggestion that GVA may be the causal agent of the disease.: Rugose wood recorded from Yemen.: Rugose wood recorded from Malta Monette and Godkin: Recovery of a closterovirus-like virus by mechanical inoculation from a corky bark-affected vine.: Clear-cut connection of GVA and rugose wood. Namba et al.: Rugose wood recorded from Albania. Virus named Grapevine virus C (GVC). 1991 1991 1991 1991 1991 1991 1991 1991 1992 1992 1993 1993 1993 1993 1994 1994 1994 1994 1994 1994 1994 1994 1995 84 . Garau et al. both associated with a stem pitting condition of grapevines rather than with leafroll.: Development and diagnostic use of a cloned probe to GVB. Minafra and Hadidi: Detection of GVA and GVB in viruliferous mealybugs by PCR.: Development of digoxigenin-labelled riboprobes for the detection of GVA and GVB in infected tissue extracts. Virus transmission by the mealybug Ps. Digiaro et al. Martelli et al. Saric and Korosec-Koruza: Rugose wood recorded from Croatia and Slovenia Ioannou: Rugose wood recorded from Cyprus. Minafra et al. Padilla: Rugose wood recorded from Spain Boscia et al. ficus induced corky bark symptoms in LN 33 Saldarelli et al. Thorough comparative study of nine GVB isolates from different countries.: Sequence of the 3' end of GVA and GVB genome. Suggestion that GVA is implicated in the aetiology of the disease. Garau et al.: A closterovirus with particles 1440-2000 nm long serologically unrelated to all other known grapevine closteroviruses found in corky bark-affected vines.: GVA and Kober stem grooving are closely associated. Merkuri et al. Torbato in Italy. Boulila et al. Both viruses qualify for the inclusion in the genus Trichovirus. Minafra et al.:Rugose wood recorded from Tunisia Milkus et al.: Synthesis of a cloned probe for GVA. Chavez and Varon de Agudelo: Rugose wood recorded from Colombia.
: Rugose wood recorded from Lebanon. Chevalier et al. Saldarelli et al. this may be the first visualization of GRSPaV. and GVD) later assigned to the genus Vitivirus.: GVB is consistently associated with corky bark and is present.: GVA is transmitted by by Ps. Rubinson et al.: Sequencing of a Californian isolate of GRSPaV. Efficient detection method based on TAS-ELISA developed. longispinus in a semi-persistent manner.: A study of the spatial distribution pattern of corky bark in aThompson seedless vineyard in Israel. Martelli and Jelkmann: Establishment of the genus Foveavirus. Choueiri et al. 1995 1995 1995 1996 1996 1996 1996 1996 1997 1997 1997 1997 1997a 1997b 1997 1997 1997 1997 1998 1998 1998 1998 85 .: Sequence and structrural organization of Grapevine rupestris stem pittingassociated virus genome (GRSPaV).: Consistent detection of GVA in Kober stem grooving-infected grapevines by immunocapture-polymerase chain reaction. La Notte et al.: Development of a PCR technique for the detection of GVA and GVB in nylon membrane-spotted sap.: Nucleotide sequence of GVA genome and taxonomic position of the virus.: GVA and GVB are transmitted by Pseudococcus affinis. GVB. Abou Ghanem et al. GRSPaV is assigned to this genus.: Rugose wood recorded from Jordan. Guidoni et al.: Nucleotide sequence of GVB genome. GVC. Faoro: Review of the cytopathology of grapevine trichovirus infections. Since particle size (600-700 nm in length) is compatible with that of Grapevine rupestris stem pitting-associated virus (GRSPaV) particles identified in 2002. GVC. Garau et al.: Estasblishment of the genus Vitivirus with GVA as type species. The virus is not seed-borne.: GVA and GVB are serologically related.: GVA and GVD are serologically distantly related. Tanne et al. and GVD removed from the genus Trichovirus and assigned to the new genus. Boscia et al. Boscia et al. though not consistently in vines showing a syndrome denoted "corky rugose wood". La Notte et al. GVA. Haidar et al. Alkowni et al: Rugose wood in Palestine. Zhang et al. Martelli et al.: Description of Grapevine virus D (GVD). Bonavia et al.: Review of the properties of putative grapevine-infecting trichoviruses (GVA.: Elimination of GVA from cv Nebbiolo clones by heat therapy improves agronomic performance of the vines and quality of the must. Minafra et al. Goszczynski et al. Suggestion that spreading is by a vector that transmits in a semipersistent manner.1995 Monette and Godkin: Detection of non mechanically transmissible capillovirus-like particles in a grapevine affected by rugose wood. Further support of the cause-effect relationship between GVA and this disease.: Antiserum to the movement protein of GVA is useful for virus detection in ELISA. Meng et al. GVB.
Samples made up of three buds from dormant canes are less laboriour to prepare than cane shavings and yield comparable results. i.: Movement proteins of GVA and GVB detected by gold immunolabelling in association with cell walls and plasmodemata of infected cells. Goszczynski and Jooste: GVA and GLRaV-3 are both consistently associated with Shiraz disease in South Africa but only GVA seems to be required for disease induction. Petrovic et al.: GVA transmission by Pseudococcus comstocki. Meng et al. Dovas and Katis: Improved RT-PCR method for the simultanous detection in grapevine extracts of vitivirus (GVA. Habili et al. Nucleotide sequence identity within groups is 91-99. observed for the first time.: Consistent association of GRSPaV with vines indexing positive for Rupestris stem pitting. Wang et al. Boscia et al. Confirmation of the cause-effect relationship between GVA and Kober stem grooving.: Rugose wood viruses in Egypt.e. GVA coat protein encapsidating GVB RNA. Ahmed et al.: Production of monoclonal antibodies to GVB.: GVA particles carry a highly structured epitope centered on a common peptide region of the coat protein sequence. GVB and Corky bark and GRSPaV and Rupestris stem pitting.: Western blots and ELISA using a polyclonal antiserum to recombinant coat protein of GRSPaV detect the virus in infected grapevine tissues almost with the same efficiecy of RTPCR. Virus detected in bleached seeds suggesting that it is present inside the seeds. Saldarelli et al. intermingled with virus particle aggregates. Galiakparov et al.8% and 78-89.: Elimination of GVA by cryopreservation. Goszczynski and Jooste: Thre groups of GVA strains (I. Dell'Orco et al. Seyval seedlings.3% among groups. Kominek et al. GVA movement protein is also present in great quantity in the cytoplasm. and III) identified in South Africa based primarily on sequence homology of the 3' end of the viral genome.: Rugose wood viruses in Iran. Saldarelli et al. II.: The function of GVA genes identified by mutation analysis of individual ORFs of a full-length infectious viral clone. 2000b 2000 2001 2001 2001 2002 2002 2003 2003 2003 2003 2003 2003 2003 2003 2003 2003 2003 86 .: Synthesis of full-length cDNA copies of GVA and GVB genomes.: Production of a polyclonal antiserum to a recombinant coat protein of GRSPaV and its use in dot immunobinding on polyvynil difluoride membranes for virus detection in grapevine tissue extracts. GCD) and foveavirus (GRSPaV) sequences in two steps.: One-sided phenotyping mixing. Nakano et al.: GRSPaV particles. GVB. Goszczynski and Jooste: Use of single-strand conformation polymorphism reveals molecular heterogenity in GVA populations.1999 1999 2000a Meng et al. occurs in Nicotiana plants doubly infected with GVA and GVB. Further support to the cause-effect relationship of GRSPaV with this disease Galikparov et al. Stewart and Nassuth: An improved extraction method allows RT-PCR detection of GRSPaV virtually thoughout the year in all grapevine tissues.: Production of an infectious RNA transcript from a full-length cDNA clone of GVA. are filamentous and measure 723 nm in length. Minafra et al. The virus was not detected in 245 seedlings from infected cv.: Rugose wood viruses in the Czeck Republic. Buzkan et al.
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GRAFT INCOMPATIBILITY .
Foliar and trunk symptoms resemble very much those induced by rugose wood/graft incompatibililty and are shown by aged as well as young (4-year-old) vines. Known viruses associated with this disorder (different GLRaV-2 strains and GVB) can be identified by ELISA using polyclonal antisera and/or monoclonal antibodies The best antigen sources for serological diagnosis are cortical shavings from mature dormant canes. Syrah decline is a severe disease occurring in France. with margins more or less extensively rolled downwards. Based on the differential responses of a panel of 18 rootstocks. associated with grapevine bushy stunt is still unidentified. Other molecular variants of GLRaV-2 were reported from New Zealand (Alphie virus). In this case.GRAFT INCOMPATIBILITY Infection by phloem-limited viruses may damage grapevines in the nursery (reduced graft take) or in the early stages of growth in the field (graft incompatibility). The heat-labile graft-transmissible agent present in the hybrid 140R. However. European grape varieties grafted on tolerant rootstocks (e. Harmony. though not consistently and. 03916. but the colour of the canopy remains green. Freedom. Argentina and probably elsewhere. 3309) develop a discolored canopy . leaves are small-sized. Detection: Indexing on Caberent sauvignon is a reliable method for detecting incompatibility conditions. 101-14) exhibit a green canopy and perform rather well. A transitory form of incompatibility was reported from Italy under the name of bushy stunt. Cases of graft union disorders have been observed in Europe (Kober 5BB incompatibility). Agents: An ordinary strain of Grapevine leafroll-associated virus 2 (GLRaV-2) is consistently associated with Kober 5BB incompatibility (Europe).g. Salt creek . California. Severely affected vines decline and may die within one or two years.g. decline and may die. and again California (rootstock stem lesions). scions show a stunted and bushy vegetation due to the contemporary proliferation of apical and axillary buds. so as to represent veritable emerging diseases. and Australia in association with young vine decline conditions.). Of these. The putative agent of bushy stunt was consistenly found in clones of the rootstock 140R in which it is latent. Varietal susceptibility: Appearance of graft union disorders depends more on the rootstock rather than the scion. appears to be involved in California's young vine decline. but the yield is reduced. is not transmitted by mealybugs and does not have a known vector. Normal growth resumes with the second or third leaf. The nature of this disease has not been ascertained but one or more graft-transmissibile agents may be involved in its aetiology. A prominent swelling forms at the scion/rooststock junction and variously extended necrotic lesions may develop on the rootstock stem. incompatibilità d'innesto (Ital. except for GRSPaV. Australia and Chile (young vine decline).) Symptoms: Newly planted vines grow weakly. only GLRaV-2 RG was identified. and together with Grapevine virus B (GVB). Other assays include nucleic acid-based tecniques such as single step or nested reverse transcription-polymerase chain reaction (RT-PCR) and 95 . Redglobe in California called Grapevine rootstock stem lesion-associated virus (GRSLaV) proved to be a molecular and biological variant of GLRaV-2 (GLRaV-2 RG). 5C. shoots are short. consistently. Chile. although none of a number of known grapevine-infecting viruses has been found in affected vines. This latter condition has been known for a long time and occurs also in rugose wood-affected vines. Infected propagative material is to be blamed for its dissemination. a member of the genus Closterovirus. GVB is mealybug-borne and can be spread at a site by these insects. 1. Transmission: GLRaV-2. 1103P. Description Main synonyms: Incompatibilité au greffage (Fr. A virus originally detected in cv. New Zealand. The same virus was detected in diseased Chilean grapes. Kober 5BB. Incompatibility may also develop in the form of a brown line of necrotic tissues at the bud union when grape cultivars hypersensitively resistant to the nepovirus ToRSV are grafted on susceptible rootstocks. in Argentinian grapes. which are usually not accompanied by wood abnormalities (pitting or grooving). the increased use of clonal material is disclosing unprecedented conditions of generalized decline that develop dramatically in certain scionrootstock combinations. up to five different grafttransmissible agents inducing incompatibility could be differentiated in California. whereas varieties grafted on susceptible roostocks (e. and the vegetation is stunted The canopy shows autumn colours off season so that leaves turn reddish in red-berried varieties or yellow in white-berried varieties much earlier than normal.
Durquety et al. Uyemoto and Rowhani: Indexing on 18 different grape rootstocks reveals the existence of at least five different agents causing graft incompatibility. meristem tip culture. Boubals: Report of a national French study group investigating the aetiology of Syrah decline. Martelli: GRSLV and GLRaV-2 are serologically related and are both recognized by a panel of 18 monoclonal antibodies.immunocapture RT-PCR. Bonfiglioli et al.: GLRaV-2 is the cause of a graft incompatibility revealed by Kober 5BB. 2. Legin and Walter: The graft-transmissibile agent that causes incompatibility of different varieties on Kober 5BB is a virus which can be eliminated by heat treatment at 37 °C for 58 days. though not consistently.: Third paper of a series on incompatibility on Kober 5BB. Boubals: Syrah decline occurs in Argentina Uyemoto et al. Currently known graft incompatibility agents can be eliminated with reasonable efficiency by heat therapy.: GLRaV-2 is associated.: Report of a new molecular variant of GLRaV-2 from New Zealand. utilization of tolerant roostocks is advisable. 1977 1979 1986 1991 1995 2000 2000 2000 2001 Jacob: Description of graft incompatibility in different scion/stock combinations. Boubals and Huglin: Report on graft incompatibility of certain varieties grafted on 57R.: Identification of an apparently new closterovirus denoted Grapevine rootstock stem lesion virus (GRSLV) causing stem necrosis of rootstocks. whenever feasible. or a combination of the two.: GLRaV-2 and GVB are consistently associated with young vine decline in California. The problem is very complex and may involve several still unidentified factors. Renault Spilmont et al. If scionwood is infected. Greif et al. 2003 2003 2003 2003 2003 2003 2004 96 . Graft-transmission of the incompatibility factor.: Updated report on the state of the art of investigations carried out in France on Syrah decline. Historical review 1942 1950 1973. GRSLV has about 75% nucleotide homology with GLRaV-2. No conclusion are drawn. the use of sensitive rootstocks is to be avoided and. decline. Suggestion that they are molecular variants of the same virus species. GRSLV re-named Redglobe strain of GLRaV-2. and death of the vines. Savino et al. Bertazzon and Angelini: Comparison of several detection methods for the broad or specific identification of Grapevine leafroll-associated virus 2 variants. Control: Prevent introduction of infected vines in the vineyard by using certified grafted plants or virusfree scionwood and rootstocks.: Description of bushy stunt and evidence that it is caused by a graft-transmissibile heat-sensitive agent carried by some clonal rootstocks.: Two papers describing incompatibility phenomena between clonal selections of different cultivars grafted on Kober 5BB Fallot et al. Strategies on how to protect healthy stocks from vector-mediated GVB reinfection in the field are yet to be developed. with a decline condition of young Thomposn seedless vines in Chile. using degenerate or virus-specific primers. Golino et al. Gomez Talquenca et al: GLRaV-2 is consistently associated with declining Cabernet sauvingon vines grafted on different roostocks in Argentina. Prodan et al.
2000. Grau. Savino V. Rowhani. 1973. Golino D. C. 137-141 Boubals D. Walter. Pantaleo.M. Locorotondo 2003. Le clone et ses reactions au greffage. Montalegre and N. Fallot. Etude de l'incompatibilité au graffege de certain cépages et du 57R.S. Benassac and R. Progrés Agricole et Viticole 94. 1991. Molecular studies on a graft incompatibility syndrome in New Zealand vineyards yields another probable variant of Grapevine leafroll-associated virus 2. Compte-rendu de la réunion du Groupe de Travail National. La trasmission de l'incompatibilité au greffage entre 5BB et Vitis vinifera.M. Le clone et ses reactions au greffage.E. Ruchaud.. J. 141. Greif C. R. M. Grapevine virology highlights 2000-2003. Renault Spilmont A. 167-173. Prota. A young grafted vine decline syndrome in Argentina vineyards. 142-143. Syrah decline in French vineyards.. 28-31. Volos 1990. Durquety and J. Extended Abstracts 14th Meeting of ICVG. V. 2003. Proceeding of the American Society of Horticultural Science 41.3. I. Locorotondo 2003. Gazeau and J. 202-210 Uyemoto J. Durquety P. C. Huglin. 1979. Identification of the latent viruses associated with young vine decline in California. Garcia Lampasona and O.. 1995. Progrés Agricole et Viticole 67. B. 2003. Angelini. 2004. 201-203. Le dépérissement de la Syrah existe aussi en Argentine. J. O. and E.. S. Fallot.. Martelli G. 1950. Discovery of different grapevine sources with graft-transmissible agents causing union-incompatibility on sensitive rootstocks. Rowhani. P.. 277. 183-189. Edwards and A. J. D. 2003. 122-129 and 171-178. Gracia. Progrés Agricole et Viticole 117. 211-216. Le clone et ses reactions au greffage. Jacob H. Garau. Proceedings 10th Meeting of ICVG. References Bertazzon N. Fiore. California Agriculture 55 (4). The relationship of grapevine leafroll-associated virus 2 with a graft incompatibility condition of grapevines. 2001.P.. and B. Durquety P. Ruchaud. Locorotondo 2003. 1977. Di Terlizzi. Boubals D..II. Le dépérissement de la Syrah. Advances in the detection of Grapevine leafroll-associated virus 2 variants.A. Rivieccio and F. Legin R. Locorotondo 2003.P. D. 2000. Gazeau. Locorotondo 2003.S. B. Di Silvio. S.III. 2003. 2003. Examples of incompatibility between grape varieties and roostocks.. 139-140. Fiori. F. and P. Gomez Talquenca G. 2003. Extended Abstracts 13th Meeting of ICVG. and A. Krag. Ruchaud. Extended Abstracts 14th Meeting of ICVG. Extended Abstracts 14th Meeting of ICVG.A. J. Nouvelles recherches sur l'incompatibilité clonale d'Abouriou greffé sur 5BB. S.M.P. Prota. 85-86. Progrés Agricole et Viticole 90. 1986. Bass. Extended Abstracts 14th Meeting of ICVG.. Progrés Agricole et Viticole 117. New closterovirs in 'Redglobe' grape causes decline of grafted plants.P.. Grenan and J. Sim and A. P. Existence dans un cépage population de clones présentant divers degrés de compatibilité avec certains porte-greffes. Locorotondo 2003. 279-283. Presence in clonal rootstocks of a grafttransmissible factor that induces stunting and bushy growth in European grapevines. S. Dauty.. Extended Abstracts 14th Meeting of ICVG. Luvisi and R.. Journal of Plant Pathology 86. 420-427 Fallot J.M. Progrés Agricole et Viticole 96. 283-290. Boursiquot. 2000. Uyemoto J. 3-10. Etude de phénomènes d'incompatibilité au greffage chez la vigne.. Autres cas chez la vinge. Rowhani.. Bonfiglioli R. Boubals D. Phytopathologia Mediterranea 34.K. Prodan S. 85-86. C. 1942. Adelaide 2000. Boscia. 144. Aetiology of decline in Thompson seedless grafted table grape plants. Progrés Agricole et Viticole 103. Walter and U. A. 97 . Extended Abstracts 14th Meeting of ICVG.K.
FLECK COMPLEX .
positive sense RNA with high cytosine content (c. Grapevine asteroid mosaic-associated virus (GAMaV). adverse influence on vigour. vinifera. Affected vines are often stunted. grapevine rupestris necrosis. Description Main synonyms: A.). GFkV genomic RNA (Mol. Records from Italy and South Africa have not been confirmed experimentally and a record from Greece was proven to refer to Grapevine rupestris vein feathering. rupestris. Symptoms are expressed in Vitis rupestris and consist of clearing of the veins of third and fourth order. twisted and puckered along the veins. is latent in European grapevine varieties. c. In V. and produce little or no fruit. and grapevine rupestris vein feathering) and viruses (Grapevine redglobe virus) that cause latent or semi-latent infections in Vitis vinifera and most American Vitis species and rootstock hybrids. Asteroid mosaic. Fleck. Sternmosaik der Rebe (Germ. V. 1. b. producing localized translucent spots. 25 kDa. but likely to occur elsewhere. vinifera in California. Grapevine red globe virus (GRGV) is a Grapevine fleck virus (GFkV)-like virus which apparently does not induce symptoms in European grapevine varieties (e. The coat protein (CP) of GFkV and GRGV particles is made up of a single protein species with Mr of c. Rupestris necrosis. which is used as indicator. Severe strains induce also varying degrees of stunting. Recorded from California and Italy . This disease.g. B. GAMaV. Although the elusive nature of the complex hinders the assessment of its economic impact. twisted and may curl upward. which are twisted and asymmetric. e. grapevine asteroid mosaic. Red globe) nor in V. sometimes with necrotic center. irregularly distributed over the leaf blade. rupestris following graft inoculation. GFkV genomic RNA constitutes about 35% of the particle weight. rupestris reacts with localized necrosis of the shoots. containing the genome.). In V. whereas the CP of GAMaV and GRVFV consists of a major protein of 21 kDa and a minor prtotein of 25 kDa. Grapevine fleck: Marbrure (Fr. Symptoms: a. Leaves are asymmetric. The disease is latent in European grapevine varieties and in most American rootstocks. d. screziatura (Ital.). Sultanina). mosaico stellare (Ital.). All have isometric particles about 30 nm in diameter with rounded contour and prominent surface structure with clusters of coat protein subunits arranged as pentamers and hexamers. Rupestris vein feathering. and GRVFV genomes are available. Fleck is an ubiquitous disease reported from most viticultural countries in the world. Marmorierung der Rebe (Germ. rooting ability of rootstocks and on graft take has been reported. Agents: All viruses of the complex.). leaf symptoms are characterized by star-shaped chlorotic spots. wt of 2. and Grapevine rupestris vein feathering virus (GRVFV) are all phloem-limited and non mechanically transmissible. rupestris. T made up of empty protein shells and B. which is a monopartite single-stranded. Grapevine asteroid mosaic: Mosaïque étoilée (Fr. 50%). Leaf symptoms usually become less severe in summer. Asteroid mosaic symptoms have been observed in several varieties of V. GFkV particles sediment as two centrifugal components.). The putative causal agent of the disease has only been found in California. GRGV. The complete sequence of GFkV and partial sequences of GRGV.g. the disease elicits creamy-yellow bands developing along the major veins of the leaves.6 x 106 ) is 7564 nt in size and contains four open reading frames (ORF) that 101 . Mild asteroid mosaic-like symptoms are shown by some European grapevine varieties (e. capped.FLECK COMPLEX The fleck complex consists of several diseases (grapevine fleck. Italy and California. Transient mild chlorotic discolourations of the primary and secondary leaf veins develop in V. reported only from Japan. leaf petioles and veinlets. GFkV. The putative causal agent of the disease so far has been found in Greece. Leaves with intense flecking are wrinkled. maculatura infettiva.
Although observations from Italy. GRGV.: "Marbrure". Comparison of symptoms with those of other mosaic diseases of grape. whereas GAMaV induces peripheral vesiculation of chloroplasts. Polyclonal antisera and monoclonal antibodies to GFkV heve been raised. South Africa and Japan suggest natural field spread of GFkV and a similar behaviour was reported in Greece for a disease formerly thought to be asteroid mosaic but now identified as grapevine rupestris vein feathering. GAMaV.encode a 215. Attempts to transmit the disease by mechanical inoculation to herbaceous test plants or by Xiphinema index were unsuccessful. Transmission: No vector is known for any of the viruses of the fleck complex. The same sanitation procedures are likely to operate successfully with the other viruses of the complex. rupestris allows with a reasonable level of confidence the discrimination of the different viruses of the complex based on the differential reaction of the indicator. but no experimental data are available. whilst GAMaV and GRVFV were assigned to the genus Marafivirus. Detection: Indexing on V. Because of its molecular characteristics.4 polypeptide with the conserved motif of replication associated proteins (ORF 1). of which it represents the type species.rupestris described in France. The genomic structure of GAMaV and GRVFV differs from the above in that both these viruses have a single ORF encoding a large polypeptide which is proteolitically processed to yield individual proteins. GFkV was identified as the representative of a new genus denoted Maculavirus. GFkV can be eliminated by heat therapy. primary dissemination of these and the other viruses of the complex is through infected propagative material. rupestris St. and two proline rich polyproteins of 31. Transmission through dodder of GFkV has been reported but it is of no epidemiological importance. Further physico-chemical. molecular and ultrastuctural studies disclosed sufficient similarities between maculaviruses. As the disease is rare and does not appear to be spreading. but cannot be used for any of the other members of the complex due to the unvailability of antisera. Vuittenez et al. ELISA is currently employed for routine detection of GFkV.: First record of fleck as an unidentified symptom different from fanleaf and transmissible from symptomless varieties to V. a disease inducing symptoms similar to those of fleck in V. The current taxonomic classification of viruses of the fleck complex is therefore the following: Family Tymoviridae Genus Marafivirus Grapevine asteroid mosaic-associated virus Grapevine rupestris vein feathering virus Genus Maculavirus Grapevine fleck virus Grapevine redgloble virus Cytopathology: GFkV infections are characterized by a severe modification of mitochondria into structures called "multivesiculate bodies". Control: Because of the latency of symptoms sanitary selection of European grapevine cultivars and most American rootstock hybrids is unrealiable.4 kDa (ORF 3) and 15. Hewitt et al. No information is available on individual susceptibility. The 3' end of GRGV genome is structurally similar to that of GFkV except for the lack of ORF 4. 2. marafiviruses and members of the genus Tymovirus to warrant the establisment of the a new family denoted Tymoviridae. its economic importance is low. Historical review 1954 1962 1966 1966 Hewitt: First description of asteroid mosaic in California. Refatti: Review paper on asteroid mosaic. These deranged organelles are thought to be sites of virus replication. the CP (ORF 2). Virus specific and degenerate primers have been designed for single or multiplex RT-PCR detection of GFkV. meristem tip or fragmented shoot apex culture. Therefore. 102 .George. GFkV is not seed transmitted.9 kDa (ORF 4) with unknown function. and GRVFV. Varietal susceptibility: GFkV and possibly all the other viruses of the complex infect naturally a large number of varieties and Vitis species.
later called grapevine phloem-limited isometric virus (GPLIV). rupestris propagating wood. Mink and Parsons: Use of a growth chamber with controlled temperature for a quicker and improved symptom expression of fleck and other virus or virus-like diseases (fanleaf.: Observation of a non mechanically transmissible virus.: Report of widespread occurrence of fleck in visually selected grapevine clones in southern Italy. rupestris.: Physicochemical characterization of GPLIV. Hévin et al. Ottenwaelter et al. Confirmation that the virus can be eliminated by heat therapy and is not related to leafroll.: Description of fleck as an independent graft-transmissible disease present in many European varieties and American rootstocks. Dolja et al. Savino et al. 1982 1983 1983 1983 1984 1985 1985 1987 1989 1990 1990 1990 1991 103 . Castellano et al. The efficiency of heat treatment for disease elimination is unsatisfactory. Verderevskaya et al. Report of multivesiculate inclusion bodies probably connected with GPLIV infection. in sieve tubes of field-grown vines with leafroll symptoms but likely to be affected by other diseases. Engelbrecht and Kasdorf: Observation of natural field spread of fleck in South Africa.1970 1972 1972 1972 1973 1973 1973 1974 1977 Refatti: Symptoms resembling asteroid mosaic as described in California are reported from Italy and South Africa.: Identification of a dsRNA of about 7 Kb pairs in diseased vines.: Observation of an isometric non mechanically transmissible virus in the phloem of diseased vines. Triolo and Resta: Tetracycline treatments are ineffective against fleck. Barlass et al. Dismissal of the prokaryote etiology hypothesis. Woodham and Krake: Dodder transmission of fleck from vine to vine. Bovey: Identification of fleck in Switzerland as a latent disease of Chasselas transmissible to V.: Purification of GPLIV from naturally diseased vines and production of a specific antiserum.: Successful elimination of fleck through heat therapy. Triolo and Materazzi: Fleck has a detrimental effect on the quality V. Castellano and Martelli: Confirmation that GPLIV is associated with multivesiculate bodies and demonstration that these derive from deranged mitochondria. Rives: Further demonstration that fleck is distinct from fanleaf based on differential responses to heat treatment. Goheen and Luhn: A novel heat therapy system based on virus inactivation in buds grafted onto healthy LN 33 rootstocks is effective against fleck. leafroll and corky bark). Rooting ability and graft take are adversely affected Yamakawa: Field spread of fleck in Japan Boulila et al. Castellano et al. Milkus: Suggestion of a prokaryotic etiology for fleck.: Successful elimination of fleck through fragmented shoot apex culture in vitro.: Fleck is not seed transmissible. Hewitt et al. Report that a virus serologically similar to GPLIV is associated with the disease.
vinifera.: Molecular reagents (degenerate primers) developed for the specific identification of viruses of the fleck complex (GFkV. Boscia et al.1991 1991a 1991b 1991 Gugerli et al. rupestris with a necrotic disease.rupestris of an isometric virus latent in V. based on the differential reactions of indicators Credi and Babini: Infection by fleck. One of these antibodies is more sensitive than the polyclonal antiserum for GFkV detection by ELISA Faoro and Gugerli: An unidentified phloem-limited isometric virus serologically differing from GFkV observed in vines showing double-membraned peripheral invaginations of chloroplast envelope.: Additional monoclonal antibodies raised in France. The disease seems to be spreading naturally.: GPLIV shown to be the agent of fleck.: A spherical virus purified from berries of Ajinashica disease-affected vines is serologically related to GPLIV (= GFkV) and has physicochemical properties comparable to those of GFkV. 1991 1993 1994 1995 1995 1996 1996 1997 1997 1998 2000 2001 2001 104 .: Two GFkV-specific monoclonal antibodies raised in Italy can successfully be used in ELISA Kuniyuki and Costa: Three strains of GFkV reported from Brazil. GRGV). GAMaV and of another virus with GFkV-like particles phylogenetically but not serologically related to GFkV present in a cv Red globe vine. June-July are the best months for ELISA detection of the virus in Alsace. Boscia et al. Marsumoto and Ohki: A spherical virus resembling GFkV identified in thin sectioned cells of V.: A non mechanically transmissible isometric virus similar but unrelated to GFkV identified in asteroid mosaic-infected grapevines. Symptoms are severe and affected vines are almost fruitless.: Use of degenerate primers designed on the methyl transferase and polymerase cistrons of members of Tymovirus and Marafivirus genera and of GFkV amplified a genome fragment of GFkV. vinifera cv. Walter and Cornuet: Confirmation by ELISA of the consistent association of GFkV with fleck disease. Elbeaino et al.: Report of a highly consistent association of GPLIV with fleck in naturally infected and graft-inoculated V. Sabanadzovic et al. rupestris. vein necrosis and vein mosaic has a detrimental effect on rootstock growth. Fortusini et al. GFkV-like multivesiculate bodies derived from deranged mitochondria are present in infected cells. Pruning wood is reduced by 51% in 420A and by 37% in Kober 5BB. Boscia et al.: Natural field spread of GFkV observed in Northern Italy Schieber et al. ELISA used successfully for virus detection in large scale survey.: Complete nucleotide sequence of GFkV genome. Virus named Grapevine redglobe virus. Kyriakopoulou: Description of a disease similar to asteroid mosaic observed in V. Sabanadzovic et al. Sultanina in Greece. This cytological feature recalls that later found in vines infected by Grapevine rupestris vein feathering virus. Molecular properties of this virus further support the notion that it warrants classification in a genus of its own. Detection of sequences of an undentified virus from a Greek grapevine. GAMaV. Virus named Grapevine asteroid mosaicassociated virus (GAMaV) Boscia et al. Virus renamed Grapevine fleck virus (GFkV).: Report of the close association with fleck symptoms in V. Adverse effect on Teleki 5A is negligible. Meristem tip culture effectively eliminates the virus. later named Grapevine rupestris vein feathering virus (GRVFV). Namba et al.
Dolja V. Martelli.A. Rowhani P. 1995..G. Savino. Boscia D. Annales de Phytopathologie. Sabanadzovic. Martelli and M. leafroll and Shiraz decline diseases and associated viruses in South African grapevines. Bovey R. G. 1972.P. Virus-like particles and ultrastructural modifications in the phloem of leafroll-affected grapevines. Phytopathologia Mediterranea 24. R. Babini. 2003b 2003 3. Journal of Phytopathology 129. Abou Ghanem-Sabanadzovic et al.G. Sabanadzovic . S. fleck.E. Molecular detection of Grapevine fleck virus-like viruses. G. Savino and M. a new genus of plant viruses having GFkV as type species and GRGV as tentative species. Skene. K. D. Some properties of a phloem-limited non mechanically-transmissible grapevine virus. GAMaV and of a virus of Greek origin which induces vein feathering in V.. Tomashevskaya.P. Argentina. Virus Genes 27.V. Identification of the agent of grapevine fleck disease.. G.A. Boscia D. 1982. 151-158.A. Greek virus recognized as a species in its own right denoted Grapevine rupestris vein feathering virus (GRVFV). V. Savino and D. Vitis 33. A non mechanically transmissible virus associated with asteroid mosaic of the grapevine. Field spread of corky bark. Journal of Ultrastructure Research 89.R. Martelli et al. N. Vitis 40: 65-68. Martelli.: Sequencing of the 3' end of the genome of GRGV. respectively.: Description of Maculavirus. 1983. and A. Sabanadzovic. Sequencing of the 3' end of three grapevine fleck virus-like viruses . 160-163.D. V.P. Castellano. A... Phytophylactica 22. and Japan) only the variant without insertion (GFkV353) was detected. Proceedings 8th Congress of the Mediterranean Phytopatological Union. Castellano M. South Africa. 23-39. Iran. T. 173-174. 97-105. Boyko. rupestris confirms the assignment of GRGV to the genus Maculavirus and of GAMaV and the Greek virus to the genus Marafivirus. 2003b. 291-295.: A sequence variant of GFkV (GFkV416) with a 63 nucleotide insertion in the replicase gene identified in Australia and New Zealand..: Description of the family Tymoviridae. Martelli.F. P. 1990. and G. Boscia. Castellano M. A. M. Engelbrecht D.M. Annals of Applied Biology 101. Savino. U. 1985. 3134..P. V.A. 105 .V. 1994. GRVFV recorded from California and confirmation that GAMaV does not occur outside of California. Savino and G. Un virus latent dans le Chasselas. 1990. 165-169.. Savino.P. Karsev. 347-354.: Development of a multiplex RT-PCR protocol for the simultaneous detection of GFkV-like viruses using plant mRNA as an internal control. Association of a phloem-limited non mechanically transmissible isometric virus with grapevine fleck disease.P. Martelli. Savino. Double stranded RNA associated with fleck disease of grapevine. 1996. 1991a. 191. 1991b.P. Martelli. Rowhani and G. 95-98. References Abou Ghanem-Sabanadzovic. Boscia D. Extended Abstracts 14th Meeting of ICVG. Martelli.2002a 2002b 2003a Martelli et al. Production of monoclonal antibodies to grapevine fleck virus. 1990. 56-64. 2003a. Kyriakopoulou and G.. V. Effect of virus and virus-like infections on the growth of grapevine rootstocks.P.E. In other countries (USA. Atabekov. O.A. Proceedings 10th Meeting of ICVG. Kyriakopoulou and G. S. Vitis 22. Martelli 2001. Barlass M. S. Progress in the study of the phloem-limited isometric virus-like particles associated with leafroll-diseased grapevines. A.G. N.J. Abou Ghanem-Sabanadzovic. S.R. N. A. and G. and G. Agadir 1990. V.. Minafra. Kasdorf. Elicio. 101-102 Boscia D. Vitis 30. Martelli. Abou Ghanem-Sabanadzovic et al.P.P. Verderevskaya and J. Boulila M. Locorotondo 2003. G. Credi R. Advances in Horticultural Science 10. M. Digiaro. Sabanadzovic and G. Volos 1990. 1984. Castellano M. Plant Pathology 44. B. comprising the genera Maculavirus and Marafivirus that include GFkV/GRGV and GAMaV/GRVFV. Numéro hors série. V.A. V. Woodham and L. Multiplex RTPCR detection of Grapervine fleck virus-like viruses in grapevine with co-amplification of control plant mRNA. Martelli. Martelli and V. Ultrastructure and nature of vesiculated bodies associated with isometric virus-like particles in diseased grapevines. 11-16 Abou Ghanem-Sabanadzovic. Boscia. Castellano. Shi et al.P. Di Terlizzi. Krake..P. Regeneration of virus-free grapevines using in vitro apical culture. Castellano. ElBeaino T. 195...C.
Rosciglione. Phytopathologia Mediterranea 24. M. Ohki. Annales de Phytopathologie. 560-564.Faoro F and P. Investigating the transmission of marbrure and fan-leaf through the seed in the grapevine.. L. Diffusione naturale del virus 1 (GLRaV-1) e del virus 3 (GLRaV-3) dell'accartocciamento fogliare e del virus della maculatura infettiva o "fleck" (GFkV) della vite. Martelli G. 1973. 9. Martelli. Proceedings 10th Meeting of ICVG. 57-83.C Edwards and T.. and J. Martelli. M. Rivista di Patologia Vegetale. 43-47. Abou Ghanem-Sabanadzovic. 2003. G. Goheen A.P. Phytopathologia Mediterranea 24. and A. Grapevine fleck disease. a new genus of plant viruses. Raski and G. 1970. 39-43. 5960. Symons.L. Cytological alterations associated with an unidentified isometric grapevine virus (UIGV). 2000. Informatore Fitopatologico 46 (12). M. 75-77. 1977. Acta Phytopathologica Academiae Scientiarum Hungaricae 9. Mycoplasma. Further characterization of grapevine leafroll disease. 2001. Matsumoto T.. latent in many varieties. Archives of Virology 147. Walter..M. Refatti E. Fortusini A. In Frazier N. 1997. N. A. Luhn. Bonnard. Hewitt W. 1991. Nucleotide sequence variation in a small region of Grapevine fleck virus replicase provide evidence for two sequence variants of the virus. 2002b. 47-64. Tsuchizaki and D... A. Boscia and G. Davis 1965.A. Lisbon 1997.P. D. Kuniyuki H. Ottenwaelter M.P.. J. 1972.IV. is transmitted by graft inoculation.V. 253-258. 1837-1846. J. Rivista di Patologia Vegetale. Castellano. 143-146. Procedures for rapid detection of virus and viruslike diseases of grapevine. Grapevine fleck virus-like viruses in Vitis. J. Berkeley.IV. University of California. Dreher. Goheen. Annals of the Phytopathologial Society of Japan 64. 1997. 1985. 567-571.P.P. Monoclonal antibodies for detection. 2002a. Volos 1990. S. D. Cinquanta and S. Jr.P. and C. A. 349-355. Yamashita. Hewitt W. C. T. Belin and B.P. serological characterization and immunopurification of grapevine fleck virus. 2009-2015 Savino V. Vitis 3. Rives. 1972. and E. Symptoms of grapevine asteroid mosaic in Greece. Rivista di Patologia Vegetale.B. Schieber O. Shi B. 1962.. B.. Ser. Sabanadzovic. N. IV. M. 204-207. 618-622.. Numéro hors série. Refatti E.H. P. Ramel and F.E. Prati. Abou Ghanem-Sabanadzovic. 12491253. 157-164. Plant Disease 75. Doazan and M. Sabanadzovic S. 1973. 281-285. Triolo E. Maculavirus. Martelli. Proceedings 10th Meeting of ICVG. George).. D. Heat therapy eliminates the ability to transmit the causal agent of "marbrure" in several V. Annals of Applied Biology 142. 1847-1853. 767-774. affected by marbour. 1995. Journal of General Virology 82. Digiaro and G. European Journal of Plant Pathology 103.-E.C. Bulletin of the California Department of Agriculture 43. Numéro hors série. S. M. Goheen. Brugger. Gugerli P. 1991. Sabanadzovic. Ottenwaelter. Hévin. 9.. N. Plant Disease Reporter 61. 1985. Parsons. Studies on virus diseases of the grapevine in California. Mink G. 1974.or chlamidia-like bodies in grape. Gooding. Hewitt W.J. Annales de Phytopathologie. Rives.. Volos 1990. Gugerli. A possible new necrotic diseases of grapevine associated with small isometric particles and novel membrane-bound large particles. vinifera clones and in V. Saldarelli. Resta.T. 1954. rupestris "du Lot" (St. Proceedings International Conference on Virus and Vector on Perennial Hosts. M. S. N. Abou Ghanem-Sabanadzovic and P. Seddas. Leclair. Costa. 106 . Gonsalves. 1966. Abou Ghanem.. The family Tymoviridae. Archives of Virology 147. Heat inactivation of viruses in grapevines. Rives M.. Some virus and virus-like diseases of grapevines. 385-388. and S.): Virus Diseases of Small Fruits and Grapevines (A Handbook). 1973. Doazan and M.. The responses of the Grapevine fleck agent to tetracycline-HCl antibiotic and Dienes' stain. with Special Reference to Vitis. Boscia. Tremea. 197-203. Luhn. S. 287-289. N. Martelli G. 9. Asteroid mosaic of grapevine. Ser. (Ed.W. Hévin M. Cory and C.S.C. Kyriakopoulou P. 31-32.. 1998. Habili and R.C. Division of Agricultural Sciences.. 1991.I. 212-214. Sabanadzovic S.. Incidence of some graft-transmissible virus-like diseases of grapevine in visually selected and heat-treated stocks from Southern Italy. Milkus B. S. P. Saldarelli and G.B. Namba S. Séparation de la marbrure et du court-noué (panachure) chez la vigne par thermothérapie. 553-565. 1996. Scattini.-J.M. Occorrencia de mais um isolado do virus do mosaico da nervuras da videira que no causa sintomas no porta-enxerto Kober 5BB. Ser. Archives of Virology 145.. Extended Abstracts 12th Meeting of ICVG. Fitopatologia Brasileira 20.B. Purification and properties of spherical virus particles associated with grapevine Ajinashica disease. Grapevine asteroid mosaic. Complete nucleotide sequence and genome organization of grapevine fleck virus.J.
Vuittenez A. 221-226. 107 . Agronomie 13. Marinesku and E. 1987. and L. George.D.G. Yamakawa Y. La maculatura infettiva della vite: influenza di isolati diversi sull'attitudine alla propagazione vegetativa di Vitis rupestris St. Krake. probablement indépendante du virus du Court-noué.. and A. 1993. Legin and J. 1966. 1989. ELISA detection of grapevine fleck virus (GFkV). 67-73. Investigations on transmission of grapevine leafroll. Archiv für Phytopathologie und Pflanzenschutz 19. La Recherche Agtonomique en Suisse 26. 3209-324. 651-657.. V. Woodham R. Phytopathologische Zeitschrift 106. Kuszala. Walter B. Annales des Epiphyties 17. Cornuet. 1983.Triolo E. Numéro hors série. Journal of the Japanese Society of Horticultural Science 58. and P. yellow speckle and fleck diseases by dodder. 297-302.. R. Verderevskaja T. Ätiologie und Diagnose der Marmorierung der Weinrebe. Semtschik. Observations sur une mosaïque de la Vigne. 193-198. Materazzi.. 1983.R.C.S. Virus reinfection of virus-free Cabernet sauvingon and Cabernet franc vines.
Agent: Alfalfa mosaic virus (AMV). Description Main synonyms: None. Control: Use of healthy material obtained by heat treatment. The spring growth shows more or less extensive yellowing of the leaf blades that does not extend to the veins. 0. however. Some of these diseases have been recorded only from Europe. but some are of economic relevance locally (e. Beczner and Lehoczky: AMV infections recorded from Hungary. has differently shaped particles.04 x 106 Da (3644 nt). Varietal susceptibility: Little information available. wts: RNA-1. rupestris and the hybrid Grézot 1 x 5C. 18% of the particle weight.MINOR VIRUS DISEASES Several graft-transmissible diseases are known. RNA-2. Hungary. 0. European diseases GRAPEVINE YELLOW MOTTLE 1. Switzerland. Bovey and Brugger: AMV recorded from Switzerland in grapevine and transmitted by grafting to V. and a tripartite RNA genome accounting for c. rings and lines are typical summer responses of infected vines. Jankulova: AMV infections recorded from Bulgaria. Chardonnay and Veltliner rouge précoce identified as reliable indicators. Novak and Lanzova: AMV infections recorded from hop and grapevine in Czechoslovakia. Transmission: AMV is efficiently transmitted by aphids in a non persistent manner and can cause epidemic outbreaks in many of its natural hosts. former Czechoslovakia. Yellow mottle has been reported from Germany. 1.73 x 106 (2593 nt).: First record of AMV infections and description of symptoms in German grapevines. Capsid proteins subunits are of one type. Plant vigour and yield do not seem appreciably affected.g. AMV. Detection: AMV is mechanically transmissible to herbaceous hosts and can also be identified by ELISA and moleculat techniques in infected vines. with Mr 24x 103 Da.62 x 106 (2037 nt). Main symptoms: Various patterns of yellow discolouration characterize the disease. 109 . Grapevine berry inner necrosis). a mechanically transmissible virus. from quasi isometric to bacilliform. Bulgaria. with the following mol. A. Bovey and Cazelles: AMV particles visualized in thin sectioned grapevine leaves. Virus elimination by treating 37 days at 37-38°C. the type species of the genus Alfamovirus. Their overall importance is minor if compared with that of the major diseases dealt with in previous chapters. and Turkey. Faint yellow speckling. with which specific viruses are associated and thought to be their possible causal agents. In grapevines. others occur in Japan. infections are scattered and occasional. Historical review 1973 1975 1976 1979 1978 1981 Bercks et al. 30 to 57 nm in size. 2. is the putative causal agent. RNA-3. There may be differential susceptibility among cultivars. suggesting that the virus spreads primarily through infected planting material.
. 166-171. Identification of alfalfa mosaic virus and tomato bushy stunt virus in hop (Humulus lupulus L.). 131-134. Rome. roughly following its contour. ed. 1993. Polyhedral Virions with Tripartite Genomes (R. is seed-transmitted and spreads with diseased propagative materials. 1975..B.-J. 1976. Grapevine line pattern virus (GLPV) a possible member of the genus Ilarvirus. Main symptoms: Leaves show bright yellow discolourations that form marginal rings. Akbas and Erdiller: AMV infections recorded from Turkey. or maple leaf-like line patterns typically confined to the petiolar area. D. Control: No information. Detection and Diagnosis of Graft Transmissible Diseases of Grapevines. New York . 1979. has differently shaped particles. Description Main synonyms: None.1985 1993 1993 Francki: Comprehensive review of the properties of AMV and other viruses with tripartite genome.). quasi spherical 25-30 nm in diameter to bacilliform 40 to 75 nm in length. and a multipartite genome. The viruses and their taxonomy. 1985.P. Bovey R. Grapevine disease in Hungary caused by alfalfa mosaic virus infection. or the upper part of the blade. Journal of Turkish Phytopathology 22. 1-18. 39. Investigation on certain viruses spread on grapevines in Bulgaria.I. Acta Phytopathologica Academiae Scientiarum Hungaricae 16. Martelli: Yellow mottle suggested as the name for the disease caused by AMV in grapevines. and J.B. Über den Nachweis des Alfalfa mosaic virus in einer Weinrebe. Brugger. 3rd International Congress of Plant Pathology. Lehoczky. . Lesemann and G. Alfalfa mosaic virus on grapevine. Martelli G. This disease is known to occur only in Hungary. Querfurth. 63-65. and J. 1993. Horticulture 7. Jubileum 75. 55-63. Researches on grapevine virus diseases and determination of their incidence in Ankara Turkiye. Agent: The putative agent. Cazelles. Jankulova M.I. Novak J. Francki. Plenum Press. Biologia Plantarum 18. Lanzova. 1973. Cordoba 1976. Francki R. Le virus de la mosaïque de la luzerne sur la vigne. GRAPEVINE LINE PATTERN 1.) and grapevine (Vitis vinifera subsp. Phytopathologische Zeitschrift 76. and O. 119-128. Monografias INIA No. References Akbas B. Bovey R. and G. Transmission: GLPV has no known vector. vinifera cultivars are susceptible. Munich 1978.18 . Proceedings 6th Meeting of ICVG. Several V. Bercks R. sativa DC. Historical rewiev 1985 Francki: Comprehensive review of the properties of AMV an other viruses with tripartite genome. Erdiller. Graft transmission to cv. 263 pp. Vigour and yield are reduced. 152-154. In: The Plant Viruses. 2.B. Varietal susceptibility: No information. (ed. scattered spots or blotches. 1981. 3. 110 . Arboriculture. Beczner L. Revue Suisse de Viticulture. FAO Publication Division. 1978.Hegi) plants in Czechoslovakia. Detection: GLPV is mechanically transmissible to herbaceous hosts. and J.
D. Castellano.: Purification and characterization of GLPV and suggestion that it is the causal agent of the disease. 1987. Plenum Press. Martelli and J. Lehoczky et al. 1-18. 61-79 Lehoczky J. Detection: Graft-transmission to V. Phytopathologia Mediterranea 31. a novel virus disease of grapevine in Hungary. Agent: Symptomatic grapevines were reported to be doubly infected by GFLV and Carnation mottle virus (CarMV) the type species of the genus Carmovirus. L. 3. In: The Plant Viruses.B..I. 111 . Polyhedral Virions with Tripartite Genomes (R. size and have low sugar content.P.. family Tombusviridae. GFLV may not be involved in the aetiology of the disease.P.. Historical review 1989 1991 Rumbos and Avgelis: Roditis leaf discoloration described in Greece. ed. The disease has been recorded only from Greece. with Mol. Description Main synonyms: None. 1992.. especially near the petiole Leaves are deformed in correspondence of discoloured sectors. 1985. Lehoczky J. J. 2. Mission. 1987. has a monopartite RNA genome accounting for c. References Francki R. 115-116. New York . However. or variously extended sectors of the leaf blade. Boscia. 1989. Kertgazdasag 19. Lazar.).A. Varietal susceptibility: No information. Beczner and G. M. The disease is graft-transmissible and spreads through infected propagating material.: Description of line pattern disease in Hungary. wt 1. The viruses and their taxonomy. 18% of the particle weight.I. RODITIS LEAF DISCOLORATION 1. Burgyan. Main symptoms: Symptoms are prominent in late summer and consist of yellow and/or reddish discolourations of the tissues along the veins. Burgyan. Transmission: No vector is known. Boscia. one of the putative agents of corky bark (rugose wood complex) has a very high association with diseased grapevines. the interveinal areas.B. L. J. Kiryat Anavim. Occurrence of the line pattern hitherto unknown virus disease of grapevine in Hungary. Beczner and G. Grapevine virus B (GVB). according to more recent findings.A. Evidence that a graft-and mechanically transmissible virus is associated with it. Proceedings 9th Meeting of ICVG.: Evidence that GLPV is transmitted through grapevine seeds. Line pattern. D. Viruses associated with the disease are readily transmitted by sap inoculation and can be detected by ELISA and molecular techniques. Farkas. Seed transmission of grapevine line pattern virus. Lehoczky et al. vinifera cv. G. CarMV is an isometric virus 30 nm in diameter. Martelli. Lehozcky J. Bunches are reduced in numbers.4 x 106 (4003 nt in size) and coat protein subunits of Mr 38 x 103 Da.1987 1989 1992 Lehoczky et al. Evidence of its grafttransmissibility. Israel. 23-30. Francki. Avgelis and Rumbos: Double infection of diseased vines by GFLV and CarMV reported. M. By converse. Farkas. Castellano. Control: No information. G.
I. Control: No information 2. Carnation mottle virus isolated from vines affected with "Roditis leaf discoloration". Proceedings 10th Meeting of ICVG. Infected grapevines are stunted. Whether this mechanism operates with grapevines has not been ascertained. and A.M. Description Main synonyms: None Main symptoms: Symptoms are chlorotic angular spots on the leaf blades. 1991. Avgelis. discoloration of tissues bordering the veins. thus is regarded as the agent of the disease..P. 2003. Avgelis. and I. N. 2000. crinkling and deformation of the leaves. Kyriakopoulou. P. A.: First record of GAMV. References Girgis S.M. Description Main synonyms: None Main symptoms: Grapevines of cv.: Evidence that GAMV is the agent of grapevine angular mosaic disease. Transmission: GAMV is pollen-borne in herbaceous hosts and was able to infect pollinated plants. P. A.E. Dovas. Journal of Phytopathology 125. Dovas. mechanical transmision to herbaceous hosts. and ELISA. 437-443. P. Katis. Bem. C. 1989. F. Tsagris.C. Volos 1990. 274-278. Extended Abstracts 14th Meeting of ICVG Locorotondo 2003. 19. Historical review 2000 2003 Girgis et al. Agent: Grapevine angular mosaic virus (GAMV). Avgelis and N. A. Girgis S. the closest being Tobacco streak virus (TSV). reproduced the field syndrome in mechanically inoculated grapevine seedlings. Varietal susceptibility: No information Detection: Indexing on cv. Plant Disease 84.I. Bem. YELLOW LINE PATTERN (Rasperry bushy dwarf virus) 1. Kyriakopoulou. 3. Girgis et al. Roditis leaf discoloration -. but differs from GLPV. S. Rumbos I. Laski Rizling from Slovenia exhibit a yellow line pattern syndrome resembling the grapevine line pattern disease described from Hungary. Flowers abortion results in straggly bunches with small wrinkled berries bearing non viable seeds.a new virus disease of grapevine: symptomatology and transmission to indicator plants. the only other ilarvirus reported from grapevine. A new ilarvirus isolated from grapevine in Greece. P. The etiology of a new virus disease: grapevine angular mosaic. Rumbos.E. GAMV is molecularly related to a number of ilarviruses. The disease has been recorded only from Greece. Baresana x Baresana. decline gradually and some die. Infected grafting material is likely to be responsible for virus dissemination.D.3. GRAPEVINE ANGULAR MOSAIC 1.D. C. a virus with a tripartite RNA genome and a 30 kDa coat protein. 112 . Sklavounos. Katis. Tzortzakakis and M.C.. 1345. F. References Avgelis A.
: First record of RBDV in grapevine. a dimeter of about 33 nm. Transmission: GINV is transmitted by grafting to grapevines and by mechanical inoculation to herbaceous hosts. 3. Mavric et al. is unknow. Locorotondo 2003. The virus spreads naturally in the vineyards.5 Kb in size) and RNA-2 with Mol. References Mavric I. 2003. Almost all Japanese table grape cultivars derived from crosses with cv. Ripening of bunches is delayed. Raspberry bushy dwarf virus. However. Campbell Early are suscetible as well as cvs Takao. representing the most important virus disease in Yamanashi Prefecture. being transmitted by the eryophid mite Colomerus vitis. the 3' terminal region of which (2469 nts) has been sequenced.2 kb in size).g. if any. Grapevine berry inner necrosis has been reported only from Japan. berries are small and show external discolorations and internal necrosis. 24% of the particle weight and consisting of two functional species RNA-1 with Mol. Healthy vines of cvs Kyoho and Pione became naturally infected in the field within one year from planting. Japanese diseases GRAPEVINE BERRY INNER NECROSIS 1.. delayed bud break and young shoots with short internodes and internal browning. GINV has filamentous particles about 750 nm in length and a single-stranded RNA genome with Mol. and RT-PCR. RBDV. M. a mechanically transmissible definitive member of the genus Trichovirus. Agent: The disease agent is Grapevine berry inner necrosis virus (GINV). Description Main synonyms: None Main symptoms: Infected grapevines have low vigor. 20 Murant A. 1976. wt of 2 x 106 Da (5. Delaware. Control : No information 2. and Kaiji are infected latently. 165 B. Extended Abstracts 14th Meeting of IGVG. and Pione whereas cvs. The vay of natural spreading in grapevine.Agent: Raspberry bushy dwarf virus (RBDV) was isolated from symptomatic vines. F. Some rootstocks (e. ELISA .8 x 106 Da (2. CMI/AAB Description of Plant Viruses. Chlorotic mottling. Kyoho. Historical review 1976 2003 Murant: Description of RBDV.5 x 106 Da. Transmission: In raspberry the virus infects progeny seedlings and pollinated plants through pollen. Varietal susceptibility: No information Detection: Mechanical transmission to herbaceous hosts.. infected propagative material can disseminate the RBDV. Varietal susceptibility: Symptom severity varies with the cultivar. and a bipartite singlestranded RNA genome accounting for c. No. Koshu. the type species of the genus Idaeovirus is a pollen and seed-borne virus with quasi spherical particles made up of a sigle type of coat protein subunits (Mr c. 113 . Virscek Marn and I. 30 x 103). rings and line patterns are shown by leaf blades. Raspberry bushy dwarf virus infection of grapevine in Slovenia. Zezlina. wt of 7. wt 0. Vitis riparia Gloire) are also susceptible.
Annals of the Phytopathological Society of Japan 58. 1992. Bulletin of Yamanashi Fruit Tree Experimental Station 10. A new virus disease. Yoshikawa et al. Kunigi Y. Nishijima T..: First account of grapevine berry inner necrosis disease in a non Japanese publication. H.. 77-78 Yanase H. Y. 114 . S. Studies on the grapevine berry innner necrosis virus disease. Disease re-named Grapevine berry inner necrosis Terai et al. 1993. 1. 57-63. Back-transmission of a grapevine filamentous virus to grapevine seedlings and induction of foliar and berry symptoms in grapevine. 617.. Purification of a filamentous virus isolated from grapevine berry inner necrosis and foliar mosaic. T. Kunugi. 2000. H. Yanase. Induction of berry necrosis in Kyoho back-inoculated with the virus isolate from grapevine mosaic diseased clones and renaming to grapevine berry inner necrosis. Studies on the grapevine berry innner necrosis virus disease.. Y. Magome. Terai and Y. Montreux 1993. Archives of Virology 142. Shinkai 2000. Tanaka H. Terai and A. Annals of the Phytopathological Society of Japan 55. Kyoho back inoculated with the filamentous virus isolated from mosaic-diseased grapevines. Mosaic symptoms on cv Kyoho. Kyoho in Japan Yanase: Purification of a filamentous virus isolated from mosaic-diseased grapevines Yanase and Terai: Induction of mosaic symptoms in grapevines inoculated with the filamentous virus Terai and Yanase: Induction of berry internal necrosis in cv. Bulletin of Yamanashi Fruit Tree Experimental Station 10. S. 536 Terai Y. Grapevine berry inner necrosis. Takahashi and Y. 2. Historical review 1984 1985 1987 1992 Tanaka: Description of a mosaic disease in cv. 1985. 423. Symptoms on vines. Extended Abstracts 11th Meeting of ICVG.. a new trichovirus: comparative studies with several known trichoviruses. Description Main synonyms: None. Y.. Asari. 2. Yanase H. and Yanase H. Annals of the Phytopathological Society of Japan 51.Detection: Indexing on cvs Kyoho or Pione. 1351-1363 GRAPEVINE STUNT 1. Iida. Terai. and Terai Y. 1987. 47-56. 1997. 362. 1984.: Experimental evidence that GINV is transmitted by the the grape erineum mite Colomerus vitis 1993 1997 2000 2000 3. varietal susceptibility and natural spread. Goto.: An account of the varietal susceptibility to the disease and natural field spread Kunigi et al. GINV is mechanically transmissible to herbaceous hosts and can be identified by ELISA and moleculat techniques in infected vines.. and H.. Terai.: Partial sequencing of GINV genome and assignement of the virus in the genus Trichovirus Nishijima et al. Transmission of grapevine berry inner necrosis virus by the grape erineum mite Colomerus vitis in Yamanashi. Control: Use of tolerant cultivars in areas where the disease spreads epidemically. Yoshikawa N. References Kunugi Y. grapevine berry inner necrosis with natural spread in Japan.Annals of the Phytopathological Society of Japan 53.
1981. Description Main synonyms: none. leaves are small. S. 137 Namba S. Food and Fertilizer Technology Center for the Asian and Pacific Region. Main symptoms: No appreciable symptoms are visible on the foliage of cv. The berries. 1-17. Namba. with scorched margins. Yora.: A small isometric virus associated with stunt disease in Japan. Yamashita. Hatamoto et al. 2. Transmission: The disease is transmitted by the leafhopper Arboridia apicalis. Historical review. References Hatamoto M. S. Annals of the Phytopathological Society of Japan 50. which makes the crop unmarketable. Inflorescences are undersized. sometimes.. 1982. Koshu nor any apparent reduction of vigour and yield. 1986.Main symptoms: Spring vegetation is delayed. p. 1984. M. Iwanami. Doi. S. Doi. Doi and M. Graft transmissibility of grapevine stunt disease. summer vegetation is apparently normal. Because of heat recovery. are pale-coloured and have a low sugar content.: Evidence that the disease is transmitted by the leafhopper Arboridia apicalis. however. 396 Hatamoto M. S. GRAPEVINE AJINASHIKA DISEASE 1. 3. Agent: An isometric. Transmission of grapevine stunt disease by the grapevine leafhopper Arboridia apicalis Nawa. Control: Use of disease-free material obtained through heat therapy. American rootstocks are infected without showing symptoms.: Purification and characterization of the virus associated with stunt disease. The disease has only been reported from Japan. FFTC Book series. No. A small spherical virus associated with grapevine stunt disease. Fujii.. The disease has only been reported from Japan. vinifera cv. Hatamoto et al. T. Detection: Grafting to Campbell Early and ELISA using extracts from infected vine tissues. non mechanically transmissible virus about 25 nm in diameter is consistently associated with diseased vines and regarded as the possible causal agent. internodes are short. Campbell Early. Annals of the Phytopathological Society of Japan 47. Yamashita and Y. S. 1986 Namba et al. S. This condition gives the name to the disease which in Japanese means "unpalatable fruits with low sugar content". 33. Y.109-126. Fujii.: Successful graft-transmission of stunt disease. Taipei. curled and. The disease is apparently restricted to the V. 115 . Namba. Spread occurs also through infected propagative material. Yamashita.. Y. Hatamoto. 85 Namba S. Doi and K. Taiwan (Reference 2951 in the Review of Plant Pathology 66. 1987). fruit setting is impaired and bunches are few and shelled. 316. 1981 1982 1984 Namba et al. Annals of the Phytopathological Society of Japan 48. phloem-limited. This virus is serologically distinct from the putative agent of ajinashika disease. Varietal susceptibility: No information. Evidence that it is not related to the presumed agent of ajinashika disease. M. Three phloem-limited viruses of grapevine: direct fluorescence detection. Yamashita and Y.. The same paper appears in Taiwan Food and Fertilizer Technology Center Technical Bulletin 92. In Plant Virus Diseases of Horticultural Crops in the Tropics and Subtropics.
12491253. Doi and K. 1979 1980 1986 1991 1991 Namba et al. S. Purification and properties of spherical virus particles associated with grapevine ajinashika disease. and R. Boscia. Historical review. Terai and Yano: Description of ajinashika disease and suggestion that it is caused by the concomitant infection of leafroll and fleck. vinifera cv. 70-73. A small spherical virus associated with the ajinashika disease of Koshu grapevine. Three phloem-limited viruses of grapevine: direct fluorescence detection. Ajinashika disease of the grapevine cultivar Koshu in Japan. consistently found in infected vines.: Further characterization of the isometric virus and claim that it is the putative agent of the disease. Hatamoto. Proceedings 10th Meeting of ICVG. 1979.Agent: The disease was reported to be caused by the concurrent infection of leafroll and fleck. Yora.. No relationship found with fleck. non mechanically transmissible virus about 25 nm in diameter. Yamashita. 1986. The disease seems to be restricted to V. Namba et al. However. Volos 1990. Namba et al. Detection: Graft transmission to cv. Niagara Falls 1980. Plant Disease 75. 116 . 1980.: Partial characterization of the isometric virus associated with the disease and its detection by ELISA in infected vines. Koshu and ELISA using extracts from infected vine tissues. Transmission: No vector is known. Terai Y. 1991. Y. Dissemination is through infected propagative material. 1991. Iwanami. Y. D. Control: Use of disease-free material obtained through heat therapy. Yamashita. S. 67-70. 2. Annals of the Phytopathological Society of Japan 45. 1-17. was suggested as the possible causal agent. and D. phloem-limited. T. Gonsalves. Yano. Koshu. Varietal susceptibility: No information. Terai Y. an isometric. 15-19. 3. References Namba S.. Proceedings 7th Meeting of ICVG. Namba S.: First mention of ajinashika disease and report of the association with it of a non mechanically transmissible virus with isometric particles.. S. Ajinashika disease: a combined effect of grapevine leafroll and grapevine fleck viruses on sugar content in the Japanese grape cultivar Koshu. Tsuchizaki. Taiwan Food and Fertilizer Technology Center Technical Bulletin 92. Doi and M. T. Yamashita. Namba S. Terai: Additional report on ajinashika disease as derived from the combined effect of leafroll and fleck.
VIRUS-LIKE DISEASES .
The disease has been reported from many European and extra-European countries Agent: The etiology of enation disease is not yet clear. omeoplasie crestiformi (Ital. North (Tunisia) and South Africa. Enations develop mostly on the underside of the leaves at the base of the shoots.). and the frequent occurrence of fanleaf virus in enation bearing vines supported the hypothesis that enation disease could be due to a severe strain of fanleaf virus. Detection: By observing the symptoms on the leaves and by indexing on LN 33 hybrid. Hewitt: Description of symptoms in California.VIRUS-LIKE DISEASES Several latent or semi-latent grapevine diseases are known. growth tends to become normal again. with a fanlike aspect and abnormal indentation. Basal leaves. the absence of symptoms does not necessarily mean that the plant is healthy. Australia and New Zealand. The varieties Panse Precoce. Description Enation disease of grapevine is one of the oldest known disorders of European grapes. with drawings of symptoms. Gigante : Research on histological and cytological aspects of enations. Control: Use of healthy material. The disease is perpetuated by vegetative propagation. However.). according to the cultivar) and is of poor quality. Later in the year. North America (California). but some are heat-labile and can be eliminated by heat therapy. Severely affected leaves drop prematurely. its description dating back to the late 1800s Main synonyms: Enationenkrankheit der Rebe (Germ. Symptom expression varies year by year. which gives a bushy aspect to the plant. apparently in relation with climatic conditions. They are outgrowths 2-3 mm high and 3-5 mm long or more. Their agents are still unknown. Grenache and Tokay appear to be quite susceptible and develop severe symptoms when infected. however. Riesling. ENATION DISEASE 1. The basal internodes are short. 119 . and is probably due to a virus-like agent. Varietal susceptibility and sensitivity: There is little information available. maladie des énations (Fr. as symptom expression is variable in successive years and graft transmission not 100 % successful. and often show longitudinal cracks between the nodes. All persist in propagative material and are transmitted by grafting. Primus. with prominent veins. some of which have a clear-cut detrimental effect on the crop. are often mis-shapen. The transmission by graft is rather erratic. Italia. Latin America (Venezuela). Main symptoms: Affected vines show a delayed opening of the buds and a slow growth of the shoots in the spring. 2. Leaves developed later in the season are usually normal. Historical review 1891 1937 1954 Buchenau: First description of enation disease of grapevine occurring in Germany. They are often thicker than normal. The infectious agent of the disease perennates in propagating material. vinifera varieties in Europe. The crop can be drastically reduced (up to about 50%.). has now been dismissed Transmission: By vegetative propagation. but attempts to transmit it by graft or mechanical inoculation gave no results. This hypothesis. malattia delle enazioni. There is no information on the possibility of curing this disease by heat treatment or meristem culture. which run more or less parallel to the main veins. Graft transmission suggests that it is a virus disease. whether they bear enations or not. irregular and mis-shapen. Symptoms have been observed on many V.
Padilla et al. Trebbiano romagnolo and reduces the yield from 13% to 23% according to the severità of symptom expression.. In the vineyards under observation. Weinberg und Keller 15.: Detailed description of macroscopic and microscopic symptoms of enation disease. Malvasia (10. The mean yield loss of diseased vines ranged from 17. Enation disease in Spain Chabbouh and Savino: Enation disease in Tunisia 1966 1966 1968 1970 1970 1971 1973 1975 1978 1979 1980 1980 1981 1983 1989 1996 1997 1997 3. Martelli et al. Presence of enations in Razaki grapevine in Crete (Greece). Nieder: Enation disease in Austria Garau et al. lowest in cv. but diseased vines which had not shown enation symptoms for several years had almost normal yields. There is some evidence that enation can be carried in the rootstocks. Refatti: Hypothesis of a correlation between fanleaf and enation disease.5%). Phytopathologia Mediterranea 17. References Avgelis A. Xafis. 1978. attempts to transmit the disease by graft. Brückbauer: Description of symptoms of enation in Germany and confirmation of graft transmission of its agent. 195 Brückbauer H.: More data on the effects of enation on the yield of cv. the proportion of diseased vines was highest in cv. historical account. Italia in Sardinia. Graniti and Martelli: Review paper on enation. and possibly caused by a virus. although the rate of symptom expression does not exceed 30% Credi: Enation diseasae affects the vegetative vigour of cv. The possible role of fanleaf virus in the etiology of enation requires further investigation.: Enation disease in France Pozdena et al. but report on observations made in Australia where no fanleaf virus could be recovered from enation-affected vines. only fanleaf virus was recovered.4 to 48. . Marinesku and Bondarchuk: Enation disease in Moldova Brückbauer: Influence of enation disease on growth and yield of grapevine in West Germany. 1968.: Enation disease in Turkey Hevin et al. McGechan: Enation disease in Australia Tekinel et al. Prota et al. Confirmation of graft transmissibility of the disease. The authors conclude that the disease is probably of European origin. Beobachtungen und Untersuchungen über die Enationenkrankheit der Rebe. with negative results. 120 . Vernaccina (1. Enationaffected vines produced less than 50 % of the yield of healthy plants.: In experiments on graft transmission of enation disease aimed at determining the best indicator.: Enation disease in Czechoslovakia Avgelis and Xafis: Enation disease in Greece Prota and Garau: Enation disease found in Sardinia. The authors discuss the hypothesis that enation is caused by a strain of fanleaf virus.: Successful transmission of enation disease from diseased to healthy grapevine by graft strongly supports the hypothesis of a viral origin. Mechanical transmission tests were also negative. the hybrid LN 33 was found to be the most sensitive and reliable indicator variety.5%). and C.3% .1966 Graniti et al. 79-112.
Main synonyms: Mosaïque des nervures (Fr.D. 1965.. Savino. and V. Effetto della malattia delle enazioni della vite sulla produzione e sullo sviluppo vegetativo della cv Trebbiano romagnolo. Pozdena J. Ita and F. Enations. Abnorme Blattbildungen.. F. In the most sensitive varieties. 293-306.. Die Eantionenkrankheit der rebe erstmal auch in Osterreich nachgewiesen. Marinesku V. Trasmissione per innesto della "malattia delle enazioni" della vite. 207-217. Adernmosaik (Germ.. Quacquarelli.Brückbauer H. (ed. Lisbon 1997. Martelli and F. O. Martelli G. Lamberti. Enations of grapevine in Sardinia. Rivista di Patologia Vegetale. 179-189. Phytopathologia Mediterranea 5. and G. Enation disease of grapevine in Italy. Tekinel N. G. 1966. 1997. Kiryat Anavim. Cordoba. U. 1937. Credi R. 1970. 225-246. 7-12. Studies on reproduction of enation symptoms by grafting in Sardinia. Important virus diseases of grapevine in New South Wales. Rivista di Patologia Vegetale S. Nieder G..P. 1976.1989. 1987. Moldavian Research Institute of Horticulture Kishinev. 241-243. G. Enation symptoms found in France. Vein mosaic appears to be of low economic importance. 17. Z. A similar disease has been reported in Australia under the name of summer mottle. Vanek. Garau R. 59-64. but these necroses do not affect the veins as it is the case with vein necrosis. Symptom expression seems to depend on climatic conditions. Martelli. Main symptoms: Pale green mosaic affecting mostly the tissues adjacent to the main veins or the smaller ones. A study of infectious degeneration (fanleaf) in vineyards of the Mediterranean region. Lisbon 1997.). Proceedings 9th Meeting of ICVG. Padilla V.G.. Lamberti and A. 169-192. producing often a vein banding effect. University of California. Proceedings International Conference on Virus and Vector on Perennial Hosts.W. Salih and Y. VEIN MOSAIC 1. Graniti A. Spain. 1983. Phytopathologia Mediterranea 20. 349-352. Graniti A. Gigante R. 1966. The research of the virus diseases of grapevine in CSSR. 1966.P.. Bolletino della Regia Stazione di Patologia Vegetale. Prota U.P.. it was clear that vein mosaic was not caused by fanleaf virus. 47-64. Gazeau...). 1973.P. In: Verderevskaya T. Proceedings 6th Meeting of ICVG.. Benayas. Bericht der deutschen botanischen Gesellschaft 9. Bitki Koruma Buletin 11. Prota U. Pflanzenharzt 36. 145-152. Berkeley. Petria 6.) Virus and mycoplasma-like diseases of fruit trees. n. Deutsches Weinbau-Jahrbuch 31.). Davis. Su una possibile correlazione fra il virus del complesso dell'arricciamento e la malattia delle enazioni della vite. Ricerche istologiche sulle omeoplasie crestiformi (Enations) delle foglie di vite affette da rachitismo. Nas. Garau R. And G. areas of the leaf blade may become necrotic. 251-252 Hewitt W. Bondarchuk. Studies on some variable characters of grapevines affected by enation disease in Sardinia..IV. 1983. Garau. Rives.S. Bulletin of the California Department of Agriculture 43. 326332. Salcan.s. (N. but when fanleaf virus transmission to herbaceous hosts became possible. 1975. Dolar. 1891. Israel. Garcia. A. 1970. 1971. McGechan J. 2. H. 1981.. Grapevine enation disease in Murcia (Spain). 121 . Chabbouh N and V. Ser. This disease is widespread and probably worldwide. Prota and M. 47. 1996. with Special Reference to Vitis. Facultet Landbouwwetenschap (Gent) 40. 1997. Frazier N. Roma.). 823-827. B.V. Extended Abstracts 12th Meeting of ICVG. I. Buchenau F. 122-124. Description The symptoms of vein mosaic have been confused for some time with those of fanleaf/yellow mosaic. IV. Cugusi. 97-98.W. Occurence of grapevine enations in the Moldavian SSR. Hevin M. and R. and M. 9.B. Some virus and virus-like diseases of grapevines.K. Agricultural Gazette of New South Wales 81. 203-206. Graniti. Monografias INIA 18. 1954. small fruit and grapevine in Moldavia. Occurrence of enation disease in Tunisia. Leclair and M. Mosaico delle nervature (Ital. Extended Abstracts 12th Meeting of ICVG. 1979. Division of Agricultural Sciences. California. Mededel. Cugusi.. 46-51. M. In: Virus Diseases of Small Fruits and Grapevines (A Handbook). Refatti E. Einfluss von Virusinfektionen auf Wachstum und Ertrag der Rebe. 1980. 48.
but this hypothesis has not been confirmed. Muscat de Hambourg. No vector known. Legin and Vuittenez: Description of vein mosaic. Canova. Control: Use of indexed material. Marinesku and Bondarchuk: Vein mosaic in Moldova. Samonina et al.R. No claim is made that these putative viruses are connected with the disease. Ehrenfelser showing symptoms of vein mosaic. Milkus et al. Detection: Indexing with V.. Occurrence of grapevine vein necrosis and grapevine vein mosaic in the Emilia-Romagna region (northern Italy). Pinot.A. Several V. Woodham and Krake: Comparison of summer mottle and vein mosaic. in the absence of any detectable virus.: Observation of a type of mosaic of grapevines which appears to be independent of fanleaf virus. 148-152.R. 2. Bonfiglioli: Vein mosaic in New Zealand (unpublished). 17-23. vinifera cvs. Viognier.: Vein mosaic in URSS. and R. Golino: Vein mosaic in California. Kuniyuki: Vein mosaic in Brazil. Alphonse Lavallée. Golino D. 1979 1980 1982 1983 1985 1993 2004 3. show symptoms (Syrah. Servant. Potential interaction between rootstocks and grapevine latent viruses. 1978. Vuittenez and Stocky: Electron microscope study of thin-sectioned tissues of leaves from Vitis riparia and Vitis vinifera cv. Woodham. 266-270. Pop: Vein mosaic in Romania. References Credi R. vein mosaic and vein necrosis. LN 33 is also sensitive. Abracheva: Vein mosaic in Bulgaria. American Journal of Enology and Viticulture 44. 1993. Transmission: By graft and vegetative propagation. 122 .Agent: Unknown. Comparison of symptoms of fleck. Vitis 17. Mycoplasma-like organisms were supposed to be the cause of vein mosaic. Varietal susceptibility and sensitivity: Vitis riparia Gloire de Montpellier is especially sensitive and is used as indicator. Symptoms are very similar to those of vein mosaic in Europe. A number of cytological modifications primarily involving chloroplasts were observed along with the presence of bundles of filamentous structures resembling closterovirus particles. Saric and Hranuelli: Vein mosaic in Croatia. riparia Gloire de Montpellier. Pearl of Csaba). Babini and A. Krake and Woodham: Description in Australia of a systemic mottling syndrome which is expressed during summer on the leaves of some varieties. Phytopathologia Mediterranea 24. Grapevine summer mottle: a new graft-transmissible disease. Chardonnay. Krake L. A. Chasselas. Historical review 1966 1973 1973 1976 1973 1973 1978 Vuittenez et al.C.: Vein mosaic in Ukraine.. and Gamay apparently show little or no symptoms. 1985. The disease can be eliminated by heat therapy.
Krylova. Control: Use of disease-free propagating material obtained by culture of fragmented shoot apices. Description Main synonyms: None. producing a feathering or banding effect. IV. Niagara Falls 1980. Vinogradr.. rupestris and LN33 are infected symptomlessly. probablement indépendante du virus du Court-noué. Annales des Epiphyties 17. 68-72. 247-252.C. Woodham and Krake: Comparative graft transmission trials demonstrate that summer mottle 123 . Evidence that the disease is graft-transmissible. Numéro hors série. Grapevine vein mosaic. de la marbrure de V. Observations sur une Mosaïque de la Vigne. Agent: Unknown. 9 . 1973. resembles in some respects the European vein mosaic and the Greek Roditis leaf discolouration (see under Minor virus diseases). Proceedings 7th Meeting of IGCV. 1966. 243-250 Samonina I. whereas the opposite occurs with summer mottle. and V. Spread is through infected propagative material but is has also been observed between adjacent vines Varietal susceptibility: No grapevine tested was immune to infection. Investiation on grapevine viruses in the SR Croatia. Rivista di Patologia Vegetale . SUMMER MOTTLE Summer mottle.V. 67-73.. Adverse effect of light and of high temperature on symptom expression of grapevine vein mosaic in Sao Paulo. URSS. an Australian disease. Barlass et al. maladie à virus de la vigne. Stocky. Bondarchuk. Saric A. 1973.R.Legin R.V. Roditis leaf discolouration and summer mottle have similarities suggesting that they may be the same disease. several European grape cultivars show symptoms. 191-204. La mosaique des nervures. e. and A. A comparison of grapevine summer mottle and vein mosaic diseases. 1. 9. Vitis 22. R. 2. 1982. Vuittenez.-Berl.g.N.V. ou la “feuille rouge”. Kuszala. Moldavii 31. Milkus. Vinodel. poorly developed and with small berries..G. Krylov and V. Symptoms show on vegetative growth that develops at temperatures in excess of 30 °C. Pop I.V. 41-42. Proceedings of the Conference on Excoriosis and Virus Diseases of Grapevine. 1977. 1973. Main symptoms: Pale green to yellowish dicolourations of the tissues adjacent to the main or secondary veins. Mataro.N. 57-63. Mostar 1977. Vuittenez A. Detection: Graft transmission to a number of cvs. Legin and J. IV . 1983. Summa Phytopathologica 11. Comparaison des symptômes et transmission par greffage d'une mosaïque nervaire de Vitis vinifera . Ultrstructure de vignes infecteés par deux maladies de type viral: la mosaique des nervures. A.. Transmission: No vector is known. and G. and Hranuelli T.. 110 R. Vuittenez A. suspected to be a virus or a viroid.: Elimination of the disease agent by culturing fragmented shoot apices. V. Rivista di Patologia Vegetale. 1976. B. Bunches of infected cvs Sideritis and Cabernet sauvignon are fewer. Krake. 149-141. Historical review 1978 1982 1983 Krake and Woodham: Description of summer mottle in Australia. However. 48-49 Marinesku V. Cabernet franc. rupestris et d'une affection nécrotique des nervures de l'hybride Rup. Symptoms of vein mosaic develop under mild weather conditions and fade during hot weather. and L. These symptoms appear in summer and persist through the autumn. Ser. Mission. S. Rivista di Patologia Vegetale S IV 9. Cabernet sauvignon. A grapevine virus disease in the Primorye territory. Woodham R. 1985. Kuniyuki H..
necrosi delle nervature (Ital. 266-270. rupestris x V. A comparison of grapevine summer mottle and vein mosaic diseases. Regeneration of virus-free grapevines using in vitro apical culture. most likely GRSPaV.M. Adernnekrose (Germ. Australia.).: Vein necrosis in Italy and Bulgaria 124 . and some infected plants may die. References Barlass M. 247-252. varieties or hybrids. Woodham. Cause-effect relationships between MLOs and the disease has never been ascertained.C. Krake L. Varietal susceptibility and sensitivity: The rootstock 110 R is most sensitive. K. Skene. Mycoplasma-like organisms have been observed in the phloem of symptomatic vines. Woodham R. RT-PCR with virus specific primers and Western blot with an antiserum to recombinant coat protein of GRSPaV allow sensitive and reliable detection of this virus in symptomatic 110 R plants. at first on the leaves at the base of the shoots. Martelli et al. 1999 Krake et al. Milkus and Kalashyan: Mycoplasma-like organisms found in phloem tissues of vines with vein necrosis. Taylor. growth is much reduced and necrosis of the leaf veins appears.: Suggestion that summer mottle and Roditis leaf discolouration are the same disease 3. Woodham and L. 1982. Control: Use of indexed planting material.. R. Main symptoms: On the rootstock 110 R. Description Vein necrosis has probably a worldwide distribution. VEIN NECROSIS Although vein necrosis is currently regarded as a disease in its own right. So far. its economic importance has not been assessed. Krake. Krake L.H. 70-74. Historical review 1973 1978 1978 Legin and Vuittenez: Discovery and description of this virus-like disease while searching for indicators for fleck.A.R.differs from vein mosaic.. Rezaian and R. Transmission: By grafting and vegetative propagation.S. and the only Vitis species that is clearly affected is the rootstock 110 R. Main synonyms: Nécrose des nervures (Fr. 1978. Annals of Applied Biology 101. Possible viroidal etiology put forward.R. Many grapevine varieties and rootstocks are infected symptomlessly. and L. Grapevine summer mottle: a new graft-transmissible disease. 2. Vitis 17. Collingwood. 1983. especially under greenhouse conditions. 1.. 291-295. M.C. but their etiological relationship with the disease has not been proven. N. Scott. Vitis 22. Necrotic reactions are best seen on the lower face of the leaf blade. berlandieri 110 Richter (110R) with vein necrosis symptoms. Also the tendrils and many shoots can necrotize.R.R.C. Little is known about sensitivity of other Vitis species. and R. later on younger leaves as they develop. The agent of vein necrosis can be eliminated by heat therapy.G. CSIRO Publishing. No vector known. Graft-transmissible Diseases of Grapevines.). 1999. Krake. Agent: Suspected to be a virus. This strongly supports the hypothesis that vein necrosis is a specificic reaction of the rootstock 110 R to GRSPaV infection.). recently an association exceeding 95% has been experimentally observed between Grapevine rupestris stem pitting-associated virus (GRSPaV) and vines of V. Detection: By grafting on 110 R.
the incidence of vein necrosis in visually selected stocks of table and wine grape varieties is on the average 71 %. 59-65. Abracheva and B. Vuittenez. 1978. V. 9. 1986.N.: In southern Italy. Savino.A. 1994. and J. 1989. 1978.: Vein necrosis in Hungary Gursoy: Vein necrosis in Turkey Rumbos: Vein necrosis in Greese Martelli et al.. Martelli G.A. 43-45 Kuhn G. Pirolo. ser. 1993.Z. and L. Boscia and G.. 301. 79-83 Legin R. Informatore Fitopatologico 28 (10).-Berl. but did not eliminate the disease entirely. 1984. Minafra and G.B. 61 Savino V. Rosciglione. Savino. 1986 1988 1989 1992 1993 1994 2004 3. Rumbos I C. P. Castellano. de la marbrure de V.R. Vein necrosis. La Notte. Martelli.P. 204-207. 58-60. IV. Lehoczky et al.P. Lehozcky J. Izvestiya Akademii nauk Moldav. Grapevine vein necrosis disease detected in rooststocks in Australia. Incidence of some graft-transmissible virus-like diseases of Grapevine in visually selected and heat-treated stocks from Southern Italy. 29-30. Vein necrosis: new virus-like disease in Turkish vineyards. J.P. D. Milkus B. Savino. Necrosi delle nervature della vite in Italia e Bulgaria. Boscia. American Journal of Enology and Viticulture 44. 1988.C. Journal of the Australian Institute of Agricultural Sciences 50. i Chim Nauka 1. rupestris et d'une affection nécrotique des nervures de l'hybride Rup. 1992.A. H. Ser.C. Kalashyan. 606-612. P.. and A. rupestris. 2004. Rivista di Patologia Vegetale.. Farkas.5 %. V. Phytopathologia Mediterranea 24. 1973. Comparaison des symptômes et transmission par greffage d'une mosaïque nervaire de Vitis vinifera.. Heat therapy reduced this proportion to 35. fleck and leafroll in Vitis vinifera and grapevine rootstocks in central Greece. Gursoy Y. and L R. Boscia and V.. Kergazdasag 18 (4).1984 1985 Woodham R. 148-152. Viruses of grapevine in Malta. 1985. Bulletin OEPP/EPPO Bulletin 22. C. Lazar. Biol. Journal of Turkish Phytopathology 17. 110 R. SSR. Mycoplasma-like bodies in phloem tissue of grapevine affected by vein necrosis.. 57-63. Detection of vein necrosis virus (GVNV) in the vines of cultivated grape varieties. References Bouyahia H. Martelli. Fitopatologia Brasileira 19. Phytoparasitica 17. Krake. Galea Souchet.: An association exceeding 95% observed between GRSPaV and 110R vines showing vein necrosis symptoms in indexing trials. 3-5 Martelli G. No veing necrosis observed in 110R top grafted on GRSPaV-free V. Woodham R. Golino D. D. Vein necrosis a disease that is latent in most grapevine cultivars of the State of Rio grande do Sul. 125 .: Vein necrosis in Malta Golino: Vein necrosis in California Khun: Vein necrosis in Brazil Bouyahia et al. Krake: Vein necrosis in Australia Savino et al. Suggestion than vein necrosis is a specificic reaction of 110R to GRAPaV. A. Is Grapevine vein necrosis a reaction to Grapevine rupestris stem pitting-associated virus? Journal of Plant Pathology 86.. Potential interaction between rootstocks and grapevine latent viruses. G.P. D.. M.
VIROID (Yellow speckle) .
plant age. and perhaps the type of infecting viroidal sequence variant. all of which belong in the family Pospiviroideae: Grapevine yellow speckle viroid 1 (GYSVd-1). AGVd has been reported from Australia. Semancik (eds. Kyoto vine with yellow speckle symptoms. and AGVd. Only GYSVd-1 and GYSVd-2 are pathogenic. 129 . picchiettatura gialla (Ital. climatic conditions. Europe. Two families are known. It was isolated in Australia from a grapevine that contained also other viroids and was distinguished from these because it replicated in cucumber and tomato. north and south America. phylogenetic analysis of hop and grapevine isolates of HSVd has provided evidence that the viroid that causes hop stunt disease in Japan is a variant of HSVd-g. The suggestion is that HSVd moved from grapevine to hop probably 50-60 ago in the Nagano and/or Fukushima prefectures in which it is not uncommon to find hop gardens next to vineyards. Hop stunt viroid grapevine strain (HSVd-g). Very often. has a genome 369 nt in size. AGVd. 2003.). Flores. vein banding-like symptoms can be observed in vines infected only by yellow speckle viroids. It was first detected in Japan and transmitted to cucumber and grapevine seedlings in which. Randles and J. Sometimes. was demonstrated to be caused by a co-infection by yellow speckle viroids and GFLV. Interestingly. GYSVd 1 and GYSVd 2 are made up of 366 and 363 nucleotides (nt). Neither of them is able to replicate in herbaceous hosts but both were succesfully inoculated to grapevine seedlings reproducing the yellow speckle syndrome. Grapevine yellow speckle viroid 2 (GYSVd-2). HSVd-g. They are made up of a non encapsidated circular RNA of 246-375 nucleotides.. These symptoms appear in the height of summer on a limited number of mature leaves and persist for the rest of the vegetative season. it did not induce symptoms. Cabernet franc and a cv. a disease characterized by chrome yellow flecks localized along the main veins of mature leaves and progressing into the interveinal areas. CSIRO Publishing. are not associated to any specific symptomatology. 370 pp. respectively from a cv. genera and species.). Description Main synonyms: Moucheture jaune (Fr. R. the non coding genomes. Pospiviroideae and Avsunviroideae whose significant discriminating traits are the presence of a central conserved region in the secondary structure and nuclear replication (Pospiviroideae) or a branched secondary structure lacking the central conserved region. a size much smaller than that the smallest viral genome. however. viroids are classified in families.VIROIDS Viroids. YELLOW SPECKLE 1. Citrus exocortis viroid grapevine strain (CEVd-g). Main symptoms: Few to many minute chrome yellow spots or flecks scattered over the leaf surface. The symptomatology varies depending on the cultivar. HSVd-g. HSVd-g has been recorded from Australia. Vein banding. presence of ribozymes. inducing a disease called yellow speckle. CEVd-g. Australian grapevine viroid (AGVd). References Hadidi A. the USA. thought to be elicited by a specific strain of GFLV.). the type species of the genus Hostuviroid. Gelbsprenkelung der Rebe (Germ. Collingwood.S.). GYSVd-1 and GYSVd -2 cause the disease individually or in combination. GYSVd-1 and GYSV-2 have a worldwide distribution The three additional viroids that have been detected in grapevines. a member of the genus Apscaviroid. J. and plastidial replication (Avsunviroideae ). and Tunisia. Like viruses. Viroids. and may have a worldwide distribution. or gathering along the main veins to give a vein banding pattern. respectively and both belong in the genus Apscaviroid. Agents: Two distinct viroids. infected vines are symptomless or show symptoms erratically. Five grapevine-infecting viroids are known. Both these viroids wer first isolated in Australia.W. has a genome 297 nt in size. are subviral pathogerns endowed with autonomous replication in their hosts. however.
: Reconstruction of the secondary structure of CEVd-g Szychowski et al. yellow speckle and fleck through dodder. CEVd-g and AGVd. In the great majority of grapevine germplasm infection is latent.CEVd-g. Control: Use of viroid-free propagative material obtained by meristem tip culture.: Evidence that viroids are widespread in grapevines. GYSVd-2. Abracheva et al.: A disease of cv. found in grapevine accessions from Europe and California.: Yellow speckle eliminated by in vitro apical culture. one of which identified as the agent of citrus exocortis. Mink and Parsons: Yellow speckle can be detected by growing vines for 2-3 weeks at 32 °C under continuous illumination. First identification of AGVd Koltunow and Rezaian: Identification and sequencing of grapevine yellow speckle viroid. Shikata et al. Rezaian et al. Krake and Woodham: Evidence that the agent of yellow speckle is implicated in the etiology of vein banding. Barlass et al.: Four viroids found in Australian grapevines. Varietal susceptibility: All Vitis species. Although CEVd is present in most. Polyacrylamide gel electrophoresis has been used extensively before the advent of nucleic acid-based assays (hybridization and RT-PCR) which constitute far better detection and identification tools. has a genome 369 nt in size. Natural dissemination takes place by mechanical inoculation through surface-contaminated cutting tools during management operations. Woodham and Krake: Artificial transmission of grapevine leafroll. Semancik et al. It was first recoverd in Spain from symptomless grapevines. and distribution of infected propagating material. Seed transmission has been demostrated for GYSVd-1. as the disease may have spread naturally. a disease formerly thought to be caused by a chromogenic strain of GFLV. Historical review 1972 1975 1978 1982 1982 1983 1983 Taylor and Woodham: First description of yellow speckle as a graft transmissible disease separate from chromogenic disorders induced by grapevine fanleaf virus (GFLV). Sano et al. Cannonau not associated with the presence of GFLV reported from Italy. This latter way of dissemination has been considered as more efficient and frequent than mechanical transmission. 1984 1985 1985 1985 1987 1987 1988 1988 1988 130 . Three different viroids found in a number of accessions in a Californian varietal collection.: A vein banding condition of cv. grafting. its grapevine strain so far has only been recorded from Australia and the USA.: The Japanese grapevine viroid identified as a strain of hop stunt viroid. Experimental transmission through dodder is possible. Woodham and Krake: Evidence of field spread of yellow speckle. 2. Transmission: No vector is known. Flores et al. For yellow speckle. Detection: Some viroids can be transmitted mechanically to herbaceous hosts but this is not an efficient detection method. Garcia Arenal et al. besides Spain. Prota et al.: First recovery of a viroid from grapevines in Japan.: Successful mechanical transmission of viroids to grapevines. a member of the genus Pospiviroid. Rcatzitelli resembling yellow speckle reported from Bulgaria.: Two new viroids. if not all citrus-growing countries. hybrids and cultivars appear to be susceptible. the authors consider the results as inconclusive.
. Semancik. 1982. Koltunow et al.: Review of viroids.: A survey of viroids of grapevine in Italy. American Journal of Enology and Viticulture 39.S. GYSVd-1 and GYSVd-2 Rezaian: Complete nucleotide sequencing of AGVd. N. 131 . Cordoba. Proceedings of the 6th Meeting of ICVG..: Structural analysis reveals that five distinct viroids infect commercial grapevine varieties. Detection of viroid and viroid-like RNAs from grapevine. Barlass M. 1976. Marrakchi.P. Annals of Applied Biology 101. 291-295. Fakhfakh.. Koltunow and Rezaian: Description and sequencing of grapevine viroid 1B (later renamed Grapevine yellow speckle viroid 2). References Abracheva P. Grapevine yellow speckle viroid 1 (GYSVd 1). Little and Rezaian: Updated review of grapevine viroids.: Improvement of meristem tip culture technique for the production of viroid-free grapevines. K. 1991 Semancik and Szychowski: There are two classes of grapevine viroids: (i) apparent viroids. Rezaian et al. Quacquarelli and V. R. Pallas and J.P. CEVd-g and AGVd via grape seeds. Skene. Minafra et al. Duran-Vila N. Savino. These viroids. Sano et al. Flores et al.R. GYSVd-2. Elleuch A. V. Monografias INIA 18. are to be named as follows: Hop stunt viroid grapevine strain (HSVd-g).: First report of AGVd in the Mediterranean. A possible virus disease of Rcatzitelli vines in Bulgaria. Arregui. 2003. Martelli. Wan and Symons: Transmission of GYSVd-1. Martelli: Brief review of grapevine viroid situation supporting the idea that vein banding is primarily induced by viroidal rather than GFLV infection. Flores R. Krake. Regeneration of virus-free grapevines using in vitro apical culture. 2095-2102. Production of viroid-free grapevines by shoot tip culture. which can readily be isolated directly from grapevines.b Szychowski et al. Juarez and J. 1991 1996 1997 1999 2001 2003 2003 3. 1988. Woodham and L. Elleuch et al.G.: Suggestion that the viroid causing stunting in hop (HSVd) originated from grapevines. G.: Evidence that two related viroids (GYSVd 1 and GYSVd 2) can cause yellow speckle disease independently. Perreault and H. (ii) enhanced viroids. J.1988 1989 1989 1989 1990 1990 Duran-Vila et al.: First record of grapevine viroids in China. M. Citrus exocortis viroid grapevine strain (CEVd-g).M. Greece. The occurence is reported of HSVd.: Extensive comparative analysis of grapevine accessions from California and Europe reveal a similar pattern of viroid distribution.. 1985. Duran-Vila. J. First report of Australian grapevine viroid from the Mediterranean region. Wang et al.C. based on phylogenetical analysis of hop and grapevine isolates of this viroid. Journal of General Virology 66. 217-220. 45-57.. which require amplification in an alternate host. 127-130. Journal of Plant Pathology 85. Molecular evidence that this viroid originated from recombination between five different viroids among which GYSVd-1 and GYSVd-2 1991a. Spain. A.M. 1978. according to an international agreement reached during the 10th Meeting of ICVG held in 1990 at Volos. Grapevine yellow speckle viroid 2 (GYSVd 2) Australian grapevine viroid (AGVd).
Dore. sanitation and situation in the Arab countries.A.R. and M. Meshi. Krake and K. Greece. 2001. 869-872. R.C. 1985. Rezaian. 270-278.S. 1987. Rezaian M. M. N. Koltunow A.S.R.H. 447-452. Relationships among grapevine viroids from sources maintained in California and Europe. 333338..A. Sano T. 1983. J. American Journal of Enology and Viticulture 38. 1989. J.R. M. Infectious diseases of grapevines. Koltunow. Semancik J. 1990. Ohno and Y. 39-41..P. Two related viroids cause grapevine yellow speckle disease independently.. detection. Parsons. Duran-Vila. J. Rapid indexing procedures for detecting yellow speckle disease in grapevines.M.W. and R. A. 40-50. Leclair.G. Uyeda. CSIRO Publishing. and L. Grapevine yellow speckle agent implicated in the aetiology of vein banding disease. G.A.M.C. Rezaian M. Johnson and M. Credi. Koltunow A. Shikata. Hernadez.A.M. China. P. 1988. 1991. 1990. 1989. Journal of Phytopathology 147.. Plant Disease Reporter 59. 1991a. L.. Credi.Flores R. Semancik.evidence for extensive recombination between viroids. 1990. 53-59. Rivera-Bustamante and A. 1990. Rezaian. Australian grapevine viroid . 1985. 25-36. Rezaian M. Di Serio and C. Woodham R. Journal of General Virology 69. Rezaian. The field investigation of virus and viroid diseases of grapevine and stone fruit trees in Shandong and Liaoning province. A. 193-198. Minafra A.Leclair. Uyeda. 1991b. Martelli and V. F.A. The sequence of a viroid from grapevine closely related to to severe isolates of citrus exocortis viroid. Nature. 195-206. 1984. Hadidi.D. Koltunow A. 1999. Okado.Woodham. 1989. Taylor R. 4203-4210. Comparative properties of viroids of grapevine origin isolated from grapevines and alternate hosts. S. T. 210-219.A. 5587-5598.L. Pallas and R.. An infectious low molecular weight RNA was detected in grapevines by molecular hybridization with hop stunt viroid cDNA. Mink G. 1991. Widespread occurrence of viroid-like RNAs in grapevines. Z. Phytopathologia Mediterranea 24. R. In: Viroids (A.Jiang. 202-205.C. M. L.A. Minafra. Grapevine yellow speckle -. Skene. Mechanical transmission and rootstock reservoirs as factors in the widespread distribution of viroids in grapevines.R. Grapevine yellow speckle disease: studies on natural spread observed in the field.A. Cugusi and M.M. T. 1972. J. Vitis 21.A.A. Martelli G. yellow speckle and fleck diseases by dodder. and L. A. Flores. Wolpert and J.. A. 1987. Journal of General Virology 66. Grapevine viroids..R. Little A. 285-291 Wang G. N. 1988. 65-73.. Nucleic Acids Research 15. Woodham. Vitis 29. Semancik. N. Proceedings of the 10th Meeting of ICVG.A. R. Sano and I. 297. 1983. Ohshima.W. Garau. Semancik (eds. Zhang and X. Flores. 1996.).. R. Greece. and M. Relationship and patterns of distribution among grapevine viroids from California and Europe.S. Doazan. Minafra. Garnier. Mimura and K. 287-288. a new member of the apple scar skin viroid group contains the left terminal region of tomato planta macho viroid. and M. J. Duran-Vila. 213-216. 370 pp. Wolpert and J. E. Prota U. China Fruits 4. 173-182.A.a newly recognized grafttransmissible disease of Vitis. Woodham R. Proceedings of the Japanese Academy of Science 60(B). Grapevine yellow speckle viroid: structural features of a new viroid group.C. and J. S. Vitis 22.C. Savino. Krake L. Volos. Phylogenetic analysis of hop and grapevine isolates of hop stunt viroid supports a grapevine origin for hop stunt disease. Zhang. Nucleic Acids Research 16. Rezaian.P. and R.S. Investigations on transmission of grapevine leafroll. Virology 170.P. American Journal of Enology and Viticulture 39. 132 .S. 849-864. Symons. I. Sano T.. 24-28. 337-345. 1997. Szychowski. Volos.. R. Australian Journal of Agricultural Research 23. R. Collingwood. Seminars in Virology 8. 2003. Goheen. Semancik J. A viroid-like RNA isolated from grapevine has high sequence homology with hop stunt viroid. Garnier. 35-40. 3411-3419. Krake.S.M. Krake. Phytopathologische Zeitschrift 106.. Koltunow and L. 1982. Journal of General Virology 70. 413-422. A. T. Volos. Arab Journal of Plant Protection 7. Semancik. Proceedings of the 10th Meeting of ICVG.. V. Szychowski J. Wan C.A. Garcia Arenal F. Nucleic Acids Research 10. and R. Szychowski J. Greece. 1990. Viroids: the non coding genomes.P. 575-578. Viroids of grapevines in Italy. 1975. Randles and J. Investigations on a vein banding disease of Grapevine in Sardinia.C.. Grapevine viroids.I. Virus Genes 22. Krake. Proceedings 10th Meeting of ICVG.. Vitis 30.P. 1988. Doazan.H. Krake. Szychowski J. Grapevine viroid 1B. Goheen and J. Isolation of three viroids and a circular RNA from grapevines. Shikata E. Hong.R. P. Transmission of viroids via grape seeds. and J..
Nonetheless. the term phytoplasma has become the genus name of these plant pathogenic agents under the provisional taxonomic status Candidatus. climate and infection pressure. The different GYs cannot be differentiated on the basis of symptoms. including roots. their movement is slow and their distribution in the plant is erratic. it was mainly in the 1990's that GYs could be differentiated. host range. Description Grapevine yellows (GYs) are a group of severe phytoplasma-induced diseases occurring worldwide on a number of Vitis vinifera cultivars. Main synonyms: Flavescence dorée-like disease. and serology. a phytoplasma belonging to group 16SrII (or peanut witches' 135 . a severe decline of the vine. vector species have not been identified for all GY diseases. Main diseases: Flavescence dorée. Nevertheless. Bois noir. However. Red-berried varieties show reddish to purple discolorations while white-berried varieties show golden to chlorotic discolorations. Australian grapevine yellows. canes. variable in shape and size (50-1000 nm in diameter) that circulate in phloem sieve tubes and can be seen in the electron microscope passing through pores of sieve plates.200 kbp). Flavescenza dorata. Their titre is usually very low in grapevine. Downwards rolling of the leaves and sectorial discolorations of the blades. though some sieve tubes may appear crowded with phytoplasma cells in the electron microscope. They are wall-less. In 1997. Golden flavescence. Phytoplasmas have the smallest genome reported for procaryotic organisms (560 2. Candidatus Phytoplasma australasia. based mainly on sequence similarity of their rRNA genes and of a few other genes. The first GY to be recorded was Flavescence dorée in France in the 1950's. inflorescence and berries. develop on leaves. Agents: Phytoplasmas were called Mycoplasma-like organisms (MLOs) from their discovery in 1967 until the International Committee of Systematic Bacteriology (ICSB) replaced this name with the term phytoplasma in 1993. buds. Characteristic symptoms in the end of summer are a partial or total lack of lignification which may affect individual canes and shoots or the whole plant. involving also the main veins. Main symptoms: Young shoots of affected Vitis vinifera are weak with short internodes and frequent necrosis of terminal buds. Amarilliamento. Severely infected stocks decline rapidly. Phytoplasma cells are vesicular rounded bodies. their genome is poorly known because they cannot be cultivated. but not seeds. all organs of the plant may be infected. Vergilbungskrankheit. Schwarzholzkrankheit. North American grapevine yellows. associated to Bois noir (BN). However. such as the ribosomal protein genes or the elongation factor Tu. in addition to other criteria such as symptomatology. Stolbur phytoplasmas. Rubbery canes fall downwards with a typical "weeping" aspect. autumn fall of leaves occurs later on diseased than on healthy vines. persistence of infection in dormant plant material. Vergilbungskrankheit. Legno nero. resulting in reduction of quantity and quality of the crop. Flavescence dorée (FD) and Palatinate Grapevine Yellows (PGY) phytoplasmas belong to group 16SrV (or Elm yellows group). Phytoplasmas in the 16SrIII group (or X-disease group) are associated with grapevine yellows in North America and Israel. Palatinate grapevine yellows. Flavescenza dorata. However.GRAPEVINE YELLOWS: GENERAL PROPERTIES 1. Several Candidatus phytoplasma species have been recently described. Vergilbungskrankheit (=Schwarzholzkrankheit) or Legno nero and also one of the agents of Australian grapevine yellows (Candidatus Phytoplasma australiense) belong to group 16SrXII (or stolbur group). shoots. Phytoplasmas form a homogeneous phylogenetic clade subdivided into about 20 groups and subgroups. Leaf blades are crispy and brittle. The main characteristics of GY diseases are important losses or damage to the yield. and transmission by specific leafhopper or planthopper vectors. phloem-restricted bacteria that belong to the class Mollicutes. Bunches wither in early summer or berries shrivel later in season. GY agents have been identified in no less than 5 groups of phytoplasma clade. Quite often. The occurrence and spread of a particular GY mainly depend on the presence of efficient vectors in the vineyards or their close vicinity. after their aetiology had been understood and molecular methods for the characterization of associated phytoplasmas had been developed. depending on the particular disease and other conditions such as variety.
Full recovery and transient remission of symptoms have been observed in low and medium sensitive cultivars. Random Fragment Length Polymorphism (RFLP) analysis of the amplicon can provide further characterization of the infecting phytoplasma. Individual symptoms can be confused with other diseases or disorders. vinifera cultivars have been reported to be sensitive or very sensitive to GYs in all countries where infection occurs. are very sensitive to all GY diseases. Other host plants: Most phytoplasmas are ubiquitous plant parasites that can be hosted by several sometimes tolerant plant species. or on random selected non-ribosomal DNA fragments. Detection: GY syndrome is characteristic. including the agents of FD and BN. vector and abundance of reservoirs). The three ascertained vectors of GY diseases are univoltine leafhopper or planthopper species. Detection can also be made by transmission electron microscopy of thin sections or by scanning electron microscopy. The best antigen source are leaf veins and petioles. Though the rate of transmission to plant material can be very low. Varietal susceptibility and sensitivity: Numerous V. Double infections have been reported. numerous plants or weeds may be overt or discrete reservoirs from which acquisition of phytoplasma by vectors is possible according to the feeding preference of each species. Simultaneous presence of symptoms on leaves. Particular procedures for extraction of phytoplasma antigens from infected vines must be used to enhance the sensitivity of detection. Phytoplasmas in the 16SrI group (or aster yellows group) are associated with endemic GY diseases in several countries of Europe and in the USA. These phenomena also depend on the disease complex (phytoplasma. on less conserved genes. Riesling and also numerous local varieties. It is probable that the feeding preference of insect vectors. and bunches is strong evidence for phytoplasma infection. using DAPI staining. but the latter technique has never been successfull on field-grown grapevines. The extraction from infected tissues of total DNA containing enough phytoplasma DNA of good quality and the elimination of inhibitors of the PCR reaction. such type of transport is risky when potential insect vector species occur on the site of planting. which allows restoring the sanitary status of affected vineyards when the vector activity is controlled. as well as their feeding activity. Vitis riparia and American rootstock varieties and hybrids do not show typical symptoms of GY. canes. ELISA detection is possible from vascular tissue of symptomatic grapevines. 136 . These methods are not specific for any phytoplasma and are too laborious for disease monitoring. Cabernet Sauvignon. Polyclonal and monoclonal antisera have been raised to a few phytoplasmas. The situation of cv Syrah is controversial.broom group) is a second agent of Australian grapevine yellows. Outstanding progress in detection was achieved with DNA-based techniques. Hence. more specific primers can be used. They can also be observed in sieve tubes by light microscopy. Transmission: Phytoplasmas are vectored by hemipters. Some cultivars such as Chardonnay. Infection of rootstocks has been detected only in the case of Flavescence dorée but it cannot be excluded for other GY agents. However. when grapevine is not a preferred host for the vector. Then. shoots. Pinot noir. Phytoplasmas also persist in vine stocks during winter. are available in the literature. Most of these PCR primers have been designed on conserved regions of the rRNA gene and are "universal" for all known phytoplasmas. influence symptom expression and differential sensitivity of cultivars. Phytoplasmas can be detected by graft transmission to sensitive vines. designed on variable regions of the rDNA. erratic transmission to grapevine may nevertheless take place. are critical. When the presence of a particular disease is suspected. Others. mainly hoppers (cicadellids or cixiids) or psyllids. Propagation material may be the infection source for long distance dissemination of GY diseases. Phytoplasmas persist in their vectors but vertical transmission to the progeny of infected gravid females is not possible or scarce. PCR amplification with universal primers is preferred. are group specific. A range of primers that can be used for amplification with Polymerase Chain Reaction (PCR) of characteristic DNA fragments of phytoplasmas. When no information on the particular phytoplasma type is available.
Branas (a and b): Description of flavescence epidemics on Baco 22A and Chardonnay in France. Detection in the upper parts of the vines is usually done from veins and petioles of the leaves of symptomatic shoots. but is also possible using young leaves shortly before symptom expression or phloem scrapings from lignified canes. evolution of the disease in space and time. under the name Flavescence dorée. FD can be detected from leaves of non-symptomatic American roostocks. Detection of phytoplasma DNA can be achieved from all plant organs or insect vectors. Schvester et al. littoralis Ball. weeds or other crops. Control methods of vector populations are being experimented with natural insecticides. denoted Flavescence dorée (FD) in agreement with the name proposed by Levadoux (1955) because of the golden yellow metallic aspect of leaves. an insect that has been introduced recently from America. Pruning or top-grafting are useless because roots and trunks are infected. on physiological changes and defence mechanisms induced by infection. BN is considered as a non epidemic form of FD. and natural enemies. soaking of dormant material before or after grafting. The disease is thought to be caused by root damage. on the possibility to limit or eradicate their population and prevent their migration or movement in vineyards. Generally. FD is transmitted by the leafhopper S. Historical review 1955 Levadoux: Description of a severe outbreak of "Flavescence dorée" on Baco 22A in southwestern France. into hot water (50 °C) for a sufficient length of time (minimum 30 mn). In any case. and on the identification of reservoirs of inoculum such as infected grapevines. Caudwell (a): Studies on Bois noir (BN) and on its relationships with FD. Gärtel: Description. but does not rule out the possible involvement of a pathogen. The author suggests this disease to be due to adverse climatic and soil conditions. phytoplasmas are unevenly distributed in GY-affected vines. using detection of PCR amplification products with DNA-DNA hybridization with a specific probe. i. 1956 1957 1961 1961 1961 1964 1965 137 . The disease can be transmitted by grafting and is probably a virus disease. efforts should address prevention by planting healthy propagation material and limitation of vector populations and their inoculative activity. Caudwell (b): Description of recovery in grapevines affected with FD. Symptoms (macroscopic and microscopic). of a disease occurring in the vineyards of the Mosel Valley and of the Rhine Valley. tolerance and potential for recovery of varieties. Nonsymptomatic grapevines that show irregular symptoms from one year to the next may also test positive.. Caudwell: Characterization of a new type of flavescence. Investigations are being conducted on sensitivity. littoralis Ball found in Italy in 1963. 2.e. Control: There are no direct curing methods of infected plants.Other molecular methods are being developed. The biology of the insect is described. even sensitive detection methods may not be fit for use in sanitary selection because of the uneven distribution of phytoplasmas in mother plants and their vegetative progeny. cultural practices. and on elicitors of defence reactions to these phloem-restricted bacterial agents. Real-time PCR has also been successfully used. Production of sanitized propagation material is possible with hot water treatment (HWT). Vidano: S. Control of GY diseases depends on the knowledge of vector insect species. hypotheses on its nature. However.
Caudwell et al. and Euscelis sp. these findings have not been confirmed.: Rickettsia-like organisms observed in roots of grapevines with "Vergilbungskrankheit" in the Saar. titanus fed on the latter V. littoralis Ball is present in Ticino (southern Switzerland). is probably of European origin. Caudwell et al. Immersing cuttings taken on FD-affected grapevines in water at 30°C prior to planting reduced the proportion of infected plants by about 83%. Mosel and Rhine regions are considered as the causal pathogens of this disease. Rafaila and Costache: Yellows of grapevine with symptoms similar to those of FD found in 1967 for the first time in Romania. Conversely. Inoculation of grapevine seedlings with S. So far. Belli et al.1966 Caudwell: Attempts to inhibit the agent of FD in vivo by heat treatment. faba are different from those obtained with feeding inoculation with infective S. Caudwell and Larrue: Spread of FD in France is related to sanitary status of planting material. mainly on cv. Potential existence of several different diseases: leafhoppers (Euscelidius sp. Provisory name "Rhine riesling problem". witches' broom and yellowing. (b): Evidence that FD and BN are two different diseases with similar symptoms.: Production of antisera to FD-MLO raised to extracts of experimentally FDinfected plants (Vicia faba) and leafhoppers (Euscelidius variegatus) and first microscope observation of MLOs trapped with ISEM. As similar organisms have been found in nematodes of the species Xiphinema index. An important FD-like disease is reported.: Discovery of Mycoplasma or PLT group-like microorganisms in the phloem elements of plants showing dwarfing. This phytoplasma was later on identified as a Clover phyllody phytoplasma. Recommendation that mother plants should be grown in areas far away from FD-infected regions. of which S. Baggiolini: S. Caudwell et al.: S. littoralis. Magarey and Wachtel: Description of a new disease of grapevine of the yellows type on the cv Rhine Riesling in South Australia.) caught in the vicinity of vineyards could transmit a yellows disease to Vicia faba plants. Rumbos et al.: S. Tanne and Nitzany: Occurrence of a yellows disease resembling FD in Israel. Osler et al. Belli et al. the hypothesis is put forward that this nematode is the vector of the disease. Symptoms on V. titanus (= littoralis) is not a vector. The author assumes that both FD and its vector were introduced from North America with varieties of Vitis labrusca between 1927 and 1950. (a): Mycoplasma-like organisms (MLOs) are observed in diseased grapevines and infective leafhoppers and in Vicia faba submitted to feeding inoculation with infective S. Granata: Description of an epidemic yellows disease on cv Inzolia in Sicily. Bois noir. The disease has been observed for the first time in 1968. Caudwell: Symptoms of grapevine yellows (Amarilliamento) reported in Chile on cv Elqui. littoralis. faba plants produced typical GY symptoms. littoralis is present in the same region of Oltrepò Pavese.: Presence of a disease similar to FD in the Oltrepò Pavese (northern Italy). Caudwell: Discussion on the origins of yellows diseases of plants. Doi et al. First record in 197576. 1967 1968 1970 1971 1971 1973 1973 1975 1977 1979 1980 1982 1982 1983 1984 1985 1985 138 . with special reference to grapevine. titanus found in 1984 in vineyards of northern Italy. Regina. Credi and Babini: Grapevine yellows are reported in Emilia Romagna (Italy).
control measures. Egger and Grasselli: Presence in Toscana. Credi et al. Italy. vector of FD. Two of these vineyards were sprayed with insecticides in 1986 and 1987. Fortusini and Belli: Description of the development of epidemics of FD-like diseases in northern Itay. Credi: Description of a FD-like disease in Emilia Romagna and the reaction of 3 different cultivars. disease distribution. in viticultural areas of northern Italy and of other Auchenorrhynchas susceptible to play a role in the transmission of yellows diseases in this region. Susceptibility and sensitivity of affected varieties (Chardonnay.: Study on the distribution of S. S. for the presence of a FD-like disease. Italy.: Contribution to the knowledge of FD (or similar diseases) in the Veneto region.: Observations. Seljak: Presence of Scaphoideus titanus in western Slovenia (Yugoslavia).: Presence of a FD-like disease in Emilia-Romagna. Recovery and crop loss are variable. titanus.: New data on the spread of FD in three vineyards of Oltrepò pavese (northern Italy) from 1985 to 1987. Quaroni et al. Carraro et al.: Presence and distribution of a FD-like disease in the region Friuli-Venezia Giulia in Italy. Credi and Callegari: Survey of vineyards in Emilia-Romagna.or xylem-limited prokaryotes in Europe. Vidano et al. resulting in a reduced diffusion of the disease. northern Italy. However. Vidano et al. whereas an unsprayed vineyard was more affected. Rui et al. The results suggest that the disease is brought into the vineyards from outside local sources. their etiology is unclear.: Occurrence of FD-like symptoms in the valley of Sarca in Trentino. Pinot nero) and comparison with other similar diseases. titanus. Chardonnay is the more affected variety. Boudon-Padieu and Larrue: ELISA detection of FD pathogen on infected reared experimental and wild natural leafhopper vectors (E. titanus is not always associated to the disease.: Survey of potential Auchenorryncha vectors of the pathogen agent of FD in Piemonte (Italy). 1987 1987 1987 1988 1988 1988 1988 1988 1988 1989 1989 139 .1985 1986 1986 1986 1986 1986 1987 1987 1987 Rumbos and Avgelis: Observations of a FD-like disease in Greece with severe symptoms on cvs Razaki and Roditis. Italy. respectively). Martelli: A review on the knowledge on grapevine diseases induced by phloem. using the scanning electron microscope. titanus. Borgo et al. Pinot bianco. that are responsible for severe decline of vines. Conti: A review of intracellular procaryotic phytopathogenic agents. Survey for the presence of S. vector of FD. of a FD-like disease on Chardonnay. Magarey et al. Mescalchin et al.: Observation of MLOs in phloem tissue of yellows-affected grapevines in Australia. of MLOs in phloem tissues of grapevines affected by "FD" in northern Italy. In addition to S. variegatus and S.: Description in Italy of the presence of FD or FD-like diseases. Hyalesthes obsoletus. Fortusini et al. Euscelidius variegatus and Euscelis incisus are taken into consideration. titanus. Italy. Borgo: General information on the presence of FD-type diseases in northern Italy.
(a and b): Identification of numerous species of Auchenorrhynchas in the vineyard ecosystem in Piemonte (northen Italy) and of ampelophilous species. Similar symptoms on Chardonnay could be obtained by insect inoculation (Euscelis incisus) and by cleft-grafting of tissues taken from yellows-infected elm. Boidron and Grenan: Construction and testing in ENTAV. Di Terlizzi et al. Credi et al. Affected grapevines in the Rhône valley tested negative.: Bench grafting experiments of buds of indicator cvs. Conti: A yellows-type disease of cv. Specific primers can be used for MLOs. Vidano et al. suggesting an unrelated agent. 1989 1990 1990 1990 1990 1991 1991 1991 1991 1991 1991 1991 1992 1992 1992 140 .: The FD-like disease developing in Friuli-Venezia Giulia since the early 1980's was not decreased significantly in vineyards sprayed with insecticides though S. Caudwell: Review on the epidemiology and characterization of FD and other GY diseases. Credi and Santucci: Development of "Flavescenza dorata" in grapevine in Emilia-Romagna and rate of infestation in different systems of cultivation. Chardonnay develops in Tuscany. Treatment prior to storage is more efficient. The authors recommend that research continues on techniques for the ecological control of weeds and of leafhoppers linked to them and on the aetiology of the different forms of "golden flavescence". The recommended temperature / time combination is 50°C / 3560 min. Caudwell and Kuszala: Detection with ELISA of FD-MLO in naturally FD-affected grapevines. Le Grau du Roi (France) of a device for secure soaking of dormant buds and canes into hot water (50°C. Symptoms also developed on plants obtained from buds taken on diseased Chardonnay and Sangiovese grafted on healthy Kober 5BB.: Occurrence of grapevine yellows in Moldavian SSR. Granata and Grimaldi: MLOs are observed with the electron microscope on affected grapevines of cv. titanus was present in all vineyards checked. titanus was not found in the area. Natural infection with MLOs of bait plants (Catharanthus roseus and Vicia faba) placed in vineyards in spring and summer. Restriction profiles after digestion with endonucleases permit the differenciation of MLOs associated to different plant diseases. S. Caudwell et al. This is a prospect for investigation on MLOs associated to plant yellows. Chardonnay and Baco 22A grafted on donor yellows-diseased plants of cvs Pinot blanc and Sangiovese. Granata and Russo: An epidemic disease with FD-type symptoms develops in Sicily on cv. Only part of the plants were infected. The aetiology is not fully ascertained though the presence of MLO is probable.1989 Rumbos: Review of the knowledge on the etiology of grapevine yellows and observations with the scanning electron microscope of root and petiole tissues from diseased and unaffected plants of cv Riesling.: Presence of yellows-like symptoms in Apulian grapevines (central Italy). Deng and Hiruki: A method to amplify 16S rRNA gene from culturable and nonculturable mollicutes.: Grapevine graftwood shoots infected with FD can be disinfected by hot water treatment in the dormant stage. Inzolia in Sicily. Several weeds with symptoms of phyllody were found to contain MLO in electron microscope studies. The annual rate of symptomatic plants was never greater than 20 % over a 6-year period. Inzolia. Ahrens and Seemüller: Oligonucleotides selected in the conserved parts of the 16S rRNA gene of MLOs can be used with polymerase chain reaction (PCR) to amplify a sequence of the gene in several plant pathogenic MLOs maintained on periwinkle (Catharanthus roseus). 45mn). Refatti et al. Marinesku et al.
Senescent or degraded forms of MLOs identified inside degenerate sieve elements. genomic tests).1992 Credi and Santucci: Of 628 attempts to transmit the agent of a yellows disease of grapevine from Sangiovese. Evolution of the disease and of its vector in the various viticultural regions of France.6%) were obtained. transmission by feeding to healthy grapevines succeeded only in 1 attempt out of 100. One MLO strain transmitted with Macrosteles quadripunctulatus appears similar or close to aster yellows MLO. Typical symptoms on several cvs. titanus population in vineyards and the rate of spread of the disease. detection (ELISA. The MLO strains found in grapevine are related with aster yellows MLOs. Davis et al. (a and b): Characterization of Italian periwinkle virescence (IPVR) MLO (obtained on bait periwinkle plants exposed in yellows-affected vineyards in Italy) is possible with DNA-DNA hybridization and PCR. Bianco et al. Arzone et al. titanus. (a and b): Observations supporting the existence of different types of grapevine yellows in northern Italy. but are different from known aster yellows cluster MLO strains.: FD can be transmitted by symptomless rootstocks. (b): Evidence of an association of MLOs with grapevine yellows in Emilia Romagna. FD can be readily distinguished from Bois noir. Quacquarelli and Barba: Review of the distribution of flavescence dorée and other grapevine yellows in EEC viticultural countries.: Transmission experiments from grapevine to grapevine and to herbaceous hosts with S.: Construction of serological and molecular detection tools to MLO transmitted to periwinkle with dodder from a yellows-diseased grapevine in Friuli-Venezia Giulia (Italy) (later 1992 1992 1992 1992 1992 1993 1993 1993 1993 1993 1993 1993 1993 1993 141 . MLOs were visible in sections of petioles of periwinkle showing symptoms and subsequent graft transmission from periwinkle to periwinkle was successful. Alma et al.: First detection of FD-MLO DNA in grapevine extracts by DNA-DNA hybridization with DNA probes cloned from FD-MLO partially purified from experimentally infected plants (Vicia faba). Attempts to characterize MLO DNA extracted from insects with molecular methods. suggesting the presence of two different diseases at least in Veneto and Friuli-Venezia Giulia. However.: Description of MLOs and associated disorders in the phloem of FD-affected LN33 experimentally inoculated with S. No spontaneous recovery. Meignoz et al. Boubals (a): Report on a meeting of the French working group on FD. titanus is not present.: ELISA with FD-antibodies permit to detect related antigens in S. Hot water treatments suppress the transmission to Chardonnay indicators. Arnò et al. Osler et al.: The survey in Piemonte of Chardonnay vineyards affected with a FD-like disease shows that no correlation can be made between the importance of the S. FD-MLO DNA can be distinguished from other MLO DNA with Dot-blot hybridization and Southern-blotting. Daire et al. S. Caveccia and Chardonnay vines to periwinkle by the dodder species Cuscuta campestris. Caudwell et al. It is more related to the agent of Elm yellows than to other yellows agents. Chen et al. titanus and other leafhopper species show that yellows disease present in northeastern Italy differs from flavescence dorée as it occurs in France. (a): Differentiation with PCR-RFLP analysis between a MLO related to aster yellows in GY-diseased plants from Lombardia and a MLO related to elm yellows in an affected grapevine in Friuli-Venezia Giulia. results of research. The titre of MLO appears very low in grapevine. Bertaccini et al. 4 positive results (0. IPVR is related to aster yellows MLO. Boubals (b): A severe epidemic of grapevine yellows occurs in Golan (Israel). titanus leafhoppers.
MLO appear to be in very low titre. BN (stolbur MLO) detected in several regions of Italy and in Israel. (a and b): PCR-RFLP analysis of 16S rDNA from MLOs show that 2 different MLOs at least occur in yellows-affected grapevines in France. Osler et al. Computer analysis for spatial patterns of diffusion show a non-random distribution and suggest transmission from other vines or from weeds. graft and dodder. 1993 Daire et al.: Important outbreak of a yellows disease on local varieties in Apulia (southeastern Italy) and infection of periwinkle as bait plants and with dodder transmission.: FD-ELISA method was used on tissues of yellows-affected grapevines from diverse countries. Credi (a and b): Observation of MLO in yellows-affected grapevines in northern Italy. Daire: PhD thesis on detection and differentiation with DNA-based methods of the MLO agents of GY diseases in France. using insects. Detection in non symptomatic V. titanus to test plants of cvs Perera and Chardonnay in Veneto and Friuli-Venezia Giulia (Italy). S. Subcommittee on the Taxonomy of Mollicutes: The trivial name MLO is replaced by the term phytoplasma as a consequence of the demonstration by diverse studies that the phytoplasmas represent a monophyletic clade of organisms more closely related to mollicutes than to walled bacteria. Maixner: Demonstration that Hyalesthes obsoletus (Auchenorrhyncha: Cixiidae) is a vector of German grapevine yellows (Vergilbungskrankheit). Specific probes cloned and specific primers designed for PCR amplification of phytoplasma DNA.: Six-year transmission trials of different types of yellows with S. c and d): Grapevine yellows (Vergilbungskrankheit) are common in the Moselle and Rhine valley. titanus is not present. at least two different phytoplasmas are infecting grapevines in Veneto: FD sensu stricto (transmitted by S. Only FD and BN have been identified in naturally affected grapevines. Prince et al. Maixner (a. Transmission has been obtained only from vines of Veneto. In northeastern Italy. International Committee of Systematic Bacteriology (ICSB). FD is related to elm yellows and BN is related to stolbur. Kuszala et al. Electron microscopic observation of MLO in periwinkle and grapevine.: Diversity of MLOs associated with grapevine yellows and transmitted to periwinkle.called FDU. healthy plants protected with plastic screens did not develop symptoms and diseased plants protected with screens showed transient recovery. S. FD detected in leaves of rootstock 3309C and is also present in samples from Veneto (Italy). Positive assays obtained only on samples from France and Veneto. FD belongs to the elm yellows group but is different from elm phytoplasmas. A MLO has been transmitted to periwinkle with dodder. b. Di Terlizzi et al. BN belongs to the stolbur group. aster yellows and Xdisease. Description of pathological changes in phloem of leaves and identification of senescent forms of MLO inside degenerate sieve elements. probably transmitted by another insect. So. Classification of grapevine MLOs into three RFLP groups: elm yellows (FD from France). then GYU phytoplasma) and comparison of specificity and sensitivity of the latter tools to detect phytoplasmas in grapevines in New York and Italy. Experiments on the use of hot water treatment on planting material. GYU was later on characterized by other authors as a phytoplasma belonging to the Xdisease (16SrIII) group. titanus is present in the Friuli-Venezia Giulia region but no transmission assay has been successful. 1993 1993 1993 1993 1993 1993 1994 1994 1994 1994 1994 142 . Carraro et al. Detection in grapevines from Virginia (USA) of MLO related to X-disease. on the effects of pollarding affected vines and of chemical sprays against vectors. A third MLO related to X-disease of Prunus detected in a grapevine from New York (USA) and in a periwinkle carrying a grapevine MLO (FDI = FDU) transmitted through dodder in Udine (Italy). (a and b): Various attempts to transmit FD-like disease in different regions of northern Italy. riparia in New York and failure of detection in some symptomatic grapevines in New-York and Italy. titanus) and a second one. Girolami and Egger: Importance of contamination of vineyards and diffusion of Grapevine yellows in northern Italy.
: Emphasis on the possible role of weeds as reservoir for MLO in vineyards. One grapevine out of 16 contained a 16SrV (= elm yellows) phytoplasma together with a 16SrI-G phytoplasma.: Grapevines from Piemonte (northern Italy) contained several different phytoplasmas. Detection is also possible during winter on wood scrapings. Laviña et al.1994 1994 Parente et al. in grapevines with yellows symptoms in Soave (Veneto.: Identification of BN (stolbur phytoplasma) in Spain. Padovan et al. Double infection is reported. 1995 1995 1995 1995 1995 1995 1995 1995 1995 1996 1996 1996 1996 1996 1996 1996 1996 143 . probably of Bois noir type. (a and b): Grapevine yellows in Latium (southwestern Italy) are not associated with S. Prince et al. aster yellows. Ten weeds were found harboring MLOs with transmission electron microscopy. (a and b): Detection of a phytoplasma closer to aster yellows phytoplasma than to elm yellows phytoplasmas in GY-affected vines in Australia Sancassani and Posenato: Simultaneous presence of FD and of other yellows type in vineyards of Veneto.: Six-year survey of evolution of yellows symptoms in a 6-12 year-old vineyard of cv Chardonnay that was never sprayed with insecticides. PCR detection of phytoplasma with universal primers.: Survey of yellows symptoms in an ampelographic collection of 1281 cultivars in Conegliano (Veneto. Murari et al. Haidar: First report of symptoms of yellows on grapevines in Lebanon. with the aim of identifying sources of tolerance or resistance to yellows diseases.: Presence and distribution of GY in Sicily. Molecular detection of aster yellowsrelated phytoplasma. Alma et al. Consideration on possible resistance and research on interaction of phytoplasma with nepovirus infection. Del Serrone and Barba (a and b): Monitoring of detection of phytoplasma in the organs of grapevine according to the vegetative state. sometimes in mixed infection. Boudon-Padieu (a and b): Reviews on knowledge and research on grapevine yellows. in the vector H. Arzone et al. Italy).: The phytoplasma associated with Australian grapevine yellows is close to but different from stolbur phytoplasma causing BN. Italy). Egger et al. The most frequent belonged to group 16SrI-G (later designated as 16SrXII or stolbur group). Koruza: Dispersal of grapevine yellows in Slovenia. (a): Specific detection with PCR of stolbur phytoplasma in grapevines affected with Vergilbungskrankheit. Related MLOs were also detected in wild grapevines. The importance of proper identification of disease type for control measures is emphasized. Albanese et al. Del Serrone et al.: Presence of grapevine yellows in northern Portugal and electron microscopic visualization of MLOs in the phloem tissue of petioles of affected plants.: MLOs associated to grapevine yellows in Virginia belong to 2 different groups : X-disease group (16SrIII) and aster yellows group (16SrI). Padovan et al. obsoletus and in weeds growing in the vineyard in Germany. stolbur and apple proliferation) can be detected. titanus which has not been found. Maixner et al. Fortusini et al.: Four different phytoplasmas (FD. Maixner et al (b): Detection of a new phytoplasma type related to FD in cv Scheurebe showing symptoms of yellows in Palatinate (Germany).
(a): Only stolbur and FD phytoplasmas were detected in samples of yellows-affected grapevines from numerous regions of France and from Italy. Boudon-Padieu and Maixner: Updating in French and English of present knowledge on grapevine yellows: etiology. biology and control. No epidemic outbreak has been observed. No positive detection with PCR on the HW-treated vegetative progeny of mature canes of symptomatic plants. Del Serrone : A review in Italian on the knowledge on etiology. Kölber et al.: Use of Hot water treatment (HWT) to eliminate phytoplasmas (FD and AY) from 3 cultivars of grapevine. transmission. spread. titanus is not present. Bosco et al. Borgo: Description and colour photographs of symptoms of grapevine yellows and knowledge on their aetiology and vectors. International Committee of Systematic Bacteriology (ICSB). Phytoplasma belonging to the stolbur subgroup have been identified. Severe symptoms with high incidence on seven cultivars are reported from four grapevine-growing regions surveyed for 5 years (1993-1996). titanus reported for the first time in this region. The possibilities of control 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1998 1998 1998 144 . S. Presence of S.1996 1997 1997 1997 Rüdel: Review on the history and diversity of Vergilbungskrankheit of grapevine in Germany.: Survey of leahoppers in vineyards of Piemonte. Davis et al. Aster yellows and X-disease types were never detected. Batlle et al. titanus is not reported.: Recurrent and severe yellows symptoms observed since the 1980's in Sardinia (Itlay) on several cultivars.: Symptoms of grapevine yellows observed in Hungary in the early 1970's. No double infection has been found to occur. 10 species were known vectors of phytoplasmas. Among 29 species of Auchenorrhynchas from the vine canopy and weeds of the border. Spain and Israel. (b): Prospects on detection and identification of grapevine phytoplasmas with new molecular tools and consequence on future epidemiology studies. 10 are confirmed phytoplasma vectors. (a): Description of available methods and strategies for detection of phytoplasmas in grapevine. Aldini et al. Tanne and Orenstein (a and b): Successful heterografting transmission of phytoplasma from symptomatic grapevine to periwinkle and easy subsequent identification of phytoplasma type. Positive reaction with a DNA probe specific to aster yellows phytoplasmas. Garau et al. Subclades are considered to represent the equivalent of distinct species. S. AY and X-disease related phytoplasmas have been transmitted and detected. Italy). Maixner et al.: Survey of hoppers in vineyards of the province of Piacenza (Emilia Romagna. Murari et al.: Identification of Flavescence dorée in Spain. The term phytoplasma will become the genus name of plant pathogenic mycoplasmas under the provisional taxonomic status Candidatus. Maixner et al.: History and control of GY diseases in Italy. vectors and detection of the agents of Grapevine yellows. Among 32 species identified. Belli et al. Subcommittee on the Taxonomy of Mollicutes: The taxonomy of phytoplasmas will refer to their molecular phylogeny. Maixner and Reinert (a): Hot water treatment (HWT) of 50°C for 60 min applied to dormant mature canes eliminated Vergilbungskrankheit phytoplasma with a small loss in survival of the cuttings. Daire et al. (b): Survey of possible auxiliaries for the biological control of leafhoppers and planthoppers in vineyards. northern Italy.
: A comprehensive review of the classification of phytoplasmas in the world. phytoplasmas can be classified into 14 groups and 38 subgroups. are compulsory in France. methods of detection of phytoplasmas in grapevine tissues and in insect vectors. BN is very frequent but H. Guadagnini et al.: Phylogenetic studies of phytoplasmas associated to numerous plant diseases. Boudon-Padieu: Review on recent progress in the knowledge on grapevine yellows.: Summary of the complex situation of GY in northeastern Italy. FD is restricted to the north of Cataloña. Lherminier et al. Sensitivity of assays must be good because of the low titre of phytoplasmas in affected plants. 1998 1998 1998 1998 1999 1999 1999 1999 2000 2000 2000 2000 2000 145 . has been identified as a member of the X-disease group. Boudon-Padieu (b): Chapter in a Handbook on viral and bacterial diseases of the grapevine. distribution in the world. Survey with PCR of phytoplasmas infecting grapevines in Friuli-Venezia Giulia and Trento province (Italy) and in Slovenia. titanus. Maixner and Reinert: Updating in German of the research made in different countries on grapevine yellows diseases. Search for alternative vectors. Frausin et al. Borgo et al. Batlle et al. Insecticide sprays against S. 1998 Lee et al. have shown that only stolbur group phytoplasma was consistently found.: Report of positive detection of aster-yellows phytoplasma in grapevine fed with Metcalfa pruinosa and in the body of the insects used for transmission. and M. titanus is more widely distributed. titanus. Reinert W. according to the most recent studies on molecular structure of rDNA. nucleic acid hybridization and serological comparisons. GY affect mainly the northern provinces.: Unsatisfactory results on the use of hot water treatments to eliminate phytoplasmas from grapevine wood.: Monitoring grapevine yellows in North-eastern Italy and attempts to rationalize routine laboratory testing for the differentiation between FD and stolbur phytoplasmas. obsoletus is rarely found in vineyards. Grapevine phytoplasmas classify into different groups. Seemüller et al.: A history of GYs in north-eastern Italy. Firrao et al.: Evaluation on the efficiency and security of hot water treatment used to eradicate phytoplasmas from propagation material. No negative effect was observed on scion buds but rootstock cuttings were more sensitive to negative effects of the treatment. In addition a phytoplasma (FDU=GYU) transmitted from GYaffected vines to Catharanthus roseus by means of dodder in Friuli-Venezia Giulia. epidemiology of these diseases. In the Veneto region only stolbur-group phytoplasma was recorded in some areas and both stolbur and elm yellows group phytoplasmas in other areas. Carraro et al.: First report of the use of oligonucleotides for in situ hybridization with transmission electron microscopy to localize phytoplasma in plant cells. though S. epidemiology and etiology of GYs in the world. symptoms. Maixner: A review of thermotherapy to cure phytoplasma infected material. Similar attemp with FD phytoplasma were not successful.: A review of the situation of GY in Spain.depend on the knowledge of the biology of phytopathogenic agents and of their vectors. Refatti et al. Phytoplasmas for which 16S rDNA sequence is available have been classified into 20 major groups and subgroups. vector of FD. FD-related phytoplasmas (elm yellows group) have been detected in grapevines from Treviso and Verona and transmitted from vine to vine using S. Only FD and BN have been identified. Probes were antisense to stolbur specific sequences in the 16S rDNA and used on stolbur-infected periwinkle. On the basis of RFLP of PCR-amplified 16S rRNA gene. with a particular reference to the work done in Germany. History.
Moretti et al.: The presence of phytoplasmas in liquid medium on which insect vectors have been fed can account for potential vectorship of the species. Crocker et al.: Use of immersion in hot water (HWT) or in chemicals to suppress phytoplasmas from grapevine propagation material in Italy. Seljak: A review of non-European hoppers introduced in Slovenia. Quartau et al.2000 Moretti and Anaclerio: Evaluation of the effects of hot water treatment on the viability of grapevine cuttings of different varieties with different temperature / time combinations. Influence of rootstock on the rate of GY symptoms in the same vineyard and same grapevine variety. Dual infection may occur (13 %). Frosini et al. Plant variety and cutting diameter influence the rate of surviving cuttings. Macrosteles quadripunctulatus and Orosius orientalis were found on weeds in vineyards or trapped on sticky traps. A high rate of recovery is observed from one year to the next.: Recommandations for the organisation of nursery in Australia and handling of hot water treatment to avoid dehydration of cuttings during treatment. Experimental evaluation of the method and application to field-trapped individuals of suspected vector species. Klein et al. Phytoplasmas were detected in all five species. Bertamini and Nedunchezhian: Study of the effect of phytoplasma infection of field-grown plants of cv Chardonnay affected with Bois noir (stolbur phytoplasma) on the physiological response of plants: photosynthesis.: In Golan Heights (Israel) Hyalesthes obsoletus. Tanne et al.: Report of experiments combining temperature and time of treatment with hot water on 5 grapevine cultivars to cure planting material from phytoplasmas. titanus is widespread in western vineyards. sugar metabolism. although its vector S.. Boudon-Padieu: Updating of knowledge and research on grapevine yellows worldwide. The latter was difficult to evaluate because of the non homogenous distribution of phytoplasmas in canes. nitrate and nitrite reductase. Marzachi et al.: Use of hot water treatment in commercial nurseries in Australia to eradicate diverse pests and pathogen agents from propagation material. Seljak and Petrovic: Overview on phytoplasma diseases of grapevine and fruit trees in Slovenia. 2000 2001 2001 2001 2001 2001 2001 2001 2001 2001 2001 2002 2002 2002 2002 2002 2003 146 . Israel. FD has not been identified.: Survey of GY in Israel. Zahavi et al. Bertaccini et al.: Development of a new technique with DNA chips to detect phytoplasmas in grapevine. Orenstein et al. Phytoplasmas seemed to disappear from the insect body after the transfer of insects to healthy plants. Primers fitted for specific detection of these 3 phytoplasmas permit cheaper and quicker diagnosis.: Metcalfa pruinosa specimen reared in the laboratory may acquire FD and clover phyllody (aster yellows group) phytoplasmas by feeding on infected Vicia faba but feeding transmission to several plant species has not been obtained. BN and aster yelllows in vineyards of southeastern Piemonte (Italy). aster yellows (11 %) and X-disease groups phytoplasmas. (a): Presence of FD. Hyalesthes obsoletus has been found but not in vineyards. Osler and Refatti: The situation of GYs in northern Italy.: Scaphoideus titanus is present in Portugal. HWT (50°C/40 mn) gave acceptable survival but none of the conditions experimented provided a total elimination of phytoplasmas. Circulifer sp. Neoaliturus sp. which are all known as vector of phytoplasmas. Clair et al. The phytoplasma agents are not specified. Waite et al. suggesting that they did not multiply in the body of the insects. confirmed the presence of three phytoplasmas associated to GY: stolbur (70%).: A 2-year survey in Golan Heights.
: Important presence of GY in Abruzzo (central Italy). An aster yellows phytoplasma is suspected.: Flight activity of Scaphoideus titanus and Hyalesthes obsoletus and their infection status towards FD and BN (stolbur) phytoplasma.: Data on 15-year studies on efficiency and effect on propagation material of hot water treatment.: Measurements of basal respiration rate of grapevine dormant wood as an indicator of the best period for application of hot water treatment.: Survey of potential vectors of phytoplasmas in vineyards of the Golan Heights. two nonpreferentially ampelophagous vectors of grapevine phytoplasmas.: Grapevines affected with yellows are found free of phytoplasmas after recovery. Clair et al. Orenstein et al. titanus found associated to grapevine yellows with a high incidence in vineyards of Rastina (South Serbia). rapidity and sensitivity of methods for use in routine diagnosis of grapevine phytoplasmas. Recovery is a progressive phenonenon developing over 3 years in the case of BN. in particular a clover phyllody type (16SrI-C) which is quite frequent.. Tassart-Subirats et al. 2003 2003 2003 2003 2003 2003 2003 2003 2003 2004 2004 2004 147 . No important damage has been reported. M'hirsi et al. Neoaliturus fenestratus.: Important outbreak of grapevine yellows on cv Chardonnay and identification of a stolbur phytoplasma in diseased plants.: Report of aster yellows phytoplasmas in grapevine affected with yellows in Tunisia. stolbur phytoplasmas are suspected to occur because of long-lasting record of stolbur disease on many host plants in Serbia. fitted to routine detection. Megophthalmus scabripennis positive for aster yellows phytoplasma.2003 2003 2003 2003 2003 2003 Boudon-Padieu et al. Gajardo et al. Study of the spatial and temporal dispersion of the four species. Kuzmanovic et al. BN (stolbur phytoplasma) has an incidence of about 30 %. Marzachi et al.: Report of stolbur phytoplasmas in grapevine affected with yellows in Albania.: Identification of aster yellows (16SrI-B and C) and of ash yellows (16Sr-VII) related phytoplasmas associated to yellows of grapevine in Chile.: Report of severe symptoms of yellows on numerous grapevine varieties in Serbia and observation of organisms resembling phytoplasmas in sieve tubes of affected shoots collected in the Zupa area. Discussion on benefits and limits of artificial feeding medium to assess the transmission capability of phytoplasma by insects. Crocker et al. Lessio et al. Consequently roguing of affected vines should be avoided when possible and especially if vectors are controlled. Hyalesthes obsoletus and Circulifer haematoceps abundant and positive for stolbur and aster yellows phytoplasmas. Leitner: The situation of Grapevine yellows in Austria is now surveyed by means of molecular detection assays.: Improvement of detection of FD and BN in field-grown grapevines with real-time PCR. Duduk et al.: Compared efficiency. (a and b): FD phytoplasma (16S rV-C) and S. Chabbouh et al. D'Ascenzo et al. respectively. Osler et al. Apart from FD phytoplasma reported by Duduk et al. Myrta et al.: Identification of grapevine yellows symptoms in Tunisia and attempts to identify pathogen agent and vector.: Development of a sensitive PCR procedure for dual identification of FD and BN phytoplasmas. Ge and Maixner (a): Transmission to feeding medium is more efficient than transmission to natural host plants and to grapevine for Hyalesthes obsoletus and Oncopsis alni. Milkus et al. Other phytoplasmas are reported.
Isolates F70. After the discovery of other GY diseases. Corsica. FLAVESCENCE DORÉE 1. Two isolates denoted FD-D and FD-C. titanus is well established. Agents: FD was first regarded as a physiological disorder. Diseased vines have a patchy distribution in the plot. titanus. littoralis Ball) (Homoptera. It is widespread in all northern provinces of Italy and an outbreak was recently reported from southern Serbia. Hence. First symptoms may 149 . southern Italy. Several isolates have been characterized with molecular criteria. S. rooted cuttings from infected canes of a few rootstock varieties can develop "vinifera-like" symptoms on the wood and leaves. In addition. indicating a vine-to-vine transmission. These isolates could be distinguished by Western-blot analysis using FD antisera and monoclonal antibodies (Mabs). Vineyards in these areas are under the threat of an FD outbreak that could occur if FDinfected planting material is introduced. FD88. and western Slovenia. symptoms may be limited to a few shoots. titanus individuals collected in infected vineyards of south-western France. were characterized in Italy and shown to be transmitted also by S. Phytoplasma acquisition by all insect instars (larvae and nymphs) may occur from infected vines at any time from the beginning of hatching (usually in the beginning of May) throughout the growing season. Infected rootstocks show little or no symptoms. and Savoie. Eggs are laid in summer on two-year old wood and trunk. If the plants are inoculated after recovery. Main symptoms: It is believed that vines that show symptoms for the first time in a vegetative season had been infected in the previous summer. Feeding transmission starts after a 4-week latency in the body of the vector. Infected rootstocks are important means of dissemination because they are symptomless. that has colonized a wide climatic area. In France. It is also present in regions from which FD has not been reported in France. Moreover. FD is endemic only in regions where S.GRAPEVINE YELLOWS: INDIVIDUAL DISEASES A. titanus. Cicadellidae). Crop losses may be very high. the term "FD sensu stricto" was applied to diseases and phytoplasmas that are transmitted by S. However. Symptoms develop on leaves and shoots in June and July to become oustanding in August and autumn. The associated phytoplasma (= MLO) was observed in phloem tissue of affected plants and in insect vectors in 1971 and later classified in the Elm yellows group (16SrV). with an incidence that increases rapidly from one year to the next. FD92 and FD2000 were experimentally transmitted to broadbean (Vicia faba L. decline progressively and die. It is highly epidemic and extremely dangerous because of the biology and ethology of its leafhopper vector. Lignification of the canes is usually incomplete. Affected vines of most varieties may recover in the second year when they are protected against reinoculation with insecticide treatments. transmission is possible from early June (about one month after the beginning of hatching) until the death of adults. Transmission occurs also by vegetative propagation and grafting. However. then as a virus disease. extremely sensitive varieties do not recover.) using S. isolated infected vines have been identified in northern vineyards of Burgundy and Alsace. littoralis Ball). north of Spain and Portugal. Transmission: The FD agent is transmitted in the persistent mode by the leafhopper Scaphoideus titanus Ball (= S. symptoms affect the whole plant. Description Flavescence dorée (FD) was the first Grapevine yellows (GY) disease to be reported. This leafhopper is a neartic species specialized on Vitis sp. Though the rate of transmission by bench grafting can be very low. Infected buds or graftwood may be collected from symptomless parts of infected mother vines or from recently infected vines that have not yet developed symptoms. Scaphoideus titanus Ball (= S. It was described in a limited area in north-eastern Cataluña (Spain). FD88. introduced into Europe at the beginning of the 20th century. titanus is a univoltine species that overwinters at the egg stage and develops from May to September on grapevine leaves with 5 aerial apterous larval instars followed by alate nymphs that feed on the leaf veins and petioles and on green shoots. Switzerland. Molecular comparisons have shown that all these isolates are more closely related among them than with other phytoplasma strains in the same group. this way of spreading is significant when infected grafted plants are planted in vineyards hosting the vector. Most of the time. FD92 and FD-D could not be distinguished from one another. Leaves persist longer on affected plants. show a general asymmetric bent posture and necrosis of terminal bud. it occurs in all southern vine-growing regions.
Cabernet Sauvignon. Alicante Bouschet. its area of origin. Grenache. The second treatment. In spite of the possibility of recovery. Italy and Spain are susceptible with various degrees of sensitivity. such as Merlot. titanus in New York State (USA). a DNA fragment specific to BN (stolbur) phytoplasma can be used for the survey and monitoring of these two GY diseases. Soaking of dormant material in hot water (HW) is recommended. Under these circumstances only chemical insecticides are efficient for eradication of the disease. has been used as the official method for large scale survey of FD in France from 1993 to 2003. In 2003 a phytoplasma resembling the FD-C isolate was found in wild Clematis in Veneto (Italy). appear more tolerant. In France and Italy budwood can be taken only from mother plants growing in plots that have shown no GY symptoms for the last two growing seasons. Antigen extraction from vascular tissue of symptomatic grapevines requires the use of high Tris molarity and strong detergents in the extraction buffer. Nevertheless. has become the preferred detection method because it is easy to perform and was improved in reliability and sensitivity. Standard conditions (50°C for 45 min) proved highly efficient for sanitizing infected rootstocks and grapevine cultivars. Sequence or RFLP analysis of the FD9 DNA fragment permit to readily differentiate all the known FD isolates. Ugni blanc (Trebbiano) and Prosecco are sensitive varieties that may recover when they are protected from new inoculations. Nested PCR and RFLP analysis of selected fragments of the 16S rDNA permit the characterization of any phytoplasma and especially to differentiate FD from BN. has allowed the identification of potential auxiliaries for biological control. which is specific for 16SrV-group phytoplasmas allowed the reliable detection of FD phytoplasma in grapevine. Natural insecticides can be used to preventively limit vector populations but their efficiency is limited in epidemic outbreaks. Control: FD is a quarantine organism in the EC. Molecular detection by PCR of selected phytoplasma DNA fragments. have been found carrying FD phytoplasma until recently. whereas the third treatment. Their rearing is still under development. Detection with laboratory methods is possible on insect vectors and infected vines. titanus in an affected vineyard is an important indication that may lead to suspect a FD infection. Other DNA-based methods. aims at killing newly hatched insects. sensitive amplification with nested PCR of the DNA fragment FD9. Planting of HW treated material is especially important in areas where the vector is present. must be destroyed. control measures are compulsory according to legal regulations. Chardonnay. An indirect DAS-ELISA using coating of wells with rabbit polyclonal antisera and detection with a cocktail of several Mabs. Symptoms are rare in Syrah. The same is true for American rootstocks which can therefore be dangerous sources of infection and dissemination of the disease. Other host plants: No host plants other than Vitis sp. at the beginning of July. 150 . Bois noir (BN) is also frequent in regions inhabited by this leafhopper. Monitoring natural enemies of S. Two additional treatments are applied during summer. Recently. Vitis riparia can be infected but shows little symptoms. Varietal susceptibility and sensitivity: All varieties grown in the various FD-affected regions in France. Detection: FD symptoms can be confused with those of other GYs and also with other diseases or disorders. Sauvignon blanc. titanus. Polyclonal antisera and Mabs have been raised to FD phytoplasma. Planting material must not be collected from mother vines in areas or vineyards affected by the disease. The presence of S. although heavily infected vines can be observed.appear on young vines as late as four years after grafting. such as real-time PCR or identification of PCR-amplified product with DNA-DNA hybridization using specific probes. is directed against winged adults migrating from nearby vineyards or wild vines. at the beginning of August. Roguing is compulsory in France. are currently under development. Other varieties. The first treatment is applied 30 days after first instar emergence at the beginning of the period of potential transmission. roguing of diseased vines is advisable when the epidemic pressure is high. The low rate of transmission by grafting and the long delay in symptom expression make the detection of infected vines in planting material difficult. In France and Italy. Material from mother plants that have developed symptoms in the following growing season. and the risk of disease spreading important. Indirect control is obtained by insecticide treatments against S. Simultaneous amplification in a multiplex procedure of both FD9 and Stol11. Nielluccio and Garganega are very sensitive varieties that usually do not recover after infection.
Branas (a and b): The new disease is thought to be caused by root damage. littoralis in its area of origin in North America.: First report of S. Schvester et al. littoralis recorded from in Italy in 1963. faba plants. littoralis on plants other than the grapevine and transmission trials of FD to other host plants. Schvester et al. Caudwell et al. an insect recently introduced from North America.: Successful transmission of FD to herbaceous hosts by increasing the feeding access period of the vector S. Baggiolini et al. Schvester et al. (a): Control of FD by insecticide treatments against the vector. littoralis. Description of the insect. Description and study of recovery phenomenon and of localised symptoms. Caudwell et al. Caudwell et al. Caudwell: PhD thesis on FD. S. Spontaneous recovery occurs after a 1-2 year crisis period. Boubals and Caudwell: FD epidemics observed in Corsica. No positive results of transmission trials with 15 species of Hemiptera (mainly Cicadellidae) other than S.: Demonstration that FD is transmitted by the leafhopper S. Carle and Moutous: Study on possible secondary or alternative vectors of FD or Bois noir (BN). littoralis. Bonfils and Schvester: Relationships of leafhoppers with FD of grapevine in western France.2. Description of macroscopic and microscopic symptoms. Vidano: Study of the ecology and biology of S. Recovered vines may be infected again but the symptoms are lighter.: Field tests for insecticide control of S. littoralis. biology and feeding damage and of the symptoms of FD in France on Baco 22A. The aetiology is unknown. 1960 1960 1961 1961 1962 1962 1963 1964 1964 1966 1966 1967 1968 1969 1970 1971 1971 1972 151 . littoralis in Corsican vineyards with winter treatment of dormant eggs with oleoparathion. S. littoralis. Caudwell et al. (a). Caudwell and Poitou: Study of the interaction between fanleaf and FD. Study of the role in the aetiology of FD of mycoplasmalike organisms (MLO) observed in insect vectors and in inoculated grapevine and V. Use of 4-5 treatments with DDT or parathion and removal or destruction of pruning wood older that two years. Caudwell: The disease is denoted Flavescence dorée (FD) in agreement with the name proposed by Levadoux (1955) because of the golden yellow metallic aspect of the leaves. (b): Biology of S. Vidano: S. littoralis from Ticino (southern Switzerland). littoralis is present. Caudwell: Study of the phenomenon of recovery of FD-affected vines. littoralis Ball. Historical review 1955 1956 1957 Levadoux: Description of an outbreak of "Flavescence dorée" on Baco 22A in southwestern France. Schvester et al. The disease can be transmitted by grafting and has probably a viral aetiology. evolution of the disease in space and time. littoralis. (a): Possibility to limit the populations of S. considered as a virus disease. Caudwell: Attempts to inhibit the agent of FD in vivo by heat treatment. First report that abandoned or wild vines may be a source of infection.: Studies on the survival of S. Immersion in lukewarm water (30°C) for 72 h of cuttings from FD-affected grapevines reduced the proportion of infected plants by about 83 % as compared with untreated control.
1972 1972 1973 1974 1975 1977 1977 1978 1979 1982
Caudwell et al. (c): Transmission of FD from Vicia faba to V. faba by leafhoppers of the genera Euscelis and Euscelidius and from V. faba to grapevine by S. littoralis. Caudwell et al. (d): Different damage caused by leafhoppers in vineyards and specific control of S. littoralis. Belli et al.: A disease similar to FD observed in vineyards of Oltrepò Pavese since 1968. Caudwell et al.: Research on liquid media to preserve and cultivate MLO agents of FD. Use of the infectivity test to evaluate MLO survival or multiplication. Osler et al.: S. littoralis found in vineyards of Oltrepò pavese affected with a FD-type disease. Caudwell and Larrue: Rearing of colonies of healthy and MLO-infected leafhoppers. Moutous et al.: Results of ovicide treatments against S. littoralis. Caudwell et al.: Successful transmission with S. littoralis of the Corsican GY, thus identified as the same disease as FD. Caudwell and Larrue: The problem of FD in France in relation with sanitary selection of grapevine planting material. Caudwell et al.: Use of serology for detecting FD MLOs by immunosorbent electron microscopy. Antisera raised to MLOs extracted from E. variegatus are used to trap MLOs in extracts from V. faba. Belli et al.: Presence of S. titanus (= S. littoralis) in vineyards of the Veneto (northern Italy) and spread of FD in this region. Anonymous: An important outbreak of FD observed in South France since the beginning of the 1980's. The situation is critical. Belli et al. (a): FD, which was reported in Italy for the first time in 1973, is now spreading northeast in the Veneto region. The vector S. titanus is abundantly present. Belli et al. (b): Problems of identification of FD in Veneto (Italy) and correlation with the presence of the vector. Bagard and Felici: The situation of FD in Corsica is very serious. Boudon-Padieu and Larrue: ELISA detection of FD MLO is possible in leafhopper vectors reared in the laboratory and in S. littoralis individuals trapped in FD-affected vineyards. Positive insects were found in South-eastern France. Kuszala: Injection of MLO-enriched extracts into the body of Euscelidius variegatus is used for evaluating the infectivity of the extracts. Males are better vectors than females. Agulhon and Laurent: Biology and spread of Scaphoideus titanus (= S. littoralis) in vineyards of the south of France. Borgo (a and b): Study on the susceptibility and tolerance of grapevine cultivars in Veneto. Boudon-Padieu et al. (a and b): Development of serological assays (ELISA, Immunofluorescence and Western-blotting) for the detection and characterization of FDantigens in infected leahopper vectors. Caudwell et al.: Present knowledge on the biology, aetiology and diagnosis of FD MLO. Pavan et al.: Dynamics of the populations of S. titanus in Veneto. Planas: Report on field trials to control FD and its vector in South France.
1984 1985 1985 1985 1986 1986
1986 1987 1987 1987
1987 1987 1987
1987 1989 1989
Seljak: Presence of S. titanus in Slovenia (western Yugoslavia). Anonymous: Compared efficiency of various insecticides in the control of S. titanus. Boudon-Padieu et al. (a and b): Polyclonal antisera raised to FD MLO maintained in the laboratory permit detection in infective S. titanus individuals from affected vineyards. Description of the infection cycle of the MLO in the body of the experimental vector E. variegatus. Du Fretay et al.: Satisfactory control of the vector of FD in 1988 in South France. Fortusini et al.: Successful transmission of FD to grapevine cuttings using S. titanus in Italy. Laurent and Agulhon: 1989. Comprehensive review of the situation and evolution of FD and the vector leafhopper in French wine-growing regions. Lherminier et al.: In situ detection of FD MLO in salivary glands of infective Euscelidius variegatus leafhoppers with immunofluorescence. Schwartz: PhD thesis on the production, screening and characterization of monoclonal antibodies to the FD-MLO. Boudon-Padieu et al.: Serological differentiation between FD-MLO and Phy-MLO, an agent causing phyllody in V. faba plants, transmitted in the laboratory by Euscelidius variegatus leafhoppers. Caudwell et al.: Hot water treatment to disinfect grapevine wood from FD-MLO. Lherminier et al. (a and b): The use of immunolabeling in the electron microscope to detect and localize MLO in plants and insects with specific polyclonal antibodies. Arzone et al.: Electron microscopic observation of MLO in sieve elements of Chardonnay and Perera grapevines showing FD symptoms, in clover (T. repens) inoculated with S. titanus fed on infected grapevines and in salivary glands of the leafhopper. Cazenove and Planas: Attempts to obtain a satisfactory control of S. titanus with natural insecticides used in biological agriculture. Anonymous: Recommendations for control of FD in biological wine-growing. Combination of winter treatments with white oil and removing of pruning wood to suppress eggs. Caudwell and Kuszala: ELISA detection of FD MLO in affected grapevines. The method requires extraction of proteins with a strong detergent. DAS ELISA is developed with polyclonal rabbit antiserum as coating antibodies and a cocktail of mouse monoclonal antibodies as detecting antibodies. Daire et al.: Random cloning of FD-MLO DNA and selection of specific probes for detection of FD MLO with DNA-DNA hybridization. Detection in field-infected grapevines is possible only on DNA extracted from a MLO-enriched fraction. Jermini et al.: In spite of the presence in Ticino (Switzerland) of S. titanus in vineyards and nurseries, FD does not seem to occur. Lozzia: A review of the presence, biology and control of S. titanus in Italy. Meignoz et al.: Electron microscopic study of the cytopathogenic effects of FD-MLO in experimentally infected cuttings of LN33. Osler et al.: S. titanus fed on yellows-diseased grapevines in northern Italy (Friuli-Venezia Giulia), test positive to FD-antibodies raised in France. Caudwell et al.: Positive indexing of rootstocks showing latent FD infection and erratic distribution of the pathogen in the material. Sanitation of rootstocks with hot water treatment (50°C for 45 mn).
1989 1989 1989 1989 1989 1990
1990 1990 1991
1991 1992 1992
1992 1992 1992 1992 1993
Daire et al. (a and b): Diversity among MLOs inducing GY diseases in France and other countries, shown with PCR-RFLP analysis of rDNA. FD shows the same pattern as elm yellows MLO. BN is related to stolbur MLO. FD MLO was detected in samples from France and Veneto (Italy). Jermini et al.: Assessment of the optimal number and disposition of sticky traps to evaluate the importance of S. titanus populations in the vineyard. Lefol: PhD thesis on recognition between the FD-MLO and the organs of the experimental leafhopper vector Euscelidius variegatus. Lefol et al.: Development of the "double-dot" method and a derived procedure in the transmission electron microscope to identify the sites of attachment of the FD-MLO on insect organs in vector and non-vector insect species. Maixner et al.: Study of S. titanus in New York and its possible relationship to an American GY. S. titanus transmitted a yellows disease to 29% of Vicia faba plants. FD antibodies raised in France tested positive on 13% of American S. titanus individuals. Seddas et al. (a and b): Immunoaffinity purification of FD-MLO from infected experimental vectors. Carraro et al.: Demonstration with transmission trials using S. titanus, ELISA and PCR analyses, that FD sensu stricto is present in Veneto and not in Friuli-Venezia Giulia, though S. titanus is also present in the vineyards of the latter region. Caudwell et al.: Identification of latent FD infection in several rootstock varieties, including 3309 Couderc and Fercal. The rate of graft transmission to scion depends on the infection status of mother vines and could reach 80 %. Farmer and Boudon-Padieu: Cloning of FD-MLO DNA, construction of an expression library and screening with FD antibodies allowed the selection of three inserts carrying information for FD membrane proteins. Lefol et al.: Route and multiplication of the FD-MLO in the body of experimentally infected Euscelidius variegatus. All main organs were infected, except for germinal cells. Lherminier et al.: Use of ELISA and immunolabeling by transmission electron microscopy for studying the distribution and movement of FD-MLO in the experimental host plant Vicia faba. Seddas: PhD thesis on the production and use of monoclonal antibodies to identify the main antigens of FD-MLO and obtain highly purified fractions of the infectious agent from vector insects and herbaceous host plants. Bertaccini et al.: Presence of phytoplasmas of two types, sometimes in double infections, in severely GY-affected grapevines in Liguria (North-western Italy). One type is related to elm yellows phytoplasma and the other to IPVR phytoplasma, shown later on to belong to the stolbur group. Seddas et al.: Immunoaffinity purified FD phytoplasmas appear as whole cells in ISEM. Experimental vectors (E. variegatus) injected with purified fractions of FD phytoplasma supported its multiplication and were able to feed-inoculate V. faba seedlings. Alma et al.: In Piemonte, several phytoplasmas were identified in yellows-affected grapevines. A 16SrV (= elm yellows) phytoplasma was detected in a double infection with a 16SrI-G phytoplasma. Phytoplasmas of the latter group (later renamed 16SrXII = stolbur) were the more numerous. Bianco et al. (a and b): Elm yellows-related (16SrV) phytoplasmas are present only in the Vicenza and Arezzo provinces of northern Italy. Two different RFLP patterns can be distinguished, suggesting the existence of two strains of FD phytoplasma, one of which is similar to the French strain.
1993 1993 1993
1994 1994 1994
FD is not present in southern Italy. Survey of hemipters in the vineyard and presence of other known phytoplasma vector species.: Monoclonal antibodies raised to immunoaffinity purified FD phytoplasmas. titanus in Italy.: Development of specific PCR primers for the detection of FD or BN phytoplasmas. Control recommendations. Rousseau: A review of different ways to reduce the populations of S.: Important outbreak of FD in Lombardia (northern Italy). Belli et al. The possibility that direct inoculations have occurred in nursery is discussed. Vindimian et al. (a and b): Evolution of FD in the Treviso province (Veneto. Italy). nymphs and adults of S. Clerc et al. the eggs of S. Bosco et al.: Monitoring of leafhoppers in vineyards in Italy. Three different RFLP patterns at least are shown in FD isolates from France. titanus in north-eastern Italy where vines are trained in the pergola system. Transmission by grafting is studied. The possible role of six species in the transmission of GY is discussed. Caudwell et al. Lherminier and Boudon-Padieu: Use of transmission electron microscopy with in situ immunolabeling to detect FD phytoplasma in grapevine and herbaceous host plants. First outbreak in the 1980's on cv Perera.: Insecticide control of S. titanus in biological viticulture. Several different peptides from membrane proteins are identified by Western-blot.: Verification of hot water treatment conditions to suppress FD phytoplasma infection in dormant wood of grapevine. according to the severity of the FD epidemics.: First report of FD outbreak in northern Cataluña (Spain). Mori et al. Marcone et al. Pavan et al. Bianco et al. Seddas et al.1996 1996 1996 1996 1996 Borgo: The first occurrence of FD sensu stricto in the province of Treviso (northern Italy) was reported in the 1980's. Increase and spread to other cultivars. especially Prosecco in the period 1993-1996. Jermini and Baillod: Combination of visual observation of instar larvae and trapping of adults on sticky traps to monitor the populations of S. Kuszala: Specific ELISA procedures to detect FD or BN phytoplasmas in extracts from diseased grapevines from the field.: In spite of the presence and rapid spread of S.: A phytoplasma of the 16SrI (aster yellows) group is detected in eggs. Posenato et al. titanus in the canton of Geneva (Switzerland). titanus laid in the bark were killed.: RFLP studies of 16SrV phytoplasmas. Serological relationships with elm yellows phytoplasma is confirmed Alma et al. In the same conditions. FD has not been identified. The biological significance of this finding is unknown. (a and b): Demonstration of curing effect of hot water treatment on FD-affected plant material. titanus reared on healthy plants. Martini et al. titanus in Trentino (Italy). The amplified FD9 fragment is specific and variable among 16SrV (elm yellows) group phytoplasmas. Daire et al.: Demonstration that two isolates of FD sensu stricto (FD-D and FD-C) are spreading in Veneto (Italy). Infection is rapidly spreading to the east. 1996 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1999 1999 2000 2000 155 . titanus in Switzerland.: First report of S. (a and b): Situation of FD and S. Spain and Italy. Batlle et al. Numerous species identified.
Boudon-Padieu (a): Description, biology, spread, FD-transmission and control of S. titanus. Clair et al.: Design of internal nucleotides on the specific DNA fragment FD9 to be used as primers in nested-PCR for improving sensitivity of detection of elm yellows (16SrV) phytoplasmas in FD-affected grapevine or in elms affected with yellows in France. Cravedi and Aldini: Biology of S. titanus in Oltrepò pavese (Italy), a region affected by a severe outbreak of FD since 1999. Morone et al.: Occurrence of FD in Piemonte (northern Italy) since 1998. Roure: The situation of FD in France. Compulsory control (insecticide sprays and roguing of affected vines) on 300 000 ha. Scattini et al.: Important spread of FD in Lombardia (Italy) in cv Sangiovese. Angelini et al.: Phylogenetic comparison of FD isolates from France and Italy among them and other 16SrV (elm yellows) strains. One isolate from France (FD92 = FD88) and one from Italy (FD-D) are widespread and identical. Bianco et al.: S. titanus in northern Italy can transmit different grapevine isolates of 16SrV (elm yellows) phytoplasmas. Credi et al.: Detection of FD in grapevine samples from the provinces of Piacenza, Parma, Reggio Emilia and Modena in Emilia Romagna (Italy). Davis and Dally: A new 16S rDNA sequence of FD is produced and it is proposed that FD phytoplasmas are placed in two distinct subgroups. However, further work has shown that the 1994 original sequence was erroneous (see Angelini et al., 2003a). Harrison et al.: An elm yellows (16SrV) group phytoplasma close to the Italian isolate FD-C is detected in Virginia creeper plants (Parthenocissus quinquefolia) in southern Florida (USA). It is the first time that a phytoplasma resembling a FD phytoplasma is found in a plant species other than Vitis sp. Marzachi et al. (b): Development of a PCR strategy for large scale detection of FD phytoplasma in plants and insects. Pasquini et al.: Harmonization of procedures for diagnosis of FD in Italy. Quartau et al.: Presence of S. titanus in Portugal. Alma: A review of the distribution and biology of S. titanus in Italy. Bianco et al.: Development of a specific TaqMan® assay for detection of FD phytoplasmas. Borgo and Angelini: Spread of FD in Italy and the situation of mother vines and cultural practice. Boudon-Padieu: Present knowledge on the epidemiology, aetiology and diagnosis of FD. Credi et al. (b): Presence of FD in the Marche region (central Italy). Martini et al.: Further comparisons between FD phytoplasma isolates from France and Italy. Mori et al.: Positive transmission of the two Italian FD isolates (FD-D and FD-C) by S. titanus. Posenato et al.: Efficiency of various insecticides to destroy nymphs of S. titanus and Metcalfa pruinosa. Viggiani: Presence of S. titanus in the Basilicata region (southern Italy).
2000 2000 2000 2000 2001
2001 2001 2001
2001 2001 2001 2002 2002 2002 2002 2002 2002 2002 2002 2002
Angelini et al.: Use of Heteroduplex Mobility Assay and full sequencing on two genomic DNA fragments (ribosomal and non ribosomal) to further study the relationships between FD isolates and related 16SrV (elm yellows) group phytoplasmas. Bianco et al. (b): Presence of FD-D and FD-C isolates of FD phytoplasma in Lombardia (northern Italy). Botti and Bertaccini: Molecular variability between FD phytoplasma isolates using the 16S rRNA gene, the ribosomal protein operon and the FD9 DNA fragment. Distinction between epidemic and non epidemic isolates. Bressan et al.: An attempt to identify a pattern of seasonal transmission of FD, taking into account the transmission efficiency of S. titanus, the pattern of symptom expression by infected grapevines, the emergence of nymphs of S. titanus and the importance of vector population and diseased grapevines. Cavallini et al.: Presence of FD together with BN in the area of Modena (north central Italy). Clair et al.: A multiplex nested-PCR procedure for easy simultaneous diagnosis of FD and BN. The method is now the official method registered in the Official Journal of the French Republic to be used by laboratories in the frame of compulsory control of FD in vineyards. Constable and Boudon-Padieu: Isolation of FD phytoplasma chromosome in Pulse Field Gel Electrophoresis and first physical map for two FD isolates. De Sousa et al.: Identification of FD-D phytoplasma in S. titanus specimen trapped in Portugal. No description of FD-affected grapevines. Duduk et al. (a and b). Identification of FD in southern Serbia and presence of S. titanus. The phytoplasma isolate is similar to FD-C. Malausa et al.: Search for natural enemies of S. titanus in New York with the scope of their introduction in France as biological control auxiliaries. Nusillard et al. Results of a 2-year study in North America of the natural enemies of S. titanus. Santinelli et al.: Presence of S. titanus in Umbria (central Italy). Torres et al. The situation of FD in Spain. Confirmation that only FD-D is present. Angelini et al.: A phytoplasma identical to FD-C isolate is found in wild Clematis in the vicinity of a FD-C affected vineyard in Veneto (Italy). Lessio and Alma: Study of the distribution of S. titanus within a vineyard and use of chromatic sticky traps. Abundance is greater in normal- than in low-density planted vineyards. S. titanus is monophagous. Leafhoppers are not able to spread significantly outside a vineyard. Females are less likely to fly far away than males.
2003 2003 2003 2003 2003 2003 2003 2004 2004
B. BOIS NOIR (VERGILBUNGSKRANKHEIT, LEGNO NERO) 1. Description Bois noir (BN) and related diseases are Grapevine yellows (GY) known in Europe and Asia Minor. The first observations in Eastern France date back to the early 1940's. BN was formally described in 1961 in comparison with Flavescence dorée (FD), because symptoms were similar. It was assumed that the two diseases were different because BN occurred in regions where the vector of FD was not present and experimental transmission with S. titanus was not successful. Because the vector insect, Hyalesthes obsoletus, does not live on vines, affected plants are usually not grouped into patches as with FD, except for situations where a high percentage of affected plants is observed. Nevertheless, distribution is non-random and spread follows a main direction or is preferentially along the border of the plot. BN has been identified in all wine-growing regions of western and eastern Europe, in Israel, Lebanon and recently in the Ukraine.
Main synonyms: The syndrome named Vergilbungskrankheit (VK) (1971) or Schwarzholzkrankheit (SHK) (2003) in Germany, Legno nero (LN) in Italy, Bois noir in Spain, has been occasionally described under the name of Flavescence dorée-like disease. Moreover, it has been sometimes confused with FD in the 1970's and 1980's. It is only since the 1990's that the aetiology was established and that it was shown that the agents of these diseases in the different countries were similar or closely related. Furthermore, the same vector insect species, H. obsoletus, has been identified in all countries, except for Spain where it appears to be very rare. Main symptoms: All the main symptoms typical of GY can be found in BN / VK / LN - infected grapevines. In addition to leaf discoloration, growth abnormalities can be observed on canes and roots. Usually, only a few canes per plant show typical symptoms with either an uneven or a total lack of lignification. On whiteberried varieties, yellow banding develops along the veins which may undergo necrosis. On red-berried varieties, sectorial reddening of blades develops in summer and progressively invades the whole leaf. When berries develop they remain immature, greenish and eventually fall down. Remaining berries are tasteless and sour. Internodes often show longitudinal rows of brown pustules along the green bark of unripe wood. Severely affected vines decline and may die after a few years. Some vines may recover or express irregular symptoms in successive years. Agents: The phytoplasmas associated to these diseases belong to the stolbur group (16SrXII), a group that was for some time considered by some to be the I-G subgroup of the larger Aster yellows group (16SrI). Although stolbur phytoplasmas are ubiquitous pathogens, known to infect solanaceous crops as well as many weeds and bushes, only a limited number of strains have been identified. Hence the different names of BN disease appear to be more geographically than genetically significant. Transmission: Vergilbungskrankheit (= Schwarzholzkrankheit) was the first disease for which the vector was identified. The known vector insect of stolbur of tomato, H. obsoletus (Homoptera, Cixiidae) can transmit stolbur phytoplasma to grapevine, but this occurs during feed probing as the insect does not keep feeding on grapevine. Hence, vine-to-vine transmission does not occur. The role of H. obsoletus has been confirmed in France, in Italy and Israel. In Spain, where the species is rare, its role in BN epidemiology must be verified. Alternative vectors have been suspected but not demonstrated. H. obsoletus is a univoltine polyphagous species that overwinters as larval instars that live on the roots of host plants. Nymph emerge in spring and transmission occurs during adult mating flights. Overall, the presence of insects on vines may last a couple of months. Eggs are laid during summer on the ground at the base of the stem of host plants. Main host plants of the insect vector are common weeds such as Convolvulus arvensis, Ranunculus sp., Urtica dioica, Cardaria draba and Calystegia sepium. However, only C. arvensis and U. dioica were found consistently infected with stolbur phytoplasma, distinct isolates of which were identified in these hosts. From 10 to 80% of trapped populations of the vector may test positive for the presence of stolbur phytoplasma. Transmission by grafting is possible but only at a low rate. Since acquisition by the vector from infected vines does not seem possible, the significance of graft transmission on the spread of the disease is probably very low. Varietal susceptibility and sensitivity: In the different affected regions and countries, numerous cultivars of V. vinifera were found susceptible to the stolbur phytoplasma, cv Chardonnay being the most sensitive. The increasing incidence of BN has been related to the increase of surfaces planted with Chardonnay in the last decade in all countries. Infection of rootstock varieties has not been reported. In all varieties there is a proportion of affected grapevines that undergo transitory recovery or remission, while other plants in the same plot are tolerant or immune to infection. Other host plants: Stolbur phytoplasma is widespread, and has a wide host range. In addition to C. arvensis, U. dioica, Ranunculus sp., C. draba and C. sepium, shrubs and trees of Prunus spp., solanaceaous crops (tomato, tobacco, pepper, egg plant) and lavender (Lavandula spp.) may be significant reservoirs. Detection: The use of monoclonal antibodies raised to stolbur phytoplasma did not show enough sensitivity for routine ELISA detection in field-grown infected grapevines. Detection is achieved mainly with PCR assays or PCR-RFLP analyses of DNA from infected host plants and insect vectors. Most often, nested PCR is necessary for sensitive detection from infected grapevines. Universal primers were first used to detect both BN and FD in areas where the two diseases may coexist. Subsequent RFLP analyses of amplified phytoplasma rDNA permit the characterization of pathogens. Stolbur specific primers have also been designed, either on variable regions of the rRNA gene or on selected specific non-ribosomal
absence of recovery. the multiplex nested-PCR assay cited in the FD chapter can be used. (b): BN is distinct from FD based on non transmissibility by S. Caudwell et al. The species has been described as the vector of stolbur "virus" by Suchov in USSR in 1948. Historical review 1961 1965 1968 1969 1971 1971 Caudwell: Study on BN and its relationships to FD. Control: As with other GY diseases. obsoletus in the south of France. titanus. Description of structures that were probably closteroviruses. littoralis. Leclant: Presence of Hyalesthes obsoletus a cixiid planthopper in the south of France. roguing should be avoided.: GY symptoms are present in Switzerland (Suisse romande and Ticino) but they are not associated with S. Findings never confirmed. for the presence of a FD-like disease. S. Credi and Callegari: Survey of vineyards in Emilia-Romagna. Rumbos et al. Superficial soil cultivation in summer or winter ploughing are methods intended to damage the larval instars and reduce vector populations. Some grapevines show symptoms repeatedly in successive years. Though pruning does not cure infected vines. Combination of RFLP profiles of two DNA fragments allow the identification of three types of stolbur phytoplasmas associated to VK in Germany and hosted by different weeds. titanus is not always associated with the disease.: Rickettsia-like organisms found in roots of VK-affected grapevines in the Saar. Italy. Borgo: General information on the presence of FD-type diseases in northern Italy. 2. Natural enemies are being investigated. Recovery and remission of symptoms for several consecutive years are frequent in vines affected by BN. These findings have never been confirmed. Chemical sprays against H. Hence. For direct differentiation between FD and BN/VK phytoplasmas. Rumbos et al. because of the complex biological cycle and multiple host plants of the insect species. suppression of affected branches may improve harvest quality and help in progressive recovery. The results suggest that the disease is brought into the vineyards from outside local sources. Leclant and Lacote: Study of the ability to transmit stolbur to herbaceous plants by H. except for declining vines.: Rickettsia-like organisms from VK-affected grapevines are cultured in chick embryos. no direct control is possible.DNA fragments. Cazelles et al. and different susceptibility and sensitivity of grapevine cultivars. Nienhaus et al. The disease occurs in the Moselle valley and the Rhine valley. history. Gärtel: Description of VK under the name of Flavescence dorée. 1972 1977 1978 1978 1988 1989 1992 159 . cytological and electron microscope study of tissues of affected grapevine. BN is considered as a non epidemic form of FD. while others seem more tolerant and do not show symptoms. (b): Epidemiological observations of BN lead to the assumption that an aerial vector is involved. Mosel and Rhine valleys are considered as the causal agents of the disease and it is hypothesized that the nematode Xiphinema index is the vector of the disease. Caudwell et al. unknown at that time. Mendgen: 1971.: Rickettsia-like organisms detected in phloem and xylem of roots of grapevines affected with VK in West Germany and in nematodes of the species Xiphinema index feeding on roots of these grapevines. Description of symptoms of VK. obsoletus are not efficient.
obsoletus is a vector of VK in Germany. and from Israel. Maixner (b. (a): Differentiation by PCR-RFLP of a MLO related to aster yellows in GY-diseased plants from Lombardia and a MLO related to elm yellows in an affected grapevine in FriuliVenezia Giulia (Italy). Bianco et al. the most frequent belonging to group 16SrI. Cazelles and Kuszala: Negative results with FD-ELISA on all tested GY-affected plants from Switzerland suggest that the GY present is BN rather than FD.: First detection of BN (stolbur phytoplasma) in Spain. subgroup I-G (later named group 16SrXII (stolbur) in the provinces of Vicenza. 1993 1993 1993 1993 1993 1993 1993 1994 1994 1994 1995 1995 1995 1996 1996 1996 1996 160 . (b): Evidence of an association of MLOs with grapevine yellows in EmiliaRomagna. H. Several weeds are host plants of the stolbur phytoplasma. Daire et al. (a and b): PCR amplification of MLO DNA shows that MLOs associated with GY in several regions of northern Italy are related to the Italian periwinkle virescence (IPVR) MLO. Bianco et al.: The agent of a GY disease in the Italian Riviera is detected with a probe specific for Eastern aster yellows. c and d) Transmission of a yellows to periwinkle with dodder in Germany. obsoletus identified as the vector.: One of the two phytoplasmas detected in yellows-diseased grapevines in Liguria is related to IPVR phytoplasma. from several regions of Italy. subgroup I-G (later renamed as 16SrXII or stolbur group). Maixner et al. PCR-RFLP analysis confirms relationship to aster yellows phytoplasmas. obsoletus.: Grapevines from Piemonte (northern Italy) contain several different phytoplasmas. Biology of the insect.1992 Fos et al. Reservoirs and possible role of other vectors remain to be elucidated. Laviña et al. MLO associated to GY in Italy are related to aster yellows MLO. aetiology and transmission of BN in France. (a): Development of stolbur-specific primers on phytoplasma rDNA and use for detection of stolbur phytoplasma in grapevines affected with VK and in the vector H. Davis et al. Italy. clustered by the authors into aster yellows group but shown later to belong to the stolbur group. The latter MLO was shown later to belong to the stolbur group. Maixner et al. Alma et al. It is suggested that vectors with preference for other host plants act as vectors for VK. The MLO strains found in grapevine are related with aster yellows MLOs. Boudon-Padieu (b): History. Davis and Prince: According to a different terminology. Bertaccini et al.: A monoclonal antibody raised to stolbur mycoplasma-like organism is used for ELISA detection of the agent in herbaceous host plants and insects. but are different from known strains of MLOs of the aster yellows cluster. Computer analysis for spatial patterns of spread shows a non-random distribution and suggests preferential transmission from other vines or from weeds. Italy) where FD is prevalent. obsoletus is confirmed as a vector of stolbur disease plants in France. Brescia and Pavia (northern Italy). (a and b): PCR-RFLP analysis of 16Sr DNA of MLOs show that the MLO associated with BN is different from the FD MLO and similar to the MLO associated with stolbur disease. Minucci et al. H. Maixner : Demonstration that H. BN MLO is detected in grapevines from France. Borgo: Presence of BN in mixed infections with FD in the province of Treviso (Veneto.: PCR-RFLP analysis of MLO-DNA detected in VK-affected grapevines in Germany show that this MLO is related to stolbur. Bertaccini et al. (b): Presence of phytoplasmas in the group 16SrI.
(a and b): The identification with PCR-RFLP of rDNA of phytoplasma detected in European and Israeli grapevines affected by GY has shown that BN occurs in all regions. subgroup I-G (identified later as belonging to the stolbur group or 16SrXII).1996 1996 Koruza: Survey of GY in three regions of Slovenia shows that the disease is of BN type. Maixner: The situation of VK in Germany: symptoms.: Occurrence of GY diseases in Hungary and identification of stolbur phytoplasma in affected grapevines. a strong annual increase. Kölber et al. using hot water treatment with conditions recommended in France. obsoletus and its role as the vector of VK. Osler et al. a high proportion of previously infected vines that did not show symptoms for 2 years at least and. First detection of BN agent in diseased grapevines in Switzerland with the same procedure. Maixner and Reinert (b): Spatio-temporal analysis of the distribution of VK in 4 vineyards in the Moselle an Rhine valleys over a 2-3 year period showed a high incidence. Estimated crop damage ranges from 20 to 40 %. (c): Identification of stolbur phytoplasma (16SrXII) as agents of GYs in Greece and Israel. Search for variability among BN phytoplasmas with RFLP analysis of the specific non ribosomal DNA was unsuccessful. (a and b): BN is poorly transmitted by bench-grafting. Maixner and Reinert: A review of the research and present knowledge on VK and other GY diseases.: Prevalence of BN (stolbur phytoplasma) in Navarra and Cataluña (north eastern Spain). titanus. arvensis is recommended to expose instar larvae and kill them with frost. Refatti et al. as well as phytoplasmas transmitted by Credi to periwinkle in Emila Romagna and previously classified in the aster yellows group. Sforza and Boudon-Padieu: Description of H. Ploughing in winter of soils infested by C. Results were satisfactory. obsoletus as a vector of BN. Kuszala: Detection in ELISA using a monoclonal antibody to stolbur phytoplasma of BNassociated antigens in diseased grapevines in France. obsoletus in Germany. Vindimian: LN but not FD is present in Trentino (northern Italy) in spite of the presence and active spread of S.: Monitoring of field populations of H. Marcone et al. search for vectors. a high proportion of vines in which symptoms re-occurred after one season or more. 1996 1996 1996 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1998 1998 1998 1998 161 .: Phytoplasmas associated to GY in Campania (southern Italy) are genetically uniform and belong to the stolbur group. Preferential spread along the rows.: Important spread of GY symptoms on several grapevine varieties in Umbria (central Italy). Weber et al. Daire et al. Maixner and Reinert (a): Trials on elimination of VK infection from vine wood in Germany. conversely. biology of H. Laviña et al.: Consistent association of stolbur phytoplasma with GY diseases in the Italian provinces of Fiuli-Venezia Giulia and Trento and in Slovenia. Sforza: PhD thesis on the epidemiology of BN in France. The maximum delay of symptom expression in young plants is 2 years. Davis et al. vector transmission and possible control measures. obsoletus and possible control measures. Weber: PhD thesis on the biology of the cixiid H. Albanese et al. Characterization of a phytoplasma belonging to group 16SrI.
obsoletus and Metcalfa pruinosa were found positive for BN phytoplasma (16SrXII-A). the vector of Palatinate grapevine yellows (PGY). Up to 30 % of the insects were infected and the highest rate was in vineyards with high populations of Convolvulus arvensis. Acquisition may occur at larval stages since emerging 5th instars were infective. obsoletus as a vector to grapevine. Seljak and Petrovic: A review of the Slovenian situation of GY diseases. Varga et al. High rate of BN / LN infection in the different vine-growing regions. and a high proportion of vines that escaped infection during the 3-5 year period of monitoring. No other phytoplasma found in the South of the country. The same pathogen detected in weeds and other crops. Females are more often infected than males. The rate of infected H. Bourquin et al. Identification of main reservoir weeds. regardless of the cultivar in Campania (southern Italy).: Epidemiology of BN in France. obsoletus. Šeruga et al. 1998 1998 1998 1999 1999 2000 2000 2000 2000 2000 2000 2000 2000 2000 2000 162 . Maixner and Reinert: Collection of H.: Widespread presence of LN in Toscana (central Italy). Darimont and Maixner: Comparison of the transmission efficiency of phytoplasma to their host plants and to grapevine by H.: BN reported only in north-western and eastern vineyards of Croatia. obsoletus.: Use of PCR to confirm the presence of stolbur phytoplasma in GY-affected grapevines in Switzerland. Phytoplasmas in the 16SrXII group (stolbur) were the most frequent. PCR detection is reliable when testing together 25 insects among which only one is infected. obsoletus and possibility to control the insect in vineyards. The study confirms a high rate of infected H. the second GY disease in Germany. formerly reported in western Europe.: First detection of stolbur phytoplasma in grapevines in Croatia. First launching of colonies of the insect under controlled conditions. the ability to recover of cvs Riesling and Pinot in spite of systemic symptoms.: Ethological onservations and morphological descriptions of adults and larvae of H. Škoric et al. Larrue et al. obsoletus can be high because of the presence of reservoir weeds in the vineyard. Evidence secured for the occurrence of tolerance (no symptom at all for 15 years) and temporary remission (no symptoms for a few years) .: Only stolbur phytoplasma detected in grapevines showing GY symptoms.1998 1998 Sforza and Bourgoin: Anatomy of reproductory organs and mating of H. Transmission to natural host plants is higher than for grapevine with both insects. Confirmation of the role of H. H. Sforza et al. obsoletus in vineyards is efficient with sticky traps placed at the soil level. Braccini et al. Maixner et al. No other grapevine phytoplasma was detected.: VK disease dramatically increased in incidence in the Moselle and Rhine valleys during the 1990's. Weber and Maixner (b): Etholology of H. Braccini and Pavan: Report of survey of Auchenorrynchas in vineyards of Toscana (central Italy). Weber and Maixner (a): Procedures for the survey of infection status of populations of H. different phytoplasmas were identified in a number of cultivars. obsoletus related to VK infection. Sforza et al. obsoletus and Oncopsis alni. Identification of two infected symptomless weeds in the vineyard. obsoletus. among which hoary cress. Marcone et al.: Several vineyards affected by BN in Burgundy (France) monitored for over 15 years and assessed for symptom severity. Up to 34% of insects were infected with stolbur phytoplasma.: In a large survey of vineyards in 8 Hungarian counties conducted in 1998.
All affected grapevines and two naturally infected periwinkles contained the same stolbur isolate. obsoletus and Circulifer haematoceps were widespread and positive for stolbur or aster yellows phytoplasmas. obsoletus. Morandell: Detection of VK in south Austria. Gugerli et al. the incidence of VK was significantly higher in conventional vineyards. Study of the spatial and temporal dispersion of the four species. dioica) as a reservoir of stolbur phytoplasma and as a host for the insect. Pentastiridius sp. obsoletus to the German viticulture. Kölber et al.: Anoder cixiid planthopper. H. Same conclusions about tolerance and transient recovery of grapevines. Laviña et al. Indicators based on climatic parameters permit to predict the flight period.: Only BN identified in Switzerland on cvs Chardonnay. Langer et al.: Observation of the occurrence of BN in Trentino in eight Chardonnay vineyards on more than 11. BN is the main disease and cvs Chardonnay and Zweigelt appear to be the more sensitive among whiteand red-berried varieties.2001 Bertamini and Nedunchezhian: Evaluation of effects of BN disease (stolbur phytoplasma) in field-grown grapevines.: Assessment of the best period for detection of BN in affected grapevines. Merlot and Pinot noir. obsoletus.: Demonstration that H.: The highest risk of phytoplasma contamination for grapevines coincides with the flight activity of adults of H. obsoletus and the presence in organic vineyards of several weeds known as host plants carrying larvae on the roots. on pigments. Alma et al. general stress responses and rapid senescence.: Incidence of BN in a small area of the Bekaa valley (Lebanon) on cvs Chardonnay and Alicante Bouschet. Bertaccini et al. respectively. Cicotti et al. Darimont and Maixner: Importance of the presence and infectivity of H. Attempt to bring up the larvae to the surface of the soil by ploughing so that they are killed by frost in winter is under evaluation. Neoaliturus fenestratus. Gatineau et al. Samples were taken on all organs of 10 affected grapevines. Detection in December was the more constant.: Survey of potential vectors of phytoplasmas in vineyards of the Golan Heights.: First detection in Albania of BN phytoplasma in grapevines of different age and varieties. Severity of symptoms vary from one year to the other. reached by other authors. Myrta et al. obsoletus is the vector of LN in Italy. obsoletus in Lebanon is recorded.: Survey of GY from 1997 to 2002 on 21 cultivars in 12 Hungarian counties.: Importance of outbreaks of BN in the province of Modena (Italy) and monitoring of potential insect vectors in addition to H.: The situation of GY diseases in Croatia shows that only BN is present in the northern and eastern vineyards. very similar to European isolates. Mescalchin and Mattedi: A description of the development of VK in Trentino (northern Italy). Orenstein et al. In spite of a similar abundance of H. Maixner et al. The authors conclude that phytoplasma infection induces non specific. while Megophthalmus scabripennis was positive for aster yellows phytoplasma.: Comparison of infection risk by VK in organic and conventional vineyards in Germany. chlorophyll fluorescence and photosynthetic activities suggest that Photo System II is affected.000 plants over 12 years. The presence of H. Choueiri et al. 2001 2001 2001 2002 2002 2002 2003 2003 2003 2003 2003 2003 2003 2003 2003 2003 163 . Curkovic Perica et al. every month for one year except in winter. Role of stinging nettle (U. which is widespread in the province. is a natural vector of stolbur phytoplasma to sugarbeet in Bourgogne (France).
H.: Search for vectors of stolbur phytoplasma in Cataluña (Spain) where H. Sabaté et al. Description Palatinate grapevine yellows (PGY) is a disease occurring in Germany in Pfalz and Moselle regions (west of Germany) mainly on cv Scheurebe. Šeruga et al. Germany). Main synonyms: FD-Pfalz was a first name because the agent was shown to be related to the FD phytoplasma. Transmission: Transmission was obtained with the leafhopper Oncopsis alni Schrank (Hemiptera.: Important expression of BN in cv Chardonnay in the Ukraine. Alder is considered as the source of PGY infection. titanus leafhoppers have failed. together with high populations of H.: Important epidemics of "Schwarzholzkrankheit" (SHK = VK) in Franken (Bayern. It is not epidemic and occurs most often in old vines in vineyards along creeks or rivers where alders (Alnus glutinosa L) are present. Another alder insect. According to the data of field survey. Cicadellidae) trapped on alders and caged on V. Transmission to plants is not reported. alni is highly monophagous and specimen do not survive long on grapevine. was found carrying the alder phytoplasma at a higher rate than O. Milkus et al. (a): Investigation on the variability of stolbur (16SrXII) phytoplasmas from extracts of different grapevines in Croatia. the psyllid Psylla alni. Main symptoms: Symptoms are similar to those of other GY diseases and may affect the whole plant. Control: No possibility of control because of erratic transmission by the vector. specific association of each isolate with a different weed is discussed. and rarely in groups. It is associated to phytoplasmas in the Elm yellows (16SrV) group. alni but failed to transmit both to alder and grapevine. Langer and Maixner: Three different types of stolbur phytoplasma isolates found in grapevines. Varietal susceptibility and sensitivity: PGY occurs in limited areas. obsoletus were positive for stolbur phytoplasma. Gilge et al. using 4 fragments of phytoplasma DNA showed that all isolates are similar. Symptoms were recorded on 26% of 30-year old Scheurebe grapevines and 96% of tested individuals of H.2003 2003 Petrovic et al. Three isolates (PGY-A. 164 . Šeruga et al. The disease was identified in 1995. 2003 2003 2004 2004 2004 C. More specific characterization can be obtained with EY-group specific primers fAY/rEY that amplify a 16S rDNA fragment or with FD9 primers that amplifiy the non ribosomal FD9 DNA fragment. Several hemipters have been found to deliver stolbur phytoplasma to the medium. affected plants are not numerous. Agents: The agent is an EY (16SrV) group phytoplasma. The latter results have not been published. Attempts to transmit PGY phytoplasma from grapevine to grapevine using reared S. occur most often at the border of plots. Living insects checked for transmissibility of phytoplasma by feeding on artificial medium. RFLP of FD9 fragment permit to distinguish PGY phytoplasmas from FD phytoplasmas and from other phytoplasmas in the same group (see the Flavescence dorée chapter). PGY phytoplasma is similar to the Alder yellows phytoplasma that is widespread in black alder in Germany. It is different from FD phytoplasma. Other host plants: Alder trees (Alnus glutinosa L. However. O. Trapping of insects on sticky traps or with D-Vac suction. arvensis and U. obsoletus is rare. (b): First identification of BN infection in the Macedonian republic. dioica. -B and C) have been identified.: Importance of a GY epidemics in the Drava region (Slovenia) in 2001 and 2002. vinifera seedlings that later developed GY symptoms.) Detection: Universal primers and RFLP analysis of amplicons can be used to identify a 16SrV phytoplasma. PALATINATE GRAPEVINE YELLOWS 1. mainly on cv Scheurebe. obsoletus and weeds in different areas in Germany. obsoletus and frequence of C.
Reinert and Maixner: All the 3 strains of PGY were detected both in alder trees and in specimen of O. Characterization of the three PGY strains showed that they were closer to FD than to strain EY1 from American elm. (b): Transmission to V. alni (Auchenorrhyncha. Reinert: PhD thesis on detection. alni trapped on alders. alni) is able to transmit the alder yellows phytoplasma to healthy seedlings of alder. The proportion of infective leafhoppers is lower with O. titanus living mostly on wild Vitis riparia around affected vineyards were found to carry an agent that was serologically related to that of FD. a phytoplasma belonging to the X-disease group (16SrIII) was detected in affected grapevines in New York. In the same period. alni. The risk of infection by PGY is lower than for VK because reservoir plants are fewer and outside vineyards. Extensive survey of yellows-affected grapevines in several areas of Germany did not show the presence of other EY group phytoplasmas. obsoletus. In 1993. Historical review 1995 Maixner et al. both insect species trapped on alder. Maixner and Reinert: Demonstration that O. using Heteroduplex mobility assay and sequencing of rDNA and FD9 fragment. (b): Detection of a new EY-related phytoplasma in GY-affected cv Scheurebe in Germany. NORTH AMERICAN GRAPEVINE YELLOWS 1. vinifera seedlings of the PGY phytoplasma using specimen of O.: Comparison between Elm yellows group (16SrV) phytoplasmas including PGY strains. Same RFLP profiles of P1-P7 rDNA amplicons obtained from yellows affected grapevines. American grapevine yellows. 165 . die back of shoot tips and abortion of developing fruit. molecular characterisation and epidemiology of PGY. Angelini et al. respectively. Main synonyms: Leaf curl and berry shrivel (LCBS).2. The differences in vector specificity between PGY and FD phytoplasmas are not supported by molecular data. Description Grapevine yellows (GY) were first reported in New York State (USA) in 1977. alder. which was associated with two different phytoplasmas in group 16SrIII and 16SrI (aster yellows). populations of S. alni and H. Variability analysis and sequencing of the FD9 DNA fragment and homology of the sequence to a part of the SecY gene. Maixner et al. Reinert and Maixner: 1997. In 1991. Oncopsis alni and Psylla alni specimen. The strong adaptation of O. affected vines often do not survive winter frost because they lack reserves. a serious outbreak of GY occurred in Virginia. Cicadellidae) (but not P. Darimont and Maixner: Comparison of the transmission efficiency of phytoplasma to their host plants and to grapevine by O. the latter findings were not confirmed. the vector of BN / VK. 4 Italian and French strains of FD and elm and alder phytoplasmas. Identification of the PGY-associated phytoplasma with phytoplasmas infecting alder in the vicinity of affected vineyards. Angelini et al. However. The alder phytoplasma resembles ALY. The PGY strains are closest to the ALY phytoplasma from alder in Italy and to one strain of FD.: Further comparison between elm yellows group (16SrV) phytoplasmas. It is different from FD phytoplasma but molecular and serological data show a close relationship. a phytoplasma isolate transmitted from alder to periwinkle in Italy. alni to alder prevents a frequent inoculation to grapevine even if the infestation of the leafhopper population is high. In New York. Infected grapevines often die within months from the onset of symptoms and significant losses of vines have been observed. Main diseases: Virginia grapevine yellows I (VGYI). As much as 5 % of leafhoppers from Palatinate and 15 % from Moselle tested positive. 1997 1999 1999 2000 2000 2000 2001 2003 D. alni. Virginia grapevine yellows III (VGYIII) Main symptoms: North American Grapevine Yellows (NAGY) is a lethal disease causing yellowing of the leaves.
observed in 1983. Larvae are more frequent on wild Vitis riparia at the border of the vineyards. However. very similar to those of LCBS. Extracts of periwinkles were used to raise two monoclonal antibodies and to design oligonucletides that were used as probes in DNA-DNA hybridisation or as PCR primers. 1985 1991 1993 1993 1993 1993 166 . In Virginia. The Virginia Grapevine Yellows (VGY) has been associated with the VGYIII phytoplasma (a strain of X-disease or 16SrIII. ELISA and immunfluorescence were positive from periwinkle.Agents: The 16SrIII phytoplasma detected in NY has not been further characterized. the survey of vineyards and transmission assays to V. present in New York and its possible relationship with a GY disease. Symptoms. Data showed that the Italian and American MLO were related. No transmission by vine planting material reported. Probes and primers yielded positive reactions using DNA extracted from grapevine. However. The X-disease phytoplasma occurring in New York was also detected with monoclonal antibodies and with probes for dot-blot hybridization. Varietal susceptibility and sensitivity: In New York. Historical review 1977 Uyemoto et al. vinifera. Pearson et al. Daire et al. X-disease phytoplasma was detected in native Vitis sp. then on White Riesling. Symptoms may occur for several years in succession in the same vines. Detection was positive in symptomless wild Vitis riparia growing near vineyards in New York and in some symptomatic grapevines. faba bean plants and feeding solutions. titanus. Chen et al. Sauvignon blanc and Cabernet franc. titanus in New York and transmission assays of a possible infectious agent.: Monitoring of S. Adults migrate from V. as well as direct PCR assays from insects. (a): A Riesling plant from New York and a periwinkle carrying the FDI MLO described above. Transmission: No vector insects have been identified for VGY phytoplasmas. X-disease phytoplasma was detected in native Vitis sp. Prince et al. subgroup I-A). Other host plants: Both phytoplasmas are widespread in the USA in weeds and cultivated species. Maixner et al.: Molecular comparisons between MLOs transmitted to periwinkle from grapevine or detected in a few naturally affected grapevines. The latter observation suggests that another non-related MLO might have been present. among which Agallia constricta. The FDI MLO was identified in a parallel work as a member of the X-disease group. and aster yellows was detected from numerous weeds and shrubs within and around the vineyard. vinifera cuttings. Maixner and Pearson: First data on the study on S. and an aster yellows phytoplasma was detected in numerous weeds and shrubs within and around the vineyards. It appeared to be spreading and was perhaps insect borne. the disease was first observed on cv De Chaunac. it is very severe on Chardonnay vines but was observed on other varieties including Riesling.: Occurrence of FD-like symptoms on White Riesling grapevines in New York. other grapevines with strong symptoms tested negative. Control: There are no control methods since vectors and reservoirs of the associated phytoplasmas are still undetermined. subgroup III-B) and with the VGYI phytoplasma (a strain of aster yellows or 16SrI. tested positive for a MLO in the X-disease group with PCR-RFLP of 16S rDNA.: Description of a disease of the yellows type affecting the variety De Chaunac in vineyards of New York State denoted Leaf curl and berry shrivel (LCBS).: Two MLOs were transmitted from affected grapevines to periwinkles in Udine (Italy) (= FDI or FDU or GYU) and New York. allowed the identification of a few candidate vectors of the aster yellows phytoplasma. Detection: With molecular assays using universal or group-specific PCR primers. show that the FDU (= FDI) strain from Italy and a MLO detected in a grapevine in Virginia (USA) are both related to X-disease. riparia to V. 2.
5%) with the phytoplasmas associated to Papaya die back (PDB) in Australia and to Phormium yellow leaf (PYL) in New Zealand. riparia vines. In addition. In this particular disease. there are reported cases where they were not associated with AGY disease or phytoplasmas. waxy appearance and remain rubbery. which is the more frequent and the Tomato big bud (TBB) phytoplasma. Mapping of diseased plants show that grapevines appear to be a terminal host of AGY. Early and recent surveys have shown that AGY are found in most viticultural regions of Australia. Agallia constricta is a suspected vector of VGYI (aster yellows phytoplasma). two syndromes with particular symptoms (Restricted Growth and Late Season Leaf Curl) have been found in some cases. with a higher incidence in warmer inland districts of New South Wales and South Australia. The association of AGY with phytoplasmas was confirmed as early as 1988. a GY disease found in Victoria (Australia). The TBB phytoplasma has been designated as Candidatus species "Candidatus Phytoplasma australasia". leaves tend to fall early. Davis et al. respectively. 1998 2003 E. 167 . The second phytoplasma. AUSTRALIAN GRAPEVINE YELLOWS 1. one node after the other. suggesting that the source is from an alternative host. also detected in wild V. Description Grapevine yellows were first reported from Australia in 1976 and called Australian grapevine yellows (AGY) in 1983. Conversely. Some plants also display uneven bud development resulting in canes and cordons that are bare in places with little or no bunch development. Beanland and Wolf: Search for possible insect vectors and reservoirs of NAGY phytoplasmas in Virginia. Shoots may display abortion of bunches or berry shrivel later in the season and death of terminal buds. Affected vines carry one or more shoots with irregular veinal of interveinal yellowing and downward rolling of the leaves that overlay one another in a shingled fashion. It belongs to the peanut witches' broom group (16SrII) and has a broad plant host range in Australia.: Two phytoplasmas are detected in yellows diseased grapevines in Virginia. is a member of the aster yellows (16SrI) group. Late Season Leaf Curl (LSLC). III-I. Main symptoms: All symptoms typical for GY can be found on AGY-diseased plants. BVGY phytoplasma has not been clustered in an established phytoplasma group. Agents : AGY disease (sensu stricto) is associated with two phytoplasmas: the AGY phytoplasma. Restricted Growth (RG). Main diseases: AGY (sensu stricto). Stems of affected shoots develop a blue. Confirmation that a X-disease (16SrIII) group phytoplasma is involved and classification of the latter phytoplasma into a new subgroup. Restricted Growth (RG)-affected grapevines show retarded growth resulting in shortened shoots and smaller leaves.: A GY disease observed in Virginia since 1983 affects large areas with a low incidence and a low spread rate. It has been suggested that all three phytoplasmas are strains of the same phytoplasma species. The AGY phytoplasma has been designated as Candidatus species "Candidatus Phytoplasma australiense". consistently associated with AGY symptoms and AGY phytoplasmas but their aetiology is not clear. subgroup I-A. overlay one another like shingles and become leathery and brittle. It belongs to the stolbur group (16SrXII) but is distinct from the BN and VK phytoplasmas. The yellow leaf tissue can become necrotic. The use of PCR has shown the association with AGY of three different phytoplasmas. clover phyllody phytoplasma (16SrI-C) being the closest . The sequence similarity of the 16S rRNA gene shows that it is close to members of the aster yellows group (16SrI).1993 Wolf et al. The 16S rRNA gene has a high sequence similarity (more than 99. The relatedness of the associated MLO to X-disease MLO is confirmed. Both VGYI and VGYIII were found in several weeds and shrubs around and inside the vineyards. Buckland Valley Grapevine Yellows (BVGY). Other species tested positive and transmitted to feeding solutions. Late Season Leaf Curl (LSLC)-affected grapevines show symptoms in late summer with tightly downward rolled leaves that often remain green. followed by the progressive death of the shoots.
140-510 nm in diameter. Other host plants: As mentioned above. this phytoplasma is associated with GY disease in vineyards in the same restricted grape-growing areas that were established with planting material from different sources. Reduction of symptom expression in vines injected with tetracycline. suggesting that this contributes to the decline in phytoplasma numbers and to the remission of symptoms.: MLOs. Magarey and Wachtel: GY symptoms are found in most viticultural areas of Australia. branches. Phytoplasma australasia). once vines are infected by AGY. RG and LSLC diseases has been observed. AGY phytoplasmas are common in several crops in Australia. argentatus. It is possible that AGY diseases have become prevalent in Australia in the last years with the increasing planting of Chardonnay. Hence.and white-berried varieties. Similarly. Cixiidae) is known as the vector of PYL disease in New Zealand but the species has not been reported from Australia. and trunks in October is more reliable.: The "Rhine Riesling problem" denoted "Australian Grapevine Yellows". Detection can be from shoots. The AGY phytoplasma was detected in the common leafhopper Orosius argentatus (Evans) but transmission to grapevine has not been reported. Magarey et al. branches. argentatus and subsequently transmitted to faba bean but the ability to transmit to grapevine was not confirmed. The incidence of disease may be temporarily high in some vineyards of Chardonnay. suggesting that the disease is the result of an aerial transmission and that the phytoplasma was not present in the original planting material. Sanitation with hot water treatment is being developed in Australia. Remission of AGY. vector of TBB in tomato crops. followed by RFLP analysis or sequencing. Phloem fluorescence of diseased vines in the UV microscope. Several observations suggest that aerial transmission prevails. The TBB phytoplasma is transmitted to other crops by O. Control: There are no methods to control AGY diseases in the vineyard. The BVGY phytoplasma has no suspected insect vector. is found in high numbers together with two other potential MLO vectors in vineyards of Victoria (Australia). Detection: Detection is obtained with PCR. The planthopper Oliarus atkinsoni Myers (Hemiptera. Magarey: The situation of GY in Australia. they are at greater risk of displaying symptoms again in the following years. Varietal susceptibility and sensitivity: Chardonnay and Riesling appear to be most often affected. All generic "universal" primers may be used. found in sieve tubes of diseased vines are associated with AGY symptoms. A "heat curing" effect of AGY disease by hot weather has been observed. 2.: Orosius argentatus. a specific primer in the intergenic 16S-23S spacer region has been designed for detection of TBB phytoplasma (Ca. Phytoplasma australiense) either in single-step or nested PCR following a first amplification with universal primers. reservoirs must be important but the epidemiology is not fully understood. Recent studies have shown that the latter phytoplasma can be acquired from infected grapevine by O. However. However. RG or LSLC.Transmission: No vectors have been identified for any AGY disease. Sampling simultaneously from shoots. but phytoplasmas were also detected in other red. trunks and roots throughout the year and infection is persistent from year to year. indicating persistence of the disease. Historical review 1982 1983 1985 1986 1986 1988 1988 1989 Magarey and Wachtel: Description of a new disease of the yellows type in cv Rhine Riesling in South Australia. Magarey: Comprehensive review of GY diseases in the world. Magarey et al. Magarey and Wachtel: Review of the AGY problem. such as Shiraz or Semillon. Osmelak et al. Specific primers have been designed on the 16S rRNA gene for detection of AGY phytoplasma (Ca. 168 .
The problem of symptomlessly infected vines. Uneven distribution of phytoplasmas in infected plants. (a and b): The phytoplasma associated with AGY is related to an aster yellows type phytoplasma associated with GY in Italy (later on identified as the stolbur phytoplasma).: Comparison of visual assessment and diagnostic assays. Papaya dieback and Phormium yellow leaf diseases. Constable et al. insecticide sprays are useless for controlling the disease. BVGY might form a new AY subgroup of AY (16Sr) group or even a new phytoplasma group. RG. Wilson et al. Constable and Symons: AGY phytoplasma is detected in most symptomatic grapevines but also TBB phytoplasma and a third uncharacterised phytoplasma. established from a different source of plant material indicate that BVGY disease occurs as a result of aerial transmission. It is occasionnally detected in symptomless vines and in vines with LSLC or RG symptoms.1995 1995 1996 Bonfiglioli et al. The TBB phytoplasma is detected occasionnally in vines with AGY symptoms and also in LSLC-affected vines. RG and LSLC are associated. resulting in increasing cumulative incidence.: The AGY phytoplasma is consistently present in the majority of symptomatic grapevines. Bonfiglioli et al. (a and b): Positive detection of phytoplasma with PCR in AGY-affected grapevines and tentative survey of phytoplasmas in Australia.: Close relationships between phytoplasmas associated with AGY.: Detection of phytoplasmas associated with AGY. Detection of BVGY in another plot in the same area. Davis et al. Candidatus Phytoplasma australiense. Constable: PhD thesis on the biology and epidemiology of AGY phytoplasmas.: TBB phytoplasmas can be acquired from diseased grapevine by Orosius argentatus and subsequently transmitted to faba bean. Constable et al. Beanland et al.: Identification of a new phytoplasma in Chardonnay grapevines in Victoria (Australia). Waite et al.: No vector for AGY found. LSLC would be followed by AGY on the following year and AGY would be followed by RG in subsequent years.: AGY phytoplasma detected in the body of Orosius argentatus leafhoppers. Kelly et al. Padovan et al.: Further comparison of AGY phytoplasma with overseas GY phytoplasmas show that it is closely related to. Liefting et al. Padovan et al. but different from the stolbur phytoplasma associated with BN in France. Spain and Israel. Constable et al. Gibb et al. and LSLC.: Study of the phylogenetic relationships of the BVGY phytoplasma with aster yellows and stolbur phytoplasmas. 1997 1997 1997 1998 1998 1998 1999 1999 1999 2000 2000 2001 2001 2002 2002 2002 169 .: Description of the AGY phytoplasma and designation of a new taxon. Beanland et al. Spring and summer are optimal for detection. later called BVGY phytoplasma.: Use of hot water treatment in commercial nurseries of Australia. Habili et al.: BVGY phytoplasma is associated to GY symptoms that may show remission then reoccurrence but also appear on vines that were not affected previously.: Description of AGY symptoms. Hence.: Assumption from field observations and monitoring that AGY. Constable et al.
The third situation is represented by poorly characterized GYs recorded from new countries. Constable et al. Varietal susceptibility and sensitivity: Chardonnay. Description Besides the cases reported above. In three instances. recurrence in other vines and new occurrence in previously unaffected vines. 2004 2004 F. a 16SrI-B phytoplasma was found in mixed infection with a stolbur phytoplasma (at that time designated 16SrI-G). The first is the detection in countries other than the USA.: Survey of AGY. The second situation is the significant occurrence of X-disease group (16SrIII) phytoplasmas in countries other than the USA. However. OTHER GRAPEVINE YELLOWS 1. It seems that aster yellows phytoplasmas have no important significance in European grapevine yellows. is often reported in connection with these cases. The total cumulative incidence for AGY could be as high as 90%. 2. of phytoplasmas belonging to the aster yellows group (16SrI) in yellows-affected grapevines.2002 2003 Crocker et al. At other times of the year. to a lesser extent. The subgroup of aster yellows was not determined but primers specific for stolbur tested negative. No variability was observed amongst BVGY phytoplasma isolates. detection is more reliable from samples taken from branches and trunks. three different situations must be described. Constable et al. later designated 16SrXII. However. a variety widely grown in all countries and very sensitive to all GY agents. They must not be confused with the 16SrI-G phytoplasma of earlier reports. RG and LSLC symptoms for 3 to 6 years in Chardonnay and Shiraz vineyards. Other varieties are cited in the different situations. In Chardonnay. namely Israel. sometimes as a second phytoplasma in co-infection with FD or BN phytoplasmas and sometimes also in symptomless grapevines. they seem to have some relevance in Israel and were recently reported from Tunisia. Saric et al. There is no information of transmission of AGY by planting material. TBB phytoplasma may have a role in the aetiology of LSLC. These aster yellows phytoplasmas belong to subgroups I-B or I-C (clover phyllody) which are ubiquitous in Europe. X-disease group phytoplasmas. Constable and Symons: Study with Heteroduplex mobility assay (HMA) of the elongation factor Tu (Tuf) gene of the AGY and the BVGY phytoplasmas. Historical review Europe 1996 Alma et al. The AGY and the TBB phytoplasmas may persistently affect grapevines throughout the year and between years. Transmission: Aster yellows and. the diseases are characterized by remission in some plants. The incidence of AGY in a plot of cv Shiraz that had been treated with hot water suggests that AGY phytoplasmas are introduced to vineyards by aerial transmission. Two isolates of AGY and Papaya dieback (PDB) phytoplasma did not show variability between them but showed some variability when compared to a reference AGY phytoplasma. 1997 170 .: Survey of phytoplasmas infecting grapevine in northern Italy. which was the first designation for stolbur phytoplasma in some laboratories. Statistical analysis showed that RG and LSLC diseases were not always associated with AGY. are very frequent in many host plant species and several insect species are known as vectors or potential vectors in different countries. (a and b): Biology and epidemiology of AGYs. Symptomatic shoots from AGY affected vines are reliable for detection in summer (January).: The management of the area for plant material in the nursery with the scope of hot water treatment.: Detection of phytoplasma in the aster yellows group in GY affected plants in Slovenia and Croatia.
Detection with PCR show erratic identification of a 16SrI-B phytoplasma. obsoletus was found in addition to the preceding 5 species. a X-disease phytoplasma was detected in some Neoaliturus specimen. Gajardo et al. 2003 Israel 1997 2000 Tanne and Orenstein: Grapevine phytoplasmas in Israel were transmitted to periwinkle with heterografting and identified in the periwinkle as 16SrIII and 16SrI phytoplasmas. Neoaliturus sp. The spatial and temporal dispersion of the four species was analysed. H. In addition to similarities of symptoms with FD. Caudwell et al.: Experimental transmission of the Chrysanthemum yellows phytoplasma (16SrI-B) to grapevine by four leafhopper species. Similar transmission of clover phyllody (16SrI-C) MLO by S. Trapping of hemipters and identification of numerous potential phytoplasma vectors.: Further survey of GY vectors in the Golan Heights. Orosius orientalis. Circulifer spp. some affected vines showed a sudden fall of leaves in summer. titanus. Klein et al. Orenstein et al.: A 16SrI-C phytoplasma was found associated with GY in relevant percentages in the same vineyards as stolbur (BN) phytoplasma in Abruzzo region (eastern central Italy). H.2001 Alma et al. Transmission to periwinkle of yellows diseases using collected insects. titanus had been obtained in 1971 in France (cf. and Anaceratagalia laevis were all found carrying an aster yellows phytoplasma.: Survey over several years of potential vectors of phytoplasmas in vineyards of the Golan Heights showed that Neoaliturus fenestratus. including S. 2001 2003 Tunisia 2003 Chabbouh et al. of phytoplasmas belonging to group 16SrI. In addition. Megophthalmus scabripennis was positive for aster yellows phytoplasma.: Formal identification of a 16SrI-B phytoplasma in GY-affected grapevines in Tunisia.: Identification in yellows-affected vines of varieties Petit Syrah.: First results on the monitoring of GY diseases in Tunisia in imported table grape varieties. obsoletus and Circulifer haematoceps were all three abundant and positive for stolbur and aster yellows phytoplasmas. M'hirsi et al. 1971b). Stolbur phytoplasma (16SrXII) was also detected.. Tanne et al. 2003 South America 1980 Caudwell: Description of a disease of the yellows type in Chile. called "Amarilliamento de Elqui". Macrosteles sexnotatus. D'Ascenzo et al. Merlot and Carmenere.: Monitoring of potential phytoplasma vectors in vineyards in Israel and molecular detection of phytoplasmas in their body. 2003 171 . subgroups I-B and I-C and also to group 16SrVIII (Ash yellows) in mixed infection with I-B.
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