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Sections

  • PREFACE
  • INTRODUCTION
  • INFECTIOUSAGENTSOF GRAPEVINES
  • GRAFT INCOMPATIBILITY
  • FLECKCOMPLEX
  • VIRUS-LIKEDISEASES
  • GRAPEVINEYELLOWS: INDIVIDUALDISEASES
  • GRAPEVINEYELLOWS: REFERENCES

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

Opinions, data and facts exposed in this number are under the responsibility of the authors and do not engage either CIHEAM or the Member-countries.

Les opinions, les données et les faits exposés dans ce numéro sont sous la responsabilité des auteurs et n'engagent ni le CIHEAM, ni les pays membres.

Martelli. Boudon-Padieu 2006 .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.

September 2006 tel. Italy. +39 080 542 45 87 e-mail: ideaprint@virgilio.it Directory of Infectious Diseases of Grapevines and Viroses and Virus-like Diseases of the Grapevine: Bibliographic Report 1998-2004 Editors: G. Martelli. E.P. Série B : N. 55 ISSN : 1016-1228 ISBN : 2-85352-338-1 © CIHEAM.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. Boudon-Padieu Bari : CIHEAM (Centre International de Hautes Etudes Agronomiques Méditerranéennes) p. 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» . 279 Options Méditerranéennes.

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 .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 .

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whose Secretatiat he has admirably conducted since its very establishment in 1962. virus-like and phytoplasma diseases of grapevines constitute a major limiting factor to the development and well-being of the world viticultural industry. Boudon-Padieu and the latest Bibliographic Report (1998-2004) on “The Viroses and Virus-like Diseases of the Grapevine” by R. in the belief of rendering a good service to the scientific community and. whose publication IAM-B is happy to support. a distinguished virologist of worldwide reputation and one of the father founders of ICVG. is therefore a goal of utmost importance to which the Mediterranean Agronomic Institute of Bari (IAM-B). The present issue of Options Méditerrnéennes. Martelli and E. except when virus-free vines are grown for comparison alongside with non-sanitized sister vines.PREFACE Virus. whereas E. Furthemore. 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. Italy 7 . by and large. Special care was taken in the organization of the subject matter. produced under the auspices 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. As a whole. hosts a new edition of the “Directory of Infectious Diseases of Grapevines” by G. to all those interested in viticultural matters. grouping diseases and setting them in a well thought out order. being a recognized authority in this field.P. Cosimo Lacirignola Director of IAM Bari. ICVG has a long-lasting tradition in these endeavours. although the technology is now available for detection and elimination of most of the pathogens occurring in propagation material. As to the authors. these diseases cause loss of vigour and often a decline of affected stocks. taking also into account the future role of the Mediterranean as a free-trade area. 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. Notwithstanding these efforts and the promoting activity of the International Council for the Study of Virus and Virus-like Diseases of the Grapevine (ICVG). students. it is most unfortunate that dissemination of infectious diseases continues. so as to produce a document which can readily be used by scientits. Bovey The “Directory” is a work of great thoroughness and accuracy. which reflect on the commercial value and productive life of the vineyards. is contributing with direct involvement in international research projects and through the activity the Mediterranean Network on Certification of Grapevines. The “Bibliographic Report” is another most precious achievement by R. they reduce graft compatibility of scions and rootstocks. technicians. to which IAM-B was happy to contribute. Improving the sanitary conditions of the industry. promoting the publication of the Bibliografic Report 1985-1997 on virus and virus-like diseases of the grapevine. up to date as to the time of publication. that fosters coperative studies in grapevine virology. Boudon-Padieu has a long lasting experience in the study of phytoplasmas. Bovey. 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. Sincere thanks are espressed to the authors of these contributions. and practitioners. or induce subtle debilitating effects which are difficult to percieve.P. G.

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Boudon-Padieu INRA Dijon.International Council for the Study of Virus and Virus-like Diseases of the Grapevine (ICVG) DIRECTORY OF INFECTIOUS DISEASES OF GRAPEVINES G. France 2006 . Italy E. Martelli University of Bari.P.

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2003. Infected propagating material is largely responsible for the spread of diseases among and within viticultural countries. in turn. Options Méditerranéenes. which were compiled under the auspices of the International Council for the Study of Viruses and Virus Diseases of the Grapevine (ICVG). attracting the attention of scientists. and P. To this effect. 2006. A bibliographic report 1979-1984. as well as on the quality and quantity of the yield. Paris.. Bovey R.. 3rd part).. 1986. Bibliographie de 1965-1970. nurserymen. grapevines are exposed to the attacks of a variety of pests and pathogens among which infectious intracellular agents (viruses. Vitis 25. 3rd part). 227-275. Options Méditerranéenes. Office International de la Vigne et du Vin. The increased attention paid to grapevine virological problems and the like. 10-172. and endangering the survival of affected vines. Vitis 18. A bibliographic report 1985-1997. The papers up to 2004 are listed and commented upon in six bibliographic reports: Caudwell A. Thus. that has been especially active at the international scale from the late 1950's onwards. ICVG has been instrumental in fostering basic and applied research in grapevine virology. A bibliographic report 1971-1978. growers. Locorotondo 2003. 11 . has produced an impressive series of papers which now number about 5. The importance of the grapevine industry and the magnitude of the problems caused by these pathogens has generated wide interest which. W. 1965. Bibliographie des viroses de la vigne des origines à 1965. causing heavy losses. The viroses and virus-like diseases of the grapevine. 1979. Les virus de la vigne. having a negative impact on the plant vigour and longevity.. and a highly valuable agricultural commodity. and G. 1999. A bibliographic report 1998-2004. Bovey R. its 14th Conference having taken place in Italy in 2003 (Bovey and Gugerli. 1-2. Martelli. 1972. 2003). The viroses and virus-like diseases of the grapevine. viroids. and phytoplasmas) recorded from the grapevine can be highly detrimental to this crop. recognizes that a number of the 60 or so infectious agents (viruses.400. Caudwell A. Hewitt and R. Since its foundation. ICVG has issued the recommendations that follow: The International Council for the Study of Virus and Virus-like Diseases of the Grapevine (ICVG). As most of the vegetatively propagated crops.B. From the very beginning.INTRODUCTION The grapevine (Vitis spp. Gugerli. 29 (Series B. ICVG has met regularly every 3 to 4 years. 303-324. Bovey. and supporting initiatives for the establishment and implementation of clean stock and certification programmes. Bovey R. and R. The viroses and virus-like diseases of the grapevine. viroids. Vitis 11. 76 pp. Hewitt W. and administrators on the detrimental effects of infectious diseases on the well-being of the industry. 205-279. Bovey R. 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. all efforts should be made to improve its sanitary conditions.P. A short history of ICVG.. among other things. has fostered intensive research. xx (Series B. shortening the productive life of vineyards. The viroses and virus-like diseases of the grapevine. 316-376.B.) undoubtedly represents one of the horticultural crops most widely grown in temperate climates. phloem-and xylemlimited prokaryotes) play a major role. Bovey. Extended Abstracts 14th Meeting of ICVG.

a moratorium will be established for these viruses. Minnesota. Virus and Virus-like Diseases of Grapevines. Compendium of Grape Diseases. 181 pp. Uyemoto. Portugal ) The International Council for the Study of Virus and Virus-like Diseases of the Grapevine (ICVG). No other stock should move outside regulatory regions. and A. R. Certified selections should be tested for specific pathogens. high standards are essential to ensure that no viticultural area is compromised by the introduction and spread of diseases. rugose wood. rugose wood (GVA.C. 1978. In addition. G. as well as on the quality and quantity of the yield.K. W. Goheen and H. Their elimination from mother vines intended for propagation should therefore be pursued. Hewitt.G. and fleck. Inadequate certification standards have repeatedly resulted in disease problems for growers and nurserymen. G. Paul. W.F. Set 1 (O. leafroll. Martelli. Class 1 should include only grape nursery stock which tests negative for the most damaging diseases/pathogens. 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.K. The certification process should specify conditions to prevent and detect subsequent infection of nursery plants by regulated pests. the virological problems of grapevines have been extensively treated and illustrated in a number of books and major review articles: Uyemoto J. valuable grape genetic resources exist which are infected with virus but are essential to the preservation of world viticultural heritage. that enables vineyards to economically and sustainably maintain quality and productivity. Wilcox. The American Phytopathological Society Press. Gärtel. and 3). St.K. is regarded as incompatible with an accepted sanitary status.D. Thus. Dias. As efforts are made to harmonize grapevine certification protocols. However. Lisbon. regional viticultural conditions. Goheen. Woodham. In order to preserve valuable grape clones and varieties. Martelli and A. The regional certification standards for Class 2 stock should be created at a local level based on the rate of endemic infection. GVB and GVD). Vuittenez. Infected propagation material is largely responsible for the spread of diseases among and within viticultural countries. Barnett and S. It would move freely between regulatory boundaries. In the future.. viroids and phytoplasmas). and the need for preservation of heritage germplasm. Lausanne. and permit tracing the certified grapevines to the originally selected and tested plants. (issued and approved in 1997 at the 12th ICVG Meeting.C. G.P. all efforts should be made to improve its sanitary conditions. Martelli and A. clonally selected primary sources. including the causal agents of fleck and rupestris stem pitting. Certification of grapevine nursery stock is a powerful and effective tool to control these agents. and phytoplasmas (flavescence dorée. Rowhani will be published in 2005 in the series APS Plant Virus Image CD-ROM. They represent a most valuable source of information. A. and other grapevine yellows). The agents that should be controlled by the Class 1 certification program are those associated with infectious degeneration and grapevine decline (nepoviruses). Gubler and J. Editions Payot. under the name of Grapevine Viruses. we propose two sanitary classes. Clemson.P. 12 .A. 100 slides. (issued and approved in 2003 at the 14th ICVG Meeting. Until that time. leafroll disease and associated closteroviruses (grapevine leafroll associated viruses 1.B.W. ensure clonal integrity. Locorotondo. technology should make it possible to exclude additional disease-causing viruses from the certified stock. 1988. Tolin.P. 29 pp. Grape Section.The presence of diseases such as infectious degeneration. (a revised edition by J. 2. bois noir. Italy) The Proceedings of all the international ICVG Conferences have been published in extenso or. which are best performed in the framework of certification programmes encompassing also clonal selection. Bovey R. Virus-like Diseases and Other Disorders). having a negative impact on plant vigour and longevity. 93 pp. Grapevine (Vitis) virus and virus-like diseases. Plant Virus Slide Series. W. lately. many of which can be highly detrimental to this crop. recognises over 70 infectious agents affecting grapevine (viruses. Improvement of the sanitary level can be achieved through selection and sanitation. Pearson R. as Extended Abstracts.C. eds) Clemson University. Uyemoto will be published in 2005). (a 2nd revised edition edited by W.F. USA. 1980. Certified grapevines are derived from pathogen tested.

1993. Hervieu. INRA Editions. R. Food and Agriculture Organization of the United Nations. 1996. Walter B. Bulletin de l'OIV 70. University of Bari. 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.A. Martelli and E. Rezaian and R. Hadidi. Taylor. Bovey and G. Director of the Mediterrenean Agronomic Institute of Bari for supporting the publication of this work. C. Editions Féret.H. St. American Phytopathological Society Press. Hamze and Mr. Giovanni P. Martelli. 2000. 5-23.137 pp.K. E.A. Protocols for detection of viruses and virus-like diseases: Les Colloques. Influence des viroses et qualité. 1997. In: Plant Virus Disease Control (A. Rome.P. a body now estinguished. CSIRO Publishing. Lacirignola. Martelli. Martelli Professor of Plant Virology. Sanitary selection of the grapevine.Université de Bourgogne Dijon. Paris. sélection pomologique. Elisabeth Boudon-Padieu Directeur de Recherches Head Biologie et Ecologie des Phytoplasmes Plante Microbe Environnement INRA . Martelli and published under the auspices of the Mediterranean Fruit Crop Improvement Council (MFCIC). n° 86. Walter B. Martelli G.P.. Notwithstanding this wealth of published information a "Directory of Major Virus and Virus-like Diseases of Grapevines" was compiled in 1992 by R. (ed. France 13 . and G. Bari.P. 2ere partie: Sélection sanitaire. and G. N.Frison E. B. Martelli G. They express their deep gratitude to Prof. Rome. respectively. Paul. Ridé. 945-971. bactéries et phytoplasmes de la Vigne.P. Sélection sanitaire de la vigne: sélection sanitaire et sélection pomologique. Italy.R. Influence des viroses et qualité. Bulletin de l'OIV 69.). 54 pp.P. 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. 192 pp. Paris. M. Walter. Ikin. Krake L. 261-276. (ed). 1999. Walter B. 225 pp. eds. and R.P. Detection and Diagnosis of Graft-transmissible Diseases of Grapevines.). of the Centre International des Hautes Etudes Agronomiques Méditerranéennes. Koganezawa. Sélection sanitaire de la vigne: sélection sanitaire et sélection pomologique. 1991. Virus certification of grapevines. Maladies à Virus. the Steering Committee of ICGV decided to update the Directory and to entrust this task to G. FAO Publication Division. 1997. Boudon-Padieu and M. Graft-transmissible Diseases of Grapevines. Walter B. Boudon-Padieu.S.CNRS . and to Dr. 1998. At the 14th ICVG Meeting.. Collingwood. Bordeaux. FAO/IBPGR Technical Guidelines for the Safe Movement of Grapevine Germplasm. Chairman of the Board of Directors and Secretary General. 1ere partie: Effets des viroses sur la culture de la vigne et ses produits. Rome /International Board for Plant Genetic Resources. M. Khetarpal. Scott. 263 pp. and B. H.

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This represents the highest number of intracelluar pathogens ever found in a single crop. The viral scenario of Vitis: viruses and their taxonomic affiliation(a) FAMILY GENUS SPECIES A. phytoplasmas (8). and insecttransmitted xylematic bacteria (1) have been recorded form grapevines. viroids (5).INFECTIOUS AGENTS OF GRAPEVINES More than 70 infectious agents among viruses (58). 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 .

U. C. Virus Taxonomy.A. and Ball.. J.. Maniloff. (a) 16 . Mayo. Viruses belonging to unassigned genera Idaeovirus Sadwavirus Sobemovirus Tobamovirus C. London.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. Elsevier/Academic Press.A.M. The names of tentative species are written in Roman characters. 8Ith Report of the International Committee on Taxonomy of Viruses.. Desselberger. 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. (eds). The updated taxonomy of all classified grapevine viruses can be found in: Fauquet. M. pp. 1258. L. 2005.

INFECTIOUS DEGENERATION .

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The consensus is that fanleaf degeneration may have existed in the Mediterranean Basin and the Near East since the earliest time of grape cultivation. records of this disease date back some 150 years. showing abnormal branching. Now the disease is known to occur worldwide. and inflorescences). reduced rooting ability of propagation material. low proportion of graft take. the yellowing fades rapidly and the canopy develops a normal green color. Leaves are variously and severely malformed. Reisigkrankheit (partly). deep lobes. sometimes appearing as rings or lines to extensive mottling of the veins and/or interveinal areas to total yellowing . Symptomatic leaves show little malformation. Fruit set is poor. Main synonyms: court-noué. however. 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. shortened productive life. phloem. parenchyma. Trabeculae. that give the leaf the appearance of an open fan. showing progressive decline. double nodes. dégénérescence infectieuse (Fr. and berries ripen irregularly. arricciamento. especially in the basal internodes. Gelbmosaik (Germ. i. mosaico giallo. Chromatic alterations of the leaves vary from a few scattered yellow spots. Yellow mosaic is induced by chromogenic virus strains. but bunches are small and few. chlorotic mottling may accompany foliar deformations. These structures are readily visible by light microscopy in lignified shoots.INFECTIOUS DEGENERATION (GRAPEVINE FANLEAF VIRUS) Several nepoviruses infect grapevines in Europe and the Mediterranean area. Occasionally. or endocellular cordons. Description Fanleaf is the oldest known and one of the most important and widespread virus disease of the grapevine. may show open petiolar sinuses. The foliage and shoots show little if any malformation. roncet. More recently. are diagnostic of grapevines infected by GFLV. puckered. and grapevine leaves with typical symptoms were found in herbaria established before the introduction of American rootstock hybrids. which exhibit widely open petiolar sinuses and abnormally gathered primary veins.). asymmetrical. degenerazione infettiva (Ital. With increased ambient temperatures during summer. fasciations. a disorder induced by Grapevine fanleaf virus (GFLV). are small-sized and set poorly. low yields and low fruit quality. and acute denticulations. urticado (Port. Tolerant cultivars produce fairly good crops whereas sensitive cultivars are severely affected.and zigzag growth. Main symptoms: Two distinct syndromes caused by different strains of the causal agent characterize this disease.).e. GFLV and several of the other grapevine-infecting European nepoviruses have distorting and chromogenic strains and may occur in mixed infections. another disease sometimes occurring in vineyards affected by infectious degeneration. bunches are straggly. tendrils. The characterizing symptoms of "Vein banding". causing degenerative diseases whose symptoms are similar to. Shoots are also malformed. Bunches are smaller and fewer in number. panachure. and the yield may be much reduced. In the European literature. and decreased resistance to adverse climatic factors.).). The foliage develops bright chrome yellow discolorations early in the spring that may affect all vegetative parts (leaves. Infectious malformations are induced by virus strains causing distortions. Foliar symptoms develop early in the spring and persist throughout the vegetative season becoming less distinct in summer. vein banding symptoms were shown to be caused by a co-infection by Grapevine yellow speckle viroid and GFLV. The name of this virus comes from the peculiar malformation of infected leaves. and xylem cells. FANLEAF 1. 19 . Their economic impact varies with the tolerance of the cultivar to the viruses. radial bars crossing the lumen of epidermal. This disorder was first reported from California as a syndrome elicited by a specific strain of GFLV. Often infected grapevines occur in patches. or indistinguishable from those of fanleaf. shoots. short internodes.

but is not a specific test. reorganization. consisting of two functional molecules 7342 nt (RNA-1) and 3774 nt (RNA-2) in size. or by somatic embryogenesis. encapsidated in different particles. amaranticolor. The best antigen sources for serological diagnosis are leaves collected in spring or cortical shavings from mature dormant canes. but it is still regarded as necessary for confirming freedom from virus infection. Virus particles are often within tubular structures that accumulate in bundles in the cytoplasm or nucleus. 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. Chenopodium quinoa. Observation of symptoms in the field is useful as a first step in selection. use of fumigants is more and more questioned for environmental reasons and is being progressively banned. Transmission by Xiphinema italiae has not been consistently documented. However. Some V. For transformation. GFLV was the first grapevine virus to be recovered by mechanical inoculation and to be thoroughly characterized physicochemically and molecularly. quinoa. Resistance to X. The virus occurs in the pollen of infected grapevines and herbaceous hosts. amaranticolor. mannose and xylose. in a persistent manner by the longidorid nematode Xiphinema index feeding on the roots of grapevines and retaining the virus for several months. serologically rather uniform and occurring as a family of minor molecular variants. vinifera x M. where they occur in a radial orientation. Control: Use of virus-tested scionwood and rootstock material in the framework of clean stock or certification programmes. Detection: ELISA using polyclonal antisera and monoclonal antibodies is a quick. by in vitro meristem tip culture. rotundifolia can be infected by GFLV when graft inoculated. Detection of trabeculae can give information on the health of American rootstocks. and soybean. and very sensitive method. which are desirable as they cause no harm to human health. both required for infectivity. but is not reliable. rotundifolia hybrid rootstocks (e. GFLV can be transmitted by mechanical inoculation from infected grapevine tissues to various herbaceous hosts (e.g. but show few symptoms. index is determined by the viral coat protein. 20 . C. C. Virus elimination is readily achieved from vegetating shoot tips by heat treatment (38-40 °C for as little as four weeks). Molecular assays using radioactive or digoxigenin-labelled probes. the endosperm of grapevine seeds. RT-PCR is estimated to be four to sixfold more sensitive than ELISA.g. with variable levels of sensitivity. cheap. In contaminated soils. In the laboratory. index (10 individuals) by ELISA and in single nematodes by RT-PCR and immunosorbent electron microscopy. RT-PCR and immunocapture RT-PCR are becoming increasingly popular. Indexing on Vitis indicators by grafting takes a lot of time and field or greenhouse space. There are conflicting reports on seed transmission in grapevines. 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. rotundifolia hybrids is thought to be controlled by a single dominant gene. American rootstocks are also susceptible and are generally very sensitive. and is transmitted through seeds of C. Muscadinia (Vitis) rotundifolia and Vitis munsoniana are highly resistant to X. index feeding. vuittenezi has been suspected but not proven. Work is under way in different laboratories to create GFLV-resistant rootstocks or cultivar through traditional breeding methods or genetic transformation technology. rupestris x M. This resistance is controlled by two unlinked recessive genes. However. These inclusions derive from membrane proliferation. index in V. Positive sense single-stranded RNA genome. a number of selectable marker genes toxic to non engineered vines are used. but resists infection when the virus is transmitted by the nematode. A satellite RNA 1114 nt in size is associated with some virus isolates. Nematode populations transmit local virus isolates with a higher efficiency than those from other geographical areas. Natural GFLV infections have been detected in weeds in Hungary and Iran. However. Varietal susceptibility: Almost all known Vitis vinifera L. Gomphrena globosa). the use of fumigants against nematode vectors gives only a temporary but economically valuable control of the disease. Dissemination over medium and long distances is through infected vegetatively propagated scionwood and rootstocks. Specific transmission by X. vinifera. GFLV has been detected in small groups of viruliferous X. Transmission: At a site. M.Agent: Grapevine fanleaf virus (GFLV) is a nepovirus with polyhedral particles of about 30 nm in diameter. and redistribution and are thought to be sites of viral polyprotein processing and RNA replication. O36-16) show interesting levels of field resistance to GFLV. Cytopathology: GFLV elicits the formation of intracellular cytopathic structures known as vesiculatevacuolate inclusion bodies which are often apposed to the nucleus. and transmission by X. are toxic to many plants but not to V. Indexing on herbaceous hosts by mechanical inoculation requires climatized greenhouses and is less reliable than ELISA. varieties are susceptible. although some like Vitis labrusca can be infected.

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. Petri: Grapevine "arricciamento" (fanleaf) has a viral origin Arnaud and Arnaud: Hypothesis of a viral origin for grapevine court-noué (fanleaf). research and hypotheses on fanleaf. For a comprehensive review on early observations. Tome 1: Les maladies dues à des végétaux (champignons. see the book by Galet.2. 1977.b Hewitt: Fanleaf and yellow mosaic recorded from California. Montpellier. 3. Historical review From the late 1800 to 1997. bactéries. Les maladies et les parasites de la vigne. Arnaud: Court-noué is considered as a soil-borne virus disease. 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. 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. 871 pp. With roots of fanleaf-infected grapevines with phylloxera feeding on them. 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. Healthy rooted cuttings or seedling of Rupestris du Lot were contaminated: 1. 1977(b) 1865 1882 1895 1896 1902 1906 1910 1912 1912 1917 1918 Cazalis-Allut. Report on first experiments on heat treatment of grapevine in order to eliminate fanleaf. 1954 1958 Hewitt: Review on grapevine virus and virus-like diseases found in California.. France. Imprimerie du "Paysan du Midi". 1929 1931 1937 1937 1946 1950a. With soil containing phylloxera. (a) See references in the Introduction.: Hypothesis that court-noué (fanleaf) is caused by a virus transmitted by phylloxera. Hypothesis about a possible role of phylloxera as a vector. No conclusive results were obtained. Branas et al. Hypothesis that fanleaf is a fungal disease. but only circumstantial evidence. With individual phylloxera (radicicolous or gallicolous) fed on infected vines. 2. (b) Galet P. viroses et phanérogames).: Experiments on the capacity of phylloxera to transmit fanleaf. 21 . Bovey: Review on grapevine virus and virus-like diseases. No direct proof of transmission by this aphid. as well as on controversies about transmission by phylloxera. the ICVG Bibliographic Reports (a) have recorded more than 1000 papers dealing with fanleaf. Branas et al.

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. including fanleaf virus: bentonite flocculation test. whereas insecticide treatment has no effect. Reproduction of fanleaf symptoms in mechanically inoculated grapevine seedlings 1964 1964 1965 Taylor and Hewitt: Description and characterization of Australian isolates of GFLV. 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. 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. Dichloropropane-dichloropropene (DD) at 1000 l/ha gave satisfactory results. Serological relationship with ArMV reported. index. Vuittenez: Mechanical transmission of fanleaf virus to Chenopodium quinoa and C.: Transmission of GFLV by Xiphinema italiae in Israel. amaranticolor is confirmed. Transmission of fanleaf virus by X. Control of the vector by soil fumigation. index in France. Cohn et al. Hewitt et al. Bercks: Research on the use of three serological methods for detecting plant viruses. Boubals and Dalmasso: Experiments on soil disinfection against X.1958 1958 1960 Vuittenez: Fumigation of fanleaf-contaminated soil with nematicides prevents infection of healthy grapevines replanted immediately. Gifford and Hewitt: Use of heat therapy and in vitro shoot tip culture to eliminate fanleaf virus from infected grapevines. Das and Raski: Studies on the relationships of GFLV with its vector X. index. Goheen and Hewitt: Description of vein banding as a GFLV-induced disease Dias: Host range and properties of fanleaf and yellow mosaic viruses.: Transmission of fanleaf virus from grapevine to herbaceous hosts by mechanical inoculation and preliminary characterization of the virus. Dias and Harrison: Relationships between the viruses causing fanleaf. Hewitt et al. Gerola et al. Description of the chipbudding method for indexing.: Description of the Davis method of heat therapy of grapevines. yellow mosaic and ArMV 1960a 1960b 1961 1961 1962 1962 1963 1963 1963a. Goheen et al. Vuittenez: New observations on the effects of soil fumigants on fanleaf in contaminated soils.b Martelli and Hewitt: Comparative studies showed that Californian and Italian GFLV isolates are the same.: Fanleaf virus is transmitted by the nematode Xiphinema index Cadman et al.: Detection of GFLV particles in thin-sectioned grapevine root tissues. 1965 1967 1968 1968 1969 1969 1970 22 . index occurred during the 6 year period of observation.: Investigations on grapevine virus diseases in California. Reproduction of fanleaf symptoms in mechanically inoculated grapevine seedlings is confirmed Galzy: Heat treatment of grapevine plantlets grown aseptically in vitro. latex test and barium sulfate test. and no reinfestation by X. Bercks and Querfurth: Use of latex-test for detecting GFLV and other nepoviruses in grapevine tissue extracts in Germany.

The sensitivity of the latex test is increased. Raski and Schmitt: Progress in the control of the fanleaf-nematode complex by soil disinfection with 1. George for detecting GFLV. Alfaro and Goheen: The different strains of fanleaf virus (fanleaf sensu stricto. PALLAS is quicker and cheaper than ELISA. The acquisition time threshold is less than 5 minutes. Uyemoto et al. George.: Detailed physico-chemical characterization of GFLV. Querfurth and Paul: Protein A-coated latex-linked antiserum (PALLAS) method for detecting GFLV and other viruses.: Use of green grafting as a quick and secure method for graft-indexing. 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. A dormant bud of the variety to be cured is grafted onto a healthy potted rootstock. Hévin et al. rupestris is more accurate than mechanical transmission to C. Walter et al. Mission or LN 33.: 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.: GFLV particles observed in the lumen of the oesophagus of X. St. but ELISA is more sensitive. Taylor: Review paper on grapevine vein banding. rupestris St. quinoa. especially with low titre antisera.1970 1970 Hewitt et al. quinoa. yellow mosaic and vein banding) are transmitted in the same way by X. the plant is placed in a heat cabinet for treatment Hévin et al. 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. During the moult of the nematode. Vuittenez: Review paper on grapevine fanleaf. Both tests are more sensitive than mechanical inoculation to C. Vineyards replanted in contaminated but treated soils remained healthy for at least 5 years. 1970 1970 1970 1971 1971 1972 1972 1973 1973 1973a 1973b 1974 1974 1975 1976 1976 1977 1979 1979 23 . this lining is shed and ingested in the intestine. After bud take.: Control of fanleaf by soil fumigation with 1.: GFLV and marbrure (fleck) are not transmitted through the seeds of grapevine. Both methods give satisfactory and similar results. anterior oesophagus and oesophageal bulb of their nematode vectors. Bercks: Serological detection of grapevine viruses in West Germany.3-dichloropropene or methyl bromide.: Comparison of indexing by mechanical inoculation to Chenopodium quinoa and by graft-transmission to V. Goheen and Luhn: New method of heat therapy.3 dichloropropene or methyl bromide. Indexing by budding on V.: Comparison between PALLAS latex test and ELISA for detecting GFLV in France. index. Raski et al. Raski et al.: Heat therapy of grape plantlets grown in vitro causes changes in some characteristics of the variety. Van Velsen and Niejalke: Green budding for indexing grapevine with the indicator cvs. Mur et al. Dias: Review paper on grapevine yellow mosaic.

Russo et al.3-dichloropropene (1. A Middle Eastern V. Bouquet: Serological detection of GFLV in its vector X.: Study on the effectiveness of soil fumigation for the control of X. Savino et al.: Detection of fanleaf virus in grapevine tissues by ELISA and ISEM in different periods of the year. Even in the cases where the virus was transmitted. in California.3-D) or methyl bromide applied 75-90 cm deep with 90 cm spacing for 1. The use of methyl bromide requires a continuous cover with polyethylene sheeting for some time after the treatment. 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. index feeding. Raski et al. index. vinifera accession represent an excellent example of host plant resistance to GFLV. vuittenezi.: Identification of a natural serological variant of GFLV from Tunisia Huss et al. a very common species in vineyards of Rheinhessen and Palatinate.: Identification of several Vitis species and interspecific hybrids resistant to fanleaf virus. Carbon disulfide gave less satisfactory results.: 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. as vector of GFLV. Brown and Roberts: Detection of fanleaf virus in its vector X. index. Methyl bromide and 1. Rüdel: Discussion on the possible role of X. index by ISEM Bovey et al. These are promising sources of germplasm for obtaining resistant rootstocks. Bouquet: M. index. although it is not resistant to the virus itself.: GFLV particles are arrayed in long parallel rows in thin-sectioned mesophyll cells of infected grapevines.: Soil fumigation with 1. rotundifolia is resistant to fanleaf virus transmission by X. Transmission trials gave a few positive results. 1980 1980 1980 1981 1981 1981 1981 1983a 1983b 1983 1983 1983 1985 1985 1986 1986 24 . vuittenezi might be a vector of GFLV is therefore uncertain. Vuittenez: Review on serological methods of detection and identification of grapevine viruses.: Experiments with systemic nematicides for controlling X. index. vuittenezi population used for the experiments could not be entirely ruled out.: Detection of fanleaf virus and other sap-transmissible viruses in grapevine tissues by ISEM.: Use of systemic nematicides for the control of X.: Comparison of polyclonal and monoclonal antibodies for detecting fanleaf virus with ELISA in various grapevine tissues. Efficiency of both methods is compared. Forty days of treatment. RNA-2 contains the cistron coding for the viral coat protein. index larvae were present in the X. Raski et al. That X. Monette: Heat therapy of GFLV and ArMV--infected grapevines with alternating temperatures.3-dichloropropene failed to eradicate either nematodes or fanleaf virus from the soil but reduced the incidence of the disease to acceptable levels. Lear et al. Morris-Krsinich et al.1979 1980 1980 Kalasian et al. Bouquet: Muscadinia rotundifolia becomes infected by GFLV when the virus is transmitted by grafting but resists infection when transmission is by X. Krake and Woodham: Possibility that the agent of yellow speckle is involved together with GFLV in the etiology of vein banding. Walker et al.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. index by ELISA. Hafez et al. index and fanleaf in grapevines. especially in wood shavings of dormant canes during winter. the possibility that a few X.

TBRV and leafroll from infected grapevines by in vitro meristem tip culture.: Use of green grafting technique for sensitive and quick GFLV detection under greenhouse conditions. No eradication was obtained.: Identification of a satellite RNA of GFLV. Serghini et al. Fuchs et al.1987 1987 1987 Huss et al.: Detection of GFLV in grapevine seeds and seedlings by ELISA. Viruliferous nematodes were crushed in standard extraction buffer and tested in batches of 1-50 by means of DAS-ELISA. SLRV. Long term fallow (about 5 years). Catalano et al. Resistance is controlled by two unliked recessive genes Walter et al.3-dichloropropene and methyl bromide for controlling X. Walker and Meredith: Identification of two accessions of Vitis vinifera resistant to GFLV. Treated vines yielded more for over 4 years. Effect on yield and economic importance. Reliable results were obtained with samples of 20-50 nematodes. Rüdel: Review on the most important virus diseases of grapevines in West Germany. RNA-3 encodes a non structural protein.: Production and use of monoclonal antibodies to GFLV.: Determination of the complete sequence of GFLV RNA-2. Treatments with soil fumigants are no longer permitted in Germany. Raski and Goheen: Comparison of 1. ArMV.: Improvement in the serological detection of GFLV and ArMV viruses using monoclonal antibodies. but less consistent results were obtained with 1-10 nematodes. The selection of resistant cultivars and rootstocks is considered of primary importance.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. Etienne et al. Mild and severe strains were discriminated on the basis of field performance of infected Vitis. quinoa and transmitted by heterografting to Vialla and Kober 5BB rootstocks. Walter and Etienne: Detection of GFLV in wood shavings of dormant canes. index and GFLV. and has strong homologies with the satellite RNA associated with ArMV. Altmayer: Elimination of GFLV. Pinck et al. Previous experience showed that 1.: Evidence of a differential efficiency of GFLV transmission by Xiphinema index populations from different geographical origins. RpRV. GFLV.: Two rootstock selections derived from crossings V. Rüdel: Severe restrictions set on the use of soil fumigants in West Germany for environmental reasons make control of "Reisigkrankheit" very difficult.: Study of interactions between GFLV and ArMV isolates grown in C. the latter being especially damaging on the variety Kerner.: Determination of the nucleotide sequence of the satellite RNA (RNA-3) of GFLV. Walter et al. 1988 1988 1988 1989 1989 1989 1989 1989 1990 1990 1990 1990 1990 1991 1991 1991 25 . Catalano et al. Lázár et al. Walter et al.: 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. rotundifolia showed good resistance to GFLV in California.: Detection of GFLV in the vector Xiphinema index by ELISA. Mild strains were shown to confer protection towards severe challenging strains in Chenopodium and grapevine. Walker et al. Martelli and Taylor: Review article on nematode-transmitted viruses. cultivation of non-host plants and organic soil amendments are recommended. vinifera x V. Positive. RpRSV and ArMV are common.

Walker et al.b Hans et al. identification of their replication determinants and evidence of replication in Chenopodium quinoa protoplasts 1993 1993 1993 1993 Martelli 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.: In naturally GFLV-infected vineyards the hypovirulent ArMV A1 isolate induces delayed infection by GFLV Saldarelli et al.: Genomic RNA-1 of GFLV is completely sequenced and its genetic organization determined.: Production of GFLV satellite RNA transcripts.: GFLV isolated in Hungary from naturally infected symptomatic plants of Aristolochia clematis and Lagenaria siceraria turbinata. which is also resistant to phylloxera.: GFLV satellite RNA detected in 5 of 34 virus isolates from different geographical locations. Walter et al. Staudt and Weischer: Vitis rotundifolia and Vitis munsoniana resist infection by GFLV transmitted by X.: Development of cDNA probes to GFLV genomic and satellite RNAs and their use for virus detection directly in grapevine extracts. Esmenjaud et al.: Two Vitis vinifera x Muscadinia rotundifolia rootstock hybrids (O39-16 and O4343) grafted with Cabernet sauvignon showed a high level of tolerance to GFLV. O39-16 in particular.: Use of modified GFLV coat protein genes for transformation of different Nicotiana species for inducing resistance. Horvath et al.: Co-inoculation of C. GFLV is a quasispecies occurring in the field as a series of minor molecular variants. index by biotin-avidin ELISA Bardonnet et al.: Detection of GFLV in X. index.:Detection of GFLV in single nematodes by RT-PCR. index. delays symptom expression by 1-2 days and adversely affects virus replication. Satellite RNA appears to have a modulating effect on virus pathogenicity. This represents the first substantiated record of a natural GFLV infection in hosts other than Vitis. GFLV is eliminated from somatic embryos obtained from tissue cultures grown at 35 °C 1991 1991 1992 1992 1992a. Both became infected in the course of a 12-year trial but had no reduced crop yields. thus qualifying for use in X.: Evidence that transgenic tobacco plants expressing the coat protein of GFLV are protected from GFLV infection. index infested soils. quinoa with biologically active transcripts of GFLV F-13 satellite RNA and GFLV strains devoid of satellite. Viry et al. Fuchs et al. Staudt: Study of the spread of GFLV in several Vitis species. Goussard and Wiid: First application of somatic embryogenesis for sanitation of grapevines. hybrids and breeding stocks after infection of the roots by means of viruliferous X. 1993 1993 1993 1993 1993 1994 1994 1994 1994 26 . Spielmann et al. Ritzenthaler et al.1991a 1991b Fuchs et al. Esmenjaud et al.: Production of biologically active transcripts from cloned cDNA of genomic RNAs of GFLV.

: Genetic transformation of Vitis vinifera with the coat protein gene of GFLV for resistance induction.: Development of a GFLV detection system based on PCR analysis of immobilized virions. Gaire et al. Mauro et al. Belin et al. Walker and Jin: V. including the two accessions reported as resistant by Walker and Meredith (1990).: Demonstration that a 28 kDa protein coded by GFLV RNA-2 is involved in the replication of this RNA. : Identification of RNA2-encoded proteins in the specific transmission of GFLV by X. all were susceptible to the virus. RNase L) genes for inducing resistance to GFLV. Lahogue and Boulard: Search for genes of resistance in grapevines. Pinck: A comprehensive review of the molecular aspects of GFLV genome and its replication strategy. 5 oligoadenylate synthase. Pfeiffer et al.: Up to date account of GFLV replication strategy. Of 531 accessions of European. Spielmann et al. Ritzenthaler et al.: Successful transformation of somatic embryos of an European grape cultivar (Russalska 3) with the normal.: Identification of the molecular signal accounting for the systemic spread of GFLV in infected hosts. except for four.: Transformation of Nicotiana species and Vitis rupestris with different virusderived (coat protein. rupestris x M.: Genetic transformation of American roostocks with the coat protein gene of GFLV for resistance induction.: Identification of membranes derived from the endoplasmic reticulum as sites of GFLV replication. Rowhani et al.: Attempts to characterize molecularly X. Krastanova et al. Belin et al. Walter and Martelli: Review article on detrimental effects of viruses on grape yields. index feeding. Fuchs: Review article on genetic transformation of grapevines for resistance to GFLV and other pathogens. Martelli and Walter: Review article on certification of grapevines. This resistance is controlled by a single dominant gene.: 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. rotundifolia hybrids show high resistance to X. Naraghi-Arani et al. De Luca et al. Gölles et al.: Demostration that the movement protein of GFLV is located on the intracellular tubular structures containing rows of virus particles. American.1995 1995 1995 1995 1995 1996 1996 Brandt and Himmler: Use of immunocapture RT-PCR for GFLV detection in host tissues. index. 1997 1998 1998 1999 1999 2000 2000 2000 2000 2001 2002 2003 2003 2003 27 . Ritzenthaler et al. The level of resistance obtained looks promising. replicase) and exogenous (2. and Asian Vitis species inoculated by green grafting with a GFLV source.: 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. Pfeiffer et al. truncated or nontranslatable coat protein gene of GFLV. index populations by PCR-RFLP and sequencing of the ITS region.

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Nepoviruses. Maniloff.. M.V. 2005) Extensive reviews of the biological. 1978. Vol. 2000. Eight Report of the International Committee on Taxonomy of Viruses (C. References Goldbach R. Rüdel M... H. Fritsch. RT-PCR) are routinely used for their detection in grapevine tissues (primarily cortical scrapings from dormant canes). Family Comoviridae. J.D. Bulletin de l'OIV 69. A tentative grouping of nepoviruses Phytopathologia Mediterranea 17. Oxon. Plenum Press.INFECTIOUS DEGENERATION (EUROPEAN AND MEDITERRANEAN NEPOVIRUSES) Besides GFLV several other nepoviruses can infect grapevine in Europe. and G.P. P. Martelli G. Walter B. T.J. V. Serological (ELISA. Taylor and Brown. the Mediterranean and Middle East. Nepoviruses.F. single-stranded RNA genome occurring as two functional species (RNA-1 and RNA-2). Several of these viruses have distorting and chromogenic strains and may occur in mixed infections with GFLV.. 945-971. and A. 2005. are now classified in the genus Nepovirus. 1996. 1981. Seventh Report of the International Committee on Taxonomy of Viruses (M.J. causing diseases whose symptoms are similar to. Gallitelli. Family Comoviridae. 3. 1997) and their satellite RNAs (Mayo et al. 1992). and A. 807-818. Simons and M. 185.. 2000) are available. J. The Plant Viruses.. Le Gall O. Amsterdam. New York. Karasev. typified by Tomato ringspot virus (ToRSV) (Martelli et al. G. T.). Bari. Leibowitz.A. (G. Harrison B.P.. 1955) and are subdivided into subgroups based on physico-chemical properties of member viruses. Wellink.M. 2005). 145-147.F. Mechanical transmission to herbaceous hosts or indexing on Vitis indicators can also be used. 286 pp. eds). Strawberry latent ringspot virus (SLRSV). D. (E. Elsevier/Noth Holland.A. Influence des viroses et qualité. or indistinguishable from those of fanleaf.F. Springer-Verlag. 1955. Palukaitis. epidemiological. These viruses can readily be eliminated by heat therapy or in vitro meristem tip culture. Wetzel and N. Murant (eds). In: Handbook of Plant Virus Infections: Comparative Diagnosis. Quacquarelli. Academic Press. All have polyhedral particles about 30 nm in diameter and a positive sense. eds) Elsevier/Academic Press. Piazzolla. Polyhedral Virions and Bipartite RNA Genomes. Kurstak. i.B.. Murant 1981. G. Ball. Vienna. Brown. T.A. 341-347. 1978. In: Virus Taxonomy. Milne. physico-chemical.P. Nepoviruses of grapevine and their nematode vectors in the EEC. has now been assigned to the newly established genus Sadwavirus (Le Gall et al. subgroup B. Van Regenmortel et al. which are separately encapsidated. 197-238. 5. 35 . Sixth Report of the International Committee on Taxonomy of Viruses (F.e.. ed). typified by Tomato black ring virus (TBRV).. 362 pp. Taylor C. Mayo M. U. 1996.V. Martelli. 23-39. CAB International. 1977.D.A. In: Virus Taxonomy.H. eds). Istituto Agronomico Mediterraneo. San Diego.. Iwanami.A. Scholtof. Harrison B. subgroup C.G. ISEM) and molecular assays (hybridization.F. 1992. a non-taxonomic clustering.E and D. C. 1ere partie: Effects des viroses sur la culture des vignes et ses produits. Le Gall et al. Savino and P. Fauquet. Satellite nucleic acids. 1977) .. CMI/AAB Descriptions of Plant Viruses No. A. Mayo. which were originally included in the Nepovirus group (Harrison and Murant. Taliansky. 10281032 Murant A. and molecular characteristics of nepoviruses (Harrison and Murant. A.P. K. In: Virus Taxonomy. ed.. K. subgroup A typified by Tobacco ringspot virus (TRSV). Nematode Vectors of Plant Viruses. Lehto. Their detrimental effects to grapevine culture and products have been summari zed by Walter and Martelli (1996). In: Grapevine Viruses and Certification in EEC Countries: State of the Art. Martelli and R. Martelli. Yoshikawa. Desselberger and L. family Comoviridae (Goldbach et al. Sélection clonale de la vigne: sélection sanitaire et sélection pomologique. Many are transmitted by longidorid nematodes (Rüdel. Murant. Sanfaçon. Murphy et al. Jones. Quaderno No. 1997. a nematode-borne virus originally classified as a tentative species in the genus Nepovirus. London. M. Nepovirus Group. 1996.

Romania. and Japan. Yugoslavia. Dalmasso et al. respectively. In Germany. Quacquarelli et al.: Physico-chemical properties of GFLV. Kerner appears to be caused by ArMV infection. Kobayashi et al. Hungary. Turkey. Component T is made up of empy protein shells. It can infect many woody and herbaceous plants and is transmitted to grapevine by the nematode Xiphinema diversicaudatum but not by X. M. Description ArMV. In other V. Transgenic plants expressing the coat protein gene of the virus have been produced. ArMV occurs often in mixed infections with GFLV in certain areas of France and Italy. accounting for 27% (component M) and 41% (component B) of the particle weight. Jankulova and Kaitasova: ArMV found in grapevine in Bulgaria. Rüdel: Transmission of ArMV to grape seedlings by Xiphinema diversicaudatum. This virus has also been found in grapevine in Switzerland. 36 . 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.: ArMV detected in Japan in grapevines imported from Europe. Canada. the vector of GFLV.2 x 106 (RNA-1) and 1. ArMV and TBRV by ELISA in Israel. Israel. 2. is serologically related to GFLV. a typical nepovirus belonging in subgroup A of the genus Nepovirus. losses up to 50 % have been recorded. and sediment as three components (T. Brückbauer and Rüdel: Symptoms of atypical Reisigkrankheit in the vineyard are associated with ArMV in West Germany Bercks et al. Coat protein has a single type of subunits with Mr 54 x 103. SLRSV and TBRV found in grapevines with atypical Reisigkrankheit in West Germany. Vuittenez et al. and. Its particles are about 30 nm in diameter. Bulgaria. TBRV. have a angular outline. Iran. Cross-protection between ArMV and GFLV has been reported.: Xiphinema diversicaudatum can transmit ArMV to grapevine.95-2.Genus NEPOVIRUS ARABIS MOSAIC VIRUS (ArMV) 1. USA (California). The genome is a positive sense ssRNA. AILV and GCMV.: Isolation of ArMV from grapevine in Italy. Murant: ArMV description in the CMI/AAB Descriptions of Plant Viruses series.: Ultrastructure of ArMV infections in plant tissues Martelli and Lehoczky: Detection of ArMV in grapevine in Hungary. 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. whereas components M and Bcontain RNA. symptoms are of the fanleaf type. and B). Belli et al. index.: Detection of ArMV by ISEM Tanne: Detection of GFLV. consisting of two separately encapsidated molecules with Mol.4 x 106 and a size of 1104 nt. vinifera varieties. and circular about 350 nt in size. ArMV. Russo et al. wt of 0. always in Germany the severe dieback disease of the cv. wt 2. The virus supports the replication of two types of satellite RNAs. and with other nepoviruses in the Reisigkrankheit complex of western Germany. Stellmach: Review paper on ArMV in grapevine. linear with Mol.: ArMV. Gerola et al.1 x 106 (RNA-2).: Interactions between nepoviruses in grapevine and herbaceous hosts.

SLRV and TBRV Belli et al. Walter et al.1982 1984 1985 Brückbauer: Possibility of distinguishing GFLV. Eppler et al.Kerner to ArMV seems to be limited in time. Stellmach: Kerner disease is probably caused by ArMV. the virus is recovered from the scion only during the first year. Study of histological changes at the graft union level. When a healthy scion is grafted onto an infected rootstock. whereas other nepoviruses.: Transformation of grapevines with the coat protein gene of ArMV. RRV. Stellmach and Berres: The susceptibility of cv. Rüdel: In the Palatinate (West Germany) ArMV is transmitted by Xiphinema diversicaudatum and occurs often in mixed infections with GFLV in grapevine.: ArMV is not seed-transmitted in grapevines Liu et al.: Detection of ArMV by PCR in herbaceous hosts and grapevines Steinkellner et al. Kaper et al. Becker et al. Steinkellner et al. Molecular assays are as good as ELISA for routine testing.: Properties of a strain of ArMV isolated from grapevine in Italy. 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 .: Association of ArMV-infected rootstocks with Kerner disease in West Germany. Hypothesis of a graft incompatibility when the rootstock is infected with ArMV.: ArMV in Switzerland Lázár et al.: Transgenic Nicotiana plants transformed with the coat protien of ArMV produce empty viral shells. such as RpRSV or GFLV can be found in both rootstock and scion.: 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. Yield losses may reach 77% in cv. The virus cannot be recovered from leaves or buds of the Kerner scions.: Use of cDNA probes for ArMV detection. Faber. Marc-Martin et al.: A hypovirulent ArMV isolate delays GFLV infection in grapevines under field conditions. MacKenzie et al. Ipach et al. : Comparison of coat proteins of ArMV and other nepoviruses. whereas the rootstock remains infected.: ArMV recorded from Romania Huss et al. ArMV.: Cross-protection experiments in Chenopodium quinoa between ArMV and GFLV Gugerli et al.: Properties of ArMV small satellite RNA.: The presence of ArMV satellite RNA can attenuate symptoms in certain hosts Bertioli et al.: Nucleotide sequenece of a small circular satelllite RNA Steinkellner et al.: Survey for the presence of ArMV in Canadian vineyards Lahogue and Boulard: Search for genes of resistance in grapevines.: Nucleotide sequence of the ArMV satellite RNA Liu et al.: Evidence that ArMV has two RNA-2 molecules and complete nucleotide sequence of both RNAs Etscheid 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.

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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.

1976 1977

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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

40

35-41. 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.. Cigasr.: A virus serologically related to GBLV isolated from Vitis labrusca cv. S. Boscia and G. Bulletin OEPP/EPPO Bulletin 32. Virus Genes 30. Description Grapevine Bulgarian latent virus (GBLV) owes it name to the fact that it was found for the first time in Bulgaria in 1971. By contrast. 2. 471-475 Gokalp K. Digiaro and G. Martelli et al. Martelli. Component T is made up of empy protein shells. Martelli. I. Coat protein has a single type of subunits with Mr 54 x 103. Sabanadzovic.: Isolation of virus CM112 from in vitro cultures of grapevine tissues. Complete nucleotide sequence of RNA-2 of two Turkish nepoviruses. A.. accounting for 39% (componet B1) and 42% (component B2) of the particle weight. 1977 1978 41 .P. 2003.2 x 106 (RNA-1) and 1. The virus occurs as different closely related but serologically distinguishable strains. wt of 2. Kizlar tahtasi showing mild fanleaf-like symptoms Gokalp et al. Historical review 1972 1972 1977 Ferreira and De Sequeira: Description and preliminary characterization of an unidentified virus denoted CM112. M. and B2). Two isolates of GBLV have been transmitted by mechanical inoculation to seedlings and rooted cuttings of several grapevine cultivars without inducing symptoms.P. M. isolated in 1970 in Portugal from symptomless vines. GRAPEVINE BULGARIAN LATENT VIRUS (GBLV) 1. wt 0.: Description of GBLV. Digiaro. Biological. Abou Ghanem-Sabanadzovic. 335-340. Cigsar I. Sanitary status of grapevine in south eastern and central Anatolia.95-2. The virus can be detected directly in grapevine leaf extracts by immunodiffusion in agar plates. Historical review 2002 2003 2005 Cigsar et al.P. whereas components B1 and B2 contain RNA. The scattered distribution of infected vines in the field suggests that the virus is spread primarily by infected propagative material. Uyemoto et al. consisting of two separately encapsidated molecules with Mol. N. De Stradis. Virus particles are about 30 nm in diameter and sediment as three components (T. is not seed borne and was reported only from south-eastern Turkey. but different from GBLV. A strain of this virus had been found previously in Portugal and described as virus CM112.1 x 106 (RNA-2).: Description and thorough characterization of GARSV identified as a new species in the subgroup B of the genus Nepovirus Abou Ghanem-Sabanadzovic et al. GBLV has also been recorded from Hungary and Yugoslavia. B1. M.: First isolation by mechanical transmission of an unknown nepovirus from cv. 2. Martelli et al. The genome is a positive sense ssRNA. 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.5 x 106 (less than 1800 nt).vector.: GBLV description in the CMI/AAB Descriptions of Plant Viruses series. Concord in New York State. The economic importance of the virus is minor. References Abou Ghanem-Sabanadzovic N. Martelli. Digiaro and G. 2005. Journal of Plant Pathology 85. The virus supports the replication of a satellite RNA with Mol. physico-chemical and serological characterization of the virus and assignment to the Nepovirus group (now genus Nepovirus). De Mendonça et al. 2002. where it is widespread and infects latently several grapevine varieties growing in widely separared areas.: Complete nucleotide sequence of GARSV RNA-2 3. D.

Gallitelli et al. M.: A comparative study of Bulgarian GBLV isolates and the Portuguese virus CM112 establishes that CM112 is a serologically close but distinguishable strain of GBLV. P.P. 1970. Niagara Falls 1980.: 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. 217-222.A. A. De Sequeira and A. Proceedings 7th Meeting of ICVG. Niagara Falls 1980. Colmar 1970. D.: Improvement of ELISA protocol for GBLV detection the whole year round. Properties of a distinctive strain of Grapevine Bulgarian latent virus. and O.C. 1985. Martelli G.A. Annals of Applied Biology 85. Quacquarelli. 1972. Gallitelli D. 1979. Physical and chemical properties of the particles and ribonucleic acid of blueberry leaf mottle virus. The ultrastructure of grapevine Bulgarina latent virus infections in natural and artificial hosts. 21-24. De Sequeira. A. Gallitelli. Gallitelli. 186 Martelli G. Monografias INIA 18 . Savino and A. Proceeding 4th Meeting of ICVG. Numéro hors série. Phytopathology 71. Russo et al. 1980. Simoes. Di Franco. Dimitrijevic B. De Sequeira.Sur l'isolement d'un virus à partir de cultures de tissus de vigne.. Annales de Phytopathologie. V. Quacquarelli and D. De Sequeira O. 245-250. D. Ganeva and M. Ramsdell D. 1980. D. 135-141. Acta Phytopathologica Academiae Scientiarum Hungaricae 16. 349-354. Proceedings 6th Meeting of ICVG. Some properties of grapevine Bulgarian latent virus. 51-58. 42 . Colmar. Savino and O. 95-101. Preliminary studies on an undescribed grapevine virus. A manually transmissible latent virus of the grapevine from Bulgaria. Pereira and V. 1992. Krastanova S. Abracheva. O.. P. Phytopathologia Mediterranea 22. V.. Savino. 468-472. Rasteniev'dni Nauki 29.. De Sequeira. 1981. The Portuguese strain supports the replication of a satellite RNA.Proceedings 7th Meeting of ICVG. Numéro hors série.P.: Ultrastructural study of GBLV infections in grapevine and C. Ferreira. 1980 1980 1980 1980 1981 1981 1982 1983 1985 1992 3. CMI/AAB Descriptions of Plant Viruses. 27-32. Possibilities for the whole-year ELISA detection of viruses infecting grapevines. 1980. Niagara Falls 1980. Martelli et al. 143-145 De Mendonça A. Dimitrijevic: GBLV found in Yugoslavia De Mendonça et al. Cordoba 1976.: Detection of virus CM112 in grapevine leaf extracts by ISEM. Vasconcelos-Costa. Krastanova et al. Varennes and De Sequeira: First application of ELISA for the detection of virus CM112. Some properties of the new latent virus from grapevine rootstocks in Yugoslavia. Occurence of grapevine Bulgarian latent virus in Hungary. France.A. V. 113-120.. No... Pocsai: Occurence of GBLV in Hungary.A. Jankulova. Martelli G.. 1972. Proceedings 7th Meeting of ICVG. An immunoradiometric assay for the titration of a Portugese strain of grapevine Bulgarian latent virus (GBLV). and J. A preliminary report. Abracheva. Ferreira A. 1977. Pocsai E. Gallitelli. Martelli G. Annales de Phytopathologie. quinoa. 1981.P. Applicability of immunosorbent electron microscopy (ISEM) for the detection and identification of CM112 virus in grapevine.N. A. and R. Stace-Smith. Mota. 269-272. M. Yankulova. 1978. M.. De Sequeira and Vasconcelos-Costa: Use of an immunoradiometric assay for the titration of the Portuguese strain of GBLV. 1983. O.P.: Detection of GBLV in grapevine leaf extracts by ISEM. Savino and A. References De Mendonça A. Russo and V. Proceeding 4th Meeting of ICVG.A.1979 Martelli et al. Ramsdell and Stace-Smith: The New York isolate of GBLV is a strain of BLMoV. Grapevine Bulgarian latent virus. Quacquarelli.A.A.. Garcia de Orta Estacao Agronomica 12.

It has been recorded also from Czechoslovakia. Virus re-named Grapevine chrome mosaic virus. a symptom virtually indistiguishable from GFLV-induced yellow mosaic. E. Martelli and Quacquarelli: Description of HCMV in the CMI/AAB Descripitons of Plant Viruses series. Plant Disease Reporter 61. Hummer. Varennes A. wt of 1. RNA-1 has Mol. Saric and Hranuelli: GCMV recorded from Croatia.K. and was originally called Hungarian chrome mosaic virus. Although GCMV particles have been detected by immunosorbent electron microscopy in Xiphinema index fed on infected hosts. 949-953. early reports that this nematode could transmit the virus have not been confirmed. Martelli et al. 1977. Detection of CM122 latent grapevine virus by enzyme-linked immunosorbent assay (ELISA). Pozsár et al. The virus is not serologically related with GFLV. The virus belongs in the same subgroup of TBRV (subgroup B) to which is distantly related serologically. transmitted to herbaceous hosts from Hungarian grapevines with symptoms similar to those of fanleaf and yellow mosaic.F.: Host range and properties of a spherical virus. The virus appears to be unrelated serologically to GFLV and is not transmitted by X. index.8 x 106 .K. Leaves of infected vines are partially or entirely bright yellow or whitish. Martelli and Quacquarelli : Physico-chemical characterization of HCMV and comparison with TBRV. Taschenberg and D. Jakó et al: HCMV affects pigment and sugar content of infected grapevine leaves.: The isometric virus associated with Hungarian chrome mosaic is serologically distantly related to Tomato black ring virus (TBRV). called Hungarian chrome mosaic virus. pretty much like GFLV. The genome is bipartite. 269-277. GCMV is transmitted through grapevine seeds. 2..Uyemoto J. Lehoczky and Tasnady: A study of the effect of HCMV on yield and sugar content of infected grapevines. The coat protein has a single type of subunits of Mr 52 x 103.: HCMV adversely affects photosynthetical carbon dioxide fixation. Description Grapevine chrome mosaic virus (GCMV) was first found in Hungary. Historical review 1966 Martelli et al. Martelli and Sarospataki: X. Croatia and Austria. RNA-2 has Mol. Agronomia Lusitana 41. Some virus strains induce leaf deformity.A. Mali et al. However.: GCMV recorded from Slovakia and report of X. 1966 1968 1968 1969 1969 1971 1972a 1972b 1972 1975 1977 43 . Tobacco pants and the rootstock 110R have been successfully transformed with the viral coat protein for induction of resistance. symptomless infection may occur. index as vector of the virus (unconfirmed results). a size of 7212 nt and accounts for 40% of the particle weight. vuittenezi transmits GCMV or GFLV. Isolation and identification of a strain of grapevine Bulgarian latent virus in Concord grapevines in New York State.6 x 106 . and O. 1982. double nodes and short internodes. Affected vines lack in vigour and may decline and die. sometimes together with X. De Sequeira. GRAPEVINE CHROME MOSAIC VIRUS (GCMV) 1. index. a size of 4441 nt and accounts for 31% of the particle weight. near Lake Balaton. Kenten: GCMV is distantly serologically related to Cacao necrosis virus. vuittenezi is very frequently found in vineyards with chrome mosaic patches. wt of 2. No evidence that X. Evaluation of short reaction times and re-use of a-globulin and conjugate. Martelli: Purification and serology of the virus isolated from Hungarian grapevines with fanleafand yellow mosaic-like symptoms.

Brault. Brault et al. Dimou et al.: Tobacco plants genetically engineered with the coat protein gene of GCMV are resistant to infection.M.: Detection of GCMV in leaf dips by ISEM.1979 1980 1980 1982 1984 1985 1985 1989 1989 1989 1990 1993 1993 1994 1994 1995 1995 1997 2000 Lehoczky et al. Bretout et al.. Le Gall et al. Acta Horticulture 235. Candresse. V. Laimer da Camara Machado and V. L. Nucleotide sequence and genetic organization of Hungarian grapevine chrome mosaic nepovirus RNA2.: Characterization of a GCMV strain and confirmation of its serological relationship with TBRV. Le Gall et al. Brandt and Himmler: Development of a IC-PCR protocol for GCMV detection in cortical scrapings from dormant grapevine canes.: GCMV and TBRV can recombine. Le Gall. D'Onghia.: Up-to-date report on virus diseases of grapevines in Hungary and description of the clean stock programme implemented in the country. No assessment of resistance made. Nucleic Acid Research 17. Candresse. O.P. O. Bretout C.: Development of molecular probes for GCMV detection. Lanneau and J.: GCMV detected by ELISA in infected field-grown vines. Ravelonandro and J.: GCMV recorded from Austria. and G. Occurrence of grapevine chrome mosaic nepovirus in Austria. Le Gall and J.: Complete nucleotide sequence of GCMV RNA-1. References Brandt S. Lehoczky: Pinot noir and Jubileum 75 are good indicators for GCMV. T. Dunez. Le Gall et al. Virus and RNAspecific molecular hydridization probes for two nepoviruses. Journal of Phytopathology 142. index extracts by ISEM. Dodd and Robinson: GCMV and TBRV are molecularly related.. 258-262. Vitis 34. 1994. Brault V. Further demonstration that the two viruses are related. This finding does not imply vectoring capacity by this nematode. Candresse. Brault et al. 231-238. index are inconclusive. Lázár et al: Seed transmission of GCMV in grapevine. Genetically engineered resistance against grapevine chrome mosaic virus. 3. 1989. M. Hilbrand. Lehoczky et al. Le Gall. Plant Molecular Biology 21. Brault V.. Kölber et al. M. Roberts and Brown: Detection of GCMV in X. Delbos. Russo et al. Dunez. Doz et al. A. Savino.: Description of a certification scheme for the production of virus-free propagating material in Hungary. T. Himmler. 1993. 1989. Detection of nepoviruses in ligneous grapevine material using IC/PCR.: Transformation of roostock 110R with the coat protein gene of GCMV. Dimou D. 89-97. 44 . 1995. The virus vector is yet to be identified. Taylor and Brown: Results of GCMV transmission trials with X.: GCMV cross-protects Chenopodium quinoa from the severe apical necrosis induced by a TBRV strain. Lázár et al. T. O. R.: Complete nucleotide sequence of GCMV RNA-2. Dunez. 127-128. M. 7809-7819.

.. NW Frazier (ed. 45 . No. ArMV) in the seeds and seedlings of grapevine by ELISA. Dunez. Acta Phytopathologica Academia Scientiarum Hungaricae 1. Hungarian chrome mosaic. 567-568. Host range and properties of a virus associated with Hungarian grapevines showing macroscopic symptoms of fanleaf and yellow mosaic. L. Vanek and V. and G. Sarospataki.. Delbos and J. Sarospataki. Mikulás..C. 1966. Prémunition: une competition entre souches faibles et souches sévères pour un voie commune d'expression de symptômes. 1966. Acta Phytopathologica Academia Scientiarum Hungaricae 3. 185-192. 1968. G. Nucleotide sequence homologies among RNA species of strains of tomato black ring virus and other nepoviruses. Martelli G.. Quacquarelli. J.H. G. Szonyegi and M. 1971. 103. Kölber and J. and A. Quacquarelli and J. Pol'nohopodarska Veda . with Special Reference to Vitis. Lehoczky and A.P. Farkas. University of California. 102. Kuszala and A. Grapevine chrome mosaic virus. 1968.P.P. Bouquet. 135-140. Le Gall O.. Agrobacterium-mediated genetic transformation of grapevine somatic embryos and regeneration of transgenic plants expressing the coat protein of grapevine chrome mosaic nepovirus (GCMV). Lehoczky. Phytopathologia Mediterranea 24. Y. 236-237. Grapevine virus diseases and clean stock program in Hungary. Preliminary report on purification and serology of a virus associated with Hungarian grapevines showing macroscopic symptoms of fanleaf and yellow mosaic. Lázár J. 1995.P. Robinson. The change of pigment and sugar content in the chrome mosaic virus infected leaves of grapevine. S. 1970. T. Dunez. 206-210. Torregrosa . 1-7. Sarospataki and J. Annals of Applied Biology 71.P.P. 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.. 1-130. Effect of the grape chrome mosaic and grape fanleaf-yellow mosaic virus infection on the photosynthetical carbon dioxide fixation in vine leaves. Martelli G. Division of Agricultural Sciences. E. L. Devergne. Davis 1965. Martelli G. The purification and some properties of cocoa necrosis virus. A. O.P. Nucleotide sequence of Hungarian grapevine chrome mosaic nepovirus RNA1. 1966.. Lehoczky and A... Brault and J. 1989.. Jakó N. 1975. Montreux 1993. Caractérization d'une souche du virus de la mosaïque jaune crome de la vigne (GCMV) isolée en Hongrie de vignes non panachées. Kölber. 1972a. Beczner. Jakó N. Extended Abstracts 11th Meeting of ICVG. Sznyegi. Berkeley. Kölber M. Annales de Phytopathologie 11. Les Colloques de l'INRA 11. 1972. Lehoczky . 1994. 1285-1289. 119-126. Candresse and J. 165-173. Dunez. S. Kiserletugyi-Kozlemenyek 64 (1-3). J. Martelli G. 3. Luntz. Certification scheme for production of virus-free grape propagating material and its results in Hungary. Danglot. Martelli G.. Martelli G. Division of Agricultural Sciences. Phytopathologia Mediterranea 8. Weinberg und Keller 15. Division of Agricultural Sciences.J. CMI/AAB Descriptions of Plant Viruses. 7795-7807. Davis 1965.Ser. Lehoczky and G. Studies on the nitrogen metabolism of vines infected with yellow mosaic virus. Vuittenez. 2000. Nucleic Acid Research 17. J. 1985. Serologische Verwandschaft eines mit dem ungarischen "chrome mosaic" vergesellschafteten Virus mit einem Stamm des "tomato black ring virus". 1731-1740. Journal of General Virology 65. L. R. J. 1990. A Handbook.. The effect of fanleaf and chrome mosaic virus diseases on yield and the fruit sugar content of grapevine. Lehoczky J. Lehoczky J. and S. Transmission by nematodes of some grapevine viruses occurring in Czecholovakia and Hungary. J.R.. Kölber M.. and D.M. Pacsa and J. Vitis 8. Hajdú. Quacquarelli.. J. Pozsár B. and A. Journal of General Virology 76. 505. L. 172-173. 123-141. Mali V. M. University of California. Martelli G. GCMV. Lehoczky. In: Virus Diseases of Small Fruits and Grapevines. Sarospataki. Detection of grapevine chrome mosaic virus in field-grown vines by ELISA. T. 58-72. 1972b. Kenten R. Detection of grapevine chrome mosaic virus in naturally infected vines by indexing. Nouvelle évidence d'une parenté sérologique éloignée entre ce virus et celui deas anneaux noirs de la tomate (tBRV). Annales de Phytopathologie. 389-401. a serotype ot tomato black ring virus. Extended Abstracts 13th Meeting of ICVG. G. Lázár J. 129-134. G. Lehoczky J. Hungarian chrome mosaic of grapevine and tomato black ring: two similar but unrelated plant viruses. Detection of some nepoviruses (GFV. Adelaide 2000. 29-44. Candresse and A. Plant Science. Le Gall O.Tasnady.) University of California. Candresse. GFV-YM. J. with Special Reference to Vitis.J. T. 1993.. Muranyi . 1979.Dodd S. 171-170. Lehoczky J. Sarospataki. A. Lehoczky. and G. Phytopathologia Mediterranea 24. 1969. 1969. Proceedings International Conference on Virus and Vector on Perennial Hosts. Quacquarelli. Doz B. 1984. Horváth. Numero hors série.. Quacquarelli. M. Bojnansky. J. Proceedings International Conference on Virus and Vector on Perennial Hosts. Lehoczky and G. 169-170. Nematodes of the family Longidoridae (Thorne 1935) Meyl 1960 found in Hungarian vineyards and virus transmission trials with Xiphinema index Thorne et Allen. 49-64. S. 1982.. V. 402-410.. Lázár. Kertgazdasag 22. 1985. Cardin. Le Gall O.

De Stradis. Martelli and V. Taylor C. The virus belongs in the subgroup A of the genus Nepovirus. 2005. 5. D.: A mechanically transmissible virus was recovered by sap inoculation from Tunisian grapevines showing mild fanleaf-like symptoms. Particles are isometric c. RNA-1 has a mol. wt of 2 x 106 daltons and apparent size of c. showing leaf and cane deformations Cigsar et al.: Description and thorough characterization of GDefV. 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. Immunosorben electron microscopy for detecting saptransmissible viruses of grapevine. GDefV is readily detected by ELISA and PCR using primers designed on the coat protein sequence. and belongs in the subgroup C of the genus Nepovirus. Digiaro and G. Grapevine deformation virus. Mostar 1977.: First isolation by mechanical transmission of an unknown nepovirus from cv.)..J. 2.Russo M.The virus has no recognized vector. M. : Complete nucleotide sequence of GDefV RNA-2 3. M (particles contaning a molecule of RNA-2 with Mol. wt of 2. 1980.. Proceedings of the Conference on Excoriosis and Virus Diseases of Grapevine. 30 nm in diameter and sediment as three components.. 335-340 Cigsar I. Saric A. is not seed-borne and reported only from south-eastern Turkey. Sabanadzovic. Nematode Vectors of Plant Viruses.800 nt. Investigations on grapevine viruses in Croatia. Boscia and G. Description Grapevine deformation virus (GDefV) was recovered from Turkish grapevines showing distinct fanleaf-like symptoms. 286 pp.P. Niagara Falls 1980. a novel nepovirus from Turkey. Martelli. Martelli. 2003. CAB International. Coat protein subunits have a Mr 53 x 103. 1997. E.P.F Brown. mol. Historical review 1991 Ouertani et al. is distantly related serologically to ArMV but not to GFLV. M.4 x 106 daltons and apparent size of c.. 149-151.800 nt) and B (particles containing a molecule of RNA-1 with Mol. The scattered distribution of infected vines in the field suggests that the virus is spread primarily by infected propagative material. distantly serologically related with ArMV.P. No vector is known and no information is available on the distribution and economic importance of the virus. A. M. Proceedings of the 7th Meeting of ICVG. GRAPEVINE TUNISIAN RINGSPOT VIRUS (GTRSV) 1. N.P.3 x 106 Da and a size of 3753 nt. wt of 1. Gokalp. Hranuelli. including all those known to infect grapevine. Digiaro. G. S. Digiaro and G. was isolated from a Tunisian grapevine with mild fanleaflike symptoms. Complete nucleotide sequence of RNA-2 of two Turkish nepoviruses. wt of 2.. 35-41. 2. Bulletin OEPP/EPPO Bulletin 32. Journal of Plant Pathology 85. The virus sediments as three components: T (empty shells). Oxon. and T. Sanitary status of grapevine in south eastern and central Anatolia. Virus Genes 30. GTRSV is serologically unrelated to any of 19 nepoviruses tested.6 x 106 Da and RNA-2. Abou Ghanem-Sabanadzovic et al. K. and D. 1977. Savino. 2002. 6. Historical review 2002 2003 2005 Cigsar et al. Abou Ghanem-Sabanadzovic. 46 . GRAPEVINE DEFORMATION VIRUS (GDefV) 1. identified as a new species in the subgroup A of the genus Nepovirus. 471-475 Cigsar I. 251-257. References Abou Ghanem-Sabanadzovic N. Martelli. The genome is bipartite. Description Grapevine Tunisian ringspot virus (GTRSV).

wt of 2. Mayo. Jones A. D. The virus is transmitted by Paralogidorus maximus Ebel et al. have angular outline. M. M. This strain differs considerably from the English type strain of the virus although is serologically closely related to it. The grapevine strain of this virus is serologically very distantly related to the two main serotypes. V. G. G. 88-118.M. 1978.: Recovery of RpRSV from grapevines of Palatinate. Raspberry ringspot virus.: Nucleotide sequence of RpRSV RNA-2 Jones et al. Journal of General Virology 73. M. Crop losses can be higher than 30 %. Murant A.T. Mögliche Beziehungen zwischen Virus und Symptomausprägung bei der Rebe. The viral genome is a bipartite positive sense ssRNA. Greco. 1994. Elbling in West Germany.: Sequencing and molecular characterization of two German isolates of RpRSV from grapevine 2003 3. Manoukian. 2189-2194. Wetzel. Symptoms are similar to those of fanleaf. accounting for 29% (component M) and 43% (component B) of the particle weight.A. McGavin. Locorotondo 2003.F. The nucleotide sequence of RNA 2 of raspberry ringspot nepovirus.J. Wardell J. Jolly. M. The virus has only been found in grapevine in western Germany. Castellano. Molecular characterization of two German Raspberry ringspot virus isolates infecting grapevines and construction of full length infectious clones. Properties of an unusual isolate of raspberry ringspot virus from grapevine in Germany and evidence of its possibile transmission by Paralogidorus maximus. Rüdel and B. Edwards and M. C.C.3. G. A. Savino. 1982. Reustle.A. D. Heat therapy of infected grapevines. W. Brown.J. 1992. Brückbauer H. Altmayer. Description Raspberry ringspot virus (RpRSV) is a nepovirus belonging in subgroup A of this genus. Ebel R.: Biological and physico-chemical characterization of the grapevine strain of RpRSV. and B). Martelli and N. 107-117. The coat protein has a single type of subunits with Mr 54 x 103. References Blok V. Brückbauer: RpRSV can be distinguished from other nepoviruses on the basis of symptoms induced on Vitis idicator plants Blok et al. A Schnabel. RASPBERRY RINGSPOT VIRUS (RpRSV) 1.J. RNA-2 is 3928 nt in size and contains a single ORF encoding a polypeptide with Mr of 124 kDa. D. and differs from the type strain for it often sediments as if it were a single centrifugal component... References Ouertani R. CMI/AAB Descriptions of Plant Viruses. Minafra. Scottish and English.. 283-300. Two strains of different virulence occur in the Palatinate.. 2003.F. Annals of Applied Biology 124.P. and sediment as three components (T. Historical review 1970 1978 1978 1982 1992 1994 Vuittenez et al. 2. 16. FS4 is a good indicator. These differences strongly suggest that the grapevine-infecting RpRSV may be a different viral species. A. Die Wein-Wissenschaft 37. Robinson. 47 . consisting of two separately encapsidated molecules with Mol. Krczal and T. Extended Abstracts 14th Meeting of ICVG.6 x 106 (RNA-2). Particles are about 30 nm in diameter. Murant: Description of RpBRSV in the CMI/AAB Plant Virus Description series Stellmach and Querfurth: Study of a strain of RpRSV isolated from cv.A.6 x 106 (RNA-1) and 1. Properties of a previously undescribed grapevine nepovirus from Tunisia. Archives of Virology 126.L. No. The type strain of RpRSV is transmitted by Longidorus macrosoma but the grapevine strain is transmitted by Paralongidorus maximus. 198. 1992. Boscia.

The genome is a bipartite positive sense single-stranded RNA occurring as two separately encapsidated functional molecules with Mol. latex test and barium sulfate test. RRV and TBRV are not transmitted by contact of roots or foliage in the vineyard. Historical review 1963 1965 1966 Stellmach and Bercks: TBRV detected in rootstock Aramon x V. TBRV produces a reduction in growth and yield. 1978. but they can be high. known to cause severe damage to the grapevine variety Joannes Seyve in Ontario. respectively.5 x 106 daltons and a size of 1327 nt. riparia 143 A in West Germany. Description Tomato black ring virus (TBRV) was first found in grapevines in Germany.. its own subgroup. Vuittenez et al. SLRV and TBRV found in grapevine in the Palatinate. Turkey. mottling of the older leaves. 2. Ontario as the cause of severe damage to cv. Meyer et al. 128-136. 279-327 TOMATO BLACK RING VIRUS (TBRV) 1.virus). 1967 1970 1970b 1976 1977 1980 1984 1984 1986 48 . Stellmach and Bercks: Further investigations on TBRV in grapevine. chlorotic spots. RNA-2 is 4662 nt in size encoding a polyprotein with Mr of 150 kDa. J. TBRV supports the replication of a satellite RNA with Mol. and Ontario (Canada).7 x 106 (RNA-1) and 1. wt of 0. Untersuchungen zur Serologie. M. The latex test is considered as the most sensitive and the least time consuming method.Stellmach G. Joannes Seyve virus. rings and lines on the leaves of recently infected plants. or directly in grapevine leaf extracts using bentonite flocculation test. is a strain of this virus. 1970. ArMV. The virus is a definitive nepovirus species classified in subgroup A of this genus. Kuszala.: RRV. The virus was detected in grapevines in the Niagara Peninsula. including TBRV : bentonite flocculation test. Lehoczky and Burgyan: Occurrence of TBRV in grapevines in Hungary. Greece. Bercks and Stellmach: ArMV. and G. Bercks and Querfurth: GFLV.: Nucleotide sequence of a TBRV satellite RNA.65 x 106 (RNA-2) accounting for 44% and 37% of the particle weight. about 30 nm in diameter have angular outline and sediment as three components (T. Virus particles are isometric. The vector to grapevine is Longidorus attenuatus. RNA-1 is 7356 nt in size and contains a single open reading frame encoding a polypeptide with Mr of 254 kDa. and an increase in graft failure. Pathologie und Thermo-Labilität mehrerer Reben-Isolate des Himbeerringflecken-Virus (raspberry-ringspot. RpRV and TBRV detected serologically in grapevine in West Germany. Bercks: Comparison of three serological tests for detecting several viruses. and B). M. Détection et étude selogique du virus latent des taches annulires du frasier (strawberry latent ringspot). Tanne: Detection of TBRV by ELISA in Israel. Vuittenez A. Rüdel and H Brückbauer. wt of 2. Joannes Seyve. either by agar gel diffusion with extracts of herbaceous hosts previously infected mechanically from grapevine. Rüdel: Transmission of TBRV to grapevine by Longidorus attenuatus. Israel. 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. Weinberg und Keller 25. Stellmach: Review paper on TBRV in grapevine. then in Yugoslavia. Losses are not known precisely. Annales de Phytopathologie 2. Their coat protein consists of a single type of subunits with Mr of about 57 kDa. Stobbs and Van Schagen: First record of TBRV from Canada. Querfurth.

Hemmer and C. and B).1986 1988 1993 Meyer et al. 3-5. Savino et al.6 x 106 (RNA-2). The virus is a definitive species in the genus Sadwavirus. M.: Nucleotide sequence of TBRV RNA-1 Abkas and Erdiller: TBRV recorded from grapevines in Turkey 3.. The virus is transmitted by Xiphinema diversicaudatum. Symptoms on affected European grapes are of the fanleaf type. Fritsch. Journal of General Virology 65. Virus particles are isometric. 1984. Description Strawberry latent ringspot virus (SLRSV) has been isolated from grapevine in the Palatinate (Germany) and in northern Italy. 1988.. Brückbauer. M. Van Schagen.: Nucleotide sequence of SLRSV satellite RNA. Bercks et al. Kreiah et al. 1517-1529. 49 . Turkiye. Assignement of SLRSV to the new genus Sadwavirus. 1986. Genus SADWAVIRUS STRAWBERRY LATENT RINGSPOT VIRUS (SLRSV) 1. Researches on grapevine virus diseases and determination of their incidence in Ankara. 55-61. du virus des anneaux noirs de la Tomate (tomato black ring). 279-327.W. It was also detected in imported vines in Turkey and Portugal. References Akbas B. Greif C.: SLRSV found in grapevine in Turkey.: Nucleotide sequence of TBRV RNA-2 Greif et al. O. Occurrence of tomato black ring virus on grapevine in southern Ontario. M. Canadian Plant Disease Survey 64. O. Historical review 1974 1977 1981 1982 1982 1987 1993 1994 2005 Murant: Description of SLRSV in the CMI/AAB Descriptions of Plant vrises series. Journal of General Virology 69. Their coat protein consists of two types of subunits with Mr 43 x 103 and 27 x 103. Fritsch. The nucleotide sequence of tomato black ring virus RNA-2. O. and G.A. Hemmer. J. Fritsch. and 1. Erdiller. respectively.: SLRSV recorded from grapevine in Italy. The virus supports the replication of a satellite RNA 1118 nt in size. Le Gall et al. about 30 nm in diameter have angular outline and sediment as three components (T. Rüdel and H. Babini and Bertaccini: Electron microscope study SLRSV infections in plant tissues. 1970. 1984.6 x 106 (RNA-1) accounting for 38% of the particle weight. Meyer M. Kuszala. wt 2. RNA-2 is 3824 nt in size and encodes a single ORF expressing a polypetide with Mr of about 99 kDa. and J. 1575-1583. Brückbauer: SLRSV can be distinguished from other nepoviruses on the basis of symptoms induced on Vitis idicator plants.. Meyer M. et du virus des taches annulaires du Framboisier (raspberry ringspot) chez des vignes du Palatinat. Détection et étude sérologique du virus latent des taches annulaires du Fraisier (strawberry latent ringspot).: Nucleotide sequence of SLRSV RNA-2.G. The genome is a bipartite positive sense singlestranded RNA occurring as two separately encapsidated functional molecules with Mol. 2.: SLRSV and other nepoviruses isolated from grapevines in Germany Credi et al. Nucleotide sequence of tomato black ring virus RNA-1. Journal of Gerneal Virology 67. Mayo and C. Vuittenez A. 1993. Kreiah et al. Journal of Turkish Phytopathology 22. Complete nucleotide sequence of a satellite RNA of tomato black ring virus. Hemmer and C. 1257-1271 Stobbs L. Annales de Phytopathologie 2.

In: Virus Taxonomy. Savino V. T Jones. Betti.V. Genus Sadwavirus. 102-104. 50 . Strawberry latent ringspot virus in grapevine. 1982.. G. 2527-2532. C. Strunk. Cooper and G. Kreiah S. CMI/AAB Descriptions of Plant Viruses No. Sequence analysis and location of capsid protein within RNA 2 of strawberry latent ringspot virus. 1977.P. Mayo. 74. Iwanami.I.M. Yilmaz.. K. D'Onghia and M.I. J. Karasev. (eds) 799-802 Elsevier/Academic Press. Turkey.. Wellink.. Weinberg und Keller 24. Phytopathologia Mediterranea 20..M. Desselberger. Mögliche Beziehungen zwischen Virus und Symptomausprägung bei der Rebe. Untersuchungen überdie Viruskrankheited der rebe unter besonderer Berüchsichtigung “atypischer Formen” der Reisigkrankheit. 1994. Le Gall O. The nucleotide sequence of a satellite RNA associated with strawberry latent ringspot virus. Eight Report of the International Committee on Taxonomy of Viruses (Fauquet. Sanfaçon. 2005. Babini. References Babini A..3.. and A. Credi R.. 1982. Gelli. L. H. 1993.. Cooper. A. 1163-1165. Martelli. Bercks R..A. Lehto. T.. Viral aggregates induced by a distinctive strain of strawberry latent ringspot virus from grapevine. Maniloff. A. Querfurth and M. Wetzel and N. Murant A. Journal of General Virology 75. FAO Plant Protection Bulletin 35.R. Brückbauer. T. Journal of General Virology. Rüdel. Bertaccini and C. M.R. Yoshikawa.A. A. Kreiah S. U. Brückbauer H. Phytopathologische Zeitschrift 104. 1987. 126. 56-63. 1981. J. A. Die Wein-Wissenschaft 37. Bertaccini. 88-118. G. A distinctive isolate of Strawberry latent ringspot virus from grapevines in Italy. Strawberry latent ringspot virus.. G. 133-180. London. and Ball. 1974.F. Strunk and J. J. H. L.A. 304-308.

New York State and Ontario) own-rooted European grapes affected by Tomato ringspot virus (ToRSV) and Tobacco ringspot virus (TRSV) decline rapidly. TRSV is transmitted by X. mottled (oak leaf pattern. positive-sense. V. PRMV. about 30 nm in diameter with angular outline. All these viruses. berlandieri or V. poor fruit setting. ingiallimento nervale (Ital. rivesi transmit ToRSV type strain (decline). its main host. Varietal susceptibility: There are great variations in the susceptibility of Vitis species and cultivars.GRAPEVINE DEGENERATION AND DECLINE (AMERICAN NEPOVIRUSES) 1. Bunches are small and straggly (Maryland's grapevine little berry) and leaves may show chrome yellow flecking along the veins (California's yellow vein). americanum sensu stricto. BLUEBERRY LEAF MOTTLE VIRUS (BLMoV) 1. TRSV. Partial sequence of the 3' 51 . Control: Use of virus-free propagating material and resistant rootstocks. Nematicidal control of vectors is possible. Adernvergilbung (Germ. whereas in V. In cold climates (e. californicum transmits ToRSV yellow vein strain. the infecting virus and the climatic conditions. rupestris plasma show field resistance to the northern US strain of ToRSV and to TRSV and PRMV. V. Longidorus diadecturus and L. labrusca is also resistant to TRSV. All other viruses are transmitted by longidorid nematodes: Xiphinema americanum sensu stricto and X. interspecific hybrids).). grapevine decline. little grape (Eng. Description Blueberry leaf mottle virus (BLMoV) is named after the disease induced in highbush blueberry (Vaccinium corymbosum). PRMV. and ToRSV separately or in combination. and/or ring spots) and distorted leaves. elongatus. This type of resistance is hypersensitivity. distortion of canes. riparia. A number of rootstocks containing V.e. In warmer climates (Maryland. X. are involved in the aetiology of North American grapevine degeneration and decline. which in blueberry is transmitted by pollen. except for BLMoV which may have been introduced from Europe. Vitis vinifera. americanum sensu lato and PRMV by X. Concord it delays bud burst. Alternative weed hosts that have epidemiological significance are known for ToRSV. are endemic in North America and thought to be native of the region. Peach rosette mosaic virus (PRMV) in V. BLMoV. Transmission: These viruses are all transmitted by grafting and mechanical inoculation.g.) Main symptoms: Symptomatological responses of grapevines vary according to the species (i.). ELISA and molecular tools are useful for testing field-infected material. jaunissement des nervures.15 x 106 (RNA-2). depérissement de la vigne (Fr. but not conclusive. sedimenting as three components. most V. malformation and mottling of the leaves. No vector is known for BLMoV.35 x 106 (RNA-1) and 2. induces fanleaf-like symptoms on leaves and canes. exhibiting stunted growth. labrusca. Their coat protein consists of a single type of subunits with Mr of about 54 x 103 Da. The genome is a bipartite. Virus particles are isometric. Blueberry leaf mottle virus (BLMoV) infects latently European grapes. California) yield but not vigour is affected. labrusca causes a severe disease characterized by delayed bud burst. BLMoV is a definitive nepovirus species assigned to subgroup C. Infected vines decline slowly over time. V.). single-stranded RNA occurring as two separately encapsidated functional molecules with Mol . 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. and poor fruit setting Agent: The above mentioned four distinct nepoviruses. interestingly. Long distance spread takes place primarily through infected propagating material. All roostocks and. straggly and shelled clusters. wt of 2. deperimento della vite. ToRSV and BLMoV are also seed transmitted in grapes. Description Main synonyms: Yellow vein. and poor fruit setting. labrusca cv. TRSV and PRMV.

. E. Healthy grapevines become infected when planted in soils from diseased vineyards. The virus is transmitted through seeds in grapevines and C. The vector is unknown. PRMV is seed-borne in both naturally infected dandelion (4% of infected seeldlings) and in artificially infected C. As the virus is endemic and seed-borne in some perennial weeds Taraxacum officinale (dandelion). 1994. Concord grapes are apparently virus-free but 9. Description Peach rosette mosaic virus (PRMV) is named after the disease induced in peach. Grapevines (Vitis labrusca) are infected in New York State (USA) by a serologically distinct strain of the virus. Phytopathology 71.Use of resistant roostock hybrids and of certified planting material is recommended. elongatus has also been reported. 145-156 Ramsdell D. wt of 2. 1977. Hancock. 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. Physical and chemical properties of the particles and ribonucleic acid of blueberry leaf mottle virus. Vines are stunted and show a progressive decline. but not eradicate. The genome is a bipartite positive sense single-stranded RNA occurring as two separately encapsidated functional molecules with Mol. Hummer. Plant Disease Reporter 61. : Partial nucleotide sequence of BLMoV RNA-1 and RNA-2 1981 1994 3.W.5% of seedlings from seeds taken from diseased vines proved to be infected. but identified as a strain of GBLV.2 x 106 (RNA-2) accounting for 44% and 37% of the particle weight.4 x 106 (RNA-1) and 2. 1981.F. and R. Virus particles are isometric. 2. respectively. Occasional.F. PRMV can also be introduced in a site by infected planting material and be spread by vectors to adjacent vines. which may lead to their death. Vectors are the Dorylaimoid nematodes Xiphinema americanum sensu lato and Longidorus diadecturus. 949-953. 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. Their coat protein consists of a single type of subunits with Mr of about 57 x 103 Da. quinoa (90% of infected seedlings). The virus is seed-transmitted in grapevines and C. mostly to vines adjacent to previously infected plants. labrusca cultivars (Concord. Clusters are straggly. vector populations . 468-472. The virus is a definitive nepovirus species assigned to subgroup C. References Bacher J. about 28 nm in diameter with angular outline. Uyemoto J. Historical review 1977 Uyemoto et al: BLMoV isolated from New York 'Concord' vines showing fanleaf-like symptoms. sedimenting as three components. possibily non specific transmission by L. Isolation and identification of a strain of grapevine Bulgarian latent virus in Concord grapevines in New York State. especially) and a number of American-French hybrids have been recorded. Stace-Smith. mottled and variously deformed leaves and delayed bud burst. D. Solanum carolinense (Carolina horse nettle) and Rumex crispus (curly dock). Warkentin. smaller and fewer than normal .Rasmdel and J. Infected grapevines show shortened and crooked shoots. Pollen grains of cv. and with extensive shelling of the berries.termini of both RNA molecules has been determined. which induces fanleaf-type symptoms and is distantly related to GBLV. but in highbush blueberry the virus is pollen-borne and suspected to be pollen-transmitted . Crop losses up to 60% and death of susceptible V. D. RNA1 is 8004 nt in size and contains a single open reading frame encoding a polypeptide with Mr of 240 kDa. Sequence analysis of the 3' temini of RNA 1 and RNA 2 of blueberry leaf mottle virus. PEACH ROSETTE MOSAIC VIRUS (PRMV) 1.K. and has no economic importance.. when a vineyard is planted susceptible cultivars may become infected by nematode vectors.K. one of its crop plant hosts.C. Virus Research 33. 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. where the disease occurs in more or less circular patches and spreads slowly. 52 . PRMV is soil-borne. Taschenberg and D.

: Longidorus diadecturus transmits PRMV to grapevines. and D.2.A Ebsary. 1972a.. Ramsdell and Myers: Description of PRMV-induced grapevine degeneration and association of X. Canadian Journal of Botany 54. 1-5 Allen W. Allen and Ebsary: Longidorus attenuatus transmits PRMV non specifically and with low efficiency. Carolinense. Rasmdell et al. Dias: Grapevine and peach strains of PRMV can be differentiated serologically. 1982. 1972b. Historical review 1972a 1972b 1974 1976 Dias: Preliminary characterization of the grapevine isolate of PRMV. Ramsdell and Myers: Field spread of PRMV is associated with the presence of infected weeds (T.. J. J. 1988.G Van Schagen and E. References Allen W.G Van Schagen and B. americanum with the disease. 29-32 Allen W. crispus) and transmission through grapevine seeds. officinale. Canadian Journal of Plant Pathology 6.R.. Transmission of raspberry ringspot. Numero hors sèrie. Dias and Cation: Biological characterization of the grapevine strain of PRMV.: Investigation on the susceptibility to PRMV infection of American and European grapevines and hybrid rootstocks. 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. 105-106. The virus is seedborne in C. Annales de Phytopathologie. 1984. S.. Ramsdell: Review article on PRMV. Canadian Journal of Plant Pathology 10. Cation. Transmission of peach rosette mosaic virus by Ontario populations of Longidorus diadecturus and Xiphinema americanum (Nematoda: Longidoridae). 97-103. 53 . Allen et al. 1978 1979 1980 1982 1983 1984 1985 1988 1988 1995 1998 1999 3. Ramsdell and Gillet: Description of PRMV in the AAB Descriptions of Plant Viruses.S Everleigh. Dias and Allen: Physico-chemical characterization of PRMV.R.F. Lammers et al. Transmission of peach rosette mosaic virus to peach grape and cucumber by Longidorus diadecturus obtained from diseased orchards in Ontario. 1228-1239. The characterization of a virus responsible for peach rosette mosaic and grape decline in Michigan. tomato black ring and peach rosette mosaic viruses by an Ontario population of Longidorus elongatus. 1976.F. Dias H. Dias H. Ramsdell and Gillet: List of grapervine cultivars and roostocks showing differential susceptibility to PRMV. quinoa and has reproduced in part the field syndrome when inoculated mechanically to Concord grape seedlings.F. Purification and some characteristics of peach rosette mosaic virus (grape isolate).: Nucleotide sequence of RNA-1 of the grapevine strain of PRMV.R. R. Dias H.: Xiphinema americanum is an efficient vector of PRMV. Ramsdell et al: Use of ELISA for PRMV detection in grapevines. Annales de Phytopathologie. and B. Canadian Journal of Plant Pathology 4. Strains of peach rosette mosaic virus differentiated by cross absorption and immunodiffusion tests.A Ebsary. Allen et al. Numero hors sèrie. 16-18.

Phytopathology 64. J. wt of 2. American Phytopathological Society Press. and B). R. Coat protein consists of a single type of subunits with Mr of about 57 x 103 Da. Characterization of the single protein and the two nucleic acids of peach rosette mosaic virus. 1999.C Ramsdell. Susceptibility of American grapevine scion cultivars and French hybrid roostocks and scion cultivars to infection by peach rosette mosaic virus. TRSV has a relatively wide natural host range. 1988.). 1974.C. Uyemoto et al. There is no evidence of seed trasmission in the grapevine.W. Phytopathology 68.E Morris. 51-52.M.W. Historical review 1970 1977 1985 1985 1986 1990 Gilmer et al. 1985. J. Myers. Myers. RNA-2 has been sequenced only in part.: A review of viruses infecting grapevines in New York vineyards.3 x 106 (RNA-2). Gillet.. Bird. 1995. 4143. AAB Descriptions of Plant Viruses. 447-450 Ramsdell D. 1979. G.C.M. Relative susceptibility of American. R. In: Compendium of Grape Diseases (R..L. 2. Goheen eds. M. Plant Disease Reporter 63. 1174-1178. Peach rosette mosaic virus decline.C. Gillet.C. Phytopathologia Mediterrenea 24.7 x 106 (RNA-1) and 1.. and W. is endemic in Central and Eastern North America. sedimenting as three components (T. American Vitis species reported to be resistant to ToRSV and TRSV. and J. .M.R. Peach rosette mosaic virus. Ramsdell D. RNA-1 is 7514 nt in size and contains a single open reading frame encoding a polypeptide with Mr of 225 kDa.M. Ramsdell D. French hybrids and European grape cultivars to infection by peach rosette mosaic virus. Buzayan et al. Pearson and A. Peach rosette mosaic virus. Plant Disease 67. A comparison between enzyme-linked immunosorbent assay (ELISA) and Chenopodium quinoa for detection of peach rosette mosaic virus in 'Concord' grapevines. respectively. 54 .C.H. TOBACCO RINGSPOT VIRUS (TRSV) 1. Andrews. Epidemiology of peach rosette mosaic virus in a Concord grape vineyard.C.: Survey of ToRSV and TRSV in Pennsylvanian vineyards.C.: TRSV agent of a new grapevine disease in New York State. Canadian Journal of Botany 58. but in European grapes responses are similar to those elicited by GFLV.C. Ramsdell D. Preventive control measures are the use of resistant roostock hybrids and of certified planting material. No. Ramsdell D. Ramsdell D.: Nucleotide sequence of TRSV satellite RNA. 1980.L. G. 57-73. 364. Gillet and C. The virus supports the replication of a circular satellite RNA 359 nt in size.C. Stace-Smith: Description of TRSV in the AAB Descriptions of Plant Viruses series. 625-627. and J.W. and R. respectively. Description Tobacco ringspot virus (TRSV) is the type species of the genus Nepovirus and the prototype of subgroup A. Gillet and L.. Ramsdell D. 1983. Cloning an sequencing of peach rosette mosaic virus RNA1. Plant Disease 79. 74-78. Powell et al.M.C. Paul. Ramsdell D. symptomatology and nematodes associated with grapevine 'degeneration' in Michigan. TRSV is soil-borne and is transmitted by Xiphinema americanum sensu stricto. St. but was recorded from grapevines only in New York State and Pennsylvania. and R. accounting for 44% and 28% of B and M particle weight. Virus particles are isometric. Allen. 1978. Rose. 154-157. Gillet and G. Virus Research 65.F Allison and D.M. 1998. Bird GW.C. Foster and Morris-Krsinich: In vitro translation of TRSV RNA-1 and TRSV RNA-2 yields major polypeptides with Mr of 225K and 116K. Symptoms elicited by TRSV are the same as those of ToRSV in native cultivars. about 30 nm in diameter with angular outline. Lammers A. 1747-1754.Dias H.. The genome is a bipartite positive sense single-stranded RNA occurring as two separately encapsidated functional molecules with Mol. Superimposed shallow and deep soil fumigation to control Xiphinema americanum and peach rosette mosaic virus reinfection a Concord vineyard.

M. Nucleotide sequence and in vitro expression of the capsid protein gene of tobacco ringspot virus. Nucleotide sequence of satellite tobacco ringspot virus RNA and its relationship to multimeric forms. Buzayan J. Disease named yellow vein. mottled and distorted leaves and short internodes. sedimenting as three components (T. Virus and virus-like diseases affecting grapevines in New York vineyards. Stace-Smith R.M. Description Tomato ringspot virus (ToRSV) is a definitive species in the genus Nepovirus and the prototype of subgroup C. Morris-Krsinich. Infected vines have small. 1986. "yellow vein" being the characterizing syndrome of its infections. which often leads to death.L..B.M. 309. Synthesis and processing of the translation products of tobacco ringspot virus in rabbit reticulocyte lysates. Two serological ToRSV variants are known to infect grapevines. Plant Disease 74. especially if self-rooted. Vines grow vigorously but bear little or no fruit. Gilmer R. V.. 2.S. References Buckley B. Gould.K. 702-704. the infecting virus strain.S. Buzayan and G. smaller and fewer than normal. AAB Descriptions of Plant Viruses. Chemical cleavage of the 5'-linked protein of tobacco ringspot virus genomic RNAs and characterization of the protein-RNA linkage. 516-519. 1996. californicum in California The virus can be introduced in a site by infected planting material and be spread by vectors to adjacent vines. Preventive control measures are the use of resistant roostock hybrids and of certified planting material. Clusters are straggly. The yellow vein strain of the virus is pollen-borne but is not transmitted through seeds.R. Historical review 1954 1956 1962 Hewitt: Report of an "unfruitful vine" condition in California to which a yellow speckling of the leaves is associated. ToRSV has a relatively wide natural host range and is endemic in North America. Bruening.R. Silva and S. Both RNAs contain a single open reading frame encoding polypeptides with Mr of 244 kDa (RNA-1) and 207 kDa (RNA-2). 335-349. wt of 2. Longenecker and L. about 30 nm in diameter with angular outline. Symptomatological responses vary according to the species (V.L. : Partial nucleotide sequence of TRSV RNA-2. Virology 151. rivesi in north American States and Canada and X. Tobacco ringspot virus. Incidence of tomato rigspot virus and tobacco ringspot virus in grapevines in Pennsylvania. and B).A. and the climatic conditions. 1977. 55 .1993 1996 Buckley et al. J. Gerlach G. Hewitt: Successful graft transmission of unfruitful vine condition.M. where it occurs iin the region of the Great Lakes and in the Pacific seaboard from California to British Columbia. Virus Research 30..M. Virus particles are isometric. ToRSV-induced decline affects European cultivars. TOMATO RINGSPOT VIRUS (ToRSV) 1. 1985.. 1990. respectively. Vines are stunted and show a progressive rapid decline. 619-627. Forer. Virology 144. 1-8. W. Cummins and G. Coat protein consists of a single type of subunits with Mr of about 58 x 103 Da. and B. Keese and A. P.: Nucleotide sequence of TRSV RNA-1 3. Kelts.K. interspecific hybrids). B. J. Foster R. Singh. In California ToRSV affects the yield rather than the vine's growth. The virus has been occasionaly recorded from grapevines outside of North America..L. No. Zalloua P.A. 131-136. contrary to the decline strain which is seed-transmitted. Vectors are the Dorylaimoid nematodes Xiphinema americanum sensu stricto and X. Uyemoto J.4 x 106 (RNA-2) accounting for 44% and 41 % of the particle weight. labrusca. Virology 219. more severely in colder than in warmer climates. Phytopathology 60. Uyemoto and L. 1993. American Journal of Enology and Viticulture 28. Abawi. Powell C.8 x 106 (RNA-1) and 2. A new grapevine disease induced by tobacco ringspot virus. Bruening. Gooding and Hewitt: A mechanically transmissible virus found to be associated with yellow vein. and with extensive shelling of the berries. J. The genome is a bipartite positive sense single-stranded RNA occurring as two separately encapsidated functional molecules with Mol.J. J. 186-199. ToRSV is soil-borne. RNA-1 is 8214 nt and RNA-2 is 7273 nt in size. vinifera.. Zallua et al. 1985.A. 1970.

: Xiphinema rivesi identified as the main vector of ToRSV in Ontario vineyards.: Complete nucleotide sequence of ToRSV RNA-2. Uyemoto and Gilmer: Spread of ToRSV through the soil of New York State vineyards recorded.: ToRSV found in grapevines in Taiwan. Baumgartnerova and Subikova: ToRSV recorded form grapevine in Slovakia. 56 . Bays and Tolin: ToRSV found in grapevines in Virginia Powell et al. Dias: Record of ToRSV in the Niagara peninsula.: Complete nucleotide sequence of ToRSV RNA-1.1963 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. Martelli: Review of nematode-borne viruses of grapevines and their epidemiology. Uyemoto: Seed transmission of the decline strain of ToRSV.: Confirmation that the grape yellow vein and the the grape decline strains of ToRSV are serological variants of the same virus. American Vitis species reported to be resistant to ToRSV and TRSV. Stace-Smith and Ramsdell: Review of nepoviruses of the Americas. Podlekis and Corbett: ToRSV is the agent of little grape disease in Maryland.: List of grapevine roostocks and cultivars showing differential susceptibility to ToRSV in Canada. Martelli and Taylor: Review of nematode-borne viruses and their vectors. Li et al: ToRSV identified in China in grapevine seedlings grown from seeds imported from France. Corbett and Podleckis: Ultrastructural study of ToRSV-infected grapevine tissues. Rott et al. Allen et al. Rott et al.: Transmission of the yellow vein strain of ToRSV by X. Bitterlin and Gonslaves: ToRSV retained and transmitted by viruliferous Xiphinema rivesi stored for two years at 1-3°C. Teliz et al. Allen et al. americanum (now X. Yang et al. Uyemoto et al. Gilmer and Uyemoto: ToRSV agent of a decline of Baco noir in New York State. Herrera and Madariaga: ToRSV recorded from grapevine in Chile. Allen and Dias: Physico-chemical characterization of ToRSV. Stace-Smith: Description of ToRSV in the CMI/AAB Descriptions of Plant Viruses series. Rowhani et al: Description of sampling strategy for detection of ToRSV. Cory and Hewitt: The yellow vein strain of ToRSV is not transmitted through seeds.: Survey of ToRSV and TRSV in Pennsylvanian vineyards. californicum).: A review of viruses infecting grapevines in New York State vineyards. Piazzolla et al. Gonsalves: ToRSV is irregularly distributed in infected vines but can be detected by ELISA.

Corbett.F.G. Li M.M. and grape yellow vein viruses by Xiphinema americanum.F. Grape yellow vein: symptomatology.. Niagara Falls 1980.M.K. 57 . 35-44.3. 475-480.. 151-189. and D. Vitis 31.G. Xiang .K. Musci. Tolin. Walker and S. and S. Martelli G.F. Forer. Gooding G. Dias and J. identification. Castellano and D. Plant Disease Reporter 56. 1987. Allen W.. Téliz D. J. Nematode-borne viruses of grapevine. G.. Nucleotide sequence of tomato ringspot virus RNA 1. Plant Disease 71.S Whei.F. 1978. Piazzolla P. 24-28. Van Schagen and B. Lee and D'A. Transmission of tomato ringspot. 275-277. Tomato ringspot virus associated with little grape disease of Vidal 256 grapevines.. Ramsdell. 1990. Nematologia Mediterranea 6. Gonsalves. Purification and serology of a virus associated with the grape yellow vein disease. Tremaine and D'A.B. Stace-Smith R. and E. Ebsary. 1966. H.V..V. Distribution of viruses and their nematode vectors. 702-704. Y. Advances in Disease Vector Research 6. and V.H. 1982. 1972. Hewitt. Phytopathologia Mediterranea 24. 408-411. 1988. Baumgartnerova H. Van Schagen. Uyemoto J. Tomato ringspot virus in "Baco noir" grapevines in New York. Powell C. Corbett M. Susceptibility of grape cultivars and rootstocks to an Ontario isolate of tomato ringspot virus. and J.A. Gonsalves D. and W. and the association of a mechanically transmissible virus with the disease. 157-164. 31-34. 1995. Nepoviruses of the Americas. Podleckis E. 710. Rott M.C. Journal of General Virology 76. Phytopathology 79. Rochon.V. M. 1505-1514.P.W.K. 1169. 1-27.. Proceedings 7th Meeting of IGVG. 861-863. 1989. Rochon.F.. Gilchrist. Bulletin of the California Department of Agriculture 43.. Plant Disease Reporter 61. A.L. 290 Stace-Smith R. 44-50. Stobbs. Bitterlin M. No. Zhang and Y. Rott M. 1982. Dias. J. Spatial distribution of Xiphinema rivesi and persistence of tomato ringspot virus and its vector in soil. and W.G. Phytopathology 72. Detection and identification of plant viruses for quarantine in China. L. Uyemoto.F. 13161320 Dias H. 1987. J. Phytopathologia Mediterranea 24.. Tomato ringspot virus.F. Gilmer R. Agricultura Tecnica 61. Journal of General Virology 72. their epidemiology and control.A. Plant Disease 72. Savino. R. Phytopathology 53. 131-166. Works of the Institute of Experimental Phytopathology and Entomology . 2004. Herrera M. 1985. Incidence of tomato rigspot virus and tobacco rigspot virus in grapevines in Pennsylvania.W. Podleckis.B. Identification of tomato rinsgspot virus in leafroll diseased grapevines. V. 1993. CMI/AAB Descriptions of Plant Viruses.R.A. Membrane-associated spherical particles in extracts and tissues of virus-infected grapevines.V. 1968..A. Gooding G. M.P. Plant Disease Reporter 59. 196-203. M. Phytopathology 56. 658-663. 1975.R. Incidence and geographic distribution of tomato ringspot virus in De Chaunac vineyards in the Niagara peninsula.E.. 1984. and B. Longenecker and L. Subikova. American Journal of Enology and Viticulture 13. Chen. 1962. peach yellow bud mosaic. 1985.G and V. and C. Bays D. A severe outbreak of virus-induced grapevine decline in Cascade grapevines in New York. Li. 1954. Beijing 2004. Incidence of tomato ringspot virus in grape in Virginia. 95-106. L.C. N. Properties of the single protein and two nucleic acids of tomato ringspot virus.. 1980. Canadian Journal of Botany 55.. 393-400 Hewitt W. Taylor.E. Canadian Journal of Plant Pathology 4. 47-64. Proceedings 15th International Plant Protection Congress. 1977. Detection of tomato ringspot virus in grapevines: irregular distribution of virus. Sampling strategies for the detection of grapevine fanleaf virus and the grapevine strain of tomato ringspot virus.B. Association of Xiphinema species with soil types ang grapevines infected with tomato ringspot virus in Ontario. Cory L.. 1963. Madariaga.K. and M. Some virus and virus-like diseases of grapevines.A. 1991.J. Rokni. 1028-1037. and H.. A comparison of Grapevine yellow vein virus and a Grapevine isolate of Tomato ringspot virus. 98-101. 1992. Allen W. Grogan and B.E. Plant Disease 74. Nucleotide sequence of tomato ringspot virus RNA-2. References Allen W. Phytopathology 58. Current Topics in Vector Research 5. 1989. Lownsberry. 663 Martelli G. 465-473.B. 1977.R. 2001. Hewitt. Some grapevine viruses in pollen and seed. Rowhani A. 133-135. Canada. Presence and incidence of grapevine viruses in central Chile. Ivanka pri Dunaji 4.

Cummins and G. J. 131-136. Abawi.K.S.K. 1062-1064. 58 . T. Spread of tomato ringspot virus in "Baco Noir" grapevines in New York. American Journal of Enology and Viticulture 28..L.C. and R.J.M. Sap-transmissiible viruses associated with grapevine yellow mottle disease in Taiwan. Gilmer.. Deng and M.Uyemoto J. 1977. Virus and virus-like diseases affecting grapevines in New York vineyards. 1986. Chen. Uyemoto J. Plant Disease Reporter 56. Yang I. 1972.R. Journal of Agricultural Research China 35. 504-510.

LEAFROLL .

.

accartocciamento fogliare (Ital. Blattrollkrankheit (Germ. in autumn. These spots enlarge with time and coalesce so that. the symptoms are similar. changes in the pattern of peroxidase and polyphenoloxidase isoenzymes. In white-berried cultivars of V.GRAPEVINE LEAFROLL 1. whereas the Mr of all other viruses ranges between 35 and 44 kDa. All GLRaVs belong in the family Closteroviridae. accartocciamento. which are differentiated from one another by a progressive number. as estimated by polyacrylamide gel electrophoresis. Virus particles are very flexuous filaments about 12 nm wide. Rollkrankheit. These negative effects are reverted if the disease is eliminated by sanitation treatments. Also plant anatomy is affected. i. enrollado (Sp. especially the phloem.5 nm. In most cases. Until 1995. 61 . A number of other physiological parameters are affected. Main synonyms: White Emperor disease (Eng. differing somewhat in aspect and in severity. Leafroll decreases grapevine yield (by 15-20% in average) and affects negatively rooting ability. and lowering of sugar content. they are inferior in quantity and quality. and low in sugar. Enrolamento de la folha (Port. 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. Description The first descriptions of grapevine leafroll date back to the mid 19th century. infection of rootstocks is symptomless. vinifera. exhibiting open structure and distinct cross banding with a pitch of about 3. brittle and rolls downwards. the whole leaf surface becomes deep purple. Leafroll is no less important than fanleaf in economic importance.).e. but the leaves become chlorotic to yellowish. enrollamiento de la hoja.). 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. These symptoms progress towards the top of the canes as the season advances. reduction of protein content. whereas GLRaV-7 is presently classified as unassigned species to the family. GLRaV-2 CP has a Mr of 24 kDa. Particle length varies from 1400 to 2200 nm according to individual viruses. Sieve elements are obliterated and crusheds.) Main symptoms: In red-berried cultivars of Vitis vinifera reddish spots develop in the lower leaves in late spring or summer. called grapevine leafroll-associated viruses (GLRaV). GLRaV-5. Starch accumulates in degenerated chloroplasts causing increased thickness and brittleness of the leaf blades.). thus impairing carbohydrate translocation from foliar parenchymas. Agents: To date.). usually leaving a narrow green band along the primary and secondary veins. GLRaV-2 in the genus Closterovirus. thus suggesting that there can be several causal agents. Also the composition and aromatic profile of the musts are modified. and GLRsV-9 in the newly established genus Ampelovirus. GLRaV-6. and with many cultivars. In the most severe cases. instead of reddish.). Its occurrence in viticultural areas is worldwide. Other such viruses may exist. the risk of disseminating the disease is great if untested rootstocks are used. potassium depletion in the leaf blade and accumulation in the petioles. Sizes deduced from the nucleotide sequence of the CP cistron are 22 kDa for GLRaV-2 . respectively. Careful observation of field symptoms in infected vines reveals that there are several types of leafroll. enroulement (Fr. have been found in leafroll-infected vines. GLRaV-8. depending on the climate and geographic location. differentiation of GLRaVs at the species level was by Roman numerals. most of the leaf surface becomes reddish. the same as the size of coat protein (CP) subunits. except for a variable decrease in vigour. and is probably the most widespread virus disease of grapevine. Hence. nine different viruses with filamentous particles. GLRaV-3. GLRaV-4. graft take and plant vigour. GLRaV-1. The leaf blade becomes thick. The fruits often mature late and irregularly.

singlestranded. which is the type species of the novel genus Ampelovirus has a genome 17. 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. soil condition and probably. 'Cabernet Franc' 'Pinot noir'. and some are commercially avaliable. the type member of the genus. and has structural organization differing from that of other sequenced closteroviruses. vinifera varieties show symptoms most clearly because of the reddening of the leaves. only vectors of GLRaV-1. Natural field spread of leafroll disease has been reported from many countries in Europe and elsewhere. Foliar tissues are not recommended for serological GLRaVs detection in American Vitis species and roostocks. grafted vines) which is largely responsible for its dissemination over medium and long distances. number an types of infecting viruses. GLRaV-5 and GLRaV-9 are phylogenetically close to one another but are serologically unrelated. positive sense RNA molecule. biologically. with none of which they are serologically related. Red-berried V. contains 13 ORFs.528 nt in size. comstocki. Varietal susceptibility and sensitivity: No immune variety or rootstock is known. maritimus. GLRaVs differ in various vays (molecularly. As to nucleic acid-based assays. classical double antibody sandwich ELISA (Chromo-ELISA) or Lumino-ELISA. Parthenolecanium corni. Indexing on red-fruited cultivars such as 'Cabernet sauvignon'. Detection: In many cases. 'Merlot'. and Neopulvinaria innumerabilis. Other GLRaVs may exist in nature as suggested by reports.5 kDa for GLRaV-4 . American rootstocks are usually symptomless carriers of GLRaVs. Mealybug vectors of GLRaV-3 are Planococcus ficus. Membranous vesicles can derive eitrher from peripheral vesiculation of mitochondria followed by disruption of the organelles (GLRaV-1. The genome of GLRaV-4 and GLRaV-9 are incompletely sequenced but show a structural organization compatible with that of the genus. citri. in agreement with the quasispecies nature of viruses. All GLRaVs can be identified by serological and nucleic acid-based techniques. Ps. calceolariae. GLRaVs show molecular variations which give rise to a population of strains. and epidemiologically) from most of the known closteroviruses.35 kDa for both GLRaV-3 and GLRaV-1. GLRaV-2. Polyclonal antisera and/or monoclonal antibodies have been raised to each single GLRaV . Composite samples should be used to minimize false negative responses that may originate from the unven distribution of GLRaVs in chronically infected vines. The genome of GLRaV-1 is 17. affinis. Ps.Transmission is semipersistent and does not appear to be vector-specific.647 nt in size and contains 10 major ORFs. 29. the only member of the group to be mechanically transmissible. Ps. Regardless of whether they belong to the genus Closterovirus or Amplelovirus. or the hybrid LN 33 is still the most popular method for identifying the disease. GLRaV-3. None of the GLRaVs is known to be seedborne. or by molecular techniques. The presence of high molecular weight double-stranded RNAs (dsRNA) in phloem tissue extracts can be used as infection marker. ultrastructurally. GLRaV-3. So far. 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. has a number of minor biological variants which can be differentiated by the reaction of inoculated Nicotiana species. roostocks. GLRaV-1 is transmitted in nature by the pseudococcid mealybugs Heliococcus bohemicus and Phenacoccus aceris and the soft scale insects Pulvinaria vitis. GLRaV-2 and GLRaV-3. climate. and some of them are used as indicators. Ps. and degenerate primers have been designed and successfully used for PCR detection of virtually all GLRaVs. leafroll can be detected by its symptoms in the field on red-fruited varieties. The genome of GLRaV-2 is 15. vinfera and cortical shavings from mature dormant canes of V. broadspectrum. viburni and Ps. Its soft scale insects vectors are Pulvinaria vitis and Neopulvinaria innumerabilis. Disappearance of dsRNAs from vines submitted to 62 . vinifera. GLRaV-5 and GLRaV-9 have been identified. longispinus. These reagents are routinely used for ISEM. but it does not discriminate between GLRaVs and was reported to be less sensitive than ELISA. GLRaV-5 and GLRaV-9 are both transmitted by Ps. American Vitis species and rootstocks are the best antigen sources for serological assays. GLRaV-3. This has been ascertained experimentally for GLRaV-1.919 nt in size. Symptom expression depends on the variety. The genome is a monopartite. Spread at a site is mediated by mealybug and soft scale insect vectors. Leaf tissues or petioles from mature symptomatic leaves of V. Pl. 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. 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. Pseudococcus longispinus. contains nine open reading frames (ORF) and has structural organization identical to that of Beet yellow virus (BYV).

Tanne and Nitzany: Leafroll in Israel Tanne et al. vinifera and there is no published information on how to protect healthy stocks from vector-mediated reinfection in the field. Ravaz and Verge: Occurrence in France of “rougeau”. 1971 1973 1974 1975 63 . The incidence if the disease was less than 20% as compared with 80 to 100% in commercial vineyards. Scheu: Leafroll is widespread in German vineyards. roostocks are suggested as the major souces of leafroll dissemination. 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”. Dimitrijevic: Leafroll in Yugoslavia. 2. dsRNAs cannot be utilized for virus identification. Hypothesis of the viral origin of leafroll. a grapevine disorder similar to leafroll.: White Emperor and leafroll are identical diseases.sanitation treatments is regarded as evidence for successful virus elimination. Control: Production and use of clonally selected and sanitized propagation material is very effective and the only preventive method for leafroll control available. Later studies showed that the virus is an occasional contaminant Lider et al. These particles are probably closteroviruses associated with leafroll. Goheen et al. Vuittenez: Leafroll in France. Harmon and Snyder: The "White Emperor" disease is graft-transmissible and is regarded a virus disease. Luhn and Goheen: Leafroll found in the original grapevine stocks imported from Europe into California in 1890. Blattny et al. Lehoczky et al. However. No sources of resistance are known in V. Chamberlain: Leafroll in New Zealand. Bovey: Leafroll in Switzerland. Fraser: Leafroll in Australia. Introduction of transgenic resistance to GLRaV-2 and GLRaV-3 is being attempted by engineering different viral genes into rootstocks and European grape cultivars.: Transmission of a virus to herbaceous plants from a leafroll-infected vine in Israel.: Leafroll in Hungary. Belli et al.: Leafroll in Czechoslovakia. As no apparent spread of the disease was observed.: Studies on the effects of leafroll on yield of grapevines in California.: Leafroll in Italy.: Leafroll virus can be inactivated in vivo by heat therapy. Hewitt: Leafroll in California Goheen et al. 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. Hoefert and Gifford: Study of the effects of leafroll infection on vine anatomy. unless they are hybridized with virus-specific probes.

Martelli et al.: ELISA testing reveals that GLRaV-3 has an uneven distribution in grapevine tissues. Sasahara et al. Rosciglione and Gugerli: GLRaV-3 is transmitted by the mealybug Planococcus ficus. Zimmermann et al. Teliz et al. Castellano et al. Hu and Gonsalves: Monoclonal antibodies produced against GLRaV-3.: Closterovirus-like particles purified from leafroll-diseased grapevines in France. A large dsRNA molecule is consistently isolated from leafroll-diseased grapes. Gugerli et al.like particles. Tanaka: Leafroll in Japan. Purification and serology of associated closterovirus-like particles. A third closterovirus type called GLRaV-3. Namba et al. Zee et al. 1981 1981 1982 1982 1983 1984 1984 1984 1985 1986 1986 1987 1987 1988 1988 1989 64 . Rosciglione and Gugerli: GLRaV-1 and GLRaV-2 with particles of 2200 nm and 1800 nm respectively.: Elimination of leafroll by in vitro meristm tip culture and apex fragmentation.: Studies on the cytopathology of leafroll-diseased grapevines.: First record of successful elimination of leafroll in grapevine by using meristem tip culture in Japan.: 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. Kliever and Lider: Study of biochemical changes found in grapevine infected with leafroll in California. Barlass at al. Antiserum against a New York isolate also reacted with GLRaV-3 from Europe. Confirmation that GLRaV-3 and the New York closterovirus isolate cross react serologically.: 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. are also present in leafroll-affected grapevines from Italy. but not in similar praparations from healthy plants. Mossop et al. found in grapevines affected by leafroll. Abracheva: Leafroll in Bulgaria. Production of polyclonal antisera for use in ELISA. Absence of such particles in healthy grapevines. previously found in grapevines in Switzerland. Hofmann: Symptoms of leafroll in affected clones of Pinot noir and performance in West Germany. Production of rabbit and hen antibodies for ELISA and ISEM to GLRaV-1 and GLRaV-3.: Closterovirus-like particles and specific dsRNA found in leafroll-diseased grapevines in New Zealand. but not in healthy controls.: Review on the detrimental effects of viral infection on grapevine physiology.: Aggregates of closterovirus-like particles observed in thin sections of phloem from leafroll-diseased grapevines. Von der Brelie and Nienhaus: Light and electron microscope study of cytopathological changes induced by leafroll in grapevines. Faoro 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.: Ultrastructural study of leafroll-infected grapevine tissues. Presence of virus-like particles in thin sections of leafroll-diseased vines. Corbett et al: Electron microscope observations by negative staining of leaf extracts from leafroll-diseased grapevines in South Africa showed the presence of closterovirus. Suggestion that a closterovirus may be the agent of the disease.

: Characterization of leafroll-associated closterovirus-like particles from grapevine using also monoclonal antibodies. Boscia et al. was detected in P. Identification of GLRaV-4 1990 1990 1990 1990 Walter et al. whereas meristem tip culture yields up to 100 % sanitation Walter and Zimmermann: Further characterization of closteroviruses associated with leafroll in France.: Production and characterization of monoclonal antibodies specific to GLRaV-3. Transmission of GLRaV-3 by the mealybug Planococcus ficus. Credi and Santucci: GLRaV-1 and GLRaV-3 cannot be detected by direct ELISA in leaves of graft-inoculated American rootstock. Gugerli et al. Good results in summer with extracts of basal leaves and in autumn or winter with wood shavings macerated in buffer.1989 1989 Tanne et al. stem pitting and corky bark spread rapidly. later in the leaves. but not GLRaV-1. GLRaV-1. symptoms are obtained within 20-70 days. Some vines indexing positive for leafroll do not react positively with any ofthe antisera. 1989 1989 1989 1990 1990 1990a.: Leafroll and associated closteroviruses in China Engelbrecht and Kasdorf: Transmission of GLRaV-3 by Planococcus ficus from grapevine to grapevine in South Africa. -2 and -3 are common whereas GLRaV-5 is rarely detected.: Leafroll and associated closteroviruses in Chile Kuhn: Leafroll in Brasil Li et al.: Transmission of GLRaV-3 from grapevine to grapevine by the mealybug Pseudococcus longispinus in Israel. In Mexico leafroll. Agran et al. Gugerli et al.: Detection of leafroll-associated closterovirus in recently infected grapevines in New York. GLRaV-2. Téliz et al. ELISA is to be applied to cortical scrapings rather than leaf tissues. GLRaV-1 and GLRaV-2 were not transmitted. especially those containing V. Auger et al.: 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.: Comparison of heat therapy and meristem tip culture for eliminating GLRaV-3 from Italian grape varieties. Heat therapy requires very long treatments and is only 20-30 % successful.: Evidence of the irregular distribution of GLRaV-3 in American rootstocks. Faoro et al. There is evidence that other GLRaVs are involved in leafroll etiology. Zimmermann et al. indicating the presence of other leafroll-associated viruses. Pseudococcus longispinus is present on weeds around diseased vineyards. The virus was detected in root tissues.: Production of monoclonal antibodies to GLRaV-1 and GLRaV-3. ficus fed on infected vines. For reliable testing. rupestris plasma.: Use of green grafting for detecting virus-like diseases of grapevine. 1990 1991 1991 1991 1991 1991 1991 1991 65 . but they are easily detected in inoculated LN33 vines and in V. Savino et al. b Hu et al. Identification of GLRaV-5. Gugerli: Review of grapevine closteroviruses.: Immunocytological detection and localization of GLRaV-1 and GLRaV-3 by immunogold labelling in grapevine thin sections. With leafroll. vinifera varieties used as inoculum source.: Further characterization of GLRaV-1 and GLRaV-3 by monoclonal antibodies.

Jordan: In a New Zealand commercial vineyard GLRaV-3 incidence increased from 9. Krake: Characterization of leafroll disease based on symptoms shown by field-infected vines and graft-transmission tests. 66 .: Comparison of different assay methods for detecting GLRaVs : ELISA. Milkus et al. Kassemeyer: Detection of GLRaVs in Germany. later identified as GLRaV-2. Segura et al. ELISA is recommended for large screening.: Leafroll and associated closteroviruses in Spain 1991 1991 1991 1991 1991 1991 1992 1993 1993 1993 1993 1993 1993 1994a. Minafra and Hadidi: Detection of GLRaV-3 in viruliferous mealybugs by PCR. Habili et al. Tzeng et al. 1994 1994 1994 1994 1994 1995 1994 1995 1995 1995 1995 Merkuri et al. Castellano et al. whereas the other assays are more suitable for analyzing samples that gave inconclusive results with ELISA. Former GLRaV 2 is re-named GLRaV-6.: Leafroll and associated closteroviruses in Moldova. Faoro and Carzaniga: Ustrastructural study of GLRaV-1 and GLRV-3 infections. Martelli et al. Belli et al. Bondarchuk et al. Gugerli and Ramel: Analysis by monolconal antibodies of a Swiss source of cv. Gozsczynski et al.: Leafroll and associated closteroviruses in Albania.1991 Hu et al. Namba et al.: Production of radioactive and non-radioactive molecular probes to GLRaV-3 from denatured dsRNA template and their use for virus identification.1% in 1992 Ioannou: Leafroll and natural spread of associated closteroviruses in Cyprus. denoted GLRaV 2a and GLRaV 2b. Observation of peripherically vesiculated mitochondria. Katis et al: Leafroll and associated closteroviruses in Greece.: Transmission of GLRaV-3 by the soft scale insect Pulvinaria vitis.: Leafroll and associated closteroviruses in Romania. Golino et al.: Transmission of GLRaV-3 by Pseudococcus affinis in California. Chasselas shows the prsence of two different GLRaV-2.b Saldarelli et al.1% in 1988 to 93. Boscia et al.: Revision of the nomenclature of GLRaVs and use of Arabic numerals in the species names. Flak and Gangl: Leafroll and associated closteroviruses in Austria. Boehm and Martins: Leafroll in Portugal.: Purification and physico-chemical characterization of grapevine corky bark associated virus.: Production of antisera to GLRaVs using electrophoretically separated coat protein subunits as antigens.: Analysis for the presence of double-stranded RNAs can be used for assessing virus elimination following sanitation treatments.: Leafroll in Taiwan. ISEM and dsRNA analysis.: Mechanical transmission of GLRaV-2 and ultrastructural study of infected tissues of Nicotiana benthamiana.: Leafroll and associated closteroviruses in Ukraine. Pop et al. Leafroll and associated closteroviruses in Yemen.

Gozsczynski et al.: Development of a spot-PCR technique for GLRaVs identification. Petersen and Charles: Pseudococcus calceolariae acts as vector of GLRaV-3 in New Zealand. MacKenzie et al.GLRaV-5 induces mitochondrial vesiculation.: Association of GLRaV-2 in Italy and France with a graft incompatibility revealed by Kober 5BB.9% in 1996. Faoro: Comprehensive review of the ultrastructure of GLRaVs infections.: Distribution and incidence of GLRaVs in Canadian viticultural districts. Rowhani and Uyemoto: Comparative trials between indexing and laboratory detection methods show that the latter are more sensitve for GLRaVs detection. Ling et al.: Transmission of GLRaV-1 by the soft scale insects Parthenolecanium corni and Neopuvinaria innumerabilis.1% in 1986 to 51.: Identification of two different mechanically transmissibile strains of GLRaV-2. Fortusini et al. Wilcox et al. Haidar et al. Zhu et al. Krastanova et al.: Identification of GLRaV-7 and production of a polyclonal antiserum. Martelli et al. La Notte et al. Cabaleiro et al. 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1998 1998 1998 1998 1998 1998 1998 67 . GLRaV-3 appears to be a typical closterovirus. Viruses are irregularly distributed in the vines.: Extensive sequencing of GLRaV-3 genome. the type specied of the genus Closterovirus.: GLRaV-3 is transmitted by Planococcus citri in a semipersisten manner. Alkowni et al.: GLRaV-2 and GLRaV-3 genes engineered in grapevine rootstocks to induce resistance. Ling et al. Al-Tamimi et al.: GLRaV-3 detected in native American vines in Western New York. Routh et al. Gugerli et al. None of 223 accessions of European. Lahogue and Boulard: Search for genes of resistance in grapevines.: Use of degenerate primers for PCR detection of GLRaV-4 and GLRaV-5.: Leafroll and associated closteroviruses in Palestine.: Serological characterization of GLRaV-6 and production of monoclonal antibodies. American. and Asian Vitis species inoculated by green grafting with a GLRaV-1 and GLRaV-3 sources were resistant. Habili and Nutter: In an Australian commercial vineyard GLRaV-3 incidence increased from 23.: The sequenced genome of GLRaV-2 has the same structural organization of that of Beet yellows virus. Guidoni et al.: Leafroll and associated closteroviruses in Jordan.: Leafroll and associated closteroviruses in Lebanon.1995 1996 1996 1996 1996 1996 Greif et al. No vector was identified.: Cloning an sequencing of the coat protein gene of GLRaV-3 and its expression in transgenic tobacco.: Comprehensive review of the properties of GLRaVs.: Elimination of GLRaV-3 by heat therapy improves agronomic performances of a Nebbiolo clone and the quality of the must. Choueiri et al.

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 . Cardinal in Italy and Greece. Martelli et al.: New vectors of GLRaV-1 are the mealybugs Heliococcus bohemicus and Phenacoccus aceris and the soft scale insect Pulvinaria vitis. Demonstration that the membranous vesicles accumulating in the cytoplasm derive from proliferation of the endoplasmic reticulum. Meng et al. a chemiluminometric enzyme-linked immuosorbent assay.: Evidence that vines sanitized from leafroll have superior qualitative and quantitative traits. Mannini and Credi. Sforza et al.: New monoclonal antibodies to GLRaV-2. Ps.: Leafroll and associated closteroviruses in South Korea.: Survey of GLRaVs in Mediterranean and Near East countries.: Identification and partial characterization of a new strain of GLRaV-2. Boscia et al. Golino et al.: Association of an unusual strain of GLRaV-2 with a graft incompatibility condition described from California as young vine decline. Seddas et al. Ps.: 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. Production of a new set of monoclonal antibodies. one of which is especially suited for ELISA testing. Golino et al.: Revision of the family Closteroviridae. viburni. Little et al. Proposal for the establishment of Vinvirus (Ampelovirus) . maritimus. GLRaV-2 and GLRaV-4. Rowhani et al.: Use of degenerate primers for PCR detection of GLRaV-1 and GLRaV-7. Gugerli: Detection of GLRaVs by Lumino-ELISA. a new genus having GLRaV-3 as type species.: Evidence that GLRaV-1 and GLRaV-3 are serologically related based on the cross-reactivity of a monoclonal antibody raised to GLRaV-1. affinis and Planococcus citri transmit with various efficiency GLRaV-3 in California but not GLRaV-1. Gonsalves: Review article on the molecular traits of GLRaVs. transmitted by mealybugs and soft scale insects. : Partial sequencing of GLRaV-7 genome. Fazeli and Rezaian: Partial sequencing of GLRaV-1 genome.: Completion of the nucleotide sequence of GLRaV-3 genome and use of the HSP90 and coat protein genes for producing transgenic rootstocks. Kim et al.: Mealybug species Pseudococcus longispinus. Establishment and description of Ampelovirus . Zhou et al. Abou Ghanem -Sabanadzovic et al. Karasev: Review article on closteroviruses.: Intriguing association of GLRaV-6 with cv.1998 2000 2000 2000 Saldarelli et al.: Identification of hypervariable regions in GLRaV-1 genome. Turturo et al.: Completion of the nucleotide sequence of GLRaV-2 genome. Monis: Identification of GLRaV-8 and production of monoclonal antibodies. Digiaro et al. Ps. a genus with monopartite RNA species. Castellano et al: Ultrastructural study of GLRaV-2 and GLRV-7 infections. Evidence of the quasispecies nature of the virus. Ling et al.

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Partial molecular characterization and RT-PCR detection of grapevine leafroll-associated virus 7. Vitis 12. The occurrence of grapevine leafroll disease among the main grapevine cultivars and breeding stocks in Taiwan. Phytopathologische Zeitschrift 80. Locorotondo 2003. Gonzales and C. 1924.. Verge. Proceedings 9th Meeting of ICVG. Maladies de l'enroulement et du bois strié de la vigne: analyse microscopique et sérologique.K.P.. Rowhani. Nienhaus.. 135-141. Adelaide 2000. Segura A. Rosciglione B. and J. Minafra and M.L. 1238-1243. Tanne E. Boscia.. 1989. Berlin. A. Harpaz. 71-73. Gugerli 1989. Saldarelli and A. 2000. Turturo C. 1981. 433-436. Field serological detection of viral antigens associated with grapevine leafroll disease. Digiaro. Journal of the Japanese Society for Horticultural Science 50. 157-160. Martelli. Kiryat Anavim 1987. Chen and D. Annals of the Phytopathological Society of Japan 42. M. A. Y. Plant Pathology 49. 176-180.. A. Regeneration of plantlets by meristem tip culture for virus-free grapevine. Use of degenerate primers for partial sequencing and RT-PCR-based assays of grapevine leafroll-associated viruses. P. with special reference to closteroand vitiviruses of the grapevine. Der Deutsche Weinbau 14. Rowhani. Turturo C. 1991.. Teliz D. 1994b. G. Saldarelli. Gonsalves and F. and G. Saldarelli. 156-167. Uyemoto. Proceedings 10th Meeting of ICVG. 82.. Extended Abstracts 13th Meeting of ICVG. 704-709.P. 1987. Histological and cytological studies on the infectious leafroll disease of the grapevine. 169175. Le rugeau de la vigne. Plant Pathology 43. 2000. Transmission of grapevine leafroll disease and an associated closterovirus to healthy grapevine by the mealybug Planococcus ficus Signoret. 67-69. 345-346. Establishment of a relationship between grapevine leafroll closteroviruses 1 and 3 by use of monoclonal antibodies. 1986. 1989 Detection of a grapevine leafroll-associated closterovirus in recently infected tissues in New York and spread of the disease in Mexico. 1998. M. 508-517. J. Progrès Agricole et Viticole 45. Sforza R.A. Vitis 33. H. Yafeng. 38. 192-196. Scheu G. Raccah. and P. 2000. Phytopathology 88. 356358.M. C.. V. Minafra. Teliz D.. Savino V. 1973. D.. 1994a. Minafra. and F. M. Ben-Dov and B. D. Saldarelli P.Ravaz L. Il rossore delle viti.K. 91-96. T.P. 1982a. 222-225 Tanne E. Transmission of grapevine leafroll virus to herbaceous plants. 1993. Jacquet. Rosciglione B. Digiaro. New scale insect vectors of grapevine closteroviruses. Indexing grapes in Japan for viruses. P. Mein Winzerbuch. D'Onghia and G.. Walter. 17-18. 80-85. 1994.. 162-163. Savino and G. 11-17. Phytoparasitica 17. and F.L. Martelli. E. Sasahara H. Takezawa and M. Zhang. Hu and D.D. Zee. Cloquemin and B.. Gonsalves. 1998. Zhang . Haidar. Plant Disease 71. 945-950. Rowhani A. Adelaide 2000. Saldarelli P. Tazaki. Greif. Sannino F.. Y. Martelli and B. Saldarelli P. Plant Disease 81. Rivista di Patologia Vegetale 1. and P. M. Arboriculture. A. Komar and C. Effect of heat therapy and meristem tip culture on the elimination of grapevine leafroll-associated closterovirus type III. W. Cabaleiro. Detection of grapevine leafroll-associated closterovirus III by molecular hybridization. Volos 1990. Hummer.P. Y. Tada.P.P. 1997. Scheu G. Greif. V.. 125-126.K. Extended Abstracts 13th Meeting of ICVG. D. Plant Pathology Bulletin 3. Kiryat Anavim 1987. Die Rollkrankheit des Rebstockes.. Routh G. 74 . Horticulture 18.M.E. Uyemoto. A. 1906. 14. Tzeng H. M. Preliminary investigations of genetic variability of Grapeviune leafroll associated virus 3 isolates.P. 1935. 110-113.C.. Martelli. Guglielmi Montano and G. 8689. 68-69. Martelli. Extended Abstracts 14th Meeting of ICVG. Rowhani A. Nitzany. 1976. Transmission of closterolike particles associated with grapevine leafroll by mealybugs (Pseudoccidae) in Israel. Jelkmann and G. 1974. Reichnährstand Verlag. C. Use of degenerate primers in a RT-PCR assay for the identification and analysis of some filamentous viruses. D. 207211.. 2000. Virus diseses of grapevine in Israel. P. 222-223. Adelaide 2000. Montreux 1993. Presence of grapevine leafroll in North West Spain. Zeitschrift für Pflanzenkrankheiten und Pflanzenschutz 89. A comparison between serological and biological assays in detecting grapevine leafroll-associated viruses. Von der Brelie D. D. I. Iri. G.J. Golino. Rott. Proceedings 9th Meeting of ICVG. Revue Suisse de Viticulture. Tzeng. Sela and I. Seddas A. Non radioactive molecular probes for the detection of three filamentous viruses of the grapevine.S. 799-801. Extended Abstracts 13th Meeting of ICVG. and J. Tanaka S.S. K.. Gugerli.. 1936.A .. European Journal of Plant Pathology 104. Extended Abstracts 11th Meeting of ICVG. 35-38. Walter. Tanne. Isolation and partial characterization of two new viruses from grapevine. Tanne E. M.

Lee.. Legin. Phytopathology 77. Extended Abstracts 14th Meeting of ICVG. 1990. Z. Walter B. Zimmermann D. B. 62-66. Walter and O.Y. R.E Goszczynski. C.. the closterovirus type member. Boscia. D. Zimmermann. Walter and M.P.. Boscia. Locorotondo 2003. 130. Agronomie 8.E. 1427-1434. R. B. A. 137-145. 1991. Zhu H. 731-741. Jiang and D. K. and G. C. Cytopathology of leafroll diseased grapevines and the purification and serology of associated closteroviruslike particles. Martin.A.H. Vernoy. 1998 Leafroll virus is common in cultivated American grapevines in Western New York. G. Zhou Z. The use of a greengrafting technique for the detection of virus-like diseases of the grapevine. Saldarelli. Gonsalves. Abou-Ghanem. Adelaide 2000. Potere. N. Nucleotide sequence and genome organization of Grapevine leafroll-associated virus 2 are similar to Beet yellows virus. Le Gall.. Purification de particules virales associées à l'enroulement de la vigne et mise au point d'un protocole ELISA permettant leur détection. Volos 1990.S. 2003. Gonsalves. A. O. Potere. P.W. D. Journal of Phytopathology 130. Bass. Journal of General Virology 79. Sommermeyer. O. Castellano. Kim. 1958. Journal of Phytopathology 128. Production and characterization of monoclonal antibodies specific for Grapevine leafroll-associated virus 3 and epitope mapping of the coat protein. R. Turturo. 1062. D. Martelli.F. Proceedings 10th Meeting of ICVG. Walter B..S. 1289-1298. Van Regenmortel. D.. Monoclonal antibodies for detection and characterization of Grapevine leafroll-associated virus 2. Zee F. Further characterization of closterovirus-like particles associated with the grapevine leafroll disease. Zimmermann D. Gonsalves.Y. D. Extended Abstracts 13th Meeting of ICVG. 1987. Transmission par greffage d'une virose type enroulement foliarie commune dans le vignobles de l'Est e du Centre-Est de la France. Zhou Z. 1990. 203. 1998. Ling. Pool and R. and D. Goszczynski and M. Comptes Rendues de l'Academie d'Agriculture de France 44. J.Vuittenez A. Vesselle. Collas and G.V.R. 2000. McFerson and D... 313-316. 75 . K. 277-288. Production and characterization of monoclonal antibodies specific to closterovirus-like particles associated with grapevine leafroll disease. 1988. Goheen. Wilcox F. Plant Disease 82. P.

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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

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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

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are mechanically transmissible.: Graft transmission of rough bark to LN 33. Vitiviruses. 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. Graniti and Martelli: Demonstration of the infectious nature of rugose wood. GVA can be eliminated to a very hight rate (up to 97%) by the procedure used for cryopreservation of grapevine shoot tips. Suggestion that it may be caused by a virus. Graniti: Detailed description of rugose wood symptoms. described from California. 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. experimental evidence has been obtained of the very close association of GRSPaV. Kober 5BB and LN 33) is the only reliable method for detecting and sorting out the diseases of the complex. it is plausible to regard 110R as a specific indicator of rupestris stem pitting in addition to V. Recently. since symptomless infections make sanitary selection not totally reliable. Faccioli: First histological study of corky bark-affected grapevines. Name of the disease changed into corky bark. Histological study of diseased vines. Control of mealybugs is difficult for they overwinter under the bark of grapevines and possess an unwettable waxy covering. as shown by 110R. "stem pitting" and "stem grooving". but not foveaviruses. all sources must be indexed and/or laboratory tested. or a combination of the two. Suggestion that rugose wood may be a disease of combination requiring the contact of scion and rootstock for the development of symptoms. Transgene expression was detected in these plants and in transformed grapevine explants. Thus. meristem tip culture. rupestris. possibly in relation with the scion/stock combination and climatic conditions. though with difficulty. rupestris. 2. 1965 1965 1967 81 . 1954 1961 1962 1963 1963 1964 1965 Hewitt: Rough bark. Martelli et al. Beukman and Goheen: Brief account of the histological modifications of corky bark-affected LN 33. to a restricted range of herbaceous hosts (mostly Nicotiana species). The best antigen sources for serological diagnosis are cortical shavings from mature dormant canes. Historical review Names like "legno riccio". In general. Goidanich and Canova: First record of corky bark in Europe. rugose wood agents can be eliminated with reasonable efficiency by heat therapy. immunocapture RT-PCR. a virus-like disease. several resistant plant lines were obtained by transformation with the coat protein and the movement protein genes of GVA and GVB.: Corky bark is remarkably heat stable and difficult to eliminate by heat therapy. if not otherwise associated with a specific syndrome. Individual viruses can be identified by ELISA or dot immunobinding on nylon membranes using polyclonal antisera and/or monoclonal antibodies when available. are synonymized with "rugose wood". Goheen et al. with vein necrosis. Hewitt et al. However. other assays include: single step or nested reverse trascription-polymerase chain reaction (RT-PCR). Detection: Indexing on indicators (V. no strategy has yet been developed for the chemical control of vectors. Latent infection can occur also in grafted vines.were observed in self-rooted cultivars and ungrafted roostock stocks (V. Thus. The intensity of wood abnormalities (pitting and grooving) vary. and that it may be a composite disease resulting from the interaction of different viruses among which GFLV.: First record of rugose wood outside of Italy. In addition. the putative agent of rupestris stem pitting. or spot-RT-PCR using degenerate or virus-specific primers. Using a Nicotiana benthamiana model system. rupestris and Kober 5BB). Immuno-capture RT-PCR is 1000-fold more sensitive than ELISA for virus detection in grapevines. Graniti and Ciccarone: First record of rugose wood from southern Italy.

Mink and Parsons: Use of growth chambers for rapid symptom expression of corky bark in Vitis indicators. Goheen and Luhn: Suggestion that corky bark and rugose wood are the same disease. Goheen: Evidence that corky bark and leafroll. Conti et al. Sarooshi et al. First record of rugose wood symptoms outside of Europe (Israel).: Recovery by mechanical inoculation of a closterovirus with particles 800 nm long. from a rugose wood-infected vine. Rosciglione et al.: Heat therapy effective against rugose wood. 1968 1969 1970 1970 1971 1971 1972 1973 1973 1975 1975 1977 1978 1979 1979 1980 1980 1981 1981 1982 1983 82 . Hewitt: Up-to-date review on grapevine virus and virus-like disease worldwide. Bovey and Brugger: Further evidence that GFLV may not be implicated in the etiology of rugose wood in Switzerland.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.Virus provisionally called grapevine stem pitting-associated virus (GSP-AV). At 38 °C the minimum inactivation period for leafroll is 56 days and for corky bark 98 days. Beukman and Gifford: Detailed account of adverse effects of corky bark on the anatomy of Vitis.1968 1968 Lehoczky et al. Teliz et al. despite similarities in the s y m p t o m s o n t h e foliage are different diseases. Goheen and Luhn: Heat treatment of dormant buds grafted onto LN 33 is effective against corky bark. Engelbrecht and Nel: Rugose wood and fanleaf are not related.b. based on graft transmission tests. Castillo et al. Rugose wood may not require a grafted plant for full symptom expression. is transmitted by the pseudoccid mealybug Pseudococcus longispinus. Anonymous: A review of rugose wood in Italy. No nepoviruses implicated in their etiology. Hewitt: Successful graft transmission of Californian rugose wood. Legin et al. Boccardo and D'Aquilio: Physicochemical characterization of GSP-AV Abracheva: Survey of over 650 grapevine cultivars and hybrids for rugose wood reaction in Bulgaria. Beukman and Goheen: Up-to-date review of corky bark.: Green grafting useful for corky bark indexing. Hewitt and Neja: First record of rugose wood in USA (California).: Observation of rugose wood symptoms in self-rooted vines. a. Graniti and Martelli: Up-to-date review of rugose wood.: Rugose wood recorded from Australia.: First experimental evidence that a filamentous virus (GVA). Lehoczky: Destructive effects of rugose wood registered in Hungary in both self-rooted and grafted European grape varieties.

Sc.: Experimental confirmation that rugose wood may not express symptoms in grafted indicators.: Two distinct dsRNAs with a mol. Engelbrecht et al.: Evaluation of the effect of rugose wood on cv. Prudencio: M. The first two disorders appear to be spreading in the vineyards. Rugose wood and corky bark are not the same disease. Tanne et al.: First indication of the possible existence of LN 33 stem grooving. Martelli: Rugose wood recorded in southern Mediterranean and Arab countries. GSP-AV re-named Grapevine virus A (GVA). Garau et al. Corbett and Wiid: Closterovirus-like particles found in extracts from vines affected by corky bark and rugose wood in South Africa. Monette et al. but not consistently in grapevines from New York. Li et al.: Three types of wood disorders of the stem-grooving type observed in South African grapevines. No closterovirus-like particles in samples with rupestris stem pitting. wt of 5.: Application of spot hybridization for the detection of GVA in grapevine sap. First report of Kober stem grooving. Italia propagated on six different rootstocks. Murant et al. Azzam et al.: First record of rugose wood from China. ficus. Savino et al. thesis describing rupestris stem pitting disease in comparison with corky bark. Similar dsRNA species were detected. Corky bark and Rupestris stem pitting.: Evidence that GSP-AV can occur in grapevines together with another similar but serologically unrelated virus with short closterovirus-like particles. Rosciglione and Castellano: Demonstration that GVA is transmitted also by Planococcus citri and P. Engelbrecht and Kasdorf: Natural field spread of corky bark in South Africa associated with the presence of P. Gallitelli et al. an additional disease of the rugose wood complex.ficus.: Heracleum latent virus and GVA are distantly serologically related. Monette and James: Detection of two biologically distinct but serologically indistinguishable isolates of GVA. similar to Kober stem grooving.1984 Milne et al. Suggestion that the disease is not related to closteroviruses associated with GLRaV and corky bark. Savino et al. denoted Grapevine virus B (GVB). Castrovilli and Gallitelli: Physicochemical comparison of two Italian isolates of GVA. ficus.: A low molecular weight dsRNA associated with rupestris stem pitting. Garau et al.: Transmission of corky bark by the mealybug P.: Assessment of crop losses induced by rugose wood to two different European grape varieties. 1985 1985 1985 1985 1985a 1985b 1985 1985 1985 1987 1988 1989 1989 1989 1989 1989 1989 1990 1990 1991 1991 83 . Kuniyuki and Costa: Rugose wood recorded from Brasil Goheen: First published description of rupestris stem pitting.: Experimental confirmation of the complex nature of rugose wood based on the differential reaction of woody indicators.3 and 4.4 x 106 associated with rupestris stem pitting in grapevines from California and Canada. Savino et al.

Suggestion that GVA is implicated in the aetiology of the disease.: Contemporary occurrence of Rupestris stem pitting and Kober stem grooving in symptomless scions of cv.:Rugose wood recorded from Tunisia Milkus et al.: Sequence of the 3' end of GVA and GVB genome. Saldarelli et al. Virus transmission by the mealybug Ps. Digiaro et al.: Synthesis of a cloned probe for GVA.: Rugose wood recorded from Ukraine Boscia et al. Namba et al. Chavez and Varon de Agudelo: Rugose wood recorded from Colombia. 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. ficus induced corky bark symptoms in LN 33 Saldarelli et al. Martelli et al. Padilla: Rugose wood recorded from Spain Boscia et al.: Presence of two distinct serotypes of GVA. 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. Both viruses qualify for the inclusion in the genus Trichovirus. Torbato in Italy. Garau et al. Virus named Grapevine virus C (GVC).: GVA and Kober stem grooving are closely associated.: Production of monoclonal antibodies to GVA and their use for ELISA detection of the virus in infected vines.: Rugose wood recorded from Malta Monette and Godkin: Recovery of a closterovirus-like virus by mechanical inoculation from a corky bark-affected vine.: A closterovirus with particles 1440-2000 nm long serologically unrelated to all other known grapevine closteroviruses found in corky bark-affected vines.1991 1991 Gugerli et al.: Purification and properties of GVB. Merkuri et al. Minafra et al.: Development and diagnostic use of a cloned probe to GVB.: Development of digoxigenin-labelled riboprobes for the detection of GVA and GVB in infected tissue extracts. Minafra and Hadidi: Detection of GVA and GVB in viruliferous mealybugs by PCR. Martelli et al. Suggestion that GVA may be the causal agent of the disease. Boscia et al. Saric and Korosec-Koruza: Rugose wood recorded from Croatia and Slovenia Ioannou: Rugose wood recorded from Cyprus.: Rugose wood recorded from Yemen. Minafra et al. Garau et al. Boulila et al.: Rugose wood recorded from Albania.: Clear-cut connection of GVA and rugose wood. both associated with a stem pitting condition of grapevines rather than with leafroll. Thorough comparative study of nine GVB isolates from different countries. 1991 1991 1991 1991 1991 1991 1991 1991 1992 1992 1993 1993 1993 1993 1994 1994 1994 1994 1994 1994 1994 1994 1995 84 .

: Sequencing of a Californian isolate of GRSPaV. Boscia et al. La Notte et al. The virus is not seed-borne.: GVB is consistently associated with corky bark and is present. GVB. Rubinson et al.: Consistent detection of GVA in Kober stem grooving-infected grapevines by immunocapture-polymerase chain reaction.: GVA is transmitted by by Ps.: Estasblishment of the genus Vitivirus with GVA as type species. Minafra et al. Suggestion that spreading is by a vector that transmits in a semipersistent manner.: Elimination of GVA from cv Nebbiolo clones by heat therapy improves agronomic performance of the vines and quality of the must.: Nucleotide sequence of GVB genome. GRSPaV is assigned to this genus. Efficient detection method based on TAS-ELISA developed. Zhang et al. Chevalier et al. Tanne et al. Saldarelli et al.: Rugose wood recorded from Lebanon. Goszczynski et al. Bonavia et al.: Nucleotide sequence of GVA genome and taxonomic position of the virus.: A study of the spatial distribution pattern of corky bark in aThompson seedless vineyard in Israel. Meng et al. longispinus in a semi-persistent manner. Boscia et al. Faoro: Review of the cytopathology of grapevine trichovirus infections. and GVD) later assigned to the genus Vitivirus. GVB.: Rugose wood recorded from Jordan. 1995 1995 1995 1996 1996 1996 1996 1996 1997 1997 1997 1997 1997a 1997b 1997 1997 1997 1997 1998 1998 1998 1998 85 . this may be the first visualization of GRSPaV. GVA.: GVA and GVB are transmitted by Pseudococcus affinis.: Review of the properties of putative grapevine-infecting trichoviruses (GVA.: GVA and GVD are serologically distantly related. Alkowni et al: Rugose wood in Palestine.1995 Monette and Godkin: Detection of non mechanically transmissible capillovirus-like particles in a grapevine affected by rugose wood. GVC. Martelli and Jelkmann: Establishment of the genus Foveavirus.: Sequence and structrural organization of Grapevine rupestris stem pittingassociated virus genome (GRSPaV). and GVD removed from the genus Trichovirus and assigned to the new genus.: Antiserum to the movement protein of GVA is useful for virus detection in ELISA. GVC.: GVA and GVB are serologically related. Garau et al. Abou Ghanem et al. La Notte et al. though not consistently in vines showing a syndrome denoted "corky rugose wood".: Development of a PCR technique for the detection of GVA and GVB in nylon membrane-spotted sap. Since particle size (600-700 nm in length) is compatible with that of Grapevine rupestris stem pitting-associated virus (GRSPaV) particles identified in 2002. Further support of the cause-effect relationship between GVA and this disease. Haidar et al. Guidoni et al. Choueiri et al.: Description of Grapevine virus D (GVD). Martelli et al.

intermingled with virus particle aggregates. Minafra et al.: 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. and III) identified in South Africa based primarily on sequence homology of the 3' end of the viral genome. Meng et al. Saldarelli et al. Nucleotide sequence identity within groups is 91-99.: GVA particles carry a highly structured epitope centered on a common peptide region of the coat protein sequence. Goszczynski and Jooste: Use of single-strand conformation polymorphism reveals molecular heterogenity in GVA populations. GVA coat protein encapsidating GVB RNA. Ahmed et al. Habili et al. Saldarelli et al. 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. GVB.: Consistent association of GRSPaV with vines indexing positive for Rupestris stem pitting. Seyval seedlings. Kominek et al. II.3% among groups.: Rugose wood viruses in Iran.: 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.: Production of monoclonal antibodies to GVB. GVA movement protein is also present in great quantity in the cytoplasm. Boscia et al. Nakano et al.8% and 78-89. The virus was not detected in 245 seedlings from infected cv.: Elimination of GVA by cryopreservation.: Synthesis of full-length cDNA copies of GVA and GVB genomes. observed for the first time.: Movement proteins of GVA and GVB detected by gold immunolabelling in association with cell walls and plasmodemata of infected cells. 2000b 2000 2001 2001 2001 2002 2002 2003 2003 2003 2003 2003 2003 2003 2003 2003 2003 2003 86 . are filamentous and measure 723 nm in length. Galiakparov et al. i. Buzkan et al. Confirmation of the cause-effect relationship between GVA and Kober stem grooving. GCD) and foveavirus (GRSPaV) sequences in two steps. Virus detected in bleached seeds suggesting that it is present inside the seeds.: Rugose wood viruses in Egypt.e.: Production of an infectious RNA transcript from a full-length cDNA clone of GVA.: GRSPaV particles. Stewart and Nassuth: An improved extraction method allows RT-PCR detection of GRSPaV virtually thoughout the year in all grapevine tissues. Further support to the cause-effect relationship of GRSPaV with this disease Galikparov et al. GVB and Corky bark and GRSPaV and Rupestris stem pitting. occurs in Nicotiana plants doubly infected with GVA and GVB. Dell'Orco et al.: The function of GVA genes identified by mutation analysis of individual ORFs of a full-length infectious viral clone. Petrovic et al. Wang et al.: One-sided phenotyping mixing.: GVA transmission by Pseudococcus comstocki. Samples made up of three buds from dormant canes are less laboriour to prepare than cane shavings and yield comparable results.1999 1999 2000a Meng et al.: Rugose wood viruses in the Czeck Republic. Goszczynski and Jooste: Thre groups of GVA strains (I. Dovas and Katis: Improved RT-PCR method for the simultanous detection in grapevine extracts of vitivirus (GVA.

179-182. a tumor-inducing virus of grapevines. Milne and C. 185-194. A. M. Monette (Ed. Martelli. Bottalico. Davis.M. P. Informatore Fitopatologico 29(2). Foster. D. 1970. comprising grapevine viruses belonging in the genera Vitivirus. Masannat.A. Boscia D. In: Filamentous Viruses of Woody Plants. 2001. 3. Buzkan. Minafra. Abou Ghanem. Martelli.P. Berkeley. 1965. M. Minafra.A.P. Golino. A. V. Bulletin OEPP/EPPO Bulletin 28. R. 1996.J. Studies on "corky rugose wood" of grapevine and the diagnosis of grapevine virus B. Castellano. 1992. Extended Abstracts 14th Meeting of ICVG. 15-25. Minafra.F. 1981. V. Archives of Virology 127. Gonsalves and D. 1981. 1965. Grape corky bark. Journal of General Virology 53. Boetalico and O. 1045-1060. V. and A.. Boscia D. A comparative study of grapevine virus B isolates. Bouyahia et al..G.W. M. The protein and nucleic acid of a closterovirus isolated from a grapevine with stem-pitting symptoms. Rivista di Patologia Vegetale Ser. M. 109-120 Boscia D. Alkowni R. Locorotondo 2003.). Adams M. N.. A. Martelli. Abou-Zurayk and G. Candresse. Savino and G. Current knowledge on viruses and virus diseases of grapevines in Tunisia. Trivandrum. Savino. 1969.A. Brunt. Elicio. K. A preliminary survey for grapevine viruses in Egypt.P. Beukman E. Savino. V. Boscia.. Trichovirus and Foveavirus. No veing necrosis observed in 110R top grafted on GRSPaV-free V. A. Abracheva P. N. Il legno riccio delle vite in Italia.. Namba. Plant Disease 75. N. 1995. Abou Ghanem. 960-964. Garau. M. D. 203-205. Journal of Plant Pathology 79. M. Archives of Virology 149.: An association exceeding 95% observed between GRSPaV and 110R vines showing vein necrosis symptoms in indexing trials. Martelli.. Beukman E. N. 104-110.D. P. Ahmed MH. Savino and G. A. and E. The new plant virus family Flexiviridae and assessment of molecular criteria for species demarcation. D'Aquilio.R. 1994. References Abou Ghanem N. Adams et al. Bar-Joseph. Phytopathologia Mediterrenea 34. Division of Agric. a novel putative trichovirus.. Chabbouh.P. Minafra. Production of monolconal antibodies to grapevine virus D and contribution to the study of its aetiological role in grapevine diseases.C. Vitis 35. 1991. Martelli. 73103. Castellano and G. Martelli. M. A. R. R. Corky bark. and A. Volos 1990. M. Castellano and G. Univ. 1993.I. A.C.F.: Establishment of the family Flexiviridae. Gonsalves and G. Boscia D. Martelli. Castellano. Savino. G. E.. Meng and Gonsalves: Comprehensive review of the characteristics of GRSaV.. Potere.): Virus Diseases of Small Fruits and Grapevines (A Handbook). Elicio. T.P. Boscia D. Bonavia M. 207-209. characterization and use of monoclonal antibodies to grapevine virus A. Saldarelli. Martelli. In Frazier N. G.F. and M. Martelli . Hilgardia 40. Z. Anatomic effects of corky bark virus in Vitis. Beukman E. Properties of Grapevine virus D. V. 178-179.V.A. C.A. La sensibilité de certaines variétés de vigne à la maladie du bois strié (legno riccio). 3-18. Digiaro and G.A. Martelli. Detection of dsRNA in grapevines showing symptoms of rupestris stem pitting disease and the variabilities encountered. 1991. 1998.P. Digiaro.M. Safi. 19-28. Cherif and G. Fauquet. Anonymous 1979. Research Signpost. Azzam O.P. Suggestion than vein necrosis is a specificic reaction of 110R to GRSPaV. Boari. Garau. of California. Production.F Antoniw. V..L. (Ed. 2004. 53-48.P. J. 164-166. Boulila M. Aslouj. California. D. 126-128.. Vitis 40. Minafra and G. rupestris. Zorloni et al. Boccardo G.M. A. Sci. Rugose wood of the grapevine in Jordan.: Experimental transmission of GVA by the mealybug Heliococcus bohemicus. Archives of Virology 130. Gifford. S. 1997. Proceedings International Conference on Virus and Vector on Perennial Hosts with Special Reference to Vitis.. Goheen. 4. 87 .P. Viruses and virus diseases of grapevine in Palestine. 11-24.. 189-195. G. M. Goheen. Digiaro and V. Filamentous viruses of the grapevine: putative trichoviruses and capilloviruses. 69-74.. Phytopathologia Mediterranea 20. G. Saldarelli. Properties of a filamentous virus isolated from grapevines affected by corky bark. P. 2003. Digiaro. Zhou. Proceedings 10th Meeting of ICVG.2003 2003 2004 2004 2004 Minafra and Boscia: Review of rugose wood-associated viruses.. Boscia D. 1997. M.P.

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GRAFT INCOMPATIBILITY .

.

Normal growth resumes with the second or third leaf. 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.g. in Argentinian grapes. 101-14) exhibit a green canopy and perform rather well. leaves are small-sized. GVB is mealybug-borne and can be spread at a site by these insects. although none of a number of known grapevine-infecting viruses has been found in affected vines. Agents: An ordinary strain of Grapevine leafroll-associated virus 2 (GLRaV-2) is consistently associated with Kober 5BB incompatibility (Europe). Argentina and probably elsewhere. Detection: Indexing on Caberent sauvignon is a reliable method for detecting incompatibility conditions. decline and may die. Severely affected vines decline and may die within one or two years. Australia and Chile (young vine decline).). The putative agent of bushy stunt was consistenly found in clones of the rootstock 140R in which it is latent. European grape varieties grafted on tolerant rootstocks (e. incompatibilità d'innesto (Ital. associated with grapevine bushy stunt is still unidentified. so as to represent veritable emerging diseases. 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). but the yield is reduced. A prominent swelling forms at the scion/rooststock junction and variously extended necrotic lesions may develop on the rootstock stem. This latter condition has been known for a long time and occurs also in rugose wood-affected vines. but the colour of the canopy remains green. whereas varieties grafted on susceptible roostocks (e. Infected propagative material is to be blamed for its dissemination. though not consistently and. which are usually not accompanied by wood abnormalities (pitting or grooving). 3309) develop a discolored canopy . 1.g. Cases of graft union disorders have been observed in Europe (Kober 5BB incompatibility). Syrah decline is a severe disease occurring in France. New Zealand. scions show a stunted and bushy vegetation due to the contemporary proliferation of apical and axillary buds. California. except for GRSPaV. with margins more or less extensively rolled downwards. Chile. The nature of this disease has not been ascertained but one or more graft-transmissibile agents may be involved in its aetiology. the increased use of clonal material is disclosing unprecedented conditions of generalized decline that develop dramatically in certain scionrootstock combinations. 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. 1103P. 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. Based on the differential responses of a panel of 18 rootstocks. Varietal susceptibility: Appearance of graft union disorders depends more on the rootstock rather than the scion. up to five different grafttransmissible agents inducing incompatibility could be differentiated in California. The same virus was detected in diseased Chilean grapes. Other molecular variants of GLRaV-2 were reported from New Zealand (Alphie virus). consistently. 5C. A transitory form of incompatibility was reported from Italy under the name of bushy stunt. and again California (rootstock stem lesions). The heat-labile graft-transmissible agent present in the hybrid 140R. 03916.) Symptoms: Newly planted vines grow weakly. and together with Grapevine virus B (GVB). Kober 5BB. Harmony. 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). Freedom. 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. Transmission: GLRaV-2. a member of the genus Closterovirus. Other assays include nucleic acid-based tecniques such as single step or nested reverse transcription-polymerase chain reaction (RT-PCR) and 95 . Of these. Description Main synonyms: Incompatibilité au greffage (Fr. However. and Australia in association with young vine decline conditions. shoots are short. Salt creek . A virus originally detected in cv. is not transmitted by mealybugs and does not have a known vector. appears to be involved in California's young vine decline. only GLRaV-2 RG was identified.

immunocapture RT-PCR.: GLRaV-2 is associated. Suggestion that they are molecular variants of the same virus species. Durquety et al. Savino et al. Martelli: GRSLV and GLRaV-2 are serologically related and are both recognized by a panel of 18 monoclonal antibodies. GRSLV has about 75% nucleotide homology with GLRaV-2. 2003 2003 2003 2003 2003 2003 2004 96 . utilization of tolerant roostocks is advisable. though not consistently. the use of sensitive rootstocks is to be avoided and. whenever feasible.: Third paper of a series on incompatibility on Kober 5BB. Currently known graft incompatibility agents can be eliminated with reasonable efficiency by heat therapy.: Description of bushy stunt and evidence that it is caused by a graft-transmissibile heat-sensitive agent carried by some clonal rootstocks. Bertazzon and Angelini: Comparison of several detection methods for the broad or specific identification of Grapevine leafroll-associated virus 2 variants.: Identification of an apparently new closterovirus denoted Grapevine rootstock stem lesion virus (GRSLV) causing stem necrosis of rootstocks. Gomez Talquenca et al: GLRaV-2 is consistently associated with declining Cabernet sauvingon vines grafted on different roostocks in Argentina.: Two papers describing incompatibility phenomena between clonal selections of different cultivars grafted on Kober 5BB Fallot et al. Graft-transmission of the incompatibility factor. decline.: GLRaV-2 is the cause of a graft incompatibility revealed by Kober 5BB. No conclusion are drawn. Bonfiglioli et al.: GLRaV-2 and GVB are consistently associated with young vine decline in California. with a decline condition of young Thomposn seedless vines in Chile. The problem is very complex and may involve several still unidentified factors.: Report of a new molecular variant of GLRaV-2 from New Zealand. Golino et al. Prodan et al. 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. or a combination of the two. If scionwood is infected. Historical review 1942 1950 1973. using degenerate or virus-specific primers. Greif et al. 2. Boubals: Syrah decline occurs in Argentina Uyemoto et al. Control: Prevent introduction of infected vines in the vineyard by using certified grafted plants or virusfree scionwood and rootstocks. GRSLV re-named Redglobe strain of GLRaV-2. 1977 1979 1986 1991 1995 2000 2000 2000 2001 Jacob: Description of graft incompatibility in different scion/stock combinations. meristem tip culture. Renault Spilmont et al. Strategies on how to protect healthy stocks from vector-mediated GVB reinfection in the field are yet to be developed. and death of the vines. Boubals and Huglin: Report on graft incompatibility of certain varieties grafted on 57R.: Updated report on the state of the art of investigations carried out in France on Syrah decline. Uyemoto and Rowhani: Indexing on 18 different grape rootstocks reveals the existence of at least five different agents causing graft incompatibility. Boubals: Report of a national French study group investigating the aetiology of Syrah decline.

Fallot. Di Silvio.II. 1986... Molecular studies on a graft incompatibility syndrome in New Zealand vineyards yields another probable variant of Grapevine leafroll-associated virus 2. Presence in clonal rootstocks of a grafttransmissible factor that induces stunting and bushy growth in European grapevines. Locorotondo 2003. 202-210 Uyemoto J. B. Discovery of different grapevine sources with graft-transmissible agents causing union-incompatibility on sensitive rootstocks.M. Uyemoto J. 201-203.3. J. Gomez Talquenca G. Prodan S.M.A. Nouvelles recherches sur l'incompatibilité clonale d'Abouriou greffé sur 5BB. 85-86.. M. Fiori. The relationship of grapevine leafroll-associated virus 2 with a graft incompatibility condition of grapevines.. S.M. 122-129 and 171-178.III.P. 283-290. Le clone et ses reactions au greffage. Phytopathologia Mediterranea 34. Bonfiglioli R. 1950. 85-86. New closterovirs in 'Redglobe' grape causes decline of grafted plants. and A. Syrah decline in French vineyards. Bass.. J...E. Rowhani. and B. La trasmission de l'incompatibilité au greffage entre 5BB et Vitis vinifera.. Extended Abstracts 14th Meeting of ICVG. Extended Abstracts 14th Meeting of ICVG. Locorotondo 2003. Gracia. Boscia.M. Luvisi and R. Prota. Prota. Le clone et ses reactions au greffage. O. and P. Rowhani. Grapevine virology highlights 2000-2003. 2003. 97 . Extended Abstracts 14th Meeting of ICVG. Grenan and J. 279-283. Le clone et ses reactions au greffage. 1977. Walter. J. Journal of Plant Pathology 86. Extended Abstracts 14th Meeting of ICVG. A young grafted vine decline syndrome in Argentina vineyards. Boursiquot. 2003. I. Extended Abstracts 14th Meeting of ICVG. Identification of the latent viruses associated with young vine decline in California. Locorotondo 2003. Di Terlizzi.. Proceedings 10th Meeting of ICVG. Angelini. S. 2000. Renault Spilmont A. P. 277. Fallot. 2003. 141. 1995. 167-173. Krag. Martelli G.K. Grau. Le dépérissement de la Syrah existe aussi en Argentine. Etude de l'incompatibilité au graffege de certain cépages et du 57R. 2004. and E. F. Garcia Lampasona and O. D. C. Progrés Agricole et Viticole 117. 144. Durquety and J. 1942.K. Durquety P.A. 142-143.S. Compte-rendu de la réunion du Groupe de Travail National. V. Progrés Agricole et Viticole 96. Progrés Agricole et Viticole 103. Aetiology of decline in Thompson seedless grafted table grape plants. Progrés Agricole et Viticole 117.P. Locorotondo 2003. Legin R. B. Extended Abstracts 13th Meeting of ICVG. Progrés Agricole et Viticole 67. Existence dans un cépage population de clones présentant divers degrés de compatibilité avec certains porte-greffes. D. J. 2001. Extended Abstracts 14th Meeting of ICVG. 183-189. 2003. 28-31. Proceeding of the American Society of Horticultural Science 41.S. Boubals D. P. Jacob H. 3-10. Boubals D. Progrés Agricole et Viticole 90.P. 2003. 420-427 Fallot J. Examples of incompatibility between grape varieties and roostocks. References Bertazzon N. Walter and U. 1979. Volos 1990. Locorotondo 2003. A. Ruchaud. Sim and A. Benassac and R. Le dépérissement de la Syrah. Dauty. S. 2000.P. Fiore. Ruchaud. 2000. Locorotondo 2003. Durquety P.. Pantaleo. Advances in the detection of Grapevine leafroll-associated virus 2 variants. 2003. S. Progrés Agricole et Viticole 94. Edwards and A. Ruchaud. 139-140. 1991. Rivieccio and F. Golino D. 211-216. Gazeau. Garau. Etude de phénomènes d'incompatibilité au greffage chez la vigne. C. Huglin. California Agriculture 55 (4).. Greif C.. Gazeau and J. Autres cas chez la vinge. 1973. Rowhani... Adelaide 2000. Savino V. C. 137-141 Boubals D. R. Montalegre and N..

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FLECK COMPLEX .

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6 x 106 ) is 7564 nt in size and contains four open reading frames (ORF) that 101 . Affected vines are often stunted. and Grapevine rupestris vein feathering virus (GRVFV) are all phloem-limited and non mechanically transmissible. Fleck. vinifera in California. The disease is latent in European grapevine varieties and in most American rootstocks. GFkV particles sediment as two centrifugal components. Leaves are asymmetric. The putative causal agent of the disease has only been found in California. e. Grapevine asteroid mosaic: Mosaïque étoilée (Fr.). maculatura infettiva. In V. d. In V. Sternmosaik der Rebe (Germ. c. sometimes with necrotic center. Sultanina). T made up of empty protein shells and B.). Rupestris necrosis. is latent in European grapevine varieties. Severe strains induce also varying degrees of stunting. rupestris. screziatura (Ital. leaf petioles and veinlets. rupestris reacts with localized necrosis of the shoots. 1. The putative causal agent of the disease so far has been found in Greece. rupestris. Mild asteroid mosaic-like symptoms are shown by some European grapevine varieties (e. Asteroid mosaic. Agents: All viruses of the complex. Red globe) nor in V. Italy and California. Symptoms: a. containing the genome. Transient mild chlorotic discolourations of the primary and secondary leaf veins develop in V. capped. GFkV. which are twisted and asymmetric. and GRVFV genomes are available. The complete sequence of GFkV and partial sequences of GRGV. 25 kDa.g. B.FLECK COMPLEX The fleck complex consists of several diseases (grapevine fleck. Marmorierung der Rebe (Germ. the disease elicits creamy-yellow bands developing along the major veins of the leaves. Symptoms are expressed in Vitis rupestris and consist of clearing of the veins of third and fourth order. The coat protein (CP) of GFkV and GRGV particles is made up of a single protein species with Mr of c. rooting ability of rootstocks and on graft take has been reported. rupestris following graft inoculation. 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. Recorded from California and Italy . Grapevine red globe virus (GRGV) is a Grapevine fleck virus (GFkV)-like virus which apparently does not induce symptoms in European grapevine varieties (e. irregularly distributed over the leaf blade. vinifera. and produce little or no fruit. producing localized translucent spots. Leaves with intense flecking are wrinkled. twisted and may curl upward. grapevine asteroid mosaic. adverse influence on vigour. Although the elusive nature of the complex hinders the assessment of its economic impact.). Fleck is an ubiquitous disease reported from most viticultural countries in the world. GRGV. twisted and puckered along the veins. which is used as indicator. This disease. Grapevine fleck: Marbrure (Fr. reported only from Japan. Description Main synonyms: A.). GAMaV.). whereas the CP of GAMaV and GRVFV consists of a major protein of 21 kDa and a minor prtotein of 25 kDa. V.). Rupestris vein feathering. b. mosaico stellare (Ital. 50%). which is a monopartite single-stranded. leaf symptoms are characterized by star-shaped chlorotic spots.g. 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. grapevine rupestris necrosis. Leaf symptoms usually become less severe in summer. Asteroid mosaic symptoms have been observed in several varieties of V. GFkV genomic RNA (Mol. GFkV genomic RNA constitutes about 35% of the particle weight. 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. wt of 2. but likely to occur elsewhere. positive sense RNA with high cytosine content (c. Grapevine asteroid mosaic-associated virus (GAMaV).

2. marafiviruses and members of the genus Tymovirus to warrant the establisment of the a new family denoted Tymoviridae. GFkV is not seed transmitted.: First record of fleck as an unidentified symptom different from fanleaf and transmissible from symptomless varieties to V. but cannot be used for any of the other members of the complex due to the unvailability of antisera. 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. but no experimental data are available. and two proline rich polyproteins of 31. These deranged organelles are thought to be sites of virus replication. Because of its molecular characteristics. Therefore. whereas GAMaV induces peripheral vesiculation of chloroplasts. The 3' end of GRGV genome is structurally similar to that of GFkV except for the lack of ORF 4. ELISA is currently employed for routine detection of GFkV. Varietal susceptibility: GFkV and possibly all the other viruses of the complex infect naturally a large number of varieties and Vitis species.4 kDa (ORF 3) and 15. primary dissemination of these and the other viruses of the complex is through infected propagative material. meristem tip or fragmented shoot apex culture. The same sanitation procedures are likely to operate successfully with the other viruses of the complex. and GRVFV. whilst GAMaV and GRVFV were assigned to the genus Marafivirus. GAMaV. molecular and ultrastuctural studies disclosed sufficient similarities between maculaviruses. GFkV was identified as the representative of a new genus denoted Maculavirus. of which it represents the type species. Transmission through dodder of GFkV has been reported but it is of no epidemiological importance. Hewitt et al. rupestris St. Control: Because of the latency of symptoms sanitary selection of European grapevine cultivars and most American rootstock hybrids is unrealiable. 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.encode a 215. 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". 102 . Comparison of symptoms with those of other mosaic diseases of grape. Historical review 1954 1962 1966 1966 Hewitt: First description of asteroid mosaic in California. 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.4 polypeptide with the conserved motif of replication associated proteins (ORF 1). No information is available on individual susceptibility. Transmission: No vector is known for any of the viruses of the fleck complex. Although observations from Italy.George. GRGV. Attempts to transmit the disease by mechanical inoculation to herbaceous test plants or by Xiphinema index were unsuccessful. a disease inducing symptoms similar to those of fleck in V. Virus specific and degenerate primers have been designed for single or multiplex RT-PCR detection of GFkV.rupestris described in France. its economic importance is low. Further physico-chemical. As the disease is rare and does not appear to be spreading. GFkV can be eliminated by heat therapy.: "Marbrure". Refatti: Review paper on asteroid mosaic. the CP (ORF 2). Detection: Indexing on V.9 kDa (ORF 4) with unknown function. Vuittenez et al.

later called grapevine phloem-limited isometric virus (GPLIV).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. Goheen and Luhn: A novel heat therapy system based on virus inactivation in buds grafted onto healthy LN 33 rootstocks is effective against fleck. Hévin et al. 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. Report of multivesiculate inclusion bodies probably connected with GPLIV infection. Barlass et al. Dismissal of the prokaryote etiology hypothesis. Triolo and Resta: Tetracycline treatments are ineffective against fleck. Woodham and Krake: Dodder transmission of fleck from vine to vine. Castellano and Martelli: Confirmation that GPLIV is associated with multivesiculate bodies and demonstration that these derive from deranged mitochondria.: Purification of GPLIV from naturally diseased vines and production of a specific antiserum. Verderevskaya et al. Castellano et al. The efficiency of heat treatment for disease elimination is unsatisfactory.: Observation of a non mechanically transmissible virus.: Report of widespread occurrence of fleck in visually selected grapevine clones in southern Italy. Dolja et al.: Physicochemical characterization of GPLIV. 1982 1983 1983 1983 1984 1985 1985 1987 1989 1990 1990 1990 1991 103 . Castellano et al. in sieve tubes of field-grown vines with leafroll symptoms but likely to be affected by other diseases. leafroll and corky bark). Milkus: Suggestion of a prokaryotic etiology for fleck. Ottenwaelter et al.: Description of fleck as an independent graft-transmissible disease present in many European varieties and American rootstocks.: Successful elimination of fleck through heat therapy. rupestris. rupestris propagating wood. Bovey: Identification of fleck in Switzerland as a latent disease of Chasselas transmissible to V.: Fleck is not seed transmissible. Rooting ability and graft take are adversely affected Yamakawa: Field spread of fleck in Japan Boulila et al. Engelbrecht and Kasdorf: Observation of natural field spread of fleck in South Africa.: Successful elimination of fleck through fragmented shoot apex culture in vitro.: Observation of an isometric non mechanically transmissible virus in the phloem of diseased vines. Report that a virus serologically similar to GPLIV is associated with the disease. Hewitt et al. Confirmation that the virus can be eliminated by heat therapy and is not related to leafroll.: Identification of a dsRNA of about 7 Kb pairs in diseased vines. Rives: Further demonstration that fleck is distinct from fanleaf based on differential responses to heat treatment. Savino et al. Triolo and Materazzi: Fleck has a detrimental effect on the quality V.

Symptoms are severe and affected vines are almost fruitless.: GPLIV shown to be the agent of fleck. vein necrosis and vein mosaic has a detrimental effect on rootstock growth. Virus named Grapevine asteroid mosaicassociated virus (GAMaV) Boscia et al. GFkV-like multivesiculate bodies derived from deranged mitochondria are present in infected cells. Sabanadzovic et al.: 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. The disease seems to be spreading naturally. Meristem tip culture effectively eliminates the virus.: Molecular reagents (degenerate primers) developed for the specific identification of viruses of the fleck complex (GFkV. vinifera cv.1991 1991a 1991b 1991 Gugerli et al.: Natural field spread of GFkV observed in Northern Italy Schieber et al.: Report of the close association with fleck symptoms in V. rupestris. Boscia et al. 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. Pruning wood is reduced by 51% in 420A and by 37% in Kober 5BB.: A non mechanically transmissible isometric virus similar but unrelated to GFkV identified in asteroid mosaic-infected grapevines. ELISA used successfully for virus detection in large scale survey. rupestris with a necrotic disease. Boscia et al.: 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.: Additional monoclonal antibodies raised in France.rupestris of an isometric virus latent in V. Walter and Cornuet: Confirmation by ELISA of the consistent association of GFkV with fleck disease. Sultanina in Greece. Namba et al. Adverse effect on Teleki 5A is negligible. later named Grapevine rupestris vein feathering virus (GRVFV). 1991 1993 1994 1995 1995 1996 1996 1997 1997 1998 2000 2001 2001 104 . Virus renamed Grapevine fleck virus (GFkV). Fortusini et al. GAMaV.: Report of a highly consistent association of GPLIV with fleck in naturally infected and graft-inoculated V. vinifera.: Two GFkV-specific monoclonal antibodies raised in Italy can successfully be used in ELISA Kuniyuki and Costa: Three strains of GFkV reported from Brazil. Elbeaino et al. Detection of sequences of an undentified virus from a Greek grapevine. Marsumoto and Ohki: A spherical virus resembling GFkV identified in thin sectioned cells of V.: Complete nucleotide sequence of GFkV genome. Sabanadzovic et al. GAMaV and of another virus with GFkV-like particles phylogenetically but not serologically related to GFkV present in a cv Red globe vine. This cytological feature recalls that later found in vines infected by Grapevine rupestris vein feathering virus. June-July are the best months for ELISA detection of the virus in Alsace. Boscia et al. Molecular properties of this virus further support the notion that it warrants classification in a genus of its own. based on the differential reactions of indicators Credi and Babini: Infection by fleck. Virus named Grapevine redglobe virus. Kyriakopoulou: Description of a disease similar to asteroid mosaic observed in V. GRGV).

: Development of a multiplex RT-PCR protocol for the simultaneous detection of GFkV-like viruses using plant mRNA as an internal control. 1991a. Boulila M. Tomashevskaya. S. 2003a. V. Castellano M. Rowhani and G.P. 347-354.E.. comprising the genera Maculavirus and Marafivirus that include GFkV/GRGV and GAMaV/GRVFV. Phytophylactica 22. fleck. In other countries (USA. Martelli.V. Vitis 33. and A. Abou Ghanem-Sabanadzovic et al. G. References Abou Ghanem-Sabanadzovic. ElBeaino T. Vitis 22. 1972.P. S.A. N. Martelli and M.P. D. Sabanadzovic. Castellano M. Martelli et al.A. 101-102 Boscia D. Karsev. 1983. Savino.D.. 105 .F. rupestris confirms the assignment of GRGV to the genus Maculavirus and of GAMaV and the Greek virus to the genus Marafivirus. Progress in the study of the phloem-limited isometric virus-like particles associated with leafroll-diseased grapevines.P. 56-64. GAMaV and of a virus of Greek origin which induces vein feathering in V. Virus-like particles and ultrastructural modifications in the phloem of leafroll-affected grapevines. Sabanadzovic . South Africa.. Kyriakopoulou and G.A. 173-174. 1994. respectively. Sequencing of the 3' end of three grapevine fleck virus-like viruses . Locorotondo 2003. Elicio. Castellano. Boscia D.G. Krake. Sabanadzovic. Castellano M. Field spread of corky bark.. Plant Pathology 44.: Description of Maculavirus. Martelli.P. Virus Genes 27. Boyko. Credi R. Atabekov. G.P. Greek virus recognized as a species in its own right denoted Grapevine rupestris vein feathering virus (GRVFV). Abou Ghanem-Sabanadzovic et al. U.. 2003b. Un virus latent dans le Chasselas.R.. 3134.. Extended Abstracts 14th Meeting of ICVG. 97-105. 23-39. Numéro hors série. Double stranded RNA associated with fleck disease of grapevine. 151-158. and G. Association of a phloem-limited non mechanically transmissible isometric virus with grapevine fleck disease.M. G..A. Some properties of a phloem-limited non mechanically-transmissible grapevine virus. leafroll and Shiraz decline diseases and associated viruses in South African grapevines. Boscia D.P.E. Savino and D. Di Terlizzi. Savino and M. 165-169. Sabanadzovic and G. Digiaro. Vitis 30. Proceedings 10th Meeting of ICVG. 11-16 Abou Ghanem-Sabanadzovic. Martelli 2001. and Japan) only the variant without insertion (GFkV353) was detected. A. 1996. Castellano. R.. 1991b.: Description of the family Tymoviridae. 1982. B. 1990. K.. Martelli. A. Kasdorf. Martelli. Savino. a new genus of plant viruses having GFkV as type species and GRGV as tentative species. V. S.G. Martelli. Argentina. Babini. 1990.P.A. Barlass M. A non mechanically transmissible virus associated with asteroid mosaic of the grapevine. Rowhani P. V.R. S. Savino. Savino and G.V. Castellano. Boscia D. Martelli.. Regeneration of virus-free grapevines using in vitro apical culture. Shi et al. Volos 1990. Effect of virus and virus-like infections on the growth of grapevine rootstocks. O. Abou Ghanem-Sabanadzovic. N. and G. Multiplex RTPCR detection of Grapervine fleck virus-like viruses in grapevine with co-amplification of control plant mRNA. Woodham and L. M. Journal of Ultrastructure Research 89. T.P. Minafra. A. and G. A. Martelli. Kyriakopoulou and G.. N. GRVFV recorded from California and confirmation that GAMaV does not occur outside of California. Proceedings 8th Congress of the Mediterranean Phytopatological Union. Iran. M. Identification of the agent of grapevine fleck disease. Savino.C. Production of monoclonal antibodies to grapevine fleck virus.A. Ultrastructure and nature of vesiculated bodies associated with isometric virus-like particles in diseased grapevines. Molecular detection of Grapevine fleck virus-like viruses. Dolja V. 95-98. 2003b 2003 3. 195.. Annales de Phytopathologie. 191. Boscia. 160-163. G. Boscia. V. Journal of Phytopathology 129. 1985. 1990.J. Bovey R.: Sequencing of the 3' end of the genome of GRGV. Annals of Applied Biology 101.P. Verderevskaya and J. Martelli and V. Advances in Horticultural Science 10.2002a 2002b 2003a Martelli et al.G.: A sequence variant of GFkV (GFkV416) with a 63 nucleotide insertion in the replicase gene identified in Australia and New Zealand. V. V. Engelbrecht D. Martelli.P. Agadir 1990. 291-295.P. Phytopathologia Mediterranea 24. 1984. 1995. Vitis 40: 65-68. Skene. P. V.

D. Hewitt W. 1985. Matsumoto T... N. D. Heat therapy eliminates the ability to transmit the causal agent of "marbrure" in several V. Jr.P. Seddas.. and A. 1997. 106 .. Rosciglione. Doazan and M. 385-388.B. Fitopatologia Brasileira 20..J. Resta. Some virus and virus-like diseases of grapevines. Division of Agricultural Sciences.. Hewitt W. rupestris "du Lot" (St. European Journal of Plant Pathology 103. Shi B.A. Occorrencia de mais um isolado do virus do mosaico da nervuras da videira que no causa sintomas no porta-enxerto Kober 5BB. J. Milkus B. Grapevine asteroid mosaic. Yamashita. D. and E. 9. A. Acta Phytopathologica Academiae Scientiarum Hungaricae 9. Numéro hors série. Fortusini A. Journal of General Virology 82.M. N. 1991. 204-207. George). Parsons. and J.P. Saldarelli and G.T. Rivista di Patologia Vegetale. Hévin. Ohki. Archives of Virology 147. Studies on virus diseases of the grapevine in California. Ser.J. Berkeley. latent in many varieties. 57-83. Boscia. Annals of the Phytopathologial Society of Japan 64. J. Heat inactivation of viruses in grapevines. Prati. Investigating the transmission of marbrure and fan-leaf through the seed in the grapevine. Scattini. A. The family Tymoviridae. M. P. is transmitted by graft inoculation. Costa..P.. Numéro hors série. Mink G. Annals of Applied Biology 142. 560-564. 1991. Doazan and M. Abou Ghanem-Sabanadzovic.. IV. Rives. G. Goheen.B. Saldarelli. Gugerli P.): Virus Diseases of Small Fruits and Grapevines (A Handbook). 1998. M. Refatti E. 1972.. Ser. Vitis 3. S. Gugerli. Gooding. Ramel and F. 212-214. with Special Reference to Vitis. 1991. Informatore Fitopatologico 46 (12). 253-258. M. Annales de Phytopathologie. Bulletin of the California Department of Agriculture 43.Faoro F and P. Incidence of some graft-transmissible virus-like diseases of grapevine in visually selected and heat-treated stocks from Southern Italy. N. Habili and R. 1970. 43-47. T. Davis 1965. Rives M. 12491253. 2003.C. Dreher. Hévin M. Boscia and G.-J. 767-774.E. 281-285. Raski and G. Asteroid mosaic of grapevine. Tsuchizaki and D.. Digiaro and G. Ottenwaelter. Martelli G. affected by marbour. Plant Disease 75. Further characterization of grapevine leafroll disease. L. University of California. Phytopathologia Mediterranea 24. Plant Disease Reporter 61. 2001. Proceedings 10th Meeting of ICVG. Cory and C. In Frazier N. Symptoms of grapevine asteroid mosaic in Greece.IV. Schieber O. Castellano. Séparation de la marbrure et du court-noué (panachure) chez la vigne par thermothérapie. 2009-2015 Savino V. 1962. 1997. Belin and B.P.. (Ed. 39-43. Bonnard. 1973. Sabanadzovic S. Archives of Virology 145. Gonsalves. 9. Martelli G. 9. Purification and properties of spherical virus particles associated with grapevine Ajinashica disease.. 1974. S. B. vinifera clones and in V... Sabanadzovic. and C. 287-289. 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. Kyriakopoulou P. N. N. Abou Ghanem-Sabanadzovic and P. 567-571.P.V. Hewitt W. The responses of the Grapevine fleck agent to tetracycline-HCl antibiotic and Dienes' stain. Sabanadzovic S.. 1954. Extended Abstracts 12th Meeting of ICVG.L. Refatti E. 75-77. 2000. Volos 1990. M. Monoclonal antibodies for detection. 31-32. M. A possible new necrotic diseases of grapevine associated with small isometric particles and novel membrane-bound large particles. Rives.-E. S. 1977.M. 349-355. 47-64. Grapevine fleck virus-like viruses in Vitis. Abou Ghanem. Walter. Cinquanta and S.S. Kuniyuki H. A. Grapevine fleck disease. 553-565.C. Ottenwaelter M. 1837-1846. C. 1966. Nucleotide sequence variation in a small region of Grapevine fleck virus replicase provide evidence for two sequence variants of the virus. M. 143-146. Archives of Virology 147. Goheen. Leclair. 1847-1853. serological characterization and immunopurification of grapevine fleck virus. Luhn. 197-203.W.. 5960. J.C. P.B. Volos 1990.or chlamidia-like bodies in grape. 1995. Abou Ghanem-Sabanadzovic. 618-622.P. Procedures for rapid detection of virus and viruslike diseases of grapevine. S. Sabanadzovic.. 2002b. Triolo E. Martelli. and S. Goheen A. Ser. 1973. 1973.IV. 1996. 1972.. Cytological alterations associated with an unidentified isometric grapevine virus (UIGV). Martelli. Luhn. 157-164.I. Namba S. Lisbon 1997.. Symons.H. Maculavirus.C Edwards and T. 1985. Martelli. 2002a. Phytopathologia Mediterranea 24. S. Rivista di Patologia Vegetale. Rivista di Patologia Vegetale. Annales de Phytopathologie. Proceedings International Conference on Virus and Vector on Perennial Hosts. Complete nucleotide sequence and genome organization of grapevine fleck virus. Proceedings 10th Meeting of ICVG. a new genus of plant viruses. Mycoplasma. Tremea.P. Brugger.

V.G. Walter B. 1993. Legin and J. La maculatura infettiva della vite: influenza di isolati diversi sull'attitudine alla propagazione vegetativa di Vitis rupestris St.D. Yamakawa Y. Kuszala. 193-198. Cornuet. 1983. Ätiologie und Diagnose der Marmorierung der Weinrebe. Vuittenez A.Triolo E. Krake. Verderevskaja T. 1987.. and L. probablement indépendante du virus du Court-noué... R. Numéro hors série. and A. and P. Materazzi. Phytopathologische Zeitschrift 106. 1966. George. Woodham R. 221-226.R. Journal of the Japanese Society of Horticultural Science 58. Investigations on transmission of grapevine leafroll. 1989.. Marinesku and E. Virus reinfection of virus-free Cabernet sauvingon and Cabernet franc vines. 67-73. Semtschik. 1983. yellow speckle and fleck diseases by dodder. Archiv für Phytopathologie und Pflanzenschutz 19. 3209-324. La Recherche Agtonomique en Suisse 26. 107 .S. ELISA detection of grapevine fleck virus (GFkV). 651-657. Observations sur une mosaïque de la Vigne. Agronomie 13. Annales des Epiphyties 17. 297-302.C.

.

however. Agent: Alfalfa mosaic virus (AMV). 18% of the particle weight. 0. Main symptoms: Various patterns of yellow discolouration characterize the disease. 1.73 x 106 (2593 nt).MINOR VIRUS DISEASES Several graft-transmissible diseases are known. Control: Use of healthy material obtained by heat treatment. Some of these diseases have been recorded only from Europe. Yellow mottle has been reported from Germany. Bovey and Brugger: AMV recorded from Switzerland in grapevine and transmitted by grafting to V. Novak and Lanzova: AMV infections recorded from hop and grapevine in Czechoslovakia. Virus elimination by treating 37 days at 37-38°C. Hungary. Capsid proteins subunits are of one type. Transmission: AMV is efficiently transmitted by aphids in a non persistent manner and can cause epidemic outbreaks in many of its natural hosts. wts: RNA-1. Varietal susceptibility: Little information available. is the putative causal agent. former Czechoslovakia. 2. Jankulova: AMV infections recorded from Bulgaria. The spring growth shows more or less extensive yellowing of the leaf blades that does not extend to the veins.04 x 106 Da (3644 nt). Their overall importance is minor if compared with that of the major diseases dealt with in previous chapters. Description Main synonyms: None. Historical review 1973 1975 1976 1979 1978 1981 Bercks et al. Switzerland. RNA-3. 109 . Bovey and Cazelles: AMV particles visualized in thin sectioned grapevine leaves. Bulgaria. Faint yellow speckling. rupestris and the hybrid Grézot 1 x 5C. There may be differential susceptibility among cultivars. and a tripartite RNA genome accounting for c. suggesting that the virus spreads primarily through infected planting material. with which specific viruses are associated and thought to be their possible causal agents. has differently shaped particles. A. Chardonnay and Veltliner rouge précoce identified as reliable indicators. 0. infections are scattered and occasional. Detection: AMV is mechanically transmissible to herbaceous hosts and can also be identified by ELISA and moleculat techniques in infected vines. European diseases GRAPEVINE YELLOW MOTTLE 1. RNA-2.: First record of AMV infections and description of symptoms in German grapevines. with Mr 24x 103 Da. a mechanically transmissible virus. but some are of economic relevance locally (e. rings and lines are typical summer responses of infected vines. In grapevines. others occur in Japan. and Turkey.62 x 106 (2037 nt). AMV. Plant vigour and yield do not seem appreciably affected. Grapevine berry inner necrosis).g. 30 to 57 nm in size. with the following mol. from quasi isometric to bacilliform. Beczner and Lehoczky: AMV infections recorded from Hungary. the type species of the genus Alfamovirus.

FAO Publication Division. and a multipartite genome. 1976. 1993. or maple leaf-like line patterns typically confined to the petiolar area. Munich 1978. Grapevine disease in Hungary caused by alfalfa mosaic virus infection. References Akbas B. Erdiller. Cazelles. 119-128. New York . Agent: The putative agent. Francki. 55-63. Brugger. Grapevine line pattern virus (GLPV) a possible member of the genus Ilarvirus. Polyhedral Virions with Tripartite Genomes (R. Jubileum 75. 1975. Acta Phytopathologica Academiae Scientiarum Hungaricae 16. Transmission: GLPV has no known vector. Journal of Turkish Phytopathology 22. Bovey R. Biologia Plantarum 18. Historical rewiev 1985 Francki: Comprehensive review of the properties of AMV an other viruses with tripartite genome. 1-18. 63-65. Horticulture 7. is seed-transmitted and spreads with diseased propagative materials.) and grapevine (Vitis vinifera subsp. Rome. Identification of alfalfa mosaic virus and tomato bushy stunt virus in hop (Humulus lupulus L. Main symptoms: Leaves show bright yellow discolourations that form marginal rings. and J.B. or the upper part of the blade. Le virus de la mosaïque de la luzerne sur la vigne. 3rd International Congress of Plant Pathology. Jankulova M. Querfurth. Phytopathologische Zeitschrift 76. 1973. Lanzova.Hegi) plants in Czechoslovakia. vinifera cultivars are susceptible.P. Detection: GLPV is mechanically transmissible to herbaceous hosts. Akbas and Erdiller: AMV infections recorded from Turkey. Martelli: Yellow mottle suggested as the name for the disease caused by AMV in grapevines. Varietal susceptibility: No information. Arboriculture. Vigour and yield are reduced. Control: No information. 3. Martelli G.18 . Novak J. Alfalfa mosaic virus on grapevine. Researches on grapevine virus diseases and determination of their incidence in Ankara Turkiye. Bovey R. 263 pp. GRAPEVINE LINE PATTERN 1. This disease is known to occur only in Hungary.B. Investigation on certain viruses spread on grapevines in Bulgaria... Cordoba 1976. and J. . 1993. (ed. Beczner L. Über den Nachweis des Alfalfa mosaic virus in einer Weinrebe. 110 . Detection and Diagnosis of Graft Transmissible Diseases of Grapevines. Francki R. Monografias INIA No. Proceedings 6th Meeting of ICVG.B. Several V. Graft transmission to cv. 152-154. and O. sativa DC. and G. has differently shaped particles. and J. The viruses and their taxonomy. 131-134. scattered spots or blotches. D. 1985. 1981.1985 1993 1993 Francki: Comprehensive review of the properties of AMV and other viruses with tripartite genome. 166-171.I.I. roughly following its contour.). ed. In: The Plant Viruses.-J. Lehoczky. 39. Plenum Press. 1979. 1978. Description Main synonyms: None. Revue Suisse de Viticulture.). Lesemann and G. quasi spherical 25-30 nm in diameter to bacilliform 40 to 75 nm in length. 2. Bercks R.

Description Main synonyms: None. Viruses associated with the disease are readily transmitted by sap inoculation and can be detected by ELISA and molecular techniques. Martelli and J. Occurrence of the line pattern hitherto unknown virus disease of grapevine in Hungary. Detection: Graft-transmission to V. However. Farkas.: Evidence that GLPV is transmitted through grapevine seeds. Lehozcky J. 61-79 Lehoczky J. M. D.P. Kertgazdasag 19. 18% of the particle weight. Castellano. G. wt 1. Varietal susceptibility: No information. Lehoczky J. ed. The disease is graft-transmissible and spreads through infected propagating material.. Martelli. one of the putative agents of corky bark (rugose wood complex) has a very high association with diseased grapevines. Boscia. Plenum Press. 111 .A. Bunches are reduced in numbers. 115-116.I. 1985. Line pattern. Beczner and G. Proceedings 9th Meeting of ICVG. Kiryat Anavim. Control: No information. 3. Evidence that a graft-and mechanically transmissible virus is associated with it. J. 2.B. Main symptoms: Symptoms are prominent in late summer and consist of yellow and/or reddish discolourations of the tissues along the veins. Lehoczky et al. 1987. Agent: Symptomatic grapevines were reported to be doubly infected by GFLV and Carnation mottle virus (CarMV) the type species of the genus Carmovirus.. a novel virus disease of grapevine in Hungary. 23-30.. The disease has been recorded only from Greece. Beczner and G. 1987. In: The Plant Viruses.A.: Description of line pattern disease in Hungary. By converse. 1992. family Tombusviridae. Grapevine virus B (GVB). with Mol.). Mission.I. The viruses and their taxonomy. 1-18. Avgelis and Rumbos: Double infection of diseased vines by GFLV and CarMV reported. References Francki R. especially near the petiole Leaves are deformed in correspondence of discoloured sectors. CarMV is an isometric virus 30 nm in diameter. 1989. Castellano. Lazar.1987 1989 1992 Lehoczky et al. Lehoczky et al. L.: Purification and characterization of GLPV and suggestion that it is the causal agent of the disease.4 x 106 (4003 nt in size) and coat protein subunits of Mr 38 x 103 Da. Evidence of its grafttransmissibility..P. Phytopathologia Mediterranea 31. D. or variously extended sectors of the leaf blade. Seed transmission of grapevine line pattern virus. Francki. Burgyan. vinifera cv. Boscia. according to more recent findings. the interveinal areas. New York . J. size and have low sugar content. Historical review 1989 1991 Rumbos and Avgelis: Roditis leaf discoloration described in Greece. GFLV may not be involved in the aetiology of the disease. Burgyan. Israel. Polyhedral Virions with Tripartite Genomes (R. G.B. Farkas. L. has a monopartite RNA genome accounting for c. Transmission: No vector is known. M. RODITIS LEAF DISCOLORATION 1.

and ELISA. A. Laski Rizling from Slovenia exhibit a yellow line pattern syndrome resembling the grapevine line pattern disease described from Hungary. Dovas. 112 .: Evidence that GAMV is the agent of grapevine angular mosaic disease. Description Main synonyms: None Main symptoms: Grapevines of cv.E. Rumbos. The etiology of a new virus disease: grapevine angular mosaic. The disease has been recorded only from Greece. F.P.C. but differs from GLPV. A new ilarvirus isolated from grapevine in Greece.M. F. P. Plant Disease 84. thus is regarded as the agent of the disease. Volos 1990. Infected grafting material is likely to be responsible for virus dissemination. Tzortzakakis and M. and I. Bem. 1989. Katis. Kyriakopoulou. Girgis S. decline gradually and some die. Varietal susceptibility: No information Detection: Indexing on cv. Kyriakopoulou. a virus with a tripartite RNA genome and a 30 kDa coat protein. 1991. and A. Infected grapevines are stunted.D. Avgelis. Avgelis. Historical review 2000 2003 Girgis et al. A. Transmission: GAMV is pollen-borne in herbaceous hosts and was able to infect pollinated plants. Bem. P. reproduced the field syndrome in mechanically inoculated grapevine seedlings. 2000.I. References Girgis S. C.: First record of GAMV. Description Main synonyms: None Main symptoms: Symptoms are chlorotic angular spots on the leaf blades. 1345. Roditis leaf discoloration -.I. Tsagris.3. Proceedings 10th Meeting of ICVG. Control: No information 2. Dovas. 2003. discoloration of tissues bordering the veins. 19. Rumbos I. GAMV is molecularly related to a number of ilarviruses. Journal of Phytopathology 125. YELLOW LINE PATTERN (Rasperry bushy dwarf virus) 1. 437-443. the only other ilarvirus reported from grapevine. Sklavounos. 274-278. Avgelis and N.. mechanical transmision to herbaceous hosts. C. GRAPEVINE ANGULAR MOSAIC 1.M.a new virus disease of grapevine: symptomatology and transmission to indicator plants. P.C. P. 3. Baresana x Baresana. Whether this mechanism operates with grapevines has not been ascertained. Flowers abortion results in straggly bunches with small wrinkled berries bearing non viable seeds. Extended Abstracts 14th Meeting of ICVG Locorotondo 2003. crinkling and deformation of the leaves. Agent: Grapevine angular mosaic virus (GAMV). A. N. Katis. Carnation mottle virus isolated from vines affected with "Roditis leaf discoloration". References Avgelis A. Girgis et al.D.E.. the closest being Tobacco streak virus (TSV). S.

M. Raspberry bushy dwarf virus infection of grapevine in Slovenia.8 x 106 Da (2. Control : No information 2. 2003. Varietal susceptibility: No information Detection: Mechanical transmission to herbaceous hosts. Healthy vines of cvs Kyoho and Pione became naturally infected in the field within one year from planting. if any. representing the most important virus disease in Yamanashi Prefecture. Some rootstocks (e. a dimeter of about 33 nm. Chlorotic mottling. a mechanically transmissible definitive member of the genus Trichovirus. Delaware. and RT-PCR. 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. Description Main synonyms: None Main symptoms: Infected grapevines have low vigor.2 kb in size). 30 x 103). 113 . wt 0. 24% of the particle weight and consisting of two functional species RNA-1 with Mol. Transmission: GINV is transmitted by grafting to grapevines and by mechanical inoculation to herbaceous hosts. 3. The virus spreads naturally in the vineyards. Varietal susceptibility: Symptom severity varies with the cultivar. Locorotondo 2003. 165 B. berries are small and show external discolorations and internal necrosis. References Mavric I.5 Kb in size) and RNA-2 with Mol. Mavric et al. RBDV. Vitis riparia Gloire) are also susceptible. GINV has filamentous particles about 750 nm in length and a single-stranded RNA genome with Mol. Raspberry bushy dwarf virus. being transmitted by the eryophid mite Colomerus vitis. the 3' terminal region of which (2469 nts) has been sequenced. and Pione whereas cvs.: First record of RBDV in grapevine. infected propagative material can disseminate the RBDV.Agent: Raspberry bushy dwarf virus (RBDV) was isolated from symptomatic vines. Virscek Marn and I. Grapevine berry inner necrosis has been reported only from Japan. F. Transmission: In raspberry the virus infects progeny seedlings and pollinated plants through pollen. The vay of natural spreading in grapevine.g. and Kaiji are infected latently.5 x 106 Da. Historical review 1976 2003 Murant: Description of RBDV. Koshu. wt of 2 x 106 Da (5. Ripening of bunches is delayed. 1976. 20 Murant A. rings and line patterns are shown by leaf blades. However. wt of 7. Campbell Early are suscetible as well as cvs Takao.. and a bipartite singlestranded RNA genome accounting for c.. No. Japanese diseases GRAPEVINE BERRY INNER NECROSIS 1. CMI/AAB Description of Plant Viruses. Extended Abstracts 14th Meeting of IGVG. Zezlina. delayed bud break and young shoots with short internodes and internal browning. Kyoho. is unknow. Almost all Japanese table grape cultivars derived from crosses with cv. ELISA . Agent: The disease agent is Grapevine berry inner necrosis virus (GINV).

Studies on the grapevine berry innner necrosis virus disease. 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. Yanase H.: Experimental evidence that GINV is transmitted by the the grape erineum mite Colomerus vitis 1993 1997 2000 2000 3. 2.Annals of the Phytopathological Society of Japan 53. Grapevine berry inner necrosis. varietal susceptibility and natural spread. Archives of Virology 142. Bulletin of Yamanashi Fruit Tree Experimental Station 10. Annals of the Phytopathological Society of Japan 51. A new virus disease.. Disease re-named Grapevine berry inner necrosis Terai et al. 423. Terai and Y. Induction of berry necrosis in Kyoho back-inoculated with the virus isolate from grapevine mosaic diseased clones and renaming to grapevine berry inner necrosis. and H. References Kunugi Y. GINV is mechanically transmissible to herbaceous hosts and can be identified by ELISA and moleculat techniques in infected vines. 362. Y. 2. 77-78 Yanase H. Terai. 1997. Asari. 47-56. Nishijima T.. 536 Terai Y. Extended Abstracts 11th Meeting of ICVG. 617. Y.. Magome.Annals of the Phytopathological Society of Japan 58. S. Takahashi and Y. 1992. Kunugi. Mosaic symptoms on cv Kyoho. Yoshikawa et al. Shinkai 2000. S. 1. and Terai Y. Studies on the grapevine berry innner necrosis virus disease. Purification of a filamentous virus isolated from grapevine berry inner necrosis and foliar mosaic. 1351-1363 GRAPEVINE STUNT 1.Detection: Indexing on cvs Kyoho or Pione. 1987. Terai. Yoshikawa N.. H. Description Main synonyms: None.. Kunigi Y. Transmission of grapevine berry inner necrosis virus by the grape erineum mite Colomerus vitis in Yamanashi. Back-transmission of a grapevine filamentous virus to grapevine seedlings and induction of foliar and berry symptoms in grapevine.: Partial sequencing of GINV genome and assignement of the virus in the genus Trichovirus Nishijima et al. Control: Use of tolerant cultivars in areas where the disease spreads epidemically.. Yanase. Montreux 1993. H. 2000. and Yanase H.: An account of the varietal susceptibility to the disease and natural field spread Kunigi et al.. grapevine berry inner necrosis with natural spread in Japan. 1985.: First account of grapevine berry inner necrosis disease in a non Japanese publication. 1984. Y. a new trichovirus: comparative studies with several known trichoviruses. T. Kyoho back inoculated with the filamentous virus isolated from mosaic-diseased grapevines. Iida. Terai and A. Annals of the Phytopathological Society of Japan 55. Tanaka H. Symptoms on vines. Bulletin of Yamanashi Fruit Tree Experimental Station 10. Historical review 1984 1985 1987 1992 Tanaka: Description of a mosaic disease in cv.. 114 . Goto. 57-63. 1993.

. 1987). p. Main symptoms: No appreciable symptoms are visible on the foliage of cv. M. 2.. Because of heat recovery. are pale-coloured and have a low sugar content. Yamashita and Y. Fujii. Yamashita. 1984. Detection: Grafting to Campbell Early and ELISA using extracts from infected vine tissues. Three phloem-limited viruses of grapevine: direct fluorescence detection. FFTC Book series. Annals of the Phytopathological Society of Japan 48. S. The berries. fruit setting is impaired and bunches are few and shelled. with scorched margins. Yora. Historical review.: A small isometric virus associated with stunt disease in Japan. S.: Purification and characterization of the virus associated with stunt disease. which makes the crop unmarketable. GRAPEVINE AJINASHIKA DISEASE 1. internodes are short. 1986. Namba. Fujii. Yamashita. vinifera cv. The same paper appears in Taiwan Food and Fertilizer Technology Center Technical Bulletin 92.. Doi and K. Transmission: The disease is transmitted by the leafhopper Arboridia apicalis. leaves are small. S. Namba. Y. Inflorescences are undersized. In Plant Virus Diseases of Horticultural Crops in the Tropics and Subtropics. Iwanami. Graft transmissibility of grapevine stunt disease. S. 316. Control: Use of disease-free material obtained through heat therapy.: Successful graft-transmission of stunt disease. Spread occurs also through infected propagative material. 396 Hatamoto M. M.109-126. Hatamoto. The disease is apparently restricted to the V. sometimes. however. 1981. S. A small spherical virus associated with grapevine stunt disease. Yamashita and Y. The disease has only been reported from Japan.: Evidence that the disease is transmitted by the leafhopper Arboridia apicalis. References Hatamoto M. The disease has only been reported from Japan. Varietal susceptibility: No information. This virus is serologically distinct from the putative agent of ajinashika disease. American rootstocks are infected without showing symptoms. Agent: An isometric. T. Doi and M. Taipei. Doi. 1981 1982 1984 Namba et al. 115 . Y. curled and. Food and Fertilizer Technology Center for the Asian and Pacific Region. Evidence that it is not related to the presumed agent of ajinashika disease. Hatamoto et al. This condition gives the name to the disease which in Japanese means "unpalatable fruits with low sugar content". Hatamoto et al. non mechanically transmissible virus about 25 nm in diameter is consistently associated with diseased vines and regarded as the possible causal agent. Annals of the Phytopathological Society of Japan 50.Main symptoms: Spring vegetation is delayed. Transmission of grapevine stunt disease by the grapevine leafhopper Arboridia apicalis Nawa. summer vegetation is apparently normal. Annals of the Phytopathological Society of Japan 47. 1-17. Koshu nor any apparent reduction of vigour and yield. 33. 85 Namba S. 1986 Namba et al. Doi. 1982. No.. 137 Namba S. 3. Description Main synonyms: none. phloem-limited. S. Taiwan (Reference 2951 in the Review of Plant Pathology 66. Campbell Early.

. Volos 1990. and D. Ajinashika disease: a combined effect of grapevine leafroll and grapevine fleck viruses on sugar content in the Japanese grape cultivar Koshu. 1980. 1979 1980 1986 1991 1991 Namba et al. 1-17. T.: First mention of ajinashika disease and report of the association with it of a non mechanically transmissible virus with isometric particles. S. Koshu. Namba S. Ajinashika disease of the grapevine cultivar Koshu in Japan. 70-73. Doi and M. and R. Hatamoto. 116 . Namba S. 15-19. Yamashita..: Further characterization of the isometric virus and claim that it is the putative agent of the disease. Control: Use of disease-free material obtained through heat therapy. 1979. A small spherical virus associated with the ajinashika disease of Koshu grapevine. Y. Yamashita. consistently found in infected vines. References Namba S. an isometric. Doi and K. Terai Y. Proceedings 10th Meeting of ICVG. Transmission: No vector is known. S. Dissemination is through infected propagative material. Annals of the Phytopathological Society of Japan 45. Three phloem-limited viruses of grapevine: direct fluorescence detection.: Partial characterization of the isometric virus associated with the disease and its detection by ELISA in infected vines. 2. Yano. 1986. Koshu and ELISA using extracts from infected vine tissues. was suggested as the possible causal agent.. Niagara Falls 1980. Varietal susceptibility: No information. Purification and properties of spherical virus particles associated with grapevine ajinashika disease. Terai Y. Yamashita. Y. Yora. non mechanically transmissible virus about 25 nm in diameter. Namba et al. Tsuchizaki. Gonsalves. 12491253. Terai: Additional report on ajinashika disease as derived from the combined effect of leafroll and fleck. Historical review. Namba et al. Plant Disease 75. Iwanami.Agent: The disease was reported to be caused by the concurrent infection of leafroll and fleck. D. T. 67-70. The disease seems to be restricted to V. 1991. Boscia. phloem-limited. However. No relationship found with fleck. Terai and Yano: Description of ajinashika disease and suggestion that it is caused by the concomitant infection of leafroll and fleck. vinifera cv. S. 1991. Taiwan Food and Fertilizer Technology Center Technical Bulletin 92. Detection: Graft transmission to cv. Proceedings 7th Meeting of ICVG. 3.

VIRUS-LIKE DISEASES .

.

Varietal susceptibility and sensitivity: There is little information available. Primus. Their agents are still unknown.). Historical review 1891 1937 1954 Buchenau: First description of enation disease of grapevine occurring in Germany. omeoplasie crestiformi (Ital.). some of which have a clear-cut detrimental effect on the crop. Grenache and Tokay appear to be quite susceptible and develop severe symptoms when infected. as symptom expression is variable in successive years and graft transmission not 100 % successful. Riesling. however.VIRUS-LIKE DISEASES Several latent or semi-latent grapevine diseases are known. They are outgrowths 2-3 mm high and 3-5 mm long or more.). This hypothesis. which run more or less parallel to the main veins. maladie des énations (Fr. Leaves developed later in the season are usually normal. vinifera varieties in Europe. The crop can be drastically reduced (up to about 50%. Latin America (Venezuela). malattia delle enazioni. Description Enation disease of grapevine is one of the oldest known disorders of European grapes. whether they bear enations or not. 2. North America (California). The basal internodes are short. with drawings of symptoms. has now been dismissed Transmission: By vegetative propagation. its description dating back to the late 1800s Main synonyms: Enationenkrankheit der Rebe (Germ. Graft transmission suggests that it is a virus disease. The varieties Panse Precoce. ENATION DISEASE 1. which gives a bushy aspect to the plant. apparently in relation with climatic conditions. with prominent veins. The infectious agent of the disease perennates in propagating material. 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. Later in the year. Detection: By observing the symptoms on the leaves and by indexing on LN 33 hybrid. Basal leaves. with a fanlike aspect and abnormal indentation. according to the cultivar) and is of poor quality. Symptom expression varies year by year. There is no information on the possibility of curing this disease by heat treatment or meristem culture. irregular and mis-shapen. Main symptoms: Affected vines show a delayed opening of the buds and a slow growth of the shoots in the spring. and is probably due to a virus-like agent. Australia and New Zealand. growth tends to become normal again. The disease has been reported from many European and extra-European countries Agent: The etiology of enation disease is not yet clear. 119 . the absence of symptoms does not necessarily mean that the plant is healthy. However. Enations develop mostly on the underside of the leaves at the base of the shoots. Symptoms have been observed on many V. Hewitt: Description of symptoms in California. but attempts to transmit it by graft or mechanical inoculation gave no results. and often show longitudinal cracks between the nodes. are often mis-shapen. Italia. but some are heat-labile and can be eliminated by heat therapy. North (Tunisia) and South Africa. The disease is perpetuated by vegetative propagation. They are often thicker than normal. Gigante : Research on histological and cytological aspects of enations. All persist in propagative material and are transmitted by grafting. Severely affected leaves drop prematurely. The transmission by graft is rather erratic. Control: Use of healthy material.

Prota et al.: Enation disease in France Pozdena et al. Trebbiano romagnolo and reduces the yield from 13% to 23% according to the severità of symptom expression.: More data on the effects of enation on the yield of cv. Martelli et al. Phytopathologia Mediterranea 17. Beobachtungen und Untersuchungen über die Enationenkrankheit der Rebe.. The possible role of fanleaf virus in the etiology of enation requires further investigation. 195 Brückbauer H. only fanleaf virus was recovered. Xafis. In the vineyards under observation. McGechan: Enation disease in Australia Tekinel et al. the proportion of diseased vines was highest in cv. Nieder: Enation disease in Austria Garau et al.4 to 48. 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. Italia in Sardinia. Malvasia (10. historical account. with negative results. There is some evidence that enation can be carried in the rootstocks. Graniti and Martelli: Review paper on enation. but diseased vines which had not shown enation symptoms for several years had almost normal yields. 120 . and C. Weinberg und Keller 15. attempts to transmit the disease by graft. Confirmation of graft transmissibility of the disease. Padilla et al. 1968. The authors discuss the hypothesis that enation is caused by a strain of fanleaf virus.3% .5%). lowest in cv. Presence of enations in Razaki grapevine in Crete (Greece). the hybrid LN 33 was found to be the most sensitive and reliable indicator variety. Mechanical transmission tests were also negative.: In experiments on graft transmission of enation disease aimed at determining the best indicator. Marinesku and Bondarchuk: Enation disease in Moldova Brückbauer: Influence of enation disease on growth and yield of grapevine in West Germany. .5%). Refatti: Hypothesis of a correlation between fanleaf and enation disease. The authors conclude that the disease is probably of European origin.1966 Graniti et al. Vernaccina (1.: Enation disease in Czechoslovakia Avgelis and Xafis: Enation disease in Greece Prota and Garau: Enation disease found in Sardinia. Brückbauer: Description of symptoms of enation in Germany and confirmation of graft transmission of its agent.: Detailed description of macroscopic and microscopic symptoms of enation disease. References Avgelis A. Enationaffected vines produced less than 50 % of the yield of healthy plants. but report on observations made in Australia where no fanleaf virus could be recovered from enation-affected vines. 79-112.: Enation disease in Turkey Hevin et al. The mean yield loss of diseased vines ranged from 17. although the rate of symptom expression does not exceed 30% Credi: Enation diseasae affects the vegetative vigour of cv. 1978.: Successful transmission of enation disease from diseased to healthy grapevine by graft strongly supports the hypothesis of a viral origin. and possibly caused by a virus.

.. 17. Martelli G. Main symptoms: Pale green mosaic affecting mostly the tissues adjacent to the main veins or the smaller ones.V. O. 1975. Deutsches Weinbau-Jahrbuch 31. Phytopathologia Mediterranea 20. A study of infectious degeneration (fanleaf) in vineyards of the Mediterranean region. IV. and M. 1981.IV. 1976. Graniti. Roma. This disease is widespread and probably worldwide. Israel. 47. 1966.. 1970. Description The symptoms of vein mosaic have been confused for some time with those of fanleaf/yellow mosaic. Bulletin of the California Department of Agriculture 43. Proceedings 9th Meeting of ICVG. 1891. Pflanzenharzt 36. Salih and Y. Studies on some variable characters of grapevines affected by enation disease in Sardinia. Benayas. 47-64. Enations of grapevine in Sardinia. Tekinel N. McGechan J. Bitki Koruma Buletin 11.. Credi R. 1954. 1973. 1987. 48.. Lisbon 1997. Prota U. Einfluss von Virusinfektionen auf Wachstum und Ertrag der Rebe. Cugusi. Rives. Prota U. Proceedings 6th Meeting of ICVG. Petria 6. B. Proceedings International Conference on Virus and Vector on Perennial Hosts. Graniti A. 2. VEIN MOSAIC 1. it was clear that vein mosaic was not caused by fanleaf virus. Division of Agricultural Sciences.. 293-306. Occurence of grapevine enations in the Moldavian SSR. 1965. G. Hevin M. Vanek. Frazier N. 179-189. Berkeley. 203-206.. and R. 1970. 349-352. In: Virus Diseases of Small Fruits and Grapevines (A Handbook). Die Eantionenkrankheit der rebe erstmal auch in Osterreich nachgewiesen. Phytopathologia Mediterranea 5.). Mosaico delle nervature (Ital.G. Extended Abstracts 12th Meeting of ICVG. 326332.. 1966. Gigante R. n. 122-124. Mededel. Garau. Enation symptoms found in France. The research of the virus diseases of grapevine in CSSR. Main synonyms: Mosaïque des nervures (Fr.). Dolar. Moldavian Research Institute of Horticulture Kishinev.B. Lamberti. University of California. 59-64. Prota and M. Salcan.. Adernmosaik (Germ. Vein mosaic appears to be of low economic importance.W. 1966. Ser. but these necroses do not affect the veins as it is the case with vein necrosis. Enations. small fruit and grapevine in Moldavia. (ed. Lisbon 1997. Garau R. 823-827. Lamberti and A.D. 1979. producing often a vein banding effect. 46-51. Studies on reproduction of enation symptoms by grafting in Sardinia. Pozdena J. H. M. 225-246.. Marinesku V. 169-192. areas of the leaf blade may become necrotic. Agricultural Gazette of New South Wales 81.K. 251-252 Hewitt W.. Kiryat Anavim. Graniti A. Nas. Trasmissione per innesto della "malattia delle enazioni" della vite. Effetto della malattia delle enazioni della vite sulla produzione e sullo sviluppo vegetativo della cv Trebbiano romagnolo.S.P. Rivista di Patologia Vegetale S. Refatti E.P.. 1971. Chabbouh N and V.s. Symptom expression seems to depend on climatic conditions. Enation disease of grapevine in Italy.). Su una possibile correlazione fra il virus del complesso dell'arricciamento e la malattia delle enazioni della vite. Monografias INIA 18. In: Verderevskaya T. but when fanleaf virus transmission to herbaceous hosts became possible. 97-98. I. A. and V. G. Garcia. and G. Buchenau F. Facultet Landbouwwetenschap (Gent) 40. 121 . (N. Extended Abstracts 12th Meeting of ICVG. 1937. Garau R. California. Savino. 1980. Occurrence of enation disease in Tunisia.P. In the most sensitive varieties.) Virus and mycoplasma-like diseases of fruit trees. Ita and F. Rivista di Patologia Vegetale. Z.Brückbauer H. 1997. Abnorme Blattbildungen. Bondarchuk. Some virus and virus-like diseases of grapevines. U. with Special Reference to Vitis. 7-12. A similar disease has been reported in Australia under the name of summer mottle. And G.P.). Ricerche istologiche sulle omeoplasie crestiformi (Enations) delle foglie di vite affette da rachitismo. Gazeau. Cordoba. 145-152. Cugusi. Bolletino della Regia Stazione di Patologia Vegetale. 1983. 207-217. 9.1989. Martelli. 1983. F. Nieder G. Davis. Important virus diseases of grapevine in New South Wales. Leclair and M. Quacquarelli. 1997.W. Martelli and F.. Padilla V. Grapevine enation disease in Murcia (Spain). 1996. 241-243. Bericht der deutschen botanischen Gesellschaft 9. Spain..

Milkus et al.R. Chasselas. Abracheva: Vein mosaic in Bulgaria.: Observation of a type of mosaic of grapevines which appears to be independent of fanleaf virus.C. and R. vinifera cvs.A. Several V. Bonfiglioli: Vein mosaic in New Zealand (unpublished). Occurrence of grapevine vein necrosis and grapevine vein mosaic in the Emilia-Romagna region (northern Italy). LN 33 is also sensitive. Vuittenez and Stocky: Electron microscope study of thin-sectioned tissues of leaves from Vitis riparia and Vitis vinifera cv. 1985. Krake L. 148-152. Control: Use of indexed material. No claim is made that these putative viruses are connected with the disease. Grapevine summer mottle: a new graft-transmissible disease.: Vein mosaic in Ukraine. A. Servant. Babini and A. Legin and Vuittenez: Description of vein mosaic.: Vein mosaic in URSS. 266-270. Transmission: By graft and vegetative propagation. Marinesku and Bondarchuk: Vein mosaic in Moldova. 17-23. Canova. Viognier.Agent: Unknown. Varietal susceptibility and sensitivity: Vitis riparia Gloire de Montpellier is especially sensitive and is used as indicator. Pearl of Csaba). The disease can be eliminated by heat therapy. Golino: Vein mosaic in California. Mycoplasma-like organisms were supposed to be the cause of vein mosaic. Vitis 17. in the absence of any detectable virus. vein mosaic and vein necrosis. Symptoms are very similar to those of vein mosaic in Europe. Kuniyuki: Vein mosaic in Brazil. Woodham and Krake: Comparison of summer mottle and vein mosaic. American Journal of Enology and Viticulture 44.. Saric and Hranuelli: Vein mosaic in Croatia. Muscat de Hambourg. and Gamay apparently show little or no symptoms. Pop: Vein mosaic in Romania. Comparison of symptoms of fleck. Alphonse Lavallée.. riparia Gloire de Montpellier. 1978. Pinot. No vector known. Chardonnay. Phytopathologia Mediterranea 24. Detection: Indexing with V. 1993. Samonina et al. 122 . References Credi R. 2. Krake and Woodham: Description in Australia of a systemic mottling syndrome which is expressed during summer on the leaves of some varieties. but this hypothesis has not been confirmed. Potential interaction between rootstocks and grapevine latent viruses. show symptoms (Syrah. 1979 1980 1982 1983 1985 1993 2004 3. Golino D. Historical review 1966 1973 1973 1976 1973 1973 1978 Vuittenez et al. Ehrenfelser showing symptoms of vein mosaic. A number of cytological modifications primarily involving chloroplasts were observed along with the presence of bundles of filamentous structures resembling closterovirus particles.R. Woodham.

Bondarchuk. whereas the opposite occurs with summer mottle. Rivista di Patologia Vegetale. Investiation on grapevine viruses in the SR Croatia. Niagara Falls 1980. 1. Proceedings of the Conference on Excoriosis and Virus Diseases of Grapevine. Transmission: No vector is known. SUMMER MOTTLE Summer mottle. Rivista di Patologia Vegetale S IV 9. Vinogradr. and L. 1973. Historical review 1978 1982 1983 Krake and Woodham: Description of summer mottle in Australia. Krylova.. 2. resembles in some respects the European vein mosaic and the Greek Roditis leaf discolouration (see under Minor virus diseases). 1982. A comparison of grapevine summer mottle and vein mosaic diseases. La mosaique des nervures. IV . producing a feathering or banding effect. These symptoms appear in summer and persist through the autumn.V. Kuszala. Kuniyuki H. IV. Stocky. Comparaison des symptômes et transmission par greffage d'une mosaïque nervaire de Vitis vinifera .C.. Bunches of infected cvs Sideritis and Cabernet sauvignon are fewer. an Australian disease. Ser. probablement indépendante du virus du Court-noué. 1985. Ultrstructure de vignes infecteés par deux maladies de type viral: la mosaique des nervures. 9 . Woodham and Krake: Comparative graft transmission trials demonstrate that summer mottle 123 . Krake. 9. S.G. Vinodel. and A. Legin and J. rupestris et d'une affection nécrotique des nervures de l'hybride Rup. ou la “feuille rouge”.. B. 1983. Symptoms show on vegetative growth that develops at temperatures in excess of 30 °C. Vitis 22.R. Pop I. 243-250 Samonina I. However. de la marbrure de V. Symptoms of vein mosaic develop under mild weather conditions and fade during hot weather.-Berl. 57-63. Vuittenez. Main symptoms: Pale green to yellowish dicolourations of the tissues adjacent to the main or secondary veins. maladie à virus de la vigne. Cabernet sauvignon. 149-141. Description Main synonyms: None. 1966. rupestris and LN33 are infected symptomlessly. Woodham R. 1973. Rivista di Patologia Vegetale . e. poorly developed and with small berries. Proceedings 7th Meeting of IGCV. 67-73.V.. Mataro. suspected to be a virus or a viroid.: Elimination of the disease agent by culturing fragmented shoot apices. A. Summa Phytopathologica 11. Annales des Epiphyties 17.Legin R.N.. A grapevine virus disease in the Primorye territory. Barlass et al.V. R. Mostar 1977. and Hranuelli T. Cabernet franc. 247-252. Roditis leaf discolouration and summer mottle have similarities suggesting that they may be the same disease. 110 R.N. Adverse effect of light and of high temperature on symptom expression of grapevine vein mosaic in Sao Paulo. Saric A. 1973. 48-49 Marinesku V. 191-204. and G. Agent: Unknown. Numéro hors série. Mission. 1977. Vuittenez A. Krylov and V. Grapevine vein mosaic. Evidence that the disease is graft-transmissible.V.g. Control: Use of disease-free propagating material obtained by culture of fragmented shoot apices. 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. Milkus. 41-42. Observations sur une Mosaïque de la Vigne. URSS. and V. Vuittenez A. 68-72. 1976. Moldavii 31. several European grape cultivars show symptoms. Detection: Graft transmission to a number of cvs.

Little is known about sensitivity of other Vitis species..A. 266-270. K. Main synonyms: Nécrose des nervures (Fr. Necrotic reactions are best seen on the lower face of the leaf blade. necrosi delle nervature (Ital.). Rezaian and R. Vitis 17. 1999. Description Vein necrosis has probably a worldwide distribution. 1. R. Historical review 1973 1978 1978 Legin and Vuittenez: Discovery and description of this virus-like disease while searching for indicators for fleck.R. 1978. 247-252. No vector known. especially under greenhouse conditions. N. Martelli et al.). Adernnekrose (Germ. The agent of vein necrosis can be eliminated by heat therapy. Agent: Suspected to be a virus. and some infected plants may die. 291-295. growth is much reduced and necrosis of the leaf veins appears. 2.G. This strongly supports the hypothesis that vein necrosis is a specificic reaction of the rootstock 110 R to GRSPaV infection. Graft-transmissible Diseases of Grapevines. VEIN NECROSIS Although vein necrosis is currently regarded as a disease in its own right. Main symptoms: On the rootstock 110 R. Annals of Applied Biology 101. Also the tendrils and many shoots can necrotize. at first on the leaves at the base of the shoots. Woodham. Krake.: Vein necrosis in Italy and Bulgaria 124 . most likely GRSPaV. berlandieri 110 Richter (110R) with vein necrosis symptoms. Woodham and L.: Suggestion that summer mottle and Roditis leaf discolouration are the same disease 3. Milkus and Kalashyan: Mycoplasma-like organisms found in phloem tissues of vines with vein necrosis. and R.R..M. Vitis 22. 1983. Krake L. recently an association exceeding 95% has been experimentally observed between Grapevine rupestris stem pitting-associated virus (GRSPaV) and vines of V. Possible viroidal etiology put forward. Taylor. Australia. A comparison of grapevine summer mottle and vein mosaic diseases. 1999 Krake et al.C. later on younger leaves as they develop. rupestris x V. varieties or hybrids. Grapevine summer mottle: a new graft-transmissible disease. So far. Krake. Krake L. 1982.differs from vein mosaic. Many grapevine varieties and rootstocks are infected symptomlessly.. Control: Use of indexed planting material.R.S. 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. Varietal susceptibility and sensitivity: The rootstock 110 R is most sensitive.R. CSIRO Publishing. M. Woodham R. References Barlass M. and the only Vitis species that is clearly affected is the rootstock 110 R. Transmission: By grafting and vegetative propagation.C.H. Regeneration of virus-free grapevines using in vitro apical culture. and L. Mycoplasma-like organisms have been observed in the phloem of symptomatic vines.). its economic importance has not been assessed. Cause-effect relationships between MLOs and the disease has never been ascertained.C. Skene. Detection: By grafting on 110 R. Scott. 70-74. Collingwood. but their etiological relationship with the disease has not been proven.

148-152. 1993. Heat therapy reduced this proportion to 35. Lehozcky J. Fitopatologia Brasileira 19. Savino. Phytoparasitica 17. Vein necrosis: new virus-like disease in Turkish vineyards.B. V.N. and L R. Kergazdasag 18 (4). Incidence of some graft-transmissible virus-like diseases of Grapevine in visually selected and heat-treated stocks from Southern Italy. Farkas. IV.. 2004. and L.: Vein necrosis in Malta Golino: Vein necrosis in California Khun: Vein necrosis in Brazil Bouyahia et al. Boscia and V. M. Boscia and G.. D. the incidence of vein necrosis in visually selected stocks of table and wine grape varieties is on the average 71 %. 58-60. P. Journal of Turkish Phytopathology 17. 61 Savino V. Viruses of grapevine in Malta. and A.1984 1985 Woodham R. 1985. 1986 1988 1989 1992 1993 1994 2004 3. Suggestion than vein necrosis is a specificic reaction of 110R to GRAPaV. Boscia. Comparaison des symptômes et transmission par greffage d'une mosaïque nervaire de Vitis vinifera. References Bouyahia H. Bulletin OEPP/EPPO Bulletin 22. fleck and leafroll in Vitis vinifera and grapevine rootstocks in central Greece. Martelli. Mycoplasma-like bodies in phloem tissue of grapevine affected by vein necrosis. 3-5 Martelli G. A.P. 204-207.. Ser. Savino. P. Galea Souchet. 79-83 Legin R.A. but did not eliminate the disease entirely. Savino. Gursoy Y.P.P.A. 1988.C. Golino D. Krake: Vein necrosis in Australia Savino et al. 1978. Lazar. No veing necrosis observed in 110R top grafted on GRSPaV-free V. Detection of vein necrosis virus (GVNV) in the vines of cultivated grape varieties. 1978. SSR. Informatore Fitopatologico 28 (10). D. i Chim Nauka 1. Grapevine vein necrosis disease detected in rooststocks in Australia.Z. Is Grapevine vein necrosis a reaction to Grapevine rupestris stem pitting-associated virus? Journal of Plant Pathology 86. G.. Martelli. 1994. ser. American Journal of Enology and Viticulture 44. Necrosi delle nervature della vite in Italia e Bulgaria.-Berl. Krake. 1989.: An association exceeding 95% observed between GRSPaV and 110R vines showing vein necrosis symptoms in indexing trials. Vein necrosis. 606-612. Lehoczky et al. 29-30. Phytopathologia Mediterranea 24.. Rumbos I C. Milkus B. 1992. V. H. 1973.: Vein necrosis in Hungary Gursoy: Vein necrosis in Turkey Rumbos: Vein necrosis in Greese Martelli et al.. 1984.R.P. Martelli G. Kalashyan. Journal of the Australian Institute of Agricultural Sciences 50. 1986. Abracheva and B. J.. Vein necrosis a disease that is latent in most grapevine cultivars of the State of Rio grande do Sul. Vuittenez. 110 R.5 %. Potential interaction between rootstocks and grapevine latent viruses. Izvestiya Akademii nauk Moldav. 301. 43-45 Kuhn G. 57-63.. Biol. 59-65. rupestris. D.. rupestris et d'une affection nécrotique des nervures de l'hybride Rup. 9. de la marbrure de V. Woodham R. Rosciglione. Castellano. and J. Rivista di Patologia Vegetale.A. 125 . Pirolo. La Notte.: In southern Italy. Minafra and G. C.C.

.

VIROID (Yellow speckle) .

.

Both these viroids wer first isolated in Australia. the type species of the genus Hostuviroid. Like viruses. AGVd has been reported from Australia. AGVd. 129 . R. are not associated to any specific symptomatology. These symptoms appear in the height of summer on a limited number of mature leaves and persist for the rest of the vegetative season.).). infected vines are symptomless or show symptoms erratically. picchiettatura gialla (Ital. Description Main synonyms: Moucheture jaune (Fr. Collingwood. It was first detected in Japan and transmitted to cucumber and grapevine seedlings in which. Flores. the non coding genomes. 2003. Interestingly. Sometimes. Australian grapevine viroid (AGVd). Randles and J. Agents: Two distinct viroids.S.. all of which belong in the family Pospiviroideae: Grapevine yellow speckle viroid 1 (GYSVd-1). a size much smaller than that the smallest viral genome. north and south America. viroids are classified in families. however. CSIRO Publishing. 370 pp. Viroids. Cabernet franc and a cv. vein banding-like symptoms can be observed in vines infected only by yellow speckle viroids. They are made up of a non encapsidated circular RNA of 246-375 nucleotides. J. however. are subviral pathogerns endowed with autonomous replication in their hosts. and plastidial replication (Avsunviroideae ).). YELLOW SPECKLE 1. Two families are known. 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. genera and species. GYSVd-1 and GYSV-2 have a worldwide distribution The three additional viroids that have been detected in grapevines. and may have a worldwide distribution. or gathering along the main veins to give a vein banding pattern. Very often. The symptomatology varies depending on the cultivar.W.VIROIDS Viroids. thought to be elicited by a specific strain of GFLV. and perhaps the type of infecting viroidal sequence variant. has a genome 369 nt in size. HSVd-g. 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. presence of ribozymes. GYSVd 1 and GYSVd 2 are made up of 366 and 363 nucleotides (nt). inducing a disease called yellow speckle. respectively and both belong in the genus Apscaviroid. Main symptoms: Few to many minute chrome yellow spots or flecks scattered over the leaf surface. Neither of them is able to replicate in herbaceous hosts but both were succesfully inoculated to grapevine seedlings reproducing the yellow speckle syndrome. Hop stunt viroid grapevine strain (HSVd-g). Citrus exocortis viroid grapevine strain (CEVd-g). Five grapevine-infecting viroids are known. 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. plant age. the USA. has a genome 297 nt in size. References Hadidi A. and AGVd. Vein banding. respectively from a cv. it did not induce symptoms. HSVd-g has been recorded from Australia. Europe. a member of the genus Apscaviroid. and Tunisia. Semancik (eds.). HSVd-g. Grapevine yellow speckle viroid 2 (GYSVd-2). 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. CEVd-g. GYSVd-1 and GYSVd -2 cause the disease individually or in combination. Only GYSVd-1 and GYSVd-2 are pathogenic. Kyoto vine with yellow speckle symptoms. climatic conditions. a disease characterized by chrome yellow flecks localized along the main veins of mature leaves and progressing into the interveinal areas. was demonstrated to be caused by a co-infection by yellow speckle viroids and GFLV. Gelbsprenkelung der Rebe (Germ.

found in grapevine accessions from Europe and California. GYSVd-2. its grapevine strain so far has only been recorded from Australia and the USA. a disease formerly thought to be caused by a chromogenic strain of GFLV. and distribution of infected propagating material. Garcia Arenal et al.: Evidence that viroids are widespread in grapevines. Flores et al. Varietal susceptibility: All Vitis species.: Successful mechanical transmission of viroids to grapevines. It was first recoverd in Spain from symptomless grapevines.CEVd-g. Mink and Parsons: Yellow speckle can be detected by growing vines for 2-3 weeks at 32 °C under continuous illumination. In the great majority of grapevine germplasm infection is latent. Krake and Woodham: Evidence that the agent of yellow speckle is implicated in the etiology of vein banding. 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. Sano et al. has a genome 369 nt in size. Transmission: No vector is known. 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). Prota et al. First identification of AGVd Koltunow and Rezaian: Identification and sequencing of grapevine yellow speckle viroid. 2. a member of the genus Pospiviroid. yellow speckle and fleck through dodder. Woodham and Krake: Artificial transmission of grapevine leafroll. Shikata et al. if not all citrus-growing countries.: Yellow speckle eliminated by in vitro apical culture. CEVd-g and AGVd.: A vein banding condition of cv.: Reconstruction of the secondary structure of CEVd-g Szychowski et al. Semancik et al. Seed transmission has been demostrated for GYSVd-1. Rezaian et al. For yellow speckle.: First recovery of a viroid from grapevines in Japan.: Two new viroids. one of which identified as the agent of citrus exocortis. Control: Use of viroid-free propagative material obtained by meristem tip culture. Abracheva et al. Although CEVd is present in most. Three different viroids found in a number of accessions in a Californian varietal collection. besides Spain. as the disease may have spread naturally. grafting. hybrids and cultivars appear to be susceptible. 1984 1985 1985 1985 1987 1987 1988 1988 1988 130 . Natural dissemination takes place by mechanical inoculation through surface-contaminated cutting tools during management operations. Barlass et al. the authors consider the results as inconclusive. Experimental transmission through dodder is possible. 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.: The Japanese grapevine viroid identified as a strain of hop stunt viroid. Woodham and Krake: Evidence of field spread of yellow speckle. Rcatzitelli resembling yellow speckle reported from Bulgaria.: Four viroids found in Australian grapevines. Detection: Some viroids can be transmitted mechanically to herbaceous hosts but this is not an efficient detection method.: A disease of cv.

: First report of AGVd in the Mediterranean. 131 .: Review of viroids. 127-130. based on phylogenetical analysis of hop and grapevine isolates of this viroid. 291-295.: Structural analysis reveals that five distinct viroids infect commercial grapevine varieties. N. Koltunow and Rezaian: Description and sequencing of grapevine viroid 1B (later renamed Grapevine yellow speckle viroid 2). 1976. according to an international agreement reached during the 10th Meeting of ICVG held in 1990 at Volos. Production of viroid-free grapevines by shoot tip culture. 1991 Semancik and Szychowski: There are two classes of grapevine viroids: (i) apparent viroids.. Grapevine yellow speckle viroid 1 (GYSVd 1). Spain. GYSVd-1 and GYSVd-2 Rezaian: Complete nucleotide sequencing of AGVd. Juarez and J.: A survey of viroids of grapevine in Italy. which can readily be isolated directly from grapevines.. Cordoba. Journal of General Virology 66. K. (ii) enhanced viroids. Minafra et al. R.: First record of grapevine viroids in China.M. American Journal of Enology and Viticulture 39. First report of Australian grapevine viroid from the Mediterranean region. Regeneration of virus-free grapevines using in vitro apical culture. 2003. Perreault and H. Pallas and J.: Suggestion that the viroid causing stunting in hop (HSVd) originated from grapevines. 2095-2102. Greece. Koltunow et al. Martelli. Krake. Journal of Plant Pathology 85. Citrus exocortis viroid grapevine strain (CEVd-g). Quacquarelli and V. Monografias INIA 18.1988 1989 1989 1989 1990 1990 Duran-Vila et al.C. J.b Szychowski et al. are to be named as follows: Hop stunt viroid grapevine strain (HSVd-g). Sano et al. Arregui. Barlass M..: Extensive comparative analysis of grapevine accessions from California and Europe reveal a similar pattern of viroid distribution. Skene. CEVd-g and AGVd via grape seeds. which require amplification in an alternate host. 1978. Martelli: Brief review of grapevine viroid situation supporting the idea that vein banding is primarily induced by viroidal rather than GFLV infection. Duran-Vila N. 1982. Annals of Applied Biology 101. The occurence is reported of HSVd. Grapevine yellow speckle viroid 2 (GYSVd 2) Australian grapevine viroid (AGVd).. References Abracheva P. Flores R. J. Woodham and L.R. M. V. Wang et al. Savino. Wan and Symons: Transmission of GYSVd-1. A.: Improvement of meristem tip culture technique for the production of viroid-free grapevines. 217-220. Little and Rezaian: Updated review of grapevine viroids. Fakhfakh. 45-57.: Evidence that two related viroids (GYSVd 1 and GYSVd 2) can cause yellow speckle disease independently. G.G. Proceedings of the 6th Meeting of ICVG. 1985.M.P. Molecular evidence that this viroid originated from recombination between five different viroids among which GYSVd-1 and GYSVd-2 1991a. Semancik. 1988.. A possible virus disease of Rcatzitelli vines in Bulgaria. GYSVd-2. Elleuch et al. These viroids. Marrakchi. Duran-Vila.P.S. Elleuch A. Flores et al. Detection of viroid and viroid-like RNAs from grapevine. 1991 1996 1997 1999 2001 2003 2003 3. Rezaian et al.

A. 1990.H.. Mimura and K.A.Flores R.. Rezaian. Goheen and J. sanitation and situation in the Arab countries. 3411-3419. 1991. Vitis 21.R. yellow speckle and fleck diseases by dodder. Uyeda.W.. 337-345. Flores.. Shikata. Parsons. Flores. Vitis 29. Szychowski J. Investigations on a vein banding disease of Grapevine in Sardinia. Prota U. Krake.A.S. 869-872. Mechanical transmission and rootstock reservoirs as factors in the widespread distribution of viroids in grapevines. 210-219. and J. An infectious low molecular weight RNA was detected in grapevines by molecular hybridization with hop stunt viroid cDNA. M.Leclair. 1972. 24-28.A. 287-288. Grapevine yellow speckle disease: studies on natural spread observed in the field.P. R. Grapevine yellow speckle -.M. Vitis 30. Grapevine yellow speckle agent implicated in the aetiology of vein banding disease. 193-198. Viroids of grapevines in Italy. Krake.W. R. 132 . Woodham R. Phytopathologische Zeitschrift 106. Rivera-Bustamante and A. Woodham. Isolation of three viroids and a circular RNA from grapevines. Transmission of viroids via grape seeds. A. Rezaian M.Woodham. T. Ohno and Y. 1987. Krake L. 195-206. Sano and I. Symons. 1985. Seminars in Virology 8. Journal of Phytopathology 147. Viroids: the non coding genomes. Proceedings of the 10th Meeting of ICVG.H. G. and J.. J. E. Wolpert and J.C. Woodham R. 53-59. and L. Rezaian. Savino. Ohshima. Rezaian M.S. Cugusi and M.P. 2003. V. A. 1990. Semancik.A. Zhang and X.P. 297. Duran-Vila. 1975. S. Dore. 285-291 Wang G. Di Serio and C. 413-422.. Koltunow A.. 1988. Vitis 22.R. 5587-5598. Rezaian. Rezaian M. L.. 1987. F. Z. Meshi.R. 25-36. Relationships among grapevine viroids from sources maintained in California and Europe. R. Journal of General Virology 69.R. Nucleic Acids Research 10.S. and R. Phylogenetic analysis of hop and grapevine isolates of hop stunt viroid supports a grapevine origin for hop stunt disease.C. Wolpert and J. Krake. Grapevine viroids. 1997. Krake and K. Zhang.R. M..A. N. A viroid-like RNA isolated from grapevine has high sequence homology with hop stunt viroid.. Two related viroids cause grapevine yellow speckle disease independently. Australian grapevine viroid . Duran-Vila. Pallas and R. Koltunow A. Semancik. 1985. Johnson and M. I. Hadidi. 1983. Martelli and V. Collingwood. China Fruits 4. Credi. Nucleic Acids Research 15. Arab Journal of Plant Protection 7. 1990. S.M. 1983.C. Infectious diseases of grapevines. Little A. Proceedings of the 10th Meeting of ICVG.G. 213-216. J. The field investigation of virus and viroid diseases of grapevine and stone fruit trees in Shandong and Liaoning province. Goheen. 39-41. Credi. Rapid indexing procedures for detecting yellow speckle disease in grapevines.C. J. Doazan. 4203-4210. Garcia Arenal F.. Semancik J. Sano T.A. 1984. Journal of General Virology 66.. The sequence of a viroid from grapevine closely related to to severe isolates of citrus exocortis viroid. Australian Journal of Agricultural Research 23.C.A. Plant Disease Reporter 59. 333338. Skene. 35-40.. Greece. 2001. P. J.A. 1989. Szychowski.. Nucleic Acids Research 16. T.D. Uyeda. Koltunow and L. American Journal of Enology and Viticulture 38.M. Garnier. 1999. Phytopathologia Mediterranea 24.evidence for extensive recombination between viroids. 65-73. American Journal of Enology and Viticulture 39. Koltunow. Taylor R. Nature. a new member of the apple scar skin viroid group contains the left terminal region of tomato planta macho viroid.). Minafra. Rezaian. J.I. Okado. R. Minafra. Grapevine yellow speckle viroid: structural features of a new viroid group. Mink G. 1990. T. 849-864. L. N. 1996. Grapevine viroids. and R. Volos. 447-452.R. M.a newly recognized grafttransmissible disease of Vitis.A. Investigations on transmission of grapevine leafroll. 1991b. Journal of General Virology 70. Proceedings of the Japanese Academy of Science 60(B). 40-50. and L. In: Viroids (A. A.Jiang.S. 173-182. Volos. Sano T. 202-205. P. 1990.S. Proceedings 10th Meeting of ICVG. A. Volos. Leclair. 575-578. and R. R. Comparative properties of viroids of grapevine origin isolated from grapevines and alternate hosts. Minafra A. 1988. detection. 1988.A. Semancik J. Widespread occurrence of viroid-like RNAs in grapevines. 1991.P. Grapevine viroid 1B. Szychowski J.A. Doazan. China. and M. Greece.M.A. Krake.P. CSIRO Publishing.A. Koltunow A. Hernadez. Shikata E. 1989. Martelli G. 370 pp. 1989.S. 270-278. N.C. Semancik. Greece. 1991a. Virology 170.M. and M. and M. Szychowski J.L. Randles and J.. Hong. R. Garnier. Wan C. Relationship and patterns of distribution among grapevine viroids from California and Europe. Semancik (eds. Garau... 1982. Virus Genes 22.

YELLOWS .

.

GY agents have been identified in no less than 5 groups of phytoplasma clade. Downwards rolling of the leaves and sectorial discolorations of the blades. involving also the main veins. Leaf blades are crispy and brittle. it was mainly in the 1990's that GYs could be differentiated. Nevertheless. Rubbery canes fall downwards with a typical "weeping" aspect. host range. in addition to other criteria such as symptomatology. Phytoplasmas form a homogeneous phylogenetic clade subdivided into about 20 groups and subgroups. phloem-restricted bacteria that belong to the class Mollicutes. The main characteristics of GY diseases are important losses or damage to the yield. buds. Candidatus Phytoplasma australasia. Their titre is usually very low in grapevine. canes. Vergilbungskrankheit. In 1997. all organs of the plant may be infected. Flavescence dorée (FD) and Palatinate Grapevine Yellows (PGY) phytoplasmas belong to group 16SrV (or Elm yellows group). resulting in reduction of quantity and quality of the crop. and serology. Main synonyms: Flavescence dorée-like disease. Phytoplasmas have the smallest genome reported for procaryotic organisms (560 2. including roots. shoots. 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). Amarilliamento. Main symptoms: Young shoots of affected Vitis vinifera are weak with short internodes and frequent necrosis of terminal buds. Flavescenza dorata. climate and infection pressure. Phytoplasmas in the 16SrIII group (or X-disease group) are associated with grapevine yellows in North America and Israel. depending on the particular disease and other conditions such as variety. Description Grapevine yellows (GYs) are a group of severe phytoplasma-induced diseases occurring worldwide on a number of Vitis vinifera cultivars. the term phytoplasma has become the genus name of these plant pathogenic agents under the provisional taxonomic status Candidatus. 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. but not seeds. Quite often. Bois noir. Several Candidatus phytoplasma species have been recently described. 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. after their aetiology had been understood and molecular methods for the characterization of associated phytoplasmas had been developed. Phytoplasma cells are vesicular rounded bodies. based mainly on sequence similarity of their rRNA genes and of a few other genes. associated to Bois noir (BN). persistence of infection in dormant plant material. Golden flavescence. Nonetheless. a severe decline of the vine. However. Legno nero. The different GYs cannot be differentiated on the basis of symptoms. develop on leaves. Vergilbungskrankheit. Schwarzholzkrankheit. However. The occurrence and spread of a particular GY mainly depend on the presence of efficient vectors in the vineyards or their close vicinity. They are wall-less. Bunches wither in early summer or berries shrivel later in season. vector species have not been identified for all GY diseases. such as the ribosomal protein genes or the elongation factor Tu. though some sieve tubes may appear crowded with phytoplasma cells in the electron microscope. However. Australian grapevine yellows. autumn fall of leaves occurs later on diseased than on healthy vines. North American grapevine yellows.200 kbp). Flavescenza dorata. and transmission by specific leafhopper or planthopper vectors.GRAPEVINE YELLOWS: GENERAL PROPERTIES 1. inflorescence and berries. their genome is poorly known because they cannot be cultivated. their movement is slow and their distribution in the plant is erratic. a phytoplasma belonging to group 16SrII (or peanut witches' 135 . Red-berried varieties show reddish to purple discolorations while white-berried varieties show golden to chlorotic discolorations. The first GY to be recorded was Flavescence dorée in France in the 1950's. 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. Main diseases: Flavescence dorée. Palatinate grapevine yellows. Stolbur phytoplasmas. Severely infected stocks decline rapidly.

A range of primers that can be used for amplification with Polymerase Chain Reaction (PCR) of characteristic DNA fragments of phytoplasmas. are available in the literature. 136 . erratic transmission to grapevine may nevertheless take place. Outstanding progress in detection was achieved with DNA-based techniques. Detection can also be made by transmission electron microscopy of thin sections or by scanning electron microscopy. are critical. These methods are not specific for any phytoplasma and are too laborious for disease monitoring. influence symptom expression and differential sensitivity of cultivars. Phytoplasmas persist in their vectors but vertical transmission to the progeny of infected gravid females is not possible or scarce. canes. Though the rate of transmission to plant material can be very low. The best antigen source are leaf veins and petioles. Particular procedures for extraction of phytoplasma antigens from infected vines must be used to enhance the sensitivity of detection. Vitis riparia and American rootstock varieties and hybrids do not show typical symptoms of GY. PCR amplification with universal primers is preferred. Phytoplasmas in the 16SrI group (or aster yellows group) are associated with endemic GY diseases in several countries of Europe and in the USA. The extraction from infected tissues of total DNA containing enough phytoplasma DNA of good quality and the elimination of inhibitors of the PCR reaction. Infection of rootstocks has been detected only in the case of Flavescence dorée but it cannot be excluded for other GY agents. vinifera cultivars have been reported to be sensitive or very sensitive to GYs in all countries where infection occurs. These phenomena also depend on the disease complex (phytoplasma. The situation of cv Syrah is controversial. Individual symptoms can be confused with other diseases or disorders. However. Then. are very sensitive to all GY diseases. Others. Detection: GY syndrome is characteristic. shoots. Double infections have been reported. designed on variable regions of the rDNA. Phytoplasmas also persist in vine stocks during winter. Cabernet Sauvignon. Most of these PCR primers have been designed on conserved regions of the rRNA gene and are "universal" for all known phytoplasmas. They can also be observed in sieve tubes by light microscopy. which allows restoring the sanitary status of affected vineyards when the vector activity is controlled. Riesling and also numerous local varieties. but the latter technique has never been successfull on field-grown grapevines. Some cultivars such as Chardonnay. including the agents of FD and BN. ELISA detection is possible from vascular tissue of symptomatic grapevines. vector and abundance of reservoirs). such type of transport is risky when potential insect vector species occur on the site of planting. Polyclonal and monoclonal antisera have been raised to a few phytoplasmas. Random Fragment Length Polymorphism (RFLP) analysis of the amplicon can provide further characterization of the infecting phytoplasma. When the presence of a particular disease is suspected. Full recovery and transient remission of symptoms have been observed in low and medium sensitive cultivars. Propagation material may be the infection source for long distance dissemination of GY diseases. The three ascertained vectors of GY diseases are univoltine leafhopper or planthopper species. Transmission: Phytoplasmas are vectored by hemipters. Phytoplasmas can be detected by graft transmission to sensitive vines. using DAPI staining. and bunches is strong evidence for phytoplasma infection. when grapevine is not a preferred host for the vector. It is probable that the feeding preference of insect vectors. on less conserved genes. mainly hoppers (cicadellids or cixiids) or psyllids. Hence. Varietal susceptibility and sensitivity: Numerous V. are group specific. Other host plants: Most phytoplasmas are ubiquitous plant parasites that can be hosted by several sometimes tolerant plant species. Simultaneous presence of symptoms on leaves. 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.broom group) is a second agent of Australian grapevine yellows. or on random selected non-ribosomal DNA fragments. Pinot noir. When no information on the particular phytoplasma type is available. more specific primers can be used. as well as their feeding activity.

evolution of the disease in space and time.e. The disease is thought to be caused by root damage. Caudwell (b): Description of recovery in grapevines affected with FD. but does not rule out the possible involvement of a pathogen. Nonsymptomatic grapevines that show irregular symptoms from one year to the next may also test positive. In any case. The biology of the insect is described. 1956 1957 1961 1961 1961 1964 1965 137 . Vidano: S. Historical review 1955 Levadoux: Description of a severe outbreak of "Flavescence dorée" on Baco 22A in southwestern France. using detection of PCR amplification products with DNA-DNA hybridization with a specific probe. Real-time PCR has also been successfully used. Gärtel: Description.Other molecular methods are being developed. Production of sanitized propagation material is possible with hot water treatment (HWT). Schvester et al. 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. under the name Flavescence dorée. efforts should address prevention by planting healthy propagation material and limitation of vector populations and their inoculative activity. on the possibility to limit or eradicate their population and prevent their migration or movement in vineyards. FD is transmitted by the leafhopper S. The author suggests this disease to be due to adverse climatic and soil conditions.. BN is considered as a non epidemic form of FD. Detection in the upper parts of the vines is usually done from veins and petioles of the leaves of symptomatic shoots. Pruning or top-grafting are useless because roots and trunks are infected. The disease can be transmitted by grafting and is probably a virus disease. on physiological changes and defence mechanisms induced by infection. Control of GY diseases depends on the knowledge of vector insect species. hypotheses on its nature. Detection of phytoplasma DNA can be achieved from all plant organs or insect vectors. into hot water (50 °C) for a sufficient length of time (minimum 30 mn). Control: There are no direct curing methods of infected plants. phytoplasmas are unevenly distributed in GY-affected vines. Generally. cultural practices. i. 2. FD can be detected from leaves of non-symptomatic American roostocks. Control methods of vector populations are being experimented with natural insecticides. littoralis Ball. of a disease occurring in the vineyards of the Mosel Valley and of the Rhine Valley. Branas (a and b): Description of flavescence epidemics on Baco 22A and Chardonnay in France. Caudwell (a): Studies on Bois noir (BN) and on its relationships with FD. Symptoms (macroscopic and microscopic). littoralis Ball found in Italy in 1963. an insect that has been introduced recently from America. and natural enemies. However. but is also possible using young leaves shortly before symptom expression or phloem scrapings from lignified canes. denoted Flavescence dorée (FD) in agreement with the name proposed by Levadoux (1955) because of the golden yellow metallic aspect of leaves. Investigations are being conducted on sensitivity. and on the identification of reservoirs of inoculum such as infected grapevines. tolerance and potential for recovery of varieties. and on elicitors of defence reactions to these phloem-restricted bacterial agents. weeds or other crops. Caudwell: Characterization of a new type of flavescence. soaking of dormant material before or after grafting.

Bois noir.: Presence of a disease similar to FD in the Oltrepò Pavese (northern Italy). The author assumes that both FD and its vector were introduced from North America with varieties of Vitis labrusca between 1927 and 1950. An important FD-like disease is reported.: S. Credi and Babini: Grapevine yellows are reported in Emilia Romagna (Italy). Symptoms on V. titanus (= littoralis) is not a vector. Rumbos et al. 1967 1968 1970 1971 1971 1973 1973 1975 1977 1979 1980 1982 1982 1983 1984 1985 1985 138 . is probably of European origin. This phytoplasma was later on identified as a Clover phyllody phytoplasma. Caudwell: Discussion on the origins of yellows diseases of plants. Mosel and Rhine regions are considered as the causal pathogens of this disease. (b): Evidence that FD and BN are two different diseases with similar symptoms.: Rickettsia-like organisms observed in roots of grapevines with "Vergilbungskrankheit" in the Saar. Immersing cuttings taken on FD-affected grapevines in water at 30°C prior to planting reduced the proportion of infected plants by about 83%. littoralis. Baggiolini: S. titanus found in 1984 in vineyards of northern Italy. Recommendation that mother plants should be grown in areas far away from FD-infected regions. littoralis Ball is present in Ticino (southern Switzerland). witches' broom and yellowing. these findings have not been confirmed. Caudwell: Symptoms of grapevine yellows (Amarilliamento) reported in Chile on cv Elqui. with special reference to grapevine. So far. Caudwell et al.: Discovery of Mycoplasma or PLT group-like microorganisms in the phloem elements of plants showing dwarfing. Inoculation of grapevine seedlings with S. titanus fed on the latter V. Granata: Description of an epidemic yellows disease on cv Inzolia in Sicily. Potential existence of several different diseases: leafhoppers (Euscelidius sp. Belli et al. the hypothesis is put forward that this nematode is the vector of the disease. Caudwell et al. and Euscelis sp. Tanne and Nitzany: Occurrence of a yellows disease resembling FD in Israel. First record in 197576. littoralis is present in the same region of Oltrepò Pavese.1966 Caudwell: Attempts to inhibit the agent of FD in vivo by heat treatment. Caudwell and Larrue: Spread of FD in France is related to sanitary status of planting material. of which S. (a): Mycoplasma-like organisms (MLOs) are observed in diseased grapevines and infective leafhoppers and in Vicia faba submitted to feeding inoculation with infective S. Belli et al. Conversely. Regina.) caught in the vicinity of vineyards could transmit a yellows disease to Vicia faba plants. faba are different from those obtained with feeding inoculation with infective S. Provisory name "Rhine riesling problem". faba plants produced typical GY symptoms. Magarey and Wachtel: Description of a new disease of grapevine of the yellows type on the cv Rhine Riesling in South Australia. Osler et al.: S.: 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. The disease has been observed for the first time in 1968. littoralis. Doi et al. mainly on cv. Rafaila and Costache: Yellows of grapevine with symptoms similar to those of FD found in 1967 for the first time in Romania. Caudwell et al. As similar organisms have been found in nematodes of the species Xiphinema index.

: Study on the distribution of S. Carraro et al. Quaroni et al. their etiology is unclear. whereas an unsprayed vineyard was more affected. titanus is not always associated to the disease. Recovery and crop loss are variable. respectively). However. Two of these vineyards were sprayed with insecticides in 1986 and 1987. Rui et al. vector of FD.: New data on the spread of FD in three vineyards of Oltrepò pavese (northern Italy) from 1985 to 1987. titanus. of MLOs in phloem tissues of grapevines affected by "FD" in northern Italy. Italy. titanus. Mescalchin et al.: Survey of potential Auchenorryncha vectors of the pathogen agent of FD in Piemonte (Italy).: Observations. Euscelidius variegatus and Euscelis incisus are taken into consideration. Hyalesthes obsoletus.: Presence and distribution of a FD-like disease in the region Friuli-Venezia Giulia in Italy. in viticultural areas of northern Italy and of other Auchenorrhynchas susceptible to play a role in the transmission of yellows diseases in this region. Borgo et al. Martelli: A review on the knowledge on grapevine diseases induced by phloem. titanus. Fortusini et al. vector of FD. Egger and Grasselli: Presence in Toscana. disease distribution. In addition to S. Seljak: Presence of Scaphoideus titanus in western Slovenia (Yugoslavia). Survey for the presence of S. control measures. variegatus and S. Italy. Conti: A review of intracellular procaryotic phytopathogenic agents. Fortusini and Belli: Description of the development of epidemics of FD-like diseases in northern Itay.: Occurrence of FD-like symptoms in the valley of Sarca in Trentino.: Contribution to the knowledge of FD (or similar diseases) in the Veneto region. Pinot bianco. Credi and Callegari: Survey of vineyards in Emilia-Romagna. Susceptibility and sensitivity of affected varieties (Chardonnay. Vidano et al.or xylem-limited prokaryotes in Europe. Italy. Pinot nero) and comparison with other similar diseases. S. Chardonnay is the more affected variety. for the presence of a FD-like disease. Italy. Magarey et al. The results suggest that the disease is brought into the vineyards from outside local sources.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. Borgo: General information on the presence of FD-type diseases in northern Italy. using the scanning electron microscope. Boudon-Padieu and Larrue: ELISA detection of FD pathogen on infected reared experimental and wild natural leafhopper vectors (E.: Observation of MLOs in phloem tissue of yellows-affected grapevines in Australia. that are responsible for severe decline of vines.: Presence of a FD-like disease in Emilia-Romagna. of a FD-like disease on Chardonnay. titanus. northern Italy.: Description in Italy of the presence of FD or FD-like diseases. resulting in a reduced diffusion of the disease. 1987 1987 1987 1988 1988 1988 1988 1988 1988 1989 1989 139 . Vidano et al. Credi: Description of a FD-like disease in Emilia Romagna and the reaction of 3 different cultivars. Credi et al.

Refatti et al.: Occurrence of grapevine yellows in Moldavian SSR. Deng and Hiruki: A method to amplify 16S rRNA gene from culturable and nonculturable mollicutes. 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". Credi et al.: 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. 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). The recommended temperature / time combination is 50°C / 3560 min. Treatment prior to storage is more efficient. This is a prospect for investigation on MLOs associated to plant yellows. Le Grau du Roi (France) of a device for secure soaking of dormant buds and canes into hot water (50°C. Di Terlizzi et al. suggesting an unrelated agent. 45mn). titanus was present in all vineyards checked. Affected grapevines in the Rhône valley tested negative. Granata and Russo: An epidemic disease with FD-type symptoms develops in Sicily on cv. Granata and Grimaldi: MLOs are observed with the electron microscope on affected grapevines of cv. Chardonnay and Baco 22A grafted on donor yellows-diseased plants of cvs Pinot blanc and Sangiovese. Inzolia.: Grapevine graftwood shoots infected with FD can be disinfected by hot water treatment in the dormant stage. S. Boidron and Grenan: Construction and testing in ENTAV. Caudwell et al. Chardonnay develops in Tuscany. (a and b): Identification of numerous species of Auchenorrhynchas in the vineyard ecosystem in Piemonte (northen Italy) and of ampelophilous species. Only part of the plants were infected.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). Credi and Santucci: Development of "Flavescenza dorata" in grapevine in Emilia-Romagna and rate of infestation in different systems of cultivation. The annual rate of symptomatic plants was never greater than 20 % over a 6-year period. 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. Caudwell and Kuszala: Detection with ELISA of FD-MLO in naturally FD-affected grapevines. Caudwell: Review on the epidemiology and characterization of FD and other GY diseases.: Bench grafting experiments of buds of indicator cvs. Symptoms also developed on plants obtained from buds taken on diseased Chardonnay and Sangiovese grafted on healthy Kober 5BB. Similar symptoms on Chardonnay could be obtained by insect inoculation (Euscelis incisus) and by cleft-grafting of tissues taken from yellows-infected elm. 1989 1990 1990 1990 1990 1991 1991 1991 1991 1991 1991 1991 1992 1992 1992 140 . Specific primers can be used for MLOs. Marinesku et al. Inzolia in Sicily. Restriction profiles after digestion with endonucleases permit the differenciation of MLOs associated to different plant diseases. Several weeds with symptoms of phyllody were found to contain MLO in electron microscope studies. The aetiology is not fully ascertained though the presence of MLO is probable. Vidano et al. Conti: A yellows-type disease of cv.

: FD can be transmitted by symptomless rootstocks.: Description of MLOs and associated disorders in the phloem of FD-affected LN33 experimentally inoculated with S.6%) were obtained.1992 Credi and Santucci: Of 628 attempts to transmit the agent of a yellows disease of grapevine from Sangiovese. Attempts to characterize MLO DNA extracted from insects with molecular methods. Typical symptoms on several cvs. (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. One MLO strain transmitted with Macrosteles quadripunctulatus appears similar or close to aster yellows MLO. Davis et al. Arzone et al. MLOs were visible in sections of petioles of periwinkle showing symptoms and subsequent graft transmission from periwinkle to periwinkle was successful. It is more related to the agent of Elm yellows than to other yellows agents. (b): Evidence of an association of MLOs with grapevine yellows in Emilia Romagna. titanus is not present. titanus leafhoppers.: 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 . The titre of MLO appears very low in grapevine. 4 positive results (0. Hot water treatments suppress the transmission to Chardonnay indicators. FD can be readily distinguished from Bois noir. (a and b): Observations supporting the existence of different types of grapevine yellows in northern Italy. Arnò et al. titanus population in vineyards and the rate of spread of the disease.: Transmission experiments from grapevine to grapevine and to herbaceous hosts with S.: ELISA with FD-antibodies permit to detect related antigens in S. (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. suggesting the presence of two different diseases at least in Veneto and Friuli-Venezia Giulia. 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. transmission by feeding to healthy grapevines succeeded only in 1 attempt out of 100. titanus. Daire 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). results of research. Chen et al. IPVR is related to aster yellows MLO. The MLO strains found in grapevine are related with aster yellows MLOs. FD-MLO DNA can be distinguished from other MLO DNA with Dot-blot hybridization and Southern-blotting. However. Boubals (a): Report on a meeting of the French working group on FD. Senescent or degraded forms of MLOs identified inside degenerate sieve elements. genomic tests). detection (ELISA. titanus and other leafhopper species show that yellows disease present in northeastern Italy differs from flavescence dorée as it occurs in France. Bianco et al. but are different from known aster yellows cluster MLO strains. Caveccia and Chardonnay vines to periwinkle by the dodder species Cuscuta campestris. Evolution of the disease and of its vector in the various viticultural regions of France. Boubals (b): A severe epidemic of grapevine yellows occurs in Golan (Israel). Alma et al. Meignoz et al. Bertaccini et al. Caudwell et al. Quacquarelli and Barba: Review of the distribution of flavescence dorée and other grapevine yellows in EEC viticultural countries. No spontaneous recovery. S.

BN belongs to the stolbur group. 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. MLO appear to be in very low titre. Positive assays obtained only on samples from France and Veneto. Only FD and BN have been identified in naturally affected grapevines.: Six-year transmission trials of different types of yellows with S. healthy plants protected with plastic screens did not develop symptoms and diseased plants protected with screens showed transient recovery. So. S. S. FD detected in leaves of rootstock 3309C and is also present in samples from Veneto (Italy). Detection in non symptomatic V. graft and dodder. Electron microscopic observation of MLO in periwinkle and grapevine.called FDU. Maixner (a. Description of pathological changes in phloem of leaves and identification of senescent forms of MLO inside degenerate sieve elements. titanus) and a second one. Transmission has been obtained only from vines of Veneto. Osler et al. titanus to test plants of cvs Perera and Chardonnay in Veneto and Friuli-Venezia Giulia (Italy). at least two different phytoplasmas are infecting grapevines in Veneto: FD sensu stricto (transmitted by S. (a and b): Various attempts to transmit FD-like disease in different regions of northern Italy. Daire: PhD thesis on detection and differentiation with DNA-based methods of the MLO agents of GY diseases in France. FD is related to elm yellows and BN is related to stolbur. Specific probes cloned and specific primers designed for PCR amplification of phytoplasma DNA. A MLO has been transmitted to periwinkle with dodder.: Important outbreak of a yellows disease on local varieties in Apulia (southeastern Italy) and infection of periwinkle as bait plants and with dodder transmission. 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. Girolami and Egger: Importance of contamination of vineyards and diffusion of Grapevine yellows in northern Italy. riparia in New York and failure of detection in some symptomatic grapevines in New-York and Italy. c and d): Grapevine yellows (Vergilbungskrankheit) are common in the Moselle and Rhine valley. Classification of grapevine MLOs into three RFLP groups: elm yellows (FD from France).: FD-ELISA method was used on tissues of yellows-affected grapevines from diverse countries. (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. Carraro et al. probably transmitted by another insect. International Committee of Systematic Bacteriology (ICSB). Computer analysis for spatial patterns of diffusion show a non-random distribution and suggest transmission from other vines or from weeds. titanus is not present. b. Credi (a and b): Observation of MLO in yellows-affected grapevines in northern Italy. Prince et al. Kuszala et al. FD belongs to the elm yellows group but is different from elm phytoplasmas. using insects. 1993 1993 1993 1993 1993 1993 1994 1994 1994 1994 1994 142 .: Diversity of MLOs associated with grapevine yellows and transmitted to periwinkle. 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). Maixner: Demonstration that Hyalesthes obsoletus (Auchenorrhyncha: Cixiidae) is a vector of German grapevine yellows (Vergilbungskrankheit). aster yellows and Xdisease. BN (stolbur MLO) detected in several regions of Italy and in Israel. Di Terlizzi et al. 1993 Daire et al. Detection in grapevines from Virginia (USA) of MLO related to X-disease. 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. on the effects of pollarding affected vines and of chemical sprays against vectors. In northeastern Italy.

Fortusini et al. Italy).: Six-year survey of evolution of yellows symptoms in a 6-12 year-old vineyard of cv Chardonnay that was never sprayed with insecticides. Related MLOs were also detected in wild grapevines. Double infection is reported. PCR detection of phytoplasma with universal primers.: Identification of BN (stolbur phytoplasma) in Spain. Detection is also possible during winter on wood scrapings.: Four different phytoplasmas (FD. Alma et al. 1995 1995 1995 1995 1995 1995 1995 1995 1995 1996 1996 1996 1996 1996 1996 1996 1996 143 . in grapevines with yellows symptoms in Soave (Veneto. Arzone et al. Laviña et al. (a and b): Grapevine yellows in Latium (southwestern Italy) are not associated with S. sometimes in mixed infection. Maixner et al (b): Detection of a new phytoplasma type related to FD in cv Scheurebe showing symptoms of yellows in Palatinate (Germany). Ten weeds were found harboring MLOs with transmission electron microscopy. Italy). The importance of proper identification of disease type for control measures is emphasized. obsoletus and in weeds growing in the vineyard in Germany. Consideration on possible resistance and research on interaction of phytoplasma with nepovirus infection. Murari et al. in the vector H.: MLOs associated to grapevine yellows in Virginia belong to 2 different groups : X-disease group (16SrIII) and aster yellows group (16SrI). Haidar: First report of symptoms of yellows on grapevines in Lebanon.1994 1994 Parente et al. (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. with the aim of identifying sources of tolerance or resistance to yellows diseases. One grapevine out of 16 contained a 16SrV (= elm yellows) phytoplasma together with a 16SrI-G phytoplasma. Prince et al. Padovan et al. The most frequent belonged to group 16SrI-G (later designated as 16SrXII or stolbur group). Boudon-Padieu (a and b): Reviews on knowledge and research on grapevine yellows. Molecular detection of aster yellowsrelated phytoplasma. Padovan et al.: Emphasis on the possible role of weeds as reservoir for MLO in vineyards.: Survey of yellows symptoms in an ampelographic collection of 1281 cultivars in Conegliano (Veneto. Del Serrone and Barba (a and b): Monitoring of detection of phytoplasma in the organs of grapevine according to the vegetative state. Egger et al. stolbur and apple proliferation) can be detected. Koruza: Dispersal of grapevine yellows in Slovenia.: Presence and distribution of GY in Sicily.: Presence of grapevine yellows in northern Portugal and electron microscopic visualization of MLOs in the phloem tissue of petioles of affected plants. (a): Specific detection with PCR of stolbur phytoplasma in grapevines affected with Vergilbungskrankheit. titanus which has not been found.: Grapevines from Piemonte (northern Italy) contained several different phytoplasmas. Albanese et al. probably of Bois noir type. aster yellows. Maixner et al. Del Serrone et al.: The phytoplasma associated with Australian grapevine yellows is close to but different from stolbur phytoplasma causing BN.

: Survey of leahoppers in vineyards of Piemonte. (b): Prospects on detection and identification of grapevine phytoplasmas with new molecular tools and consequence on future epidemiology studies. Belli et al. Phytoplasma belonging to the stolbur subgroup have been identified. Del Serrone : A review in Italian on the knowledge on etiology. No double infection has been found to occur. spread. Severe symptoms with high incidence on seven cultivars are reported from four grapevine-growing regions surveyed for 5 years (1993-1996). Spain and Israel. Subclades are considered to represent the equivalent of distinct species. (a): Description of available methods and strategies for detection of phytoplasmas in grapevine. titanus is not reported. The possibilities of control 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1998 1998 1998 144 . (b): Survey of possible auxiliaries for the biological control of leafhoppers and planthoppers in vineyards. titanus reported for the first time in this region. No epidemic outbreak has been observed. Aster yellows and X-disease types were never detected. biology and control.: Survey of hoppers in vineyards of the province of Piacenza (Emilia Romagna. Italy). Subcommittee on the Taxonomy of Mollicutes: The taxonomy of phytoplasmas will refer to their molecular phylogeny. Presence of S. S.: Use of Hot water treatment (HWT) to eliminate phytoplasmas (FD and AY) from 3 cultivars of grapevine. No positive detection with PCR on the HW-treated vegetative progeny of mature canes of symptomatic plants. vectors and detection of the agents of Grapevine yellows. S. Boudon-Padieu and Maixner: Updating in French and English of present knowledge on grapevine yellows: etiology. Maixner et al. The term phytoplasma will become the genus name of plant pathogenic mycoplasmas under the provisional taxonomic status Candidatus. Garau et al. 10 species were known vectors of phytoplasmas. (a): Only stolbur and FD phytoplasmas were detected in samples of yellows-affected grapevines from numerous regions of France and from Italy. titanus is not present. AY and X-disease related phytoplasmas have been transmitted and detected. Daire et al. International Committee of Systematic Bacteriology (ICSB). Tanne and Orenstein (a and b): Successful heterografting transmission of phytoplasma from symptomatic grapevine to periwinkle and easy subsequent identification of phytoplasma type.: History and control of GY diseases in Italy.: Symptoms of grapevine yellows observed in Hungary in the early 1970's.1996 1997 1997 1997 Rüdel: Review on the history and diversity of Vergilbungskrankheit of grapevine in Germany. northern Italy. Positive reaction with a DNA probe specific to aster yellows phytoplasmas. Aldini et al. transmission. Maixner et al. Among 32 species identified.: Identification of Flavescence dorée in Spain.: Recurrent and severe yellows symptoms observed since the 1980's in Sardinia (Itlay) on several cultivars. Among 29 species of Auchenorrhynchas from the vine canopy and weeds of the border. Murari et al. Kölber et al. 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. 10 are confirmed phytoplasma vectors. Bosco et al. Batlle et al. Borgo: Description and colour photographs of symptoms of grapevine yellows and knowledge on their aetiology and vectors. Davis et al.

have shown that only stolbur group phytoplasma was consistently found. FD-related phytoplasmas (elm yellows group) have been detected in grapevines from Treviso and Verona and transmitted from vine to vine using S. Search for alternative vectors.: A review of the situation of GY in Spain. Probes were antisense to stolbur specific sequences in the 16S rDNA and used on stolbur-infected periwinkle.: Summary of the complex situation of GY in northeastern Italy. though S. Batlle et al. has been identified as a member of the X-disease group. Maixner: A review of thermotherapy to cure phytoplasma infected material. Borgo et al. Sensitivity of assays must be good because of the low titre of phytoplasmas in affected plants. epidemiology and etiology of GYs in the world. vector of FD. Grapevine phytoplasmas classify into different groups. and M.: Evaluation on the efficiency and security of hot water treatment used to eradicate phytoplasmas from propagation material. Reinert W. distribution in the world.: A comprehensive review of the classification of phytoplasmas in the world. titanus is more widely distributed. Insecticide sprays against S. nucleic acid hybridization and serological comparisons. Lherminier et al. In addition a phytoplasma (FDU=GYU) transmitted from GYaffected vines to Catharanthus roseus by means of dodder in Friuli-Venezia Giulia.: Report of positive detection of aster-yellows phytoplasma in grapevine fed with Metcalfa pruinosa and in the body of the insects used for transmission. Seemüller et al. No negative effect was observed on scion buds but rootstock cuttings were more sensitive to negative effects of the treatment. Guadagnini et al. Maixner and Reinert: Updating in German of the research made in different countries on grapevine yellows diseases.: Monitoring grapevine yellows in North-eastern Italy and attempts to rationalize routine laboratory testing for the differentiation between FD and stolbur phytoplasmas. Frausin et al. In the Veneto region only stolbur-group phytoplasma was recorded in some areas and both stolbur and elm yellows group phytoplasmas in other areas.depend on the knowledge of the biology of phytopathogenic agents and of their vectors. Survey with PCR of phytoplasmas infecting grapevines in Friuli-Venezia Giulia and Trento province (Italy) and in Slovenia. Boudon-Padieu: Review on recent progress in the knowledge on grapevine yellows.: First report of the use of oligonucleotides for in situ hybridization with transmission electron microscopy to localize phytoplasma in plant cells. titanus.: Unsatisfactory results on the use of hot water treatments to eliminate phytoplasmas from grapevine wood. Phytoplasmas for which 16S rDNA sequence is available have been classified into 20 major groups and subgroups. GY affect mainly the northern provinces. methods of detection of phytoplasmas in grapevine tissues and in insect vectors. symptoms. Similar attemp with FD phytoplasma were not successful.: Phylogenetic studies of phytoplasmas associated to numerous plant diseases. 1998 Lee et al. 1998 1998 1998 1998 1999 1999 1999 1999 2000 2000 2000 2000 2000 145 . Boudon-Padieu (b): Chapter in a Handbook on viral and bacterial diseases of the grapevine. On the basis of RFLP of PCR-amplified 16S rRNA gene. with a particular reference to the work done in Germany. phytoplasmas can be classified into 14 groups and 38 subgroups.: A history of GYs in north-eastern Italy. are compulsory in France. Firrao et al. BN is very frequent but H. according to the most recent studies on molecular structure of rDNA. Refatti et al. Carraro et al. Only FD and BN have been identified. obsoletus is rarely found in vineyards. FD is restricted to the north of Cataloña. epidemiology of these diseases. titanus. History.

Circulifer sp. although its vector S. Experimental evaluation of the method and application to field-trapped individuals of suspected vector species. titanus is widespread in western vineyards. sugar metabolism.: Survey of GY in Israel. Marzachi et al.: 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. Phytoplasmas seemed to disappear from the insect body after the transfer of insects to healthy plants. Zahavi et al.: Use of hot water treatment in commercial nurseries in Australia to eradicate diverse pests and pathogen agents from propagation material. Clair et al. nitrate and nitrite reductase.: In Golan Heights (Israel) Hyalesthes obsoletus. Tanne et al. BN and aster yelllows in vineyards of southeastern Piemonte (Italy). Primers fitted for specific detection of these 3 phytoplasmas permit cheaper and quicker diagnosis. Orenstein et al.: A 2-year survey in Golan Heights. Israel. Macrosteles quadripunctulatus and Orosius orientalis were found on weeds in vineyards or trapped on sticky traps. The phytoplasma agents are not specified.: Use of immersion in hot water (HWT) or in chemicals to suppress phytoplasmas from grapevine propagation material in Italy. A high rate of recovery is observed from one year to the next. suggesting that they did not multiply in the body of the insects. Influence of rootstock on the rate of GY symptoms in the same vineyard and same grapevine variety. Hyalesthes obsoletus has been found but not in vineyards. Boudon-Padieu: Updating of knowledge and research on grapevine yellows worldwide. Phytoplasmas were detected in all five species. Crocker et al. Osler and Refatti: The situation of GYs in northern Italy.. Quartau et al. 2000 2001 2001 2001 2001 2001 2001 2001 2001 2001 2001 2002 2002 2002 2002 2002 2003 146 .: Recommandations for the organisation of nursery in Australia and handling of hot water treatment to avoid dehydration of cuttings during treatment. HWT (50°C/40 mn) gave acceptable survival but none of the conditions experimented provided a total elimination of phytoplasmas. (a): Presence of FD. 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. Plant variety and cutting diameter influence the rate of surviving cuttings. confirmed the presence of three phytoplasmas associated to GY: stolbur (70%). Seljak: A review of non-European hoppers introduced in Slovenia. The latter was difficult to evaluate because of the non homogenous distribution of phytoplasmas in canes. aster yellows (11 %) and X-disease groups phytoplasmas. Frosini 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.: Development of a new technique with DNA chips to detect phytoplasmas in grapevine.: The presence of phytoplasmas in liquid medium on which insect vectors have been fed can account for potential vectorship of the species.: Report of experiments combining temperature and time of treatment with hot water on 5 grapevine cultivars to cure planting material from phytoplasmas. Neoaliturus sp. Klein et al. Moretti et al. Dual infection may occur (13 %). Seljak and Petrovic: Overview on phytoplasma diseases of grapevine and fruit trees in Slovenia. FD has not been identified. Waite et al. which are all known as vector of phytoplasmas.: Scaphoideus titanus is present in Portugal. Bertaccini et al.

fitted to routine detection.: Compared efficiency. Neoaliturus fenestratus. two nonpreferentially ampelophagous vectors of grapevine phytoplasmas. Other phytoplasmas are reported.: Improvement of detection of FD and BN in field-grown grapevines with real-time PCR. Consequently roguing of affected vines should be avoided when possible and especially if vectors are controlled. titanus found associated to grapevine yellows with a high incidence in vineyards of Rastina (South Serbia). Megophthalmus scabripennis positive for aster yellows phytoplasma. Study of the spatial and temporal dispersion of the four species. Marzachi et al. No important damage has been reported.: Data on 15-year studies on efficiency and effect on propagation material of hot water treatment.: Survey of potential vectors of phytoplasmas in vineyards of the Golan Heights.: Flight activity of Scaphoideus titanus and Hyalesthes obsoletus and their infection status towards FD and BN (stolbur) phytoplasma.: Report of stolbur phytoplasmas in grapevine affected with yellows in Albania. BN (stolbur phytoplasma) has an incidence of about 30 %. Myrta et al.: Measurements of basal respiration rate of grapevine dormant wood as an indicator of the best period for application of hot water treatment.2003 2003 2003 2003 2003 2003 Boudon-Padieu et al. Clair et al. respectively. Kuzmanovic et al.: Development of a sensitive PCR procedure for dual identification of FD and BN phytoplasmas.: 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. Hyalesthes obsoletus and Circulifer haematoceps abundant and positive for stolbur and aster yellows phytoplasmas.: Identification of grapevine yellows symptoms in Tunisia and attempts to identify pathogen agent and vector.: Grapevines affected with yellows are found free of phytoplasmas after recovery. Discussion on benefits and limits of artificial feeding medium to assess the transmission capability of phytoplasma by insects. Apart from FD phytoplasma reported by Duduk et al. Milkus et al. An aster yellows phytoplasma is suspected. Lessio et al. rapidity and sensitivity of methods for use in routine diagnosis of grapevine phytoplasmas. M'hirsi et al. Leitner: The situation of Grapevine yellows in Austria is now surveyed by means of molecular detection assays. D'Ascenzo et al. 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. Orenstein et al. stolbur phytoplasmas are suspected to occur because of long-lasting record of stolbur disease on many host plants in Serbia. Crocker et al. in particular a clover phyllody type (16SrI-C) which is quite frequent. Osler et al. Duduk et al. Chabbouh et al. Tassart-Subirats et al.: Report of aster yellows phytoplasmas in grapevine affected with yellows in Tunisia.: Identification of aster yellows (16SrI-B and C) and of ash yellows (16Sr-VII) related phytoplasmas associated to yellows of grapevine in Chile. (a and b): FD phytoplasma (16S rV-C) and S. 2003 2003 2003 2003 2003 2003 2003 2003 2003 2004 2004 2004 147 . Gajardo et al.. Recovery is a progressive phenonenon developing over 3 years in the case of BN.: Important presence of GY in Abruzzo (central Italy).: Important outbreak of grapevine yellows on cv Chardonnay and identification of a stolbur phytoplasma in diseased plants.

.

Cicadellidae). This leafhopper is a neartic species specialized on Vitis sp. littoralis Ball). it occurs in all southern vine-growing regions. However. Leaves persist longer on affected plants. Eggs are laid in summer on two-year old wood and trunk. transmission is possible from early June (about one month after the beginning of hatching) until the death of adults. If the plants are inoculated after recovery. These isolates could be distinguished by Western-blot analysis using FD antisera and monoclonal antibodies (Mabs). the term "FD sensu stricto" was applied to diseases and phytoplasmas that are transmitted by S. However. Switzerland. In France. Scaphoideus titanus Ball (= S. FD is endemic only in regions where S. Vineyards in these areas are under the threat of an FD outbreak that could occur if FDinfected planting material is introduced. littoralis Ball) (Homoptera. were characterized in Italy and shown to be transmitted also by S.) using S. FD92 and FD2000 were experimentally transmitted to broadbean (Vicia faba L.GRAPEVINE YELLOWS: INDIVIDUAL DISEASES A. Though the rate of transmission by bench grafting can be very low. Transmission: The FD agent is transmitted in the persistent mode by the leafhopper Scaphoideus titanus Ball (= S. Hence. It is widespread in all northern provinces of Italy and an outbreak was recently reported from southern Serbia. FLAVESCENCE DORÉE 1. Isolates F70. then as a virus disease. and Savoie. indicating a vine-to-vine transmission. 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. Two isolates denoted FD-D and FD-C. titanus individuals collected in infected vineyards of south-western France. this way of spreading is significant when infected grafted plants are planted in vineyards hosting the vector. titanus. Feeding transmission starts after a 4-week latency in the body of the vector. Agents: FD was first regarded as a physiological disorder. FD88. rooted cuttings from infected canes of a few rootstock varieties can develop "vinifera-like" symptoms on the wood and leaves. Infected rootstocks are important means of dissemination because they are symptomless. FD92 and FD-D could not be distinguished from one another. Infected rootstocks show little or no symptoms. It was described in a limited area in north-eastern Cataluña (Spain). After the discovery of other GY diseases. Transmission occurs also by vegetative propagation and grafting. isolated infected vines have been identified in northern vineyards of Burgundy and Alsace. First symptoms may 149 . It is also present in regions from which FD has not been reported in France. 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. Crop losses may be very high. In addition. Lignification of the canes is usually incomplete. Description Flavescence dorée (FD) was the first Grapevine yellows (GY) disease to be reported. introduced into Europe at the beginning of the 20th century. 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. Diseased vines have a patchy distribution in the plot. symptoms may be limited to a few shoots. S. Several isolates have been characterized with molecular criteria. 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). extremely sensitive varieties do not recover. 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. and western Slovenia. decline progressively and die. FD88. with an incidence that increases rapidly from one year to the next. Affected vines of most varieties may recover in the second year when they are protected against reinoculation with insecticide treatments. Moreover. Corsica. show a general asymmetric bent posture and necrosis of terminal bud. Molecular comparisons have shown that all these isolates are more closely related among them than with other phytoplasma strains in the same group. that has colonized a wide climatic area. southern Italy. Most of the time. north of Spain and Portugal. titanus. titanus is well established. It is highly epidemic and extremely dangerous because of the biology and ethology of its leafhopper vector. Symptoms develop on leaves and shoots in June and July to become oustanding in August and autumn. symptoms affect the whole plant.

Their rearing is still under development. titanus. titanus in an affected vineyard is an important indication that may lead to suspect a FD infection. sensitive amplification with nested PCR of the DNA fragment FD9. In spite of the possibility of recovery. Control: FD is a quarantine organism in the EC. whereas the third treatment. Planting material must not be collected from mother vines in areas or vineyards affected by the disease. such as real-time PCR or identification of PCR-amplified product with DNA-DNA hybridization using specific probes. are currently under development. has become the preferred detection method because it is easy to perform and was improved in reliability and sensitivity. Material from mother plants that have developed symptoms in the following growing season. The second treatment. Vitis riparia can be infected but shows little symptoms. Grenache. Symptoms are rare in Syrah. Sequence or RFLP analysis of the FD9 DNA fragment permit to readily differentiate all the known FD isolates. which is specific for 16SrV-group phytoplasmas allowed the reliable detection of FD phytoplasma in grapevine. The low rate of transmission by grafting and the long delay in symptom expression make the detection of infected vines in planting material difficult. Italy and Spain are susceptible with various degrees of sensitivity. 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. and the risk of disease spreading important. Planting of HW treated material is especially important in areas where the vector is present. titanus in New York State (USA). 150 . Bois noir (BN) is also frequent in regions inhabited by this leafhopper. its area of origin. appear more tolerant. Roguing is compulsory in France. Ugni blanc (Trebbiano) and Prosecco are sensitive varieties that may recover when they are protected from new inoculations. Other DNA-based methods. aims at killing newly hatched insects. Sauvignon blanc. An indirect DAS-ELISA using coating of wells with rabbit polyclonal antisera and detection with a cocktail of several Mabs. such as Merlot. at the beginning of August. Nevertheless. Molecular detection by PCR of selected phytoplasma DNA fragments. Other varieties. although heavily infected vines can be observed. Indirect control is obtained by insecticide treatments against S. Polyclonal antisera and Mabs have been raised to FD phytoplasma. Detection with laboratory methods is possible on insect vectors and infected vines. has been used as the official method for large scale survey of FD in France from 1993 to 2003. Under these circumstances only chemical insecticides are efficient for eradication of the disease. Detection: FD symptoms can be confused with those of other GYs and also with other diseases or disorders. Recently. Simultaneous amplification in a multiplex procedure of both FD9 and Stol11. Cabernet Sauvignon. Two additional treatments are applied during summer. a DNA fragment specific to BN (stolbur) phytoplasma can be used for the survey and monitoring of these two GY diseases. 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. control measures are compulsory according to legal regulations. must be destroyed. roguing of diseased vines is advisable when the epidemic pressure is high. Antigen extraction from vascular tissue of symptomatic grapevines requires the use of high Tris molarity and strong detergents in the extraction buffer. have been found carrying FD phytoplasma until recently.appear on young vines as late as four years after grafting. Natural insecticides can be used to preventively limit vector populations but their efficiency is limited in epidemic outbreaks. Standard conditions (50°C for 45 min) proved highly efficient for sanitizing infected rootstocks and grapevine cultivars. has allowed the identification of potential auxiliaries for biological control. The presence of S. Monitoring natural enemies of S. Other host plants: No host plants other than Vitis sp. Soaking of dormant material in hot water (HW) is recommended. Chardonnay. In France and Italy. at the beginning of July. Varietal susceptibility and sensitivity: All varieties grown in the various FD-affected regions in France. Nielluccio and Garganega are very sensitive varieties that usually do not recover after infection. In 2003 a phytoplasma resembling the FD-C isolate was found in wild Clematis in Veneto (Italy). The first treatment is applied 30 days after first instar emergence at the beginning of the period of potential transmission. The same is true for American rootstocks which can therefore be dangerous sources of infection and dissemination of the disease. Alicante Bouschet. is directed against winged adults migrating from nearby vineyards or wild vines.

First report that abandoned or wild vines may be a source of infection. Bonfils and Schvester: Relationships of leafhoppers with FD of grapevine in western France.: Demonstration that FD is transmitted by the leafhopper S. Vidano: S. (a). littoralis Ball. 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. Caudwell: Study of the phenomenon of recovery of FD-affected vines.2. littoralis recorded from in Italy in 1963. Caudwell et al. Schvester et al. Baggiolini et al. an insect recently introduced from North America. Vidano: Study of the ecology and biology of S. Caudwell et al. Use of 4-5 treatments with DDT or parathion and removal or destruction of pruning wood older that two years. Schvester et al. Branas (a and b): The new disease is thought to be caused by root damage. Study of the role in the aetiology of FD of mycoplasmalike organisms (MLO) observed in insect vectors and in inoculated grapevine and V. Caudwell: PhD thesis on FD. littoralis on plants other than the grapevine and transmission trials of FD to other host plants. biology and feeding damage and of the symptoms of FD in France on Baco 22A. littoralis in its area of origin in North America. Description of the insect. littoralis in Corsican vineyards with winter treatment of dormant eggs with oleoparathion. Boubals and Caudwell: FD epidemics observed in Corsica. Schvester et al. S. The aetiology is unknown. Caudwell and Poitou: Study of the interaction between fanleaf and FD. littoralis is present. Schvester et al. littoralis from Ticino (southern Switzerland). (b): Biology of S. Spontaneous recovery occurs after a 1-2 year crisis period. Description of macroscopic and microscopic symptoms. 1960 1960 1961 1961 1962 1962 1963 1964 1964 1966 1966 1967 1968 1969 1970 1971 1971 1972 151 . Caudwell: Attempts to inhibit the agent of FD in vivo by heat treatment. Caudwell et al. Description and study of recovery phenomenon and of localised symptoms. considered as a virus disease. No positive results of transmission trials with 15 species of Hemiptera (mainly Cicadellidae) other than S.: Successful transmission of FD to herbaceous hosts by increasing the feeding access period of the vector S. (a): Possibility to limit the populations of S. The disease can be transmitted by grafting and has probably a viral aetiology. S. littoralis. Historical review 1955 1956 1957 Levadoux: Description of an outbreak of "Flavescence dorée" on Baco 22A in southwestern France.: Studies on the survival of S. Carle and Moutous: Study on possible secondary or alternative vectors of FD or Bois noir (BN).: First report of S. (a): Control of FD by insecticide treatments against the vector. littoralis.: Field tests for insecticide control of S. Caudwell et al. evolution of the disease in space and time. littoralis. faba plants. Recovered vines may be infected again but the symptoms are lighter. littoralis. littoralis. 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.

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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

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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.

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(a and b): Evolution of FD in the Treviso province (Veneto. titanus in Italy. titanus in Switzerland. titanus laid in the bark were killed. Kuszala: Specific ELISA procedures to detect FD or BN phytoplasmas in extracts from diseased grapevines from the field.: RFLP studies of 16SrV phytoplasmas. nymphs and adults of S. Batlle et al. Posenato et al. Clerc et al. Several different peptides from membrane proteins are identified by Western-blot. Infection is rapidly spreading to the east. Pavan et al. Rousseau: A review of different ways to reduce the populations of S. 1996 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1999 1999 2000 2000 155 . Bosco et al.: First report of S.: First report of FD outbreak in northern Cataluña (Spain). FD has not been identified. (a and b): Situation of FD and S. Italy). the eggs of S. titanus in north-eastern Italy where vines are trained in the pergola system. The biological significance of this finding is unknown.: Monoclonal antibodies raised to immunoaffinity purified FD phytoplasmas. The possible role of six species in the transmission of GY is discussed. titanus in Trentino (Italy).: A phytoplasma of the 16SrI (aster yellows) group is detected in eggs. Caudwell et al. titanus reared on healthy plants. The amplified FD9 fragment is specific and variable among 16SrV (elm yellows) group phytoplasmas.: Development of specific PCR primers for the detection of FD or BN phytoplasmas. especially Prosecco in the period 1993-1996. Serological relationships with elm yellows phytoplasma is confirmed Alma et al. Spain and Italy. Three different RFLP patterns at least are shown in FD isolates from France.: Monitoring of leafhoppers in vineyards in Italy. In the same conditions. Daire et al. Vindimian et al. FD is not present in southern Italy. The possibility that direct inoculations have occurred in nursery is discussed. Increase and spread to other cultivars. Jermini and Baillod: Combination of visual observation of instar larvae and trapping of adults on sticky traps to monitor the populations of S. (a and b): Demonstration of curing effect of hot water treatment on FD-affected plant material.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. titanus in the canton of Geneva (Switzerland). Control recommendations. Bianco et al.: Verification of hot water treatment conditions to suppress FD phytoplasma infection in dormant wood of grapevine.: Insecticide control of S. First outbreak in the 1980's on cv Perera. according to the severity of the FD epidemics.: Important outbreak of FD in Lombardia (northern Italy). Lherminier and Boudon-Padieu: Use of transmission electron microscopy with in situ immunolabeling to detect FD phytoplasma in grapevine and herbaceous host plants. Seddas et al. Mori et al.: In spite of the presence and rapid spread of S. Transmission by grafting is studied. Numerous species identified. titanus in biological viticulture. Belli et al. Survey of hemipters in the vineyard and presence of other known phytoplasma vector species. Marcone et al. Martini et al.: Demonstration that two isolates of FD sensu stricto (FD-D and FD-C) are spreading in Veneto (Italy).

2000 2000

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).

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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.

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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.

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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

158

DNA fragments. Chemical sprays against H.: Rickettsia-like organisms found in roots of VK-affected grapevines in the Saar.: 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. no direct control is possible. 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. roguing should be avoided. while others seem more tolerant and do not show symptoms. littoralis. Leclant and Lacote: Study of the ability to transmit stolbur to herbaceous plants by H. Findings never confirmed. Leclant: Presence of Hyalesthes obsoletus a cixiid planthopper in the south of France. BN is considered as a non epidemic form of FD. unknown at that time. history. Nienhaus et al. For direct differentiation between FD and BN/VK phytoplasmas. Mendgen: 1971. Description of symptoms of VK. Cazelles et al. for the presence of a FD-like disease. Hence. cytological and electron microscope study of tissues of affected grapevine. The disease occurs in the Moselle valley and the Rhine valley. Rumbos et al. Control: As with other GY diseases. Though pruning does not cure infected vines. obsoletus in the south of France. and different susceptibility and sensitivity of grapevine cultivars. titanus. 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. Credi and Callegari: Survey of vineyards in Emilia-Romagna. Caudwell et al. The results suggest that the disease is brought into the vineyards from outside local sources. The species has been described as the vector of stolbur "virus" by Suchov in USSR in 1948. (b): Epidemiological observations of BN lead to the assumption that an aerial vector is involved. because of the complex biological cycle and multiple host plants of the insect species.: Rickettsia-like organisms from VK-affected grapevines are cultured in chick embryos. Superficial soil cultivation in summer or winter ploughing are methods intended to damage the larval instars and reduce vector populations. Italy. Rumbos et al. the multiplex nested-PCR assay cited in the FD chapter can be used. Historical review 1961 1965 1968 1969 1971 1971 Caudwell: Study on BN and its relationships to FD. Borgo: General information on the presence of FD-type diseases in northern Italy. Some grapevines show symptoms repeatedly in successive years. 1972 1977 1978 1978 1988 1989 1992 159 . 2. except for declining vines. Gärtel: Description of VK under the name of Flavescence dorée. Caudwell et al. S. Natural enemies are being investigated.: GY symptoms are present in Switzerland (Suisse romande and Ticino) but they are not associated with S. (b): BN is distinct from FD based on non transmissibility by S. These findings have never been confirmed. titanus is not always associated with the disease. absence of recovery. obsoletus are not efficient. Recovery and remission of symptoms for several consecutive years are frequent in vines affected by BN. Description of structures that were probably closteroviruses. suppression of affected branches may improve harvest quality and help in progressive recovery.

Computer analysis for spatial patterns of spread shows a non-random distribution and suggests preferential transmission from other vines or from weeds.1992 Fos et al. PCR-RFLP analysis confirms relationship to aster yellows phytoplasmas. obsoletus is confirmed as a vector of stolbur disease plants in France.: PCR-RFLP analysis of MLO-DNA detected in VK-affected grapevines in Germany show that this MLO is related to stolbur. (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. clustered by the authors into aster yellows group but shown later to belong to the stolbur group. Daire et al. Reservoirs and possible role of other vectors remain to be elucidated. Minucci et al. Bertaccini et al. Several weeds are host plants of the stolbur phytoplasma. Borgo: Presence of BN in mixed infections with FD in the province of Treviso (Veneto. Davis et al. obsoletus is a vector of VK in Germany.: One of the two phytoplasmas detected in yellows-diseased grapevines in Liguria is related to IPVR phytoplasma. Italy. and from Israel. The latter MLO was shown later to belong to the stolbur group. Bianco et al. Alma et al. Davis and Prince: According to a different terminology. c and d) Transmission of a yellows to periwinkle with dodder in Germany.: A monoclonal antibody raised to stolbur mycoplasma-like organism is used for ELISA detection of the agent in herbaceous host plants and insects. Boudon-Padieu (b): History. H. (b): Presence of phytoplasmas in the group 16SrI. Italy) where FD is prevalent.: The agent of a GY disease in the Italian Riviera is detected with a probe specific for Eastern aster yellows. Maixner : Demonstration that H. Laviña et al. 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. Bianco et al. the most frequent belonging to group 16SrI. MLO associated to GY in Italy are related to aster yellows MLO. Maixner (b. Maixner et al. BN MLO is detected in grapevines from France. from several regions of Italy. The MLO strains found in grapevine are related with aster yellows MLOs. (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).: Grapevines from Piemonte (northern Italy) contain several different phytoplasmas.: First detection of BN (stolbur phytoplasma) in Spain. (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. Maixner et al. Biology of the insect. 1993 1993 1993 1993 1993 1993 1993 1994 1994 1994 1995 1995 1995 1996 1996 1996 1996 160 . Bertaccini et al. It is suggested that vectors with preference for other host plants act as vectors for VK. Brescia and Pavia (northern Italy). (b): Evidence of an association of MLOs with grapevine yellows in EmiliaRomagna. obsoletus. obsoletus identified as the vector. H. (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. subgroup I-G (later named group 16SrXII (stolbur) in the provinces of Vicenza. but are different from known strains of MLOs of the aster yellows cluster. subgroup I-G (later renamed as 16SrXII or stolbur group). aetiology and transmission of BN in France.

search for vectors. biology of H. 1996 1996 1996 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1998 1998 1998 1998 161 . conversely. Vindimian: LN but not FD is present in Trentino (northern Italy) in spite of the presence and active spread of S. Weber: PhD thesis on the biology of the cixiid H.: Monitoring of field populations of H. Maixner: The situation of VK in Germany: symptoms. obsoletus and its role as the vector of VK. Sforza: PhD thesis on the epidemiology of BN in France. Estimated crop damage ranges from 20 to 40 %. Maixner and Reinert (a): Trials on elimination of VK infection from vine wood in Germany. Sforza and Boudon-Padieu: Description of H. 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. Kölber et al.: Prevalence of BN (stolbur phytoplasma) in Navarra and Cataluña (north eastern Spain). First detection of BN agent in diseased grapevines in Switzerland with the same procedure.1996 1996 Koruza: Survey of GY in three regions of Slovenia shows that the disease is of BN type. Weber et al.: Consistent association of stolbur phytoplasma with GY diseases in the Italian provinces of Fiuli-Venezia Giulia and Trento and in Slovenia. Marcone et al. Davis et al. subgroup I-G (identified later as belonging to the stolbur group or 16SrXII). arvensis is recommended to expose instar larvae and kill them with frost.: Important spread of GY symptoms on several grapevine varieties in Umbria (central Italy). Refatti et al. a high proportion of previously infected vines that did not show symptoms for 2 years at least and. Albanese et al. Characterization of a phytoplasma belonging to group 16SrI. a high proportion of vines in which symptoms re-occurred after one season or more. obsoletus as a vector of BN. Osler et al. Kuszala: Detection in ELISA using a monoclonal antibody to stolbur phytoplasma of BNassociated antigens in diseased grapevines in France. Laviña et al. obsoletus in Germany. Results were satisfactory.: Phytoplasmas associated to GY in Campania (southern Italy) are genetically uniform and belong to the stolbur group. Maixner and Reinert: A review of the research and present knowledge on VK and other GY diseases. vector transmission and possible control measures.: Occurrence of GY diseases in Hungary and identification of stolbur phytoplasma in affected grapevines. Ploughing in winter of soils infested by C. a strong annual increase. Search for variability among BN phytoplasmas with RFLP analysis of the specific non ribosomal DNA was unsuccessful. obsoletus and possible control measures. Preferential spread along the rows. titanus. (a and b): BN is poorly transmitted by bench-grafting. using hot water treatment with conditions recommended in France. (c): Identification of stolbur phytoplasma (16SrXII) as agents of GYs in Greece and Israel. as well as phytoplasmas transmitted by Credi to periwinkle in Emila Romagna and previously classified in the aster yellows group. Daire et al. The maximum delay of symptom expression in young plants is 2 years. (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.

H. Marcone et al.: Only stolbur phytoplasma detected in grapevines showing GY symptoms. Females are more often infected than males. Phytoplasmas in the 16SrXII group (stolbur) were the most frequent. Varga et al. 1998 1998 1998 1999 1999 2000 2000 2000 2000 2000 2000 2000 2000 2000 2000 162 . Confirmation of the role of H. Larrue et al.: Several vineyards affected by BN in Burgundy (France) monitored for over 15 years and assessed for symptom severity. The study confirms a high rate of infected H. First launching of colonies of the insect under controlled conditions.: Widespread presence of LN in Toscana (central Italy). Šeruga et al.: First detection of stolbur phytoplasma in grapevines in Croatia. Identification of main reservoir weeds. Braccini et al. Maixner et al.: Epidemiology of BN in France. No other grapevine phytoplasma was detected.1998 1998 Sforza and Bourgoin: Anatomy of reproductory organs and mating of H. Škoric et al. Up to 34% of insects were infected with stolbur phytoplasma. Sforza et al. different phytoplasmas were identified in a number of cultivars. obsoletus. PCR detection is reliable when testing together 25 insects among which only one is infected. Identification of two infected symptomless weeds in the vineyard. Weber and Maixner (b): Etholology of H. Evidence secured for the occurrence of tolerance (no symptom at all for 15 years) and temporary remission (no symptoms for a few years) . the vector of Palatinate grapevine yellows (PGY). Darimont and Maixner: Comparison of the transmission efficiency of phytoplasma to their host plants and to grapevine by H. the ability to recover of cvs Riesling and Pinot in spite of systemic symptoms. obsoletus and Oncopsis alni. Transmission to natural host plants is higher than for grapevine with both insects. Braccini and Pavan: Report of survey of Auchenorrynchas in vineyards of Toscana (central Italy).: Use of PCR to confirm the presence of stolbur phytoplasma in GY-affected grapevines in Switzerland. obsoletus can be high because of the presence of reservoir weeds in the vineyard. obsoletus and Metcalfa pruinosa were found positive for BN phytoplasma (16SrXII-A). obsoletus as a vector to grapevine. obsoletus and possibility to control the insect in vineyards. Acquisition may occur at larval stages since emerging 5th instars were infective. The rate of infected H. Maixner and Reinert: Collection of H. obsoletus in vineyards is efficient with sticky traps placed at the soil level. the second GY disease in Germany. Seljak and Petrovic: A review of the Slovenian situation of GY diseases. The same pathogen detected in weeds and other crops. formerly reported in western Europe. Bourquin et al. and a high proportion of vines that escaped infection during the 3-5 year period of monitoring.: BN reported only in north-western and eastern vineyards of Croatia. regardless of the cultivar in Campania (southern Italy). Up to 30 % of the insects were infected and the highest rate was in vineyards with high populations of Convolvulus arvensis.: Ethological onservations and morphological descriptions of adults and larvae of H. among which hoary cress. obsoletus. Sforza et al. obsoletus. High rate of BN / LN infection in the different vine-growing regions. obsoletus related to VK infection. Weber and Maixner (a): Procedures for the survey of infection status of populations of H. No other phytoplasma found in the South of the country.: In a large survey of vineyards in 8 Hungarian counties conducted in 1998.: VK disease dramatically increased in incidence in the Moselle and Rhine valleys during the 1990's.

Laviña et al. Kölber et al. Detection in December was the more constant. The authors conclude that phytoplasma infection induces non specific.: First detection in Albania of BN phytoplasma in grapevines of different age and varieties.: Survey of GY from 1997 to 2002 on 21 cultivars in 12 Hungarian counties.000 plants over 12 years.: Anoder cixiid planthopper. obsoletus and the presence in organic vineyards of several weeds known as host plants carrying larvae on the roots. Alma et al. every month for one year except in winter.: Demonstration that H. Indicators based on climatic parameters permit to predict the flight period. 2001 2001 2001 2002 2002 2002 2003 2003 2003 2003 2003 2003 2003 2003 2003 2003 163 .: Incidence of BN in a small area of the Bekaa valley (Lebanon) on cvs Chardonnay and Alicante Bouschet. the incidence of VK was significantly higher in conventional vineyards. In spite of a similar abundance of H. Role of stinging nettle (U. Samples were taken on all organs of 10 affected grapevines. Curkovic Perica et al. general stress responses and rapid senescence. which is widespread in the province.2001 Bertamini and Nedunchezhian: Evaluation of effects of BN disease (stolbur phytoplasma) in field-grown grapevines. BN is the main disease and cvs Chardonnay and Zweigelt appear to be the more sensitive among whiteand red-berried varieties. obsoletus. chlorophyll fluorescence and photosynthetic activities suggest that Photo System II is affected. reached by other authors. is a natural vector of stolbur phytoplasma to sugarbeet in Bourgogne (France). while Megophthalmus scabripennis was positive for aster yellows phytoplasma. respectively. Merlot and Pinot noir.: Observation of the occurrence of BN in Trentino in eight Chardonnay vineyards on more than 11. Study of the spatial and temporal dispersion of the four species.: Assessment of the best period for detection of BN in affected grapevines. Bertaccini et al. Orenstein et al. Pentastiridius sp. Langer et al. H. obsoletus.: The situation of GY diseases in Croatia shows that only BN is present in the northern and eastern vineyards. Severity of symptoms vary from one year to the other. Myrta et al.: Only BN identified in Switzerland on cvs Chardonnay.: Comparison of infection risk by VK in organic and conventional vineyards in Germany.: The highest risk of phytoplasma contamination for grapevines coincides with the flight activity of adults of H. All affected grapevines and two naturally infected periwinkles contained the same stolbur isolate. Gatineau et al. The presence of H. obsoletus and Circulifer haematoceps were widespread and positive for stolbur or aster yellows phytoplasmas. Mescalchin and Mattedi: A description of the development of VK in Trentino (northern Italy). Choueiri et al. Neoaliturus fenestratus. Darimont and Maixner: Importance of the presence and infectivity of H.: Importance of outbreaks of BN in the province of Modena (Italy) and monitoring of potential insect vectors in addition to H. Morandell: Detection of VK in south Austria. Gugerli et al. on pigments. obsoletus to the German viticulture.: Survey of potential vectors of phytoplasmas in vineyards of the Golan Heights. Cicotti et al. obsoletus is the vector of LN in Italy. Maixner et al. very similar to European isolates. Same conclusions about tolerance and transient recovery of grapevines. obsoletus in Lebanon is recorded. 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. dioica) as a reservoir of stolbur phytoplasma and as a host for the insect.

: Important expression of BN in cv Chardonnay in the Ukraine. 2003 2003 2004 2004 2004 C. dioica. 164 . However. and rarely in groups. Several hemipters have been found to deliver stolbur phytoplasma to the medium. Another alder insect. obsoletus were positive for stolbur phytoplasma. affected plants are not numerous. Šeruga et al. the psyllid Psylla alni. Transmission to plants is not reported. Cicadellidae) trapped on alders and caged on V.: Important epidemics of "Schwarzholzkrankheit" (SHK = VK) in Franken (Bayern. Trapping of insects on sticky traps or with D-Vac suction. H. The disease was identified in 1995. PALATINATE GRAPEVINE YELLOWS 1. PGY phytoplasma is similar to the Alder yellows phytoplasma that is widespread in black alder in Germany. obsoletus and frequence of C. Other host plants: Alder trees (Alnus glutinosa L.: Importance of a GY epidemics in the Drava region (Slovenia) in 2001 and 2002. Attempts to transmit PGY phytoplasma from grapevine to grapevine using reared S. Control: No possibility of control because of erratic transmission by the vector. (b): First identification of BN infection in the Macedonian republic. obsoletus and weeds in different areas in Germany. Germany). obsoletus is rare. alni is highly monophagous and specimen do not survive long on grapevine. Langer and Maixner: Three different types of stolbur phytoplasma isolates found in grapevines.) Detection: Universal primers and RFLP analysis of amplicons can be used to identify a 16SrV phytoplasma. occur most often at the border of plots. Transmission: Transmission was obtained with the leafhopper Oncopsis alni Schrank (Hemiptera. It is different from FD phytoplasma. Living insects checked for transmissibility of phytoplasma by feeding on artificial medium. (a): Investigation on the variability of stolbur (16SrXII) phytoplasmas from extracts of different grapevines in Croatia. It is not epidemic and occurs most often in old vines in vineyards along creeks or rivers where alders (Alnus glutinosa L) are present. Description Palatinate grapevine yellows (PGY) is a disease occurring in Germany in Pfalz and Moselle regions (west of Germany) mainly on cv Scheurebe. Main synonyms: FD-Pfalz was a first name because the agent was shown to be related to the FD phytoplasma. vinifera seedlings that later developed GY symptoms. specific association of each isolate with a different weed is discussed. Alder is considered as the source of PGY infection. 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). Šeruga et al. was found carrying the alder phytoplasma at a higher rate than O.2003 2003 Petrovic et al. Agents: The agent is an EY (16SrV) group phytoplasma.: Search for vectors of stolbur phytoplasma in Cataluña (Spain) where H. arvensis and U. Main symptoms: Symptoms are similar to those of other GY diseases and may affect the whole plant. titanus leafhoppers have failed. Sabaté et al. Milkus et al. mainly on cv Scheurebe. Varietal susceptibility and sensitivity: PGY occurs in limited areas. The latter results have not been published. alni but failed to transmit both to alder and grapevine. -B and C) have been identified. Gilge et al. Symptoms were recorded on 26% of 30-year old Scheurebe grapevines and 96% of tested individuals of H. using 4 fragments of phytoplasma DNA showed that all isolates are similar. According to the data of field survey. O. It is associated to phytoplasmas in the Elm yellows (16SrV) group. 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. together with high populations of H. Three isolates (PGY-A.

which was associated with two different phytoplasmas in group 16SrIII and 16SrI (aster yellows). The strong adaptation of O. respectively. In 1991. affected vines often do not survive winter frost because they lack reserves. alder. As much as 5 % of leafhoppers from Palatinate and 15 % from Moselle tested positive.: Further comparison between elm yellows group (16SrV) phytoplasmas. the latter findings were not confirmed. (b): Detection of a new EY-related phytoplasma in GY-affected cv Scheurebe in Germany. die back of shoot tips and abortion of developing fruit. molecular characterisation and epidemiology of PGY. alni. The risk of infection by PGY is lower than for VK because reservoir plants are fewer and outside vineyards. (b): Transmission to V. a phytoplasma isolate transmitted from alder to periwinkle in Italy. In 1993. alni (Auchenorrhyncha. Maixner et al. In New York. Virginia grapevine yellows III (VGYIII) Main symptoms: North American Grapevine Yellows (NAGY) is a lethal disease causing yellowing of the leaves. Darimont and Maixner: Comparison of the transmission efficiency of phytoplasma to their host plants and to grapevine by O. 165 . Cicadellidae) (but not P. obsoletus. Angelini et al. American grapevine yellows. The PGY strains are closest to the ALY phytoplasma from alder in Italy and to one strain of FD. Maixner and Reinert: Demonstration that O. both insect species trapped on alder. using Heteroduplex mobility assay and sequencing of rDNA and FD9 fragment. It is different from FD phytoplasma but molecular and serological data show a close relationship. alni trapped on alders. Main diseases: Virginia grapevine yellows I (VGYI). a phytoplasma belonging to the X-disease group (16SrIII) was detected in affected grapevines in New York. The differences in vector specificity between PGY and FD phytoplasmas are not supported by molecular data. Main synonyms: Leaf curl and berry shrivel (LCBS). Characterization of the three PGY strains showed that they were closer to FD than to strain EY1 from American elm. Extensive survey of yellows-affected grapevines in several areas of Germany did not show the presence of other EY group phytoplasmas. Reinert: PhD thesis on detection. alni) is able to transmit the alder yellows phytoplasma to healthy seedlings of alder. the vector of BN / VK. 4 Italian and French strains of FD and elm and alder phytoplasmas. populations of S. vinifera seedlings of the PGY phytoplasma using specimen of O. Reinert and Maixner: All the 3 strains of PGY were detected both in alder trees and in specimen of O. Variability analysis and sequencing of the FD9 DNA fragment and homology of the sequence to a part of the SecY gene. alni. Angelini et al. Historical review 1995 Maixner et al. The alder phytoplasma resembles ALY.: Comparison between Elm yellows group (16SrV) phytoplasmas including PGY strains. However. Reinert and Maixner: 1997. alni to alder prevents a frequent inoculation to grapevine even if the infestation of the leafhopper population is high. 1997 1999 1999 2000 2000 2000 2001 2003 D. Description Grapevine yellows (GY) were first reported in New York State (USA) in 1977. Infected grapevines often die within months from the onset of symptoms and significant losses of vines have been observed. a serious outbreak of GY occurred in Virginia. In the same period. Oncopsis alni and Psylla alni specimen.2. alni and H. titanus living mostly on wild Vitis riparia around affected vineyards were found to carry an agent that was serologically related to that of FD. The proportion of infective leafhoppers is lower with O. Same RFLP profiles of P1-P7 rDNA amplicons obtained from yellows affected grapevines. Identification of the PGY-associated phytoplasma with phytoplasmas infecting alder in the vicinity of affected vineyards. NORTH AMERICAN GRAPEVINE YELLOWS 1.

titanus in New York and transmission assays of a possible infectious agent. Sauvignon blanc and Cabernet franc. The X-disease phytoplasma occurring in New York was also detected with monoclonal antibodies and with probes for dot-blot hybridization. However. 2. allowed the identification of a few candidate vectors of the aster yellows phytoplasma. Symptoms may occur for several years in succession in the same vines. the disease was first observed on cv De Chaunac. it is very severe on Chardonnay vines but was observed on other varieties including Riesling. among which Agallia constricta. vinifera cuttings.: Molecular comparisons between MLOs transmitted to periwinkle from grapevine or detected in a few naturally affected grapevines. observed in 1983. Detection was positive in symptomless wild Vitis riparia growing near vineyards in New York and in some symptomatic grapevines. No transmission by vine planting material reported. riparia to V. Chen et al. Maixner et al.: Occurrence of FD-like symptoms on White Riesling grapevines in New York. Control: There are no control methods since vectors and reservoirs of the associated phytoplasmas are still undetermined. titanus. However. In Virginia. Data showed that the Italian and American MLO were related. present in New York and its possible relationship with a GY disease. the survey of vineyards and transmission assays to V. X-disease phytoplasma was detected in native Vitis sp.: Two MLOs were transmitted from affected grapevines to periwinkles in Udine (Italy) (= FDI or FDU or GYU) and New York. and an aster yellows phytoplasma was detected in numerous weeds and shrubs within and around the vineyards. Probes and primers yielded positive reactions using DNA extracted from grapevine. Larvae are more frequent on wild Vitis riparia at the border of the vineyards. Varietal susceptibility and sensitivity: In New York. subgroup I-A). Transmission: No vector insects have been identified for VGY phytoplasmas. (a): A Riesling plant from New York and a periwinkle carrying the FDI MLO described above. Daire et al. as well as direct PCR assays from insects. Symptoms. Maixner and Pearson: First data on the study on S.: Monitoring of S. 1985 1991 1993 1993 1993 1993 166 . Detection: With molecular assays using universal or group-specific PCR primers.Agents: The 16SrIII phytoplasma detected in NY has not been further characterized. The FDI MLO was identified in a parallel work as a member of the X-disease group. subgroup III-B) and with the VGYI phytoplasma (a strain of aster yellows or 16SrI. other grapevines with strong symptoms tested negative. Pearson et al. tested positive for a MLO in the X-disease group with PCR-RFLP of 16S rDNA. X-disease phytoplasma was detected in native Vitis sp. ELISA and immunfluorescence were positive from periwinkle. vinifera. 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. show that the FDU (= FDI) strain from Italy and a MLO detected in a grapevine in Virginia (USA) are both related to X-disease. faba bean plants and feeding solutions. The Virginia Grapevine Yellows (VGY) has been associated with the VGYIII phytoplasma (a strain of X-disease or 16SrIII. very similar to those of LCBS. It appeared to be spreading and was perhaps insect borne. The latter observation suggests that another non-related MLO might have been present. Other host plants: Both phytoplasmas are widespread in the USA in weeds and cultivated species. Adults migrate from V. Historical review 1977 Uyemoto et al.: 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). then on White Riesling. and aster yellows was detected from numerous weeds and shrubs within and around the vineyard. Prince et al.

a GY disease found in Victoria (Australia). 167 . It has been suggested that all three phytoplasmas are strains of the same phytoplasma species. In this particular disease. Mapping of diseased plants show that grapevines appear to be a terminal host of AGY. Main diseases: AGY (sensu stricto). which is the more frequent and the Tomato big bud (TBB) phytoplasma.: Two phytoplasmas are detected in yellows diseased grapevines in Virginia. The sequence similarity of the 16S rRNA gene shows that it is close to members of the aster yellows group (16SrI). AUSTRALIAN GRAPEVINE YELLOWS 1. BVGY phytoplasma has not been clustered in an established phytoplasma group. Agallia constricta is a suspected vector of VGYI (aster yellows phytoplasma). is a member of the aster yellows (16SrI) group.: A GY disease observed in Virginia since 1983 affects large areas with a low incidence and a low spread rate. Other species tested positive and transmitted to feeding solutions. consistently associated with AGY symptoms and AGY phytoplasmas but their aetiology is not clear. 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. The use of PCR has shown the association with AGY of three different phytoplasmas. It belongs to the peanut witches' broom group (16SrII) and has a broad plant host range in Australia. Agents : AGY disease (sensu stricto) is associated with two phytoplasmas: the AGY phytoplasma. The yellow leaf tissue can become necrotic. Early and recent surveys have shown that AGY are found in most viticultural regions of Australia. Late Season Leaf Curl (LSLC)-affected grapevines show symptoms in late summer with tightly downward rolled leaves that often remain green. Davis et al. also detected in wild V. riparia vines. 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. Shoots may display abortion of bunches or berry shrivel later in the season and death of terminal buds. 1998 2003 E. Some plants also display uneven bud development resulting in canes and cordons that are bare in places with little or no bunch development. leaves tend to fall early. Both VGYI and VGYIII were found in several weeds and shrubs around and inside the vineyards. there are reported cases where they were not associated with AGY disease or phytoplasmas. Main symptoms: All symptoms typical for GY can be found on AGY-diseased plants. Description Grapevine yellows were first reported from Australia in 1976 and called Australian grapevine yellows (AGY) in 1983.1993 Wolf et al.5%) with the phytoplasmas associated to Papaya die back (PDB) in Australia and to Phormium yellow leaf (PYL) in New Zealand. one node after the other. The 16S rRNA gene has a high sequence similarity (more than 99. The AGY phytoplasma has been designated as Candidatus species "Candidatus Phytoplasma australiense". Late Season Leaf Curl (LSLC). In addition. Restricted Growth (RG)-affected grapevines show retarded growth resulting in shortened shoots and smaller leaves. Conversely. III-I. The relatedness of the associated MLO to X-disease MLO is confirmed. Confirmation that a X-disease (16SrIII) group phytoplasma is involved and classification of the latter phytoplasma into a new subgroup. It belongs to the stolbur group (16SrXII) but is distinct from the BN and VK phytoplasmas. clover phyllody phytoplasma (16SrI-C) being the closest . Stems of affected shoots develop a blue. followed by the progressive death of the shoots. subgroup I-A. Restricted Growth (RG). The TBB phytoplasma has been designated as Candidatus species "Candidatus Phytoplasma australasia". Beanland and Wolf: Search for possible insect vectors and reservoirs of NAGY phytoplasmas in Virginia. waxy appearance and remain rubbery. respectively. The second phytoplasma. overlay one another like shingles and become leathery and brittle. suggesting that the source is from an alternative host. Buckland Valley Grapevine Yellows (BVGY).

suggesting that this contributes to the decline in phytoplasma numbers and to the remission of symptoms. Sanitation with hot water treatment is being developed in Australia. The AGY phytoplasma was detected in the common leafhopper Orosius argentatus (Evans) but transmission to grapevine has not been reported. Cixiidae) is known as the vector of PYL disease in New Zealand but the species has not been reported from Australia. The incidence of disease may be temporarily high in some vineyards of Chardonnay. a specific primer in the intergenic 16S-23S spacer region has been designed for detection of TBB phytoplasma (Ca. However. Magarey et al. All generic "universal" primers may be used. Hence.: Orosius argentatus. Specific primers have been designed on the 16S rRNA gene for detection of AGY phytoplasma (Ca. but phytoplasmas were also detected in other red. once vines are infected by AGY. AGY phytoplasmas are common in several crops in Australia. Detection can be from shoots. Varietal susceptibility and sensitivity: Chardonnay and Riesling appear to be most often affected. argentatus. Phloem fluorescence of diseased vines in the UV microscope. Control: There are no methods to control AGY diseases in the vineyard. Osmelak et al. RG and LSLC diseases has been observed. Magarey: The situation of GY in Australia. Magarey and Wachtel: GY symptoms are found in most viticultural areas of Australia. Recent studies have shown that the latter phytoplasma can be acquired from infected grapevine by O.: MLOs. followed by RFLP analysis or sequencing. Magarey and Wachtel: Review of the AGY problem. branches. 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. suggesting that the disease is the result of an aerial transmission and that the phytoplasma was not present in the original planting material. Magarey: Comprehensive review of GY diseases in the world. and trunks in October is more reliable. Detection: Detection is obtained with PCR. vector of TBB in tomato crops. A "heat curing" effect of AGY disease by hot weather has been observed. The TBB phytoplasma is transmitted to other crops by O. argentatus and subsequently transmitted to faba bean but the ability to transmit to grapevine was not confirmed. 168 . Magarey et al.: The "Rhine Riesling problem" denoted "Australian Grapevine Yellows". branches. Reduction of symptom expression in vines injected with tetracycline. However. Remission of AGY. Sampling simultaneously from shoots.Transmission: No vectors have been identified for any AGY disease. The planthopper Oliarus atkinsoni Myers (Hemiptera. Phytoplasma australiense) either in single-step or nested PCR following a first amplification with universal primers. they are at greater risk of displaying symptoms again in the following years. found in sieve tubes of diseased vines are associated with AGY symptoms. Several observations suggest that aerial transmission prevails. 140-510 nm in diameter. 2. Other host plants: As mentioned above.and white-berried varieties. is found in high numbers together with two other potential MLO vectors in vineyards of Victoria (Australia). trunks and roots throughout the year and infection is persistent from year to year. It is possible that AGY diseases have become prevalent in Australia in the last years with the increasing planting of Chardonnay. RG or LSLC. Similarly. Phytoplasma australasia). The BVGY phytoplasma has no suspected insect vector. such as Shiraz or Semillon. reservoirs must be important but the epidemiology is not fully understood. 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. indicating persistence of the disease.

LSLC would be followed by AGY on the following year and AGY would be followed by RG in subsequent years.: Study of the phylogenetic relationships of the BVGY phytoplasma with aster yellows and stolbur phytoplasmas.1995 1995 1996 Bonfiglioli et al. (a and b): Positive detection of phytoplasma with PCR in AGY-affected grapevines and tentative survey of phytoplasmas in Australia.: Use of hot water treatment in commercial nurseries of Australia. Wilson et al. later called BVGY phytoplasma. and LSLC.: The AGY phytoplasma is consistently present in the majority of symptomatic grapevines.: Detection of phytoplasmas associated with AGY. The TBB phytoplasma is detected occasionnally in vines with AGY symptoms and also in LSLC-affected vines. Spain and Israel. insecticide sprays are useless for controlling the disease. RG and LSLC are associated. 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. Bonfiglioli et al.: AGY phytoplasma detected in the body of Orosius argentatus leafhoppers. RG. Habili et al. (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). Constable et al. established from a different source of plant material indicate that BVGY disease occurs as a result of aerial transmission. Constable et al.: Identification of a new phytoplasma in Chardonnay grapevines in Victoria (Australia). Papaya dieback and Phormium yellow leaf diseases. Constable and Symons: AGY phytoplasma is detected in most symptomatic grapevines but also TBB phytoplasma and a third uncharacterised phytoplasma. Constable: PhD thesis on the biology and epidemiology of AGY phytoplasmas.: BVGY phytoplasma is associated to GY symptoms that may show remission then reoccurrence but also appear on vines that were not affected previously. Candidatus Phytoplasma australiense. Beanland et al. Padovan et al. Liefting et al.: Further comparison of AGY phytoplasma with overseas GY phytoplasmas show that it is closely related to.: No vector for AGY found.: Comparison of visual assessment and diagnostic assays. Waite et al. Beanland et al.: Close relationships between phytoplasmas associated with AGY. Hence. The problem of symptomlessly infected vines.: TBB phytoplasmas can be acquired from diseased grapevine by Orosius argentatus and subsequently transmitted to faba bean. Detection of BVGY in another plot in the same area. BVGY might form a new AY subgroup of AY (16Sr) group or even a new phytoplasma group. but different from the stolbur phytoplasma associated with BN in France. Uneven distribution of phytoplasmas in infected plants. resulting in increasing cumulative incidence. Spring and summer are optimal for detection. Padovan et al.: Assumption from field observations and monitoring that AGY. Kelly et al.: Description of AGY symptoms. It is occasionnally detected in symptomless vines and in vines with LSLC or RG symptoms. Gibb et al. Davis et al. Constable et al. Constable et al.

In Chardonnay.: Survey of AGY. X-disease group phytoplasmas. are very frequent in many host plant species and several insect species are known as vectors or potential vectors in different countries. a variety widely grown in all countries and very sensitive to all GY agents. recurrence in other vines and new occurrence in previously unaffected vines. 2. These aster yellows phytoplasmas belong to subgroups I-B or I-C (clover phyllody) which are ubiquitous in Europe. Saric et al. Statistical analysis showed that RG and LSLC diseases were not always associated with AGY. It seems that aster yellows phytoplasmas have no important significance in European grapevine yellows. TBB phytoplasma may have a role in the aetiology of LSLC. The first is the detection in countries other than the USA. However. At other times of the year. namely Israel. 2004 2004 F. 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. The total cumulative incidence for AGY could be as high as 90%. they seem to have some relevance in Israel and were recently reported from Tunisia. RG and LSLC symptoms for 3 to 6 years in Chardonnay and Shiraz vineyards. three different situations must be described. a 16SrI-B phytoplasma was found in mixed infection with a stolbur phytoplasma (at that time designated 16SrI-G). No variability was observed amongst BVGY phytoplasma isolates. Constable et al. of phytoplasmas belonging to the aster yellows group (16SrI) in yellows-affected grapevines. sometimes as a second phytoplasma in co-infection with FD or BN phytoplasmas and sometimes also in symptomless grapevines. Other varieties are cited in the different situations. the diseases are characterized by remission in some plants. detection is more reliable from samples taken from branches and trunks. 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. There is no information of transmission of AGY by planting material. 1997 170 .2002 2003 Crocker et al. The AGY and the TBB phytoplasmas may persistently affect grapevines throughout the year and between years. (a and b): Biology and epidemiology of AGYs. to a lesser extent. OTHER GRAPEVINE YELLOWS 1. In three instances.: The management of the area for plant material in the nursery with the scope of hot water treatment. Varietal susceptibility and sensitivity: Chardonnay.: Survey of phytoplasmas infecting grapevine in northern Italy. later designated 16SrXII. which was the first designation for stolbur phytoplasma in some laboratories. Symptomatic shoots from AGY affected vines are reliable for detection in summer (January). The second situation is the significant occurrence of X-disease group (16SrIII) phytoplasmas in countries other than the USA. Constable and Symons: Study with Heteroduplex mobility assay (HMA) of the elongation factor Tu (Tuf) gene of the AGY and the BVGY phytoplasmas. is often reported in connection with these cases.: Detection of phytoplasma in the aster yellows group in GY affected plants in Slovenia and Croatia. Transmission: Aster yellows and. Description Besides the cases reported above. Constable et al. Historical review Europe 1996 Alma et al. The third situation is represented by poorly characterized GYs recorded from new countries. However. The subgroup of aster yellows was not determined but primers specific for stolbur tested negative. They must not be confused with the 16SrI-G phytoplasma of earlier reports.

2001 2003 Tunisia 2003 Chabbouh et al. Orosius orientalis.: Identification in yellows-affected vines of varieties Petit Syrah. titanus. Similar transmission of clover phyllody (16SrI-C) MLO by S. 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. called "Amarilliamento de Elqui". 1971b). Gajardo et al. titanus had been obtained in 1971 in France (cf.: Further survey of GY vectors in the Golan Heights. Macrosteles sexnotatus.: 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. Tanne et al. subgroups I-B and I-C and also to group 16SrVIII (Ash yellows) in mixed infection with I-B. H.: Survey over several years of potential vectors of phytoplasmas in vineyards of the Golan Heights showed that Neoaliturus fenestratus. Stolbur phytoplasma (16SrXII) was also detected. Caudwell et al. H. a X-disease phytoplasma was detected in some Neoaliturus specimen.: Formal identification of a 16SrI-B phytoplasma in GY-affected grapevines in Tunisia. obsoletus was found in addition to the preceding 5 species. 2003 South America 1980 Caudwell: Description of a disease of the yellows type in Chile. including S. Trapping of hemipters and identification of numerous potential phytoplasma vectors.2001 Alma 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). Detection with PCR show erratic identification of a 16SrI-B phytoplasma. Merlot and Carmenere. In addition to similarities of symptoms with FD.: Experimental transmission of the Chrysanthemum yellows phytoplasma (16SrI-B) to grapevine by four leafhopper species. of phytoplasmas belonging to group 16SrI. Klein et al.: Monitoring of potential phytoplasma vectors in vineyards in Israel and molecular detection of phytoplasmas in their body. Megophthalmus scabripennis was positive for aster yellows phytoplasma. Transmission to periwinkle of yellows diseases using collected insects. and Anaceratagalia laevis were all found carrying an aster yellows phytoplasma.. The spatial and temporal dispersion of the four species was analysed. Circulifer spp. M'hirsi et al. Orenstein et al. some affected vines showed a sudden fall of leaves in summer. 2003 171 . In addition. D'Ascenzo et al. Neoaliturus sp.

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