Options Méditerranéennes Série B n.55 Directory of Infectious Diseases of Grapevines and Viroses and Virus-like Diseases of the Grapevine: Bibliographic Report 1998-2004

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Martelli.P. Boudon-Padieu 2006 . E.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.

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

TABLE OF CONTENTENTS PREFACE INTRODUCTION INFECTIOUS AGENTS OF GRAPEVINES INFECTIOUS DEGENERATION (GRAPEVINE FANLEAF VIRUS) INFECTIOUS DEGENERATION (EUROPEAN AND MEDITERRANEAN NEPOVIRUSES) GRAPEVINE DEGENERATION AND DECLINE (AMERICAN NEPOVIRUSES) GRAPEVINE LEAFROLL RUGOSE WOOD COMPLEX GRAFT INCOMPATIBILITY FLECK COMPLEX MINOR VIRUS DISEASES VIRUS-LIKE DISEASES VIROIDS GRAPEVINE YELLOWS: GENERAL PROPERTIES GRAPEVINE YELLOWS: INDIVIDUAL DISEASES GRAPEVINE YELLOWS: REFERENCES PIERCE'S DISEASE THE VIROSES AND VIRUS-LIKE DISEASES OF THE GRAPEVINE: BIBLIOGRAPHIC REPORT .1998-2004 INTRODUCTION SUBJECT INDEX REFERENCES 7 11 15 17 33 49 59 76 93 99 107 117 127 133 149 173 195 203 205 207 219 5 .

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

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

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

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

Université de Bourgogne Dijon. Food and Agriculture Organization of the United Nations. CSIRO Publishing. Influence des viroses et qualité.P. 263 pp.137 pp. Ridé. bactéries et phytoplasmes de la Vigne. 1997. Chairman of the Board of Directors and Secretary General. Martelli. Walter B. Sélection sanitaire de la vigne: sélection sanitaire et sélection pomologique. and G. Martelli G.P. Taylor. Detection and Diagnosis of Graft-transmissible Diseases of Grapevines.K. 1996. University of Bari.P. Paris. Bulletin de l'OIV 70. 5-23. Scott. Sanitary selection of the grapevine. Bulletin de l'OIV 69. Walter B. 2ere partie: Sélection sanitaire. Bovey and G. Walter. M.P. Martelli and E. Notwithstanding this wealth of published information a "Directory of Major Virus and Virus-like Diseases of Grapevines" was compiled in 1992 by R.P. FAO/IBPGR Technical Guidelines for the Safe Movement of Grapevine Germplasm.). of the Centre International des Hautes Etudes Agronomiques Méditerranéennes. Walter B. Giovanni P. Paris. 1993. 2000. the Steering Committee of ICGV decided to update the Directory and to entrust this task to G. (ed.. Lacirignola.). Hamze and Mr. and G. H. Martelli and published under the auspices of the Mediterranean Fruit Crop Improvement Council (MFCIC). Influence des viroses et qualité. M. B. 225 pp. Director of the Mediterrenean Agronomic Institute of Bari for supporting the publication of this work. Martelli Professor of Plant Virology. Sélection sanitaire de la vigne: sélection sanitaire et sélection pomologique. 54 pp. Virus certification of grapevines.. INRA Editions. C. Collingwood. Elisabeth Boudon-Padieu Directeur de Recherches Head Biologie et Ecologie des Phytoplasmes Plante Microbe Environnement INRA . 1991.P. Italy.A. At the 14th ICVG Meeting. Paul. 192 pp. R. 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. Protocols for detection of viruses and virus-like diseases: Les Colloques. Hervieu. respectively. and R. Maladies à Virus. Krake L. Hadidi. Rome /International Board for Plant Genetic Resources. Walter B. Bari. St. 261-276.CNRS . (ed). Rezaian and R. and to Dr. eds. 945-971. a body now estinguished. In: Plant Virus Disease Control (A. sélection pomologique. Boudon-Padieu. Editions Féret. They express their deep gratitude to Prof. N. 1998.A.R. Boudon-Padieu and M.H. Martelli. Khetarpal. Rome. Ikin. Graft-transmissible Diseases of Grapevines. Bordeaux. E. 1ere partie: Effets des viroses sur la culture de la vigne et ses produits. and B. 1999. 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. Martelli G.Frison E. Koganezawa. American Phytopathological Society Press. 1997. n° 86. Rome. France 13 . FAO Publication Division.S.

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

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

INFECTIOUS DEGENERATION .

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

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

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

Dichloropropane-dichloropropene (DD) at 1000 l/ha gave satisfactory results. Reproduction of fanleaf symptoms in mechanically inoculated grapevine seedlings is confirmed Galzy: Heat treatment of grapevine plantlets grown aseptically in vitro. Goheen and Hewitt: Description of vein banding as a GFLV-induced disease Dias: Host range and properties of fanleaf and yellow mosaic viruses. Serological relationship with ArMV reported.: Transmission of GFLV by Xiphinema italiae in Israel. index. Description of the chipbudding method for indexing. Control of the vector by soil fumigation. Bercks: Research on the use of three serological methods for detecting plant viruses. Boubals and Dalmasso: Experiments on soil disinfection against X.: Description of the Davis method of heat therapy of grapevines. and no reinfestation by X. Gerola et al. amaranticolor is confirmed. Vuittenez: New observations on the effects of soil fumigants on fanleaf in contaminated soils. Discussion on the possible role of weeds in the epidemiology of the disease. Dias and Harrison: Relationships between the viruses causing fanleaf. Das and Raski: Studies on the relationships of GFLV with its vector X.b Martelli and Hewitt: Comparative studies showed that Californian and Italian GFLV isolates are the same. Gifford and Hewitt: Use of heat therapy and in vitro shoot tip culture to eliminate fanleaf virus from infected grapevines.: Fanleaf virus is transmitted by the nematode Xiphinema index Cadman et al. Brückbauer and Rüdel: The virus (or viruses) of Reisigkrankheit (GFLV and/or other nepoviruses) are seed transmitted in some herbaceous indicator plants. latex test and barium sulfate test.: Investigations on grapevine virus diseases in California. 1965 1967 1968 1968 1969 1969 1970 22 . Transmission of fanleaf virus by X. whereas insecticide treatment has no effect. index. Bercks and Querfurth: Use of latex-test for detecting GFLV and other nepoviruses in grapevine tissue extracts in Germany. Yield was increased by 400 % in comparison with that of untreated controls. yellow mosaic and ArMV 1960a 1960b 1961 1961 1962 1962 1963 1963 1963a. Cohn et al.: Transmission of fanleaf virus from grapevine to herbaceous hosts by mechanical inoculation and preliminary characterization of the virus. Hewitt et al. Reproduction of fanleaf symptoms in mechanically inoculated grapevine seedlings 1964 1964 1965 Taylor and Hewitt: Description and characterization of Australian isolates of GFLV. 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. Graniti and Russo: A light microscope and cytochemical study of endocellular cordons. Goheen et al. Vuittenez: Mechanical transmission of fanleaf virus to Chenopodium quinoa and C.: Detection of GFLV particles in thin-sectioned grapevine root tissues. index in France. index occurred during the 6 year period of observation. Hewitt et al.1958 1958 1960 Vuittenez: Fumigation of fanleaf-contaminated soil with nematicides prevents infection of healthy grapevines replanted immediately.

1970 1970 Hewitt et al. but ELISA is more sensitive.: 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. George for detecting GFLV. George. Raski et al. quinoa. rupestris is more accurate than mechanical transmission to C.3 dichloropropene or methyl bromide. Uyemoto et al. A dormant bud of the variety to be cured is grafted onto a healthy potted rootstock. Bercks: Serological detection of grapevine viruses in West Germany.: Control of fanleaf by soil fumigation with 1.3-dichloropropene or methyl bromide. index. rupestris St. The acquisition time threshold is less than 5 minutes. Mission or LN 33. 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. Dias: Review paper on grapevine yellow mosaic. The sensitivity of the latex test is increased. index. this lining is shed and ingested in the intestine.: Use of green grafting as a quick and secure method for graft-indexing. Hévin et al. 1970 1970 1970 1971 1971 1972 1972 1973 1973 1973a 1973b 1974 1974 1975 1976 1976 1977 1979 1979 23 . St. Vuittenez: Review paper on grapevine fanleaf.: Detailed physico-chemical characterization of GFLV. 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. Alfaro and Goheen: The different strains of fanleaf virus (fanleaf sensu stricto. Querfurth and Paul: Protein A-coated latex-linked antiserum (PALLAS) method for detecting GFLV and other viruses.: Heat therapy of grape plantlets grown in vitro causes changes in some characteristics of the variety. yellow mosaic and vein banding) are transmitted in the same way by X. Van Velsen and Niejalke: Green budding for indexing grapevine with the indicator cvs. Both tests are more sensitive than mechanical inoculation to C. Vineyards replanted in contaminated but treated soils remained healthy for at least 5 years.: Comparison between PALLAS latex test and ELISA for detecting GFLV in France.: GFLV and marbrure (fleck) are not transmitted through the seeds of grapevine. Raski and Schmitt: Progress in the control of the fanleaf-nematode complex by soil disinfection with 1. especially with low titre antisera. Mur et al. PALLAS is quicker and cheaper than ELISA. After bud take. Walter et al. anterior oesophagus and oesophageal bulb of their nematode vectors. Indexing by budding on V. quinoa. the plant is placed in a heat cabinet for treatment Hévin et al. Both methods give satisfactory and similar results. During the moult of the nematode. Goheen and Luhn: New method of heat therapy. Raski et al.: Comparison of indexing by mechanical inoculation to Chenopodium quinoa and by graft-transmission to V. Taylor: Review paper on grapevine vein banding.: GFLV particles observed in the lumen of the oesophagus of X.

: Study on the effectiveness of soil fumigation for the control of X.1979 1980 1980 Kalasian et al. Efficiency of both methods is compared.3-dichloropropene failed to eradicate either nematodes or fanleaf virus from the soil but reduced the incidence of the disease to acceptable levels. Monette: Heat therapy of GFLV and ArMV--infected grapevines with alternating temperatures. Rüdel: Discussion on the possible role of X. Transmission trials gave a few positive results.: Experiments with systemic nematicides for controlling X. Bouquet: M. vuittenezi might be a vector of GFLV is therefore uncertain. Walker et al.: Soil fumigation with 1. vuittenezi population used for the experiments could not be entirely ruled out. Raski et al. A Middle Eastern V. although it is not resistant to the virus itself. 1980 1980 1980 1981 1981 1981 1981 1983a 1983b 1983 1983 1983 1985 1985 1986 1986 24 . 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. Vuittenez: Review on serological methods of detection and identification of grapevine viruses. vinifera accession represent an excellent example of host plant resistance to GFLV.3-dichloropropene (1. index. index and fanleaf in grapevines. Morris-Krsinich et al. These are promising sources of germplasm for obtaining resistant rootstocks. That X. index.: Detection of fanleaf virus and other sap-transmissible viruses in grapevine tissues by ISEM. a very common species in vineyards of Rheinhessen and Palatinate. Lear et al. Hafez et al. Raski et al. index.3-D (1400 l/ha) and 50-75 cm deep with 165 cm spacing for methyl bromide (448 kg/ha) gave a good control of X.: GFLV particles are arrayed in long parallel rows in thin-sectioned mesophyll cells of infected grapevines.: Comparison of polyclonal and monoclonal antibodies for detecting fanleaf virus with ELISA in various grapevine tissues. Methyl bromide and 1. Russo et al. the possibility that a few X. index feeding. index by ISEM Bovey et al. as vector of GFLV. Forty days of treatment. index larvae were present in the X. Bouquet: Serological detection of GFLV in its vector X. Even in the cases where the virus was transmitted. Bouquet: Muscadinia rotundifolia becomes infected by GFLV when the virus is transmitted by grafting but resists infection when transmission is by X. RNA-2 contains the cistron coding for the viral coat protein. vuittenezi. Brown and Roberts: Detection of fanleaf virus in its vector X. Savino et al. rotundifolia is resistant to fanleaf virus transmission by X.: Identification of several Vitis species and interspecific hybrids resistant to fanleaf virus. index by ELISA.: Use of systemic nematicides for the control of X. Krake and Woodham: Possibility that the agent of yellow speckle is involved together with GFLV in the etiology of vein banding. especially in wood shavings of dormant canes during winter. index. Carbon disulfide gave less satisfactory results. in California.3-D) or methyl bromide applied 75-90 cm deep with 90 cm spacing for 1.: Identification of a natural serological variant of GFLV from Tunisia Huss et al.: 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. The use of methyl bromide requires a continuous cover with polyethylene sheeting for some time after the treatment.: Detection of fanleaf virus in grapevine tissues by ELISA and ISEM in different periods of the year.

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

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

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

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

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

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

39

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

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

Phytopathologia Mediterranea 22. 217-222.Proceedings 7th Meeting of ICVG. Ferreira A.: A comparative study of Bulgarian GBLV isolates and the Portuguese virus CM112 establishes that CM112 is a serologically close but distinguishable strain of GBLV. 1972. Gallitelli et al. Phytopathology 71.: Improvement of ELISA protocol for GBLV detection the whole year round. 135-141.1979 Martelli et al. De Sequeira and Vasconcelos-Costa: Use of an immunoradiometric assay for the titration of the Portuguese strain of GBLV. 1981. A manually transmissible latent virus of the grapevine from Bulgaria. A. Abracheva.: Detection of GBLV in grapevine leaf extracts by ISEM. Annals of Applied Biology 85. Monografias INIA 18 . Annales de Phytopathologie. 1981. 95-101. Some properties of the new latent virus from grapevine rootstocks in Yugoslavia.Sur l'isolement d'un virus à partir de cultures de tissus de vigne. Numéro hors série. 269-272. Savino and A. Proceedings 6th Meeting of ICVG. Quacquarelli.. Martelli G. Krastanova S. CMI/AAB Descriptions of Plant Viruses. 1985. Gallitelli. V. Proceeding 4th Meeting of ICVG.. Garcia de Orta Estacao Agronomica 12. Quacquarelli and D. 1980. V. Vasconcelos-Costa. Grapevine Bulgarian latent virus. Occurence of grapevine Bulgarian latent virus in Hungary. Savino and O. Niagara Falls 1980. Ferreira.: Ultrastructural study of GBLV infections in grapevine and C.P.. Rasteniev'dni Nauki 29..P. Numéro hors série.C. The Portuguese strain supports the replication of a satellite RNA. P.. Jankulova.A. A. References De Mendonça A. Proceedings 7th Meeting of ICVG. Gallitelli. Colmar 1970. Acta Phytopathologica Academiae Scientiarum Hungaricae 16. Krastanova et al. 468-472. No. Pocsai E. 113-120. Possibilities for the whole-year ELISA detection of viruses infecting grapevines. D. De Sequeira. D. 245-250. Proceeding 4th Meeting of ICVG. Martelli G. Quacquarelli. Properties of a distinctive strain of Grapevine Bulgarian latent virus..A. Savino and A. Dimitrijevic: GBLV found in Yugoslavia De Mendonça et al. 1992. 51-58. V.P. O. Pocsai: Occurence of GBLV in Hungary. De Sequeira and A. Niagara Falls 1980. De Sequeira.N. M. and J. Di Franco.. 1977. O. 1980.. 1979. Some properties of grapevine Bulgarian latent virus. 1980 1980 1980 1980 1981 1981 1982 1983 1985 1992 3. 1970.A. Pereira and V.P. and O. 1972. Yankulova. Preliminary studies on an undescribed grapevine virus. Savino. Ganeva and M. Varennes and De Sequeira: First application of ELISA for the detection of virus CM112. 42 . and R. Ramsdell and Stace-Smith: The New York isolate of GBLV is a strain of BLMoV. D.: Detection of virus CM112 in grapevine leaf extracts by ISEM. Ramsdell D. M. Mota. Cordoba 1976. 349-354. Stace-Smith. Annales de Phytopathologie. De Sequeira O.. 1983. Niagara Falls 1980. 186 Martelli G.A. Physical and chemical properties of the particles and ribonucleic acid of blueberry leaf mottle virus.A. The ultrastructure of grapevine Bulgarina latent virus infections in natural and artificial hosts. M. 21-24. Gallitelli D. 1980. Colmar. Russo et al. Martelli G. Simoes. 1978. 143-145 De Mendonça A. Gallitelli.A. De Sequeira.: 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. An immunoradiometric assay for the titration of a Portugese strain of grapevine Bulgarian latent virus (GBLV). 27-32. Proceedings 7th Meeting of ICVG. A preliminary report. P. Applicability of immunosorbent electron microscopy (ISEM) for the detection and identification of CM112 virus in grapevine. quinoa. Martelli et al. France. Abracheva. Dimitrijevic B.A.. Russo and V. A.

Martelli et al. Martelli and Quacquarelli : Physico-chemical characterization of HCMV and comparison with TBRV. GRAPEVINE CHROME MOSAIC VIRUS (GCMV) 1. Isolation and identification of a strain of grapevine Bulgarian latent virus in Concord grapevines in New York State. a symptom virtually indistiguishable from GFLV-induced yellow mosaic. RNA-1 has Mol. Croatia and Austria. Detection of CM122 latent grapevine virus by enzyme-linked immunosorbent assay (ELISA). Jakó et al: HCMV affects pigment and sugar content of infected grapevine leaves.: GCMV recorded from Slovakia and report of X. Varennes A. Some virus strains induce leaf deformity. 269-277. Plant Disease Reporter 61. vuittenezi is very frequently found in vineyards with chrome mosaic patches. The virus is not serologically related with GFLV. 1977. E.Uyemoto J. index. Hummer.F.. vuittenezi transmits GCMV or GFLV. 1982. Martelli: Purification and serology of the virus isolated from Hungarian grapevines with fanleafand yellow mosaic-like symptoms. 2. early reports that this nematode could transmit the virus have not been confirmed. The virus belongs in the same subgroup of TBRV (subgroup B) to which is distantly related serologically. index. Martelli and Quacquarelli: Description of HCMV in the CMI/AAB Descripitons of Plant Viruses series. Saric and Hranuelli: GCMV recorded from Croatia. Although GCMV particles have been detected by immunosorbent electron microscopy in Xiphinema index fed on infected hosts. sometimes together with X. Taschenberg and D. and O. Pozsár et al. No evidence that X. double nodes and short internodes. Kenten: GCMV is distantly serologically related to Cacao necrosis virus. symptomless infection may occur. Tobacco pants and the rootstock 110R have been successfully transformed with the viral coat protein for induction of resistance.K. Virus re-named Grapevine chrome mosaic virus. The virus appears to be unrelated serologically to GFLV and is not transmitted by X. Historical review 1966 Martelli et al. a size of 4441 nt and accounts for 31% of the particle weight. transmitted to herbaceous hosts from Hungarian grapevines with symptoms similar to those of fanleaf and yellow mosaic. It has been recorded also from Czechoslovakia.8 x 106 . 949-953.6 x 106 . wt of 1.: Host range and properties of a spherical virus. The genome is bipartite. wt of 2. pretty much like GFLV. near Lake Balaton. Mali et al. a size of 7212 nt and accounts for 40% of the particle weight. Martelli and Sarospataki: X.: The isometric virus associated with Hungarian chrome mosaic is serologically distantly related to Tomato black ring virus (TBRV). However.: HCMV adversely affects photosynthetical carbon dioxide fixation. The coat protein has a single type of subunits of Mr 52 x 103. Evaluation of short reaction times and re-use of a-globulin and conjugate.A. called Hungarian chrome mosaic virus. De Sequeira. Agronomia Lusitana 41.K. Leaves of infected vines are partially or entirely bright yellow or whitish. 1966 1968 1968 1969 1969 1971 1972a 1972b 1972 1975 1977 43 . Lehoczky and Tasnady: A study of the effect of HCMV on yield and sugar content of infected grapevines. index as vector of the virus (unconfirmed results). and was originally called Hungarian chrome mosaic virus. Description Grapevine chrome mosaic virus (GCMV) was first found in Hungary. GCMV is transmitted through grapevine seeds. Affected vines lack in vigour and may decline and die. RNA-2 has Mol.

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

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

5. Martelli and V.). GDefV is readily detected by ELISA and PCR using primers designed on the coat protein sequence. 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. G. Sabanadzovic. Journal of Plant Pathology 85. 2002. 2003. showing leaf and cane deformations Cigsar et al. Mostar 1977.. Virus Genes 30. D. including all those known to infect grapevine. Grapevine deformation virus. Investigations on grapevine viruses in Croatia.J.: Description and thorough characterization of GDefV. Digiaro. 1997. Abou Ghanem-Sabanadzovic. Digiaro and G. is distantly related serologically to ArMV but not to GFLV. M (particles contaning a molecule of RNA-2 with Mol. Bulletin OEPP/EPPO Bulletin 32. Coat protein subunits have a Mr 53 x 103. Immunosorben electron microscopy for detecting saptransmissible viruses of grapevine. 1980. Proceedings of the 7th Meeting of ICVG. is not seed-borne and reported only from south-eastern Turkey.: First isolation by mechanical transmission of an unknown nepovirus from cv. No vector is known and no information is available on the distribution and economic importance of the virus. mol. 2005. Gokalp..800 nt. A.6 x 106 Da and RNA-2. Description Grapevine deformation virus (GDefV) was recovered from Turkish grapevines showing distinct fanleaf-like symptoms. wt of 2. Saric A. The scattered distribution of infected vines in the field suggests that the virus is spread primarily by infected propagative material. 335-340 Cigsar I.3 x 106 Da and a size of 3753 nt.4 x 106 daltons and apparent size of c. CAB International. K.P. M. 251-257.P. Nematode Vectors of Plant Viruses. 1977. De Stradis. Savino. wt of 1. The genome is bipartite.. 149-151. Hranuelli..800 nt) and B (particles containing a molecule of RNA-1 with Mol. 46 . Complete nucleotide sequence of RNA-2 of two Turkish nepoviruses. S. 471-475 Cigsar I. distantly serologically related with ArMV. Proceedings of the Conference on Excoriosis and Virus Diseases of Grapevine. : Complete nucleotide sequence of GDefV RNA-2 3. The virus belongs in the subgroup A of the genus Nepovirus. Particles are isometric c. E. GRAPEVINE TUNISIAN RINGSPOT VIRUS (GTRSV) 1. GRAPEVINE DEFORMATION VIRUS (GDefV) 1. 35-41. Martelli. Digiaro and G. N. and T. Sanitary status of grapevine in south eastern and central Anatolia.P. Boscia and G. Historical review 2002 2003 2005 Cigsar et al. and D. M. Martelli. 30 nm in diameter and sediment as three components. GTRSV is serologically unrelated to any of 19 nepoviruses tested. Abou Ghanem-Sabanadzovic et al. Oxon. RNA-1 has a mol. wt of 2. 6. 286 pp.Russo M.The virus has no recognized vector. and belongs in the subgroup C of the genus Nepovirus.F Brown. M. Niagara Falls 1980. was isolated from a Tunisian grapevine with mild fanleaflike symptoms. wt of 2 x 106 daltons and apparent size of c.P. Martelli. References Abou Ghanem-Sabanadzovic N.: A mechanically transmissible virus was recovered by sap inoculation from Tunisian grapevines showing mild fanleaf-like symptoms. a novel nepovirus from Turkey. Description Grapevine Tunisian ringspot virus (GTRSV).. Historical review 1991 Ouertani et al. Taylor C. 2. The virus sediments as three components: T (empty shells). 2. identified as a new species in the subgroup A of the genus Nepovirus.

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

Vuittenez et al. Bercks and Querfurth: GFLV. J. The virus was detected in grapevines in the Niagara Peninsula. The genome is a bipartite positive sense single-stranded RNA occurring as two separately encapsidated functional molecules with Mol. RNA-2 is 4662 nt in size encoding a polyprotein with Mr of 150 kDa.virus). ArMV.: Nucleotide sequence of a TBRV satellite RNA. RNA-1 is 7356 nt in size and contains a single open reading frame encoding a polypeptide with Mr of 254 kDa. and B). including TBRV : bentonite flocculation test. but they can be high. 279-327 TOMATO BLACK RING VIRUS (TBRV) 1. Bercks and Stellmach: ArMV. Stellmach and Bercks: Further investigations on TBRV in grapevine. rings and lines on the leaves of recently infected plants. Tanne: Detection of TBRV by ELISA in Israel. wt of 0. Annales de Phytopathologie 2. wt of 2. Querfurth. Description Tomato black ring virus (TBRV) was first found in grapevines in Germany.Stellmach G. Pathologie und Thermo-Labilität mehrerer Reben-Isolate des Himbeerringflecken-Virus (raspberry-ringspot. riparia 143 A in West Germany. RRV and TBRV are not transmitted by contact of roots or foliage in the vineyard. 128-136. 1967 1970 1970b 1976 1977 1980 1984 1984 1986 48 .: RRV. Kuszala. Rüdel: Transmission of TBRV to grapevine by Longidorus attenuatus. Détection et étude selogique du virus latent des taches annulires du frasier (strawberry latent ringspot). its own subgroup. then in Yugoslavia. and Ontario (Canada). Their coat protein consists of a single type of subunits with Mr of about 57 kDa. respectively. Lehoczky and Burgyan: Occurrence of TBRV in grapevines in Hungary.65 x 106 (RNA-2) accounting for 44% and 37% of the particle weight. known to cause severe damage to the grapevine variety Joannes Seyve in Ontario. Stobbs and Van Schagen: First record of TBRV from Canada.. M. Greece. Joannes Seyve virus. Rüdel and H Brückbauer. TBRV produces a reduction in growth and yield. M. chlorotic spots. The vector to grapevine is Longidorus attenuatus. latex test and barium sulfate test. Losses are not known precisely. Ontario as the cause of severe damage to cv. The latex test is considered as the most sensitive and the least time consuming method. Weinberg und Keller 25. Israel. Bercks: Comparison of three serological tests for detecting several viruses. 2. 1970. Historical review 1963 1965 1966 Stellmach and Bercks: TBRV detected in rootstock Aramon x V. is a strain of this virus. Vuittenez A. either by agar gel diffusion with extracts of herbaceous hosts previously infected mechanically from grapevine.5 x 106 daltons and a size of 1327 nt. TBRV supports the replication of a satellite RNA with Mol. and G. Virus particles are isometric. 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. about 30 nm in diameter have angular outline and sediment as three components (T. 1978.7 x 106 (RNA-1) and 1. Stellmach: Review paper on TBRV in grapevine. Joannes Seyve. Meyer et al. RpRV and TBRV detected serologically in grapevine in West Germany. Turkey. SLRV and TBRV found in grapevine in the Palatinate. mottling of the older leaves. and an increase in graft failure. Untersuchungen zur Serologie. The virus is a definitive nepovirus species classified in subgroup A of this genus. or directly in grapevine leaf extracts using bentonite flocculation test.

The genome is a bipartite positive sense singlestranded RNA occurring as two separately encapsidated functional molecules with Mol. Historical review 1974 1977 1981 1982 1982 1987 1993 1994 2005 Murant: Description of SLRSV in the CMI/AAB Descriptions of Plant vrises series. and B). Brückbauer: SLRSV can be distinguished from other nepoviruses on the basis of symptoms induced on Vitis idicator plants. 1984. respectively.: SLRSV recorded from grapevine in Italy. 1993. Greif C. Turkiye.: SLRSV found in grapevine in Turkey..: Nucleotide sequence of TBRV RNA-1 Abkas and Erdiller: TBRV recorded from grapevines in Turkey 3. 2. Rüdel and H. Kreiah et al. Meyer M. Canadian Plant Disease Survey 64. References Akbas B. Assignement of SLRSV to the new genus Sadwavirus. Le Gall et al. Journal of Gerneal Virology 67. Babini and Bertaccini: Electron microscope study SLRSV infections in plant tissues. Journal of Turkish Phytopathology 22. O. Kreiah et al. and G. and J.. and 1.A. Hemmer. M. Complete nucleotide sequence of a satellite RNA of tomato black ring virus. Fritsch..6 x 106 (RNA-1) accounting for 38% of the particle weight. Journal of General Virology 65. Occurrence of tomato black ring virus on grapevine in southern Ontario. 1517-1529. about 30 nm in diameter have angular outline and sediment as three components (T.: Nucleotide sequence of SLRSV satellite RNA. RNA-2 is 3824 nt in size and encodes a single ORF expressing a polypetide with Mr of about 99 kDa. Hemmer and C. Van Schagen. 1986. wt 2. 279-327. Savino et al. du virus des anneaux noirs de la Tomate (tomato black ring). Journal of General Virology 69.W. Meyer M.: Nucleotide sequence of TBRV RNA-2 Greif et al. 1970. Hemmer and C. 1257-1271 Stobbs L. Détection et étude sérologique du virus latent des taches annulaires du Fraisier (strawberry latent ringspot). Fritsch. The nucleotide sequence of tomato black ring virus RNA-2. J. Kuszala. Bercks et al. Erdiller. Researches on grapevine virus diseases and determination of their incidence in Ankara. Fritsch.6 x 106 (RNA-2). The virus supports the replication of a satellite RNA 1118 nt in size. Nucleotide sequence of tomato black ring virus RNA-1. Their coat protein consists of two types of subunits with Mr 43 x 103 and 27 x 103. Annales de Phytopathologie 2.: SLRSV and other nepoviruses isolated from grapevines in Germany Credi et al. O.G. Vuittenez A. It was also detected in imported vines in Turkey and Portugal. Genus SADWAVIRUS STRAWBERRY LATENT RINGSPOT VIRUS (SLRSV) 1. Description Strawberry latent ringspot virus (SLRSV) has been isolated from grapevine in the Palatinate (Germany) and in northern Italy. 49 . et du virus des taches annulaires du Framboisier (raspberry ringspot) chez des vignes du Palatinat. The virus is transmitted by Xiphinema diversicaudatum. Virus particles are isometric. O. Mayo and C. The virus is a definitive species in the genus Sadwavirus. 1575-1583. Symptoms on affected European grapes are of the fanleaf type. 55-61.1986 1988 1993 Meyer et al. M. 3-5. 1988. 1984.: Nucleotide sequence of SLRSV RNA-2. M. Brückbauer.

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

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

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

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

E Morris. Ramsdell D. 1980.: A review of viruses infecting grapevines in New York vineyards..W. Bird GW. Gillet and C. 1979. Bird. but was recorded from grapevines only in New York State and Pennsylvania. Gillet.7 x 106 (RNA-1) and 1.C. Powell et al. Peach rosette mosaic virus. Superimposed shallow and deep soil fumigation to control Xiphinema americanum and peach rosette mosaic virus reinfection a Concord vineyard. about 30 nm in diameter with angular outline. 74-78. 1978.C. Lammers A. but in European grapes responses are similar to those elicited by GFLV. G. G. and J. 54 . R. 1174-1178. Pearson and A. Symptoms elicited by TRSV are the same as those of ToRSV in native cultivars. and R. RNA-2 has been sequenced only in part. Gillet and G.L.). 1995.. .F Allison and D.: TRSV agent of a new grapevine disease in New York State. Phytopathology 64. Goheen eds. 1985. Ramsdell D. 625-627. Peach rosette mosaic virus. and B). Coat protein consists of a single type of subunits with Mr of about 57 x 103 Da.M. Plant Disease 67. 154-157.M. Virus particles are isometric. Plant Disease 79. Relative susceptibility of American. Plant Disease Reporter 63.C. Peach rosette mosaic virus decline. 4143. TRSV has a relatively wide natural host range.C. Gillet. 447-450 Ramsdell D. Preventive control measures are the use of resistant roostock hybrids and of certified planting material.C. 2. Characterization of the single protein and the two nucleic acids of peach rosette mosaic virus. Allen. Phytopathology 68.C. M. 1988.C Ramsdell. Ramsdell D. Ramsdell D. and R.C. accounting for 44% and 28% of B and M particle weight. Ramsdell D. AAB Descriptions of Plant Viruses.Dias H. 1999.. Description Tobacco ringspot virus (TRSV) is the type species of the genus Nepovirus and the prototype of subgroup A. Canadian Journal of Botany 58.M. Ramsdell D.M. The genome is a bipartite positive sense single-stranded RNA occurring as two separately encapsidated functional molecules with Mol.C.W. J. American Vitis species reported to be resistant to ToRSV and TRSV. RNA-1 is 7514 nt in size and contains a single open reading frame encoding a polypeptide with Mr of 225 kDa.H. and J.: Nucleotide sequence of TRSV satellite RNA.R. 364. Susceptibility of American grapevine scion cultivars and French hybrid roostocks and scion cultivars to infection by peach rosette mosaic virus.3 x 106 (RNA-2). 1974.M.W. sedimenting as three components (T. Paul. Buzayan et al. American Phytopathological Society Press. The virus supports the replication of a circular satellite RNA 359 nt in size. 1983. symptomatology and nematodes associated with grapevine 'degeneration' in Michigan. Stace-Smith: Description of TRSV in the AAB Descriptions of Plant Viruses series. French hybrids and European grape cultivars to infection by peach rosette mosaic virus.: Survey of ToRSV and TRSV in Pennsylvanian vineyards.L.C. Rose. wt of 2. Virus Research 65. is endemic in Central and Eastern North America. J. 1747-1754. TOBACCO RINGSPOT VIRUS (TRSV) 1. Ramsdell D. Uyemoto et al. R. 51-52. A comparison between enzyme-linked immunosorbent assay (ELISA) and Chenopodium quinoa for detection of peach rosette mosaic virus in 'Concord' grapevines. Foster and Morris-Krsinich: In vitro translation of TRSV RNA-1 and TRSV RNA-2 yields major polypeptides with Mr of 225K and 116K. TRSV is soil-borne and is transmitted by Xiphinema americanum sensu stricto. No. Gillet and L. Epidemiology of peach rosette mosaic virus in a Concord grape vineyard.. and W. Phytopathologia Mediterrenea 24. St.. Cloning an sequencing of peach rosette mosaic virus RNA1. Myers. In: Compendium of Grape Diseases (R. Historical review 1970 1977 1985 1985 1986 1990 Gilmer et al. respectively. 57-73. Andrews. respectively.C. Myers.C. 1998.M. There is no evidence of seed trasmission in the grapevine.

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

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

. Phytopathology 72.G and V.F..E. Savino. Gooding G. their epidemiology and control.C. 1977. 1989. Gonsalves D. 44-50... 1995. Téliz D. 1169. Association of Xiphinema species with soil types ang grapevines infected with tomato ringspot virus in Ontario. Nematode-borne viruses of grapevine. Taylor.3. Li. Rokni. 1505-1514. and C. 702-704. 2004. Plant Disease 74. Longenecker and L. 1982..V.G.V. 1991. Uyemoto J. Lee and D'A. 658-663.W. 1966. Ebsary. and H. Plant Disease Reporter 61. 1972. M. Xiang . Current Topics in Vector Research 5. 24-28. 1993. 1975. 1-27.F. Dias and J. Some grapevine viruses in pollen and seed. 1987. M. Hewitt. Li M. A severe outbreak of virus-induced grapevine decline in Cascade grapevines in New York. Incidence of tomato rigspot virus and tobacco rigspot virus in grapevines in Pennsylvania. 98-101.M. 290 Stace-Smith R. and B.J.S Whei.K.W. V. and D.. Podleckis. Corbett. and M. Phytopathology 58. 1992.. 133-135. Phytopathology 79.B. Purification and serology of a virus associated with the grape yellow vein disease. Van Schagen and B.E. 465-473. Castellano and D. Piazzolla P. Susceptibility of grape cultivars and rootstocks to an Ontario isolate of tomato ringspot virus. Rott M.A. Phytopathology 53. Tremaine and D'A. L. Stobbs. 1963.G. J. Presence and incidence of grapevine viruses in central Chile. 2001. 57 . Herrera M. Detection and identification of plant viruses for quarantine in China. Forer. Canadian Journal of Botany 55.A. Rochon. 95-106. Journal of General Virology 72. Madariaga. Subikova. Powell C.R. Lownsberry. Grogan and B. Phytopathology 56. Ramsdell. Bitterlin M.A. J. 35-44. J.F. Corbett M. 1980.M.. Uyemoto. Phytopathologia Mediterranea 24. 1984. and J. Martelli G. Gonsalves. Chen. Membrane-associated spherical particles in extracts and tissues of virus-infected grapevines. and the association of a mechanically transmissible virus with the disease. Incidence of tomato ringspot virus in grape in Virginia. Advances in Disease Vector Research 6. 393-400 Hewitt W. Properties of the single protein and two nucleic acids of tomato ringspot virus. A comparison of Grapevine yellow vein virus and a Grapevine isolate of Tomato ringspot virus. N. Sampling strategies for the detection of grapevine fanleaf virus and the grapevine strain of tomato ringspot virus. Allen W.V. and E. Transmission of tomato ringspot. Hewitt. 1989. American Journal of Enology and Viticulture 13. H. Cory L. Baumgartnerova H.G. Incidence and geographic distribution of tomato ringspot virus in De Chaunac vineyards in the Niagara peninsula. Beijing 2004. Agricultura Tecnica 61. No. 408-411. 1985.R.. Plant Disease 72. Stace-Smith R.P.R. Journal of General Virology 76. 1990. Bulletin of the California Department of Agriculture 43. 196-203. Dias.A. 1988. and S. Nematologia Mediterranea 6. Proceedings 15th International Plant Protection Congress. 1028-1037. Ivanka pri Dunaji 4. Musci.K. Grape yellow vein: symptomatology. 663 Martelli G. and W.. Van Schagen. Rowhani A. 1987. G. Walker and S. 475-480. CMI/AAB Descriptions of Plant Viruses. Rochon. Allen W. Vitis 31.F. Canada.. 1982. 1962. peach yellow bud mosaic. R. Plant Disease 71. L. 13161320 Dias H.. Proceedings 7th Meeting of IGVG. M.B. and V. Gilmer R. 1985.B.B.V. Canadian Journal of Plant Pathology 4. References Allen W. Gooding G. Bays D. Some virus and virus-like diseases of grapevines. Nucleotide sequence of tomato ringspot virus RNA 1.E. 1954.A.. 1977. Nucleotide sequence of tomato ringspot virus RNA-2. Plant Disease Reporter 59. Distribution of viruses and their nematode vectors.L.. Plant Disease Reporter 56. Zhang and Y. Gilchrist.H. Spatial distribution of Xiphinema rivesi and persistence of tomato ringspot virus and its vector in soil.F. 131-166.K. Identification of tomato rinsgspot virus in leafroll diseased grapevines. 861-863. A. Detection of tomato ringspot virus in grapevines: irregular distribution of virus.F. Podleckis E. Y. 31-34. 157-164. Works of the Institute of Experimental Phytopathology and Entomology . 47-64. Tomato ringspot virus associated with little grape disease of Vidal 256 grapevines. Tomato ringspot virus in "Baco noir" grapevines in New York. 710. 275-277.P.F.. Tolin. 151-189. 1968. Nepoviruses of the Americas.C. Tomato ringspot virus. Phytopathologia Mediterranea 24. 1978. Niagara Falls 1980. and W. Rott M. identification.. and grape yellow vein viruses by Xiphinema americanum.K.

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

LEAFROLL .

.

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

GLRaVs differ in various vays (molecularly. 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. which is the type species of the novel genus Ampelovirus has a genome 17. Parthenolecanium corni. but it does not discriminate between GLRaVs and was reported to be less sensitive than ELISA. All GLRaVs can be identified by serological and nucleic acid-based techniques. soil condition and probably. Leaf tissues or petioles from mature symptomatic leaves of V. has a number of minor biological variants which can be differentiated by the reaction of inoculated Nicotiana species. Indexing on red-fruited cultivars such as 'Cabernet sauvignon'. Ps. contains 13 ORFs. Regardless of whether they belong to the genus Closterovirus or Amplelovirus. GLRaV-5 and GLRaV-9 have been identified.528 nt in size. American rootstocks are usually symptomless carriers of GLRaVs. roostocks. or the hybrid LN 33 is still the most popular method for identifying the disease. So far. and some of them are used as indicators. 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. GLRaV-2. GLRaV-3. maritimus. 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. climate. This has been ascertained experimentally for GLRaV-1. and has structural organization differing from that of other sequenced closteroviruses. 29. vinfera and cortical shavings from mature dormant canes of V. broadspectrum. with none of which they are serologically related. viburni and Ps. ultrastructurally. calceolariae. Pseudococcus longispinus. comstocki. Mealybug vectors of GLRaV-3 are Planococcus ficus. Its soft scale insects vectors are Pulvinaria vitis and Neopulvinaria innumerabilis. leafroll can be detected by its symptoms in the field on red-fruited varieties. The genome is a monopartite. Ps. Varietal susceptibility and sensitivity: No immune variety or rootstock is known. As to nucleic acid-based assays. Spread at a site is mediated by mealybug and soft scale insect vectors. and some are commercially avaliable.919 nt in size. 'Merlot'. the only member of the group to be mechanically transmissible. or by molecular techniques. and degenerate primers have been designed and successfully used for PCR detection of virtually all GLRaVs. vinifera varieties show symptoms most clearly because of the reddening of the leaves. biologically. classical double antibody sandwich ELISA (Chromo-ELISA) or Lumino-ELISA. grafted vines) which is largely responsible for its dissemination over medium and long distances. Red-berried V. Natural field spread of leafroll disease has been reported from many countries in Europe and elsewhere. Polyclonal antisera and/or monoclonal antibodies have been raised to each single GLRaV . These reagents are routinely used for ISEM.5 kDa for GLRaV-4 . Pl. affinis. GLRaVs show molecular variations which give rise to a population of strains. 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. contains nine open reading frames (ORF) and has structural organization identical to that of Beet yellow virus (BYV). vinifera. number an types of infecting viruses. American Vitis species and rootstocks are the best antigen sources for serological assays. GLRaV-2 and GLRaV-3. Composite samples should be used to minimize false negative responses that may originate from the unven distribution of GLRaVs in chronically infected vines. GLRaV-5 and GLRaV-9 are phylogenetically close to one another but are serologically unrelated. The presence of high molecular weight double-stranded RNAs (dsRNA) in phloem tissue extracts can be used as infection marker. Detection: In many cases. and epidemiologically) from most of the known closteroviruses. GLRaV-5 and GLRaV-9 are both transmitted by Ps. Disappearance of dsRNAs from vines submitted to 62 . Other GLRaVs may exist in nature as suggested by reports. Ps. GLRaV-1 is transmitted in nature by the pseudococcid mealybugs Heliococcus bohemicus and Phenacoccus aceris and the soft scale insects Pulvinaria vitis. and Neopulvinaria innumerabilis. only vectors of GLRaV-1. in agreement with the quasispecies nature of viruses. 'Cabernet Franc' 'Pinot noir'. singlestranded. The genome of GLRaV-4 and GLRaV-9 are incompletely sequenced but show a structural organization compatible with that of the genus. the type member of the genus. Ps. GLRaV-3. citri.Transmission is semipersistent and does not appear to be vector-specific. GLRaV-3. The genome of GLRaV-2 is 15. Symptom expression depends on the variety. positive sense RNA molecule. Foliar tissues are not recommended for serological GLRaVs detection in American Vitis species and roostocks. None of the GLRaVs is known to be seedborne. The genome of GLRaV-1 is 17.35 kDa for both GLRaV-3 and GLRaV-1.647 nt in size and contains 10 major ORFs. Membranous vesicles can derive eitrher from peripheral vesiculation of mitochondria followed by disruption of the organelles (GLRaV-1. longispinus.

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

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

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

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

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

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

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

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

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

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

3 and 4. Murant et al. Suggestion that the disease is not related to closteroviruses associated with GLRaV and corky bark.: First record of rugose wood from China.: Evidence that GSP-AV can occur in grapevines together with another similar but serologically unrelated virus with short closterovirus-like particles. wt of 5. Savino et al.: Transmission of corky bark by the mealybug P. Azzam et al. Monette et al. denoted Grapevine virus B (GVB).: Experimental confirmation of the complex nature of rugose wood based on the differential reaction of woody indicators.: Heracleum latent virus and GVA are distantly serologically related. No closterovirus-like particles in samples with rupestris stem pitting. an additional disease of the rugose wood complex. Garau et al. Monette and James: Detection of two biologically distinct but serologically indistinguishable isolates of GVA.Sc. thesis describing rupestris stem pitting disease in comparison with corky bark. Corbett and Wiid: Closterovirus-like particles found in extracts from vines affected by corky bark and rugose wood in South Africa. but not consistently in grapevines from New York.ficus.: Three types of wood disorders of the stem-grooving type observed in South African grapevines.: First indication of the possible existence of LN 33 stem grooving. Similar dsRNA species were detected.: Assessment of crop losses induced by rugose wood to two different European grape varieties. Italia propagated on six different rootstocks. Savino et al.: Application of spot hybridization for the detection of GVA in grapevine sap. Gallitelli et al. Rosciglione and Castellano: Demonstration that GVA is transmitted also by Planococcus citri and P. ficus. The first two disorders appear to be spreading in the vineyards. Engelbrecht and Kasdorf: Natural field spread of corky bark in South Africa associated with the presence of P. Martelli: Rugose wood recorded in southern Mediterranean and Arab countries.1984 Milne et al. Engelbrecht et al. Kuniyuki and Costa: Rugose wood recorded from Brasil Goheen: First published description of rupestris stem pitting. Savino et al. Castrovilli and Gallitelli: Physicochemical comparison of two Italian isolates of GVA. Rugose wood and corky bark are not the same disease. Tanne et al.4 x 106 associated with rupestris stem pitting in grapevines from California and Canada.: Evaluation of the effect of rugose wood on cv. Garau et al. GSP-AV re-named Grapevine virus A (GVA). ficus.: A low molecular weight dsRNA associated with rupestris stem pitting. Prudencio: M. 1985 1985 1985 1985 1985a 1985b 1985 1985 1985 1987 1988 1989 1989 1989 1989 1989 1989 1990 1990 1991 1991 83 .: Experimental confirmation that rugose wood may not express symptoms in grafted indicators. similar to Kober stem grooving. Corky bark and Rupestris stem pitting. Li et al. First report of Kober stem grooving.: Two distinct dsRNAs with a mol.

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

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

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

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

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V). Musci and G. A. Savino V.. distribution. Coote. Plant Disease 80. Phytopathologia Mediterranea 24. M. Castellano. Gafny. Vitis 22. Meng. 36 pp. Immunodetection and subcellular localization of the proteins encoded by ORF 3 of grapevine viruses A and B. Valle. M. 186-188. Grape corky bark and stem pitting in Mexico.. 204-207. 91 . Phytopathology 87. D. Goheen. A. P. 964-970. Proceedings 10th Meeting of ICVG. 2645-2652. Phytopathologia Mediterranea 24. 1996. B. and E. I. Tanne E. Komazaki. Rugose wood complex of grapevine: can grafting to Vitis indicators discriminate between diseases? Proceedings 9th Meeting of ICVG. Proceedings 7th Meeting of ICVG. R. 91-94. 1980b. C. RT-PCR detection of rupestris stem pitting associated virus within fieldgrown grapevines throughout the year.A. J. 1991. 2000a. Rubinson E. Mavric and D. Rosciglione B. Vitis 34. 1966. Martelli. Minafra. Gonsalves.. Davis. Golino and D. Nakano M. R.A. Mealybug transmission of grapevine virus A. 1985a. Comparative effects of corky bark and rupestris stem pitting diseases on selected germplasm lines of grapes. Valle and A. Ben Dov and B. 1535-1542. S.Monette P. Extended Abstracts 14th Meeting of ICVG. Saldarelli P.P. Nakaune and S. 367-374. Padilla V. Cannizzaro. C.Sc. Azzam. 1993. Luévano. Tanne and R. Incidence of some graft-transmissible virus-like diseases of grapevine in visually selected and heat-treated stocks from Southern Italy. 218..M. Saldarelli P. Castellano and G. Influencia del complejo de la madeira rizada de la vid en el cv Napoleon negra. Performance and compatibility of "Muscat Gordo Blanco" grape on eight rootstocks. Téliz D.. Phytopathology 81. 34-38. Locorotondo 2003. Murant A.. D.P. Gonsalves. 331-347. Boscia and G. 1997.A. 2003. Analysis of progress and spatial patterrn of corky bark in grapes. 1989. Dell'Orco and A. Bevington and B. Journal of General Virology 77. 1990.. Boscia. Plant Disease 87. II. Mealybug transmission of grapevine viruses in Japan. D. Martelli.B. 51-64. and M. Non-radioactive molecular probes for the detection of three filamentous viruses of the grapevine. Infectious cDNA clones of two grapevine viruses. Phytopathologia Mediterranea 24. 33-36. Volos. O. 1041-1045. Hu. Occurrence and spread of viruses associated with leafroll (GLR) and stem pitting (GSP) diseases in the north-western part of Yugoslavia. Garau and G. Occurrence. The detection of disease specific double-stranded RNA in corky bark affected grapevine.. I. Castellano. and A. Vitis 33. D. Plant Disease 85. Heracleum latent virus (HLV) and heracleum virus 6 (HLV6). 1991. Archives of Virology 145. Rivista di Patologia Vegetale 3 (Ser. R. 1985.L. Report of the Scottish Crop Institute 1984. M.. Goheen and S. natural spread. Grape corky bark and stem pitting in Mexico. and Z. 397-405. P. M.. 1994. Further evidence that mealybugs can transmit Grapevine virus A (GVA) to herbaceous hosts. Nassuth. 2003. Proceedings 7th Meeting of ICVG. Rosciglione B. Phytoparasitica 17. Martelli. 1982. 1993. Galiakparov. Saldarelli P. Greece. Saldarelli P. V. Marcus.. 157-160. Tanne E.P. Transmission of the corky-bark disease by the mealybug Planococcus ficus. 15-22 Savino V. 2001. Duncan and I.G.. Minafra. Y. Evaluation of symptoms in 17 rootstocks. 2000b. H. Godkin.. Dubitzky and B. Boscia and G. N. D.E. 1985. 1985. Vitivinicultura 4 (7-8). Raccah.S. 1980a.P. Department of Plant Pathology. 68-72. University of California. Raccah. Scientia Horticulturae 16. M. 1983.P. Savino and G. Radian. G. Ravnikar. Archives of Virology 145. Prudencio S. Niagara Falls 1980 . 182.. Saric A. 416 Sarooshi R. Roberts. Effect of legno riccio (stem pitting) on "Italia" vines grafted onto rootstocks of different origin. Minafra and G. the putative movement protein. Purification and properties of closteroviruslike particles associated with grapevine corky bark disease. Boscia. A cloned probe to grapevine virus B.P. Stewart S. Martelli. Saldarelli P. K. Martelli. First detection of rupestris stem pitting associated virus particles by antibody to a recombinant coat protein.F. Savino V. Volos 1990.P.. 1991.A. G. 55. 247-250. 1985b. M. Martelli. Petrovic N. Martelli. Namba S.H. I... effect on yield and evaluation of symptoms in 128 grape cultivars.P. Niagara Falls 1980. 617-620 Tanne E. Guglielmi Montano and G. Serological detection of grapevine virus A using antiserum to a non structural protein. A.. Kiryat Anavim 1987. The nucleotide sequence and genomic organization of grapevine virus B. E. D. A. Detection of capillovirus-like particles in a grapevine affected with rugose wood. 510514. Proceedings 10th Meeting of ICVG. 65-70. Maixner. thesis.. Minafra. 241-242. 1995. E. Martelli. Sela. 1989. Téliz D. Korosec-Koruza. and S. Meir.

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GRAFT INCOMPATIBILITY .

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

immunocapture RT-PCR. Suggestion that they are molecular variants of the same virus species.: GLRaV-2 is associated. with a decline condition of young Thomposn seedless vines in Chile. whenever feasible. Boubals: Report of a national French study group investigating the aetiology of Syrah decline.: GLRaV-2 is the cause of a graft incompatibility revealed by Kober 5BB.: Two papers describing incompatibility phenomena between clonal selections of different cultivars grafted on Kober 5BB Fallot et al. GRSLV has about 75% nucleotide homology with GLRaV-2.: Third paper of a series on incompatibility on Kober 5BB. Savino et al. meristem tip culture. Renault Spilmont et al. utilization of tolerant roostocks is advisable. and death of the vines. using degenerate or virus-specific primers. or a combination of the two. 1977 1979 1986 1991 1995 2000 2000 2000 2001 Jacob: Description of graft incompatibility in different scion/stock combinations. Control: Prevent introduction of infected vines in the vineyard by using certified grafted plants or virusfree scionwood and rootstocks. 2. Historical review 1942 1950 1973. Greif et al.: Updated report on the state of the art of investigations carried out in France on Syrah decline. Graft-transmission of the incompatibility factor. Strategies on how to protect healthy stocks from vector-mediated GVB reinfection in the field are yet to be developed.: Description of bushy stunt and evidence that it is caused by a graft-transmissibile heat-sensitive agent carried by some clonal rootstocks. the use of sensitive rootstocks is to be avoided and. though not consistently. Golino et al. Boubals and Huglin: Report on graft incompatibility of certain varieties grafted on 57R. decline. Currently known graft incompatibility agents can be eliminated with reasonable efficiency by heat therapy. The problem is very complex and may involve several still unidentified factors. Durquety et al. No conclusion are drawn.: Identification of an apparently new closterovirus denoted Grapevine rootstock stem lesion virus (GRSLV) causing stem necrosis of rootstocks. 2003 2003 2003 2003 2003 2003 2004 96 . Bonfiglioli et al.: GLRaV-2 and GVB are consistently associated with young vine decline in California. Boubals: Syrah decline occurs in Argentina Uyemoto et al. If scionwood is infected. Uyemoto and Rowhani: Indexing on 18 different grape rootstocks reveals the existence of at least five different agents causing graft incompatibility. GRSLV re-named Redglobe strain of GLRaV-2. Legin and Walter: The graft-transmissibile agent that causes incompatibility of different varieties on Kober 5BB is a virus which can be eliminated by heat treatment at 37 °C for 58 days. Bertazzon and Angelini: Comparison of several detection methods for the broad or specific identification of Grapevine leafroll-associated virus 2 variants. Gomez Talquenca et al: GLRaV-2 is consistently associated with declining Cabernet sauvingon vines grafted on different roostocks in Argentina.: Report of a new molecular variant of GLRaV-2 from New Zealand. Prodan et al. Martelli: GRSLV and GLRaV-2 are serologically related and are both recognized by a panel of 18 monoclonal antibodies.

. Ruchaud..M. Garau. The relationship of grapevine leafroll-associated virus 2 with a graft incompatibility condition of grapevines. Grapevine virology highlights 2000-2003. Boubals D. 167-173. Progrés Agricole et Viticole 103. Autres cas chez la vinge. Le dépérissement de la Syrah. F. Progrés Agricole et Viticole 96. A young grafted vine decline syndrome in Argentina vineyards. V. Examples of incompatibility between grape varieties and roostocks.E. D. Durquety P. Montalegre and N. 3-10. 277. 2004. and A. S. Proceeding of the American Society of Horticultural Science 41. Garcia Lampasona and O. References Bertazzon N... 2003. Boubals D. 2003. Volos 1990. 1979. Locorotondo 2003.. Jacob H.. Locorotondo 2003. 28-31. Extended Abstracts 14th Meeting of ICVG. Sim and A. Grau. Durquety and J. 1995. Ruchaud. Progrés Agricole et Viticole 94. Benassac and R. Prodan S. 2003. C. Boursiquot. Grenan and J. Adelaide 2000. Legin R. Savino V. 2003.A. and E. Rowhani. Syrah decline in French vineyards.3. Ruchaud. California Agriculture 55 (4). Molecular studies on a graft incompatibility syndrome in New Zealand vineyards yields another probable variant of Grapevine leafroll-associated virus 2. Uyemoto J. 141. 211-216. J. S.M. 1991. Locorotondo 2003. Progrés Agricole et Viticole 117. Rivieccio and F. 202-210 Uyemoto J.A. C. S.K. 1986. Greif C. B. Etude de phénomènes d'incompatibilité au greffage chez la vigne. Rowhani. Progrés Agricole et Viticole 67. 2000. 97 . Gazeau. Phytopathologia Mediterranea 34. 1942.P. Extended Abstracts 14th Meeting of ICVG. M.M. 1973. 1977. Locorotondo 2003. 2000. Le dépérissement de la Syrah existe aussi en Argentine.S. Journal of Plant Pathology 86. Extended Abstracts 14th Meeting of ICVG. Proceedings 10th Meeting of ICVG. Extended Abstracts 14th Meeting of ICVG. J. Gracia. 283-290. New closterovirs in 'Redglobe' grape causes decline of grafted plants. 2003. Compte-rendu de la réunion du Groupe de Travail National... 2001. Prota. Le clone et ses reactions au greffage. P. Pantaleo. Huglin. La trasmission de l'incompatibilité au greffage entre 5BB et Vitis vinifera.. Fiore. 85-86. Gazeau and J. Presence in clonal rootstocks of a grafttransmissible factor that induces stunting and bushy growth in European grapevines. 122-129 and 171-178. Extended Abstracts 14th Meeting of ICVG. B.. P. Durquety P. 137-141 Boubals D. Discovery of different grapevine sources with graft-transmissible agents causing union-incompatibility on sensitive rootstocks.P. Walter. Le clone et ses reactions au greffage. Locorotondo 2003. Fiori.. Angelini. Renault Spilmont A. Aetiology of decline in Thompson seedless grafted table grape plants.III. 139-140. R. 420-427 Fallot J. 2000. Boscia.K. 1950..M. J. A..S. C. Martelli G. Krag. I. 201-203. Etude de l'incompatibilité au graffege de certain cépages et du 57R. Nouvelles recherches sur l'incompatibilité clonale d'Abouriou greffé sur 5BB. 279-283.II. Golino D. Fallot. Bonfiglioli R. Progrés Agricole et Viticole 90. and P.. Existence dans un cépage population de clones présentant divers degrés de compatibilité avec certains porte-greffes. Luvisi and R. Advances in the detection of Grapevine leafroll-associated virus 2 variants.. S. Progrés Agricole et Viticole 117. O. Rowhani. Extended Abstracts 13th Meeting of ICVG. and B. Dauty. Extended Abstracts 14th Meeting of ICVG. Bass. 2003. Walter and U. 142-143. Di Terlizzi. Gomez Talquenca G. Le clone et ses reactions au greffage. 85-86. Identification of the latent viruses associated with young vine decline in California. Edwards and A. Di Silvio. Prota. 144. 183-189. Fallot.P. J.P. Locorotondo 2003. D.

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FLECK COMPLEX .

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GFkV particles sediment as two centrifugal components.). irregularly distributed over the leaf blade. and GRVFV genomes are available. Grapevine red globe virus (GRGV) is a Grapevine fleck virus (GFkV)-like virus which apparently does not induce symptoms in European grapevine varieties (e. The putative causal agent of the disease has only been found in California. Description Main synonyms: A. Agents: All viruses of the complex. 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. vinifera. rooting ability of rootstocks and on graft take has been reported. Severe strains induce also varying degrees of stunting. d. e. Leaf symptoms usually become less severe in summer. V. c. which is a monopartite single-stranded. screziatura (Ital. 1. The disease is latent in European grapevine varieties and in most American rootstocks. This disease. Leaves with intense flecking are wrinkled. T made up of empty protein shells and B. 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. Although the elusive nature of the complex hinders the assessment of its economic impact. grapevine asteroid mosaic. twisted and puckered along the veins. wt of 2. adverse influence on vigour.). GFkV genomic RNA (Mol. leaf petioles and veinlets. positive sense RNA with high cytosine content (c. Affected vines are often stunted. Symptoms are expressed in Vitis rupestris and consist of clearing of the veins of third and fourth order. Symptoms: a. All have isometric particles about 30 nm in diameter with rounded contour and prominent surface structure with clusters of coat protein subunits arranged as pentamers and hexamers. Rupestris necrosis. twisted and may curl upward. Asteroid mosaic. Marmorierung der Rebe (Germ. which are twisted and asymmetric. Asteroid mosaic symptoms have been observed in several varieties of V. maculatura infettiva. Sultanina). whereas the CP of GAMaV and GRVFV consists of a major protein of 21 kDa and a minor prtotein of 25 kDa. producing localized translucent spots. Grapevine asteroid mosaic: Mosaïque étoilée (Fr. rupestris. Grapevine fleck: Marbrure (Fr. Fleck is an ubiquitous disease reported from most viticultural countries in the world. is latent in European grapevine varieties. Mild asteroid mosaic-like symptoms are shown by some European grapevine varieties (e. Rupestris vein feathering. GFkV genomic RNA constitutes about 35% of the particle weight. grapevine rupestris necrosis. Fleck. B.). The coat protein (CP) of GFkV and GRGV particles is made up of a single protein species with Mr of c.6 x 106 ) is 7564 nt in size and contains four open reading frames (ORF) that 101 . vinifera in California. Italy and California. Sternmosaik der Rebe (Germ. b. Leaves are asymmetric.). Recorded from California and Italy . rupestris reacts with localized necrosis of the shoots. containing the genome. capped. the disease elicits creamy-yellow bands developing along the major veins of the leaves. rupestris following graft inoculation.).FLECK COMPLEX The fleck complex consists of several diseases (grapevine fleck. In V. GRGV. and Grapevine rupestris vein feathering virus (GRVFV) are all phloem-limited and non mechanically transmissible.g. reported only from Japan. which is used as indicator. but likely to occur elsewhere. mosaico stellare (Ital. In V.). Red globe) nor in V. The complete sequence of GFkV and partial sequences of GRGV. GFkV. The putative causal agent of the disease so far has been found in Greece.g. leaf symptoms are characterized by star-shaped chlorotic spots. Grapevine asteroid mosaic-associated virus (GAMaV). GAMaV. 25 kDa. and produce little or no fruit. 50%). sometimes with necrotic center. rupestris. Transient mild chlorotic discolourations of the primary and secondary leaf veins develop in V.

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

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

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

Sequencing of the 3' end of three grapevine fleck virus-like viruses . Barlass M. GAMaV and of a virus of Greek origin which induces vein feathering in V. Savino. 1985.. G. Vitis 40: 65-68. Vitis 22. 1984. Multiplex RTPCR detection of Grapervine fleck virus-like viruses in grapevine with co-amplification of control plant mRNA. 3134.2002a 2002b 2003a Martelli et al. S. comprising the genera Maculavirus and Marafivirus that include GFkV/GRGV and GAMaV/GRVFV. R. Sabanadzovic and G.A. Double stranded RNA associated with fleck disease of grapevine.: Sequencing of the 3' end of the genome of GRGV. Bovey R. GRVFV recorded from California and confirmation that GAMaV does not occur outside of California. Martelli and V.. 165-169. Ultrastructure and nature of vesiculated bodies associated with isometric virus-like particles in diseased grapevines.. South Africa. Savino and M.. 195.. Identification of the agent of grapevine fleck disease. Verderevskaya and J. 23-39. Some properties of a phloem-limited non mechanically-transmissible grapevine virus. M.P. 105 . A non mechanically transmissible virus associated with asteroid mosaic of the grapevine. Boscia.P. 1983.: A sequence variant of GFkV (GFkV416) with a 63 nucleotide insertion in the replicase gene identified in Australia and New Zealand. Abou Ghanem-Sabanadzovic et al. M. 2003a. and G. and A.P. Castellano M. In other countries (USA.A.A. Vitis 30. Digiaro. Advances in Horticultural Science 10.P. respectively. Annales de Phytopathologie. Martelli.A. A. Association of a phloem-limited non mechanically transmissible isometric virus with grapevine fleck disease. T. 1991a.P.J. V. Tomashevskaya.G. Savino. Sabanadzovic. 1991b. Abou Ghanem-Sabanadzovic.P. Greek virus recognized as a species in its own right denoted Grapevine rupestris vein feathering virus (GRVFV).. 1990. Atabekov. A... G.G. Engelbrecht D. A. Extended Abstracts 14th Meeting of ICVG. G. Rowhani and G. Savino. A. B. V. Journal of Phytopathology 129. rupestris confirms the assignment of GRGV to the genus Maculavirus and of GAMaV and the Greek virus to the genus Marafivirus. Castellano. 1990. Castellano M. Dolja V. Shi et al. N.F. 160-163. Woodham and L. Martelli et al. Martelli. V. Krake. Boyko. N. N. 1994. 1972. 173-174. Elicio. V. Iran. Abou Ghanem-Sabanadzovic et al. Babini. leafroll and Shiraz decline diseases and associated viruses in South African grapevines. Virus-like particles and ultrastructural modifications in the phloem of leafroll-affected grapevines.. Martelli and M.: Development of a multiplex RT-PCR protocol for the simultaneous detection of GFkV-like viruses using plant mRNA as an internal control.A. 2003b. 347-354.D. Volos 1990. 97-105. U. Martelli. Savino and G. Martelli. Savino. a new genus of plant viruses having GFkV as type species and GRGV as tentative species. Sabanadzovic.. P.. Proceedings 8th Congress of the Mediterranean Phytopatological Union. Molecular detection of Grapevine fleck virus-like viruses. Savino and D. ElBeaino T.R. Kyriakopoulou and G. 291-295.: Description of Maculavirus.E. Boscia. 1982. 1990. Martelli. Castellano.P.G.P.C. Boscia D. Martelli. 101-102 Boscia D. V. 1996. Boulila M. Field spread of corky bark. Vitis 33.P. Martelli. Agadir 1990.E. Argentina. Phytophylactica 22. fleck.. Rowhani P. V.: Description of the family Tymoviridae. Skene. Boscia D. Boscia D. Numéro hors série. D.P.A. Annals of Applied Biology 101. Castellano. and G. 191.P. Virus Genes 27. and Japan) only the variant without insertion (GFkV353) was detected. Progress in the study of the phloem-limited isometric virus-like particles associated with leafroll-diseased grapevines. Phytopathologia Mediterranea 24. References Abou Ghanem-Sabanadzovic. Di Terlizzi. Effect of virus and virus-like infections on the growth of grapevine rootstocks. Karsev. Un virus latent dans le Chasselas. 56-64.P. S. O.. Production of monoclonal antibodies to grapevine fleck virus. Sabanadzovic . Credi R. 11-16 Abou Ghanem-Sabanadzovic.R. Castellano M. G. Locorotondo 2003. 151-158. S. Martelli. and G. Plant Pathology 44. 1995. Regeneration of virus-free grapevines using in vitro apical culture. S. Proceedings 10th Meeting of ICVG.M. K. Kyriakopoulou and G. Journal of Ultrastructure Research 89. 2003b 2003 3. V.V. Kasdorf.V. Martelli 2001. 95-98. Minafra.

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

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

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g. with which specific viruses are associated and thought to be their possible causal agents. Yellow mottle has been reported from Germany. Bovey and Cazelles: AMV particles visualized in thin sectioned grapevine leaves. suggesting that the virus spreads primarily through infected planting material. Bovey and Brugger: AMV recorded from Switzerland in grapevine and transmitted by grafting to V. 109 . however. Description Main synonyms: None. Varietal susceptibility: Little information available. former Czechoslovakia. Main symptoms: Various patterns of yellow discolouration characterize the disease. Some of these diseases have been recorded only from Europe. Detection: AMV is mechanically transmissible to herbaceous hosts and can also be identified by ELISA and moleculat techniques in infected vines. 18% of the particle weight. The spring growth shows more or less extensive yellowing of the leaf blades that does not extend to the veins. Historical review 1973 1975 1976 1979 1978 1981 Bercks et al. Grapevine berry inner necrosis). In grapevines.MINOR VIRUS DISEASES Several graft-transmissible diseases are known. There may be differential susceptibility among cultivars. a mechanically transmissible virus. 1. Capsid proteins subunits are of one type. 30 to 57 nm in size.62 x 106 (2037 nt).04 x 106 Da (3644 nt). Plant vigour and yield do not seem appreciably affected. Bulgaria. Chardonnay and Veltliner rouge précoce identified as reliable indicators. and Turkey. European diseases GRAPEVINE YELLOW MOTTLE 1. Their overall importance is minor if compared with that of the major diseases dealt with in previous chapters. with the following mol. the type species of the genus Alfamovirus. Agent: Alfalfa mosaic virus (AMV). Virus elimination by treating 37 days at 37-38°C. Control: Use of healthy material obtained by heat treatment. has differently shaped particles. and a tripartite RNA genome accounting for c. rings and lines are typical summer responses of infected vines. but some are of economic relevance locally (e. with Mr 24x 103 Da. from quasi isometric to bacilliform. rupestris and the hybrid Grézot 1 x 5C. Beczner and Lehoczky: AMV infections recorded from Hungary. Transmission: AMV is efficiently transmitted by aphids in a non persistent manner and can cause epidemic outbreaks in many of its natural hosts. A. RNA-2. Jankulova: AMV infections recorded from Bulgaria.: First record of AMV infections and description of symptoms in German grapevines. is the putative causal agent. Faint yellow speckling. 0. 0.73 x 106 (2593 nt). RNA-3. 2. Hungary. Switzerland. Novak and Lanzova: AMV infections recorded from hop and grapevine in Czechoslovakia. others occur in Japan. infections are scattered and occasional. AMV. wts: RNA-1.

110 . Brugger. Über den Nachweis des Alfalfa mosaic virus in einer Weinrebe. Proceedings 6th Meeting of ICVG. 2. 119-128. 131-134. GRAPEVINE LINE PATTERN 1. Grapevine line pattern virus (GLPV) a possible member of the genus Ilarvirus. Bovey R. Novak J. Plenum Press.1985 1993 1993 Francki: Comprehensive review of the properties of AMV and other viruses with tripartite genome. Cordoba 1976. 1993. (ed. Munich 1978. Acta Phytopathologica Academiae Scientiarum Hungaricae 16. 3. 263 pp. Polyhedral Virions with Tripartite Genomes (R. is seed-transmitted and spreads with diseased propagative materials. Biologia Plantarum 18.).Hegi) plants in Czechoslovakia. Le virus de la mosaïque de la luzerne sur la vigne. 55-63. vinifera cultivars are susceptible. 166-171. Lanzova. Francki R. scattered spots or blotches. ed. and J. 1975. Francki. 1-18. Lehoczky. Akbas and Erdiller: AMV infections recorded from Turkey. Vigour and yield are reduced.. sativa DC. or the upper part of the blade. Bercks R. The viruses and their taxonomy. . Revue Suisse de Viticulture. Arboriculture. Researches on grapevine virus diseases and determination of their incidence in Ankara Turkiye.I. Monografias INIA No. and J. FAO Publication Division. Investigation on certain viruses spread on grapevines in Bulgaria. Phytopathologische Zeitschrift 76.B. Jankulova M. Control: No information. and G. Rome. has differently shaped particles. Main symptoms: Leaves show bright yellow discolourations that form marginal rings. and a multipartite genome. Alfalfa mosaic virus on grapevine.18 . Jubileum 75.. Description Main synonyms: None. Historical rewiev 1985 Francki: Comprehensive review of the properties of AMV an other viruses with tripartite genome. quasi spherical 25-30 nm in diameter to bacilliform 40 to 75 nm in length. Journal of Turkish Phytopathology 22. 152-154. In: The Plant Viruses. Detection: GLPV is mechanically transmissible to herbaceous hosts. 1976. Identification of alfalfa mosaic virus and tomato bushy stunt virus in hop (Humulus lupulus L. and J. 1973. Horticulture 7. Martelli G. Agent: The putative agent. Detection and Diagnosis of Graft Transmissible Diseases of Grapevines. 1981.B.-J. 1993. Martelli: Yellow mottle suggested as the name for the disease caused by AMV in grapevines. D. 39.P. This disease is known to occur only in Hungary. Several V. References Akbas B. Transmission: GLPV has no known vector. Grapevine disease in Hungary caused by alfalfa mosaic virus infection. roughly following its contour. 1985.I. Erdiller. Graft transmission to cv. Cazelles. and O. New York . Bovey R.) and grapevine (Vitis vinifera subsp. 3rd International Congress of Plant Pathology. Beczner L. Lesemann and G. Varietal susceptibility: No information. 63-65.). 1978.B. 1979. or maple leaf-like line patterns typically confined to the petiolar area. Querfurth.

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

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

and a bipartite singlestranded RNA genome accounting for c. Healthy vines of cvs Kyoho and Pione became naturally infected in the field within one year from planting. Vitis riparia Gloire) are also susceptible. No. 3. 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. Locorotondo 2003. and RT-PCR. Grapevine berry inner necrosis has been reported only from Japan. Varietal susceptibility: No information Detection: Mechanical transmission to herbaceous hosts. the 3' terminal region of which (2469 nts) has been sequenced. representing the most important virus disease in Yamanashi Prefecture. Mavric et al. if any.. Japanese diseases GRAPEVINE BERRY INNER NECROSIS 1. 30 x 103). Historical review 1976 2003 Murant: Description of RBDV. The virus spreads naturally in the vineyards. Some rootstocks (e. F. infected propagative material can disseminate the RBDV..: First record of RBDV in grapevine. Chlorotic mottling. 1976. 113 . 165 B.Agent: Raspberry bushy dwarf virus (RBDV) was isolated from symptomatic vines. Description Main synonyms: None Main symptoms: Infected grapevines have low vigor. Agent: The disease agent is Grapevine berry inner necrosis virus (GINV). wt of 2 x 106 Da (5.8 x 106 Da (2. Koshu. 2003. However. berries are small and show external discolorations and internal necrosis. CMI/AAB Description of Plant Viruses. GINV has filamentous particles about 750 nm in length and a single-stranded RNA genome with Mol. wt 0.5 x 106 Da. Control : No information 2. RBDV. ELISA . Kyoho.2 kb in size). Ripening of bunches is delayed. Zezlina. Raspberry bushy dwarf virus.5 Kb in size) and RNA-2 with Mol. 24% of the particle weight and consisting of two functional species RNA-1 with Mol. Varietal susceptibility: Symptom severity varies with the cultivar.g. Extended Abstracts 14th Meeting of IGVG. delayed bud break and young shoots with short internodes and internal browning. a mechanically transmissible definitive member of the genus Trichovirus. The vay of natural spreading in grapevine. rings and line patterns are shown by leaf blades. Virscek Marn and I. Almost all Japanese table grape cultivars derived from crosses with cv. wt of 7. and Pione whereas cvs. Raspberry bushy dwarf virus infection of grapevine in Slovenia. a dimeter of about 33 nm. Delaware. is unknow. Transmission: GINV is transmitted by grafting to grapevines and by mechanical inoculation to herbaceous hosts. Transmission: In raspberry the virus infects progeny seedlings and pollinated plants through pollen. being transmitted by the eryophid mite Colomerus vitis. and Kaiji are infected latently. M. Campbell Early are suscetible as well as cvs Takao. References Mavric I. 20 Murant A.

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

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

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

VIRUS-LIKE DISEASES .

.

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

Malvasia (10. historical account. Refatti: Hypothesis of a correlation between fanleaf and enation disease. In the vineyards under observation. Enationaffected vines produced less than 50 % of the yield of healthy plants. although the rate of symptom expression does not exceed 30% Credi: Enation diseasae affects the vegetative vigour of cv. Vernaccina (1. Beobachtungen und Untersuchungen über die Enationenkrankheit der Rebe.: Enation disease in Czechoslovakia Avgelis and Xafis: Enation disease in Greece Prota and Garau: Enation disease found in Sardinia.5%). The authors discuss the hypothesis that enation is caused by a strain of fanleaf virus. 1978. Presence of enations in Razaki grapevine in Crete (Greece). attempts to transmit the disease by graft. Martelli et al. The mean yield loss of diseased vines ranged from 17..: In experiments on graft transmission of enation disease aimed at determining the best indicator. 79-112. McGechan: Enation disease in Australia Tekinel et al. with negative results. and possibly caused by a virus. 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 3.1966 Graniti et al.: More data on the effects of enation on the yield of cv. Graniti and Martelli: Review paper on enation. 1968. Prota et al. References Avgelis A. the hybrid LN 33 was found to be the most sensitive and reliable indicator variety. but report on observations made in Australia where no fanleaf virus could be recovered from enation-affected vines. Xafis. Confirmation of graft transmissibility of the disease. 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. only fanleaf virus was recovered. 120 .5%). Marinesku and Bondarchuk: Enation disease in Moldova Brückbauer: Influence of enation disease on growth and yield of grapevine in West Germany.: Enation disease in France Pozdena et al. Mechanical transmission tests were also negative.: Enation disease in Turkey Hevin et al. lowest in cv. Padilla et al. Phytopathologia Mediterranea 17. Weinberg und Keller 15. but diseased vines which had not shown enation symptoms for several years had almost normal yields. Nieder: Enation disease in Austria Garau et al. and C. the proportion of diseased vines was highest in cv. Italia in Sardinia. There is some evidence that enation can be carried in the rootstocks. The possible role of fanleaf virus in the etiology of enation requires further investigation.4 to 48.: Successful transmission of enation disease from diseased to healthy grapevine by graft strongly supports the hypothesis of a viral origin. 195 Brückbauer H. Trebbiano romagnolo and reduces the yield from 13% to 23% according to the severità of symptom expression.3% . The authors conclude that the disease is probably of European origin. .

Pflanzenharzt 36. Ita and F. 59-64. Symptom expression seems to depend on climatic conditions. Agricultural Gazette of New South Wales 81. 1996. Martelli G. Main symptoms: Pale green mosaic affecting mostly the tissues adjacent to the main veins or the smaller ones.W. 1997. 47-64. Nas. California. 251-252 Hewitt W. Lisbon 1997. Occurrence of enation disease in Tunisia. Salih and Y.IV. 7-12.. Cugusi. Bitki Koruma Buletin 11. 169-192. Bondarchuk. Adernmosaik (Germ. Hevin M. A. Extended Abstracts 12th Meeting of ICVG. G. small fruit and grapevine in Moldavia. and R. Deutsches Weinbau-Jahrbuch 31. Tekinel N. Refatti E. Lisbon 1997. Important virus diseases of grapevine in New South Wales. Vanek. 48. 121 .). Die Eantionenkrankheit der rebe erstmal auch in Osterreich nachgewiesen. Lamberti and A. Graniti. Petria 6.P. 145-152. Davis. Einfluss von Virusinfektionen auf Wachstum und Ertrag der Rebe. 122-124. Pozdena J. Monografias INIA 18. M. Spain. 1987. Prota U. 1937. A similar disease has been reported in Australia under the name of summer mottle. 97-98.P. Prota and M. and G. Quacquarelli. areas of the leaf blade may become necrotic. Phytopathologia Mediterranea 5. Lamberti.1989. Martelli.V. Vein mosaic appears to be of low economic importance. McGechan J. H. Israel. A study of infectious degeneration (fanleaf) in vineyards of the Mediterranean region. 179-189. Berkeley.D. Bulletin of the California Department of Agriculture 43. 17.G. 1983.. Mededel. Martelli and F..) Virus and mycoplasma-like diseases of fruit trees. 1970.s. Cordoba. Leclair and M. but these necroses do not affect the veins as it is the case with vein necrosis. Roma. 9. Enation disease of grapevine in Italy. Graniti A. but when fanleaf virus transmission to herbaceous hosts became possible. 46-51. 207-217. Garau. 1891. Facultet Landbouwwetenschap (Gent) 40. with Special Reference to Vitis. 2. Phytopathologia Mediterranea 20. In: Verderevskaya T. Effetto della malattia delle enazioni della vite sulla produzione e sullo sviluppo vegetativo della cv Trebbiano romagnolo. Extended Abstracts 12th Meeting of ICVG. O. IV. Frazier N. Moldavian Research Institute of Horticulture Kishinev. 1954. 47. In: Virus Diseases of Small Fruits and Grapevines (A Handbook). Description The symptoms of vein mosaic have been confused for some time with those of fanleaf/yellow mosaic. Proceedings 9th Meeting of ICVG. Padilla V. (N. Cugusi. (ed. 1971. Dolar. Su una possibile correlazione fra il virus del complesso dell'arricciamento e la malattia delle enazioni della vite. 349-352. and V. Garau R. Proceedings International Conference on Virus and Vector on Perennial Hosts. Division of Agricultural Sciences. 293-306. University of California. Ser. Studies on reproduction of enation symptoms by grafting in Sardinia. Buchenau F.S. 326332.. Grapevine enation disease in Murcia (Spain). F. n. 1997. Salcan. In the most sensitive varieties. 1981.. B.Brückbauer H. Mosaico delle nervature (Ital. Garcia. Enations of grapevine in Sardinia.P.W. Marinesku V. Gazeau. Some virus and virus-like diseases of grapevines. Occurence of grapevine enations in the Moldavian SSR. This disease is widespread and probably worldwide. Z.). Graniti A. G. Credi R. Enations. Benayas.. Rivista di Patologia Vegetale. Savino. Prota U. Studies on some variable characters of grapevines affected by enation disease in Sardinia.. 1973. producing often a vein banding effect. 1976. 1966.P.. Gigante R.. Kiryat Anavim..K. 823-827. Bericht der deutschen botanischen Gesellschaft 9. Abnorme Blattbildungen. 1979.. 1970. Nieder G. Garau R.). 1966. Rivista di Patologia Vegetale S. VEIN MOSAIC 1. Bolletino della Regia Stazione di Patologia Vegetale. and M.. 1983.. Rives. 241-243. U. Ricerche istologiche sulle omeoplasie crestiformi (Enations) delle foglie di vite affette da rachitismo. Proceedings 6th Meeting of ICVG. 1975. 225-246. Enation symptoms found in France. Main synonyms: Mosaïque des nervures (Fr. 1966. I. And G.. it was clear that vein mosaic was not caused by fanleaf virus. 1980. The research of the virus diseases of grapevine in CSSR.B. Chabbouh N and V.). Trasmissione per innesto della "malattia delle enazioni" della vite. 1965. 203-206.

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

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

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

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

.

VIROID (Yellow speckle) .

.

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

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

131 . Grapevine yellow speckle viroid 2 (GYSVd 2) Australian grapevine viroid (AGVd). Savino. Krake. J. 1991 1996 1997 1999 2001 2003 2003 3. American Journal of Enology and Viticulture 39.M. Sano et al. A possible virus disease of Rcatzitelli vines in Bulgaria. Molecular evidence that this viroid originated from recombination between five different viroids among which GYSVd-1 and GYSVd-2 1991a. 1991 Semancik and Szychowski: There are two classes of grapevine viroids: (i) apparent viroids. Barlass M. Annals of Applied Biology 101. Greece. Wang et al. are to be named as follows: Hop stunt viroid grapevine strain (HSVd-g).1988 1989 1989 1989 1990 1990 Duran-Vila et al. A.: Structural analysis reveals that five distinct viroids infect commercial grapevine varieties. Koltunow et al. V. First report of Australian grapevine viroid from the Mediterranean region. GYSVd-1 and GYSVd-2 Rezaian: Complete nucleotide sequencing of AGVd. Fakhfakh. R.G.P. K. Rezaian et al. Duran-Vila.C. Woodham and L. Quacquarelli and V.M. Martelli. J. 291-295. M. Journal of Plant Pathology 85.. 45-57. The occurence is reported of HSVd.: Evidence that two related viroids (GYSVd 1 and GYSVd 2) can cause yellow speckle disease independently. 2003. Little and Rezaian: Updated review of grapevine viroids. Minafra et al. Grapevine yellow speckle viroid 1 (GYSVd 1).R. Duran-Vila N. 1988.: Review of viroids. Marrakchi.b Szychowski et al. according to an international agreement reached during the 10th Meeting of ICVG held in 1990 at Volos.. 127-130.: First report of AGVd in the Mediterranean.: First record of grapevine viroids in China. N. which require amplification in an alternate host. Production of viroid-free grapevines by shoot tip culture. Martelli: Brief review of grapevine viroid situation supporting the idea that vein banding is primarily induced by viroidal rather than GFLV infection. Juarez and J. Elleuch et al. Perreault and H. CEVd-g and AGVd via grape seeds. These viroids. 1982. based on phylogenetical analysis of hop and grapevine isolates of this viroid. G. Arregui. Monografias INIA 18. (ii) enhanced viroids. 1985. Journal of General Virology 66. 217-220. which can readily be isolated directly from grapevines. GYSVd-2. Semancik.: A survey of viroids of grapevine in Italy. References Abracheva P.P. Flores R.. Proceedings of the 6th Meeting of ICVG..: Improvement of meristem tip culture technique for the production of viroid-free grapevines.: Extensive comparative analysis of grapevine accessions from California and Europe reveal a similar pattern of viroid distribution. Flores et al. 1978. Koltunow and Rezaian: Description and sequencing of grapevine viroid 1B (later renamed Grapevine yellow speckle viroid 2). Detection of viroid and viroid-like RNAs from grapevine. Skene.. Elleuch A. 2095-2102. Regeneration of virus-free grapevines using in vitro apical culture. Pallas and J. Spain. Wan and Symons: Transmission of GYSVd-1. Citrus exocortis viroid grapevine strain (CEVd-g).: Suggestion that the viroid causing stunting in hop (HSVd) originated from grapevines.S. Cordoba. 1976.

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

YELLOWS .

.

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

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

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

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

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

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

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

Experiments on the use of hot water treatment on planting material. Maixner (a. So. Transmission has been obtained only from vines of Veneto. Electron microscopic observation of MLO in periwinkle and grapevine.: Six-year transmission trials of different types of yellows with S. Kuszala et al. Positive assays obtained only on samples from France and Veneto. 1993 Daire et al. Di Terlizzi et al. titanus is not present. titanus is present in the Friuli-Venezia Giulia region but no transmission assay has been successful. Specific probes cloned and specific primers designed for PCR amplification of phytoplasma DNA. 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). FD belongs to the elm yellows group but is different from elm phytoplasmas. riparia in New York and failure of detection in some symptomatic grapevines in New-York and Italy. Girolami and Egger: Importance of contamination of vineyards and diffusion of Grapevine yellows in northern Italy. titanus) and a second one. MLO appear to be in very low titre. BN belongs to the stolbur group. probably transmitted by another insect.called FDU. at least two different phytoplasmas are infecting grapevines in Veneto: FD sensu stricto (transmitted by S. Subcommittee on the Taxonomy of Mollicutes: The trivial name MLO is replaced by the term phytoplasma as a consequence of the demonstration by diverse studies that the phytoplasmas represent a monophyletic clade of organisms more closely related to mollicutes than to walled bacteria. then GYU phytoplasma) and comparison of specificity and sensitivity of the latter tools to detect phytoplasmas in grapevines in New York and Italy. Computer analysis for spatial patterns of diffusion show a non-random distribution and suggest transmission from other vines or from weeds. Description of pathological changes in phloem of leaves and identification of senescent forms of MLO inside degenerate sieve elements. graft and dodder. (a and b): Various attempts to transmit FD-like disease in different regions of northern Italy. Maixner: Demonstration that Hyalesthes obsoletus (Auchenorrhyncha: Cixiidae) is a vector of German grapevine yellows (Vergilbungskrankheit). In northeastern Italy. healthy plants protected with plastic screens did not develop symptoms and diseased plants protected with screens showed transient recovery. BN (stolbur MLO) detected in several regions of Italy and in Israel. 1993 1993 1993 1993 1993 1993 1994 1994 1994 1994 1994 142 .: FD-ELISA method was used on tissues of yellows-affected grapevines from diverse countries. Credi (a and b): Observation of MLO in yellows-affected grapevines in northern Italy.: Diversity of MLOs associated with grapevine yellows and transmitted to periwinkle. FD detected in leaves of rootstock 3309C and is also present in samples from Veneto (Italy). (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. S. Carraro et al. Only FD and BN have been identified in naturally affected grapevines. GYU was later on characterized by other authors as a phytoplasma belonging to the Xdisease (16SrIII) group. International Committee of Systematic Bacteriology (ICSB). using insects. Detection in grapevines from Virginia (USA) of MLO related to X-disease. aster yellows and Xdisease. b. titanus to test plants of cvs Perera and Chardonnay in Veneto and Friuli-Venezia Giulia (Italy). Detection in non symptomatic V. Daire: PhD thesis on detection and differentiation with DNA-based methods of the MLO agents of GY diseases in France. S.: Important outbreak of a yellows disease on local varieties in Apulia (southeastern Italy) and infection of periwinkle as bait plants and with dodder transmission. Osler et al. c and d): Grapevine yellows (Vergilbungskrankheit) are common in the Moselle and Rhine valley. FD is related to elm yellows and BN is related to stolbur. Prince et al. on the effects of pollarding affected vines and of chemical sprays against vectors. Classification of grapevine MLOs into three RFLP groups: elm yellows (FD from France). A MLO has been transmitted to periwinkle with dodder.

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

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

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

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

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

.

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

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

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

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).

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

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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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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

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

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

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

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

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

titanus leafhoppers have failed. Transmission to plants is not reported. Sabaté et al. Description Palatinate grapevine yellows (PGY) is a disease occurring in Germany in Pfalz and Moselle regions (west of Germany) mainly on cv Scheurebe. Cicadellidae) trapped on alders and caged on V. affected plants are not numerous. the psyllid Psylla alni. According to the data of field survey. PALATINATE GRAPEVINE YELLOWS 1. using 4 fragments of phytoplasma DNA showed that all isolates are similar. (b): First identification of BN infection in the Macedonian republic. vinifera seedlings that later developed GY symptoms. Main symptoms: Symptoms are similar to those of other GY diseases and may affect the whole plant. -B and C) have been identified.: Important expression of BN in cv Chardonnay in the Ukraine. The latter results have not been published.: Importance of a GY epidemics in the Drava region (Slovenia) in 2001 and 2002.2003 2003 Petrovic et al. obsoletus is rare. together with high populations of H. Living insects checked for transmissibility of phytoplasma by feeding on artificial medium. Varietal susceptibility and sensitivity: PGY occurs in limited areas. Langer and Maixner: Three different types of stolbur phytoplasma isolates found in grapevines. Transmission: Transmission was obtained with the leafhopper Oncopsis alni Schrank (Hemiptera. Another alder insect. occur most often at the border of plots. Control: No possibility of control because of erratic transmission by the vector.: Important epidemics of "Schwarzholzkrankheit" (SHK = VK) in Franken (Bayern.) Detection: Universal primers and RFLP analysis of amplicons can be used to identify a 16SrV phytoplasma. It is not epidemic and occurs most often in old vines in vineyards along creeks or rivers where alders (Alnus glutinosa L) are present. specific association of each isolate with a different weed is discussed. Gilge et al. arvensis and U. The disease was identified in 1995. alni is highly monophagous and specimen do not survive long on grapevine. Symptoms were recorded on 26% of 30-year old Scheurebe grapevines and 96% of tested individuals of H.: Search for vectors of stolbur phytoplasma in Cataluña (Spain) where H. Milkus et al. Šeruga et al. However. Trapping of insects on sticky traps or with D-Vac suction. Attempts to transmit PGY phytoplasma from grapevine to grapevine using reared S. 2003 2003 2004 2004 2004 C. dioica. Germany). and rarely in groups. obsoletus and frequence of C. Alder is considered as the source of PGY infection. obsoletus and weeds in different areas in Germany. (a): Investigation on the variability of stolbur (16SrXII) phytoplasmas from extracts of different grapevines in Croatia. obsoletus were positive for stolbur phytoplasma. 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). mainly on cv Scheurebe. Several hemipters have been found to deliver stolbur phytoplasma to the medium. It is different from FD phytoplasma. It is associated to phytoplasmas in the Elm yellows (16SrV) group. O. Other host plants: Alder trees (Alnus glutinosa L. alni but failed to transmit both to alder and grapevine. Šeruga et al. Agents: The agent is an EY (16SrV) group phytoplasma. Main synonyms: FD-Pfalz was a first name because the agent was shown to be related to the FD phytoplasma. 164 . 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. PGY phytoplasma is similar to the Alder yellows phytoplasma that is widespread in black alder in Germany. Three isolates (PGY-A. was found carrying the alder phytoplasma at a higher rate than O. H.

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

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

is a member of the aster yellows (16SrI) group. It belongs to the stolbur group (16SrXII) but is distinct from the BN and VK phytoplasmas. riparia vines. BVGY phytoplasma has not been clustered in an established phytoplasma group. The use of PCR has shown the association with AGY of three different phytoplasmas. The relatedness of the associated MLO to X-disease MLO is confirmed. The second phytoplasma. which is the more frequent and the Tomato big bud (TBB) phytoplasma. The association of AGY with phytoplasmas was confirmed as early as 1988. Late Season Leaf Curl (LSLC). consistently associated with AGY symptoms and AGY phytoplasmas but their aetiology is not clear. respectively. Restricted Growth (RG). AUSTRALIAN GRAPEVINE YELLOWS 1. The TBB phytoplasma has been designated as Candidatus species "Candidatus Phytoplasma australasia". with a higher incidence in warmer inland districts of New South Wales and South Australia. 167 . Some plants also display uneven bud development resulting in canes and cordons that are bare in places with little or no bunch development. The yellow leaf tissue can become necrotic. Conversely. In this particular disease. 1998 2003 E. 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.: Two phytoplasmas are detected in yellows diseased grapevines in Virginia. Both VGYI and VGYIII were found in several weeds and shrubs around and inside the vineyards. suggesting that the source is from an alternative host. Stems of affected shoots develop a blue. also detected in wild V. subgroup I-A. waxy appearance and remain rubbery. Main symptoms: All symptoms typical for GY can be found on AGY-diseased plants. Early and recent surveys have shown that AGY are found in most viticultural regions of Australia. Beanland and Wolf: Search for possible insect vectors and reservoirs of NAGY phytoplasmas in Virginia. there are reported cases where they were not associated with AGY disease or phytoplasmas. Description Grapevine yellows were first reported from Australia in 1976 and called Australian grapevine yellows (AGY) in 1983. The 16S rRNA gene has a high sequence similarity (more than 99. Mapping of diseased plants show that grapevines appear to be a terminal host of AGY. It belongs to the peanut witches' broom group (16SrII) and has a broad plant host range in Australia. Late Season Leaf Curl (LSLC)-affected grapevines show symptoms in late summer with tightly downward rolled leaves that often remain green.: A GY disease observed in Virginia since 1983 affects large areas with a low incidence and a low spread rate. Confirmation that a X-disease (16SrIII) group phytoplasma is involved and classification of the latter phytoplasma into a new subgroup. In addition.5%) with the phytoplasmas associated to Papaya die back (PDB) in Australia and to Phormium yellow leaf (PYL) in New Zealand. The AGY phytoplasma has been designated as Candidatus species "Candidatus Phytoplasma australiense". Restricted Growth (RG)-affected grapevines show retarded growth resulting in shortened shoots and smaller leaves. Shoots may display abortion of bunches or berry shrivel later in the season and death of terminal buds. two syndromes with particular symptoms (Restricted Growth and Late Season Leaf Curl) have been found in some cases. Davis et al. Other species tested positive and transmitted to feeding solutions. one node after the other. followed by the progressive death of the shoots. It has been suggested that all three phytoplasmas are strains of the same phytoplasma species. leaves tend to fall early. III-I. Buckland Valley Grapevine Yellows (BVGY). Agallia constricta is a suspected vector of VGYI (aster yellows phytoplasma).1993 Wolf et al. Main diseases: AGY (sensu stricto). a GY disease found in Victoria (Australia). clover phyllody phytoplasma (16SrI-C) being the closest . overlay one another like shingles and become leathery and brittle. Agents : AGY disease (sensu stricto) is associated with two phytoplasmas: the AGY phytoplasma. The sequence similarity of the 16S rRNA gene shows that it is close to members of the aster yellows group (16SrI).

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

: BVGY phytoplasma is associated to GY symptoms that may show remission then reoccurrence but also appear on vines that were not affected previously. later called BVGY phytoplasma.: Study of the phylogenetic relationships of the BVGY phytoplasma with aster yellows and stolbur phytoplasmas.: Use of hot water treatment in commercial nurseries of Australia. RG. Papaya dieback and Phormium yellow leaf diseases. BVGY might form a new AY subgroup of AY (16Sr) group or even a new phytoplasma group.: The AGY phytoplasma is consistently present in the majority of symptomatic grapevines. Liefting et al. RG and LSLC are associated. (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). Detection of BVGY in another plot in the same area. 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.: No vector for AGY found. The TBB phytoplasma is detected occasionnally in vines with AGY symptoms and also in LSLC-affected vines. but different from the stolbur phytoplasma associated with BN in France. Constable et al.: Description of the AGY phytoplasma and designation of a new taxon.: Comparison of visual assessment and diagnostic assays. Spain and Israel. Hence. 1997 1997 1997 1998 1998 1998 1999 1999 1999 2000 2000 2001 2001 2002 2002 2002 169 . Candidatus Phytoplasma australiense. Padovan et al.: Identification of a new phytoplasma in Chardonnay grapevines in Victoria (Australia). insecticide sprays are useless for controlling the disease. Constable et al. Gibb et al.: Detection of phytoplasmas associated with AGY. Habili et al. Spring and summer are optimal for detection.: Further comparison of AGY phytoplasma with overseas GY phytoplasmas show that it is closely related to. Beanland et al. It is occasionnally detected in symptomless vines and in vines with LSLC or RG symptoms.: AGY phytoplasma detected in the body of Orosius argentatus leafhoppers. Bonfiglioli et al. Padovan et al.: Close relationships between phytoplasmas associated with AGY. Uneven distribution of phytoplasmas in infected plants. LSLC would be followed by AGY on the following year and AGY would be followed by RG in subsequent years. (a and b): Positive detection of phytoplasma with PCR in AGY-affected grapevines and tentative survey of phytoplasmas in Australia. The problem of symptomlessly infected vines. Constable et al.: Description of AGY symptoms. Wilson et al. Kelly et al. Waite et al.1995 1995 1996 Bonfiglioli et al. Davis et al.: Assumption from field observations and monitoring that AGY. and LSLC. established from a different source of plant material indicate that BVGY disease occurs as a result of aerial transmission.: TBB phytoplasmas can be acquired from diseased grapevine by Orosius argentatus and subsequently transmitted to faba bean. resulting in increasing cumulative incidence. Constable et al. Beanland et al.

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

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

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