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Options Méditerranéennes Série B n.55 Directory of Infectious Diseases of Grapevines and Viroses and Virus-like Diseases of the Grapevine: Bibliographic Report 1998-2004
Opinions, data and facts exposed in this number are under the responsibility of the authors and do not engage either CIHEAM or the Member-countries.
Les opinions, les données et les faits exposés dans ce numéro sont sous la responsabilité des auteurs et n'engagent ni le CIHEAM, ni les pays membres.
Martelli.CIHEAM Centre International de Hautes Etudes Agronomiques Méditerranéennes Options méditerranéennes Directeur de la publication: Bertrand Hervieu Directory of Infectious Diseases of Grapevines and Viroses and Virus-like Diseases of the Grapevine: Bibliographic Report 1998-2004 Editors: G.P. Boudon-Padieu 2006 . E.
Série B : N.September 2006 tel. 55 ISSN : 1016-1228 ISBN : 2-85352-338-1 © CIHEAM. 279 Options Méditerranéennes. Martelli. +39 080 542 45 87 e-mail: ideaprint@virgilio. Boudon-Padieu Bari : CIHEAM (Centre International de Hautes Etudes Agronomiques Méditerranéennes) p. 2006 Reproduction partielle ou totale interdite sans l'autorisation d'«Options Méditerranéennes» Reproduction in whole or in part is not permitted without the consent of «Options Méditerranéennes» .it Directory of Infectious Diseases of Grapevines and Viroses and Virus-like Diseases of the Grapevine: Bibliographic Report 1998-2004 Editors: G. Italy.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. E.
1998-2004 INTRODUCTION SUBJECT INDEX REFERENCES 7 11 15 17 33 49 59 76 93 99 107 117 127 133 149 173 195 203 205 207 219 5 .TABLE OF CONTENTENTS PREFACE INTRODUCTION INFECTIOUS AGENTS OF GRAPEVINES INFECTIOUS DEGENERATION (GRAPEVINE FANLEAF VIRUS) INFECTIOUS DEGENERATION (EUROPEAN AND MEDITERRANEAN NEPOVIRUSES) GRAPEVINE DEGENERATION AND DECLINE (AMERICAN NEPOVIRUSES) GRAPEVINE LEAFROLL RUGOSE WOOD COMPLEX GRAFT INCOMPATIBILITY FLECK COMPLEX MINOR VIRUS DISEASES VIRUS-LIKE DISEASES VIROIDS GRAPEVINE YELLOWS: GENERAL PROPERTIES GRAPEVINE YELLOWS: INDIVIDUAL DISEASES GRAPEVINE YELLOWS: REFERENCES PIERCE'S DISEASE THE VIROSES AND VIRUS-LIKE DISEASES OF THE GRAPEVINE: BIBLIOGRAPHIC REPORT .
and practitioners. grouping diseases and setting them in a well thought out order. it is most unfortunate that dissemination of infectious diseases continues. As to the authors. 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. students. so as to produce a document which can readily be used by scientits.P. 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. to all those interested in viticultural matters. except when virus-free vines are grown for comparison alongside with non-sanitized sister vines. whereas E. promoting the publication of the Bibliografic Report 1985-1997 on virus and virus-like diseases of the grapevine. these diseases cause loss of vigour and often a decline of affected stocks. in the belief of rendering a good service to the scientific community and. they reduce graft compatibility of scions and rootstocks. or induce subtle debilitating effects which are difficult to percieve. produced under the auspices of ICVG. G. Cosimo Lacirignola Director of IAM Bari. that fosters coperative studies in grapevine virology. whose publication IAM-B is happy to support.PREFACE Virus. 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. hosts a new edition of the “Directory of Infectious Diseases of Grapevines” by G. 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. Boudon-Padieu has a long lasting experience in the study of phytoplasmas. Bovey The “Directory” is a work of great thoroughness and accuracy. virus-like and phytoplasma diseases of grapevines constitute a major limiting factor to the development and well-being of the world viticultural industry. 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. The present issue of Options Méditerrnéennes. technicians. which reflect on the commercial value and productive life of the vineyards. Sincere thanks are espressed to the authors of these contributions. to which IAM-B was happy to contribute. Furthemore. although the technology is now available for detection and elimination of most of the pathogens occurring in propagation material. being a recognized authority in this field. ICVG has a long-lasting tradition in these endeavours. Bovey.P. Improving the sanitary conditions of the industry. Italy 7 . As a whole. is therefore a goal of utmost importance to which the Mediterranean Agronomic Institute of Bari (IAM-B). a distinguished virologist of worldwide reputation and one of the father founders of ICVG. whose Secretatiat he has admirably conducted since its very establishment in 1962. Notwithstanding these efforts and the promoting activity of the International Council for the Study of Virus and Virus-like Diseases of the Grapevine (ICVG). up to date as to the time of publication. and to the quality and quantity of the crop. Martelli and E. by and large. taking also into account the future role of the Mediterranean as a free-trade area. The “Bibliographic Report” is another most precious achievement by R. Special care was taken in the organization of the subject matter.
France 2006 . Martelli University of Bari. Boudon-Padieu INRA Dijon.International Council for the Study of Virus and Virus-like Diseases of the Grapevine (ICVG) DIRECTORY OF INFECTIOUS DISEASES OF GRAPEVINES G. Italy E.P.
INTRODUCTION The grapevine (Vitis spp. Infected propagating material is largely responsible for the spread of diseases among and within viticultural countries. grapevines are exposed to the attacks of a variety of pests and pathogens among which infectious intracellular agents (viruses. Bovey R.. 3rd part). 2006. has fostered intensive research.. nurserymen. growers. Vitis 18. The papers up to 2004 are listed and commented upon in six bibliographic reports: Caudwell A. Hewitt and R. 2003). and R. 2003. ICVG has met regularly every 3 to 4 years. A short history of ICVG. and phytoplasmas) recorded from the grapevine can be highly detrimental to this crop. viroids. 10-172. 1-2. W. 205-279. To this effect.. and endangering the survival of affected vines. Paris. and G.B. and P. 29 (Series B. viroids. Bibliographie des viroses de la vigne des origines à 1965.P. attracting the attention of scientists. ICVG has been instrumental in fostering basic and applied research in grapevine virology. causing heavy losses. and a highly valuable agricultural commodity. Caudwell A. has produced an impressive series of papers which now number about 5. 1965. A bibliographic report 1985-1997. having a negative impact on the plant vigour and longevity. as well as on the quality and quantity of the yield. Thus. Bovey R. shortening the productive life of vineyards. and supporting initiatives for the establishment and implementation of clean stock and certification programmes. ICVG has issued the recommendations that follow: The International Council for the Study of Virus and Virus-like Diseases of the Grapevine (ICVG).400. A bibliographic report 1979-1984. Since its foundation. As most of the vegetatively propagated crops. Bibliographie de 1965-1970. 1979. among other things. The increased attention paid to grapevine virological problems and the like.) undoubtedly represents one of the horticultural crops most widely grown in temperate climates. ICVG was established in 1962 by a group of American and European plant pathologists who realized the importance of creating an organization for promoting research on grapevine virology and favouring the exchange of information among students. Bovey R. Extended Abstracts 14th Meeting of ICVG. that has been especially active at the international scale from the late 1950's onwards. Gugerli. Bovey R. A bibliographic report 1971-1978. recognizes that a number of the 60 or so infectious agents (viruses. 1972. Les virus de la vigne. 227-275.B. The viroses and virus-like diseases of the grapevine. 303-324. 1999. Options Méditerranéenes. all efforts should be made to improve its sanitary conditions. The importance of the grapevine industry and the magnitude of the problems caused by these pathogens has generated wide interest which. Martelli. Locorotondo 2003. The viroses and virus-like diseases of the grapevine. Bovey. Vitis 25. 316-376. Bovey. 1986. and administrators on the detrimental effects of infectious diseases on the well-being of the industry. 76 pp. phloem-and xylemlimited prokaryotes) play a major role. 3rd part). which were compiled under the auspices of the International Council for the Study of Viruses and Virus Diseases of the Grapevine (ICVG). Office International de la Vigne et du Vin. A bibliographic report 1998-2004.. its 14th Conference having taken place in Italy in 2003 (Bovey and Gugerli. From the very beginning. Hewitt W. Vitis 11. The viroses and virus-like diseases of the grapevine. xx (Series B. in turn. The viroses and virus-like diseases of the grapevine. Options Méditerranéenes. 11 ..
is regarded as incompatible with an accepted sanitary status. (a revised edition by J. Uyemoto will be published in 2005).A. Tolin. leafroll.P. In addition.G.C. recognises over 70 infectious agents affecting grapevine (viruses. and phytoplasmas (flavescence dorée. Until that time. and fleck. rugose wood. Certification of grapevine nursery stock is a powerful and effective tool to control these agents. high standards are essential to ensure that no viticultural area is compromised by the introduction and spread of diseases.C.W. Hewitt. Paul. Italy) The Proceedings of all the international ICVG Conferences have been published in extenso or. 1978. Barnett and S. that enables vineyards to economically and sustainably maintain quality and productivity.P. lately. Virus-like Diseases and Other Disorders).K. 12 . Minnesota. Martelli and A. 1988. G. In order to preserve valuable grape clones and varieties. (issued and approved in 1997 at the 12th ICVG Meeting. Class 1 should include only grape nursery stock which tests negative for the most damaging diseases/pathogens. (a 2nd revised edition edited by W. Bovey R. Inadequate certification standards have repeatedly resulted in disease problems for growers and nurserymen. Clemson. rugose wood (GVA. The certification process should specify conditions to prevent and detect subsequent infection of nursery plants by regulated pests. (issued and approved in 2003 at the 14th ICVG Meeting. clonally selected primary sources.D. Virus and Virus-like Diseases of Grapevines. Goheen. W. all efforts should be made to improve its sanitary conditions. Uyemoto. and permit tracing the certified grapevines to the originally selected and tested plants. 181 pp. the virological problems of grapevines have been extensively treated and illustrated in a number of books and major review articles: Uyemoto J. viroids and phytoplasmas). and other grapevine yellows). Dias. In the future. Grape Section. The agents that should be controlled by the Class 1 certification program are those associated with infectious degeneration and grapevine decline (nepoviruses).K. As efforts are made to harmonize grapevine certification protocols. Portugal ) The International Council for the Study of Virus and Virus-like Diseases of the Grapevine (ICVG). Rowhani will be published in 2005 in the series APS Plant Virus Image CD-ROM. R.The presence of diseases such as infectious degeneration. Lausanne. we propose two sanitary classes. Lisbon. 2. USA. which are best performed in the framework of certification programmes encompassing also clonal selection.F. No other stock should move outside regulatory regions. a moratorium will be established for these viruses. Gärtel. 29 pp. Plant Virus Slide Series. bois noir. Martelli and A. Martelli. many of which can be highly detrimental to this crop. The regional certification standards for Class 2 stock should be created at a local level based on the rate of endemic infection. valuable grape genetic resources exist which are infected with virus but are essential to the preservation of world viticultural heritage. leafroll disease and associated closteroviruses (grapevine leafroll associated viruses 1. including the causal agents of fleck and rupestris stem pitting. Certified grapevines are derived from pathogen tested. St. GVB and GVD). and 3).K. Woodham. 93 pp. Goheen and H. Gubler and J. G. A. However. Thus. W. G.. Grapevine (Vitis) virus and virus-like diseases. as well as on the quality and quantity of the yield. and the need for preservation of heritage germplasm. technology should make it possible to exclude additional disease-causing viruses from the certified stock. Vuittenez. Certified selections should be tested for specific pathogens. Editions Payot.P. Their elimination from mother vines intended for propagation should therefore be pursued. Compendium of Grape Diseases. having a negative impact on plant vigour and longevity.F. They represent a most valuable source of information.B. ensure clonal integrity. It would move freely between regulatory boundaries. and A. Infected propagation material is largely responsible for the spread of diseases among and within viticultural countries. as Extended Abstracts. Locorotondo. The American Phytopathological Society Press. 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. Improvement of the sanitary level can be achieved through selection and sanitation. 100 slides. Wilcox.C. under the name of Grapevine Viruses. Set 1 (O. W. 1980. Pearson R. eds) Clemson University. regional viticultural conditions.
P. Bovey and G. St. Sélection sanitaire de la vigne: sélection sanitaire et sélection pomologique. Martelli and E. Taylor. Maladies à Virus. N. C. (ed). FAO Publication Division. M. 225 pp. Martelli.P. Protocols for detection of viruses and virus-like diseases: Les Colloques. Bordeaux. Boudon-Padieu and M. Martelli G. Ridé. Walter B. 192 pp. a body now estinguished. Rome. Walter B. Elisabeth Boudon-Padieu Directeur de Recherches Head Biologie et Ecologie des Phytoplasmes Plante Microbe Environnement INRA . n° 86. Rezaian and R. Boudon-Padieu. INRA Editions. Director of the Mediterrenean Agronomic Institute of Bari for supporting the publication of this work. 1998. 1997. bactéries et phytoplasmes de la Vigne. 1999.). Scott. 54 pp. Martelli and published under the auspices of the Mediterranean Fruit Crop Improvement Council (MFCIC).. the Steering Committee of ICGV decided to update the Directory and to entrust this task to G. Khetarpal. 945-971. of the Centre International des Hautes Etudes Agronomiques Méditerranéennes. Ikin. Editions Féret. FAO/IBPGR Technical Guidelines for the Safe Movement of Grapevine Germplasm. 2000. In: Plant Virus Disease Control (A.P. and G. Paris.Frison E. 5-23.P. 261-276.S.137 pp. which was established in the framework of the International Project RAB/88 sponsored by the United States Development Programme and the Food and Agriculture Organization of the United Nations. H. eds. Italy. Koganezawa.R.CNRS .K. 2ere partie: Sélection sanitaire. CSIRO Publishing. Chairman of the Board of Directors and Secretary General. Lacirignola. Bulletin de l'OIV 70. American Phytopathological Society Press. Hamze and Mr. Influence des viroses et qualité. Collingwood. 263 pp. Krake L. and to Dr. Rome /International Board for Plant Genetic Resources. and R. At the 14th ICVG Meeting. 1996. 1997. B. The authors hope that this endeavour may serve as a useful guideline and working tool for both experienced researchers and those who are now approaching the intriguing but intricate field of grapevine virology. 1ere partie: Effets des viroses sur la culture de la vigne et ses produits.A.P. Graft-transmissible Diseases of Grapevines. University of Bari. respectively. Influence des viroses et qualité. Martelli Professor of Plant Virology. and B. Hadidi. Walter B. 1991. Bulletin de l'OIV 69. Sanitary selection of the grapevine. (ed. Walter. Paul. Rome.). M. Food and Agriculture Organization of the United Nations. Paris. Walter B. 1993.. Hervieu. and G. Notwithstanding this wealth of published information a "Directory of Major Virus and Virus-like Diseases of Grapevines" was compiled in 1992 by R. Bari. Detection and Diagnosis of Graft-transmissible Diseases of Grapevines.P.Université de Bourgogne Dijon. Martelli G. Giovanni P. sélection pomologique. They express their deep gratitude to Prof. Sélection sanitaire de la vigne: sélection sanitaire et sélection pomologique. France 13 . Martelli.A.H. E. R. Virus certification of grapevines.
phytoplasmas (8). This represents the highest number of intracelluar pathogens ever found in a single crop. viroids (5). and insecttransmitted xylematic bacteria (1) have been recorded form grapevines. 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 .INFECTIOUS AGENTS OF GRAPEVINES More than 70 infectious agents among viruses (58). The viral scenario of Vitis: viruses and their taxonomic affiliation(a) FAMILY GENUS SPECIES A.
. Virus Taxonomy. The updated taxonomy of all classified grapevine viruses can be found in: Fauquet. J. L. C. London. and Ball. The names of tentative species are written in Roman characters.A. Maniloff. M.M. 8Ith Report of the International Committee on Taxonomy of Viruses.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. Mayo. Elsevier/Academic Press. 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 . Viruses belonging to unassigned genera Idaeovirus Sadwavirus Sobemovirus Tobamovirus C. 1258. pp. Desselberger.A. (eds). U. 2005..
INFECTIOUS DEGENERATION .
Leaves are variously and severely malformed. arricciamento. The consensus is that fanleaf degeneration may have existed in the Mediterranean Basin and the Near East since the earliest time of grape cultivation. showing abnormal branching. Their economic impact varies with the tolerance of the cultivar to the viruses. Now the disease is known to occur worldwide. phloem. especially in the basal internodes.). Reisigkrankheit (partly). and xylem cells. Trabeculae. and inflorescences). asymmetrical. deep lobes. Description Fanleaf is the oldest known and one of the most important and widespread virus disease of the grapevine. puckered. Shoots are also malformed. that give the leaf the appearance of an open fan. and berries ripen irregularly. tendrils. Gelbmosaik (Germ. vein banding symptoms were shown to be caused by a co-infection by Grapevine yellow speckle viroid and GFLV. Yellow mosaic is induced by chromogenic virus strains. and acute denticulations. and decreased resistance to adverse climatic factors. The characterizing symptoms of "Vein banding". low proportion of graft take. sometimes appearing as rings or lines to extensive mottling of the veins and/or interveinal areas to total yellowing . 19 .INFECTIOUS DEGENERATION (GRAPEVINE FANLEAF VIRUS) Several nepoviruses infect grapevines in Europe and the Mediterranean area. Symptomatic leaves show little malformation. causing degenerative diseases whose symptoms are similar to. i. mosaico giallo.). Occasionally. however. panachure.e. short internodes. Often infected grapevines occur in patches. dégénérescence infectieuse (Fr. records of this disease date back some 150 years. This disorder was first reported from California as a syndrome elicited by a specific strain of GFLV. The foliage and shoots show little if any malformation. which exhibit widely open petiolar sinuses and abnormally gathered primary veins. urticado (Port. reduced rooting ability of propagation material. roncet. bunches are straggly. Main symptoms: Two distinct syndromes caused by different strains of the causal agent characterize this disease. or endocellular cordons. degenerazione infettiva (Ital. shortened productive life. another disease sometimes occurring in vineyards affected by infectious degeneration. Chromatic alterations of the leaves vary from a few scattered yellow spots. Tolerant cultivars produce fairly good crops whereas sensitive cultivars are severely affected. Infectious malformations are induced by virus strains causing distortions. are small-sized and set poorly. FANLEAF 1. GFLV and several of the other grapevine-infecting European nepoviruses have distorting and chromogenic strains and may occur in mixed infections. may show open petiolar sinuses. Main synonyms: court-noué. the yellowing fades rapidly and the canopy develops a normal green color. Foliar symptoms develop early in the spring and persist throughout the vegetative season becoming less distinct in summer. Bunches are smaller and fewer in number. radial bars crossing the lumen of epidermal. low yields and low fruit quality. a disorder induced by Grapevine fanleaf virus (GFLV). parenchyma. showing progressive decline. double nodes. With increased ambient temperatures during summer. fasciations. and grapevine leaves with typical symptoms were found in herbaria established before the introduction of American rootstock hybrids. The foliage develops bright chrome yellow discolorations early in the spring that may affect all vegetative parts (leaves. 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. shoots. More recently. are diagnostic of grapevines infected by GFLV. The name of this virus comes from the peculiar malformation of infected leaves. chlorotic mottling may accompany foliar deformations.).and zigzag growth.). Fruit set is poor. or indistinguishable from those of fanleaf. but bunches are small and few. In the European literature. These structures are readily visible by light microscopy in lignified shoots. and the yield may be much reduced.
Virus elimination is readily achieved from vegetating shoot tips by heat treatment (38-40 °C for as little as four weeks). with variable levels of sensitivity. The virus occurs in the pollen of infected grapevines and herbaceous hosts. reorganization. but resists infection when the virus is transmitted by the nematode. index is determined by the viral coat protein. GFLV can be transmitted by mechanical inoculation from infected grapevine tissues to various herbaceous hosts (e. Cytopathology: GFLV elicits the formation of intracellular cytopathic structures known as vesiculatevacuolate inclusion bodies which are often apposed to the nucleus. where they occur in a radial orientation. quinoa. but is not a specific test. vinifera x M. rotundifolia hybrid rootstocks (e. use of fumigants is more and more questioned for environmental reasons and is being progressively banned. Indexing on Vitis indicators by grafting takes a lot of time and field or greenhouse space. 20 .Agent: Grapevine fanleaf virus (GFLV) is a nepovirus with polyhedral particles of about 30 nm in diameter. mannose and xylose. Work is under way in different laboratories to create GFLV-resistant rootstocks or cultivar through traditional breeding methods or genetic transformation technology. A satellite RNA 1114 nt in size is associated with some virus isolates. O36-16) show interesting levels of field resistance to GFLV. There are conflicting reports on seed transmission in grapevines. GFLV has been detected in small groups of viruliferous X. and transmission by X. and redistribution and are thought to be sites of viral polyprotein processing and RNA replication. or by somatic embryogenesis. Transmission: At a site. serologically rather uniform and occurring as a family of minor molecular variants. varieties are susceptible. which are desirable as they cause no harm to human health. in a persistent manner by the longidorid nematode Xiphinema index feeding on the roots of grapevines and retaining the virus for several months. index (10 individuals) by ELISA and in single nematodes by RT-PCR and immunosorbent electron microscopy. Indexing on herbaceous hosts by mechanical inoculation requires climatized greenhouses and is less reliable than ELISA. However. GFLV was the first grapevine virus to be recovered by mechanical inoculation and to be thoroughly characterized physicochemically and molecularly. C. These inclusions derive from membrane proliferation. In the laboratory. and very sensitive method. RT-PCR and immunocapture RT-PCR are becoming increasingly popular. Detection: ELISA using polyclonal antisera and monoclonal antibodies is a quick. The best antigen sources for serological diagnosis are leaves collected in spring or cortical shavings from mature dormant canes. rotundifolia can be infected by GFLV when graft inoculated. Chenopodium quinoa. index feeding. Nematode populations transmit local virus isolates with a higher efficiency than those from other geographical areas. consisting of two functional molecules 7342 nt (RNA-1) and 3774 nt (RNA-2) in size. Virus particles are often within tubular structures that accumulate in bundles in the cytoplasm or nucleus. Observation of symptoms in the field is useful as a first step in selection. M. index in V. Specific transmission by X. Dissemination over medium and long distances is through infected vegetatively propagated scionwood and rootstocks. RT-PCR is estimated to be four to sixfold more sensitive than ELISA. cheap. Resistance to X. Transmission by Xiphinema italiae has not been consistently documented. amaranticolor. 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. Muscadinia (Vitis) rotundifolia and Vitis munsoniana are highly resistant to X. and soybean. Gomphrena globosa). This resistance is controlled by two unlinked recessive genes. For transformation. both required for infectivity. 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. vinifera. Molecular assays using radioactive or digoxigenin-labelled probes. by in vitro meristem tip culture. vuittenezi has been suspected but not proven. C. rotundifolia hybrids is thought to be controlled by a single dominant gene. but is not reliable. Control: Use of virus-tested scionwood and rootstock material in the framework of clean stock or certification programmes. but it is still regarded as necessary for confirming freedom from virus infection. Detection of trabeculae can give information on the health of American rootstocks. Positive sense single-stranded RNA genome.g. and is transmitted through seeds of C. amaranticolor. but show few symptoms. Some V. In contaminated soils. Varietal susceptibility: Almost all known Vitis vinifera L. a number of selectable marker genes toxic to non engineered vines are used. the endosperm of grapevine seeds. However. are toxic to many plants but not to V. American rootstocks are also susceptible and are generally very sensitive. However. rupestris x M. although some like Vitis labrusca can be infected. the use of fumigants against nematode vectors gives only a temporary but economically valuable control of the disease. encapsidated in different particles.g. Natural GFLV infections have been detected in weeds in Hungary and Iran.
research and hypotheses on fanleaf. 871 pp.2. No direct proof of transmission by this aphid. bactéries.b Hewitt: Fanleaf and yellow mosaic recorded from California. as well as on controversies about transmission by phylloxera. With roots of fanleaf-infected grapevines with phylloxera feeding on them. 1929 1931 1937 1937 1946 1950a. 2. but only circumstantial evidence. (a) See references in the Introduction. Historical review From the late 1800 to 1997. 3. Imprimerie du "Paysan du Midi". see the book by Galet. 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. 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. 21 . Healthy rooted cuttings or seedling of Rupestris du Lot were contaminated: 1. 1954 1958 Hewitt: Review on grapevine virus and virus-like diseases found in California. France. Montpellier. No conclusive results were obtained. With soil containing phylloxera. Les maladies et les parasites de la vigne. (b) Galet P. 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. viroses et phanérogames). 1977(b) 1865 1882 1895 1896 1902 1906 1910 1912 1912 1917 1918 Cazalis-Allut. Report on first experiments on heat treatment of grapevine in order to eliminate fanleaf. Hypothesis about a possible role of phylloxera as a vector. Branas et al. 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.: Experiments on the capacity of phylloxera to transmit fanleaf. With individual phylloxera (radicicolous or gallicolous) fed on infected vines. Hypothesis that fanleaf is a fungal disease. For a comprehensive review on early observations. Petri: Grapevine "arricciamento" (fanleaf) has a viral origin Arnaud and Arnaud: Hypothesis of a viral origin for grapevine court-noué (fanleaf). Tome 1: Les maladies dues à des végétaux (champignons..: Hypothesis that court-noué (fanleaf) is caused by a virus transmitted by phylloxera. Bovey: Review on grapevine virus and virus-like diseases. Arnaud: Court-noué is considered as a soil-borne virus disease. Branas et al. the ICVG Bibliographic Reports (a) have recorded more than 1000 papers dealing with fanleaf. 1977.
: Fanleaf virus is transmitted by the nematode Xiphinema index Cadman et al. Discussion on the possible role of weeds in the epidemiology of the disease. Vuittenez: Mechanical transmission of fanleaf virus to Chenopodium quinoa and C. index occurred during the 6 year period of observation. Boubals and Dalmasso: Experiments on soil disinfection against X. Bercks and Querfurth: Use of latex-test for detecting GFLV and other nepoviruses in grapevine tissue extracts in Germany. Cohn et al. 1965 1967 1968 1968 1969 1969 1970 22 . Dichloropropane-dichloropropene (DD) at 1000 l/ha gave satisfactory results. Hewitt et al. Dias and Harrison: Relationships between the viruses causing fanleaf. Hewitt et al. Bercks: Research on the use of three serological methods for detecting plant viruses. including fanleaf virus: bentonite flocculation test. Brückbauer and Rüdel: The virus (or viruses) of Reisigkrankheit (GFLV and/or other nepoviruses) are seed transmitted in some herbaceous indicator plants. Yield was increased by 400 % in comparison with that of untreated controls. amaranticolor is confirmed.: Detection of GFLV particles in thin-sectioned grapevine root tissues. latex test and barium sulfate test. whereas insecticide treatment has no effect. Graniti and Russo: A light microscope and cytochemical study of endocellular cordons. Reproduction of fanleaf symptoms in mechanically inoculated grapevine seedlings is confirmed Galzy: Heat treatment of grapevine plantlets grown aseptically in vitro.b Martelli and Hewitt: Comparative studies showed that Californian and Italian GFLV isolates are the same. Description of the chipbudding method for indexing. and no reinfestation by X.1958 1958 1960 Vuittenez: Fumigation of fanleaf-contaminated soil with nematicides prevents infection of healthy grapevines replanted immediately. Gifford and Hewitt: Use of heat therapy and in vitro shoot tip culture to eliminate fanleaf virus from infected grapevines. index. Serological relationship with ArMV reported. Das and Raski: Studies on the relationships of GFLV with its vector X. index. Goheen and Hewitt: Description of vein banding as a GFLV-induced disease Dias: Host range and properties of fanleaf and yellow mosaic viruses. Control of the vector by soil fumigation. Reproduction of fanleaf symptoms in mechanically inoculated grapevine seedlings 1964 1964 1965 Taylor and Hewitt: Description and characterization of Australian isolates of GFLV. Gerola et al.: Transmission of GFLV by Xiphinema italiae in Israel. Goheen et al. index in France. 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.: Investigations on grapevine virus diseases in California. yellow mosaic and ArMV 1960a 1960b 1961 1961 1962 1962 1963 1963 1963a. Vuittenez: New observations on the effects of soil fumigants on fanleaf in contaminated soils. Transmission of fanleaf virus by X.: Transmission of fanleaf virus from grapevine to herbaceous hosts by mechanical inoculation and preliminary characterization of the virus.: Description of the Davis method of heat therapy of grapevines.
: Detailed physico-chemical characterization of GFLV. Hévin et al.3-dichloropropene or methyl bromide. but ELISA is more sensitive. 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. Walter et al. 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. St. Both tests are more sensitive than mechanical inoculation to C. the plant is placed in a heat cabinet for treatment Hévin et al. Goheen and Luhn: New method of heat therapy. index.: Control of fanleaf by soil fumigation with 1. Van Velsen and Niejalke: Green budding for indexing grapevine with the indicator cvs. Mur et al. Alfaro and Goheen: The different strains of fanleaf virus (fanleaf sensu stricto.: 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. Mission or LN 33. quinoa. Indexing by budding on V. Raski et al. A dormant bud of the variety to be cured is grafted onto a healthy potted rootstock. rupestris is more accurate than mechanical transmission to C.: Comparison between PALLAS latex test and ELISA for detecting GFLV in France.: Use of green grafting as a quick and secure method for graft-indexing. Dias: Review paper on grapevine yellow mosaic. Taylor: Review paper on grapevine vein banding. Bercks: Serological detection of grapevine viruses in West Germany. The acquisition time threshold is less than 5 minutes. index. especially with low titre antisera. this lining is shed and ingested in the intestine.: Heat therapy of grape plantlets grown in vitro causes changes in some characteristics of the variety. Raski et al. PALLAS is quicker and cheaper than ELISA. Uyemoto et al. George. George for detecting GFLV.: GFLV particles observed in the lumen of the oesophagus of X. Querfurth and Paul: Protein A-coated latex-linked antiserum (PALLAS) method for detecting GFLV and other viruses. rupestris St. Vuittenez: Review paper on grapevine fanleaf.3 dichloropropene or methyl bromide.: GFLV and marbrure (fleck) are not transmitted through the seeds of grapevine. During the moult of the nematode. yellow mosaic and vein banding) are transmitted in the same way by X.1970 1970 Hewitt et al. The sensitivity of the latex test is increased. 1970 1970 1970 1971 1971 1972 1972 1973 1973 1973a 1973b 1974 1974 1975 1976 1976 1977 1979 1979 23 . anterior oesophagus and oesophageal bulb of their nematode vectors. After bud take. quinoa. Vineyards replanted in contaminated but treated soils remained healthy for at least 5 years.: Comparison of indexing by mechanical inoculation to Chenopodium quinoa and by graft-transmission to V. Raski and Schmitt: Progress in the control of the fanleaf-nematode complex by soil disinfection with 1. Both methods give satisfactory and similar results.
Carbon disulfide gave less satisfactory results. although it is not resistant to the virus itself. The use of methyl bromide requires a continuous cover with polyethylene sheeting for some time after the treatment. vinifera accession represent an excellent example of host plant resistance to GFLV.: Identification of a natural serological variant of GFLV from Tunisia Huss et al. Bouquet: Serological detection of GFLV in its vector X. Efficiency of both methods is compared. Rüdel: Discussion on the possible role of X.: Soil fumigation with 1.3-D) or methyl bromide applied 75-90 cm deep with 90 cm spacing for 1. Bouquet: M. Morris-Krsinich et al. Forty days of treatment.: Use of systemic nematicides for the control of X. especially in wood shavings of dormant canes during winter. Krake and Woodham: Possibility that the agent of yellow speckle is involved together with GFLV in the etiology of vein banding. vuittenezi population used for the experiments could not be entirely ruled out. That X. a very common species in vineyards of Rheinhessen and Palatinate. Brown and Roberts: Detection of fanleaf virus in its vector X. index.: Comparison of polyclonal and monoclonal antibodies for detecting fanleaf virus with ELISA in various grapevine tissues. Hafez et al. the possibility that a few X. rotundifolia is resistant to fanleaf virus transmission by X. index by ISEM Bovey et al. index. A Middle Eastern V. in California. Methyl bromide and 1. as vector of GFLV. Raski et al.: Detection of fanleaf virus in grapevine tissues by ELISA and ISEM in different periods of the year.: Experiments with systemic nematicides for controlling X. index and fanleaf in grapevines. Vuittenez: Review on serological methods of detection and identification of grapevine viruses. index by ELISA. index feeding. Raski et al.3-dichloropropene failed to eradicate either nematodes or fanleaf virus from the soil but reduced the incidence of the disease to acceptable levels.3-dichloropropene (1. index. Transmission trials gave a few positive results. Lear et al.: Identification of several Vitis species and interspecific hybrids resistant to fanleaf virus. index larvae were present in the X.: Detection of fanleaf virus and other sap-transmissible viruses in grapevine tissues by ISEM.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. Monette: Heat therapy of GFLV and ArMV--infected grapevines with alternating temperatures. These are promising sources of germplasm for obtaining resistant rootstocks. vuittenezi might be a vector of GFLV is therefore uncertain. Russo et al.: GFLV particles are arrayed in long parallel rows in thin-sectioned mesophyll cells of infected grapevines.: Study on the effectiveness of soil fumigation for the control of X. RNA-2 contains the cistron coding for the viral coat protein. Walker et al. 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.1979 1980 1980 Kalasian et al. with temperatures of 39 °C for 6 h followed by 22 °C for 18 h made it possible to eliminate both viruses from the developing shoot tips (2 mm) of in vitro cultured plantlets. index. 1980 1980 1980 1981 1981 1981 1981 1983a 1983b 1983 1983 1983 1985 1985 1986 1986 24 . Savino et al. vuittenezi.: 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.
Mild strains were shown to confer protection towards severe challenging strains in Chenopodium and grapevine. Positive. Walter and Etienne: Detection of GFLV in wood shavings of dormant canes. RpRSV and ArMV are common.1987 1987 1987 Huss et al.3-dichloropropene and methyl bromide for controlling X. Lázár et al. quinoa and transmitted by heterografting to Vialla and Kober 5BB rootstocks. Etienne et al.: Use of green grafting technique for sensitive and quick GFLV detection under greenhouse conditions. Raski and Goheen: Comparison of 1. Previous experience showed that 1.: Determination of the nucleotide sequence of the satellite RNA (RNA-3) of GFLV. Pinck et al. Walter et al. Rüdel: Review on the most important virus diseases of grapevines in West Germany. Viruliferous nematodes were crushed in standard extraction buffer and tested in batches of 1-50 by means of DAS-ELISA. Altmayer: Elimination of GFLV.: Production and use of monoclonal antibodies to GFLV. SLRV. Long term fallow (about 5 years). Fuchs 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.: Detection of GFLV in grapevine seeds and seedlings by ELISA. Rüdel: Severe restrictions set on the use of soil fumigants in West Germany for environmental reasons make control of "Reisigkrankheit" very difficult. Walter et al. RpRV. TBRV and leafroll from infected grapevines by in vitro meristem tip culture. cultivation of non-host plants and organic soil amendments are recommended. rotundifolia showed good resistance to GFLV in California. No eradication was obtained. Effect on yield and economic importance. Reliable results were obtained with samples of 20-50 nematodes. GFLV. Mild and severe strains were discriminated on the basis of field performance of infected Vitis.: Study of interactions between GFLV and ArMV isolates grown in C. Catalano et al. RNA-3 encodes a non structural protein. Treated vines yielded more for over 4 years. 1988 1988 1988 1989 1989 1989 1989 1989 1990 1990 1990 1990 1990 1991 1991 1991 25 . Serghini et al. index and GFLV. Resistance is controlled by two unliked recessive genes Walter et al. Catalano et al.: Detection of GFLV in the vector Xiphinema index by ELISA.: Identification of a satellite RNA of GFLV. Walker et al. vinifera x V.: Determination of the complete sequence of GFLV RNA-2.: Evidence of a differential efficiency of GFLV transmission by Xiphinema index populations from different geographical origins.: Two rootstock selections derived from crossings V. ArMV. The selection of resistant cultivars and rootstocks is considered of primary importance. and has strong homologies with the satellite RNA associated with ArMV.: Improvement in the serological detection of GFLV and ArMV viruses using monoclonal antibodies.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. Martelli and Taylor: Review article on nematode-transmitted viruses. Walker and Meredith: Identification of two accessions of Vitis vinifera resistant to GFLV. but less consistent results were obtained with 1-10 nematodes. the latter being especially damaging on the variety Kerner. Treatments with soil fumigants are no longer permitted in Germany.
identification of their replication determinants and evidence of replication in Chenopodium quinoa protoplasts 1993 1993 1993 1993 Martelli et al. delays symptom expression by 1-2 days and adversely affects virus replication.: 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. quinoa with biologically active transcripts of GFLV F-13 satellite RNA and GFLV strains devoid of satellite. Ritzenthaler et al. Walter et al. Fuchs et al. Satellite RNA appears to have a modulating effect on virus pathogenicity. Staudt and Weischer: Vitis rotundifolia and Vitis munsoniana resist infection by GFLV transmitted by X. Viry et al.: Production of biologically active transcripts from cloned cDNA of genomic RNAs of GFLV.: Detection of GFLV in X.: Production of GFLV satellite RNA transcripts.: GFLV isolated in Hungary from naturally infected symptomatic plants of Aristolochia clematis and Lagenaria siceraria turbinata.: Two Vitis vinifera x Muscadinia rotundifolia rootstock hybrids (O39-16 and O4343) grafted with Cabernet sauvignon showed a high level of tolerance to GFLV. Esmenjaud et al. GFLV is eliminated from somatic embryos obtained from tissue cultures grown at 35 °C 1991 1991 1992 1992 1992a. thus qualifying for use in X.: Use of modified GFLV coat protein genes for transformation of different Nicotiana species for inducing resistance.: Evidence that transgenic tobacco plants expressing the coat protein of GFLV are protected from GFLV infection. Both became infected in the course of a 12-year trial but had no reduced crop yields. Walker et al. Spielmann et al. Esmenjaud et al.: Co-inoculation of C. index infested soils. index by biotin-avidin ELISA Bardonnet et al. which is also resistant to phylloxera. 1993 1993 1993 1993 1993 1994 1994 1994 1994 26 . This represents the first substantiated record of a natural GFLV infection in hosts other than Vitis. Horvath et al. O39-16 in particular.:Detection of GFLV in single nematodes by RT-PCR. index.b Hans et al.: GFLV satellite RNA detected in 5 of 34 virus isolates from different geographical locations.: Genomic RNA-1 of GFLV is completely sequenced and its genetic organization determined. Staudt: Study of the spread of GFLV in several Vitis species.: Development of cDNA probes to GFLV genomic and satellite RNAs and their use for virus detection directly in grapevine extracts. index. 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.1991a 1991b Fuchs et al.: In naturally GFLV-infected vineyards the hypovirulent ArMV A1 isolate induces delayed infection by GFLV Saldarelli et al. hybrids and breeding stocks after infection of the roots by means of viruliferous X.
Gölles et al. Martelli and Walter: Review article on certification of grapevines. index. The level of resistance obtained looks promising. Rowhani et al. Spielmann et al.: Successful transformation of somatic embryos of an European grape cultivar (Russalska 3) with the normal. Belin et al. index populations by PCR-RFLP and sequencing of the ITS region. rotundifolia hybrids show high resistance to X. Pfeiffer et al.: Attempts to characterize molecularly X.: Demonstration that a 28 kDa protein coded by GFLV RNA-2 is involved in the replication of this RNA.: 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. Of 531 accessions of European.: Demostration that the movement protein of GFLV is located on the intracellular tubular structures containing rows of virus particles.: Transformation of Nicotiana species and Vitis rupestris with different virusderived (coat protein. Walter and Martelli: Review article on detrimental effects of viruses on grape yields.1995 1995 1995 1995 1995 1996 1996 Brandt and Himmler: Use of immunocapture RT-PCR for GFLV detection in host tissues. De Luca et al. Belin et al. RNase L) genes for inducing resistance to GFLV. Gaire et al. Lahogue and Boulard: Search for genes of resistance in grapevines. all were susceptible to the virus. except for four.: 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. 1997 1998 1998 1999 1999 2000 2000 2000 2000 2001 2002 2003 2003 2003 27 . Naraghi-Arani et al. Ritzenthaler et al.: Development of a GFLV detection system based on PCR analysis of immobilized virions. index feeding. Pfeiffer et al.: Genetic transformation of Vitis vinifera with the coat protein gene of GFLV for resistance induction.: Up to date account of GFLV replication strategy.: Genetic transformation of American roostocks with the coat protein gene of GFLV for resistance induction. Pinck: A comprehensive review of the molecular aspects of GFLV genome and its replication strategy. 5 oligoadenylate synthase. and Asian Vitis species inoculated by green grafting with a GFLV source. This resistance is controlled by a single dominant gene. Walker and Jin: V. American. : Identification of RNA2-encoded proteins in the specific transmission of GFLV by X. Ritzenthaler et al.: Identification of membranes derived from the endoplasmic reticulum as sites of GFLV replication. Mauro et al. truncated or nontranslatable coat protein gene of GFLV. including the two accessions reported as resistant by Walker and Meredith (1990). Fuchs: Review article on genetic transformation of grapevines for resistance to GFLV and other pathogens.: Identification of the molecular signal accounting for the systemic spread of GFLV in infected hosts. Krastanova et al. replicase) and exogenous (2. rupestris x M.
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U.A. All have polyhedral particles about 30 nm in diameter and a positive sense. 2005). Kurstak.. 145-147. Sixth Report of the International Committee on Taxonomy of Viruses (F. 1996. D. Ball. Nepovirus Group.. References Goldbach R. K. family Comoviridae (Goldbach et al.. Family Comoviridae. ed). 5.B. J. C. Piazzolla. has now been assigned to the newly established genus Sadwavirus (Le Gall et al. In: Virus Taxonomy. In: Grapevine Viruses and Certification in EEC Countries: State of the Art. V. 341-347.H. In: Handbook of Plant Virus Infections: Comparative Diagnosis. Polyhedral Virions and Bipartite RNA Genomes. i. Nepoviruses of grapevine and their nematode vectors in the EEC. subgroup B. Influence des viroses et qualité. which were originally included in the Nepovirus group (Harrison and Murant. Wetzel and N. Desselberger and L.). Leibowitz. Simons and M. 3.F.D. 197-238. Fritsch. typified by Tomato black ring virus (TBRV). G. Taylor C. San Diego. 1ere partie: Effects des viroses sur la culture des vignes et ses produits.. 2005. Sélection clonale de la vigne: sélection sanitaire et sélection pomologique. RT-PCR) are routinely used for their detection in grapevine tissues (primarily cortical scrapings from dormant canes). Jones.M. 807-818. Yoshikawa.D. CAB International. and A. 2000. Several of these viruses have distorting and chromogenic strains and may occur in mixed infections with GFLV. and molecular characteristics of nepoviruses (Harrison and Murant. H. Nematode Vectors of Plant Viruses. A. Wellink. Walter B. Milne.E and D. Martelli G. Amsterdam. Le Gall O. T. Harrison B. Quacquarelli. A tentative grouping of nepoviruses Phytopathologia Mediterranea 17. typified by Tomato ringspot virus (ToRSV) (Martelli et al. a nematode-borne virus originally classified as a tentative species in the genus Nepovirus.J. These viruses can readily be eliminated by heat therapy or in vitro meristem tip culture. 1977. CMI/AAB Descriptions of Plant Viruses No. and A. 362 pp. 35 . Harrison B. 1978.F. Martelli and R. Mechanical transmission to herbaceous hosts or indexing on Vitis indicators can also be used. Nepoviruses. Quaderno No. subgroup C.. M. eds) Elsevier/Academic Press. Springer-Verlag. 1997.P.P. are now classified in the genus Nepovirus. 10281032 Murant A. Academic Press. (G. the Mediterranean and Middle East. Bari. Oxon..V. Vol. Le Gall et al. Brown. Taliansky.J. 1977) . Savino and P. subgroup A typified by Tobacco ringspot virus (TRSV). In: Virus Taxonomy. Gallitelli. 2005) Extensive reviews of the biological. which are separately encapsidated. Scholtof. In: Virus Taxonomy. 23-39..G. M. G..A. Van Regenmortel et al. Many are transmitted by longidorid nematodes (Rüdel.. 1996.INFECTIOUS DEGENERATION (EUROPEAN AND MEDITERRANEAN NEPOVIRUSES) Besides GFLV several other nepoviruses can infect grapevine in Europe. 1955. epidemiological. Plenum Press. K. Their detrimental effects to grapevine culture and products have been summari zed by Walter and Martelli (1996). London. eds).. Murphy et al.P.A. Seventh Report of the International Committee on Taxonomy of Viruses (M. New York. Fauquet. Murant 1981. 185. (E. Mayo M. 1978. ISEM) and molecular assays (hybridization. causing diseases whose symptoms are similar to. 2000) are available. Martelli. Elsevier/Noth Holland.P. Nepoviruses. ed. 1955) and are subdivided into subgroups based on physico-chemical properties of member viruses.A. 945-971. Taylor and Brown. eds).V. physico-chemical. and G. 1992). 1996. Satellite nucleic acids.. Lehto.F. Strawberry latent ringspot virus (SLRSV). Istituto Agronomico Mediterraneo. Karasev. Maniloff.A. Sanfaçon. The Plant Viruses. 1997) and their satellite RNAs (Mayo et al.e. Rüdel M. 1992. Bulletin de l'OIV 69. Vienna. 1981. single-stranded RNA genome occurring as two functional species (RNA-1 and RNA-2). Murant. Family Comoviridae. T. Mayo. A. a non-taxonomic clustering. T.F. Iwanami. 286 pp. P.. Murant (eds). Martelli. J. Palukaitis.. or indistinguishable from those of fanleaf. Serological (ELISA. Eight Report of the International Committee on Taxonomy of Viruses (C.
Jankulova and Kaitasova: ArMV found in grapevine in Bulgaria. In other V. Murant: ArMV description in the CMI/AAB Descriptions of Plant Viruses series. Description ArMV. Romania. Hungary. consisting of two separately encapsidated molecules with Mol. Iran. SLRSV and TBRV found in grapevines with atypical Reisigkrankheit in West Germany.: Isolation of ArMV from grapevine in Italy. TBRV. and with other nepoviruses in the Reisigkrankheit complex of western Germany.4 x 106 and a size of 1104 nt. losses up to 50 % have been recorded. the vector of GFLV. index.: Ultrastructure of ArMV infections in plant tissues Martelli and Lehoczky: Detection of ArMV in grapevine in Hungary. ArMV occurs often in mixed infections with GFLV in certain areas of France and Italy. AILV and GCMV. Transgenic plants expressing the coat protein gene of the virus have been produced. 2.Genus NEPOVIRUS ARABIS MOSAIC VIRUS (ArMV) 1. Bulgaria. Dalmasso et al.: ArMV. and. symptoms are of the fanleaf type.: ArMV detected in Japan in grapevines imported from Europe. always in Germany the severe dieback disease of the cv. The genome is a positive sense ssRNA. have a angular outline. Gerola et al. and circular about 350 nt in size. vinifera varieties. ArMV and TBRV by ELISA in Israel. ArMV. The virus supports the replication of two types of satellite RNAs. Brückbauer and Rüdel: Symptoms of atypical Reisigkrankheit in the vineyard are associated with ArMV in West Germany Bercks et al. a typical nepovirus belonging in subgroup A of the genus Nepovirus.: Xiphinema diversicaudatum can transmit ArMV to grapevine.: Detection of ArMV by ISEM Tanne: Detection of GFLV.95-2.1 x 106 (RNA-2). Canada. Rüdel: Transmission of ArMV to grape seedlings by Xiphinema diversicaudatum. This virus has also been found in grapevine in Switzerland.: Physico-chemical properties of GFLV. Component T is made up of empy protein shells. Turkey.2 x 106 (RNA-1) and 1. Yugoslavia. whereas components M and Bcontain RNA. and sediment as three components (T. Quacquarelli et al. M. USA (California). and B). Vuittenez et al. Its particles are about 30 nm in diameter. It can infect many woody and herbaceous plants and is transmitted to grapevine by the nematode Xiphinema diversicaudatum but not by X. In Germany. Coat protein has a single type of subunits with Mr 54 x 103. accounting for 27% (component M) and 41% (component B) of the particle weight. Belli et al. 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. Russo et al. Cross-protection between ArMV and GFLV has been reported. wt of 0. and Japan. Kerner appears to be caused by ArMV infection. 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. Israel. 36 . linear with Mol. is serologically related to GFLV. wt 2.: Interactions between nepoviruses in grapevine and herbaceous hosts. respectively. Kobayashi et al. Stellmach: Review paper on ArMV in grapevine.
Marc-Martin et al. Stellmach and Berres: The susceptibility of cv. whereas other nepoviruses.: Properties of a strain of ArMV isolated from grapevine in Italy. Molecular assays are as good as ELISA for routine testing.: Cross-protection experiments in Chenopodium quinoa between ArMV and GFLV Gugerli et al. MacKenzie et al. ArMV.: Nucleotide sequenece of a small circular satelllite RNA Steinkellner et al.: Evidence that ArMV has two RNA-2 molecules and complete nucleotide sequence of both RNAs Etscheid et al.: Properties of ArMV small satellite RNA. Hypothesis of a graft incompatibility when the rootstock is infected with ArMV. The virus cannot be recovered from leaves or buds of the Kerner scions. Ipach et al.: Transgenic Nicotiana plants transformed with the coat protien of ArMV produce empty viral shells. When a healthy scion is grafted onto an infected rootstock. Steinkellner et al.Kerner to ArMV seems to be limited in time. Rüdel: In the Palatinate (West Germany) ArMV is transmitted by Xiphinema diversicaudatum and occurs often in mixed infections with GFLV in grapevine. SLRV and TBRV Belli et al. Study of histological changes at the graft union level.: Nucleotide sequence of the ArMV satellite RNA Liu et al.: ArMV in Switzerland Lázár et al. Kaper et al. 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 .: Survey for the presence of ArMV in Canadian vineyards Lahogue and Boulard: Search for genes of resistance in grapevines.1982 1984 1985 Brückbauer: Possibility of distinguishing GFLV.: Association of ArMV-infected rootstocks with Kerner disease in West Germany. Faber.: Use of cDNA probes for ArMV detection. such as RpRSV or GFLV can be found in both rootstock and scion. the virus is recovered from the scion only during the first year.: 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.: ArMV is not seed-transmitted in grapevines Liu et al.: A hypovirulent ArMV isolate delays GFLV infection in grapevines under field conditions. RRV. : Comparison of coat proteins of ArMV and other nepoviruses.: Transformation of grapevines with the coat protein gene of ArMV. whereas the rootstock remains infected. Eppler et al. Becker et al. Yield losses may reach 77% in cv.: Detection of ArMV by PCR in herbaceous hosts and grapevines Steinkellner et al. Walter et al.: The presence of ArMV satellite RNA can attenuate symptoms in certain hosts Bertioli et al. Stellmach: Kerner disease is probably caused by ArMV.: ArMV recorded from Romania Huss et al.
European, American, and Asian Vitis species inoculated by green grafting with a ArMV source, 42 were apparently resistant. 1998 2000 2003 2004 Akbas and Erdiller: ArMV recorded from Turkey Goelles et al. : Transgenic grapevines expressing ArMV coat protein gene Fuchs: Review on transgenic resistance of grapevines to pathogens Pourrahim et al.: ArMV identified in Iranian grapevines by mechanical transmission to herbaceous hosts and ELISA
3. References Akbas B. and G. Erdiller, 1998. Grapevine virus diseases in Karaman, Konya and Nevsheir provinces. Proceedings VII Congress of the Turkish Phytopathological Society, Ankara 1998, 149-153. Becker A., J. Jäger and B. Altmayer, 1989. Association of arabis mosaic virus-infected rootstocks with the dieback of the Vitis vinifera cv."Kerner" in Germany. Proceedings 9th Meeting of ICVG, Kiryat Anavim, 1987, 57-61. Belli G., A. Fortusini and G. Vegetti, 1982. Il virus del mosaico dell' Arabis isolato da vite in Italia. Rivista di Patologia Vegetale, S IV, 18, 175-177. Belli G., G. Vegetti, S. Cinquanta, C. Soncini, S. Prati and D. Tolentino, 1984. Properties of a strain of arabis mosaic virus isolated from grapevine in Italy. Rivista di Patologia Vegetale, S IV, 20, 56-64. Bercks R., H. Brückbauer, G. Querfurth and M. Rüdel, 1977. Untersuchungen über die Viruskrankheiten der Rebe unter besonderer Berücksichtigung "atypischer Formen" der Reisigkrankheit. Weinberg und Keller 24, 133-180. Bertioli D.J., R.D. Harris, M.L. Edwards, J.I.Cooper and W.S. Hawes, 1991. Transgenic plants and insect cells expressing the coat protein of Arabis mosaic virus produce empty virus-like particles. Journal of General Virology 72, 1801-1809. Brückbauer H. and M. Rüdel, 1976. Untersuchungen über eine "atypische" Form der Reisigkrankheit bei der Rebsorte Silvaner. Weinberg und Keller 23, 53-79. Dalmasso A., M.C. Munck-Cardin and R. Legin, 1972. Résultats préliminaires d'essais de transmission de sérotypes de la mosaïque de l'Arabis trouvés sur vigne, par l'intermédiaire de Xiphinema diversicaudatum. Annales de Phytopathlogie 4, 410 Eppler A., V. Lesan and A. Lázár, 1989 Viruses and virus diseases in some vineyards in Romania. Meded. Facultet Landbouwwetenschappen Rijkuniversiteit Gent 54, 491-497. Etscheid M., M.E. Tousignat and J.M. Kaper, 1995. Small satellite of arabis mosaic virus autolytic processing of in vitro transcripts of (+) and (-) polarity and infectivity of (+) strand transcripts. Journal of General Virology 76, 271-282. Flak W. and H. Gangl, 1994. Grobkartierung des Rebenvirosenbefalls in der Weinbauregion Bungerland mittels ELISA. Mitteilung Klosterneuburg 44, 163-167. Fuchs M., 2003. Transgenic resistance: state of the art. Extended Abstracts 14th Meeting of ICVG, Locorotondo 2003, 221-223. Gerola F.M. M. Bassi and E. Betto, 1964. Shape and localization of arabis mosaic virus in experimentally infected cells of Chenopodium amaranticolor. Phytopathologische Zeitscrift 51, 192-194. Goelles R., R. Moser, H. Puhringer, H. Katinger, M.L. da Camara Machado, A. Minafra, V. Savino, P. Saldarelli, G.P. Martelli and M. Laimer da Camara Machado, 2000. Transgenic grapevines expressing coat protein gene sequences of grapevine fanleaf virus, arabis mosaic virus, grapevine virus A and grapevine virus B. Acta Horticulturae 528, 305-311. Gugerli P., J.J. Brugger and P. Basler, 1990. Les maladies de l'enroulement, du bois strié et de l'ècorce liégeuse de la vigne (grapevine leafroll, rugose wod and corky bark). Revue Suisse de Viticulture, Arboriculture et Horticulture 22, 35-36. Kaper J.M., M.E. Tousignant and M. Steger, 1988. Nucleotide sequence predicts circularity and selfcleavage of 300-nucleotide satellite of arabis mosaic virus. Biochemical and Biophysical Research Communications 154, 318-322. Lahogue F. and G. Boulard, 1996. Recherche de gènes de résistance naturelle à deux viroses de la vigne: le court-noué et l'enroulement. Vitis 35, 43-48. Liu Y.Y., C.U.T. Hellen, J.I. Coper, D.J. Bertioli, D. Coates and G. Bauer, 1990. The nucleotide sequence of a satellite RNA associated with arabis mosaic nepovirus. Journal of General Virology 71, 1259-1263. Liu Y.Y., J.I. Cooper, M.L. Edwards and C.U.T Hellen, 1991. A satellite RNA of arabis mosaic virus and its pathological impact. Annals of Applied Biology 118, 557-587.
Loudes A.M., C. Ritzenthaler, M. Pinck, M.A. Serghini and L.. Pinck, 1995. The 119 kDa and 124 kDa polyprotiens of arabis mosaic nepovirus (isolate S) are encoded by two distinct RNA2 species. Journal of General Virology 76, 899-906. MacKenzie D.J., R.C. Johnson and C. Warner, 1996. Incidence of four important viral pathogens in Canadian vineyards. Plant Disease 80, 955-958. Murant A.F, 1970. Arabis mosaic virus. CMI/AAB Descriptions of Plant Viruses, No. 16 Pourrahim R., A. Ahoonmanesh, Sh. Farzdfar, F.Rakhshandehro and A.R. Golanaraghi. Occurrence of Arabis mosaic virus and Grapevine leafroll-associated virus 3 in Iran. Plant Disease 88, 424. Quacquarelli A., D. Gallitelli, V. Savino, P. Piazzolla and G.P. Martelli, 1979. Some properties of grapevine fanleaf and other nepoviruses infecting the grapevine. Proceedings 6th Meeting of ICVG, Cordoba, 1976. Monografias INIA No. 18 , 41-49. Rüdel M., 1978. Übertragung des Arabis-Mosaik-Virus (AMV) durch den Nematoden Xiphinema diversicaudatum (Micoletzki) Thorne auf Rebensämlinge (Vorläufige Mitteilung). Die WeinWissenschaft 33, 243-247. Rüdel M., 1985. Grapevine damage induced by particular virus-vector combinations. Phytopathologia Mediterranea 24, 183-185. Steinkellner H., G. Himmler, M. Laimer, D. Mattanovich, G. Bisztray and H. Katinger, 1989. Konstruktion von cDNA von Arabis Mosaik Virus und deren Anwendung für Diagnose. Mitteilung Klosterneuburg 39, 242-246. Steinkellner H., G. Himmler, R. Sagl, D. Mattanovich, and H. Katinger, 1992. Amino acid sequence comparison of nepovirus coat proteins. Virus Genes 6, 197-202. Steinkellner H., A. da Camara Machado, M. Laimer-da Camada Machado, M. Gölles and H. Katinger, 1993. Studies on coat protein mediated cross protection of nepoviruses. Extended Abstracts 11th Meeting of ICVG, Montreux 1993, 175. Stellmach G., 1970. Arabis mosaic in Vitis. In: Virus Diseases of Small Fruits and Grapevines- A Handbook (N.W. Frazier, ed). University of California, Division of Agricultural Sciences Berkeley, 233-234. Stellmach G., 1987. Die Kerner Krankheit: Theoretische und praktische Aspekte einer tödlichen Rebenvirose. Die Wein-Wissenschaft 42, 421-427. Stellmach G. and R.-E. Berres, 1986. Begrenzte Infektionsanfälligkeit der Vitis vinifera- Sorte "Kerner" gegenüber dem Arabismosaik-Virus? Zeitschrift für Pflanzenkrankheiten und Pflanzenschutz 93, 356-360. Walter B, P. Bass, P. Cornuet and P.M. Guillaume, 1993. Preliminary results of cross protection experiments against grapevine fanleaf virus (GFLV) in the vineyards. Extended Abstracts 11th Meeting of ICVG, Montreux 1993, 167-168 ARTICHOKE ITALIAN LATENT VIRUS (AILV) 1. Description Artichoke Italian latent virus (AILV), a member of subgroup B of the genus Nepovirus, was isolated in Bulgaria from vines with fanleaf-like symptoms. AILV has isometric particles with angular outline, sedimenting as three components: T (empty shells), M (particles contaning a molecule of RNA-2 with Mol. wt of 1.5 x 106 daltons accounting for 34% of the particle weight) and B (particles containing a molecule of RNA-1 with Mol. wt of 2.4 x 106 daltons, accounting for 41% of the particle weight). Coat protein is made up of a single type of subunits with Mr 54 x 103. AILV is transmitted by the Dorylaimoid nematode Longidorus apulus in vegetable crops but no field transmission to grapevines has been recorded. The virus has limited distribution and economic importance. 2. Historical review 1976 Jankulova et al.: AILV isolated in southern Bulgaria in 1976 from a grapevine with fanleaf-like symptoms. Properties of the virus, cultured in Chenopodium quinoa determined and positive serological reaction with an antiserum to an Italian strain of AILV ascertained. Savino et al.: Comparison of a Bulgarian grapevine isolate of AILV with an Italian isolate from artichoke and two Bulgarian isolates from sowthistle and gladiolus. Martelli et al.: AILV description in the CMI/AAB Descriptions of Plant Viruses series.
3. References Jankulova M., V. Savino, D. Gallitelli, A. Quacquarelli and G.P. Martelli, 1979. Isolation of artichoke Italian latent virus from the grapevine in Bulgaria. Proceedings of the 6th Meeting of ICVG, Cordoba 1976. Monografias INIA 18, 143-148. Martelli G.P., G.L.. Rana and V. Savino, 1977. Artichoke Italian latent virus. CMI/AAB Descriptions of Plant Viruses, No. 176. Savino V., D. Gallitelli, M. Jankulova and G.L. Rana, 1976. A comparison of four isolates of Artichoke Italian latent virus (AILV). Phytopathologia Mediterranea 16, 41-50. CHERRY LEAFROLL VIRUS (CLRV) 1. Description Cherry leafroll virus (CLRV) is a cosmopolitan virus. In Chile it was recovered from vines with fanleaflike symptoms and in Germany from vines with yellow mosaic-like symptoms. Although CLRV is a definitive nepovirus species classified in subgroup C of the genus Nepovirus it differs from most of the other members in the genus being transmitted by pollen rather than nematodes. The vector to grapevine, if any, is unknown. CLRV occurs in nature as multiple strains but is not serologically related to any of the known nepoviruses. Virus particles are isometric, about 30 nm in diameter have angular outline and sediment as three components (T, M, and B). Their coat protein consists of a single type of subunits with Mr of about 54 kDa. The genome is a bipartite, positive sense, single-stranded RNA which has been sequenced only in part. Genomic RNA consists of two separately encapsidated functional molecules with Mol. wt of 2.8 x 106 (RNA-1) , accounting for 46% of the particle weight, and 2.3 x 106 (RNA-2), accounting for 41% of the particle weight. In grapevines CLRV is readily detected by DAS-ELISA. The best woody indicator for the German isolate is reported to be Pinot noir. 2. Historical review 1985 1993 2001 2003 Jones: Description of Cherry leafroll virus in the AAB Descriptions of Plant viruses series. Scott et al.: Partial nucleotide sequence of CLRV RNA-2. Herrera and Madariaga: First record of CLRV from Chile. Field infection is estimated to be 0.2% Ipach et al.: Isolation of CLRV from German vines showing yellow mosaic-like symptoms and reduced crop.
3. References Herrera M.G and V.M. Madariaga, 2001. Presence and incidence of grapeivne viruses in central Chile. Agricultura Tecnica 61, 393-400 Ipach U., L. Kling and D.E. Lesemann, 2003. First record of Cherry leafroll virus on grapevine in Germany. Extended Abstracts 14th Meeting of ICVG, Locorotondo, 2003, 17-18 Scott N.W., J.I. Cooper, and M.L Edwards, 1993. The identification, cloning, and sequence analysis of the coat protein coding region of a birch isolate of cherry leafroll nepovirus. Archives of Virology 131, 209-215. GRAPEVINE ANATOLIAN RINGSPOT VIRUS (GARSV) 1. Description Grapevine Anatolian ringspot Virus (GARSV) was isolated from Turkish grapevines with mild fanleaflike symptoms. The virus belongs in subgroup B of the genus Nepovirus but is not serologically related to any of the known grapevine nepoviruses. Virus particles are isometric c. 30 nm in diameter and sediment as three centrifugal components. RNA-1 has a Mol. wt of 2.2 x 106 Da and RNA-2 a Mol. wt of 1.4 x 106 Da and a size of 4607 nt. Coat protein subunits have a Mr 56 x 103 Da. GARSV can be readily detected by ELISA and PCR using primers designed on the coat protein sequence. The virus has no recognized
S. Biological. GRAPEVINE BULGARIAN LATENT VIRUS (GBLV) 1. Digiaro and G. Digiaro. 2002. Complete nucleotide sequence of RNA-2 of two Turkish nepoviruses. M. D.vector.5 x 106 (less than 1800 nt). De Mendonça et al. Sabanadzovic.. M. The genome is a positive sense ssRNA. 1977 1978 41 . Coat protein has a single type of subunits with Mr 54 x 103.P. Historical review 2002 2003 2005 Cigsar et al. Boscia and G. A strain of this virus had been found previously in Portugal and described as virus CM112. De Stradis. 2005. Virus Genes 30. 2003. M. 471-475 Gokalp K. GBLV is a typical nepovirus belonging in subgroup C of this genus but its vector is not known. Bulletin OEPP/EPPO Bulletin 32. Martelli.: Complete nucleotide sequence of GARSV RNA-2 3.1 x 106 (RNA-2). Virus particles are about 30 nm in diameter and sediment as three components (T. Cigasr. Concord in New York State. Martelli.P. Martelli. I. Journal of Plant Pathology 85. Uyemoto et al. but different from GBLV. isolated in 1970 in Portugal from symptomless vines.2 x 106 (RNA-1) and 1. Kizlar tahtasi showing mild fanleaf-like symptoms Gokalp 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.: Description and thorough characterization of GARSV identified as a new species in the subgroup B of the genus Nepovirus Abou Ghanem-Sabanadzovic et al. Cigsar I. N. is not seed borne and was reported only from south-eastern Turkey. Component T is made up of empy protein shells. physico-chemical and serological characterization of the virus and assignment to the Nepovirus group (now genus Nepovirus). accounting for 39% (componet B1) and 42% (component B2) of the particle weight. Sanitary status of grapevine in south eastern and central Anatolia. B1. and B2). wt of 2. Historical review 1972 1972 1977 Ferreira and De Sequeira: Description and preliminary characterization of an unidentified virus denoted CM112. Digiaro and G. 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. By contrast. 335-340. 2. The virus occurs as different closely related but serologically distinguishable strains. Two isolates of GBLV have been transmitted by mechanical inoculation to seedlings and rooted cuttings of several grapevine cultivars without inducing symptoms. Abou Ghanem-Sabanadzovic. consisting of two separately encapsidated molecules with Mol. The virus supports the replication of a satellite RNA with Mol. Properties of a previously undescribed nepovirus fron south-east Anatolia. The scattered distribution of infected vines in the field suggests that the virus is spread primarily by infected propagative material. References Abou Ghanem-Sabanadzovic N.P.: A virus serologically related to GBLV isolated from Vitis labrusca cv. 35-41.: First isolation by mechanical transmission of an unknown nepovirus from cv. whereas components B1 and B2 contain RNA.: Description of GBLV.: GBLV description in the CMI/AAB Descriptions of Plant Viruses series. wt 0. 2. Martelli et al.95-2. Martelli et al. The virus can be detected directly in grapevine leaf extracts by immunodiffusion in agar plates.: Isolation of virus CM112 from in vitro cultures of grapevine tissues. A. GBLV has also been recorded from Hungary and Yugoslavia. The economic importance of the virus is minor. where it is widespread and infects latently several grapevine varieties growing in widely separared areas..
21-24. Phytopathology 71.P.A. Savino and A.A. De Sequeira O. 1980. 1985. Ramsdell D. Colmar 1970. A. No. 143-145 De Mendonça A. V. A. Annales de Phytopathologie.: A comparative study of Bulgarian GBLV isolates and the Portuguese virus CM112 establishes that CM112 is a serologically close but distinguishable strain of GBLV. A manually transmissible latent virus of the grapevine from Bulgaria. Quacquarelli.: 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. Ganeva and M. Pocsai E. Niagara Falls 1980.. Cordoba 1976. Quacquarelli. O. 1980 1980 1980 1980 1981 1981 1982 1983 1985 1992 3.1979 Martelli et al. Vasconcelos-Costa.C. Dimitrijevic: GBLV found in Yugoslavia De Mendonça et al. De Sequeira. Quacquarelli and D.A. and O. 95-101. Phytopathologia Mediterranea 22. 135-141. The ultrastructure of grapevine Bulgarina latent virus infections in natural and artificial hosts. CMI/AAB Descriptions of Plant Viruses.N. D.Proceedings 7th Meeting of ICVG. Ferreira A.. D.. 1983. Some properties of grapevine Bulgarian latent virus. 113-120. Krastanova et al. 1980. Annals of Applied Biology 85. 1972. P. 1981. An immunoradiometric assay for the titration of a Portugese strain of grapevine Bulgarian latent virus (GBLV). 245-250. Krastanova S. 468-472.A.. Annales de Phytopathologie. Pereira and V. 1992. 1981. Gallitelli. Stace-Smith. De Sequeira. Pocsai: Occurence of GBLV in Hungary. Savino. Proceeding 4th Meeting of ICVG. Di Franco. M. 1979. References De Mendonça A. Possibilities for the whole-year ELISA detection of viruses infecting grapevines. Acta Phytopathologica Academiae Scientiarum Hungaricae 16. Ferreira. Ramsdell and Stace-Smith: The New York isolate of GBLV is a strain of BLMoV. Abracheva.P. O. M. Martelli et al. De Sequeira and Vasconcelos-Costa: Use of an immunoradiometric assay for the titration of the Portuguese strain of GBLV. 42 .A. Simoes. Abracheva. Applicability of immunosorbent electron microscopy (ISEM) for the detection and identification of CM112 virus in grapevine. 1972. M. Proceedings 6th Meeting of ICVG. Russo and V.P. and J. V. Gallitelli et al. A preliminary report. Mota. Dimitrijevic B. and R. A. De Sequeira. Properties of a distinctive strain of Grapevine Bulgarian latent virus. Varennes and De Sequeira: First application of ELISA for the detection of virus CM112. De Sequeira and A. Niagara Falls 1980. Colmar. 349-354. 217-222. Gallitelli D. Proceedings 7th Meeting of ICVG. P. Numéro hors série. Physical and chemical properties of the particles and ribonucleic acid of blueberry leaf mottle virus. quinoa. 27-32.: Detection of GBLV in grapevine leaf extracts by ISEM. Grapevine Bulgarian latent virus. Proceedings 7th Meeting of ICVG.Sur l'isolement d'un virus à partir de cultures de tissus de vigne.: Ultrastructural study of GBLV infections in grapevine and C. 1977. 1978. Gallitelli. Proceeding 4th Meeting of ICVG.P.A. Martelli G. Yankulova. D.A. V..: Detection of virus CM112 in grapevine leaf extracts by ISEM. Martelli G. Savino and A. 1980. Numéro hors série. 186 Martelli G. Rasteniev'dni Nauki 29. The Portuguese strain supports the replication of a satellite RNA. Gallitelli. Occurence of grapevine Bulgarian latent virus in Hungary... 51-58. 269-272. Russo et al. France.. Jankulova. Savino and O. Monografias INIA 18 . Some properties of the new latent virus from grapevine rootstocks in Yugoslavia. Garcia de Orta Estacao Agronomica 12. 1970..: Improvement of ELISA protocol for GBLV detection the whole year round. Niagara Falls 1980. Preliminary studies on an undescribed grapevine virus. Martelli G..
Kenten: GCMV is distantly serologically related to Cacao necrosis virus. early reports that this nematode could transmit the virus have not been confirmed. vuittenezi is very frequently found in vineyards with chrome mosaic patches. Pozsár et al. Tobacco pants and the rootstock 110R have been successfully transformed with the viral coat protein for induction of resistance.: HCMV adversely affects photosynthetical carbon dioxide fixation. 269-277. RNA-2 has Mol. GRAPEVINE CHROME MOSAIC VIRUS (GCMV) 1. sometimes together with X. The virus belongs in the same subgroup of TBRV (subgroup B) to which is distantly related serologically.: Host range and properties of a spherical virus.K. double nodes and short internodes. Croatia and Austria. Martelli and Sarospataki: X.A. Mali et al. However. called Hungarian chrome mosaic virus. It has been recorded also from Czechoslovakia. 1966 1968 1968 1969 1969 1971 1972a 1972b 1972 1975 1977 43 . Although GCMV particles have been detected by immunosorbent electron microscopy in Xiphinema index fed on infected hosts. Description Grapevine chrome mosaic virus (GCMV) was first found in Hungary. 949-953.: GCMV recorded from Slovakia and report of X. near Lake Balaton. Isolation and identification of a strain of grapevine Bulgarian latent virus in Concord grapevines in New York State. Lehoczky and Tasnady: A study of the effect of HCMV on yield and sugar content of infected grapevines.8 x 106 . and was originally called Hungarian chrome mosaic virus. Varennes A. The virus appears to be unrelated serologically to GFLV and is not transmitted by X. The coat protein has a single type of subunits of Mr 52 x 103. index as vector of the virus (unconfirmed results). a size of 4441 nt and accounts for 31% of the particle weight. The virus is not serologically related with GFLV. wt of 1. Affected vines lack in vigour and may decline and die. Jakó et al: HCMV affects pigment and sugar content of infected grapevine leaves. pretty much like GFLV. vuittenezi transmits GCMV or GFLV. wt of 2. 1982.: The isometric virus associated with Hungarian chrome mosaic is serologically distantly related to Tomato black ring virus (TBRV). Martelli and Quacquarelli: Description of HCMV in the CMI/AAB Descripitons of Plant Viruses series. Martelli: Purification and serology of the virus isolated from Hungarian grapevines with fanleafand yellow mosaic-like symptoms. Evaluation of short reaction times and re-use of a-globulin and conjugate. GCMV is transmitted through grapevine seeds. a symptom virtually indistiguishable from GFLV-induced yellow mosaic. Saric and Hranuelli: GCMV recorded from Croatia.K. symptomless infection may occur. 1977. Some virus strains induce leaf deformity. The genome is bipartite. Martelli et al.Uyemoto J. a size of 7212 nt and accounts for 40% of the particle weight. index. Historical review 1966 Martelli et al. index.F. Leaves of infected vines are partially or entirely bright yellow or whitish. E. Plant Disease Reporter 61. De Sequeira.. Taschenberg and D. Detection of CM122 latent grapevine virus by enzyme-linked immunosorbent assay (ELISA).6 x 106 . Hummer. transmitted to herbaceous hosts from Hungarian grapevines with symptoms similar to those of fanleaf and yellow mosaic. Virus re-named Grapevine chrome mosaic virus. No evidence that X. Martelli and Quacquarelli : Physico-chemical characterization of HCMV and comparison with TBRV. and O. RNA-1 has Mol. 2. Agronomia Lusitana 41.
Hilbrand. Lanneau and J. Le Gall. Candresse. Ravelonandro and J. Brault V. Vitis 34. M. Brault et al. Acta Horticulture 235. index are inconclusive. Lehoczky et al. 231-238. Bretout et al. Le Gall.: 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.: Description of a certification scheme for the production of virus-free propagating material in Hungary. T. Doz et al. Lázár et al: Seed transmission of GCMV in grapevine. Dunez.: GCMV detected by ELISA in infected field-grown vines. Lehoczky: Pinot noir and Jubileum 75 are good indicators for GCMV. No assessment of resistance made. Plant Molecular Biology 21.. The virus vector is yet to be identified. 89-97. Dunez.. Le Gall and J. Occurrence of grapevine chrome mosaic nepovirus in Austria. 258-262.: GCMV cross-protects Chenopodium quinoa from the severe apical necrosis induced by a TBRV strain. Brault V. 3. Nucleic Acid Research 17. Dimou et al. Lázár et al..: Transformation of roostock 110R with the coat protein gene of GCMV. T. Bretout C. Taylor and Brown: Results of GCMV transmission trials with X.: GCMV recorded from Austria. V. O.: Characterization of a GCMV strain and confirmation of its serological relationship with TBRV. 1989.: Up-to-date report on virus diseases of grapevines in Hungary and description of the clean stock programme implemented in the country.: Tobacco plants genetically engineered with the coat protein gene of GCMV are resistant to infection. Dunez.1979 1980 1980 1982 1984 1985 1985 1989 1989 1989 1990 1993 1993 1994 1994 1995 1995 1997 2000 Lehoczky et al. Brault et al. 7809-7819. O.: Complete nucleotide sequence of GCMV RNA-1.: GCMV and TBRV can recombine. R. M. Le Gall et al. References Brandt S. Nucleotide sequence and genetic organization of Hungarian grapevine chrome mosaic nepovirus RNA2. Genetically engineered resistance against grapevine chrome mosaic virus. O. L. Le Gall et al.: Detection of GCMV in leaf dips by ISEM. Himmler. Russo et al. Delbos. Further demonstration that the two viruses are related. Savino. Candresse. Brault. 1993. M. This finding does not imply vectoring capacity by this nematode. Detection of nepoviruses in ligneous grapevine material using IC/PCR. 127-128. Virus and RNAspecific molecular hydridization probes for two nepoviruses. Kölber et al. 1994. Candresse.M. Le Gall et al. 44 . 1989. Roberts and Brown: Detection of GCMV in X. T.: Development of molecular probes for GCMV detection. Journal of Phytopathology 142. and G. Dodd and Robinson: GCMV and TBRV are molecularly related. index extracts by ISEM. A. 1995. Laimer da Camara Machado and V. Dimou D. D'Onghia.P.
ArMV) in the seeds and seedlings of grapevine by ELISA. 1995. 1-7. Dunez. Detection of grapevine chrome mosaic virus in field-grown vines by ELISA.. Prémunition: une competition entre souches faibles et souches sévères pour un voie commune d'expression de symptômes. Martelli G.P. Beczner. J. Nucleotide sequence homologies among RNA species of strains of tomato black ring virus and other nepoviruses. Extended Abstracts 13th Meeting of ICVG. Martelli G. Candresse. with Special Reference to Vitis. Nucleic Acid Research 17. J.. 1285-1289. Hungarian chrome mosaic of grapevine and tomato black ring: two similar but unrelated plant viruses. Jakó N. 1971. Quacquarelli and J. Szonyegi and M. Doz B.. Lehoczky. Agrobacterium-mediated genetic transformation of grapevine somatic embryos and regeneration of transgenic plants expressing the coat protein of grapevine chrome mosaic nepovirus (GCMV). T. Sarospataki.C. 49-64. 172-173. 7795-7807. The purification and some properties of cocoa necrosis virus. 171-170. Quacquarelli. M. J. J. Kölber..Tasnady. 1969. Lehoczky and A. 129-134. Brault and J. Proceedings International Conference on Virus and Vector on Perennial Hosts. Kertgazdasag 22. Mikulás. Mali V. Lázár J. J. Sarospataki. Luntz. 1968. 1984. 1990. J. Phytopathologia Mediterranea 8. Vanek and V. Division of Agricultural Sciences. Caractérization d'une souche du virus de la mosaïque jaune crome de la vigne (GCMV) isolée en Hongrie de vignes non panachées. Lehoczky J. 236-237. University of California. 1970. Acta Phytopathologica Academia Scientiarum Hungaricae 3. Dunez. Jakó N. 169-170.. No. 3.R. Sarospataki and J. 1969.. M. Kenten R. L. Martelli G. Lehoczky J. Extended Abstracts 11th Meeting of ICVG. Host range and properties of a virus associated with Hungarian grapevines showing macroscopic symptoms of fanleaf and yellow mosaic. L.J. and A. 1731-1740.. Delbos and J. Annales de Phytopathologie. Hungarian chrome mosaic. Candresse and A. Lehoczky and A. Torregrosa . 1968. 123-141.) University of California. 1972. 1985. 1989. 389-401. Candresse and J. Bouquet. Kölber and J. 505. Phytopathologia Mediterranea 24. Preliminary report on purification and serology of a virus associated with Hungarian grapevines showing macroscopic symptoms of fanleaf and yellow mosaic. Sznyegi. G. Davis 1965. G. Effect of the grape chrome mosaic and grape fanleaf-yellow mosaic virus infection on the photosynthetical carbon dioxide fixation in vine leaves.P. 1966. Lehoczky. Sarospataki. Lehoczky and G. Phytopathologia Mediterranea 24. V.P. Robinson. In: Virus Diseases of Small Fruits and Grapevines. 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. Detection of grapevine chrome mosaic virus in naturally infected vines by indexing. T. G. Annals of Applied Biology 71..Dodd S. Journal of General Virology 65. The change of pigment and sugar content in the chrome mosaic virus infected leaves of grapevine. Adelaide 2000. 1993. 29-44. 1972a. Davis 1965. L. Grapevine virus diseases and clean stock program in Hungary.. 1985. Journal of General Virology 76.. A. Muranyi . 1982. Le Gall O. Acta Phytopathologica Academia Scientiarum Hungaricae 1. 58-72. Lehoczky and G. and A. 1994. Proceedings International Conference on Virus and Vector on Perennial Hosts. 45 . Lehoczky. Grapevine chrome mosaic virus. Kiserletugyi-Kozlemenyek 64 (1-3). 402-410. Montreux 1993... 102. J. 103.Ser. Sarospataki. Division of Agricultural Sciences. Quacquarelli.. 1975. Vitis 8. R. Cardin. Nouvelle évidence d'une parenté sérologique éloignée entre ce virus et celui deas anneaux noirs de la tomate (tBRV). Bojnansky. J. Pacsa and J. Vuittenez. Martelli G. A Handbook. 1966. S. L. NW Frazier (ed.M. Les Colloques de l'INRA 11. 135-140. University of California.. Division of Agricultural Sciences. Hajdú.P. 1979... GCMV. and G. Horváth. Dunez. and G.P. Nematodes of the family Longidoridae (Thorne 1935) Meyl 1960 found in Hungarian vineyards and virus transmission trials with Xiphinema index Thorne et Allen. and D. S. 1-130. Kölber M. Berkeley. E..J. 119-126.. 567-568. O. Lázár J. The effect of fanleaf and chrome mosaic virus diseases on yield and the fruit sugar content of grapevine.. Studies on the nitrogen metabolism of vines infected with yellow mosaic virus. Devergne. T. 185-192. 1966. Lehoczky J. Le Gall O. Martelli G. Weinberg und Keller 15. Pol'nohopodarska Veda . Y. Certification scheme for production of virus-free grape propagating material and its results in Hungary. Martelli G.H. Farkas. Lázár. Martelli G. Pozsár B..P. a serotype ot tomato black ring virus. A. CMI/AAB Descriptions of Plant Viruses. 2000. Annales de Phytopathologie 11. Kölber M. GFV-YM. Detection of some nepoviruses (GFV. 1972b. Serologische Verwandschaft eines mit dem ungarischen "chrome mosaic" vergesellschafteten Virus mit einem Stamm des "tomato black ring virus". Transmission by nematodes of some grapevine viruses occurring in Czecholovakia and Hungary. with Special Reference to Vitis. and S. Plant Science. S. Numero hors série. Lehoczky . Nucleotide sequence of Hungarian grapevine chrome mosaic nepovirus RNA1. Danglot. G. Quacquarelli. 165-173. 206-210. Lehoczky J. Kuszala and A. Le Gall O. Quacquarelli.P.
: Complete nucleotide sequence of GDefV RNA-2 3. The scattered distribution of infected vines in the field suggests that the virus is spread primarily by infected propagative material. 2003. 1980. S. Bulletin OEPP/EPPO Bulletin 32. Virus Genes 30. 251-257. and D. Journal of Plant Pathology 85. Oxon.. Digiaro and G. Boscia and G. Grapevine deformation virus. is distantly related serologically to ArMV but not to GFLV. Immunosorben electron microscopy for detecting saptransmissible viruses of grapevine.F Brown. Hranuelli. Particles are isometric c. Abou Ghanem-Sabanadzovic.4 x 106 daltons and apparent size of c. K. A.P. 2. 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. Martelli and V. 471-475 Cigsar I.P. Coat protein subunits have a Mr 53 x 103. 2. N. References Abou Ghanem-Sabanadzovic N. 335-340 Cigsar I. Martelli. 2005..: First isolation by mechanical transmission of an unknown nepovirus from cv.800 nt. M.J. 1977. identified as a new species in the subgroup A of the genus Nepovirus. The genome is bipartite. 46 . Sabanadzovic. Abou Ghanem-Sabanadzovic et al. 286 pp. Nematode Vectors of Plant Viruses.6 x 106 Da and RNA-2.The virus has no recognized vector. mol..). 149-151. Description Grapevine deformation virus (GDefV) was recovered from Turkish grapevines showing distinct fanleaf-like symptoms. M. 30 nm in diameter and sediment as three components. D. Taylor C.Russo M. Proceedings of the Conference on Excoriosis and Virus Diseases of Grapevine. M (particles contaning a molecule of RNA-2 with Mol. GDefV is readily detected by ELISA and PCR using primers designed on the coat protein sequence. is not seed-borne and reported only from south-eastern Turkey. wt of 2 x 106 daltons and apparent size of c. Martelli. and belongs in the subgroup C of the genus Nepovirus. The virus sediments as three components: T (empty shells).3 x 106 Da and a size of 3753 nt. a novel nepovirus from Turkey. Historical review 2002 2003 2005 Cigsar et al.: A mechanically transmissible virus was recovered by sap inoculation from Tunisian grapevines showing mild fanleaf-like symptoms. Proceedings of the 7th Meeting of ICVG. and T. M. De Stradis. No vector is known and no information is available on the distribution and economic importance of the virus.. wt of 2. 2002. including all those known to infect grapevine. E. GRAPEVINE DEFORMATION VIRUS (GDefV) 1.800 nt) and B (particles containing a molecule of RNA-1 with Mol.P. 6. Sanitary status of grapevine in south eastern and central Anatolia. Gokalp. Saric A. Martelli. wt of 1. Complete nucleotide sequence of RNA-2 of two Turkish nepoviruses.P. wt of 2.. Digiaro and G. Niagara Falls 1980. 35-41. Historical review 1991 Ouertani et al. showing leaf and cane deformations Cigsar et al. 1997.: Description and thorough characterization of GDefV. Digiaro. Investigations on grapevine viruses in Croatia. was isolated from a Tunisian grapevine with mild fanleaflike symptoms. The virus belongs in the subgroup A of the genus Nepovirus. distantly serologically related with ArMV. Description Grapevine Tunisian ringspot virus (GTRSV). G. GRAPEVINE TUNISIAN RINGSPOT VIRUS (GTRSV) 1. GTRSV is serologically unrelated to any of 19 nepoviruses tested. Savino. RNA-1 has a mol. CAB International. Mostar 1977. 5.
L. Krczal and T.. Brown.: Biological and physico-chemical characterization of the grapevine strain of RpRSV.. and differs from the type strain for it often sediments as if it were a single centrifugal component. These differences strongly suggest that the grapevine-infecting RpRSV may be a different viral species. Ebel R. Wardell J. The nucleotide sequence of RNA 2 of raspberry ringspot nepovirus. 1982. Murant A. Greco. A. 107-117. and sediment as three components (T. Minafra. Historical review 1970 1978 1978 1982 1992 1994 Vuittenez et al. Archives of Virology 126. Altmayer. and B). The coat protein has a single type of subunits with Mr 54 x 103.F. M. Properties of an unusual isolate of raspberry ringspot virus from grapevine in Germany and evidence of its possibile transmission by Paralogidorus maximus. Raspberry ringspot virus. RNA-2 is 3928 nt in size and contains a single ORF encoding a polypeptide with Mr of 124 kDa. The grapevine strain of this virus is serologically very distantly related to the two main serotypes. Scottish and English.J. Die Wein-Wissenschaft 37. Edwards and M. Description Raspberry ringspot virus (RpRSV) is a nepovirus belonging in subgroup A of this genus.A. Heat therapy of infected grapevines. References Blok V.6 x 106 (RNA-1) and 1. 283-300. The type strain of RpRSV is transmitted by Longidorus macrosoma but the grapevine strain is transmitted by Paralongidorus maximus. This strain differs considerably from the English type strain of the virus although is serologically closely related to it. Boscia. wt of 2. 1978. 16.F. Extended Abstracts 14th Meeting of ICVG. Journal of General Virology 73. D. C. Brückbauer H. have angular outline. W. G. 88-118. The virus has only been found in grapevine in western Germany.. FS4 is a good indicator. M. accounting for 29% (component M) and 43% (component B) of the particle weight.A. Murant: Description of RpBRSV in the CMI/AAB Plant Virus Description series Stellmach and Querfurth: Study of a strain of RpRSV isolated from cv. Molecular characterization of two German Raspberry ringspot virus isolates infecting grapevines and construction of full length infectious clones. No. Mayo. Particles are about 30 nm in diameter. Crop losses can be higher than 30 %.. The virus is transmitted by Paralogidorus maximus Ebel et al. Savino. 2189-2194.: Recovery of RpRSV from grapevines of Palatinate. McGavin. M. Wetzel. References Ouertani R. 2003.J.T. Rüdel and B. Castellano. Two strains of different virulence occur in the Palatinate. G. D. Annals of Applied Biology 124. Manoukian. A. 198. 1992. Properties of a previously undescribed grapevine nepovirus from Tunisia.3. A Schnabel. Elbling in West Germany. CMI/AAB Descriptions of Plant Viruses. Robinson. M.P. Jolly. consisting of two separately encapsidated molecules with Mol. Mögliche Beziehungen zwischen Virus und Symptomausprägung bei der Rebe. Symptoms are similar to those of fanleaf. RASPBERRY RINGSPOT VIRUS (RpRSV) 1. The viral genome is a bipartite positive sense ssRNA. 1992.: Nucleotide sequence of RpRSV RNA-2 Jones et al. D. Reustle.J. 2. V.A. 1994. 47 . Brückbauer: RpRSV can be distinguished from other nepoviruses on the basis of symptoms induced on Vitis idicator plants Blok et al.M. Locorotondo 2003. Jones A.: Sequencing and molecular characterization of two German isolates of RpRSV from grapevine 2003 3.6 x 106 (RNA-2). Martelli and N.C. G.
and an increase in graft failure. Joannes Seyve virus. then in Yugoslavia. Historical review 1963 1965 1966 Stellmach and Bercks: TBRV detected in rootstock Aramon x V. Joannes Seyve. about 30 nm in diameter have angular outline and sediment as three components (T. wt of 0.virus). The virus was detected in grapevines in the Niagara Peninsula. Vuittenez et al. The virus is a definitive nepovirus species classified in subgroup A of this genus. Rüdel and H Brückbauer. Tanne: Detection of TBRV by ELISA in Israel. is a strain of this virus. Kuszala. Losses are not known precisely.5 x 106 daltons and a size of 1327 nt. rings and lines on the leaves of recently infected plants. Greece. wt of 2. Stellmach and Bercks: Further investigations on TBRV in grapevine.65 x 106 (RNA-2) accounting for 44% and 37% of the particle weight. or directly in grapevine leaf extracts using bentonite flocculation test. The genome is a bipartite positive sense single-stranded RNA occurring as two separately encapsidated functional molecules with Mol. SLRV and TBRV found in grapevine in the Palatinate. Bercks: Comparison of three serological tests for detecting several viruses. Bercks and Querfurth: GFLV. Description Tomato black ring virus (TBRV) was first found in grapevines in Germany. RpRV and TBRV detected serologically in grapevine in West Germany. Israel.7 x 106 (RNA-1) and 1. J. known to cause severe damage to the grapevine variety Joannes Seyve in Ontario. Querfurth. ArMV. Ontario as the cause of severe damage to cv. Pathologie und Thermo-Labilität mehrerer Reben-Isolate des Himbeerringflecken-Virus (raspberry-ringspot. and Ontario (Canada). Stellmach: Review paper on TBRV in grapevine. respectively. Annales de Phytopathologie 2. mottling of the older leaves. Bercks and Stellmach: ArMV. either by agar gel diffusion with extracts of herbaceous hosts previously infected mechanically from grapevine. 128-136.. Vuittenez A. Weinberg und Keller 25. and B). Lehoczky and Burgyan: Occurrence of TBRV in grapevines in Hungary. 1967 1970 1970b 1976 1977 1980 1984 1984 1986 48 . Their coat protein consists of a single type of subunits with Mr of about 57 kDa.Stellmach G. Meyer et al. Rüdel: Transmission of TBRV to grapevine by Longidorus attenuatus. 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. TBRV supports the replication of a satellite RNA with Mol. The latex test is considered as the most sensitive and the least time consuming method. RRV and TBRV are not transmitted by contact of roots or foliage in the vineyard.: Nucleotide sequence of a TBRV satellite RNA. Stobbs and Van Schagen: First record of TBRV from Canada. 279-327 TOMATO BLACK RING VIRUS (TBRV) 1. Untersuchungen zur Serologie. The vector to grapevine is Longidorus attenuatus. Virus particles are isometric. its own subgroup. Turkey. including TBRV : bentonite flocculation test. but they can be high. M. Détection et étude selogique du virus latent des taches annulires du frasier (strawberry latent ringspot). TBRV produces a reduction in growth and yield. chlorotic spots. RNA-2 is 4662 nt in size encoding a polyprotein with Mr of 150 kDa.: RRV. 1970. 1978. RNA-1 is 7356 nt in size and contains a single open reading frame encoding a polypeptide with Mr of 254 kDa. 2. and G. latex test and barium sulfate test. M. riparia 143 A in West Germany.
2. 1517-1529. The virus is transmitted by Xiphinema diversicaudatum. Kreiah et al. References Akbas B.G.6 x 106 (RNA-1) accounting for 38% of the particle weight.A. RNA-2 is 3824 nt in size and encodes a single ORF expressing a polypetide with Mr of about 99 kDa. and 1. 1988. Meyer M. Bercks et al. Symptoms on affected European grapes are of the fanleaf type. Mayo and C. and B). Greif C. 1970.W. Kuszala. The virus supports the replication of a satellite RNA 1118 nt in size. Fritsch. Fritsch. 3-5. M. Fritsch. about 30 nm in diameter have angular outline and sediment as three components (T. Assignement of SLRSV to the new genus Sadwavirus. Journal of General Virology 65. Le Gall et al. Babini and Bertaccini: Electron microscope study SLRSV infections in plant tissues. Occurrence of tomato black ring virus on grapevine in southern Ontario. J. Journal of General Virology 69. 1257-1271 Stobbs L. Description Strawberry latent ringspot virus (SLRSV) has been isolated from grapevine in the Palatinate (Germany) and in northern Italy.6 x 106 (RNA-2). Complete nucleotide sequence of a satellite RNA of tomato black ring virus. Journal of Turkish Phytopathology 22. 1986. Annales de Phytopathologie 2. Savino et al. It was also detected in imported vines in Turkey and Portugal.: SLRSV recorded from grapevine in Italy.: Nucleotide sequence of TBRV RNA-2 Greif et al. Hemmer.1986 1988 1993 Meyer et al. du virus des anneaux noirs de la Tomate (tomato black ring). Détection et étude sérologique du virus latent des taches annulaires du Fraisier (strawberry latent ringspot).: Nucleotide sequence of SLRSV satellite RNA.: Nucleotide sequence of TBRV RNA-1 Abkas and Erdiller: TBRV recorded from grapevines in Turkey 3. Virus particles are isometric. Historical review 1974 1977 1981 1982 1982 1987 1993 1994 2005 Murant: Description of SLRSV in the CMI/AAB Descriptions of Plant vrises series. Their coat protein consists of two types of subunits with Mr 43 x 103 and 27 x 103. and J. Meyer M.. The genome is a bipartite positive sense singlestranded RNA occurring as two separately encapsidated functional molecules with Mol. 1984.: SLRSV and other nepoviruses isolated from grapevines in Germany Credi et al. Nucleotide sequence of tomato black ring virus RNA-1. 55-61. M. 49 .. Erdiller. The virus is a definitive species in the genus Sadwavirus. wt 2. Brückbauer: SLRSV can be distinguished from other nepoviruses on the basis of symptoms induced on Vitis idicator plants. 1575-1583. 1993. Hemmer and C.: SLRSV found in grapevine in Turkey. O.. et du virus des taches annulaires du Framboisier (raspberry ringspot) chez des vignes du Palatinat. O. Genus SADWAVIRUS STRAWBERRY LATENT RINGSPOT VIRUS (SLRSV) 1. Kreiah et al. Turkiye. Van Schagen. O. The nucleotide sequence of tomato black ring virus RNA-2. Vuittenez A. respectively. 1984. Hemmer and C. Brückbauer. M. 279-327. Canadian Plant Disease Survey 64. and G.: Nucleotide sequence of SLRSV RNA-2. Journal of Gerneal Virology 67. Rüdel and H. Researches on grapevine virus diseases and determination of their incidence in Ankara.
A. Sanfaçon... London. Eight Report of the International Committee on Taxonomy of Viruses (Fauquet.. T. 133-180. Viral aggregates induced by a distinctive strain of strawberry latent ringspot virus from grapevine.V. 2005. Turkey. Untersuchungen überdie Viruskrankheited der rebe unter besonderer Berüchsichtigung “atypischer Formen” der Reisigkrankheit.P. A. G.A.. Lehto. Bercks R. Phytopathologia Mediterranea 20.. Babini. 88-118. Murant A. Mögliche Beziehungen zwischen Virus und Symptomausprägung bei der Rebe. Wellink. Bertaccini. Kreiah S. 2527-2532. Le Gall O. 1982. Querfurth and M. Sequence analysis and location of capsid protein within RNA 2 of strawberry latent ringspot virus. 74. Strunk and J. 1163-1165. 1993. J. CMI/AAB Descriptions of Plant Viruses No. Iwanami. Genus Sadwavirus. 126. References Babini A. Weinberg und Keller 24. Yilmaz. Strunk. T Jones. Martelli. Savino V. Kreiah S. J. D'Onghia and M. G. Die Wein-Wissenschaft 37. Strawberry latent ringspot virus in grapevine.. Strawberry latent ringspot virus. 56-63. U. Journal of General Virology 75. Rüdel. In: Virus Taxonomy. Brückbauer H. Yoshikawa. L. 1977.. The nucleotide sequence of a satellite RNA associated with strawberry latent ringspot virus.M. FAO Plant Protection Bulletin 35.M. 1981. Maniloff.. 304-308. and Ball. Betti. Mayo.I. G. J. 50 . and A. Gelli. Phytopathologische Zeitschrift 104. 102-104.3. 1987.A. 1974. C. Bertaccini and C. H. Desselberger.R.. Cooper and G.. 1982. Credi R.I. Brückbauer. A distinctive isolate of Strawberry latent ringspot virus from grapevines in Italy. 1994. A. K. Karasev. M.F. A. Cooper. (eds) 799-802 Elsevier/Academic Press. Journal of General Virology. A. L..R.. Wetzel and N. T. H.
V. Alternative weed hosts that have epidemiological significance are known for ToRSV. are involved in the aetiology of North American grapevine degeneration and decline. poor fruit setting. This type of resistance is hypersensitivity. and/or ring spots) and distorted leaves. PRMV. straggly and shelled clusters. sedimenting as three components. induces fanleaf-like symptoms on leaves and canes. interspecific hybrids). Partial sequence of the 3' 51 . deperimento della vite. distortion of canes. are endemic in North America and thought to be native of the region. New York State and Ontario) own-rooted European grapes affected by Tomato ringspot virus (ToRSV) and Tobacco ringspot virus (TRSV) decline rapidly. BLUEBERRY LEAF MOTTLE VIRUS (BLMoV) 1. Long distance spread takes place primarily through infected propagating material. grapevine decline. labrusca. In cold climates (e. and poor fruit setting. Peach rosette mosaic virus (PRMV) in V. BLMoV. americanum sensu lato and PRMV by X. Nematicidal control of vectors is possible. California) yield but not vigour is affected. most V. TRSV is transmitted by X. ingiallimento nervale (Ital. malformation and mottling of the leaves. labrusca is also resistant to TRSV. the infecting virus and the climatic conditions. Virus particles are isometric. berlandieri or V. TRSV. ToRSV and BLMoV are also seed transmitted in grapes.g.). Vitis vinifera. Control: Use of virus-free propagating material and resistant rootstocks.). Their coat protein consists of a single type of subunits with Mr of about 54 x 103 Da. its main host. labrusca cv.GRAPEVINE DEGENERATION AND DECLINE (AMERICAN NEPOVIRUSES) 1. Description Main synonyms: Yellow vein. depérissement de la vigne (Fr. wt of 2. All these viruses. The genome is a bipartite. Description Blueberry leaf mottle virus (BLMoV) is named after the disease induced in highbush blueberry (Vaccinium corymbosum). Longidorus diadecturus and L. riparia.).e.15 x 106 (RNA-2). americanum sensu stricto. ELISA and molecular tools are useful for testing field-infected material. Varietal susceptibility: There are great variations in the susceptibility of Vitis species and cultivars. X. and poor fruit setting Agent: The above mentioned four distinct nepoviruses. PRMV. TRSV and PRMV. interestingly.) Main symptoms: Symptomatological responses of grapevines vary according to the species (i. jaunissement des nervures. californicum transmits ToRSV yellow vein strain. Concord it delays bud burst. rivesi transmit ToRSV type strain (decline). mottled (oak leaf pattern. Infected vines decline slowly over time. single-stranded RNA occurring as two separately encapsidated functional molecules with Mol . positive-sense. In warmer climates (Maryland. No vector is known for BLMoV.35 x 106 (RNA-1) and 2. about 30 nm in diameter with angular outline. labrusca causes a severe disease characterized by delayed bud burst. All roostocks and. V. A number of rootstocks containing V. BLMoV is a definitive nepovirus species assigned to subgroup C. Adernvergilbung (Germ. All other viruses are transmitted by longidorid nematodes: Xiphinema americanum sensu stricto and X. Bunches are small and straggly (Maryland's grapevine little berry) and leaves may show chrome yellow flecking along the veins (California's yellow vein). elongatus. but not conclusive. and ToRSV separately or in combination. rupestris plasma show field resistance to the northern US strain of ToRSV and to TRSV and PRMV. exhibiting stunted growth. which in blueberry is transmitted by pollen. 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. whereas in V. except for BLMoV which may have been introduced from Europe. little grape (Eng. Blueberry leaf mottle virus (BLMoV) infects latently European grapes. V. Transmission: These viruses are all transmitted by grafting and mechanical inoculation.
Healthy grapevines become infected when planted in soils from diseased vineyards. PRMV can also be introduced in a site by infected planting material and be spread by vectors to adjacent vines. The vector is unknown.Rasmdel and J. Vectors are the Dorylaimoid nematodes Xiphinema americanum sensu lato and Longidorus diadecturus. Physical and chemical properties of the particles and ribonucleic acid of blueberry leaf mottle virus. possibily non specific transmission by L. especially) and a number of American-French hybrids have been recorded. labrusca cultivars (Concord. mostly to vines adjacent to previously infected plants. quinoa (90% of infected seedlings). Taschenberg and D. wt of 2. The virus is transmitted through seeds in grapevines and C.C. PRMV is soil-borne. Hancock. quinoa.K. Stace-Smith.Use of resistant roostock hybrids and of certified planting material is recommended.4 x 106 (RNA-1) and 2. Clusters are straggly. Plant Disease Reporter 61. sedimenting as three components. 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. Crop losses up to 60% and death of susceptible V. PEACH ROSETTE MOSAIC VIRUS (PRMV) 1.. Occasional. 52 . which may lead to their death. D. 468-472. Infected grapevines show shortened and crooked shoots. D.5% of seedlings from seeds taken from diseased vines proved to be infected. PRMV is seed-borne in both naturally infected dandelion (4% of infected seeldlings) and in artificially infected C. Virus particles are isometric. and with extensive shelling of the berries. but not eradicate. 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. Concord grapes are apparently virus-free but 9. Grapevines (Vitis labrusca) are infected in New York State (USA) by a serologically distinct strain of the virus. smaller and fewer than normal . Warkentin. but identified as a strain of GBLV. Virus Research 33. As the virus is endemic and seed-borne in some perennial weeds Taraxacum officinale (dandelion). Hummer. when a vineyard is planted susceptible cultivars may become infected by nematode vectors. The genome is a bipartite positive sense single-stranded RNA occurring as two separately encapsidated functional molecules with Mol. about 28 nm in diameter with angular outline. Vines are stunted and show a progressive decline. which induces fanleaf-type symptoms and is distantly related to GBLV.F. Sequence analysis of the 3' temini of RNA 1 and RNA 2 of blueberry leaf mottle virus. but in highbush blueberry the virus is pollen-borne and suspected to be pollen-transmitted .K. elongatus has also been reported. mottled and variously deformed leaves and delayed bud burst. and R. 1994.. vector populations . Roguing of infected trees and preplanting autumn fumigation with high rates of fumigant injected at two depths (15-20 cm and 75-90 cm) can effectively reduce. where the disease occurs in more or less circular patches and spreads slowly. respectively. The virus is a definitive nepovirus species assigned to subgroup C. 145-156 Ramsdell D. Their coat protein consists of a single type of subunits with Mr of about 57 x 103 Da. 1977. 949-953. E. 1981.termini of both RNA molecules has been determined. References Bacher J. The virus is seed-transmitted in grapevines and C. : Partial nucleotide sequence of BLMoV RNA-1 and RNA-2 1981 1994 3. 2. Historical review 1977 Uyemoto et al: BLMoV isolated from New York 'Concord' vines showing fanleaf-like symptoms. and has no economic importance. Isolation and identification of a strain of grapevine Bulgarian latent virus in Concord grapevines in New York State.W. Phytopathology 71. Description Peach rosette mosaic virus (PRMV) is named after the disease induced in peach. Solanum carolinense (Carolina horse nettle) and Rumex crispus (curly dock).2 x 106 (RNA-2) accounting for 44% and 37% of the particle weight. RNA1 is 8004 nt in size and contains a single open reading frame encoding a polypeptide with Mr of 240 kDa.F. one of its crop plant hosts. Uyemoto J. Pollen grains of cv.
and B. Annales de Phytopathologie. Lammers et al.: Longidorus diadecturus transmits PRMV to grapevines. 1972a.A Ebsary. Ramsdell and Myers: Field spread of PRMV is associated with the presence of infected weeds (T. tomato black ring and peach rosette mosaic viruses by an Ontario population of Longidorus elongatus. Rasmdell et al. Dias: Grapevine and peach strains of PRMV can be differentiated serologically. officinale. 16-18. Ramsdell and Myers: Description of PRMV-induced grapevine degeneration and association of X. 1978 1979 1980 1982 1983 1984 1985 1988 1988 1995 1998 1999 3. Transmission of peach rosette mosaic virus to peach grape and cucumber by Longidorus diadecturus obtained from diseased orchards in Ontario. Strains of peach rosette mosaic virus differentiated by cross absorption and immunodiffusion tests.R. Canadian Journal of Plant Pathology 4.R. 1984.F. 29-32 Allen W. R. 1972b. Numero hors sèrie..G Van Schagen and E. The characterization of a virus responsible for peach rosette mosaic and grape decline in Michigan. 53 . 1228-1239.G Van Schagen and B. Dias H. Canadian Journal of Botany 54. J. Purification and some characteristics of peach rosette mosaic virus (grape isolate).F. Annales de Phytopathologie.: Nucleotide sequence of RNA-1 of the grapevine strain of PRMV. 1988. Allen et al. 1982.. Historical review 1972a 1972b 1974 1976 Dias: Preliminary characterization of the grapevine isolate of PRMV. J. 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. S.S Everleigh. Transmission of raspberry ringspot. Allen et al.. The virus is seedborne in C. References Allen W. quinoa and has reproduced in part the field syndrome when inoculated mechanically to Concord grape seedlings. americanum with the disease. Canadian Journal of Plant Pathology 6. Dias and Allen: Physico-chemical characterization of PRMV.. Ramsdell and Gillet: Description of PRMV in the AAB Descriptions of Plant Viruses.R. Ramsdell: Review article on PRMV.: Xiphinema americanum is an efficient vector of PRMV. crispus) and transmission through grapevine seeds. Transmission of peach rosette mosaic virus by Ontario populations of Longidorus diadecturus and Xiphinema americanum (Nematoda: Longidoridae). Ramsdell et al: Use of ELISA for PRMV detection in grapevines. Carolinense. Canadian Journal of Plant Pathology 10. Dias H. Ramsdell and Gillet: List of grapervine cultivars and roostocks showing differential susceptibility to PRMV. Dias and Cation: Biological characterization of the grapevine strain of PRMV. 1-5 Allen W. and D. 105-106.2. Numero hors sèrie. Dias H. Cation. 97-103. Allen and Ebsary: Longidorus attenuatus transmits PRMV non specifically and with low efficiency.F. 1976.: Investigation on the susceptibility to PRMV infection of American and European grapevines and hybrid rootstocks.A Ebsary.
Peach rosette mosaic virus decline.C. and J. Ramsdell D. A comparison between enzyme-linked immunosorbent assay (ELISA) and Chenopodium quinoa for detection of peach rosette mosaic virus in 'Concord' grapevines.R. R. 1985. Lammers A. Bird GW. and W. The virus supports the replication of a circular satellite RNA 359 nt in size. M.C. J. Coat protein consists of a single type of subunits with Mr of about 57 x 103 Da. Ramsdell D. Gillet and L.7 x 106 (RNA-1) and 1.M.C. 1747-1754. Historical review 1970 1977 1985 1985 1986 1990 Gilmer et al. 1174-1178. Bird. French hybrids and European grape cultivars to infection by peach rosette mosaic virus.M. Susceptibility of American grapevine scion cultivars and French hybrid roostocks and scion cultivars to infection by peach rosette mosaic virus. Stace-Smith: Description of TRSV in the AAB Descriptions of Plant Viruses series. Characterization of the single protein and the two nucleic acids of peach rosette mosaic virus. RNA-1 is 7514 nt in size and contains a single open reading frame encoding a polypeptide with Mr of 225 kDa.C Ramsdell. Phytopathology 64. is endemic in Central and Eastern North America.W. Rose. Ramsdell D. Superimposed shallow and deep soil fumigation to control Xiphinema americanum and peach rosette mosaic virus reinfection a Concord vineyard.F Allison and D.M. 625-627. 1979. about 30 nm in diameter with angular outline.C. 364. . Ramsdell D.C. TOBACCO RINGSPOT VIRUS (TRSV) 1. Virus particles are isometric. Plant Disease Reporter 63. respectively. accounting for 44% and 28% of B and M particle weight. Uyemoto et al. 57-73. Phytopathology 68.W. Ramsdell D. 4143. 1974. TRSV is soil-borne and is transmitted by Xiphinema americanum sensu stricto.3 x 106 (RNA-2). 1980. RNA-2 has been sequenced only in part. Plant Disease 79.C.C. Canadian Journal of Botany 58. 54 .. 1998. sedimenting as three components (T. but in European grapes responses are similar to those elicited by GFLV. R. There is no evidence of seed trasmission in the grapevine. Symptoms elicited by TRSV are the same as those of ToRSV in native cultivars.M. Andrews..C.: Survey of ToRSV and TRSV in Pennsylvanian vineyards. 1988. Myers. 51-52. Preventive control measures are the use of resistant roostock hybrids and of certified planting material. Relative susceptibility of American.L. Peach rosette mosaic virus. Gillet and G. G. Peach rosette mosaic virus.M. 154-157.M. 447-450 Ramsdell D.. Ramsdell D.E Morris. J.: TRSV agent of a new grapevine disease in New York State. St. 1995.: Nucleotide sequence of TRSV satellite RNA. American Phytopathological Society Press. and R.W. Plant Disease 67. 1999.C.Dias H. 1978.). Description Tobacco ringspot virus (TRSV) is the type species of the genus Nepovirus and the prototype of subgroup A. Buzayan et al. Gillet and C. No... 2. G. Cloning an sequencing of peach rosette mosaic virus RNA1. Gillet. American Vitis species reported to be resistant to ToRSV and TRSV. Myers. Epidemiology of peach rosette mosaic virus in a Concord grape vineyard.L. Virus Research 65.C. and B). Pearson and A. In: Compendium of Grape Diseases (R. Powell et al. TRSV has a relatively wide natural host range. symptomatology and nematodes associated with grapevine 'degeneration' in Michigan. Goheen eds. respectively.C. Foster and Morris-Krsinich: In vitro translation of TRSV RNA-1 and TRSV RNA-2 yields major polypeptides with Mr of 225K and 116K. AAB Descriptions of Plant Viruses. Allen. 1983. Gillet. The genome is a bipartite positive sense single-stranded RNA occurring as two separately encapsidated functional molecules with Mol. and R. wt of 2. Ramsdell D.: A review of viruses infecting grapevines in New York vineyards. and J.H. but was recorded from grapevines only in New York State and Pennsylvania. Paul. 74-78. Phytopathologia Mediterrenea 24.
S. Bruening. In California ToRSV affects the yield rather than the vine's growth. Bruening. Symptomatological responses vary according to the species (V. Virology 219.4 x 106 (RNA-2) accounting for 44% and 41 % of the particle weight. Disease named yellow vein.B. Virus and virus-like diseases affecting grapevines in New York vineyards. Foster R.A. and B). Zallua et al. Gilmer R. Vines grow vigorously but bear little or no fruit. 619-627. 186-199. 516-519. 2. Clusters are straggly. Plant Disease 74.. Morris-Krsinich. Infected vines have small.. RNA-1 is 8214 nt and RNA-2 is 7273 nt in size. Hewitt: Successful graft transmission of unfruitful vine condition. Keese and A. Virus particles are isometric.S.J. and with extensive shelling of the berries. Chemical cleavage of the 5'-linked protein of tobacco ringspot virus genomic RNAs and characterization of the protein-RNA linkage. American Journal of Enology and Viticulture 28.A. Kelts. 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. V. TOMATO RINGSPOT VIRUS (ToRSV) 1. J. No. B. Incidence of tomato rigspot virus and tobacco ringspot virus in grapevines in Pennsylvania. vinifera. Gooding and Hewitt: A mechanically transmissible virus found to be associated with yellow vein. ToRSV-induced decline affects European cultivars. AAB Descriptions of Plant Viruses. Powell C.L. Vines are stunted and show a progressive rapid decline.M. Virus Research 30.. Abawi. 1-8. wt of 2. where it occurs iin the region of the Great Lakes and in the Pacific seaboard from California to British Columbia. Two serological ToRSV variants are known to infect grapevines. californicum in California The virus can be introduced in a site by infected planting material and be spread by vectors to adjacent vines. Silva and S. Tobacco ringspot virus. 1985.: Nucleotide sequence of TRSV RNA-1 3.K. Virology 144. 335-349. Description Tomato ringspot virus (ToRSV) is a definitive species in the genus Nepovirus and the prototype of subgroup C.M. 131-136. Longenecker and L.. Gould. 1985. P. A new grapevine disease induced by tobacco ringspot virus. smaller and fewer than normal.. and B. interspecific hybrids). Singh.K.L. Vectors are the Dorylaimoid nematodes Xiphinema americanum sensu stricto and X. The virus has been occasionaly recorded from grapevines outside of North America. Uyemoto and L.A. The yellow vein strain of the virus is pollen-borne but is not transmitted through seeds. J. Phytopathology 60. mottled and distorted leaves and short internodes. Coat protein consists of a single type of subunits with Mr of about 58 x 103 Da. Gerlach G.M. 1986.R.8 x 106 (RNA-1) and 2. ToRSV has a relatively wide natural host range and is endemic in North America. sedimenting as three components (T.L. Buzayan J. J. Zalloua P. References Buckley B. respectively.. : Partial nucleotide sequence of TRSV RNA-2. Nucleotide sequence and in vitro expression of the capsid protein gene of tobacco ringspot virus. more severely in colder than in warmer climates. Both RNAs contain a single open reading frame encoding polypeptides with Mr of 244 kDa (RNA-1) and 207 kDa (RNA-2). Stace-Smith R. 1996. Virology 151. 1993. Uyemoto J. 1990. and the climatic conditions. 1970. rivesi in north American States and Canada and X. especially if self-rooted. contrary to the decline strain which is seed-transmitted. Synthesis and processing of the translation products of tobacco ringspot virus in rabbit reticulocyte lysates.M. labrusca. Preventive control measures are the use of resistant roostock hybrids and of certified planting material. The genome is a bipartite positive sense single-stranded RNA occurring as two separately encapsidated functional molecules with Mol. Nucleotide sequence of satellite tobacco ringspot virus RNA and its relationship to multimeric forms. 55 . 1977. the infecting virus strain. which often leads to death. Forer. J.M. 309. "yellow vein" being the characterizing syndrome of its infections. W. ToRSV is soil-borne. Cummins and G. 702-704.1993 1996 Buckley et al. Buzayan and G. about 30 nm in diameter with angular outline.
Allen and Dias: Physico-chemical characterization of ToRSV. Gilmer and Uyemoto: ToRSV agent of a decline of Baco noir in New York State. Uyemoto: Seed transmission of the decline strain of ToRSV. Allen et al.: Xiphinema rivesi identified as the main vector of ToRSV in Ontario vineyards. Rott et al. Stace-Smith: Description of ToRSV in the CMI/AAB Descriptions of Plant Viruses series. Podlekis and Corbett: ToRSV is the agent of little grape disease in Maryland.: Complete nucleotide sequence of ToRSV RNA-2. Rowhani et al: Description of sampling strategy for detection of ToRSV. Allen et al. Baumgartnerova and Subikova: ToRSV recorded form grapevine in Slovakia. Stace-Smith and Ramsdell: Review of nepoviruses of the Americas. Uyemoto and Gilmer: Spread of ToRSV through the soil of New York State vineyards recorded.: Complete nucleotide sequence of ToRSV RNA-1. Bays and Tolin: ToRSV found in grapevines in Virginia Powell et al.: Survey of ToRSV and TRSV in Pennsylvanian vineyards. Herrera and Madariaga: ToRSV recorded from grapevine in Chile. Rott et al. Martelli and Taylor: Review of nematode-borne viruses and their vectors. Martelli: Review of nematode-borne viruses of grapevines and their epidemiology. Teliz et al. American Vitis species reported to be resistant to ToRSV and TRSV. Piazzolla et al.: ToRSV found in grapevines in Taiwan.: List of grapevine roostocks and cultivars showing differential susceptibility to ToRSV in Canada. americanum (now X.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.: Transmission of the yellow vein strain of ToRSV by X. Li et al: ToRSV identified in China in grapevine seedlings grown from seeds imported from France.: Confirmation that the grape yellow vein and the the grape decline strains of ToRSV are serological variants of the same virus. Bitterlin and Gonslaves: ToRSV retained and transmitted by viruliferous Xiphinema rivesi stored for two years at 1-3°C. 56 . Yang et al. Cory and Hewitt: The yellow vein strain of ToRSV is not transmitted through seeds. Dias: Record of ToRSV in the Niagara peninsula.: A review of viruses infecting grapevines in New York State vineyards. Corbett and Podleckis: Ultrastructural study of ToRSV-infected grapevine tissues. Uyemoto et al. Gonsalves: ToRSV is irregularly distributed in infected vines but can be detected by ELISA. californicum).
Gonsalves D.F. Sampling strategies for the detection of grapevine fanleaf virus and the grapevine strain of tomato ringspot virus.H. Current Topics in Vector Research 5. Bitterlin M.V. 1963. and V. 57 . 1984. Identification of tomato rinsgspot virus in leafroll diseased grapevines.M. Rochon.A. their epidemiology and control. Gooding G. 408-411.B. 1992. J. Tomato ringspot virus. Presence and incidence of grapevine viruses in central Chile. M. 1982. H. Canadian Journal of Botany 55. Van Schagen. and W.R. Nepoviruses of the Americas. 131-166.V. L. Association of Xiphinema species with soil types ang grapevines infected with tomato ringspot virus in Ontario. Agricultura Tecnica 61. Xiang . 1962. J. Journal of General Virology 72. Savino..V. 1991.G. Spatial distribution of Xiphinema rivesi and persistence of tomato ringspot virus and its vector in soil. Gilmer R. Detection of tomato ringspot virus in grapevines: irregular distribution of virus. and H.F. V. R.L.. Li. Some grapevine viruses in pollen and seed.G.A. 1505-1514.C. Phytopathologia Mediterranea 24.. Chen. Gilchrist. Y.A. 98-101.B. Uyemoto J. Rott M. 658-663. 157-164. 2004. 1954. Incidence and geographic distribution of tomato ringspot virus in De Chaunac vineyards in the Niagara peninsula. Allen W. Corbett M.B. Ivanka pri Dunaji 4. Nucleotide sequence of tomato ringspot virus RNA 1. Distribution of viruses and their nematode vectors.E.W. Dias and J. Niagara Falls 1980. 275-277. Detection and identification of plant viruses for quarantine in China. Phytopathology 79. Susceptibility of grape cultivars and rootstocks to an Ontario isolate of tomato ringspot virus.. Piazzolla P. Phytopathology 72. Rott M. Rowhani A. A severe outbreak of virus-induced grapevine decline in Cascade grapevines in New York.. 290 Stace-Smith R. 1978. 47-64. Uyemoto. Canadian Journal of Plant Pathology 4.S Whei. Madariaga. Purification and serology of a virus associated with the grape yellow vein disease. Plant Disease 74. Tomato ringspot virus in "Baco noir" grapevines in New York. Dias. 1972. and W. Advances in Disease Vector Research 6.K. American Journal of Enology and Viticulture 13. 393-400 Hewitt W.. N.F. Longenecker and L.. and E.F.E.. Beijing 2004. Phytopathology 58. 1985. M.A. Incidence of tomato rigspot virus and tobacco rigspot virus in grapevines in Pennsylvania. 475-480. 1169.. Téliz D. Plant Disease Reporter 56. Rokni. and D.K. 1989. 1990. Van Schagen and B.G and V. and J. 1975. 1989. Herrera M. Canada. Lownsberry. Phytopathologia Mediterranea 24. Vitis 31.R. 663 Martelli G. 1985. 35-44. Gonsalves. Works of the Institute of Experimental Phytopathology and Entomology . 1968. Ebsary. Gooding G. 1-27. peach yellow bud mosaic. 196-203.P. Phytopathology 56. Castellano and D. M. 465-473. 44-50.G.3. and C.. identification.W. 1980.V. Plant Disease 71. Journal of General Virology 76.. Phytopathology 53. 151-189. 133-135. Bays D. 710.A. Zhang and Y. Martelli G. Forer. 1987. Nematologia Mediterranea 6. 13161320 Dias H. 1987. Plant Disease Reporter 61. Stobbs. Proceedings 15th International Plant Protection Congress.R.. Li M. 1977. G. Tremaine and D'A. Hewitt. 861-863.P. 702-704. 31-34.E. Cory L. A.F. Podleckis. J. Grape yellow vein: symptomatology. Walker and S. Lee and D'A. Transmission of tomato ringspot. Rochon.. 1995. Tomato ringspot virus associated with little grape disease of Vidal 256 grapevines. Musci. 1988. A comparison of Grapevine yellow vein virus and a Grapevine isolate of Tomato ringspot virus.B. L.C. 1966. References Allen W. Corbett. Hewitt.M. 1028-1037. Nucleotide sequence of tomato ringspot virus RNA-2.. and M.. 95-106. Properties of the single protein and two nucleic acids of tomato ringspot virus.J. and B. Incidence of tomato ringspot virus in grape in Virginia. Baumgartnerova H.F. 1982. Membrane-associated spherical particles in extracts and tissues of virus-infected grapevines. Powell C. Plant Disease 72. and the association of a mechanically transmissible virus with the disease. Stace-Smith R. CMI/AAB Descriptions of Plant Viruses. Taylor. 1977. Ramsdell. Proceedings 7th Meeting of IGVG.K.K. Some virus and virus-like diseases of grapevines. Podleckis E. Plant Disease Reporter 59. Bulletin of the California Department of Agriculture 43.. No. Grogan and B. 2001. 1993. Subikova. Allen W. Nematode-borne viruses of grapevine. and grape yellow vein viruses by Xiphinema americanum. Tolin.F. and S. 24-28.
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GLRaV-1.5 nm. 61 . i. the symptoms are similar. The fruits often mature late and irregularly. usually leaving a narrow green band along the primary and secondary veins. Leafroll decreases grapevine yield (by 15-20% in average) and affects negatively rooting ability. and with many cultivars. thus suggesting that there can be several causal agents. whereas the Mr of all other viruses ranges between 35 and 44 kDa. they are inferior in quantity and quality. graft take and plant vigour. GLRaV-3. These negative effects are reverted if the disease is eliminated by sanitation treatments. Leafroll is no less important than fanleaf in economic importance. Sizes deduced from the nucleotide sequence of the CP cistron are 22 kDa for GLRaV-2 . the same as the size of coat protein (CP) subunits. potassium depletion in the leaf blade and accumulation in the petioles. Particle length varies from 1400 to 2200 nm according to individual viruses. accartocciamento fogliare (Ital. and lowering of sugar content. reduction of protein content. GLRaV-5. vinifera. Description The first descriptions of grapevine leafroll date back to the mid 19th century. in autumn. and is probably the most widespread virus disease of grapevine. In the most severe cases. These symptoms progress towards the top of the canes as the season advances.). thus impairing carbohydrate translocation from foliar parenchymas.). Also the composition and aromatic profile of the musts are modified. Sieve elements are obliterated and crusheds. called grapevine leafroll-associated viruses (GLRaV). Enrolamento de la folha (Port. All GLRaVs belong in the family Closteroviridae. Virus particles are very flexuous filaments about 12 nm wide. GLRaV-2 in the genus Closterovirus. Agents: To date. differentiation of GLRaVs at the species level was by Roman numerals.) Main symptoms: In red-berried cultivars of Vitis vinifera reddish spots develop in the lower leaves in late spring or summer.GRAPEVINE LEAFROLL 1. enroulement (Fr. which are differentiated from one another by a progressive number. Main synonyms: White Emperor disease (Eng. The leaf blade becomes thick. enrollado (Sp. In most cases. accartocciamento. the risk of disseminating the disease is great if untested rootstocks are used. Starch accumulates in degenerated chloroplasts causing increased thickness and brittleness of the leaf blades. exhibiting open structure and distinct cross banding with a pitch of about 3. Also plant anatomy is affected. GLRaV-2 CP has a Mr of 24 kDa. have been found in leafroll-infected vines. most of the leaf surface becomes reddish. GLRaV-6. but the leaves become chlorotic to yellowish. depending on the climate and geographic location. These spots enlarge with time and coalesce so that. differing somewhat in aspect and in severity. Hence. Careful observation of field symptoms in infected vines reveals that there are several types of leafroll. and low in sugar. and GLRsV-9 in the newly established genus Ampelovirus. A number of other physiological parameters are affected. GLRaV-4. GLRaV-8. Until 1995. whereas GLRaV-7 is presently classified as unassigned species to the family. Blattrollkrankheit (Germ. 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. In white-berried cultivars of V. respectively.). nine different viruses with filamentous particles.). as estimated by polyacrylamide gel electrophoresis. instead of reddish.). Other such viruses may exist. infection of rootstocks is symptomless. especially the phloem. Rollkrankheit.e. except for a variable decrease in vigour. the whole leaf surface becomes deep purple. 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. brittle and rolls downwards. changes in the pattern of peroxidase and polyphenoloxidase isoenzymes. Its occurrence in viticultural areas is worldwide. enrollamiento de la hoja.
comstocki. The genome of GLRaV-1 is 17. but it does not discriminate between GLRaVs and was reported to be less sensitive than ELISA. The genome is a monopartite. American rootstocks are usually symptomless carriers of GLRaVs. 29. and epidemiologically) from most of the known closteroviruses. GLRaVs differ in various vays (molecularly. vinifera. All GLRaVs can be identified by serological and nucleic acid-based techniques. with none of which they are serologically related. 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. 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. which is the type species of the novel genus Ampelovirus has a genome 17. Pseudococcus longispinus. biologically. singlestranded. Foliar tissues are not recommended for serological GLRaVs detection in American Vitis species and roostocks. viburni and Ps. Composite samples should be used to minimize false negative responses that may originate from the unven distribution of GLRaVs in chronically infected vines. GLRaV-2. the only member of the group to be mechanically transmissible. and some are commercially avaliable. Its soft scale insects vectors are Pulvinaria vitis and Neopulvinaria innumerabilis. roostocks. classical double antibody sandwich ELISA (Chromo-ELISA) or Lumino-ELISA. longispinus. Ps. GLRaVs show molecular variations which give rise to a population of strains. None of the GLRaVs is known to be seedborne. and degenerate primers have been designed and successfully used for PCR detection of virtually all GLRaVs. Regardless of whether they belong to the genus Closterovirus or Amplelovirus. Detection: In many cases. GLRaV-3. or by molecular techniques. Pl. broadspectrum. The genome of GLRaV-4 and GLRaV-9 are incompletely sequenced but show a structural organization compatible with that of the genus. vinifera varieties show symptoms most clearly because of the reddening of the leaves. only vectors of GLRaV-1. and Neopulvinaria innumerabilis. Symptom expression depends on the variety. American Vitis species and rootstocks are the best antigen sources for serological assays. GLRaV-3. GLRaV-5 and GLRaV-9 have been identified.5 kDa for GLRaV-4 .35 kDa for both GLRaV-3 and GLRaV-1. or the hybrid LN 33 is still the most popular method for identifying the disease. climate. 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. Spread at a site is mediated by mealybug and soft scale insect vectors. Indexing on red-fruited cultivars such as 'Cabernet sauvignon'. Other GLRaVs may exist in nature as suggested by reports. the type member of the genus. contains 13 ORFs. Mealybug vectors of GLRaV-3 are Planococcus ficus. GLRaV-1 is transmitted in nature by the pseudococcid mealybugs Heliococcus bohemicus and Phenacoccus aceris and the soft scale insects Pulvinaria vitis. 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. Natural field spread of leafroll disease has been reported from many countries in Europe and elsewhere. vinfera and cortical shavings from mature dormant canes of V. Leaf tissues or petioles from mature symptomatic leaves of V. The presence of high molecular weight double-stranded RNAs (dsRNA) in phloem tissue extracts can be used as infection marker. Ps. As to nucleic acid-based assays. GLRaV-3.Transmission is semipersistent and does not appear to be vector-specific. This has been ascertained experimentally for GLRaV-1. Parthenolecanium corni. Varietal susceptibility and sensitivity: No immune variety or rootstock is known. affinis. Membranous vesicles can derive eitrher from peripheral vesiculation of mitochondria followed by disruption of the organelles (GLRaV-1.528 nt in size. maritimus. citri. Disappearance of dsRNAs from vines submitted to 62 . has a number of minor biological variants which can be differentiated by the reaction of inoculated Nicotiana species. contains nine open reading frames (ORF) and has structural organization identical to that of Beet yellow virus (BYV). So far. These reagents are routinely used for ISEM. GLRaV-5 and GLRaV-9 are both transmitted by Ps. ultrastructurally. The genome of GLRaV-2 is 15. Polyclonal antisera and/or monoclonal antibodies have been raised to each single GLRaV . in agreement with the quasispecies nature of viruses. and some of them are used as indicators. calceolariae. 'Cabernet Franc' 'Pinot noir'. 'Merlot'. and has structural organization differing from that of other sequenced closteroviruses. GLRaV-5 and GLRaV-9 are phylogenetically close to one another but are serologically unrelated. grafted vines) which is largely responsible for its dissemination over medium and long distances.919 nt in size. Red-berried V. soil condition and probably. Ps.647 nt in size and contains 10 major ORFs. leafroll can be detected by its symptoms in the field on red-fruited varieties. Ps. GLRaV-2 and GLRaV-3. number an types of infecting viruses. positive sense RNA molecule.
vinifera and there is no published information on how to protect healthy stocks from vector-mediated reinfection in the field.: Studies on the effects of leafroll on yield of grapevines in California. Belli et al. a grapevine disorder similar to leafroll Scheu: Demonstration of graft transmission of leafroll from diseased to healthy Vitis vinifera. Goheen et al. Fraser: Leafroll in Australia. Introduction of transgenic resistance to GLRaV-2 and GLRaV-3 is being attempted by engineering different viral genes into rootstocks and European grape cultivars.: Transmission of a virus to herbaceous plants from a leafroll-infected vine in Israel. Scheu: Leafroll is widespread in German vineyards. Chamberlain: Leafroll in New Zealand. unless they are hybridized with virus-specific probes. Lehoczky et al. Later studies showed that the virus is an occasional contaminant Lider et al. Control: Production and use of clonally selected and sanitized propagation material is very effective and the only preventive method for leafroll control available. 2. However. Ravaz and Verge: Occurrence in France of “rougeau”. roostocks are suggested as the major souces of leafroll dissemination. Hoefert and Gifford: Study of the effects of leafroll infection on vine anatomy. dsRNAs cannot be utilized for virus identification. Vuittenez: Leafroll in France. Hewitt: Leafroll in California Goheen et al.: Leafroll virus can be inactivated in vivo by heat therapy. Hypothesis of the viral origin of leafroll. Blattny et al. Tanne and Nitzany: Leafroll in Israel Tanne et al. Mendgen: Presence of filamentous particles in grapevines with symptoms of flavescence dorée in West Germany.: Leafroll in Czechoslovakia.: White Emperor and leafroll are identical diseases.: Leafroll in Italy.sanitation treatments is regarded as evidence for successful virus elimination. Harmon and Snyder: The "White Emperor" disease is graft-transmissible and is regarded a virus disease. Bovey: Leafroll in Switzerland. Dimitrijevic: Leafroll in Yugoslavia. a grapevine disorder similar to leafroll. No sources of resistance are known in V. 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”. These particles are probably closteroviruses associated with leafroll. The incidence if the disease was less than 20% as compared with 80 to 100% in commercial vineyards. As no apparent spread of the disease was observed. Luhn and Goheen: Leafroll found in the original grapevine stocks imported from Europe into California in 1890.: Leafroll in Hungary. 1971 1973 1974 1975 63 .
: Closterovirus-like particles purified from leafroll-diseased grapevines in France. Absence of such particles in healthy grapevines.: Ultrastructural study of leafroll-infected grapevine tissues. Teliz et al. Abracheva: Leafroll in Bulgaria. previously found in grapevines in Switzerland. Zee et al. found in grapevines affected by leafroll. Gugerli et al. Presence of virus-like particles in thin sections of leafroll-diseased vines.: ELISA testing reveals that GLRaV-3 has an uneven distribution in grapevine tissues. Tanaka: Leafroll in Japan. Rosciglione and Gugerli: GLRaV-1 and GLRaV-2 with particles of 2200 nm and 1800 nm respectively. Namba et al. Zimmermann et al.: Studies on the cytopathology of leafroll-diseased grapevines. Production of rabbit and hen antibodies for ELISA and ISEM to GLRaV-1 and GLRaV-3.1975 1976 1976 1977 1979 Martelli and Piro: Evidence from a herbarium that leafroll occurred in Sicily in the second half of the 19th century. A large dsRNA molecule is consistently isolated from leafroll-diseased grapes. Von der Brelie and Nienhaus: Light and electron microscope study of cytopathological changes induced by leafroll in grapevines. Hu and Gonsalves: Monoclonal antibodies produced against GLRaV-3. Purification and serology of associated closterovirus-like particles. but not in similar praparations from healthy plants.: Closterovirus-like particles and specific dsRNA found in leafroll-diseased grapevines in New Zealand. Hofmann: Symptoms of leafroll in affected clones of Pinot noir and performance in West Germany. Sasahara et al. 1981 1981 1982 1982 1983 1984 1984 1984 1985 1986 1986 1987 1987 1988 1988 1989 64 .: Aggregates of closterovirus-like particles observed in thin sections of phloem from leafroll-diseased grapevines. Barlass at al. Martelli et al. but not in healthy controls. Mossop et al. A third closterovirus type called GLRaV-3. Castellano et al.: Elimination of leafroll by in vitro meristm tip culture and apex fragmentation.: Review on the detrimental effects of viral infection on grapevine physiology. Confirmation that GLRaV-3 and the New York closterovirus isolate cross react serologically.like particles. Kliever and Lider: Study of biochemical changes found in grapevine infected with leafroll in California. Suggestion that a closterovirus may be the agent of the disease. are also present in leafroll-affected grapevines from Italy. Corbett et al: Electron microscope observations by negative staining of leaf extracts from leafroll-diseased grapevines in South Africa showed the presence of closterovirus. Production of polyclonal antisera for use in ELISA.: 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.: First record of successful elimination of leafroll in grapevine by using meristem tip culture in Japan. Faoro et al.: 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. Antiserum against a New York isolate also reacted with GLRaV-3 from Europe.
GLRaV-1.: Production and characterization of monoclonal antibodies specific to GLRaV-3.: Leafroll and associated closteroviruses in China Engelbrecht and Kasdorf: Transmission of GLRaV-3 by Planococcus ficus from grapevine to grapevine in South Africa. symptoms are obtained within 20-70 days. 1990 1991 1991 1991 1991 1991 1991 1991 65 . but not GLRaV-1. vinifera varieties used as inoculum source. In Mexico leafroll. rupestris plasma. Transmission of GLRaV-3 by the mealybug Planococcus ficus. stem pitting and corky bark spread rapidly. Faoro et al.: Evidence of the irregular distribution of GLRaV-3 in American rootstocks. especially those containing V. whereas meristem tip culture yields up to 100 % sanitation Walter and Zimmermann: Further characterization of closteroviruses associated with leafroll in France.: Transmission of GLRaV-3 from grapevine to grapevine by the mealybug Pseudococcus longispinus in Israel. Boscia et al. 1989 1989 1989 1990 1990 1990a. Some vines indexing positive for leafroll do not react positively with any ofthe antisera.: Leafroll and associated closteroviruses in Chile Kuhn: Leafroll in Brasil Li et al.1989 1989 Tanne et al. Credi and Santucci: GLRaV-1 and GLRaV-3 cannot be detected by direct ELISA in leaves of graft-inoculated American rootstock. GLRaV-1 and GLRaV-2 were not transmitted. ficus fed on infected vines. Auger et al.: Further characterization of GLRaV-1 and GLRaV-3 by monoclonal antibodies. later in the leaves. For reliable testing. but they are easily detected in inoculated LN33 vines and in V. Identification of GLRaV-4 1990 1990 1990 1990 Walter et al. With leafroll. Agran et al.: Characterization of leafroll-associated closterovirus-like particles from grapevine using also monoclonal antibodies. Gugerli et al. b Hu et al. Savino 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. -2 and -3 are common whereas GLRaV-5 is rarely detected. Zimmermann et al. Gugerli et al. indicating the presence of other leafroll-associated viruses. Pseudococcus longispinus is present on weeds around diseased vineyards. Identification of GLRaV-5.: Detection of leafroll-associated closterovirus in recently infected grapevines in New York. There is evidence that other GLRaVs are involved in leafroll etiology. Good results in summer with extracts of basal leaves and in autumn or winter with wood shavings macerated in buffer. was detected in P.: Immunocytological detection and localization of GLRaV-1 and GLRaV-3 by immunogold labelling in grapevine thin sections. Heat therapy requires very long treatments and is only 20-30 % successful. Téliz et al.: Use of green grafting for detecting virus-like diseases of grapevine.: Comparison of heat therapy and meristem tip culture for eliminating GLRaV-3 from Italian grape varieties.: Production of monoclonal antibodies to GLRaV-1 and GLRaV-3. Gugerli: Review of grapevine closteroviruses. ELISA is to be applied to cortical scrapings rather than leaf tissues. GLRaV-2. The virus was detected in root tissues.
Minafra and Hadidi: Detection of GLRaV-3 in viruliferous mealybugs by PCR. Boehm and Martins: Leafroll in Portugal. Golino et al. Namba et al.: Leafroll and associated closteroviruses in Romania. Kassemeyer: Detection of GLRaVs in Germany. Belli et al.: Leafroll in Taiwan.b Saldarelli et al.: Mechanical transmission of GLRaV-2 and ultrastructural study of infected tissues of Nicotiana benthamiana. Gugerli and Ramel: Analysis by monolconal antibodies of a Swiss source of cv. Milkus et al. 1994 1994 1994 1994 1994 1995 1994 1995 1995 1995 1995 Merkuri et al.: Production of radioactive and non-radioactive molecular probes to GLRaV-3 from denatured dsRNA template and their use for virus identification.1991 Hu et al. Chasselas shows the prsence of two different GLRaV-2. Jordan: In a New Zealand commercial vineyard GLRaV-3 incidence increased from 9.: Leafroll and associated closteroviruses in Moldova. Habili et al.: Production of antisera to GLRaVs using electrophoretically separated coat protein subunits as antigens. Boscia et al. Gozsczynski et al. whereas the other assays are more suitable for analyzing samples that gave inconclusive results with ELISA. ISEM and dsRNA analysis. Pop et al. Krake: Characterization of leafroll disease based on symptoms shown by field-infected vines and graft-transmission tests.: Transmission of GLRaV-3 by the soft scale insect Pulvinaria vitis. Bondarchuk et al. Castellano et al. Katis et al: Leafroll and associated closteroviruses in Greece. later identified as GLRaV-2.: Leafroll and associated closteroviruses in Albania.: Revision of the nomenclature of GLRaVs and use of Arabic numerals in the species names. denoted GLRaV 2a and GLRaV 2b.: Comparison of different assay methods for detecting GLRaVs : ELISA. 66 .: Leafroll and associated closteroviruses in Ukraine.: Leafroll and associated closteroviruses in Spain 1991 1991 1991 1991 1991 1991 1992 1993 1993 1993 1993 1993 1993 1994a. Former GLRaV 2 is re-named GLRaV-6.: Purification and physico-chemical characterization of grapevine corky bark associated virus. Segura et al. ELISA is recommended for large screening.1% in 1992 Ioannou: Leafroll and natural spread of associated closteroviruses in Cyprus. Flak and Gangl: Leafroll and associated closteroviruses in Austria.1% in 1988 to 93.: Analysis for the presence of double-stranded RNAs can be used for assessing virus elimination following sanitation treatments. Observation of peripherically vesiculated mitochondria. Tzeng et al. Leafroll and associated closteroviruses in Yemen.: Transmission of GLRaV-3 by Pseudococcus affinis in California. Faoro and Carzaniga: Ustrastructural study of GLRaV-1 and GLRV-3 infections. Martelli et al.
Haidar et al.: Comprehensive review of the properties of GLRaVs.: Transmission of GLRaV-1 by the soft scale insects Parthenolecanium corni and Neopuvinaria innumerabilis. Krastanova et al. Martelli et al. Zhu et al.: Leafroll and associated closteroviruses in Palestine. Ling et al.: Elimination of GLRaV-3 by heat therapy improves agronomic performances of a Nebbiolo clone and the quality of the must. None of 223 accessions of European. Viruses are irregularly distributed in the vines.: Cloning an sequencing of the coat protein gene of GLRaV-3 and its expression in transgenic tobacco.1% in 1986 to 51. MacKenzie et al. Lahogue and Boulard: Search for genes of resistance in grapevines. Fortusini et al.: Leafroll and associated closteroviruses in Lebanon.: The sequenced genome of GLRaV-2 has the same structural organization of that of Beet yellows virus. Al-Tamimi et al. Gozsczynski et al. 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1998 1998 1998 1998 1998 1998 1998 67 .1995 1996 1996 1996 1996 1996 Greif et al. Alkowni et al.: Leafroll and associated closteroviruses in Jordan.: GLRaV-3 detected in native American vines in Western New York. GLRaV-3 appears to be a typical closterovirus. Habili and Nutter: In an Australian commercial vineyard GLRaV-3 incidence increased from 23. the type specied of the genus Closterovirus. American. Wilcox et al. Routh et al. Gugerli et al.: Use of degenerate primers for PCR detection of GLRaV-4 and GLRaV-5.: GLRaV-3 is transmitted by Planococcus citri in a semipersisten manner.9% in 1996.: Distribution and incidence of GLRaVs in Canadian viticultural districts. Cabaleiro et al.: Identification of GLRaV-7 and production of a polyclonal antiserum. Rowhani and Uyemoto: Comparative trials between indexing and laboratory detection methods show that the latter are more sensitve for GLRaVs detection. Ling et al. and Asian Vitis species inoculated by green grafting with a GLRaV-1 and GLRaV-3 sources were resistant.: Association of GLRaV-2 in Italy and France with a graft incompatibility revealed by Kober 5BB.: Serological characterization of GLRaV-6 and production of monoclonal antibodies. No vector was identified.: Identification of two different mechanically transmissibile strains of GLRaV-2.: Development of a spot-PCR technique for GLRaVs identification. Choueiri et al. Guidoni et al.: Extensive sequencing of GLRaV-3 genome.GLRaV-5 induces mitochondrial vesiculation.: GLRaV-2 and GLRaV-3 genes engineered in grapevine rootstocks to induce resistance. Petersen and Charles: Pseudococcus calceolariae acts as vector of GLRaV-3 in New Zealand. Faoro: Comprehensive review of the ultrastructure of GLRaVs infections. La Notte et al.
: Revision of the family Closteroviridae. Gugerli: Detection of GLRaVs by Lumino-ELISA. Ling et al. Fazeli and Rezaian: Partial sequencing of GLRaV-1 genome. Sforza et al. Gonsalves: Review article on the molecular traits of GLRaVs. : Partial sequencing of GLRaV-7 genome. Evidence of the quasispecies nature of the virus. Ps. GLRaV-2 and GLRaV-4.: Association of an unusual strain of GLRaV-2 with a graft incompatibility condition described from California as young vine decline.1998 2000 2000 2000 Saldarelli et al. transmitted by mealybugs and soft scale insects. Proposal for the establishment of Vinvirus (Ampelovirus) .: Evidence that vines sanitized from leafroll have superior qualitative and quantitative traits. Rowhani et al. a genus with monopartite RNA species. affinis and Planococcus citri transmit with various efficiency GLRaV-3 in California but not GLRaV-1.: 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. a new genus having GLRaV-3 as type species. Karasev: Review article on closteroviruses.: Mealybug species Pseudococcus longispinus. maritimus. Zhou et al. Seddas et al. Boscia et al.: Use of degenerate primers for PCR detection of GLRaV-1 and GLRaV-7. Monis: Identification of GLRaV-8 and production of monoclonal antibodies. one of which is especially suited for ELISA testing. Cardinal in Italy and Greece. Establishment and description of Ampelovirus . Little et al. viburni. Abou Ghanem -Sabanadzovic et al. Martelli et al.: Identification of hypervariable regions in GLRaV-1 genome.: Leafroll and associated closteroviruses in South Korea. Castellano et al: Ultrastructural study of GLRaV-2 and GLRV-7 infections. Demonstration that the membranous vesicles accumulating in the cytoplasm derive from proliferation of the endoplasmic reticulum.: Completion of the nucleotide sequence of GLRaV-3 genome and use of the HSP90 and coat protein genes for producing transgenic rootstocks.: Completion of the nucleotide sequence of GLRaV-2 genome. Turturo et al. Digiaro et al. Golino et al.: Identification and partial characterization of a new strain of GLRaV-2.: New monoclonal antibodies to GLRaV-2. Production of a new set of monoclonal antibodies.: Intriguing association of GLRaV-6 with cv.: Survey of GLRaVs in Mediterranean and Near East countries. Kim 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 . a chemiluminometric enzyme-linked immuosorbent assay. Meng et al.: Evidence that GLRaV-1 and GLRaV-3 are serologically related based on the cross-reactivity of a monoclonal antibody raised to GLRaV-1. Ps. Golino et al.: New vectors of GLRaV-1 are the mealybugs Heliococcus bohemicus and Phenacoccus aceris and the soft scale insect Pulvinaria vitis. Mannini and Credi. Ps.
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A. 71-73. Plant Pathology 49. 110-113. M. Adelaide 2000.C.D.. 1989.A . Zhang . Martelli and B. Proceedings 9th Meeting of ICVG. Walter. Jelkmann and G. Sannino F. 356358. Teliz D. Isolation and partial characterization of two new viruses from grapevine.P. Plant Disease 71. Berlin. Martelli. 68-69. Saldarelli P. Yafeng. 1935. Proceedings 10th Meeting of ICVG. Cloquemin and B.P.. D. 1986.L. Ben-Dov and B. Gugerli 1989. Savino V. The occurrence of grapevine leafroll disease among the main grapevine cultivars and breeding stocks in Taiwan. Martelli. Savino and G.. Scheu G. 1997. Die Rollkrankheit des Rebstockes. 162-163. Minafra. Digiaro. I. and P. Zeitschrift für Pflanzenkrankheiten und Pflanzenschutz 89. Adelaide 2000. M. 17-18. A. Tanne E. 222-223. 8689.P. Golino. Annals of the Phytopathological Society of Japan 42. Turturo C. Sela and I.. Use of degenerate primers for partial sequencing and RT-PCR-based assays of grapevine leafroll-associated viruses. Tanne. Chen and D. V.L. Proceedings 9th Meeting of ICVG.. Rowhani A. Locorotondo 2003. 1993. 508-517. V. D.E. Rowhani. and P. Saldarelli P. Journal of the Japanese Society for Horticultural Science 50. 2000. Teliz D. Tanaka S. 1981. M. Presence of grapevine leafroll in North West Spain. 135-141. Uyemoto.. W. G..S. Minafra. Zee. Von der Brelie D.A. 222-225 Tanne E. 125-126. Il rossore delle viti. Phytopathology 88. Kiryat Anavim 1987. D'Onghia and G. Y. 1994. Martelli. European Journal of Plant Pathology 104. Le rugeau de la vigne. 1989 Detection of a grapevine leafroll-associated closterovirus in recently infected tissues in New York and spread of the disease in Mexico. Transmission of grapevine leafroll disease and an associated closterovirus to healthy grapevine by the mealybug Planococcus ficus Signoret. Vitis 33. Segura A. Plant Disease 81. Saldarelli and A. Gonsalves. Digiaro. Hummer. Rosciglione B. Nitzany.. Walter. 176-180. 1906. 82. Minafra and M. T.. 1973.. Uyemoto. Turturo C. Plant Pathology 43. Der Deutsche Weinbau 14.P. Kiryat Anavim 1987. K. 2000. Zhang. Saldarelli P. Seddas A. Regeneration of plantlets by meristem tip culture for virus-free grapevine. Volos 1990. Phytoparasitica 17. 14. C. Extended Abstracts 14th Meeting of ICVG. 1974. Effect of heat therapy and meristem tip culture on the elimination of grapevine leafroll-associated closterovirus type III. 704-709. Reichnährstand Verlag. Raccah. 1994a. 192-196.P. P. and F. Routh G. P. 945-950.. Extended Abstracts 13th Meeting of ICVG. Rowhani A.. E. Vitis 12. Arboriculture. M. Maladies de l'enroulement et du bois strié de la vigne: analyse microscopique et sérologique. Hu and D. Extended Abstracts 13th Meeting of ICVG. Sasahara H. 1924. Adelaide 2000. M. Jacquet. Sforza R.. Guglielmi Montano and G.K. D. Cabaleiro. Y.. 1936. Preliminary investigations of genetic variability of Grapeviune leafroll associated virus 3 isolates. Extended Abstracts 13th Meeting of ICVG. Scheu G. 157-160. 2000. Partial molecular characterization and RT-PCR detection of grapevine leafroll-associated virus 7. and G.J. Rosciglione B. 1976. Progrès Agricole et Viticole 45. 35-38. 799-801. J. Revue Suisse de Viticulture.M. Y. Use of degenerate primers in a RT-PCR assay for the identification and analysis of some filamentous viruses. Non radioactive molecular probes for the detection of three filamentous viruses of the grapevine.P. Komar and C. Iri. 74 . Montreux 1993. Transmission of closterolike particles associated with grapevine leafroll by mealybugs (Pseudoccidae) in Israel.M. P. G. Gonzales and C.P. and J. Tanne E.. Haidar. Verge. A. Greif. Nienhaus. Virus diseses of grapevine in Israel. Harpaz. C. 1994b. 1982a. 67-69. Tada. Indexing grapes in Japan for viruses.S.. Rowhani.. Tzeng H. Detection of grapevine leafroll-associated closterovirus III by molecular hybridization. Field serological detection of viral antigens associated with grapevine leafroll disease. Extended Abstracts 11th Meeting of ICVG.Ravaz L. 207211.K. Rivista di Patologia Vegetale 1.K. Tzeng. Tazaki. 80-85. Histological and cytological studies on the infectious leafroll disease of the grapevine. and J. Rott. 2000. with special reference to closteroand vitiviruses of the grapevine. A comparison between serological and biological assays in detecting grapevine leafroll-associated viruses. Martelli. Takezawa and M. Transmission of grapevine leafroll virus to herbaceous plants. 156-167. 169175. D. Horticulture 18. Gonsalves and F. Plant Pathology Bulletin 3. 1998. 345-346. 1998. and F. New scale insect vectors of grapevine closteroviruses.. 433-436. M. Establishment of a relationship between grapevine leafroll closteroviruses 1 and 3 by use of monoclonal antibodies. 38. Saldarelli. Gugerli. 1991. 1238-1243. A. Phytopathologische Zeitschrift 80. Greif. 1987.. 91-96.. H.. A. 11-17. Boscia. Saldarelli. Mein Winzerbuch. D.
Journal of General Virology 79. Plant Disease 82. 1991. Production and characterization of monoclonal antibodies specific to closterovirus-like particles associated with grapevine leafroll disease. Bass.. R. Zee F. Transmission par greffage d'une virose type enroulement foliarie commune dans le vignobles de l'Est e du Centre-Est de la France. P. Cytopathology of leafroll diseased grapevines and the purification and serology of associated closteroviruslike particles. Purification de particules virales associées à l'enroulement de la vigne et mise au point d'un protocole ELISA permettant leur détection. Boscia. and G. Zhou Z.. Vesselle.Vuittenez A. D. 2000. 130. D. 62-66.V. McFerson and D. Z. B. K. 1289-1298. the closterovirus type member. D. Martin. Saldarelli. Walter B... Walter B. 1988. Zimmermann D. 1062.S. The use of a greengrafting technique for the detection of virus-like diseases of the grapevine. Zhou Z. Gonsalves. G. Extended Abstracts 13th Meeting of ICVG. Walter and M. B. Monoclonal antibodies for detection and characterization of Grapevine leafroll-associated virus 2. O. Turturo. Le Gall.S. Nucleotide sequence and genome organization of Grapevine leafroll-associated virus 2 are similar to Beet yellows virus. Zimmermann. 1990. Gonsalves. C. 1998 Leafroll virus is common in cultivated American grapevines in Western New York. Castellano. Phytopathology 77. 277-288. and D. Martelli. 1990. Proceedings 10th Meeting of ICVG. A. 731-741.. Potere. K. Production and characterization of monoclonal antibodies specific for Grapevine leafroll-associated virus 3 and epitope mapping of the coat protein. 2003. Further characterization of closterovirus-like particles associated with the grapevine leafroll disease. Comptes Rendues de l'Academie d'Agriculture de France 44. Goheen.. N. Journal of Phytopathology 128.W. Collas and G.A. Sommermeyer. Adelaide 2000.Y.. Wilcox F. Lee.Y. Van Regenmortel.F. R. Journal of Phytopathology 130. P.E Goszczynski. C.H.P. Locorotondo 2003. Legin. 137-145. Zhu H. Jiang and D. 75 . D. Kim. Gonsalves. Pool and R. Boscia. Vernoy. 203. J. 1998. Zimmermann D. O.R.E. Goszczynski and M. Walter and O. 1958. A. 1987. Volos 1990. Agronomie 8. Potere. R. Abou-Ghanem. 313-316.. Ling. D. Extended Abstracts 14th Meeting of ICVG. 1427-1434.
RUGOSE WOOD COMPLEX .
RUGOSE WOOD COMPLEX
The rugose wood complex consists of several diseases (Grapevine rupestris stem pitting, Grapevine kober stem grooving, Grapevine corky bark, Grapevine LN33 stem grooving) that are usually latent in ungrafted Vitis vinifera and American Vitis species and rootstock hybrids, but develop in grafted vines. Woody cylinder alterations resembling rugose wood symptoms are reported in the French literature of the early 1900s as possible physiological disorders. Rugose wood was first identified and described from southern Italy in the early 1960s as a graft-transmissible disease, and was considered to be a local problem until its discovery in Hungary in 1967. Now it is known to occur worldwide. 1. Description Main synonyms: Stem pitting, stem grooving (Eng.); legno riccio (Ital.); bois strié, cannelures du tronc (Fr.); madera rizada (Sp.); lenho rugoso (Port.); corky bark: rough bark (Eng.); suberosi corticale (Ital.); écorce liégeuse (Fr.); Korkrindenkrankheit (Germ.). Symptoms: Affected vines appear less vigorous than normal and may show delayed bud opening in spring. Some decline and die within a few years from planting. Grafted vines often show a swelling above the bud union and a marked difference between the relative diameter of scion and rootstock. With certain cultivars, the bark above the graft union is exceedingly thick and corky, has a spongy texture and a rough appearance, a condition known as "corky rugose wood". The woody cylinder is typically marked by pits and/or grooves which correspond to peg-and ridge-like protrusions on the cambial face of the bark. These alterations may occur on scion, rootstock or both. The severity of wood symptoms vary according to scion/stock combinations. Climatic conditions may have a bearing on symptom expression for under cool and wet climates symptoms are milder or absent. Cases of latent infection in grafted vines are not rare. By contrast, self-rooted European grapes and, sometimes, American roostocks, can show wood alterations. No specific symptoms are seen on the foliage, although certain cultivars show rolling, yellowing or reddening of the leaves similar to those induced by leafroll. Bunches may be fewer and smaller than normal and the crop reduced by 20-30%. The four diseases of the rugose wood complex can be recognized and sorted out by graft transmission to the indicators Vitis rupestris, LN 33 and Kober 5BB: a. Rupestris stem pitting. Distinct basipetal pitting limited to a band extending downwards from the point of inoculation in V. rupestris. LN 33 and Kober 5BB remain symptomless. b. Corky bark. Grooving and pitting of the entire surface of the stem of V. rupestris and LN 33, but no symptoms in Kober 5BB. Severe stunting of LN 33 is accompanied by rolling and reddening of the leaves and by most typical internodal swelling of the canes. c. Kober stem grooving. Marked grooving appear on the stem of Kober 5BB; no symptoms in V. rupestris and LN 33. d. LN 33 stem grooving. Grooves occur on the stem of LN 33, much the same as with corky bark, but no internodal swelling of the shoots nor foliar discolorations are present. V. rupestris and Kober 5BB show no symptoms. Agents: Putative agents of individual diseases of the rugose wood complex are members of the genera Vitivirus or Foveavirus, family Flexiviridae, i.e. viruses with flexuous filamentous particles from about 730 to 800 x 12 nm, with distinct transverse cross banding. Vitiviruses and foveaviruses are phloem-restricted in grapevines, but whereas vitiviruses are mechanically transmissibile to herbaceous hosts, though with difficulty, foveaviruses are not. The genome of all viruses consists of a single species of single-stranded positive sense RNA with Mol. wt 2.6-3.05 x 106 that accounts for c. 5% of the particle weight. Coat protein subunits have a single size and Mr of 22-28 kDa. Rugose wood-associated viruses have a worldwide distribution. Records exist from Europe, the Mediterranean basin, Near and Far East, Australasia, South Africa, and North and South Americas. Grapevine rupestris stem pitting-associated virus (GRSPaV), a definitive member of the genus Foveavirus, is the associated agent of Grapevine rupestris stem pitting disease. Virus particles are about 730 nm in length and are not readily observed with the electron microscope. GRSPaV occurs in nature as
a family of molecular variants. Viral RNA, which has been totally sequenced, has a Mol. wt of about 3.05 x 106 Da and a size of 8726 nt. The viral genome comprises 5 or 6 ORFs encoding, in the order, the replication-associated proteins (244 kDa), movement proteins (triple gene block, 25, 13 and 8 kDa) and the coat protein (28 kDa). The 6th ORF, when present, encodes a 14 kDa proteins with unknown function. GRSPaV seems to be more closely related to potexviruses than carlaviruses both of which have a similar genomic organization. These relationships have evolutionary implications and suggest that GRSPaV may have evolved from an ancient recombiantion event between a carlavirus and a potexvirus, in which ORF 4 and 5 but not the 3' non coding region of the carlavirus were replaced by those of the potexvirus. Grapevine virus A (GVA), the type species of the genus Vitivirus, is the putative agent of Grapevine kober stem grooving. Virus particles are flexuous filaments about 800 nm long. Viral RNA has a Mol. wt of about 2.6 x 106 Da and a size of 7349 nt. The viral genome consists of 5 ORFs encoding, in the order, the replication-associated proteins (195 kDa), a 20 kDa protein with unknown function, the movement protein (31 kDa), the coat protein (22 kDa) and a 10 kDa product which has nucleotide binding properties, is a pathogenicity factor and a gene silencing suppressor. Minor biological and serological variants of the virus are known. Grapevine virus B (GVB) is a vitivirus distantly related serologically to GVA and one of the etiological agents associated with Grapevine corky bark. GVB is also involved in young grapevine decline, a graft incompatibility condition recorded from California. Its totally sequenced RNA has a Mol. wt of about 2.7 x 106 Da, a size of 7599 nt and the same gene sequence and structural organization as GVA. This virus occurs in nature as a family of molecular variants, but biological variants are also known, two groups of which can be differentiated by the reaction of herbaceous hosts. Virus particles coated by both GVA and GVB coat protein occur in cells infected contemporarily by both viruses (phenotypic mixing). Grapevine virus C (GVC) is a little known and poorly characterized virus reported from Canada. Virus particles have a vitivirus morphology and an estimated length of about 725 nm. GVC is serologically distinct from GVA and GVB. Grapevine virus D (GVD), a vitivirus distantly related serologically to GVA and GVB is associated with with corky rugose wood, a field syndrome characterized by the presence of a striking corky condition of affected vines, just above the graft union. Virus particles are flexuous filaments about 825 nm long. The viral genome, which was sequenced only in part, has an estimated size of c. 7600 nt and a 3' terminus structurally comparable to that of GVA and GBV. Cytopathology: Whereas no information is available on the cytopathology of GRSPaV infections, vitivirus-induced cellular modifications have been extensively studied, primarily in herbaceous hosts. Cytopathological features common to all four vitiviruses (GVA, GVB, GVC, and GVD) consist of: (i) virus particle aggregates of various size, forming bundles, whorls, banded bodies, stacked layers that sometimes fill the entire cell lumen; (ii) variously extended wall thickenings originating from deposits of callose-like substances; (iii) proliferation and accumulation of cytoplasmic membranes; (iv) vesiscular evaginations of the tonoplast protruding into the vacuole and containing finely fibrillar material resembling dsRNA. GVA and GVB movement proteins were found to associate with cell walls and plasmodesmata, as detected by gold immunolabelling. Transmission: For many years after its discovery there were no records of natural spread of rugose wood in the field. GVA and GVB are now known to be transmitted from grapevine to grapevine by pseudococcid mealybugs and/or scale insects in a semipersistent manner. GVA vectors are the mealybugs Planococcus citri, Pl. ficus, Pseudococcus longispinus, Ps. affinis, Heliococcus bohemicus, and the scale insect Neopulvinaria innumerabilis, whereas GVB is transmitted by Ps. longispinus, Ps. affinis, and Pl. ficus. GRSPaV has no known vectors, but is suspected to be pollen-borne. There are, however, conflicting reports on its presence within seed and no evidence that it occurs in seedlings from infected vines. None of the putative agents of rugose wood has alternative hosts in nature and, because of the relatively limited range of vector movement, is not disseminated over long distances by natural means. Transport of infected propagative material represents the major means of dispersal. The presence of rugose wood and its causal agents in phylloxera-free countries with a millenial history of ownrooted grapevine cultivation, suggests that the disease originated in the Old World and was distributed worldwide by commercial trading and planting of infected grafted plants. Varietal susceptibility: Most if not all V.vinifera varieties and American rootstocks are susceptible. Although customarily grapevines are infected symptomlessly when ungrafted, rugose wood symptoms
: Graft transmission of rough bark to LN 33. Historical review Names like "legno riccio". 1965 1965 1967 81 . 1954 1961 1962 1963 1963 1964 1965 Hewitt: Rough bark. Martelli et al. with vein necrosis. or spot-RT-PCR using degenerate or virus-specific primers. Control of mealybugs is difficult for they overwinter under the bark of grapevines and possess an unwettable waxy covering. several resistant plant lines were obtained by transformation with the coat protein and the movement protein genes of GVA and GVB. GVA can be eliminated to a very hight rate (up to 97%) by the procedure used for cryopreservation of grapevine shoot tips. other assays include: single step or nested reverse trascription-polymerase chain reaction (RT-PCR). Graniti: Detailed description of rugose wood symptoms. Detection: Indexing on indicators (V. it is plausible to regard 110R as a specific indicator of rupestris stem pitting in addition to V. if not otherwise associated with a specific syndrome.were observed in self-rooted cultivars and ungrafted roostock stocks (V. Goidanich and Canova: First record of corky bark in Europe. Thus. since symptomless infections make sanitary selection not totally reliable. to a restricted range of herbaceous hosts (mostly Nicotiana species). Latent infection can occur also in grafted vines. Kober 5BB and LN 33) is the only reliable method for detecting and sorting out the diseases of the complex. Immuno-capture RT-PCR is 1000-fold more sensitive than ELISA for virus detection in grapevines. or a combination of the two.: First record of rugose wood outside of Italy. rugose wood agents can be eliminated with reasonable efficiency by heat therapy. all sources must be indexed and/or laboratory tested. the putative agent of rupestris stem pitting. Transgene expression was detected in these plants and in transformed grapevine explants. Vitiviruses. Faccioli: First histological study of corky bark-affected grapevines. a virus-like disease.: Corky bark is remarkably heat stable and difficult to eliminate by heat therapy. Beukman and Goheen: Brief account of the histological modifications of corky bark-affected LN 33. In addition. Suggestion that it may be caused by a virus. Suggestion that rugose wood may be a disease of combination requiring the contact of scion and rootstock for the development of symptoms. experimental evidence has been obtained of the very close association of GRSPaV. though with difficulty. 2. no strategy has yet been developed for the chemical control of vectors. described from California. rupestris. Thus. and that it may be a composite disease resulting from the interaction of different viruses among which GFLV. rupestris and Kober 5BB). The best antigen sources for serological diagnosis are cortical shavings from mature dormant canes. Graniti and Ciccarone: First record of rugose wood from southern Italy. as shown by 110R. Goheen et al. The intensity of wood abnormalities (pitting and grooving) vary. rupestris. 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. Individual viruses can be identified by ELISA or dot immunobinding on nylon membranes using polyclonal antisera and/or monoclonal antibodies when available. meristem tip culture. However. In general. Name of the disease changed into corky bark. Recently. are mechanically transmissible. Using a Nicotiana benthamiana model system. but not foveaviruses. Hewitt et al. possibly in relation with the scion/stock combination and climatic conditions. Graniti and Martelli: Demonstration of the infectious nature of rugose wood. Histological study of diseased vines. "stem pitting" and "stem grooving". immunocapture RT-PCR. are synonymized with "rugose wood". 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.
Hewitt and Neja: First record of rugose wood in USA (California). 1968 1969 1970 1970 1971 1971 1972 1973 1973 1975 1975 1977 1978 1979 1979 1980 1980 1981 1981 1982 1983 82 . Boccardo and D'Aquilio: Physicochemical characterization of GSP-AV Abracheva: Survey of over 650 grapevine cultivars and hybrids for rugose wood reaction in Bulgaria. Bovey and Brugger: Further evidence that GFLV may not be implicated in the etiology of rugose wood in Switzerland. Beukman and Goheen: Up-to-date review of corky bark.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.1968 1968 Lehoczky et al. Hewitt: Successful graft transmission of Californian rugose wood.: First experimental evidence that a filamentous virus (GVA).: Green grafting useful for corky bark indexing. Goheen and Luhn: Heat treatment of dormant buds grafted onto LN 33 is effective against corky bark. Goheen and Luhn: Suggestion that corky bark and rugose wood are the same disease.Virus provisionally called grapevine stem pitting-associated virus (GSP-AV). No nepoviruses implicated in their etiology. Rugose wood may not require a grafted plant for full symptom expression. Sarooshi et al. At 38 °C the minimum inactivation period for leafroll is 56 days and for corky bark 98 days. Graniti and Martelli: Up-to-date review of rugose wood. based on graft transmission tests. Rosciglione et al. Lehoczky: Destructive effects of rugose wood registered in Hungary in both self-rooted and grafted European grape varieties. is transmitted by the pseudoccid mealybug Pseudococcus longispinus.b. Hewitt: Up-to-date review on grapevine virus and virus-like disease worldwide. Engelbrecht and Nel: Rugose wood and fanleaf are not related. Mink and Parsons: Use of growth chambers for rapid symptom expression of corky bark in Vitis indicators. Teliz et al. from a rugose wood-infected vine. Goheen: Evidence that corky bark and leafroll. Anonymous: A review of rugose wood in Italy. despite similarities in the s y m p t o m s o n t h e foliage are different diseases.: Rugose wood recorded from Australia. First record of rugose wood symptoms outside of Europe (Israel). Conti et al. a.: Heat therapy effective against rugose wood. Castillo et al. Beukman and Gifford: Detailed account of adverse effects of corky bark on the anatomy of Vitis. Legin et al.: Observation of rugose wood symptoms in self-rooted vines.: Recovery by mechanical inoculation of a closterovirus with particles 800 nm long.
: First indication of the possible existence of LN 33 stem grooving. Gallitelli et al. wt of 5. Similar dsRNA species were detected.Sc.3 and 4.: Evaluation of the effect of rugose wood on cv. The first two disorders appear to be spreading in the vineyards. Savino et al.: Application of spot hybridization for the detection of GVA in grapevine sap. Kuniyuki and Costa: Rugose wood recorded from Brasil Goheen: First published description of rupestris stem pitting. Martelli: Rugose wood recorded in southern Mediterranean and Arab countries.: Experimental confirmation that rugose wood may not express symptoms in grafted indicators.: A low molecular weight dsRNA associated with rupestris stem pitting. Monette et al. denoted Grapevine virus B (GVB). Prudencio: M. thesis describing rupestris stem pitting disease in comparison with corky bark. No closterovirus-like particles in samples with rupestris stem pitting. an additional disease of the rugose wood complex.: Evidence that GSP-AV can occur in grapevines together with another similar but serologically unrelated virus with short closterovirus-like particles. Corky bark and Rupestris stem pitting. Castrovilli and Gallitelli: Physicochemical comparison of two Italian isolates of GVA. GSP-AV re-named Grapevine virus A (GVA). Engelbrecht and Kasdorf: Natural field spread of corky bark in South Africa associated with the presence of P. Monette and James: Detection of two biologically distinct but serologically indistinguishable isolates of GVA. Tanne et al. ficus. Savino et al.1984 Milne et al. Azzam et al.: First record of rugose wood from China. Suggestion that the disease is not related to closteroviruses associated with GLRaV and corky bark. Murant et al. Garau et al. similar to Kober stem grooving.ficus.: Transmission of corky bark by the mealybug P.: Experimental confirmation of the complex nature of rugose wood based on the differential reaction of woody indicators. Rosciglione and Castellano: Demonstration that GVA is transmitted also by Planococcus citri and P. Italia propagated on six different rootstocks.: Two distinct dsRNAs with a mol. but not consistently in grapevines from New York. Rugose wood and corky bark are not the same disease.: Three types of wood disorders of the stem-grooving type observed in South African grapevines. Li et al. Engelbrecht et al.4 x 106 associated with rupestris stem pitting in grapevines from California and Canada. Savino et al.: Heracleum latent virus and GVA are distantly serologically related. Garau et al. Corbett and Wiid: Closterovirus-like particles found in extracts from vines affected by corky bark and rugose wood in South Africa. First report of Kober stem grooving. 1985 1985 1985 1985 1985a 1985b 1985 1985 1985 1987 1988 1989 1989 1989 1989 1989 1989 1990 1990 1991 1991 83 . ficus.: Assessment of crop losses induced by rugose wood to two different European grape varieties.
Martelli et al.: Development and diagnostic use of a cloned probe to GVB.: Sequence of the 3' end of GVA and GVB genome. Namba et al. both associated with a stem pitting condition of grapevines rather than with leafroll. 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.: Rugose wood recorded from Yemen. Monette and James: A closterovirus with short particles (725 nm) isolated from a corky barkaffected vine induces necrotic local lesions and systemic symptoms in Nicotiana benthamiana. Both viruses qualify for the inclusion in the genus Trichovirus. Thorough comparative study of nine GVB isolates from different countries.: Production of monoclonal antibodies to GVA and their use for ELISA detection of the virus in infected vines.: Purification and properties of GVB. Saldarelli et al.: Contemporary occurrence of Rupestris stem pitting and Kober stem grooving in symptomless scions of cv.1991 1991 Gugerli et al. Minafra and Hadidi: Detection of GVA and GVB in viruliferous mealybugs by PCR. Minafra et al.: Development of digoxigenin-labelled riboprobes for the detection of GVA and GVB in infected tissue extracts. Torbato in Italy.: Rugose wood recorded from Ukraine Boscia et al. Martelli et al. Saric and Korosec-Koruza: Rugose wood recorded from Croatia and Slovenia Ioannou: Rugose wood recorded from Cyprus. Virus named Grapevine virus C (GVC). Suggestion that GVA is implicated in the aetiology of the disease.: Rugose wood recorded from Malta Monette and Godkin: Recovery of a closterovirus-like virus by mechanical inoculation from a corky bark-affected vine.: A closterovirus with particles 1440-2000 nm long serologically unrelated to all other known grapevine closteroviruses found in corky bark-affected vines.: Clear-cut connection of GVA and rugose wood.: Synthesis of a cloned probe for GVA. ficus induced corky bark symptoms in LN 33 Saldarelli et al. Suggestion that GVA may be the causal agent of the disease. Padilla: Rugose wood recorded from Spain Boscia et al.: Presence of two distinct serotypes of GVA. Virus transmission by the mealybug Ps.: GVA and Kober stem grooving are closely associated. Merkuri et al. Minafra et al. Chavez and Varon de Agudelo: Rugose wood recorded from Colombia. Garau et al. Garau et al.: Rugose wood recorded from Albania. Boscia et al. Digiaro et al.:Rugose wood recorded from Tunisia Milkus et al. 1991 1991 1991 1991 1991 1991 1991 1991 1992 1992 1993 1993 1993 1993 1994 1994 1994 1994 1994 1994 1994 1994 1995 84 . Boulila et al.
: Nucleotide sequence of GVB genome. Since particle size (600-700 nm in length) is compatible with that of Grapevine rupestris stem pitting-associated virus (GRSPaV) particles identified in 2002.: GVA and GVB are serologically related. GVC. GRSPaV is assigned to this genus. The virus is not seed-borne. Suggestion that spreading is by a vector that transmits in a semipersistent manner.: GVB is consistently associated with corky bark and is present. Martelli et al. Guidoni et al. Boscia et al. Saldarelli et al.: Rugose wood recorded from Lebanon. Goszczynski et al. Efficient detection method based on TAS-ELISA developed. Minafra et al. La Notte et al. Bonavia et al. Haidar et al.: Sequencing of a Californian isolate of GRSPaV. Boscia et al.: A study of the spatial distribution pattern of corky bark in aThompson seedless vineyard in Israel.: Description of Grapevine virus D (GVD).: Development of a PCR technique for the detection of GVA and GVB in nylon membrane-spotted sap. 1995 1995 1995 1996 1996 1996 1996 1996 1997 1997 1997 1997 1997a 1997b 1997 1997 1997 1997 1998 1998 1998 1998 85 . La Notte et al.: GVA and GVB are transmitted by Pseudococcus affinis. Alkowni et al: Rugose wood in Palestine.: GVA and GVD are serologically distantly related. Martelli and Jelkmann: Establishment of the genus Foveavirus. GVB.: Sequence and structrural organization of Grapevine rupestris stem pittingassociated virus genome (GRSPaV). GVC. Rubinson et al.: Antiserum to the movement protein of GVA is useful for virus detection in ELISA. Abou Ghanem et al. Faoro: Review of the cytopathology of grapevine trichovirus infections. Zhang et al. this may be the first visualization of GRSPaV. Further support of the cause-effect relationship between GVA and this disease.: Review of the properties of putative grapevine-infecting trichoviruses (GVA.: Rugose wood recorded from Jordan. though not consistently in vines showing a syndrome denoted "corky rugose wood". and GVD) later assigned to the genus Vitivirus.: Consistent detection of GVA in Kober stem grooving-infected grapevines by immunocapture-polymerase chain reaction.: Nucleotide sequence of GVA genome and taxonomic position of the virus. and GVD removed from the genus Trichovirus and assigned to the new genus. longispinus in a semi-persistent manner.: Estasblishment of the genus Vitivirus with GVA as type species. GVB.: GVA is transmitted by by Ps. Tanne et al.: Elimination of GVA from cv Nebbiolo clones by heat therapy improves agronomic performance of the vines and quality of the must. Choueiri et al. Meng et al. GVA. Chevalier et al. Garau et al.1995 Monette and Godkin: Detection of non mechanically transmissible capillovirus-like particles in a grapevine affected by rugose wood.
Samples made up of three buds from dormant canes are less laboriour to prepare than cane shavings and yield comparable results. The virus was not detected in 245 seedlings from infected cv.: 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. Buzkan et al.: Production of an infectious RNA transcript from a full-length cDNA clone of GVA.8% and 78-89. Virus detected in bleached seeds suggesting that it is present inside the seeds. Seyval seedlings. Goszczynski and Jooste: Use of single-strand conformation polymorphism reveals molecular heterogenity in GVA populations.: The function of GVA genes identified by mutation analysis of individual ORFs of a full-length infectious viral clone. Boscia et al. Ahmed et al. intermingled with virus particle aggregates.: GRSPaV particles. GCD) and foveavirus (GRSPaV) sequences in two steps. Habili et al.: GVA particles carry a highly structured epitope centered on a common peptide region of the coat protein sequence.e. Nucleotide sequence identity within groups is 91-99.: Synthesis of full-length cDNA copies of GVA and GVB genomes. Saldarelli et al. Goszczynski and Jooste: GVA and GLRaV-3 are both consistently associated with Shiraz disease in South Africa but only GVA seems to be required for disease induction. II.: Rugose wood viruses in the Czeck Republic.: One-sided phenotyping mixing.: Rugose wood viruses in Iran. Petrovic et al. are filamentous and measure 723 nm in length. Kominek et al.: Production of monoclonal antibodies to GVB.: GVA transmission by Pseudococcus comstocki. GVB. Nakano et al. Further support to the cause-effect relationship of GRSPaV with this disease Galikparov 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. Stewart and Nassuth: An improved extraction method allows RT-PCR detection of GRSPaV virtually thoughout the year in all grapevine tissues. observed for the first time.: Elimination of GVA by cryopreservation. GVA coat protein encapsidating GVB RNA.: Consistent association of GRSPaV with vines indexing positive for Rupestris stem pitting.3% among groups. occurs in Nicotiana plants doubly infected with GVA and GVB. Goszczynski and Jooste: Thre groups of GVA strains (I.: Rugose wood viruses in Egypt. Dell'Orco et al. Dovas and Katis: Improved RT-PCR method for the simultanous detection in grapevine extracts of vitivirus (GVA. Wang et al. GVB and Corky bark and GRSPaV and Rupestris stem pitting. Saldarelli et al.1999 1999 2000a Meng et al. GVA movement protein is also present in great quantity in the cytoplasm. 2000b 2000 2001 2001 2001 2002 2002 2003 2003 2003 2003 2003 2003 2003 2003 2003 2003 2003 86 . Minafra et al. Galiakparov et al. Confirmation of the cause-effect relationship between GVA and Kober stem grooving. and III) identified in South Africa based primarily on sequence homology of the 3' end of the viral genome.: Movement proteins of GVA and GVB detected by gold immunolabelling in association with cell walls and plasmodemata of infected cells. i. Meng et al.
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GRAFT INCOMPATIBILITY .
However. Freedom. consistently. 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. up to five different grafttransmissible agents inducing incompatibility could be differentiated in California. 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. 1103P. 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. The nature of this disease has not been ascertained but one or more graft-transmissibile agents may be involved in its aetiology. only GLRaV-2 RG was identified. New Zealand. shoots are short. in Argentinian grapes. with margins more or less extensively rolled downwards. a member of the genus Closterovirus. Description Main synonyms: Incompatibilité au greffage (Fr. the increased use of clonal material is disclosing unprecedented conditions of generalized decline that develop dramatically in certain scionrootstock combinations. Other assays include nucleic acid-based tecniques such as single step or nested reverse transcription-polymerase chain reaction (RT-PCR) and 95 . A transitory form of incompatibility was reported from Italy under the name of bushy stunt. Detection: Indexing on Caberent sauvignon is a reliable method for detecting incompatibility conditions.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). This latter condition has been known for a long time and occurs also in rugose wood-affected vines. Based on the differential responses of a panel of 18 rootstocks. In this case. though not consistently and. is not transmitted by mealybugs and does not have a known vector. scions show a stunted and bushy vegetation due to the contemporary proliferation of apical and axillary buds. Syrah decline is a severe disease occurring in France. The same virus was detected in diseased Chilean grapes. Normal growth resumes with the second or third leaf. Other molecular variants of GLRaV-2 were reported from New Zealand (Alphie virus). Cases of graft union disorders have been observed in Europe (Kober 5BB incompatibility). Of these. except for GRSPaV. The putative agent of bushy stunt was consistenly found in clones of the rootstock 140R in which it is latent. Chile. Agents: An ordinary strain of Grapevine leafroll-associated virus 2 (GLRaV-2) is consistently associated with Kober 5BB incompatibility (Europe).) Symptoms: Newly planted vines grow weakly. Australia and Chile (young vine decline). A virus originally detected in cv. 1. 3309) develop a discolored canopy . Transmission: GLRaV-2. Severely affected vines decline and may die within one or two years.g. appears to be involved in California's young vine decline. European grape varieties grafted on tolerant rootstocks (e. and together with Grapevine virus B (GVB). The heat-labile graft-transmissible agent present in the hybrid 140R. Harmony. California. leaves are small-sized.g. 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. so as to represent veritable emerging diseases. 5C.). 03916. decline and may die. Infected propagative material is to be blamed for its dissemination. associated with grapevine bushy stunt is still unidentified. Salt creek . GVB is mealybug-borne and can be spread at a site by these insects. which are usually not accompanied by wood abnormalities (pitting or grooving). A prominent swelling forms at the scion/rooststock junction and variously extended necrotic lesions may develop on the rootstock stem. Kober 5BB. incompatibilità d'innesto (Ital. but the yield is reduced. and Australia in association with young vine decline conditions. Argentina and probably elsewhere. Varietal susceptibility: Appearance of graft union disorders depends more on the rootstock rather than the scion. but the colour of the canopy remains green. although none of a number of known grapevine-infecting viruses has been found in affected vines. 101-14) exhibit a green canopy and perform rather well. 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). whereas varieties grafted on susceptible roostocks (e. and again California (rootstock stem lesions).
: Updated report on the state of the art of investigations carried out in France on Syrah decline. with a decline condition of young Thomposn seedless vines in Chile. meristem tip culture. Martelli: GRSLV and GLRaV-2 are serologically related and are both recognized by a panel of 18 monoclonal antibodies. Boubals: Report of a national French study group investigating the aetiology of Syrah decline.: Description of bushy stunt and evidence that it is caused by a graft-transmissibile heat-sensitive agent carried by some clonal rootstocks. Bonfiglioli et al. Greif et al. whenever feasible.: Report of a new molecular variant of GLRaV-2 from New Zealand.: GLRaV-2 is the cause of a graft incompatibility revealed by Kober 5BB. 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.: Identification of an apparently new closterovirus denoted Grapevine rootstock stem lesion virus (GRSLV) causing stem necrosis of rootstocks. Graft-transmission of the incompatibility factor. Bertazzon and Angelini: Comparison of several detection methods for the broad or specific identification of Grapevine leafroll-associated virus 2 variants. Currently known graft incompatibility agents can be eliminated with reasonable efficiency by heat therapy. Boubals: Syrah decline occurs in Argentina Uyemoto et al. Uyemoto and Rowhani: Indexing on 18 different grape rootstocks reveals the existence of at least five different agents causing graft incompatibility. 1977 1979 1986 1991 1995 2000 2000 2000 2001 Jacob: Description of graft incompatibility in different scion/stock combinations. though not consistently. Prodan et al. and death of the vines. 2. Boubals and Huglin: Report on graft incompatibility of certain varieties grafted on 57R. If scionwood is infected. The problem is very complex and may involve several still unidentified factors.: Third paper of a series on incompatibility on Kober 5BB. the use of sensitive rootstocks is to be avoided and. using degenerate or virus-specific primers. Renault Spilmont et al.: GLRaV-2 is associated. or a combination of the two. GRSLV re-named Redglobe strain of GLRaV-2. No conclusion are drawn. decline. GRSLV has about 75% nucleotide homology with GLRaV-2. Historical review 1942 1950 1973.: GLRaV-2 and GVB are consistently associated with young vine decline in California. 2003 2003 2003 2003 2003 2003 2004 96 . Gomez Talquenca et al: GLRaV-2 is consistently associated with declining Cabernet sauvingon vines grafted on different roostocks in Argentina. Savino et al. Control: Prevent introduction of infected vines in the vineyard by using certified grafted plants or virusfree scionwood and rootstocks.immunocapture RT-PCR. utilization of tolerant roostocks is advisable. Golino et al.: Two papers describing incompatibility phenomena between clonal selections of different cultivars grafted on Kober 5BB Fallot et al. Strategies on how to protect healthy stocks from vector-mediated GVB reinfection in the field are yet to be developed. Suggestion that they are molecular variants of the same virus species. Durquety et al.
144. S. Progrés Agricole et Viticole 96.. Autres cas chez la vinge. 167-173.. 183-189. Boubals D. and E. Etude de phénomènes d'incompatibilité au greffage chez la vigne. 1977. 420-427 Fallot J. Locorotondo 2003.A.S. Ruchaud.II. Renault Spilmont A. Journal of Plant Pathology 86. Dauty. 2001.. Fiori. Grau. Durquety P. Phytopathologia Mediterranea 34. Adelaide 2000.III. Greif C. and B. 139-140.M. V. 2000.. S. Gracia. Rowhani. and A. Locorotondo 2003. Luvisi and R. 2003. Fallot. Walter and U. References Bertazzon N.. 1986. Legin R. A. Le clone et ses reactions au greffage..M. Le dépérissement de la Syrah existe aussi en Argentine. B. Di Terlizzi. Bonfiglioli R. 2003. Extended Abstracts 14th Meeting of ICVG. 85-86. Golino D. 97 . S. Extended Abstracts 14th Meeting of ICVG. La trasmission de l'incompatibilité au greffage entre 5BB et Vitis vinifera.. Prota. Bass. 1979.3. Extended Abstracts 14th Meeting of ICVG. I. 2000. Savino V.P. Progrés Agricole et Viticole 103.. 141. Garcia Lampasona and O.. Extended Abstracts 14th Meeting of ICVG.M. 2003.S. Fiore.E. Etude de l'incompatibilité au graffege de certain cépages et du 57R. Boscia. Gomez Talquenca G. J. Fallot. 28-31. Gazeau and J. D. Proceeding of the American Society of Horticultural Science 41.. M. Examples of incompatibility between grape varieties and roostocks. Grapevine virology highlights 2000-2003. Extended Abstracts 14th Meeting of ICVG. Edwards and A. 1995. 2000. Locorotondo 2003. Progrés Agricole et Viticole 117. 201-203.M.K. Benassac and R. Discovery of different grapevine sources with graft-transmissible agents causing union-incompatibility on sensitive rootstocks. Aetiology of decline in Thompson seedless grafted table grape plants. O. Compte-rendu de la réunion du Groupe de Travail National. Martelli G. J. Ruchaud. 211-216. Walter. P. Huglin. Rivieccio and F. California Agriculture 55 (4). 122-129 and 171-178. Le clone et ses reactions au greffage. Rowhani. Jacob H. Locorotondo 2003. A young grafted vine decline syndrome in Argentina vineyards. Ruchaud.K.P. 142-143. B. Durquety and J. Molecular studies on a graft incompatibility syndrome in New Zealand vineyards yields another probable variant of Grapevine leafroll-associated virus 2.. Presence in clonal rootstocks of a grafttransmissible factor that induces stunting and bushy growth in European grapevines.. F.. Volos 1990. Identification of the latent viruses associated with young vine decline in California. 3-10. Boursiquot. Proceedings 10th Meeting of ICVG. R. Existence dans un cépage population de clones présentant divers degrés de compatibilité avec certains porte-greffes. Angelini. Uyemoto J. 2003. Advances in the detection of Grapevine leafroll-associated virus 2 variants. 1950. 137-141 Boubals D. Progrés Agricole et Viticole 117. J. Locorotondo 2003. The relationship of grapevine leafroll-associated virus 2 with a graft incompatibility condition of grapevines. Pantaleo. Locorotondo 2003. Montalegre and N..P. Le clone et ses reactions au greffage. Extended Abstracts 14th Meeting of ICVG. 85-86. Le dépérissement de la Syrah. New closterovirs in 'Redglobe' grape causes decline of grafted plants. C.. Progrés Agricole et Viticole 94. Syrah decline in French vineyards. 2003. C. Krag. 277. Prota. Garau. Prodan S. 1973. Gazeau. 1991. Rowhani. D. 2004. Grenan and J. Progrés Agricole et Viticole 67. P.P. Sim and A. Durquety P. 1942. 202-210 Uyemoto J. and P. 283-290.A. Boubals D. 2003. Progrés Agricole et Viticole 90. Di Silvio. Extended Abstracts 13th Meeting of ICVG. S. C. J. Nouvelles recherches sur l'incompatibilité clonale d'Abouriou greffé sur 5BB. 279-283.
FLECK COMPLEX .
twisted and may curl upward. which are twisted and asymmetric. rooting ability of rootstocks and on graft take has been reported. This disease. wt of 2. GAMaV. positive sense RNA with high cytosine content (c. Asteroid mosaic. Rupestris vein feathering. Mild asteroid mosaic-like symptoms are shown by some European grapevine varieties (e. irregularly distributed over the leaf blade. GFkV genomic RNA constitutes about 35% of the particle weight. containing the genome. adverse influence on vigour. Fleck is an ubiquitous disease reported from most viticultural countries in the world. GRGV. b. Symptoms are expressed in Vitis rupestris and consist of clearing of the veins of third and fourth order. 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. The complete sequence of GFkV and partial sequences of GRGV. Leaf symptoms usually become less severe in summer. 25 kDa.). twisted and puckered along the veins. GFkV. Sultanina). capped. and produce little or no fruit. The coat protein (CP) of GFkV and GRGV particles is made up of a single protein species with Mr of c. Symptoms: a. Agents: All viruses of the complex. Marmorierung der Rebe (Germ. In V. Severe strains induce also varying degrees of stunting. Sternmosaik der Rebe (Germ. T made up of empty protein shells and B. and Grapevine rupestris vein feathering virus (GRVFV) are all phloem-limited and non mechanically transmissible. leaf symptoms are characterized by star-shaped chlorotic spots. screziatura (Ital. GFkV genomic RNA (Mol. maculatura infettiva. mosaico stellare (Ital. B.g. Grapevine fleck: Marbrure (Fr. In V. Italy and California. Affected vines are often stunted. 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. GFkV particles sediment as two centrifugal components.). The putative causal agent of the disease has only been found in California. Recorded from California and Italy . 50%). V. Grapevine asteroid mosaic: Mosaïque étoilée (Fr. grapevine asteroid mosaic. d. Description Main synonyms: A. the disease elicits creamy-yellow bands developing along the major veins of the leaves. Grapevine red globe virus (GRGV) is a Grapevine fleck virus (GFkV)-like virus which apparently does not induce symptoms in European grapevine varieties (e. rupestris following graft inoculation. sometimes with necrotic center. 1. Transient mild chlorotic discolourations of the primary and secondary leaf veins develop in V. Asteroid mosaic symptoms have been observed in several varieties of V. rupestris. 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 in California. The disease is latent in European grapevine varieties and in most American rootstocks. leaf petioles and veinlets. but likely to occur elsewhere.).). Red globe) nor in V. Rupestris necrosis.). producing localized translucent spots.). whereas the CP of GAMaV and GRVFV consists of a major protein of 21 kDa and a minor prtotein of 25 kDa. c.g. which is a monopartite single-stranded. is latent in European grapevine varieties. rupestris reacts with localized necrosis of the shoots. which is used as indicator. Grapevine asteroid mosaic-associated virus (GAMaV). and GRVFV genomes are available. Although the elusive nature of the complex hinders the assessment of its economic impact. grapevine rupestris necrosis. e.FLECK COMPLEX The fleck complex consists of several diseases (grapevine fleck. Leaves are asymmetric. Fleck. vinifera. The putative causal agent of the disease so far has been found in Greece. rupestris. reported only from Japan.6 x 106 ) is 7564 nt in size and contains four open reading frames (ORF) that 101 . Leaves with intense flecking are wrinkled.
encode a 215. Comparison of symptoms with those of other mosaic diseases of grape. whilst GAMaV and GRVFV were assigned to the genus Marafivirus. Although observations from Italy. 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. GRGV. The 3' end of GRGV genome is structurally similar to that of GFkV except for the lack of ORF 4. Transmission through dodder of GFkV has been reported but it is of no epidemiological importance.rupestris described in France. and two proline rich polyproteins of 31. As the disease is rare and does not appear to be spreading. Therefore. Attempts to transmit the disease by mechanical inoculation to herbaceous test plants or by Xiphinema index were unsuccessful. Varietal susceptibility: GFkV and possibly all the other viruses of the complex infect naturally a large number of varieties and Vitis species. The same sanitation procedures are likely to operate successfully with the other viruses of the complex. ELISA is currently employed for routine detection of GFkV. meristem tip or fragmented shoot apex culture. of which it represents the type species.: First record of fleck as an unidentified symptom different from fanleaf and transmissible from symptomless varieties to V.4 kDa (ORF 3) and 15.9 kDa (ORF 4) with unknown function. 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". 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. molecular and ultrastuctural studies disclosed sufficient similarities between maculaviruses. primary dissemination of these and the other viruses of the complex is through infected propagative material. Polyclonal antisera and monoclonal antibodies to GFkV heve been raised. Detection: Indexing on V.George. Historical review 1954 1962 1966 1966 Hewitt: First description of asteroid mosaic in California. Refatti: Review paper on asteroid mosaic. but cannot be used for any of the other members of the complex due to the unvailability of antisera. GAMaV. Further physico-chemical.4 polypeptide with the conserved motif of replication associated proteins (ORF 1). GFkV was identified as the representative of a new genus denoted Maculavirus. and GRVFV. GFkV is not seed transmitted. 102 . marafiviruses and members of the genus Tymovirus to warrant the establisment of the a new family denoted Tymoviridae. its economic importance is low. No information is available on individual susceptibility. Vuittenez et al. These deranged organelles are thought to be sites of virus replication. 2. Because of its molecular characteristics. the CP (ORF 2). a disease inducing symptoms similar to those of fleck in V. Transmission: No vector is known for any of the viruses of the fleck complex. rupestris allows with a reasonable level of confidence the discrimination of the different viruses of the complex based on the differential reaction of the indicator. but no experimental data are available. whereas GAMaV induces peripheral vesiculation of chloroplasts. GFkV can be eliminated by heat therapy.: "Marbrure". Virus specific and degenerate primers have been designed for single or multiplex RT-PCR detection of GFkV. Control: Because of the latency of symptoms sanitary selection of European grapevine cultivars and most American rootstock hybrids is unrealiable. rupestris St. Hewitt et al.
Triolo and Materazzi: Fleck has a detrimental effect on the quality V. Milkus: Suggestion of a prokaryotic etiology for fleck. Dolja et al.: Fleck is not seed transmissible. Savino et al. in sieve tubes of field-grown vines with leafroll symptoms but likely to be affected by other diseases. The efficiency of heat treatment for disease elimination is unsatisfactory. Bovey: Identification of fleck in Switzerland as a latent disease of Chasselas transmissible to V.: Observation of an isometric non mechanically transmissible virus in the phloem of diseased vines. Verderevskaya et al. rupestris propagating wood.: Successful elimination of fleck through heat therapy.: Identification of a dsRNA of about 7 Kb pairs in diseased vines. Woodham and Krake: Dodder transmission of fleck from vine to vine.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.: Observation of a non mechanically transmissible virus. Goheen and Luhn: A novel heat therapy system based on virus inactivation in buds grafted onto healthy LN 33 rootstocks is effective against fleck. Rooting ability and graft take are adversely affected Yamakawa: Field spread of fleck in Japan Boulila et al. Triolo and Resta: Tetracycline treatments are ineffective against fleck. later called grapevine phloem-limited isometric virus (GPLIV). Ottenwaelter et al. Hévin et al. Report that a virus serologically similar to GPLIV is associated with the disease. Report of multivesiculate inclusion bodies probably connected with GPLIV infection. 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.: Successful elimination of fleck through fragmented shoot apex culture in vitro. leafroll and corky bark).: Purification of GPLIV from naturally diseased vines and production of a specific antiserum. Confirmation that the virus can be eliminated by heat therapy and is not related to leafroll. Castellano et al. Castellano and Martelli: Confirmation that GPLIV is associated with multivesiculate bodies and demonstration that these derive from deranged mitochondria. rupestris. Hewitt et al.: Report of widespread occurrence of fleck in visually selected grapevine clones in southern Italy. Barlass et al. Engelbrecht and Kasdorf: Observation of natural field spread of fleck in South Africa.: Description of fleck as an independent graft-transmissible disease present in many European varieties and American rootstocks.: Physicochemical characterization of GPLIV. Rives: Further demonstration that fleck is distinct from fanleaf based on differential responses to heat treatment. Dismissal of the prokaryote etiology hypothesis. Castellano et al. 1982 1983 1983 1983 1984 1985 1985 1987 1989 1990 1990 1990 1991 103 .
Sabanadzovic et al.rupestris of an isometric virus latent in V. vinifera cv. ELISA used successfully for virus detection in large scale survey.: GPLIV shown to be the agent of fleck. Adverse effect on Teleki 5A is negligible. GAMaV. Virus named Grapevine asteroid mosaicassociated virus (GAMaV) Boscia et al. Detection of sequences of an undentified virus from a Greek grapevine. Walter and Cornuet: Confirmation by ELISA of the consistent association of GFkV with fleck disease.: Additional monoclonal antibodies raised in France. Sabanadzovic et al. vinifera. based on the differential reactions of indicators Credi and Babini: Infection by fleck. 1991 1993 1994 1995 1995 1996 1996 1997 1997 1998 2000 2001 2001 104 .: Molecular reagents (degenerate primers) developed for the specific identification of viruses of the fleck complex (GFkV. Meristem tip culture effectively eliminates the virus. Fortusini et al. Virus renamed Grapevine fleck virus (GFkV).: Report of the close association with fleck symptoms in V. Elbeaino et al. GFkV-like multivesiculate bodies derived from deranged mitochondria are present in infected cells. This cytological feature recalls that later found in vines infected by Grapevine rupestris vein feathering virus. Kyriakopoulou: Description of a disease similar to asteroid mosaic observed in V. rupestris. Symptoms are severe and affected vines are almost fruitless.: Report of a highly consistent association of GPLIV with fleck in naturally infected and graft-inoculated V.1991 1991a 1991b 1991 Gugerli et al. Pruning wood is reduced by 51% in 420A and by 37% in Kober 5BB. Boscia et al. GAMaV and of another virus with GFkV-like particles phylogenetically but not serologically related to GFkV present in a cv Red globe vine. GRGV). Molecular properties of this virus further support the notion that it warrants classification in a genus of its own. Boscia et al. Virus named Grapevine redglobe virus.: 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. 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.: 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.: Two GFkV-specific monoclonal antibodies raised in Italy can successfully be used in ELISA Kuniyuki and Costa: Three strains of GFkV reported from Brazil. Boscia et al. June-July are the best months for ELISA detection of the virus in Alsace. Namba et al. Marsumoto and Ohki: A spherical virus resembling GFkV identified in thin sectioned cells of V. Sultanina in Greece. vein necrosis and vein mosaic has a detrimental effect on rootstock growth.: A non mechanically transmissible isometric virus similar but unrelated to GFkV identified in asteroid mosaic-infected grapevines. rupestris with a necrotic disease. later named Grapevine rupestris vein feathering virus (GRVFV).: Natural field spread of GFkV observed in Northern Italy Schieber et al.: Complete nucleotide sequence of GFkV genome. The disease seems to be spreading naturally.
and Japan) only the variant without insertion (GFkV353) was detected. Journal of Ultrastructure Research 89.P. Extended Abstracts 14th Meeting of ICVG. 1984.: Description of Maculavirus. S.. Advances in Horticultural Science 10. Digiaro. Molecular detection of Grapevine fleck virus-like viruses. Martelli. 1990.P. fleck. S.: A sequence variant of GFkV (GFkV416) with a 63 nucleotide insertion in the replicase gene identified in Australia and New Zealand. 11-16 Abou Ghanem-Sabanadzovic. Kyriakopoulou and G. Savino. Savino and M. Di Terlizzi.P. A. Boulila M. Verderevskaya and J. Identification of the agent of grapevine fleck disease. comprising the genera Maculavirus and Marafivirus that include GFkV/GRGV and GAMaV/GRVFV. and A. G.P.: Sequencing of the 3' end of the genome of GRGV. 97-105. GAMaV and of a virus of Greek origin which induces vein feathering in V. Martelli and M...F. Annals of Applied Biology 101.P. Castellano. Sabanadzovic. Woodham and L.: Development of a multiplex RT-PCR protocol for the simultaneous detection of GFkV-like viruses using plant mRNA as an internal control. a new genus of plant viruses having GFkV as type species and GRGV as tentative species. ElBeaino T. K. Boscia D. 1983. N. Field spread of corky bark.A.P. Agadir 1990. Sabanadzovic and G. Castellano. Boscia.D. Tomashevskaya. and G.E.A.G.C. Rowhani P. Phytopathologia Mediterranea 24. Argentina. V.R.P. Plant Pathology 44. Vitis 40: 65-68.V. B. Martelli and V. Karsev. Rowhani and G. V. A. Atabekov. Greek virus recognized as a species in its own right denoted Grapevine rupestris vein feathering virus (GRVFV). Savino. U. 173-174. Martelli. Boscia D. Vitis 30. Numéro hors série. Production of monoclonal antibodies to grapevine fleck virus. 191.A. and G. Journal of Phytopathology 129. Martelli. O. Kyriakopoulou and G. 1985. 1991a. Martelli.V. N. Volos 1990. Martelli 2001. D. Annales de Phytopathologie. 2003b.. M. Effect of virus and virus-like infections on the growth of grapevine rootstocks. A non mechanically transmissible virus associated with asteroid mosaic of the grapevine. Ultrastructure and nature of vesiculated bodies associated with isometric virus-like particles in diseased grapevines. V.. Savino.R. Bovey R. Abou Ghanem-Sabanadzovic. P. A.P. Elicio. 2003b 2003 3. 1982.P. respectively. Virus Genes 27. 1990. Castellano M.. Savino and D.P. Barlass M. 95-98. S. leafroll and Shiraz decline diseases and associated viruses in South African grapevines.J. Un virus latent dans le Chasselas. Some properties of a phloem-limited non mechanically-transmissible grapevine virus.. V. Martelli. Savino. 2003a. 1990. Boyko. V. Martelli et al. M. Credi R. Krake.. In other countries (USA. Phytophylactica 22. Iran. Babini. Proceedings 10th Meeting of ICVG. Boscia. Sabanadzovic. 151-158. R. Castellano M. Virus-like particles and ultrastructural modifications in the phloem of leafroll-affected grapevines.A. 3134. T. 291-295.G. 1972.A.. Savino and G. 160-163. Proceedings 8th Congress of the Mediterranean Phytopatological Union. 23-39. rupestris confirms the assignment of GRGV to the genus Maculavirus and of GAMaV and the Greek virus to the genus Marafivirus. Vitis 33. Double stranded RNA associated with fleck disease of grapevine. 105 . References Abou Ghanem-Sabanadzovic.: Description of the family Tymoviridae. South Africa. 195.P.. and G. N. Abou Ghanem-Sabanadzovic et al. Progress in the study of the phloem-limited isometric virus-like particles associated with leafroll-diseased grapevines. Shi et al. GRVFV recorded from California and confirmation that GAMaV does not occur outside of California. 1996. Abou Ghanem-Sabanadzovic et al. Dolja V.. Engelbrecht D. Regeneration of virus-free grapevines using in vitro apical culture. 347-354. Vitis 22. 165-169. Sabanadzovic ..G. G. Multiplex RTPCR detection of Grapervine fleck virus-like viruses in grapevine with co-amplification of control plant mRNA. Kasdorf. Boscia D. Martelli. G.. S. A. Martelli. V.M. 1995.P. V. 56-64. 101-102 Boscia D. Locorotondo 2003. Sequencing of the 3' end of three grapevine fleck virus-like viruses . Skene. Minafra. 1994. 1991b. Castellano M. Association of a phloem-limited non mechanically transmissible isometric virus with grapevine fleck disease.2002a 2002b 2003a Martelli et al. G. Castellano.E.A. Martelli.
W. 143-146. Gugerli P. Investigating the transmission of marbrure and fan-leaf through the seed in the grapevine. 1970. Tremea. Martelli G.. M. Rives. Ottenwaelter. Gonsalves.J. 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. M. Habili and R. 1972. D. Boscia.IV. Saldarelli. IV. 31-32. Seddas. Gooding. Lisbon 1997. A. Asteroid mosaic of grapevine. Rives. Resta. Ramel and F. and C. 567-571.. L.. Rivista di Patologia Vegetale. and J. 1972.T. Goheen. Belin and B. Rives M. Procedures for rapid detection of virus and viruslike diseases of grapevine. Schieber O. G. 1998. S. Annales de Phytopathologie.H. rupestris "du Lot" (St. with Special Reference to Vitis. Journal of General Virology 82.P. T. 75-77. Informatore Fitopatologico 46 (12).. 1991. Ser. Heat therapy eliminates the ability to transmit the causal agent of "marbrure" in several V. Phytopathologia Mediterranea 24. 9.M. Triolo E. Acta Phytopathologica Academiae Scientiarum Hungaricae 9. Parsons. 618-622.. Studies on virus diseases of the grapevine in California. serological characterization and immunopurification of grapevine fleck virus. 1974. Grapevine fleck virus-like viruses in Vitis. Fitopatologia Brasileira 20. 1991. S. is transmitted by graft inoculation. Heat inactivation of viruses in grapevines.B.B. Gugerli. Hévin M. J..): Virus Diseases of Small Fruits and Grapevines (A Handbook). 287-289. Digiaro and G.A. 1996. D. 1973. A. Castellano. 2002a. N. Further characterization of grapevine leafroll disease. Incidence of some graft-transmissible virus-like diseases of grapevine in visually selected and heat-treated stocks from Southern Italy. Fortusini A. Annals of Applied Biology 142. 767-774.C.L. Abou Ghanem-Sabanadzovic. 2003. 385-388.. 2001. Bonnard. and A.C.. Raski and G. George). Mink G. Cory and C. Purification and properties of spherical virus particles associated with grapevine Ajinashica disease. S.M. P.P. M.. C. Complete nucleotide sequence and genome organization of grapevine fleck virus. Maculavirus.J.. N. Bulletin of the California Department of Agriculture 43. Phytopathologia Mediterranea 24. Proceedings 10th Meeting of ICVG. Refatti E.or chlamidia-like bodies in grape. 1966. Mycoplasma. S. 106 . 1847-1853.C Edwards and T. Doazan and M. Archives of Virology 147.. 1954. (Ed. Ser. 9. N. Prati. a new genus of plant viruses. D. Nucleotide sequence variation in a small region of Grapevine fleck virus replicase provide evidence for two sequence variants of the virus. 1985. 281-285. 2000. Annals of the Phytopathologial Society of Japan 64. and E. Luhn. M. Brugger. N. Division of Agricultural Sciences. Martelli. 47-64. Kuniyuki H. M.. latent in many varieties. Saldarelli and G.. Grapevine fleck disease. 1977. Hévin. 1997.C. Kyriakopoulou P. Refatti E. Symons. Dreher.P. 1973. In Frazier N. 1985. Hewitt W. Sabanadzovic. 560-564. Doazan and M. M.IV.E. Annales de Phytopathologie. Rivista di Patologia Vegetale. 553-565.V. Leclair.Faoro F and P. 12491253. Davis 1965. 1962. Ser. vinifera clones and in V. Hewitt W. Archives of Virology 147. Plant Disease 75. 9. Matsumoto T.P. University of California.I. Jr.P. Yamashita. Milkus B. Tsuchizaki and D.P. Cytological alterations associated with an unidentified isometric grapevine virus (UIGV). 2009-2015 Savino V.S. J.. Goheen. Grapevine asteroid mosaic.B. Rivista di Patologia Vegetale. Sabanadzovic S. The responses of the Grapevine fleck agent to tetracycline-HCl antibiotic and Dienes' stain. A possible new necrotic diseases of grapevine associated with small isometric particles and novel membrane-bound large particles. Walter. A. Martelli.P. B. 39-43. Hewitt W. Symptoms of grapevine asteroid mosaic in Greece. Berkeley. 2002b. N.. 253-258. 1991. Costa. Shi B. Abou Ghanem. Extended Abstracts 12th Meeting of ICVG. Vitis 3. J.. Boscia and G. Plant Disease Reporter 61.. 197-203. 1973. The family Tymoviridae. Some virus and virus-like diseases of grapevines. Abou Ghanem-Sabanadzovic. Numéro hors série. 349-355. Archives of Virology 145. Volos 1990. Rosciglione. Martelli. Martelli G. 43-47. European Journal of Plant Pathology 103.. 204-207. Abou Ghanem-Sabanadzovic and P. Cinquanta and S. S. Proceedings 10th Meeting of ICVG. Volos 1990. Goheen A.. Scattini. 1997.. Ottenwaelter M. Namba S. Séparation de la marbrure et du court-noué (panachure) chez la vigne par thermothérapie.-E. Proceedings International Conference on Virus and Vector on Perennial Hosts. 1837-1846. Ohki.-J. Numéro hors série. P. Sabanadzovic S. 212-214. 1995. 57-83. and S. Sabanadzovic. Luhn. 157-164. affected by marbour. Monoclonal antibodies for detection. Occorrencia de mais um isolado do virus do mosaico da nervuras da videira que no causa sintomas no porta-enxerto Kober 5BB. 5960.
George. Krake. Walter B.Triolo E. Investigations on transmission of grapevine leafroll. 221-226. 1983. 107 . R. Kuszala. 67-73. Legin and J. La Recherche Agtonomique en Suisse 26.. ELISA detection of grapevine fleck virus (GFkV). La maculatura infettiva della vite: influenza di isolati diversi sull'attitudine alla propagazione vegetativa di Vitis rupestris St. Archiv für Phytopathologie und Pflanzenschutz 19. yellow speckle and fleck diseases by dodder..G. Yamakawa Y. 193-198. Numéro hors série. 3209-324. 1989.R.C. 1966. Semtschik. Ätiologie und Diagnose der Marmorierung der Weinrebe. probablement indépendante du virus du Court-noué. Phytopathologische Zeitschrift 106. Virus reinfection of virus-free Cabernet sauvingon and Cabernet franc vines. 1987. Materazzi. and A. Cornuet. 297-302. and L. Verderevskaja T. Observations sur une mosaïque de la Vigne. Marinesku and E.D.S. and P.. Journal of the Japanese Society of Horticultural Science 58. Woodham R. Vuittenez A. V. Annales des Epiphyties 17.. 1993. Agronomie 13. 1983. 651-657.
A. rings and lines are typical summer responses of infected vines. 18% of the particle weight. There may be differential susceptibility among cultivars. Beczner and Lehoczky: AMV infections recorded from Hungary. Main symptoms: Various patterns of yellow discolouration characterize the disease. a mechanically transmissible virus. Bovey and Cazelles: AMV particles visualized in thin sectioned grapevine leaves. has differently shaped particles. Jankulova: AMV infections recorded from Bulgaria. with Mr 24x 103 Da.MINOR VIRUS DISEASES Several graft-transmissible diseases are known. Faint yellow speckling. The spring growth shows more or less extensive yellowing of the leaf blades that does not extend to the veins. Their overall importance is minor if compared with that of the major diseases dealt with in previous chapters. Description Main synonyms: None. 2. Agent: Alfalfa mosaic virus (AMV). In grapevines. 0. however. Control: Use of healthy material obtained by heat treatment. Novak and Lanzova: AMV infections recorded from hop and grapevine in Czechoslovakia. Detection: AMV is mechanically transmissible to herbaceous hosts and can also be identified by ELISA and moleculat techniques in infected vines. with the following mol. Plant vigour and yield do not seem appreciably affected. Historical review 1973 1975 1976 1979 1978 1981 Bercks et al. Some of these diseases have been recorded only from Europe.62 x 106 (2037 nt). Capsid proteins subunits are of one type. RNA-2. Hungary. European diseases GRAPEVINE YELLOW MOTTLE 1. 109 . and Turkey.04 x 106 Da (3644 nt). wts: RNA-1. Grapevine berry inner necrosis).: First record of AMV infections and description of symptoms in German grapevines. Bovey and Brugger: AMV recorded from Switzerland in grapevine and transmitted by grafting to V. AMV. Transmission: AMV is efficiently transmitted by aphids in a non persistent manner and can cause epidemic outbreaks in many of its natural hosts. RNA-3. Yellow mottle has been reported from Germany. the type species of the genus Alfamovirus.g. Switzerland. is the putative causal agent. 1. and a tripartite RNA genome accounting for c. Bulgaria. but some are of economic relevance locally (e. suggesting that the virus spreads primarily through infected planting material. infections are scattered and occasional. 0. Chardonnay and Veltliner rouge précoce identified as reliable indicators. Varietal susceptibility: Little information available. Virus elimination by treating 37 days at 37-38°C. others occur in Japan. rupestris and the hybrid Grézot 1 x 5C.73 x 106 (2593 nt). former Czechoslovakia. from quasi isometric to bacilliform. with which specific viruses are associated and thought to be their possible causal agents. 30 to 57 nm in size.
1976. Le virus de la mosaïque de la luzerne sur la vigne. Querfurth.I. Vigour and yield are reduced. Über den Nachweis des Alfalfa mosaic virus in einer Weinrebe. Phytopathologische Zeitschrift 76.18 .1985 1993 1993 Francki: Comprehensive review of the properties of AMV and other viruses with tripartite genome. Alfalfa mosaic virus on grapevine.) and grapevine (Vitis vinifera subsp. or maple leaf-like line patterns typically confined to the petiolar area. 1993. Journal of Turkish Phytopathology 22. Lehoczky. and O. 152-154. Akbas and Erdiller: AMV infections recorded from Turkey. scattered spots or blotches. Francki. (ed. 119-128. Historical rewiev 1985 Francki: Comprehensive review of the properties of AMV an other viruses with tripartite genome. Control: No information. or the upper part of the blade.P. Identification of alfalfa mosaic virus and tomato bushy stunt virus in hop (Humulus lupulus L. vinifera cultivars are susceptible. Francki R. 2. Investigation on certain viruses spread on grapevines in Bulgaria. has differently shaped particles. Arboriculture. Munich 1978. 1-18. Bovey R.Hegi) plants in Czechoslovakia. Lesemann and G.B. quasi spherical 25-30 nm in diameter to bacilliform 40 to 75 nm in length. References Akbas B.I. and J. 3. Several V. Plenum Press. Acta Phytopathologica Academiae Scientiarum Hungaricae 16. 1985. Erdiller. Horticulture 7. and G. Proceedings 6th Meeting of ICVG. . Martelli: Yellow mottle suggested as the name for the disease caused by AMV in grapevines. In: The Plant Viruses. 110 . Detection and Diagnosis of Graft Transmissible Diseases of Grapevines. Description Main synonyms: None. and J. Grapevine line pattern virus (GLPV) a possible member of the genus Ilarvirus. Jankulova M. 131-134. 39.). Jubileum 75. 63-65. Novak J.). Detection: GLPV is mechanically transmissible to herbaceous hosts. This disease is known to occur only in Hungary. Biologia Plantarum 18. Grapevine disease in Hungary caused by alfalfa mosaic virus infection. Lanzova. Agent: The putative agent. New York . 1978. Revue Suisse de Viticulture. Brugger. Varietal susceptibility: No information. The viruses and their taxonomy. sativa DC. and a multipartite genome. Polyhedral Virions with Tripartite Genomes (R. Cordoba 1976. 1993. 3rd International Congress of Plant Pathology. 1981. is seed-transmitted and spreads with diseased propagative materials. 1973. Monografias INIA No.B.-J. Rome. Bovey R. Beczner L. Graft transmission to cv. Researches on grapevine virus diseases and determination of their incidence in Ankara Turkiye. GRAPEVINE LINE PATTERN 1. 1979. ed. Main symptoms: Leaves show bright yellow discolourations that form marginal rings. D. 55-63. FAO Publication Division. 1975.. Cazelles. 263 pp.B. Martelli G. and J. Bercks R. roughly following its contour. 166-171. Transmission: GLPV has no known vector..
Line pattern.P. Transmission: No vector is known. G. wt 1. one of the putative agents of corky bark (rugose wood complex) has a very high association with diseased grapevines. family Tombusviridae.A. Israel. Beczner and G. Castellano.: Description of line pattern disease in Hungary. according to more recent findings. Castellano.. 111 . Kiryat Anavim. Viruses associated with the disease are readily transmitted by sap inoculation and can be detected by ELISA and molecular techniques. D. M. Main symptoms: Symptoms are prominent in late summer and consist of yellow and/or reddish discolourations of the tissues along the veins. Lehoczky J. Bunches are reduced in numbers. 1992. 3. Martelli and J. The viruses and their taxonomy. D. Evidence of its grafttransmissibility. has a monopartite RNA genome accounting for c. Kertgazdasag 19. 1989. Martelli. Agent: Symptomatic grapevines were reported to be doubly infected by GFLV and Carnation mottle virus (CarMV) the type species of the genus Carmovirus. L. The disease is graft-transmissible and spreads through infected propagating material.1987 1989 1992 Lehoczky et al. Boscia. Proceedings 9th Meeting of ICVG. J. Occurrence of the line pattern hitherto unknown virus disease of grapevine in Hungary. Farkas. 1985. 115-116. Lehoczky et al. Burgyan. Plenum Press.). References Francki R. New York . Boscia.I. G. L. Varietal susceptibility: No information. RODITIS LEAF DISCOLORATION 1. Lehozcky J. Control: No information.B. Seed transmission of grapevine line pattern virus.A. the interveinal areas. Polyhedral Virions with Tripartite Genomes (R. Lazar. 18% of the particle weight. Farkas. Detection: Graft-transmission to V. However. CarMV is an isometric virus 30 nm in diameter. 1987. Evidence that a graft-and mechanically transmissible virus is associated with it.: Evidence that GLPV is transmitted through grapevine seeds. with Mol.B. Description Main synonyms: None. a novel virus disease of grapevine in Hungary.. GFLV may not be involved in the aetiology of the disease.I. 1987.. Historical review 1989 1991 Rumbos and Avgelis: Roditis leaf discoloration described in Greece. The disease has been recorded only from Greece. Grapevine virus B (GVB).. J.: Purification and characterization of GLPV and suggestion that it is the causal agent of the disease. Burgyan. Avgelis and Rumbos: Double infection of diseased vines by GFLV and CarMV reported. 61-79 Lehoczky J. In: The Plant Viruses. M. especially near the petiole Leaves are deformed in correspondence of discoloured sectors. vinifera cv. 1-18. Phytopathologia Mediterranea 31.P. Francki. By converse.4 x 106 (4003 nt in size) and coat protein subunits of Mr 38 x 103 Da. Lehoczky et al. 23-30. size and have low sugar content. or variously extended sectors of the leaf blade. Beczner and G. ed. 2. Mission.
Description Main synonyms: None Main symptoms: Grapevines of cv.E. and I. Rumbos. 19. Agent: Grapevine angular mosaic virus (GAMV).a new virus disease of grapevine: symptomatology and transmission to indicator plants. A. Carnation mottle virus isolated from vines affected with "Roditis leaf discoloration". The disease has been recorded only from Greece.I. 1345.: First record of GAMV. A. Bem. N. Tsagris. Proceedings 10th Meeting of ICVG. Whether this mechanism operates with grapevines has not been ascertained. References Girgis S. P. Infected grapevines are stunted. crinkling and deformation of the leaves. GRAPEVINE ANGULAR MOSAIC 1.I. mechanical transmision to herbaceous hosts. The etiology of a new virus disease: grapevine angular mosaic.C. Varietal susceptibility: No information Detection: Indexing on cv.M. Avgelis. Kyriakopoulou. Volos 1990. Katis. and A. Avgelis and N. Infected grafting material is likely to be responsible for virus dissemination. Journal of Phytopathology 125. reproduced the field syndrome in mechanically inoculated grapevine seedlings. A new ilarvirus isolated from grapevine in Greece. and ELISA. 3. C.: Evidence that GAMV is the agent of grapevine angular mosaic disease. a virus with a tripartite RNA genome and a 30 kDa coat protein. References Avgelis A. F. 1991. Historical review 2000 2003 Girgis et al. A. but differs from GLPV.E. Flowers abortion results in straggly bunches with small wrinkled berries bearing non viable seeds. 112 .. C. 2003. Rumbos I. Bem.D.D. Control: No information 2. Girgis et al. 437-443. Dovas. Girgis S.3. Description Main synonyms: None Main symptoms: Symptoms are chlorotic angular spots on the leaf blades. Tzortzakakis and M. discoloration of tissues bordering the veins. P. decline gradually and some die. GAMV is molecularly related to a number of ilarviruses. Plant Disease 84. Kyriakopoulou. Avgelis. 2000. Katis. Roditis leaf discoloration -. S. YELLOW LINE PATTERN (Rasperry bushy dwarf virus) 1. Transmission: GAMV is pollen-borne in herbaceous hosts and was able to infect pollinated plants. Extended Abstracts 14th Meeting of ICVG Locorotondo 2003. Dovas. Baresana x Baresana. 274-278. Laski Rizling from Slovenia exhibit a yellow line pattern syndrome resembling the grapevine line pattern disease described from Hungary. F. 1989. P. thus is regarded as the agent of the disease. P.M. Sklavounos.C.P. the closest being Tobacco streak virus (TSV).. the only other ilarvirus reported from grapevine.
berries are small and show external discolorations and internal necrosis. the 3' terminal region of which (2469 nts) has been sequenced. Description Main synonyms: None Main symptoms: Infected grapevines have low vigor.Agent: Raspberry bushy dwarf virus (RBDV) was isolated from symptomatic vines. Transmission: In raspberry the virus infects progeny seedlings and pollinated plants through pollen. Transmission: GINV is transmitted by grafting to grapevines and by mechanical inoculation to herbaceous hosts. F. and Pione whereas cvs. RBDV. Varietal susceptibility: No information Detection: Mechanical transmission to herbaceous hosts. GINV has filamentous particles about 750 nm in length and a single-stranded RNA genome with Mol. wt 0. is unknow. Extended Abstracts 14th Meeting of IGVG. No. The vay of natural spreading in grapevine. CMI/AAB Description of Plant Viruses. Grapevine berry inner necrosis has been reported only from Japan.g. 3. Agent: The disease agent is Grapevine berry inner necrosis virus (GINV). being transmitted by the eryophid mite Colomerus vitis. Virscek Marn and I. Healthy vines of cvs Kyoho and Pione became naturally infected in the field within one year from planting. 165 B. 20 Murant A. 2003. representing the most important virus disease in Yamanashi Prefecture. Raspberry bushy dwarf virus infection of grapevine in Slovenia. Varietal susceptibility: Symptom severity varies with the cultivar.5 x 106 Da.2 kb in size). 30 x 103). rings and line patterns are shown by leaf blades. and a bipartite singlestranded RNA genome accounting for c. and RT-PCR. Some rootstocks (e. References Mavric I. a dimeter of about 33 nm.: First record of RBDV in grapevine. Japanese diseases GRAPEVINE BERRY INNER NECROSIS 1. wt of 7. M. infected propagative material can disseminate the RBDV. Delaware. 113 . Zezlina. a mechanically transmissible definitive member of the genus Trichovirus. wt of 2 x 106 Da (5. Control : No information 2. Locorotondo 2003. delayed bud break and young shoots with short internodes and internal browning. if any. 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. 1976. Mavric et al. Campbell Early are suscetible as well as cvs Takao. ELISA .8 x 106 Da (2.. Historical review 1976 2003 Murant: Description of RBDV. Vitis riparia Gloire) are also susceptible. Almost all Japanese table grape cultivars derived from crosses with cv. However. Chlorotic mottling. The virus spreads naturally in the vineyards. Kyoho. Raspberry bushy dwarf virus. Ripening of bunches is delayed..5 Kb in size) and RNA-2 with Mol. 24% of the particle weight and consisting of two functional species RNA-1 with Mol. Koshu. and Kaiji are infected latently.
423.: Partial sequencing of GINV genome and assignement of the virus in the genus Trichovirus Nishijima et al. 617. Bulletin of Yamanashi Fruit Tree Experimental Station 10.: An account of the varietal susceptibility to the disease and natural field spread Kunigi et al.. 1984.: Experimental evidence that GINV is transmitted by the the grape erineum mite Colomerus vitis 1993 1997 2000 2000 3. Montreux 1993. Yoshikawa N. Bulletin of Yamanashi Fruit Tree Experimental Station 10. References Kunugi Y. Extended Abstracts 11th Meeting of ICVG. a new trichovirus: comparative studies with several known trichoviruses. Description Main synonyms: None. 1351-1363 GRAPEVINE STUNT 1. Studies on the grapevine berry innner necrosis virus disease. Transmission of grapevine berry inner necrosis virus by the grape erineum mite Colomerus vitis in Yamanashi. Terai and A. 1992. Nishijima T.. Goto. 536 Terai Y. S. 2000. Y. varietal susceptibility and natural spread. grapevine berry inner necrosis with natural spread in Japan. Yanase. Studies on the grapevine berry innner necrosis virus disease. Induction of berry necrosis in Kyoho back-inoculated with the virus isolate from grapevine mosaic diseased clones and renaming to grapevine berry inner necrosis. Control: Use of tolerant cultivars in areas where the disease spreads epidemically. A new virus disease. Iida..Annals of the Phytopathological Society of Japan 53. 1. S.. H. Asari. Yanase H. Magome.. Annals of the Phytopathological Society of Japan 55.Annals of the Phytopathological Society of Japan 58. Terai. Disease re-named Grapevine berry inner necrosis Terai et al. 362. Kunigi Y. 47-56. 1993. 1987. Grapevine berry inner necrosis. 2. Terai. Shinkai 2000. 114 . and Yanase H. Y. Y. and Terai Y.Detection: Indexing on cvs Kyoho or Pione. Tanaka H. Archives of Virology 142. Terai and Y.: First account of grapevine berry inner necrosis disease in a non Japanese publication. H. 2. GINV is mechanically transmissible to herbaceous hosts and can be identified by ELISA and moleculat techniques in infected vines. Mosaic symptoms on cv Kyoho.. T. 57-63. 77-78 Yanase H. Back-transmission of a grapevine filamentous virus to grapevine seedlings and induction of foliar and berry symptoms in grapevine.. Purification of a filamentous virus isolated from grapevine berry inner necrosis and foliar mosaic. and H. Kyoho back inoculated with the filamentous virus isolated from mosaic-diseased grapevines. Yoshikawa et al. 1997. Annals of the Phytopathological Society of Japan 51.. Takahashi and Y. Historical review 1984 1985 1987 1992 Tanaka: Description of a mosaic disease in cv. Symptoms on vines. Kunugi. 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. 1985.
M. Varietal susceptibility: No information. Doi and M. 396 Hatamoto M. Agent: An isometric. Spread occurs also through infected propagative material. Taiwan (Reference 2951 in the Review of Plant Pathology 66. 316. 33. Inflorescences are undersized. p. Doi and K. are pale-coloured and have a low sugar content. sometimes. curled and. Transmission: The disease is transmitted by the leafhopper Arboridia apicalis. The disease has only been reported from Japan. This virus is serologically distinct from the putative agent of ajinashika disease. The berries. 1984. Doi. Control: Use of disease-free material obtained through heat therapy. 1986 Namba et al. Namba. Evidence that it is not related to the presumed agent of ajinashika disease. S. FFTC Book series. This condition gives the name to the disease which in Japanese means "unpalatable fruits with low sugar content". Yamashita. Annals of the Phytopathological Society of Japan 50. Transmission of grapevine stunt disease by the grapevine leafhopper Arboridia apicalis Nawa. 85 Namba S.. Y. 3. Graft transmissibility of grapevine stunt disease. Historical review. non mechanically transmissible virus about 25 nm in diameter is consistently associated with diseased vines and regarded as the possible causal agent.. 137 Namba S. 1981 1982 1984 Namba et al. Yamashita and Y. Hatamoto et al. Yamashita. Annals of the Phytopathological Society of Japan 47. Food and Fertilizer Technology Center for the Asian and Pacific Region. M. Fujii. Because of heat recovery. 1982. Koshu nor any apparent reduction of vigour and yield. Three phloem-limited viruses of grapevine: direct fluorescence detection. S. 2. Namba. Yamashita and Y. GRAPEVINE AJINASHIKA DISEASE 1. S.. Yora. Description Main synonyms: none. Taipei. fruit setting is impaired and bunches are few and shelled. with scorched margins. S. Detection: Grafting to Campbell Early and ELISA using extracts from infected vine tissues. Hatamoto et al. however. Doi. 115 . Y. Annals of the Phytopathological Society of Japan 48. In Plant Virus Diseases of Horticultural Crops in the Tropics and Subtropics. vinifera cv. Iwanami. 1981.: Evidence that the disease is transmitted by the leafhopper Arboridia apicalis.Main symptoms: Spring vegetation is delayed. S. phloem-limited. T. References Hatamoto M. internodes are short. summer vegetation is apparently normal. 1986. The same paper appears in Taiwan Food and Fertilizer Technology Center Technical Bulletin 92. The disease is apparently restricted to the V. 1-17. A small spherical virus associated with grapevine stunt disease.: Successful graft-transmission of stunt disease. Campbell Early. No. Fujii. American rootstocks are infected without showing symptoms. which makes the crop unmarketable. leaves are small. Hatamoto. Main symptoms: No appreciable symptoms are visible on the foliage of cv. S. 1987).: Purification and characterization of the virus associated with stunt disease.109-126.. The disease has only been reported from Japan.: A small isometric virus associated with stunt disease in Japan.
Y. and D. an isometric.. Iwanami. Purification and properties of spherical virus particles associated with grapevine ajinashika disease. A small spherical virus associated with the ajinashika disease of Koshu grapevine. Namba S.. 12491253.Agent: The disease was reported to be caused by the concurrent infection of leafroll and fleck. No relationship found with fleck. 3. 1979. Terai and Yano: Description of ajinashika disease and suggestion that it is caused by the concomitant infection of leafroll and fleck. 70-73.: Further characterization of the isometric virus and claim that it is the putative agent of the disease. 2. Terai Y. Proceedings 10th Meeting of ICVG. Yamashita. and R. Varietal susceptibility: No information. S. Proceedings 7th Meeting of ICVG. Koshu. S. Taiwan Food and Fertilizer Technology Center Technical Bulletin 92. References Namba S. Namba S. Three phloem-limited viruses of grapevine: direct fluorescence detection. Transmission: No vector is known. Doi and M. However. Ajinashika disease of the grapevine cultivar Koshu in Japan. consistently found in infected vines. non mechanically transmissible virus about 25 nm in diameter. Namba et al. Historical review.: First mention of ajinashika disease and report of the association with it of a non mechanically transmissible virus with isometric particles. Plant Disease 75. Gonsalves. Koshu and ELISA using extracts from infected vine tissues. T. Yamashita.: Partial characterization of the isometric virus associated with the disease and its detection by ELISA in infected vines. Yora. 15-19. Control: Use of disease-free material obtained through heat therapy. Dissemination is through infected propagative material. Terai: Additional report on ajinashika disease as derived from the combined effect of leafroll and fleck. 116 . 1-17. 1980. 1991. 1991. Terai Y. phloem-limited. Tsuchizaki. 1979 1980 1986 1991 1991 Namba et al. Annals of the Phytopathological Society of Japan 45. Ajinashika disease: a combined effect of grapevine leafroll and grapevine fleck viruses on sugar content in the Japanese grape cultivar Koshu. Yamashita. Y. S. Yano. was suggested as the possible causal agent.. Volos 1990. T. Hatamoto. Doi and K. vinifera cv. 67-70. Detection: Graft transmission to cv. Niagara Falls 1980. The disease seems to be restricted to V. D. 1986. Namba et al. Boscia.
VIRUS-LIKE DISEASES .
growth tends to become normal again. and often show longitudinal cracks between the nodes. which gives a bushy aspect to the plant. However. The varieties Panse Precoce. Symptom expression varies year by year. The infectious agent of the disease perennates in propagating material. the absence of symptoms does not necessarily mean that the plant is healthy. some of which have a clear-cut detrimental effect on the crop. The transmission by graft is rather erratic. Symptoms have been observed on many V. with prominent veins. according to the cultivar) and is of poor quality. Gigante : Research on histological and cytological aspects of enations. Varietal susceptibility and sensitivity: There is little information available. whether they bear enations or not. Leaves developed later in the season are usually normal. Hewitt: Description of symptoms in California. ENATION DISEASE 1. Description Enation disease of grapevine is one of the oldest known disorders of European grapes. Grenache and Tokay appear to be quite susceptible and develop severe symptoms when infected. This hypothesis. malattia delle enazioni. They are outgrowths 2-3 mm high and 3-5 mm long or more. Detection: By observing the symptoms on the leaves and by indexing on LN 33 hybrid. but some are heat-labile and can be eliminated by heat therapy. They are often thicker than normal. but attempts to transmit it by graft or mechanical inoculation gave no results. vinifera varieties in Europe. which run more or less parallel to the main veins. 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. The basal internodes are short. Main symptoms: Affected vines show a delayed opening of the buds and a slow growth of the shoots in the spring. Riesling. Their agents are still unknown.). with drawings of symptoms. Historical review 1891 1937 1954 Buchenau: First description of enation disease of grapevine occurring in Germany.). Enations develop mostly on the underside of the leaves at the base of the shoots. There is no information on the possibility of curing this disease by heat treatment or meristem culture. Later in the year. has now been dismissed Transmission: By vegetative propagation. Primus. Latin America (Venezuela). Severely affected leaves drop prematurely. Graft transmission suggests that it is a virus disease. apparently in relation with climatic conditions.). as symptom expression is variable in successive years and graft transmission not 100 % successful. The crop can be drastically reduced (up to about 50%. maladie des énations (Fr. North (Tunisia) and South Africa. All persist in propagative material and are transmitted by grafting. Basal leaves. The disease is perpetuated by vegetative propagation. Australia and New Zealand. 119 . Italia. however. omeoplasie crestiformi (Ital. and is probably due to a virus-like agent. irregular and mis-shapen. Control: Use of healthy material. The disease has been reported from many European and extra-European countries Agent: The etiology of enation disease is not yet clear. its description dating back to the late 1800s Main synonyms: Enationenkrankheit der Rebe (Germ. 2. are often mis-shapen.VIRUS-LIKE DISEASES Several latent or semi-latent grapevine diseases are known. North America (California). with a fanlike aspect and abnormal indentation.
Weinberg und Keller 15.: Successful transmission of enation disease from diseased to healthy grapevine by graft strongly supports the hypothesis of a viral origin. the hybrid LN 33 was found to be the most sensitive and reliable indicator variety. Xafis. Brückbauer: Description of symptoms of enation in Germany and confirmation of graft transmission of its agent.: In experiments on graft transmission of enation disease aimed at determining the best indicator. Martelli et al. with negative results. attempts to transmit the disease by graft. The authors discuss the hypothesis that enation is caused by a strain of fanleaf virus. Malvasia (10. Confirmation of graft transmissibility of the disease. 1968. Padilla et al. but diseased vines which had not shown enation symptoms for several years had almost normal yields. References Avgelis A.4 to 48. Nieder: Enation disease in Austria Garau et al. lowest in cv. 195 Brückbauer H. and possibly caused by a virus. historical account. There is some evidence that enation can be carried in the rootstocks. and C. Prota et al.: Enation disease in Czechoslovakia Avgelis and Xafis: Enation disease in Greece Prota and Garau: Enation disease found in Sardinia.1966 Graniti et al. Beobachtungen und Untersuchungen über die Enationenkrankheit der Rebe. Enationaffected vines produced less than 50 % of the yield of healthy plants.5%).: Enation disease in France Pozdena et al.: Detailed description of macroscopic and microscopic symptoms of enation disease. Trebbiano romagnolo and reduces the yield from 13% to 23% according to the severità of symptom expression. Vernaccina (1. The possible role of fanleaf virus in the etiology of enation requires further investigation. Italia in Sardinia.: Enation disease in Turkey Hevin et al. Mechanical transmission tests were also negative. 120 . but report on observations made in Australia where no fanleaf virus could be recovered from enation-affected vines. McGechan: Enation disease in Australia Tekinel et al.: More data on the effects of enation on the yield of cv. 79-112. . Phytopathologia Mediterranea 17. Graniti and Martelli: Review paper on enation. Presence of enations in Razaki grapevine in Crete (Greece). Refatti: Hypothesis of a correlation between fanleaf and enation disease.3% . 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. The authors conclude that the disease is probably of European origin. although the rate of symptom expression does not exceed 30% Credi: Enation diseasae affects the vegetative vigour of cv. In the vineyards under observation. Marinesku and Bondarchuk: Enation disease in Moldova Brückbauer: Influence of enation disease on growth and yield of grapevine in West Germany. 1978.5%). the proportion of diseased vines was highest in cv. only fanleaf virus was recovered. The mean yield loss of diseased vines ranged from 17..
G. Credi R. Effetto della malattia delle enazioni della vite sulla produzione e sullo sviluppo vegetativo della cv Trebbiano romagnolo. Prota U. 1971. Prota U. 326332. Z. Dolar. 2. it was clear that vein mosaic was not caused by fanleaf virus. Enations of grapevine in Sardinia. A similar disease has been reported in Australia under the name of summer mottle. Proceedings 9th Meeting of ICVG. Occurence of grapevine enations in the Moldavian SSR. Garau. Bitki Koruma Buletin 11. Davis.G. Ita and F.P. Rivista di Patologia Vegetale S. and M.. Division of Agricultural Sciences. Garau R. and R. Studies on some variable characters of grapevines affected by enation disease in Sardinia. Abnorme Blattbildungen. Pflanzenharzt 36.. 1970. M. Pozdena J. IV. Garcia. Cugusi.). Rivista di Patologia Vegetale. with Special Reference to Vitis. B. 17. 46-51. Israel. Agricultural Gazette of New South Wales 81.). 1954.. 1983.s. 1997. Martelli G.). Salcan. Kiryat Anavim. Berkeley. n. 1987. 97-98. Mededel. Ricerche istologiche sulle omeoplasie crestiformi (Enations) delle foglie di vite affette da rachitismo. (ed. 1981. Padilla V. Graniti. A. Lisbon 1997. (N.. Tekinel N. 1979. And G.. Chabbouh N and V. Trasmissione per innesto della "malattia delle enazioni" della vite.). Enation symptoms found in France. In: Verderevskaya T. Rives. Vein mosaic appears to be of low economic importance.. 121 . 1965. Refatti E. Phytopathologia Mediterranea 20. 349-352. 251-252 Hewitt W. Bolletino della Regia Stazione di Patologia Vegetale. Enations. Symptom expression seems to depend on climatic conditions. Garau R. 203-206. Main synonyms: Mosaïque des nervures (Fr. Leclair and M. Studies on reproduction of enation symptoms by grafting in Sardinia. and V. Martelli. 1996. Vanek. Quacquarelli. 1975. California. 1973. G. This disease is widespread and probably worldwide. Deutsches Weinbau-Jahrbuch 31. Bericht der deutschen botanischen Gesellschaft 9. 59-64.P. 47. Grapevine enation disease in Murcia (Spain).B. 1976. U. Phytopathologia Mediterranea 5. Extended Abstracts 12th Meeting of ICVG. Lamberti and A. Monografias INIA 18.Brückbauer H.W. 122-124. Buchenau F. 48.P. Important virus diseases of grapevine in New South Wales. I.IV. University of California. Savino. and G. Proceedings International Conference on Virus and Vector on Perennial Hosts..) Virus and mycoplasma-like diseases of fruit trees. Einfluss von Virusinfektionen auf Wachstum und Ertrag der Rebe. Prota and M. 7-12. 1980. 179-189.S. A study of infectious degeneration (fanleaf) in vineyards of the Mediterranean region. Spain. Die Eantionenkrankheit der rebe erstmal auch in Osterreich nachgewiesen.. Su una possibile correlazione fra il virus del complesso dell'arricciamento e la malattia delle enazioni della vite. areas of the leaf blade may become necrotic. VEIN MOSAIC 1. 1937. Cordoba. Hevin M.W. Enation disease of grapevine in Italy. 207-217. 9. Main symptoms: Pale green mosaic affecting mostly the tissues adjacent to the main veins or the smaller ones. Moldavian Research Institute of Horticulture Kishinev. 1966. Nas. 1891. Gigante R. 169-192. Marinesku V..V. Bulletin of the California Department of Agriculture 43.. Cugusi. Description The symptoms of vein mosaic have been confused for some time with those of fanleaf/yellow mosaic. Ser. 1997. Lamberti. but these necroses do not affect the veins as it is the case with vein necrosis. 241-243.1989. but when fanleaf virus transmission to herbaceous hosts became possible. 225-246. Nieder G.. Salih and Y. Facultet Landbouwwetenschap (Gent) 40. Adernmosaik (Germ. Frazier N. Proceedings 6th Meeting of ICVG. producing often a vein banding effect. Lisbon 1997. Some virus and virus-like diseases of grapevines.. Martelli and F. 1966. 1970. Graniti A. Petria 6. 47-64. 145-152. Occurrence of enation disease in Tunisia. 1966. Roma. In: Virus Diseases of Small Fruits and Grapevines (A Handbook). 293-306.P. Bondarchuk. Gazeau. O.. Extended Abstracts 12th Meeting of ICVG.K. The research of the virus diseases of grapevine in CSSR. Graniti A. small fruit and grapevine in Moldavia. H. In the most sensitive varieties. 1983. 823-827. Mosaico delle nervature (Ital. Benayas. F. McGechan J.D..
in the absence of any detectable virus. 2. Vuittenez and Stocky: Electron microscope study of thin-sectioned tissues of leaves from Vitis riparia and Vitis vinifera cv. Babini and A. Pop: Vein mosaic in Romania. Canova. Woodham and Krake: Comparison of summer mottle and vein mosaic.: Vein mosaic in URSS. Krake L. Chasselas. 1979 1980 1982 1983 1985 1993 2004 3. Ehrenfelser showing symptoms of vein mosaic. Potential interaction between rootstocks and grapevine latent viruses. Phytopathologia Mediterranea 24. Servant.: Vein mosaic in Ukraine.. A number of cytological modifications primarily involving chloroplasts were observed along with the presence of bundles of filamentous structures resembling closterovirus particles. 266-270. Samonina et al. 1993. American Journal of Enology and Viticulture 44. A. Varietal susceptibility and sensitivity: Vitis riparia Gloire de Montpellier is especially sensitive and is used as indicator. LN 33 is also sensitive. Saric and Hranuelli: Vein mosaic in Croatia. vein mosaic and vein necrosis. Control: Use of indexed material. 1985. Abracheva: Vein mosaic in Bulgaria. and R. Muscat de Hambourg. Detection: Indexing with V. vinifera cvs.A.: Observation of a type of mosaic of grapevines which appears to be independent of fanleaf virus. Marinesku and Bondarchuk: Vein mosaic in Moldova. Pinot. References Credi R. Krake and Woodham: Description in Australia of a systemic mottling syndrome which is expressed during summer on the leaves of some varieties. Pearl of Csaba). and Gamay apparently show little or no symptoms.R. Viognier. 17-23. 148-152. 1978. Woodham. Golino: Vein mosaic in California. No vector known.. The disease can be eliminated by heat therapy. Transmission: By graft and vegetative propagation. Several V. No claim is made that these putative viruses are connected with the disease. Chardonnay.R. but this hypothesis has not been confirmed. Legin and Vuittenez: Description of vein mosaic. Vitis 17.C. Grapevine summer mottle: a new graft-transmissible disease. Alphonse Lavallée. Milkus et al. Occurrence of grapevine vein necrosis and grapevine vein mosaic in the Emilia-Romagna region (northern Italy).Agent: Unknown. Kuniyuki: Vein mosaic in Brazil. Comparison of symptoms of fleck. riparia Gloire de Montpellier. Symptoms are very similar to those of vein mosaic in Europe. show symptoms (Syrah. Historical review 1966 1973 1973 1976 1973 1973 1978 Vuittenez et al. Bonfiglioli: Vein mosaic in New Zealand (unpublished). Mycoplasma-like organisms were supposed to be the cause of vein mosaic. 122 . Golino D.
Saric A. Agent: Unknown. B. Mostar 1977. R. 1. Pop I. 149-141. Control: Use of disease-free propagating material obtained by culture of fragmented shoot apices. 1966.. probablement indépendante du virus du Court-noué. Adverse effect of light and of high temperature on symptom expression of grapevine vein mosaic in Sao Paulo. Kuszala. 9 . and G. Kuniyuki H. 1973. Milkus. Symptoms of vein mosaic develop under mild weather conditions and fade during hot weather. Vuittenez A. several European grape cultivars show symptoms. suspected to be a virus or a viroid. Krylova.. URSS.. Historical review 1978 1982 1983 Krake and Woodham: Description of summer mottle in Australia. Detection: Graft transmission to a number of cvs. Bondarchuk. 243-250 Samonina I. Woodham and Krake: Comparative graft transmission trials demonstrate that summer mottle 123 .V. 1983. Evidence that the disease is graft-transmissible.-Berl. and L. 48-49 Marinesku V.V. Barlass et al. Legin and J. Mataro. Main symptoms: Pale green to yellowish dicolourations of the tissues adjacent to the main or secondary veins. Vuittenez A. Vitis 22. Investiation on grapevine viruses in the SR Croatia. However. Proceedings 7th Meeting of IGCV.N. rupestris and LN33 are infected symptomlessly. 9. 67-73. whereas the opposite occurs with summer mottle. Stocky. ou la “feuille rouge”. and V. La mosaique des nervures. 68-72.. 1977. A grapevine virus disease in the Primorye territory.: Elimination of the disease agent by culturing fragmented shoot apices. These symptoms appear in summer and persist through the autumn.R. Comparaison des symptômes et transmission par greffage d'une mosaïque nervaire de Vitis vinifera . Moldavii 31. Woodham R. resembles in some respects the European vein mosaic and the Greek Roditis leaf discolouration (see under Minor virus diseases). Rivista di Patologia Vegetale . IV.Legin R. poorly developed and with small berries. Mission. 41-42. SUMMER MOTTLE Summer mottle. Observations sur une Mosaïque de la Vigne. IV . 57-63. 110 R.G. Bunches of infected cvs Sideritis and Cabernet sauvignon are fewer. A. producing a feathering or banding effect. Annales des Epiphyties 17. Description Main synonyms: None.V. an Australian disease.N. Transmission: No vector is known. Ser. Cabernet sauvignon. 2. 1985. 247-252. and A. de la marbrure de V. Symptoms show on vegetative growth that develops at temperatures in excess of 30 °C.. and Hranuelli T. 191-204. Rivista di Patologia Vegetale S IV 9.V. Numéro hors série. Vuittenez..g.C. 1973. Roditis leaf discolouration and summer mottle have similarities suggesting that they may be the same disease. Krylov and V. Ultrstructure de vignes infecteés par deux maladies de type viral: la mosaique des nervures. Vinodel. 1976. Spread is through infected propagative material but is has also been observed between adjacent vines Varietal susceptibility: No grapevine tested was immune to infection. e. maladie à virus de la vigne. S. 1982. Proceedings of the Conference on Excoriosis and Virus Diseases of Grapevine. Niagara Falls 1980. Summa Phytopathologica 11. Cabernet franc. Krake. V. 1973. Grapevine vein mosaic. rupestris et d'une affection nécrotique des nervures de l'hybride Rup. Vinogradr. A comparison of grapevine summer mottle and vein mosaic diseases. Rivista di Patologia Vegetale.
differs from vein mosaic. 247-252. Many grapevine varieties and rootstocks are infected symptomlessly. Taylor. Possible viroidal etiology put forward. rupestris x V. 1999. Cause-effect relationships between MLOs and the disease has never been ascertained. most likely GRSPaV. growth is much reduced and necrosis of the leaf veins appears. Woodham and L. 291-295.R. Krake. 1982. Also the tendrils and many shoots can necrotize..C. Martelli et al.: Vein necrosis in Italy and Bulgaria 124 .C. Main synonyms: Nécrose des nervures (Fr. Graft-transmissible Diseases of Grapevines.A. 2. especially under greenhouse conditions.: Suggestion that summer mottle and Roditis leaf discolouration are the same disease 3. later on younger leaves as they develop. Transmission: By grafting and vegetative propagation. Detection: By grafting on 110 R. Collingwood.S.R. at first on the leaves at the base of the shoots. M. Control: Use of indexed planting material. CSIRO Publishing. This strongly supports the hypothesis that vein necrosis is a specificic reaction of the rootstock 110 R to GRSPaV infection. Woodham. No vector known.). 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. Historical review 1973 1978 1978 Legin and Vuittenez: Discovery and description of this virus-like disease while searching for indicators for fleck. References Barlass M. The agent of vein necrosis can be eliminated by heat therapy..). berlandieri 110 Richter (110R) with vein necrosis symptoms. 266-270. Adernnekrose (Germ.R. A comparison of grapevine summer mottle and vein mosaic diseases. Description Vein necrosis has probably a worldwide distribution. 70-74. Vitis 17. Australia. Agent: Suspected to be a virus. and some infected plants may die. 1. but their etiological relationship with the disease has not been proven.M. Regeneration of virus-free grapevines using in vitro apical culture.H. Necrotic reactions are best seen on the lower face of the leaf blade. Mycoplasma-like organisms have been observed in the phloem of symptomatic vines.C. recently an association exceeding 95% has been experimentally observed between Grapevine rupestris stem pitting-associated virus (GRSPaV) and vines of V.. Rezaian and R. Little is known about sensitivity of other Vitis species.). 1978. Woodham R. Krake.G. and L. N. R. Krake L. varieties or hybrids. K. VEIN NECROSIS Although vein necrosis is currently regarded as a disease in its own right. 1999 Krake et al. Grapevine summer mottle: a new graft-transmissible disease. So far. Scott. and the only Vitis species that is clearly affected is the rootstock 110 R. Krake L. Main symptoms: On the rootstock 110 R. and R. Annals of Applied Biology 101. its economic importance has not been assessed. Milkus and Kalashyan: Mycoplasma-like organisms found in phloem tissues of vines with vein necrosis. Varietal susceptibility and sensitivity: The rootstock 110 R is most sensitive. Skene. 1983.R. necrosi delle nervature (Ital. Vitis 22.
P.. Kalashyan. Golino D. Kergazdasag 18 (4)..C. Bulletin OEPP/EPPO Bulletin 22.N. Journal of the Australian Institute of Agricultural Sciences 50.A. rupestris et d'une affection nécrotique des nervures de l'hybride Rup. 110 R.: An association exceeding 95% observed between GRSPaV and 110R vines showing vein necrosis symptoms in indexing trials.. Phytopathologia Mediterranea 24. D.C. Gursoy Y. 1985. fleck and leafroll in Vitis vinifera and grapevine rootstocks in central Greece. 1988. Galea Souchet. Milkus B. 125 . Grapevine vein necrosis disease detected in rooststocks in Australia.P. D. Fitopatologia Brasileira 19. 1992. Martelli. J.. Necrosi delle nervature della vite in Italia e Bulgaria. P. SSR. Suggestion than vein necrosis is a specificic reaction of 110R to GRAPaV. V. Castellano. G.A. Minafra and G.Z. Phytoparasitica 17. V. C.: In southern Italy. 1994. Viruses of grapevine in Malta. La Notte. A. Lehoczky et al.P. 1978. Vuittenez.B.1984 1985 Woodham R. 29-30. Lehozcky J. Comparaison des symptômes et transmission par greffage d'une mosaïque nervaire de Vitis vinifera. 1986 1988 1989 1992 1993 1994 2004 3.. 1986. Vein necrosis.A. Martelli. M. Martelli G. Vein necrosis: new virus-like disease in Turkish vineyards. Journal of Turkish Phytopathology 17.-Berl. Is Grapevine vein necrosis a reaction to Grapevine rupestris stem pitting-associated virus? Journal of Plant Pathology 86. 1989. Rumbos I C. American Journal of Enology and Viticulture 44. Woodham R.5 %.P. Pirolo. 43-45 Kuhn G. Ser. and L. 1993. 57-63. i Chim Nauka 1. Biol. No veing necrosis observed in 110R top grafted on GRSPaV-free V. IV.. Incidence of some graft-transmissible virus-like diseases of Grapevine in visually selected and heat-treated stocks from Southern Italy. Savino. ser. 61 Savino V. References Bouyahia H. Potential interaction between rootstocks and grapevine latent viruses.: Vein necrosis in Malta Golino: Vein necrosis in California Khun: Vein necrosis in Brazil Bouyahia et al. Krake. Informatore Fitopatologico 28 (10). 3-5 Martelli G. rupestris. the incidence of vein necrosis in visually selected stocks of table and wine grape varieties is on the average 71 %. 301. 204-207. 79-83 Legin R.. Rivista di Patologia Vegetale. 2004. 59-65.R. 148-152. Rosciglione. Vein necrosis a disease that is latent in most grapevine cultivars of the State of Rio grande do Sul. 1973.. Detection of vein necrosis virus (GVNV) in the vines of cultivated grape varieties. D. and J. but did not eliminate the disease entirely.P. and L R. and A. Abracheva and B. Boscia. Savino. H. 1984. 1978. 606-612. Krake: Vein necrosis in Australia Savino et al. Farkas. Savino. Lazar.: Vein necrosis in Hungary Gursoy: Vein necrosis in Turkey Rumbos: Vein necrosis in Greese Martelli et al. 9. de la marbrure de V. Mycoplasma-like bodies in phloem tissue of grapevine affected by vein necrosis. 58-60. Izvestiya Akademii nauk Moldav. Heat therapy reduced this proportion to 35. Boscia and G. Boscia and V..
VIROID (Yellow speckle) .
a disease characterized by chrome yellow flecks localized along the main veins of mature leaves and progressing into the interveinal areas. 370 pp. 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. These symptoms appear in the height of summer on a limited number of mature leaves and persist for the rest of the vegetative season. it did not induce symptoms. inducing a disease called yellow speckle. genera and species.). Australian grapevine viroid (AGVd). AGVd has been reported from Australia. however. 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. Two families are known. Hop stunt viroid grapevine strain (HSVd-g). vein banding-like symptoms can be observed in vines infected only by yellow speckle viroids. YELLOW SPECKLE 1. R. the non coding genomes. GYSVd-1 and GYSV-2 have a worldwide distribution The three additional viroids that have been detected in grapevines. It was first detected in Japan and transmitted to cucumber and grapevine seedlings in which. Viroids. Like viruses. are subviral pathogerns endowed with autonomous replication in their hosts. CSIRO Publishing. Five grapevine-infecting viroids are known. Only GYSVd-1 and GYSVd-2 are pathogenic. the type species of the genus Hostuviroid. and perhaps the type of infecting viroidal sequence variant.S. Collingwood. picchiettatura gialla (Ital. CEVd-g.).). Sometimes. respectively and both belong in the genus Apscaviroid.. 129 . AGVd. 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. Very often. Both these viroids wer first isolated in Australia. however. climatic conditions. are not associated to any specific symptomatology. Description Main synonyms: Moucheture jaune (Fr. Kyoto vine with yellow speckle symptoms. HSVd-g. and AGVd. The symptomatology varies depending on the cultivar. Flores. GYSVd-1 and GYSVd -2 cause the disease individually or in combination. Cabernet franc and a cv. Main symptoms: Few to many minute chrome yellow spots or flecks scattered over the leaf surface. Agents: Two distinct viroids. Neither of them is able to replicate in herbaceous hosts but both were succesfully inoculated to grapevine seedlings reproducing the yellow speckle syndrome. Gelbsprenkelung der Rebe (Germ. plant age. Semancik (eds. infected vines are symptomless or show symptoms erratically. and Tunisia. a size much smaller than that the smallest viral genome. Vein banding.). HSVd-g has been recorded from Australia. or gathering along the main veins to give a vein banding pattern. a member of the genus Apscaviroid. all of which belong in the family Pospiviroideae: Grapevine yellow speckle viroid 1 (GYSVd-1). Grapevine yellow speckle viroid 2 (GYSVd-2). north and south America. Interestingly. Europe.W. J. thought to be elicited by a specific strain of GFLV.VIROIDS Viroids. has a genome 369 nt in size. viroids are classified in families. They are made up of a non encapsidated circular RNA of 246-375 nucleotides. GYSVd 1 and GYSVd 2 are made up of 366 and 363 nucleotides (nt). and plastidial replication (Avsunviroideae ). HSVd-g. Randles and J. Citrus exocortis viroid grapevine strain (CEVd-g). 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. References Hadidi A. was demonstrated to be caused by a co-infection by yellow speckle viroids and GFLV. respectively from a cv. presence of ribozymes. 2003. the USA. and may have a worldwide distribution. has a genome 297 nt in size.
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.: Evidence that viroids are widespread in grapevines. Rezaian et al.CEVd-g. Sano et al. Although CEVd is present in most. yellow speckle and fleck through dodder. besides Spain. Mink and Parsons: Yellow speckle can be detected by growing vines for 2-3 weeks at 32 °C under continuous illumination. Cannonau not associated with the presence of GFLV reported from Italy. Woodham and Krake: Artificial transmission of grapevine leafroll. Transmission: No vector is known. as the disease may have spread naturally. This latter way of dissemination has been considered as more efficient and frequent than mechanical transmission. Three different viroids found in a number of accessions in a Californian varietal collection. Experimental transmission through dodder is possible. its grapevine strain so far has only been recorded from Australia and the USA. In the great majority of grapevine germplasm infection is latent.: Four viroids found in Australian grapevines.: Two new viroids. 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). a disease formerly thought to be caused by a chromogenic strain of GFLV. It was first recoverd in Spain from symptomless grapevines. CEVd-g and AGVd. 2. Prota et al. a member of the genus Pospiviroid.: A disease of cv. Garcia Arenal et al.: First recovery of a viroid from grapevines in Japan. Barlass et al. Rcatzitelli resembling yellow speckle reported from Bulgaria. Natural dissemination takes place by mechanical inoculation through surface-contaminated cutting tools during management operations. Detection: Some viroids can be transmitted mechanically to herbaceous hosts but this is not an efficient detection method. the authors consider the results as inconclusive.: Successful mechanical transmission of viroids to grapevines.: The Japanese grapevine viroid identified as a strain of hop stunt viroid. First identification of AGVd Koltunow and Rezaian: Identification and sequencing of grapevine yellow speckle viroid.: A vein banding condition of cv. Control: Use of viroid-free propagative material obtained by meristem tip culture. found in grapevine accessions from Europe and California. and distribution of infected propagating material. has a genome 369 nt in size. Semancik et al. grafting. hybrids and cultivars appear to be susceptible. Krake and Woodham: Evidence that the agent of yellow speckle is implicated in the etiology of vein banding. Flores et al. if not all citrus-growing countries. Seed transmission has been demostrated for GYSVd-1. Shikata et al. one of which identified as the agent of citrus exocortis. 1984 1985 1985 1985 1987 1987 1988 1988 1988 130 . Woodham and Krake: Evidence of field spread of yellow speckle. Abracheva et al.: Reconstruction of the secondary structure of CEVd-g Szychowski et al. Varietal susceptibility: All Vitis species.: Yellow speckle eliminated by in vitro apical culture. GYSVd-2. For yellow speckle.
Duran-Vila.M. Journal of Plant Pathology 85. Wang et al. J. Minafra et al. 127-130. American Journal of Enology and Viticulture 39. Flores R. Koltunow et al. First report of Australian grapevine viroid from the Mediterranean region. V.S. Semancik. A. Krake.: First report of AGVd in the Mediterranean. 1978.. References Abracheva P.P.: Evidence that two related viroids (GYSVd 1 and GYSVd 2) can cause yellow speckle disease independently. Regeneration of virus-free grapevines using in vitro apical culture. Grapevine yellow speckle viroid 2 (GYSVd 2) Australian grapevine viroid (AGVd).. Juarez and J. Citrus exocortis viroid grapevine strain (CEVd-g). Spain. The occurence is reported of HSVd. Molecular evidence that this viroid originated from recombination between five different viroids among which GYSVd-1 and GYSVd-2 1991a.. Rezaian et al. 1982..: Suggestion that the viroid causing stunting in hop (HSVd) originated from grapevines. Greece. 2095-2102. R.b Szychowski et al. based on phylogenetical analysis of hop and grapevine isolates of this viroid. which can readily be isolated directly from grapevines. are to be named as follows: Hop stunt viroid grapevine strain (HSVd-g). Martelli: Brief review of grapevine viroid situation supporting the idea that vein banding is primarily induced by viroidal rather than GFLV infection. Elleuch A. which require amplification in an alternate host.. 45-57.R. Marrakchi.: Extensive comparative analysis of grapevine accessions from California and Europe reveal a similar pattern of viroid distribution. A possible virus disease of Rcatzitelli vines in Bulgaria. Journal of General Virology 66. Woodham and L. Perreault and H. GYSVd-1 and GYSVd-2 Rezaian: Complete nucleotide sequencing of AGVd.: Improvement of meristem tip culture technique for the production of viroid-free grapevines. Flores et al. Little and Rezaian: Updated review of grapevine viroids. 1985. GYSVd-2. Duran-Vila N. These viroids.G. Fakhfakh. Monografias INIA 18. Cordoba. Annals of Applied Biology 101. Martelli.C.: Review of viroids. CEVd-g and AGVd via grape seeds. Wan and Symons: Transmission of GYSVd-1.: First record of grapevine viroids in China. 2003. Quacquarelli and V. K. 1991 Semancik and Szychowski: There are two classes of grapevine viroids: (i) apparent viroids. Koltunow and Rezaian: Description and sequencing of grapevine viroid 1B (later renamed Grapevine yellow speckle viroid 2). Production of viroid-free grapevines by shoot tip culture. N. Proceedings of the 6th Meeting of ICVG.: Structural analysis reveals that five distinct viroids infect commercial grapevine varieties. 1976. according to an international agreement reached during the 10th Meeting of ICVG held in 1990 at Volos.M. Skene. Pallas and J.1988 1989 1989 1989 1990 1990 Duran-Vila et al. 291-295. Barlass M. Grapevine yellow speckle viroid 1 (GYSVd 1). 1988. J. Arregui. Elleuch et al. Savino. G. (ii) enhanced viroids.P. 1991 1996 1997 1999 2001 2003 2003 3.: A survey of viroids of grapevine in Italy. 217-220. Sano et al. 131 . Detection of viroid and viroid-like RNAs from grapevine. M.
Ohno and Y. Mechanical transmission and rootstock reservoirs as factors in the widespread distribution of viroids in grapevines. P. 40-50. F.R. Australian grapevine viroid . Grapevine viroids.A. Garnier. 1988. Grapevine yellow speckle agent implicated in the aetiology of vein banding disease. 575-578. 1975. Krake. Greece. Proceedings 10th Meeting of ICVG. Shikata E. Prota U. Goheen. R. Grapevine viroids. The field investigation of virus and viroid diseases of grapevine and stone fruit trees in Shandong and Liaoning province. J. S. Two related viroids cause grapevine yellow speckle disease independently.. 1991. Duran-Vila.A. and M. Krake L. T. Krake. 35-40. 413-422. Woodham R. and R. 1987. Semancik.A.. Martelli and V.P. R. Skene.R. 297. Little A. A. 193-198. Woodham R.Leclair. Proceedings of the Japanese Academy of Science 60(B). CSIRO Publishing.P.R. Wan C.A.G.a newly recognized grafttransmissible disease of Vitis. The sequence of a viroid from grapevine closely related to to severe isolates of citrus exocortis viroid. Vitis 29. Rezaian. J. 1989.A. 65-73.M. Hernadez. Widespread occurrence of viroid-like RNAs in grapevines. 1984. Volos. Ohshima.P. Koltunow A. Minafra. Collingwood. Doazan. Koltunow A. R. Hong. L. Volos. 1990. L. 333338.S. Meshi..A.C.M. 1996.C..W.S.). Semancik (eds. 213-216. Relationship and patterns of distribution among grapevine viroids from California and Europe. G. Vitis 21. China. Virus Genes 22. Greece. Phytopathologische Zeitschrift 106. A viroid-like RNA isolated from grapevine has high sequence homology with hop stunt viroid. Comparative properties of viroids of grapevine origin isolated from grapevines and alternate hosts. An infectious low molecular weight RNA was detected in grapevines by molecular hybridization with hop stunt viroid cDNA. 1989. and L.C. Nature.D. 287-288.M.. Taylor R. M. Szychowski J. A. 1990. Johnson and M. 2003. Cugusi and M. Rezaian. 1999.. 1988.L. Vitis 22. 849-864. Krake. Zhang. R. and R. N.C. Viroids of grapevines in Italy.evidence for extensive recombination between viroids. J. and M.P. Proceedings of the 10th Meeting of ICVG. Goheen and J.P. and J. Wolpert and J. Virology 170. Koltunow A. Woodham. 869-872. 2001. Credi.A. Phytopathologia Mediterranea 24. Rezaian M.A. M. 1990. Pallas and R. 370 pp... Proceedings of the 10th Meeting of ICVG. 1985.. Sano and I.Jiang. Plant Disease Reporter 59. P. Rezaian M.S. Z. 25-36. Symons. J. Duran-Vila. Journal of General Virology 70. Nucleic Acids Research 16. 1982. sanitation and situation in the Arab countries. Nucleic Acids Research 15. American Journal of Enology and Viticulture 39.Woodham. M. 195-206. S. 270-278.. Rezaian M. A. Randles and J. Doazan. Phylogenetic analysis of hop and grapevine isolates of hop stunt viroid supports a grapevine origin for hop stunt disease. 173-182. Savino.S. Hadidi.A. yellow speckle and fleck diseases by dodder.Flores R.M. Wolpert and J. Grapevine yellow speckle -. American Journal of Enology and Viticulture 38. 337-345. Rezaian. Dore. and R. 1990. Minafra A. Semancik.. Credi. 5587-5598. Relationships among grapevine viroids from sources maintained in California and Europe. Szychowski. V.C. J. Krake. Transmission of viroids via grape seeds. Volos. Vitis 30. a new member of the apple scar skin viroid group contains the left terminal region of tomato planta macho viroid. T. 1988.H. Rapid indexing procedures for detecting yellow speckle disease in grapevines. Journal of General Virology 66. Okado.M. Koltunow and L. Garau. detection. Uyeda.W. T. Australian Journal of Agricultural Research 23. and J.R. 1987. 1990. R..R. Grapevine yellow speckle disease: studies on natural spread observed in the field. Journal of Phytopathology 147. 24-28. 1989. and L. A. Garnier. I. 39-41. Semancik J. Szychowski J. Mimura and K. Sano T. Flores. and M. Mink G. Grapevine viroid 1B. Rivera-Bustamante and A.. Shikata. Martelli G. 1997. 1983.A. 1983. A. Zhang and X. In: Viroids (A. Isolation of three viroids and a circular RNA from grapevines. Semancik.C. Sano T. Nucleic Acids Research 10.A. 3411-3419. Minafra. Greece. China Fruits 4. Koltunow. N. 285-291 Wang G. 1991. Flores. Rezaian.S. Parsons.. 202-205. Szychowski J.. Investigations on transmission of grapevine leafroll. 53-59. R. Viroids: the non coding genomes. Arab Journal of Plant Protection 7. E. Journal of General Virology 69.R.H. Garcia Arenal F.A. 1972. Krake and K..I. Di Serio and C. Seminars in Virology 8. Leclair. Infectious diseases of grapevines. N. 132 .S.A. Uyeda. Semancik J. 1991b.. Investigations on a vein banding disease of Grapevine in Sardinia. 1991a. 4203-4210. 210-219. 447-452. 1985. Grapevine yellow speckle viroid: structural features of a new viroid group.
all organs of the plant may be infected. based mainly on sequence similarity of their rRNA genes and of a few other genes. Downwards rolling of the leaves and sectorial discolorations of the blades. Palatinate grapevine yellows. in addition to other criteria such as symptomatology. The occurrence and spread of a particular GY mainly depend on the presence of efficient vectors in the vineyards or their close vicinity. Phytoplasmas form a homogeneous phylogenetic clade subdivided into about 20 groups and subgroups. 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. Golden flavescence. such as the ribosomal protein genes or the elongation factor Tu. but not seeds. Legno nero. Characteristic symptoms in the end of summer are a partial or total lack of lignification which may affect individual canes and shoots or the whole plant. involving also the main veins. their movement is slow and their distribution in the plant is erratic. Quite often. Candidatus Phytoplasma australasia. Main symptoms: Young shoots of affected Vitis vinifera are weak with short internodes and frequent necrosis of terminal buds. it was mainly in the 1990's that GYs could be differentiated. canes. Phytoplasmas in the 16SrIII group (or X-disease group) are associated with grapevine yellows in North America and Israel. Phytoplasma cells are vesicular rounded bodies. climate and infection pressure. Flavescence dorée (FD) and Palatinate Grapevine Yellows (PGY) phytoplasmas belong to group 16SrV (or Elm yellows group). They are wall-less. after their aetiology had been understood and molecular methods for the characterization of associated phytoplasmas had been developed. 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. 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. The different GYs cannot be differentiated on the basis of symptoms. Australian grapevine yellows. Vergilbungskrankheit. Phytoplasmas have the smallest genome reported for procaryotic organisms (560 2. In 1997. However. and transmission by specific leafhopper or planthopper vectors.200 kbp). However. However. Nonetheless. buds. depending on the particular disease and other conditions such as variety. their genome is poorly known because they cannot be cultivated. phloem-restricted bacteria that belong to the class Mollicutes. resulting in reduction of quantity and quality of the crop. Main diseases: Flavescence dorée. Their titre is usually very low in grapevine. inflorescence and berries. Rubbery canes fall downwards with a typical "weeping" aspect. persistence of infection in dormant plant material. Bunches wither in early summer or berries shrivel later in season. a severe decline of the vine. Schwarzholzkrankheit. Leaf blades are crispy and brittle. host range. The main characteristics of GY diseases are important losses or damage to the yield. vector species have not been identified for all GY diseases. including roots. autumn fall of leaves occurs later on diseased than on healthy vines. Bois noir. Red-berried varieties show reddish to purple discolorations while white-berried varieties show golden to chlorotic discolorations. Several Candidatus phytoplasma species have been recently described. associated to Bois noir (BN). Severely infected stocks decline rapidly.GRAPEVINE YELLOWS: GENERAL PROPERTIES 1. Main synonyms: Flavescence dorée-like disease. Flavescenza dorata. Amarilliamento. 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). Vergilbungskrankheit. develop on leaves. and serology. Stolbur phytoplasmas. The first GY to be recorded was Flavescence dorée in France in the 1950's. though some sieve tubes may appear crowded with phytoplasma cells in the electron microscope. the term phytoplasma has become the genus name of these plant pathogenic agents under the provisional taxonomic status Candidatus. shoots. Nevertheless. Flavescenza dorata. a phytoplasma belonging to group 16SrII (or peanut witches' 135 . North American grapevine yellows.
When no information on the particular phytoplasma type is available. Varietal susceptibility and sensitivity: Numerous V. The best antigen source are leaf veins and petioles. influence symptom expression and differential sensitivity of cultivars. A range of primers that can be used for amplification with Polymerase Chain Reaction (PCR) of characteristic DNA fragments of phytoplasmas. are available in the literature. These phenomena also depend on the disease complex (phytoplasma. Most of these PCR primers have been designed on conserved regions of the rRNA gene and are "universal" for all known phytoplasmas. Random Fragment Length Polymorphism (RFLP) analysis of the amplicon can provide further characterization of the infecting phytoplasma. Transmission: Phytoplasmas are vectored by hemipters. as well as their feeding activity. canes. However. Pinot noir. They can also be observed in sieve tubes by light microscopy. or on random selected non-ribosomal DNA fragments. Polyclonal and monoclonal antisera have been raised to a few phytoplasmas. Hence. including the agents of FD and BN. are very sensitive to all GY diseases. 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. The three ascertained vectors of GY diseases are univoltine leafhopper or planthopper species. when grapevine is not a preferred host for the vector. Detection can also be made by transmission electron microscopy of thin sections or by scanning electron microscopy. more specific primers can be used. such type of transport is risky when potential insect vector species occur on the site of planting. Full recovery and transient remission of symptoms have been observed in low and medium sensitive cultivars. Infection of rootstocks has been detected only in the case of Flavescence dorée but it cannot be excluded for other GY agents. vinifera cultivars have been reported to be sensitive or very sensitive to GYs in all countries where infection occurs. Simultaneous presence of symptoms on leaves.broom group) is a second agent of Australian grapevine yellows. ELISA detection is possible from vascular tissue of symptomatic grapevines. Vitis riparia and American rootstock varieties and hybrids do not show typical symptoms of GY. on less conserved genes. Phytoplasmas in the 16SrI group (or aster yellows group) are associated with endemic GY diseases in several countries of Europe and in the USA. Detection: GY syndrome is characteristic. Then. Other host plants: Most phytoplasmas are ubiquitous plant parasites that can be hosted by several sometimes tolerant plant species. Double infections have been reported. Some cultivars such as Chardonnay. shoots. which allows restoring the sanitary status of affected vineyards when the vector activity is controlled. are critical. mainly hoppers (cicadellids or cixiids) or psyllids. It is probable that the feeding preference of insect vectors. Cabernet Sauvignon. vector and abundance of reservoirs). but the latter technique has never been successfull on field-grown 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. PCR amplification with universal primers is preferred. Outstanding progress in detection was achieved with DNA-based techniques. These methods are not specific for any phytoplasma and are too laborious for disease monitoring. Individual symptoms can be confused with other diseases or disorders. When the presence of a particular disease is suspected. Though the rate of transmission to plant material can be very low. Riesling and also numerous local varieties. designed on variable regions of the rDNA. using DAPI staining. are group specific. Propagation material may be the infection source for long distance dissemination of GY diseases. Phytoplasmas can be detected by graft transmission to sensitive vines. Phytoplasmas also persist in vine stocks during winter. Others. The extraction from infected tissues of total DNA containing enough phytoplasma DNA of good quality and the elimination of inhibitors of the PCR reaction. 136 . The situation of cv Syrah is controversial. Particular procedures for extraction of phytoplasma antigens from infected vines must be used to enhance the sensitivity of detection. erratic transmission to grapevine may nevertheless take place.
an insect that has been introduced recently from America..Other molecular methods are being developed. even sensitive detection methods may not be fit for use in sanitary selection because of the uneven distribution of phytoplasmas in mother plants and their vegetative progeny. under the name Flavescence dorée. Control of GY diseases depends on the knowledge of vector insect species. BN is considered as a non epidemic form of FD. efforts should address prevention by planting healthy propagation material and limitation of vector populations and their inoculative activity. soaking of dormant material before or after grafting. Schvester et al. Caudwell: Characterization of a new type of flavescence. Control: There are no direct curing methods of infected plants. weeds or other crops. hypotheses on its nature. The biology of the insect is described. littoralis Ball. Nonsymptomatic grapevines that show irregular symptoms from one year to the next may also test positive. FD can be detected from leaves of non-symptomatic American roostocks. However. Investigations are being conducted on sensitivity. The disease can be transmitted by grafting and is probably a virus disease. The author suggests this disease to be due to adverse climatic and soil conditions. In any case. and natural enemies. FD is transmitted by the leafhopper S. The disease is thought to be caused by root damage. Detection in the upper parts of the vines is usually done from veins and petioles of the leaves of symptomatic shoots. Gärtel: Description. on physiological changes and defence mechanisms induced by infection. cultural practices. Generally. but does not rule out the possible involvement of a pathogen. 1956 1957 1961 1961 1961 1964 1965 137 . Caudwell (b): Description of recovery in grapevines affected with FD. i. Real-time PCR has also been successfully used.e. 2. Vidano: S. evolution of the disease in space and time. denoted Flavescence dorée (FD) in agreement with the name proposed by Levadoux (1955) because of the golden yellow metallic aspect of leaves. but is also possible using young leaves shortly before symptom expression or phloem scrapings from lignified canes. Historical review 1955 Levadoux: Description of a severe outbreak of "Flavescence dorée" on Baco 22A in southwestern France. into hot water (50 °C) for a sufficient length of time (minimum 30 mn). Symptoms (macroscopic and microscopic). Control methods of vector populations are being experimented with natural insecticides. littoralis Ball found in Italy in 1963. tolerance and potential for recovery of varieties. and on the identification of reservoirs of inoculum such as infected grapevines. Pruning or top-grafting are useless because roots and trunks are infected. phytoplasmas are unevenly distributed in GY-affected vines. and on elicitors of defence reactions to these phloem-restricted bacterial agents. Detection of phytoplasma DNA can be achieved from all plant organs or insect vectors. Caudwell (a): Studies on Bois noir (BN) and on its relationships with FD. Production of sanitized propagation material is possible with hot water treatment (HWT). using detection of PCR amplification products with DNA-DNA hybridization with a specific probe. on the possibility to limit or eradicate their population and prevent their migration or movement in vineyards. of a disease occurring in the vineyards of the Mosel Valley and of the Rhine Valley. Branas (a and b): Description of flavescence epidemics on Baco 22A and Chardonnay in France.
faba plants produced typical GY symptoms.: Production of antisera to FD-MLO raised to extracts of experimentally FDinfected plants (Vicia faba) and leafhoppers (Euscelidius variegatus) and first microscope observation of MLOs trapped with ISEM. This phytoplasma was later on identified as a Clover phyllody phytoplasma. Caudwell and Larrue: Spread of FD in France is related to sanitary status of planting material. Belli et al. titanus fed on the latter V. Belli et al. titanus found in 1984 in vineyards of northern Italy. Rumbos et al. Caudwell: Discussion on the origins of yellows diseases of plants. (b): Evidence that FD and BN are two different diseases with similar symptoms. Rafaila and Costache: Yellows of grapevine with symptoms similar to those of FD found in 1967 for the first time in Romania. Caudwell et al. Bois noir. littoralis is present in the same region of Oltrepò Pavese. and Euscelis sp. As similar organisms have been found in nematodes of the species Xiphinema index.: S. Caudwell et al. Caudwell et al. littoralis Ball is present in Ticino (southern Switzerland).1966 Caudwell: Attempts to inhibit the agent of FD in vivo by heat treatment. 1967 1968 1970 1971 1971 1973 1973 1975 1977 1979 1980 1982 1982 1983 1984 1985 1985 138 .: Rickettsia-like organisms observed in roots of grapevines with "Vergilbungskrankheit" in the Saar. Conversely. Caudwell: Symptoms of grapevine yellows (Amarilliamento) reported in Chile on cv Elqui. is probably of European origin. First record in 197576. Recommendation that mother plants should be grown in areas far away from FD-infected regions. The disease has been observed for the first time in 1968. Granata: Description of an epidemic yellows disease on cv Inzolia in Sicily. (a): Mycoplasma-like organisms (MLOs) are observed in diseased grapevines and infective leafhoppers and in Vicia faba submitted to feeding inoculation with infective S. witches' broom and yellowing. Osler et al. So far. Symptoms on V. Magarey and Wachtel: Description of a new disease of grapevine of the yellows type on the cv Rhine Riesling in South Australia. Credi and Babini: Grapevine yellows are reported in Emilia Romagna (Italy).: S. An important FD-like disease is reported. with special reference to grapevine. Provisory name "Rhine riesling problem". Doi et al.: Presence of a disease similar to FD in the Oltrepò Pavese (northern Italy). these findings have not been confirmed. Potential existence of several different diseases: leafhoppers (Euscelidius sp. Immersing cuttings taken on FD-affected grapevines in water at 30°C prior to planting reduced the proportion of infected plants by about 83%. of which S. Baggiolini: S. mainly on cv. faba are different from those obtained with feeding inoculation with infective S. Tanne and Nitzany: Occurrence of a yellows disease resembling FD in Israel. Mosel and Rhine regions are considered as the causal pathogens of this disease. Regina. littoralis. littoralis.) caught in the vicinity of vineyards could transmit a yellows disease to Vicia faba plants. the hypothesis is put forward that this nematode is the vector of the disease. The author assumes that both FD and its vector were introduced from North America with varieties of Vitis labrusca between 1927 and 1950. Inoculation of grapevine seedlings with S. titanus (= littoralis) is not a vector.: Discovery of Mycoplasma or PLT group-like microorganisms in the phloem elements of plants showing dwarfing.
Conti: A review of intracellular procaryotic phytopathogenic agents. S.: Occurrence of FD-like symptoms in the valley of Sarca in Trentino. However. of MLOs in phloem tissues of grapevines affected by "FD" in northern Italy.: Contribution to the knowledge of FD (or similar diseases) in the Veneto region.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.: Survey of potential Auchenorryncha vectors of the pathogen agent of FD in Piemonte (Italy). titanus is not always associated to the disease. for the presence of a FD-like disease. vector of FD. titanus.: Observation of MLOs in phloem tissue of yellows-affected grapevines in Australia. Survey for the presence of S. Fortusini and Belli: Description of the development of epidemics of FD-like diseases in northern Itay.: Observations. control measures. Italy.: Description in Italy of the presence of FD or FD-like diseases. Credi: Description of a FD-like disease in Emilia Romagna and the reaction of 3 different cultivars. Egger and Grasselli: Presence in Toscana. disease distribution. titanus. Hyalesthes obsoletus. Two of these vineyards were sprayed with insecticides in 1986 and 1987. respectively). In addition to S. whereas an unsprayed vineyard was more affected. Magarey et al. Mescalchin et al. Vidano et al. Pinot nero) and comparison with other similar diseases. Italy. Susceptibility and sensitivity of affected varieties (Chardonnay. resulting in a reduced diffusion of the disease. that are responsible for severe decline of vines. 1987 1987 1987 1988 1988 1988 1988 1988 1988 1989 1989 139 . Recovery and crop loss are variable. Chardonnay is the more affected variety. variegatus and S. northern Italy. Euscelidius variegatus and Euscelis incisus are taken into consideration.or xylem-limited prokaryotes in Europe. Fortusini et al. Quaroni et al. Italy. Borgo et al. The results suggest that the disease is brought into the vineyards from outside local sources.: Presence of a FD-like disease in Emilia-Romagna. in viticultural areas of northern Italy and of other Auchenorrhynchas susceptible to play a role in the transmission of yellows diseases in this region. titanus.: Study on the distribution of S. their etiology is unclear. Vidano et al.: Presence and distribution of a FD-like disease in the region Friuli-Venezia Giulia in Italy. Carraro et al. Martelli: A review on the knowledge on grapevine diseases induced by phloem. using the scanning electron microscope. titanus. Credi et al. Rui et al. vector of FD. Italy. Pinot bianco. Boudon-Padieu and Larrue: ELISA detection of FD pathogen on infected reared experimental and wild natural leafhopper vectors (E. Credi and Callegari: Survey of vineyards in Emilia-Romagna.: New data on the spread of FD in three vineyards of Oltrepò pavese (northern Italy) from 1985 to 1987. Borgo: General information on the presence of FD-type diseases in northern Italy. of a FD-like disease on Chardonnay. Seljak: Presence of Scaphoideus titanus in western Slovenia (Yugoslavia).
Di Terlizzi et al.: Occurrence of grapevine yellows in Moldavian SSR.: Presence of yellows-like symptoms in Apulian grapevines (central Italy). The annual rate of symptomatic plants was never greater than 20 % over a 6-year period. Treatment prior to storage is more efficient. Deng and Hiruki: A method to amplify 16S rRNA gene from culturable and nonculturable mollicutes. 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). Several weeds with symptoms of phyllody were found to contain MLO in electron microscope studies.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. Credi and Santucci: Development of "Flavescenza dorata" in grapevine in Emilia-Romagna and rate of infestation in different systems of cultivation. Caudwell: Review on the epidemiology and characterization of FD and other GY diseases. Conti: A yellows-type disease of cv. Chardonnay and Baco 22A grafted on donor yellows-diseased plants of cvs Pinot blanc and Sangiovese. 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". Symptoms also developed on plants obtained from buds taken on diseased Chardonnay and Sangiovese grafted on healthy Kober 5BB. Granata and Russo: An epidemic disease with FD-type symptoms develops in Sicily on cv. Similar symptoms on Chardonnay could be obtained by insect inoculation (Euscelis incisus) and by cleft-grafting of tissues taken from yellows-infected elm. Natural infection with MLOs of bait plants (Catharanthus roseus and Vicia faba) placed in vineyards in spring and summer. Caudwell and Kuszala: Detection with ELISA of FD-MLO in naturally FD-affected grapevines. (a and b): Identification of numerous species of Auchenorrhynchas in the vineyard ecosystem in Piemonte (northen Italy) and of ampelophilous species. Vidano et al. 1989 1990 1990 1990 1990 1991 1991 1991 1991 1991 1991 1991 1992 1992 1992 140 . Boidron and Grenan: Construction and testing in ENTAV. 45mn). titanus was present in all vineyards checked. Only part of the plants were infected. Granata and Grimaldi: MLOs are observed with the electron microscope on affected grapevines of cv.: Bench grafting experiments of buds of indicator cvs. S. Inzolia. Inzolia in Sicily. suggesting an unrelated agent. Affected grapevines in the Rhône valley tested negative. This is a prospect for investigation on MLOs associated to plant yellows. The recommended temperature / time combination is 50°C / 3560 min. Chardonnay develops in Tuscany. Marinesku et al. Credi et al. Le Grau du Roi (France) of a device for secure soaking of dormant buds and canes into hot water (50°C. titanus was not found in the area.: 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.: Grapevine graftwood shoots infected with FD can be disinfected by hot water treatment in the dormant stage. Restriction profiles after digestion with endonucleases permit the differenciation of MLOs associated to different plant diseases. Caudwell et al. Specific primers can be used for MLOs. Refatti et al. The aetiology is not fully ascertained though the presence of MLO is probable.
No spontaneous recovery. (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. Attempts to characterize MLO DNA extracted from insects with molecular methods. titanus population in vineyards and the rate of spread of the disease. The titre of MLO appears very low in grapevine. However. suggesting the presence of two different diseases at least in Veneto and Friuli-Venezia Giulia. Osler et al. detection (ELISA. Typical symptoms on several cvs. results of research. Daire et al. Arnò et al. FD-MLO DNA can be distinguished from other MLO DNA with Dot-blot hybridization and Southern-blotting. MLOs were visible in sections of petioles of periwinkle showing symptoms and subsequent graft transmission from periwinkle to periwinkle was successful. Davis et al. FD can be readily distinguished from Bois noir. Evolution of the disease and of its vector in the various viticultural regions of France.: 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. Arzone et al. The MLO strains found in grapevine are related with aster yellows MLOs. S. titanus. Caudwell et al. Senescent or degraded forms of MLOs identified inside degenerate sieve elements.: Transmission experiments from grapevine to grapevine and to herbaceous hosts with S. titanus leafhoppers. titanus and other leafhopper species show that yellows disease present in northeastern Italy differs from flavescence dorée as it occurs in France. genomic tests). Boubals (b): A severe epidemic of grapevine yellows occurs in Golan (Israel). One MLO strain transmitted with Macrosteles quadripunctulatus appears similar or close to aster yellows MLO.1992 Credi and Santucci: Of 628 attempts to transmit the agent of a yellows disease of grapevine from Sangiovese. It is more related to the agent of Elm yellows than to other yellows agents.: 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). Quacquarelli and Barba: Review of the distribution of flavescence dorée and other grapevine yellows in EEC viticultural countries. Meignoz et al. but are different from known aster yellows cluster MLO strains.: ELISA with FD-antibodies permit to detect related antigens in S.: Construction of serological and molecular detection tools to MLO transmitted to periwinkle with dodder from a yellows-diseased grapevine in Friuli-Venezia Giulia (Italy) (later 1992 1992 1992 1992 1992 1993 1993 1993 1993 1993 1993 1993 1993 1993 141 .: Description of MLOs and associated disorders in the phloem of FD-affected LN33 experimentally inoculated with S.: FD can be transmitted by symptomless rootstocks. Chen et al. transmission by feeding to healthy grapevines succeeded only in 1 attempt out of 100. (b): Evidence of an association of MLOs with grapevine yellows in Emilia Romagna. Bertaccini et al. Hot water treatments suppress the transmission to Chardonnay indicators. IPVR is related to aster yellows MLO. titanus is not present. (a and b): Observations supporting the existence of different types of grapevine yellows in northern Italy. Caveccia and Chardonnay vines to periwinkle by the dodder species Cuscuta campestris. Boubals (a): Report on a meeting of the French working group on FD. (a and b): Characterization of Italian periwinkle virescence (IPVR) MLO (obtained on bait periwinkle plants exposed in yellows-affected vineyards in Italy) is possible with DNA-DNA hybridization and PCR. Bianco et al.6%) were obtained. Alma et al. 4 positive results (0.
1993 1993 1993 1993 1993 1993 1994 1994 1994 1994 1994 142 . titanus) and a second one. Computer analysis for spatial patterns of diffusion show a non-random distribution and suggest transmission from other vines or from weeds. FD is related to elm yellows and BN is related to stolbur. Carraro et al.: Diversity of MLOs associated with grapevine yellows and transmitted to periwinkle. FD belongs to the elm yellows group but is different from elm phytoplasmas. Maixner (a. titanus is present in the Friuli-Venezia Giulia region but no transmission assay has been successful. Detection in non symptomatic V. c and d): Grapevine yellows (Vergilbungskrankheit) are common in the Moselle and Rhine valley. Only FD and BN have been identified in naturally affected grapevines. So. Credi (a and b): Observation of MLO in yellows-affected grapevines in northern 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. Maixner: Demonstration that Hyalesthes obsoletus (Auchenorrhyncha: Cixiidae) is a vector of German grapevine yellows (Vergilbungskrankheit). probably transmitted by another insect. In northeastern Italy. S. riparia in New York and failure of detection in some symptomatic grapevines in New-York and Italy. Detection in grapevines from Virginia (USA) of MLO related to X-disease. titanus to test plants of cvs Perera and Chardonnay in Veneto and Friuli-Venezia Giulia (Italy). 1993 Daire et al. Transmission has been obtained only from vines of Veneto. A MLO has been transmitted to periwinkle with dodder. Osler et al. 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). Experiments on the use of hot water treatment on planting material. on the effects of pollarding affected vines and of chemical sprays against vectors. aster yellows and Xdisease. Electron microscopic observation of MLO in periwinkle and grapevine. GYU was later on characterized by other authors as a phytoplasma belonging to the Xdisease (16SrIII) group. S. BN (stolbur MLO) detected in several regions of Italy and in Israel. Specific probes cloned and specific primers designed for PCR amplification of phytoplasma DNA. using insects. healthy plants protected with plastic screens did not develop symptoms and diseased plants protected with screens showed transient recovery. FD detected in leaves of rootstock 3309C and is also present in samples from Veneto (Italy). Positive assays obtained only on samples from France and Veneto. MLO appear to be in very low titre. titanus is not present. Daire: PhD thesis on detection and differentiation with DNA-based methods of the MLO agents of GY diseases in France. at least two different phytoplasmas are infecting grapevines in Veneto: FD sensu stricto (transmitted by S. BN belongs to the stolbur group. Description of pathological changes in phloem of leaves and identification of senescent forms of MLO inside degenerate sieve elements. Di Terlizzi et al. 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.: Important outbreak of a yellows disease on local varieties in Apulia (southeastern Italy) and infection of periwinkle as bait plants and with dodder transmission. Girolami and Egger: Importance of contamination of vineyards and diffusion of Grapevine yellows in northern Italy. Prince et al.: FD-ELISA method was used on tissues of yellows-affected grapevines from diverse countries. graft and dodder. Classification of grapevine MLOs into three RFLP groups: elm yellows (FD from France).called FDU. b. Kuszala et al. International Committee of Systematic Bacteriology (ICSB). then GYU phytoplasma) and comparison of specificity and sensitivity of the latter tools to detect phytoplasmas in grapevines in New York and Italy. (a and b): Various attempts to transmit FD-like disease in different regions of northern Italy.: Six-year transmission trials of different types of yellows with S.
: Survey of yellows symptoms in an ampelographic collection of 1281 cultivars in Conegliano (Veneto. in the vector H. aster yellows. Arzone et al. stolbur and apple proliferation) can be detected. Del Serrone et al. Laviña et al. (a): Specific detection with PCR of stolbur phytoplasma in grapevines affected with Vergilbungskrankheit. Detection is also possible during winter on wood scrapings. Alma et al. Consideration on possible resistance and research on interaction of phytoplasma with nepovirus infection. in grapevines with yellows symptoms in Soave (Veneto. Maixner et al (b): Detection of a new phytoplasma type related to FD in cv Scheurebe showing symptoms of yellows in Palatinate (Germany). titanus which has not been found. Albanese et al. Double infection is reported. Koruza: Dispersal of grapevine yellows in Slovenia. Italy).: The phytoplasma associated with Australian grapevine yellows is close to but different from stolbur phytoplasma causing BN. Molecular detection of aster yellowsrelated phytoplasma.: Six-year survey of evolution of yellows symptoms in a 6-12 year-old vineyard of cv Chardonnay that was never sprayed with insecticides. The most frequent belonged to group 16SrI-G (later designated as 16SrXII or stolbur group). PCR detection of phytoplasma with universal primers. probably of Bois noir type.: MLOs associated to grapevine yellows in Virginia belong to 2 different groups : X-disease group (16SrIII) and aster yellows group (16SrI). sometimes in mixed infection. (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. Haidar: First report of symptoms of yellows on grapevines in Lebanon.: Identification of BN (stolbur phytoplasma) in Spain. Italy).: Presence and distribution of GY in Sicily. obsoletus and in weeds growing in the vineyard in Germany. The importance of proper identification of disease type for control measures is emphasized. Del Serrone and Barba (a and b): Monitoring of detection of phytoplasma in the organs of grapevine according to the vegetative state. Related MLOs were also detected in wild grapevines. Prince et al. with the aim of identifying sources of tolerance or resistance to yellows diseases. One grapevine out of 16 contained a 16SrV (= elm yellows) phytoplasma together with a 16SrI-G phytoplasma.: Grapevines from Piemonte (northern Italy) contained several different phytoplasmas. Egger et al. (a and b): Grapevine yellows in Latium (southwestern Italy) are not associated with S. Maixner et al. Padovan et al.: Presence of grapevine yellows in northern Portugal and electron microscopic visualization of MLOs in the phloem tissue of petioles of affected plants.: Four different phytoplasmas (FD. Boudon-Padieu (a and b): Reviews on knowledge and research on grapevine yellows. 1995 1995 1995 1995 1995 1995 1995 1995 1995 1996 1996 1996 1996 1996 1996 1996 1996 143 .: Emphasis on the possible role of weeds as reservoir for MLO in vineyards.1994 1994 Parente et al. Padovan et al. Fortusini et al. Ten weeds were found harboring MLOs with transmission electron microscopy. Murari et al.
10 are confirmed phytoplasma vectors. Borgo: Description and colour photographs of symptoms of grapevine yellows and knowledge on their aetiology and vectors. Kölber et al.: Recurrent and severe yellows symptoms observed since the 1980's in Sardinia (Itlay) on several cultivars. Maixner et al. Del Serrone : A review in Italian on the knowledge on etiology. Boudon-Padieu and Maixner: Updating in French and English of present knowledge on grapevine yellows: etiology. Aldini et al.: Use of Hot water treatment (HWT) to eliminate phytoplasmas (FD and AY) from 3 cultivars of grapevine. (b): Survey of possible auxiliaries for the biological control of leafhoppers and planthoppers in vineyards.1996 1997 1997 1997 Rüdel: Review on the history and diversity of Vergilbungskrankheit of grapevine in Germany. spread. 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. transmission. No epidemic outbreak has been observed. Maixner et al. S. northern Italy. Aster yellows and X-disease types were never detected.: History and control of GY diseases in Italy. Subcommittee on the Taxonomy of Mollicutes: The taxonomy of phytoplasmas will refer to their molecular phylogeny. Tanne and Orenstein (a and b): Successful heterografting transmission of phytoplasma from symptomatic grapevine to periwinkle and easy subsequent identification of phytoplasma type. Phytoplasma belonging to the stolbur subgroup have been identified. titanus is not reported. Daire et al. (b): Prospects on detection and identification of grapevine phytoplasmas with new molecular tools and consequence on future epidemiology studies.: Survey of hoppers in vineyards of the province of Piacenza (Emilia Romagna.: Identification of Flavescence dorée in Spain. (a): Only stolbur and FD phytoplasmas were detected in samples of yellows-affected grapevines from numerous regions of France and from Italy. Positive reaction with a DNA probe specific to aster yellows phytoplasmas. Davis et al. No double infection has been found to occur.: Symptoms of grapevine yellows observed in Hungary in the early 1970's. Batlle et al. 10 species were known vectors of phytoplasmas. Murari et al. (a): Description of available methods and strategies for detection of phytoplasmas in grapevine. No positive detection with PCR on the HW-treated vegetative progeny of mature canes of symptomatic plants.: Survey of leahoppers in vineyards of Piemonte. titanus reported for the first time in this region. International Committee of Systematic Bacteriology (ICSB). Bosco et al. Among 29 species of Auchenorrhynchas from the vine canopy and weeds of the border. Severe symptoms with high incidence on seven cultivars are reported from four grapevine-growing regions surveyed for 5 years (1993-1996). titanus is not present. AY and X-disease related phytoplasmas have been transmitted and detected. Belli et al. The term phytoplasma will become the genus name of plant pathogenic mycoplasmas under the provisional taxonomic status Candidatus. The possibilities of control 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1998 1998 1998 144 . biology and control. vectors and detection of the agents of Grapevine yellows. Presence of S. Garau et al. Subclades are considered to represent the equivalent of distinct species. Spain and Israel. S. Among 32 species identified. Italy).
Reinert W. Sensitivity of assays must be good because of the low titre of phytoplasmas in affected plants. Frausin et al. epidemiology and etiology of GYs in the world. with a particular reference to the work done in Germany. epidemiology of these diseases. has been identified as a member of the X-disease group. and M. Borgo et al. titanus. History.: First report of the use of oligonucleotides for in situ hybridization with transmission electron microscopy to localize phytoplasma in plant cells. GY affect mainly the northern provinces. Lherminier et al. Boudon-Padieu (b): Chapter in a Handbook on viral and bacterial diseases of the grapevine. symptoms. Guadagnini et al. vector of FD.: A comprehensive review of the classification of phytoplasmas in the world. according to the most recent studies on molecular structure of rDNA. obsoletus is rarely found in vineyards. BN is very frequent but H.: Summary of the complex situation of GY in northeastern Italy. though S. 1998 Lee et al. No negative effect was observed on scion buds but rootstock cuttings were more sensitive to negative effects of the treatment. In the Veneto region only stolbur-group phytoplasma was recorded in some areas and both stolbur and elm yellows group phytoplasmas in other areas. titanus is more widely distributed. In addition a phytoplasma (FDU=GYU) transmitted from GYaffected vines to Catharanthus roseus by means of dodder in Friuli-Venezia Giulia. Probes were antisense to stolbur specific sequences in the 16S rDNA and used on stolbur-infected periwinkle. On the basis of RFLP of PCR-amplified 16S rRNA gene. Insecticide sprays against S. distribution in the world. Batlle et al. Carraro et al.: Report of positive detection of aster-yellows phytoplasma in grapevine fed with Metcalfa pruinosa and in the body of the insects used for transmission. Search for alternative vectors.: Monitoring grapevine yellows in North-eastern Italy and attempts to rationalize routine laboratory testing for the differentiation between FD and stolbur phytoplasmas. phytoplasmas can be classified into 14 groups and 38 subgroups.: A review of the situation of GY in Spain. are compulsory in France. Maixner and Reinert: Updating in German of the research made in different countries on grapevine yellows diseases.: A history of GYs in north-eastern Italy. titanus.: Phylogenetic studies of phytoplasmas associated to numerous plant diseases.depend on the knowledge of the biology of phytopathogenic agents and of their vectors. Survey with PCR of phytoplasmas infecting grapevines in Friuli-Venezia Giulia and Trento province (Italy) and in Slovenia. nucleic acid hybridization and serological comparisons. FD is restricted to the north of Cataloña. Grapevine phytoplasmas classify into different groups. Boudon-Padieu: Review on recent progress in the knowledge on grapevine yellows. Seemüller et al. Firrao et al. Refatti et al. Only FD and BN have been identified.: Unsatisfactory results on the use of hot water treatments to eliminate phytoplasmas from grapevine wood. FD-related phytoplasmas (elm yellows group) have been detected in grapevines from Treviso and Verona and transmitted from vine to vine using S. Similar attemp with FD phytoplasma were not successful. methods of detection of phytoplasmas in grapevine tissues and in insect vectors. Maixner: A review of thermotherapy to cure phytoplasma infected material. have shown that only stolbur group phytoplasma was consistently found. Phytoplasmas for which 16S rDNA sequence is available have been classified into 20 major groups and subgroups. 1998 1998 1998 1998 1999 1999 1999 1999 2000 2000 2000 2000 2000 145 .: Evaluation on the efficiency and security of hot water treatment used to eradicate phytoplasmas from propagation material.
Waite et al.: A 2-year survey in Golan Heights.. Plant variety and cutting diameter influence the rate of surviving cuttings. Primers fitted for specific detection of these 3 phytoplasmas permit cheaper and quicker diagnosis. nitrate and nitrite reductase.: Report of experiments combining temperature and time of treatment with hot water on 5 grapevine cultivars to cure planting material from phytoplasmas. Klein et al.: Development of a new technique with DNA chips to detect phytoplasmas in grapevine. Influence of rootstock on the rate of GY symptoms in the same vineyard and same grapevine variety. titanus is widespread in western vineyards. Boudon-Padieu: Updating of knowledge and research on grapevine yellows worldwide. Israel. Zahavi et al. Frosini et al. Macrosteles quadripunctulatus and Orosius orientalis were found on weeds in vineyards or trapped on sticky traps. The latter was difficult to evaluate because of the non homogenous distribution of phytoplasmas in canes. Orenstein et al. FD has not been identified. Moretti et al. Circulifer sp. Tanne et al. Experimental evaluation of the method and application to field-trapped individuals of suspected vector species. Seljak and Petrovic: Overview on phytoplasma diseases of grapevine and fruit trees in Slovenia.: Survey of GY in Israel. aster yellows (11 %) and X-disease groups phytoplasmas. Clair et al. Quartau et al. 2000 2001 2001 2001 2001 2001 2001 2001 2001 2001 2001 2002 2002 2002 2002 2002 2003 146 . Osler and Refatti: The situation of GYs in northern Italy. which are all known as vector of phytoplasmas.: Use of immersion in hot water (HWT) or in chemicals to suppress phytoplasmas from grapevine propagation material in Italy. BN and aster yelllows in vineyards of southeastern Piemonte (Italy).: The presence of phytoplasmas in liquid medium on which insect vectors have been fed can account for potential vectorship of the species. confirmed the presence of three phytoplasmas associated to GY: stolbur (70%). suggesting that they did not multiply in the body of the insects. Crocker et al. (a): Presence of FD. HWT (50°C/40 mn) gave acceptable survival but none of the conditions experimented provided a total elimination of phytoplasmas. Hyalesthes obsoletus has been found but not in vineyards.: In Golan Heights (Israel) Hyalesthes obsoletus.: Scaphoideus titanus is present in Portugal.: 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.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.: Use of hot water treatment in commercial nurseries in Australia to eradicate diverse pests and pathogen agents from propagation material. Dual infection may occur (13 %). Neoaliturus sp. Marzachi et al. The phytoplasma agents are not specified. although its vector S. Seljak: A review of non-European hoppers introduced in Slovenia.: 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. Phytoplasmas were detected in all five species. Bertaccini et al. sugar metabolism. 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.
Tassart-Subirats et al.: Grapevines affected with yellows are found free of phytoplasmas after recovery.: Important outbreak of grapevine yellows on cv Chardonnay and identification of a stolbur phytoplasma in diseased plants. Gajardo et al.: Flight activity of Scaphoideus titanus and Hyalesthes obsoletus and their infection status towards FD and BN (stolbur) phytoplasma. Leitner: The situation of Grapevine yellows in Austria is now surveyed by means of molecular detection assays.: Compared efficiency. Study of the spatial and temporal dispersion of the four species. rapidity and sensitivity of methods for use in routine diagnosis of grapevine phytoplasmas. Hyalesthes obsoletus and Circulifer haematoceps abundant and positive for stolbur and aster yellows phytoplasmas. in particular a clover phyllody type (16SrI-C) which is quite frequent. 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.: Report of stolbur phytoplasmas in grapevine affected with yellows in Albania. Discussion on benefits and limits of artificial feeding medium to assess the transmission capability of phytoplasma by insects. Apart from FD phytoplasma reported by Duduk et al.: 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. titanus found associated to grapevine yellows with a high incidence in vineyards of Rastina (South Serbia). Chabbouh et al. D'Ascenzo et al. An aster yellows phytoplasma is suspected. M'hirsi et al.2003 2003 2003 2003 2003 2003 Boudon-Padieu et al.: Report of aster yellows phytoplasmas in grapevine affected with yellows in Tunisia. Other phytoplasmas are reported.: Data on 15-year studies on efficiency and effect on propagation material of hot water treatment.: Survey of potential vectors of phytoplasmas in vineyards of the Golan Heights. Recovery is a progressive phenonenon developing over 3 years in the case of BN. fitted to routine detection. two nonpreferentially ampelophagous vectors of grapevine phytoplasmas. 2003 2003 2003 2003 2003 2003 2003 2003 2003 2004 2004 2004 147 . Duduk et al.: Identification of grapevine yellows symptoms in Tunisia and attempts to identify pathogen agent and vector.: Measurements of basal respiration rate of grapevine dormant wood as an indicator of the best period for application of hot water treatment. stolbur phytoplasmas are suspected to occur because of long-lasting record of stolbur disease on many host plants in Serbia. Crocker et al.: Development of a sensitive PCR procedure for dual identification of FD and BN phytoplasmas. Megophthalmus scabripennis positive for aster yellows phytoplasma. (a and b): FD phytoplasma (16S rV-C) and S. Neoaliturus fenestratus.: Important presence of GY in Abruzzo (central Italy).: Improvement of detection of FD and BN in field-grown grapevines with real-time PCR. Consequently roguing of affected vines should be avoided when possible and especially if vectors are controlled. Milkus et al. Myrta et al. Orenstein et al. respectively. Osler et al.. Clair et al. Lessio et al.: Identification of aster yellows (16SrI-B and C) and of ash yellows (16Sr-VII) related phytoplasmas associated to yellows of grapevine in Chile. Marzachi et al. No important damage has been reported. Kuzmanovic et al. BN (stolbur phytoplasma) has an incidence of about 30 %.
In France. Eggs are laid in summer on two-year old wood and trunk. Symptoms develop on leaves and shoots in June and July to become oustanding in August and autumn. symptoms may be limited to a few shoots. If the plants are inoculated after recovery. S. Infected rootstocks are important means of dissemination because they are symptomless. it occurs in all southern vine-growing regions. 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. Hence. Infected rootstocks show little or no symptoms. with an incidence that increases rapidly from one year to the next. Affected vines of most varieties may recover in the second year when they are protected against reinoculation with insecticide treatments. Transmission: The FD agent is transmitted in the persistent mode by the leafhopper Scaphoideus titanus Ball (= S. Two isolates denoted FD-D and FD-C. After the discovery of other GY diseases. It is also present in regions from which FD has not been reported in France. titanus. Switzerland. Scaphoideus titanus Ball (= S. However. and Savoie. north of Spain and Portugal. littoralis Ball). and western Slovenia. decline progressively and die. that has colonized a wide climatic area. Cicadellidae). Isolates F70. titanus. FD88. Lignification of the canes is usually incomplete. Most of the time. Leaves persist longer on affected plants. Description Flavescence dorée (FD) was the first Grapevine yellows (GY) disease to be reported. It is highly epidemic and extremely dangerous because of the biology and ethology of its leafhopper vector. titanus is well established. FD is endemic only in regions where S. It is widespread in all northern provinces of Italy and an outbreak was recently reported from southern Serbia. FD88. Corsica. southern Italy. then as a virus disease. show a general asymmetric bent posture and necrosis of terminal bud. the term "FD sensu stricto" was applied to diseases and phytoplasmas that are transmitted by S. were characterized in Italy and shown to be transmitted also by S. Though the rate of transmission by bench grafting can be very low. 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. symptoms affect the whole plant.) using S. Agents: FD was first regarded as a physiological disorder. extremely sensitive varieties do not recover. introduced into Europe at the beginning of the 20th century. However. First symptoms may 149 .GRAPEVINE YELLOWS: INDIVIDUAL DISEASES A. 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. FLAVESCENCE DORÉE 1. FD92 and FD2000 were experimentally transmitted to broadbean (Vicia faba L. This leafhopper is a neartic species specialized on Vitis sp. isolated infected vines have been identified in northern vineyards of Burgundy and Alsace. Vineyards in these areas are under the threat of an FD outbreak that could occur if FDinfected planting material is introduced. Diseased vines have a patchy distribution in the plot. Molecular comparisons have shown that all these isolates are more closely related among them than with other phytoplasma strains in the same group. littoralis Ball) (Homoptera. Transmission occurs also by vegetative propagation and grafting. indicating a vine-to-vine transmission. titanus individuals collected in infected vineyards of south-western France. Moreover. transmission is possible from early June (about one month after the beginning of hatching) until the death of adults. Crop losses may be very high. this way of spreading is significant when infected grafted plants are planted in vineyards hosting the vector. It was described in a limited area in north-eastern Cataluña (Spain). In addition. 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. Several isolates have been characterized with molecular criteria. The associated phytoplasma (= MLO) was observed in phloem tissue of affected plants and in insect vectors in 1971 and later classified in the Elm yellows group (16SrV). Feeding transmission starts after a 4-week latency in the body of the vector. These isolates could be distinguished by Western-blot analysis using FD antisera and monoclonal antibodies (Mabs). FD92 and FD-D could not be distinguished from one another. rooted cuttings from infected canes of a few rootstock varieties can develop "vinifera-like" symptoms on the wood and leaves.
In 2003 a phytoplasma resembling the FD-C isolate was found in wild Clematis in Veneto (Italy). at the beginning of July. has been used as the official method for large scale survey of FD in France from 1993 to 2003. whereas the third treatment. In France and Italy. a DNA fragment specific to BN (stolbur) phytoplasma can be used for the survey and monitoring of these two GY diseases. The low rate of transmission by grafting and the long delay in symptom expression make the detection of infected vines in planting material difficult. Polyclonal antisera and Mabs have been raised to FD phytoplasma. must be destroyed. Ugni blanc (Trebbiano) and Prosecco are sensitive varieties that may recover when they are protected from new inoculations. Other DNA-based methods. sensitive amplification with nested PCR of the DNA fragment FD9. 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. Symptoms are rare in Syrah. Cabernet Sauvignon. Nielluccio and Garganega are very sensitive varieties that usually do not recover after infection. roguing of diseased vines is advisable when the epidemic pressure is high. Other varieties. In France and Italy budwood can be taken only from mother plants growing in plots that have shown no GY symptoms for the last two growing seasons. Antigen extraction from vascular tissue of symptomatic grapevines requires the use of high Tris molarity and strong detergents in the extraction buffer. Vitis riparia can be infected but shows little symptoms. although heavily infected vines can be observed. appear more tolerant. Detection: FD symptoms can be confused with those of other GYs and also with other diseases or disorders. Roguing is compulsory in France. Sauvignon blanc. An indirect DAS-ELISA using coating of wells with rabbit polyclonal antisera and detection with a cocktail of several Mabs. have been found carrying FD phytoplasma until recently. Grenache. Simultaneous amplification in a multiplex procedure of both FD9 and Stol11. Their rearing is still under development. has become the preferred detection method because it is easy to perform and was improved in reliability and sensitivity. The presence of S.appear on young vines as late as four years after grafting. and the risk of disease spreading important. In spite of the possibility of recovery. Monitoring natural enemies of S. Molecular detection by PCR of selected phytoplasma DNA fragments. aims at killing newly hatched insects. Chardonnay. Material from mother plants that have developed symptoms in the following growing season. Indirect control is obtained by insecticide treatments against S. is directed against winged adults migrating from nearby vineyards or wild vines. control measures are compulsory according to legal regulations. Varietal susceptibility and sensitivity: All varieties grown in the various FD-affected regions in France. Other host plants: No host plants other than Vitis sp. are currently under development. Natural insecticides can be used to preventively limit vector populations but their efficiency is limited in epidemic outbreaks. has allowed the identification of potential auxiliaries for biological control. Bois noir (BN) is also frequent in regions inhabited by this leafhopper. Soaking of dormant material in hot water (HW) is recommended. Recently. The second treatment. its area of origin. The first treatment is applied 30 days after first instar emergence at the beginning of the period of potential transmission. 150 . such as real-time PCR or identification of PCR-amplified product with DNA-DNA hybridization using specific probes. Standard conditions (50°C for 45 min) proved highly efficient for sanitizing infected rootstocks and grapevine cultivars. Nevertheless. at the beginning of August. Planting material must not be collected from mother vines in areas or vineyards affected by the disease. Under these circumstances only chemical insecticides are efficient for eradication of the disease. Italy and Spain are susceptible with various degrees of sensitivity. The same is true for American rootstocks which can therefore be dangerous sources of infection and dissemination of the disease. Planting of HW treated material is especially important in areas where the vector is present. which is specific for 16SrV-group phytoplasmas allowed the reliable detection of FD phytoplasma in grapevine. titanus in New York State (USA). Detection with laboratory methods is possible on insect vectors and infected vines. Alicante Bouschet. Two additional treatments are applied during summer. Sequence or RFLP analysis of the FD9 DNA fragment permit to readily differentiate all the known FD isolates. titanus. titanus in an affected vineyard is an important indication that may lead to suspect a FD infection. Control: FD is a quarantine organism in the EC. such as Merlot.
Carle and Moutous: Study on possible secondary or alternative vectors of FD or Bois noir (BN). Use of 4-5 treatments with DDT or parathion and removal or destruction of pruning wood older that two years. Caudwell: Attempts to inhibit the agent of FD in vivo by heat treatment. an insect recently introduced from North America. Caudwell et al. littoralis. Description and study of recovery phenomenon and of localised symptoms. littoralis on plants other than the grapevine and transmission trials of FD to other host plants. considered as a virus disease.: First report of S. littoralis is present. Schvester et al. Caudwell: Study of the phenomenon of recovery of FD-affected vines. 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. Schvester et al. littoralis recorded from in Italy in 1963. Vidano: Study of the ecology and biology of S. Boubals and Caudwell: FD epidemics observed in Corsica. Description of macroscopic and microscopic symptoms. littoralis in Corsican vineyards with winter treatment of dormant eggs with oleoparathion. The disease can be transmitted by grafting and has probably a viral aetiology. S. Bonfils and Schvester: Relationships of leafhoppers with FD of grapevine in western France. (a): Control of FD by insecticide treatments against the vector. Historical review 1955 1956 1957 Levadoux: Description of an outbreak of "Flavescence dorée" on Baco 22A in southwestern France. Caudwell and Poitou: Study of the interaction between fanleaf and FD. Caudwell: PhD thesis on FD. Branas (a and b): The new disease is thought to be caused by root damage. littoralis. biology and feeding damage and of the symptoms of FD in France on Baco 22A. Vidano: S. No positive results of transmission trials with 15 species of Hemiptera (mainly Cicadellidae) other than S.2.: Demonstration that FD is transmitted by the leafhopper S. littoralis in its area of origin in North America. (a). Description of the insect. littoralis from Ticino (southern Switzerland). 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. Caudwell et al. Caudwell et al. littoralis. Immersion in lukewarm water (30°C) for 72 h of cuttings from FD-affected grapevines reduced the proportion of infected plants by about 83 % as compared with untreated control. Schvester et al. evolution of the disease in space and time. S. 1960 1960 1961 1961 1962 1962 1963 1964 1964 1966 1966 1967 1968 1969 1970 1971 1971 1972 151 .: Field tests for insecticide control of S. Recovered vines may be infected again but the symptoms are lighter. First report that abandoned or wild vines may be a source of infection. Spontaneous recovery occurs after a 1-2 year crisis period. littoralis Ball. littoralis. (b): Biology of S. The aetiology is unknown.: Successful transmission of FD to herbaceous hosts by increasing the feeding access period of the vector S. (a): Possibility to limit the populations of S. faba plants.: Studies on the survival of S. 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.
1984 1985 1985 1985 1986 1986
1986 1987 1987 1987
1987 1987 1987
1987 1989 1989
Seljak: Presence of S. titanus in Slovenia (western Yugoslavia). Anonymous: Compared efficiency of various insecticides in the control of S. titanus. Boudon-Padieu et al. (a and b): Polyclonal antisera raised to FD MLO maintained in the laboratory permit detection in infective S. titanus individuals from affected vineyards. Description of the infection cycle of the MLO in the body of the experimental vector E. variegatus. Du Fretay et al.: Satisfactory control of the vector of FD in 1988 in South France. Fortusini et al.: Successful transmission of FD to grapevine cuttings using S. titanus in Italy. Laurent and Agulhon: 1989. Comprehensive review of the situation and evolution of FD and the vector leafhopper in French wine-growing regions. Lherminier et al.: In situ detection of FD MLO in salivary glands of infective Euscelidius variegatus leafhoppers with immunofluorescence. Schwartz: PhD thesis on the production, screening and characterization of monoclonal antibodies to the FD-MLO. Boudon-Padieu et al.: Serological differentiation between FD-MLO and Phy-MLO, an agent causing phyllody in V. faba plants, transmitted in the laboratory by Euscelidius variegatus leafhoppers. Caudwell et al.: Hot water treatment to disinfect grapevine wood from FD-MLO. Lherminier et al. (a and b): The use of immunolabeling in the electron microscope to detect and localize MLO in plants and insects with specific polyclonal antibodies. Arzone et al.: Electron microscopic observation of MLO in sieve elements of Chardonnay and Perera grapevines showing FD symptoms, in clover (T. repens) inoculated with S. titanus fed on infected grapevines and in salivary glands of the leafhopper. Cazenove and Planas: Attempts to obtain a satisfactory control of S. titanus with natural insecticides used in biological agriculture. Anonymous: Recommendations for control of FD in biological wine-growing. Combination of winter treatments with white oil and removing of pruning wood to suppress eggs. Caudwell and Kuszala: ELISA detection of FD MLO in affected grapevines. The method requires extraction of proteins with a strong detergent. DAS ELISA is developed with polyclonal rabbit antiserum as coating antibodies and a cocktail of mouse monoclonal antibodies as detecting antibodies. Daire et al.: Random cloning of FD-MLO DNA and selection of specific probes for detection of FD MLO with DNA-DNA hybridization. Detection in field-infected grapevines is possible only on DNA extracted from a MLO-enriched fraction. Jermini et al.: In spite of the presence in Ticino (Switzerland) of S. titanus in vineyards and nurseries, FD does not seem to occur. Lozzia: A review of the presence, biology and control of S. titanus in Italy. Meignoz et al.: Electron microscopic study of the cytopathogenic effects of FD-MLO in experimentally infected cuttings of LN33. Osler et al.: S. titanus fed on yellows-diseased grapevines in northern Italy (Friuli-Venezia Giulia), test positive to FD-antibodies raised in France. Caudwell et al.: Positive indexing of rootstocks showing latent FD infection and erratic distribution of the pathogen in the material. Sanitation of rootstocks with hot water treatment (50°C for 45 mn).
1989 1989 1989 1989 1989 1990
1990 1990 1991
1991 1992 1992
1992 1992 1992 1992 1993
Daire et al. (a and b): Diversity among MLOs inducing GY diseases in France and other countries, shown with PCR-RFLP analysis of rDNA. FD shows the same pattern as elm yellows MLO. BN is related to stolbur MLO. FD MLO was detected in samples from France and Veneto (Italy). Jermini et al.: Assessment of the optimal number and disposition of sticky traps to evaluate the importance of S. titanus populations in the vineyard. Lefol: PhD thesis on recognition between the FD-MLO and the organs of the experimental leafhopper vector Euscelidius variegatus. Lefol et al.: Development of the "double-dot" method and a derived procedure in the transmission electron microscope to identify the sites of attachment of the FD-MLO on insect organs in vector and non-vector insect species. Maixner et al.: Study of S. titanus in New York and its possible relationship to an American GY. S. titanus transmitted a yellows disease to 29% of Vicia faba plants. FD antibodies raised in France tested positive on 13% of American S. titanus individuals. Seddas et al. (a and b): Immunoaffinity purification of FD-MLO from infected experimental vectors. Carraro et al.: Demonstration with transmission trials using S. titanus, ELISA and PCR analyses, that FD sensu stricto is present in Veneto and not in Friuli-Venezia Giulia, though S. titanus is also present in the vineyards of the latter region. Caudwell et al.: Identification of latent FD infection in several rootstock varieties, including 3309 Couderc and Fercal. The rate of graft transmission to scion depends on the infection status of mother vines and could reach 80 %. Farmer and Boudon-Padieu: Cloning of FD-MLO DNA, construction of an expression library and screening with FD antibodies allowed the selection of three inserts carrying information for FD membrane proteins. Lefol et al.: Route and multiplication of the FD-MLO in the body of experimentally infected Euscelidius variegatus. All main organs were infected, except for germinal cells. Lherminier et al.: Use of ELISA and immunolabeling by transmission electron microscopy for studying the distribution and movement of FD-MLO in the experimental host plant Vicia faba. Seddas: PhD thesis on the production and use of monoclonal antibodies to identify the main antigens of FD-MLO and obtain highly purified fractions of the infectious agent from vector insects and herbaceous host plants. Bertaccini et al.: Presence of phytoplasmas of two types, sometimes in double infections, in severely GY-affected grapevines in Liguria (North-western Italy). One type is related to elm yellows phytoplasma and the other to IPVR phytoplasma, shown later on to belong to the stolbur group. Seddas et al.: Immunoaffinity purified FD phytoplasmas appear as whole cells in ISEM. Experimental vectors (E. variegatus) injected with purified fractions of FD phytoplasma supported its multiplication and were able to feed-inoculate V. faba seedlings. Alma et al.: In Piemonte, several phytoplasmas were identified in yellows-affected grapevines. A 16SrV (= elm yellows) phytoplasma was detected in a double infection with a 16SrI-G phytoplasma. Phytoplasmas of the latter group (later renamed 16SrXII = stolbur) were the more numerous. Bianco et al. (a and b): Elm yellows-related (16SrV) phytoplasmas are present only in the Vicenza and Arezzo provinces of northern Italy. Two different RFLP patterns can be distinguished, suggesting the existence of two strains of FD phytoplasma, one of which is similar to the French strain.
1993 1993 1993
1994 1994 1994
(a and b): Evolution of FD in the Treviso province (Veneto.: A phytoplasma of the 16SrI (aster yellows) group is detected in eggs. Serological relationships with elm yellows phytoplasma is confirmed Alma et al.: RFLP studies of 16SrV phytoplasmas. titanus laid in the bark were killed. Daire et al. according to the severity of the FD epidemics. The possibility that direct inoculations have occurred in nursery is discussed. Bianco et al. Belli et al. the eggs of S.: Important outbreak of FD in Lombardia (northern Italy). nymphs and adults of S. Pavan et al. Control recommendations. Spain and Italy. (a and b): Demonstration of curing effect of hot water treatment on FD-affected plant material. Survey of hemipters in the vineyard and presence of other known phytoplasma vector species.: Insecticide control of S. Clerc et al. The possible role of six species in the transmission of GY is discussed.: Monitoring of leafhoppers in vineyards in Italy. Batlle et al. Rousseau: A review of different ways to reduce the populations of S. titanus in Italy. Three different RFLP patterns at least are shown in FD isolates from France. The biological significance of this finding is unknown. titanus in north-eastern Italy where vines are trained in the pergola system. titanus in Switzerland. titanus reared on healthy plants.: Monoclonal antibodies raised to immunoaffinity purified FD phytoplasmas. FD is not present in southern Italy. Numerous species identified. First outbreak in the 1980's on cv Perera. 1996 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1999 1999 2000 2000 155 . especially Prosecco in the period 1993-1996. Lherminier and Boudon-Padieu: Use of transmission electron microscopy with in situ immunolabeling to detect FD phytoplasma in grapevine and herbaceous host plants. Vindimian et al.: In spite of the presence and rapid spread of S. Several different peptides from membrane proteins are identified by Western-blot. Kuszala: Specific ELISA procedures to detect FD or BN phytoplasmas in extracts from diseased grapevines from the field. Martini et al. Caudwell et al.: Development of specific PCR primers for the detection of FD or BN phytoplasmas. Bosco et al. In the same conditions. The amplified FD9 fragment is specific and variable among 16SrV (elm yellows) group phytoplasmas.: First report of FD outbreak in northern Cataluña (Spain).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.: Demonstration that two isolates of FD sensu stricto (FD-D and FD-C) are spreading in Veneto (Italy). Mori et al. Italy). titanus in Trentino (Italy). (a and b): Situation of FD and S. titanus in biological viticulture. Jermini and Baillod: Combination of visual observation of instar larvae and trapping of adults on sticky traps to monitor the populations of S. Transmission by grafting is studied.: Verification of hot water treatment conditions to suppress FD phytoplasma infection in dormant wood of grapevine.: First report of S. Marcone et al. Posenato et al. Seddas et al. Infection is rapidly spreading to the east. FD has not been identified. Increase and spread to other cultivars. titanus in the canton of Geneva (Switzerland).
Boudon-Padieu (a): Description, biology, spread, FD-transmission and control of S. titanus. Clair et al.: Design of internal nucleotides on the specific DNA fragment FD9 to be used as primers in nested-PCR for improving sensitivity of detection of elm yellows (16SrV) phytoplasmas in FD-affected grapevine or in elms affected with yellows in France. Cravedi and Aldini: Biology of S. titanus in Oltrepò pavese (Italy), a region affected by a severe outbreak of FD since 1999. Morone et al.: Occurrence of FD in Piemonte (northern Italy) since 1998. Roure: The situation of FD in France. Compulsory control (insecticide sprays and roguing of affected vines) on 300 000 ha. Scattini et al.: Important spread of FD in Lombardia (Italy) in cv Sangiovese. Angelini et al.: Phylogenetic comparison of FD isolates from France and Italy among them and other 16SrV (elm yellows) strains. One isolate from France (FD92 = FD88) and one from Italy (FD-D) are widespread and identical. Bianco et al.: S. titanus in northern Italy can transmit different grapevine isolates of 16SrV (elm yellows) phytoplasmas. Credi et al.: Detection of FD in grapevine samples from the provinces of Piacenza, Parma, Reggio Emilia and Modena in Emilia Romagna (Italy). Davis and Dally: A new 16S rDNA sequence of FD is produced and it is proposed that FD phytoplasmas are placed in two distinct subgroups. However, further work has shown that the 1994 original sequence was erroneous (see Angelini et al., 2003a). Harrison et al.: An elm yellows (16SrV) group phytoplasma close to the Italian isolate FD-C is detected in Virginia creeper plants (Parthenocissus quinquefolia) in southern Florida (USA). It is the first time that a phytoplasma resembling a FD phytoplasma is found in a plant species other than Vitis sp. Marzachi et al. (b): Development of a PCR strategy for large scale detection of FD phytoplasma in plants and insects. Pasquini et al.: Harmonization of procedures for diagnosis of FD in Italy. Quartau et al.: Presence of S. titanus in Portugal. Alma: A review of the distribution and biology of S. titanus in Italy. Bianco et al.: Development of a specific TaqMan® assay for detection of FD phytoplasmas. Borgo and Angelini: Spread of FD in Italy and the situation of mother vines and cultural practice. Boudon-Padieu: Present knowledge on the epidemiology, aetiology and diagnosis of FD. Credi et al. (b): Presence of FD in the Marche region (central Italy). Martini et al.: Further comparisons between FD phytoplasma isolates from France and Italy. Mori et al.: Positive transmission of the two Italian FD isolates (FD-D and FD-C) by S. titanus. Posenato et al.: Efficiency of various insecticides to destroy nymphs of S. titanus and Metcalfa pruinosa. Viggiani: Presence of S. titanus in the Basilicata region (southern Italy).
2000 2000 2000 2000 2001
2001 2001 2001
2001 2001 2001 2002 2002 2002 2002 2002 2002 2002 2002 2002
Angelini et al.: Use of Heteroduplex Mobility Assay and full sequencing on two genomic DNA fragments (ribosomal and non ribosomal) to further study the relationships between FD isolates and related 16SrV (elm yellows) group phytoplasmas. Bianco et al. (b): Presence of FD-D and FD-C isolates of FD phytoplasma in Lombardia (northern Italy). Botti and Bertaccini: Molecular variability between FD phytoplasma isolates using the 16S rRNA gene, the ribosomal protein operon and the FD9 DNA fragment. Distinction between epidemic and non epidemic isolates. Bressan et al.: An attempt to identify a pattern of seasonal transmission of FD, taking into account the transmission efficiency of S. titanus, the pattern of symptom expression by infected grapevines, the emergence of nymphs of S. titanus and the importance of vector population and diseased grapevines. Cavallini et al.: Presence of FD together with BN in the area of Modena (north central Italy). Clair et al.: A multiplex nested-PCR procedure for easy simultaneous diagnosis of FD and BN. The method is now the official method registered in the Official Journal of the French Republic to be used by laboratories in the frame of compulsory control of FD in vineyards. Constable and Boudon-Padieu: Isolation of FD phytoplasma chromosome in Pulse Field Gel Electrophoresis and first physical map for two FD isolates. De Sousa et al.: Identification of FD-D phytoplasma in S. titanus specimen trapped in Portugal. No description of FD-affected grapevines. Duduk et al. (a and b). Identification of FD in southern Serbia and presence of S. titanus. The phytoplasma isolate is similar to FD-C. Malausa et al.: Search for natural enemies of S. titanus in New York with the scope of their introduction in France as biological control auxiliaries. Nusillard et al. Results of a 2-year study in North America of the natural enemies of S. titanus. Santinelli et al.: Presence of S. titanus in Umbria (central Italy). Torres et al. The situation of FD in Spain. Confirmation that only FD-D is present. Angelini et al.: A phytoplasma identical to FD-C isolate is found in wild Clematis in the vicinity of a FD-C affected vineyard in Veneto (Italy). Lessio and Alma: Study of the distribution of S. titanus within a vineyard and use of chromatic sticky traps. Abundance is greater in normal- than in low-density planted vineyards. S. titanus is monophagous. Leafhoppers are not able to spread significantly outside a vineyard. Females are less likely to fly far away than males.
2003 2003 2003 2003 2003 2003 2003 2004 2004
B. BOIS NOIR (VERGILBUNGSKRANKHEIT, LEGNO NERO) 1. Description Bois noir (BN) and related diseases are Grapevine yellows (GY) known in Europe and Asia Minor. The first observations in Eastern France date back to the early 1940's. BN was formally described in 1961 in comparison with Flavescence dorée (FD), because symptoms were similar. It was assumed that the two diseases were different because BN occurred in regions where the vector of FD was not present and experimental transmission with S. titanus was not successful. Because the vector insect, Hyalesthes obsoletus, does not live on vines, affected plants are usually not grouped into patches as with FD, except for situations where a high percentage of affected plants is observed. Nevertheless, distribution is non-random and spread follows a main direction or is preferentially along the border of the plot. BN has been identified in all wine-growing regions of western and eastern Europe, in Israel, Lebanon and recently in the Ukraine.
Main synonyms: The syndrome named Vergilbungskrankheit (VK) (1971) or Schwarzholzkrankheit (SHK) (2003) in Germany, Legno nero (LN) in Italy, Bois noir in Spain, has been occasionally described under the name of Flavescence dorée-like disease. Moreover, it has been sometimes confused with FD in the 1970's and 1980's. It is only since the 1990's that the aetiology was established and that it was shown that the agents of these diseases in the different countries were similar or closely related. Furthermore, the same vector insect species, H. obsoletus, has been identified in all countries, except for Spain where it appears to be very rare. Main symptoms: All the main symptoms typical of GY can be found in BN / VK / LN - infected grapevines. In addition to leaf discoloration, growth abnormalities can be observed on canes and roots. Usually, only a few canes per plant show typical symptoms with either an uneven or a total lack of lignification. On whiteberried varieties, yellow banding develops along the veins which may undergo necrosis. On red-berried varieties, sectorial reddening of blades develops in summer and progressively invades the whole leaf. When berries develop they remain immature, greenish and eventually fall down. Remaining berries are tasteless and sour. Internodes often show longitudinal rows of brown pustules along the green bark of unripe wood. Severely affected vines decline and may die after a few years. Some vines may recover or express irregular symptoms in successive years. Agents: The phytoplasmas associated to these diseases belong to the stolbur group (16SrXII), a group that was for some time considered by some to be the I-G subgroup of the larger Aster yellows group (16SrI). Although stolbur phytoplasmas are ubiquitous pathogens, known to infect solanaceous crops as well as many weeds and bushes, only a limited number of strains have been identified. Hence the different names of BN disease appear to be more geographically than genetically significant. Transmission: Vergilbungskrankheit (= Schwarzholzkrankheit) was the first disease for which the vector was identified. The known vector insect of stolbur of tomato, H. obsoletus (Homoptera, Cixiidae) can transmit stolbur phytoplasma to grapevine, but this occurs during feed probing as the insect does not keep feeding on grapevine. Hence, vine-to-vine transmission does not occur. The role of H. obsoletus has been confirmed in France, in Italy and Israel. In Spain, where the species is rare, its role in BN epidemiology must be verified. Alternative vectors have been suspected but not demonstrated. H. obsoletus is a univoltine polyphagous species that overwinters as larval instars that live on the roots of host plants. Nymph emerge in spring and transmission occurs during adult mating flights. Overall, the presence of insects on vines may last a couple of months. Eggs are laid during summer on the ground at the base of the stem of host plants. Main host plants of the insect vector are common weeds such as Convolvulus arvensis, Ranunculus sp., Urtica dioica, Cardaria draba and Calystegia sepium. However, only C. arvensis and U. dioica were found consistently infected with stolbur phytoplasma, distinct isolates of which were identified in these hosts. From 10 to 80% of trapped populations of the vector may test positive for the presence of stolbur phytoplasma. Transmission by grafting is possible but only at a low rate. Since acquisition by the vector from infected vines does not seem possible, the significance of graft transmission on the spread of the disease is probably very low. Varietal susceptibility and sensitivity: In the different affected regions and countries, numerous cultivars of V. vinifera were found susceptible to the stolbur phytoplasma, cv Chardonnay being the most sensitive. The increasing incidence of BN has been related to the increase of surfaces planted with Chardonnay in the last decade in all countries. Infection of rootstock varieties has not been reported. In all varieties there is a proportion of affected grapevines that undergo transitory recovery or remission, while other plants in the same plot are tolerant or immune to infection. Other host plants: Stolbur phytoplasma is widespread, and has a wide host range. In addition to C. arvensis, U. dioica, Ranunculus sp., C. draba and C. sepium, shrubs and trees of Prunus spp., solanaceaous crops (tomato, tobacco, pepper, egg plant) and lavender (Lavandula spp.) may be significant reservoirs. Detection: The use of monoclonal antibodies raised to stolbur phytoplasma did not show enough sensitivity for routine ELISA detection in field-grown infected grapevines. Detection is achieved mainly with PCR assays or PCR-RFLP analyses of DNA from infected host plants and insect vectors. Most often, nested PCR is necessary for sensitive detection from infected grapevines. Universal primers were first used to detect both BN and FD in areas where the two diseases may coexist. Subsequent RFLP analyses of amplified phytoplasma rDNA permit the characterization of pathogens. Stolbur specific primers have also been designed, either on variable regions of the rRNA gene or on selected specific non-ribosomal
DNA fragments. cytological and electron microscope study of tissues of affected grapevine. Caudwell et al. 2. Historical review 1961 1965 1968 1969 1971 1971 Caudwell: Study on BN and its relationships to FD. the multiplex nested-PCR assay cited in the FD chapter can be used. Cazelles et al.: GY symptoms are present in Switzerland (Suisse romande and Ticino) but they are not associated with S. Leclant and Lacote: Study of the ability to transmit stolbur to herbaceous plants by H. Hence.: 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. Gärtel: Description of VK under the name of Flavescence dorée. The results suggest that the disease is brought into the vineyards from outside local sources. because of the complex biological cycle and multiple host plants of the insect species. Chemical sprays against H. Borgo: General information on the presence of FD-type diseases in northern Italy. BN is considered as a non epidemic form of FD. Control: As with other GY diseases. no direct control is possible. Findings never confirmed. history. Description of structures that were probably closteroviruses. 1972 1977 1978 1978 1988 1989 1992 159 . Italy. The species has been described as the vector of stolbur "virus" by Suchov in USSR in 1948. littoralis. Some grapevines show symptoms repeatedly in successive years. and different susceptibility and sensitivity of grapevine cultivars. absence of recovery. (b): Epidemiological observations of BN lead to the assumption that an aerial vector is involved. except for declining vines. Credi and Callegari: Survey of vineyards in Emilia-Romagna. obsoletus are not efficient. roguing should be avoided. 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. Natural enemies are being investigated. unknown at that time. Caudwell et al. suppression of affected branches may improve harvest quality and help in progressive recovery. for the presence of a FD-like disease. titanus.: Rickettsia-like organisms found in roots of VK-affected grapevines in the Saar. Recovery and remission of symptoms for several consecutive years are frequent in vines affected by BN. Description of symptoms of VK. S. Though pruning does not cure infected vines. Rumbos et al. The disease occurs in the Moselle valley and the Rhine valley. Rumbos et al. These findings have never been confirmed. Mendgen: 1971. (b): BN is distinct from FD based on non transmissibility by S. obsoletus in the south of France.: Rickettsia-like organisms from VK-affected grapevines are cultured in chick embryos. Superficial soil cultivation in summer or winter ploughing are methods intended to damage the larval instars and reduce vector populations. Leclant: Presence of Hyalesthes obsoletus a cixiid planthopper in the south of France. titanus is not always associated with the disease. For direct differentiation between FD and BN/VK phytoplasmas. while others seem more tolerant and do not show symptoms. Nienhaus 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.
: Grapevines from Piemonte (northern Italy) contain several different phytoplasmas. and from Israel. subgroup I-G (later renamed as 16SrXII or stolbur group).: The agent of a GY disease in the Italian Riviera is detected with a probe specific for Eastern aster yellows. Davis et al. from several regions of Italy. Daire et al. MLO associated to GY in Italy are related to aster yellows MLO.: A monoclonal antibody raised to stolbur mycoplasma-like organism is used for ELISA detection of the agent in herbaceous host plants and insects. (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. Brescia and Pavia (northern Italy).: One of the two phytoplasmas detected in yellows-diseased grapevines in Liguria is related to IPVR phytoplasma. Laviña et al. obsoletus. Biology of the insect. Boudon-Padieu (b): History. (b): Evidence of an association of MLOs with grapevine yellows in EmiliaRomagna. Maixner et al. c and d) Transmission of a yellows to periwinkle with dodder in Germany. Maixner et al. Italy) where FD is prevalent. It is suggested that vectors with preference for other host plants act as vectors for VK. (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. The MLO strains found in grapevine are related with aster yellows MLOs. Bertaccini et al. Reservoirs and possible role of other vectors remain to be elucidated. (b): Presence of phytoplasmas in the group 16SrI. Maixner : Demonstration that H. subgroup I-G (later named group 16SrXII (stolbur) in the provinces of Vicenza. Several weeds are host plants of the stolbur phytoplasma. Minucci et al. Bianco et al. Davis and Prince: According to a different terminology. clustered by the authors into aster yellows group but shown later to belong to the stolbur group.: PCR-RFLP analysis of MLO-DNA detected in VK-affected grapevines in Germany show that this MLO is related to stolbur. Borgo: Presence of BN in mixed infections with FD in the province of Treviso (Veneto. aetiology and transmission of BN in France. (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. obsoletus is a vector of VK in Germany. Computer analysis for spatial patterns of spread shows a non-random distribution and suggests preferential transmission from other vines or from weeds.: First detection of BN (stolbur phytoplasma) in Spain. Maixner (b. H. obsoletus identified as the vector. obsoletus is confirmed as a vector of stolbur disease plants in France. Alma et al. Bianco et al.1992 Fos et al. but are different from known strains of MLOs of the aster yellows cluster. (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). Bertaccini et al. 1993 1993 1993 1993 1993 1993 1993 1994 1994 1994 1995 1995 1995 1996 1996 1996 1996 160 . The latter MLO was shown later to belong to the stolbur group. BN MLO is detected in grapevines from France. 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. PCR-RFLP analysis confirms relationship to aster yellows phytoplasmas. the most frequent belonging to group 16SrI. H. Italy.
subgroup I-G (identified later as belonging to the stolbur group or 16SrXII). Daire et al. Davis et al. search for vectors. Marcone et al.: Occurrence of GY diseases in Hungary and identification of stolbur phytoplasma in affected grapevines. arvensis is recommended to expose instar larvae and kill them with frost. as well as phytoplasmas transmitted by Credi to periwinkle in Emila Romagna and previously classified in the aster yellows group.: Monitoring of field populations of H. First detection of BN agent in diseased grapevines in Switzerland with the same procedure. Weber: PhD thesis on the biology of the cixiid H.: Prevalence of BN (stolbur phytoplasma) in Navarra and Cataluña (north eastern Spain). Albanese et al. Refatti et al. Kuszala: Detection in ELISA using a monoclonal antibody to stolbur phytoplasma of BNassociated antigens in diseased grapevines in France. The maximum delay of symptom expression in young plants is 2 years. 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.: Important spread of GY symptoms on several grapevine varieties in Umbria (central Italy). Search for variability among BN phytoplasmas with RFLP analysis of the specific non ribosomal DNA was unsuccessful. a strong annual increase. obsoletus as a vector of BN. Sforza and Boudon-Padieu: Description of H.: Phytoplasmas associated to GY in Campania (southern Italy) are genetically uniform and belong to the stolbur group. Kölber et al. titanus. Maixner and Reinert (a): Trials on elimination of VK infection from vine wood in Germany. a high proportion of vines in which symptoms re-occurred after one season or more. Characterization of a phytoplasma belonging to group 16SrI. Weber et al. Maixner and Reinert: A review of the research and present knowledge on VK and other GY diseases. Preferential spread along the rows. 1996 1996 1996 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1998 1998 1998 1998 161 . Laviña et al. conversely. biology of H. Results were satisfactory. (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. using hot water treatment with conditions recommended in France. (c): Identification of stolbur phytoplasma (16SrXII) as agents of GYs in Greece and Israel.1996 1996 Koruza: Survey of GY in three regions of Slovenia shows that the disease is of BN type. Osler et al. obsoletus and possible control measures. a high proportion of previously infected vines that did not show symptoms for 2 years at least and.: Consistent association of stolbur phytoplasma with GY diseases in the Italian provinces of Fiuli-Venezia Giulia and Trento and in Slovenia. vector transmission and possible control measures. obsoletus in Germany. Vindimian: LN but not FD is present in Trentino (northern Italy) in spite of the presence and active spread of S. Sforza: PhD thesis on the epidemiology of BN in France. Ploughing in winter of soils infested by C. obsoletus and its role as the vector of VK. (a and b): BN is poorly transmitted by bench-grafting. Estimated crop damage ranges from 20 to 40 %. Maixner: The situation of VK in Germany: symptoms.
: In a large survey of vineyards in 8 Hungarian counties conducted in 1998. obsoletus as a vector to grapevine. Up to 30 % of the insects were infected and the highest rate was in vineyards with high populations of Convolvulus arvensis. Maixner et al.: Only stolbur phytoplasma detected in grapevines showing GY symptoms. PCR detection is reliable when testing together 25 insects among which only one is infected.: VK disease dramatically increased in incidence in the Moselle and Rhine valleys during the 1990's. Braccini and Pavan: Report of survey of Auchenorrynchas in vineyards of Toscana (central Italy).: BN reported only in north-western and eastern vineyards of Croatia. formerly reported in western Europe. Maixner and Reinert: Collection of H.1998 1998 Sforza and Bourgoin: Anatomy of reproductory organs and mating of H. Marcone et al. Identification of main reservoir weeds.: Ethological onservations and morphological descriptions of adults and larvae of H. High rate of BN / LN infection in the different vine-growing regions.: Epidemiology of BN in France. Šeruga et al. the second GY disease in Germany. First launching of colonies of the insect under controlled conditions. Females are more often infected than males. No other grapevine phytoplasma was detected. The same pathogen detected in weeds and other crops.: Use of PCR to confirm the presence of stolbur phytoplasma in GY-affected grapevines in Switzerland. and a high proportion of vines that escaped infection during the 3-5 year period of monitoring. Weber and Maixner (b): Etholology of H. obsoletus. The study confirms a high rate of infected H. No other phytoplasma found in the South of the country. Phytoplasmas in the 16SrXII group (stolbur) were the most frequent. Up to 34% of insects were infected with stolbur phytoplasma. obsoletus and Oncopsis alni. among which hoary cress. Transmission to natural host plants is higher than for grapevine with both insects. Škoric et al. Braccini et al. the ability to recover of cvs Riesling and Pinot in spite of systemic symptoms. obsoletus related to VK infection. H. The rate of infected H.: Widespread presence of LN in Toscana (central Italy). Darimont and Maixner: Comparison of the transmission efficiency of phytoplasma to their host plants and to grapevine by H. obsoletus and Metcalfa pruinosa were found positive for BN phytoplasma (16SrXII-A).: First detection of stolbur phytoplasma in grapevines in Croatia. regardless of the cultivar in Campania (southern Italy). 1998 1998 1998 1999 1999 2000 2000 2000 2000 2000 2000 2000 2000 2000 2000 162 . Sforza et al. obsoletus.: Several vineyards affected by BN in Burgundy (France) monitored for over 15 years and assessed for symptom severity. obsoletus. obsoletus can be high because of the presence of reservoir weeds in the vineyard. obsoletus in vineyards is efficient with sticky traps placed at the soil level. Varga et al. Seljak and Petrovic: A review of the Slovenian situation of GY diseases. Larrue et al. Bourquin et al. Weber and Maixner (a): Procedures for the survey of infection status of populations of H. Acquisition may occur at larval stages since emerging 5th instars were infective. different phytoplasmas were identified in a number of cultivars. Identification of two infected symptomless weeds in the vineyard. the vector of Palatinate grapevine yellows (PGY). Sforza 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) . obsoletus and possibility to control the insect in vineyards. Confirmation of the role of H.
on pigments. In spite of a similar abundance of H. The presence of H.2001 Bertamini and Nedunchezhian: Evaluation of effects of BN disease (stolbur phytoplasma) in field-grown grapevines. Bertaccini et al. obsoletus. obsoletus and the presence in organic vineyards of several weeds known as host plants carrying larvae on the roots. Gugerli et al. while Megophthalmus scabripennis was positive for aster yellows phytoplasma. 2001 2001 2001 2002 2002 2002 2003 2003 2003 2003 2003 2003 2003 2003 2003 2003 163 . Morandell: Detection of VK in south Austria.: Survey of potential vectors of phytoplasmas in vineyards of the Golan Heights.000 plants over 12 years. reached by other authors. dioica) as a reservoir of stolbur phytoplasma and as a host for the insect. Samples were taken on all organs of 10 affected grapevines. Cicotti et al. chlorophyll fluorescence and photosynthetic activities suggest that Photo System II is affected. Role of stinging nettle (U. Curkovic Perica et al. very similar to European isolates. Study of the spatial and temporal dispersion of the four species. Darimont and Maixner: Importance of the presence and infectivity of H. is a natural vector of stolbur phytoplasma to sugarbeet in Bourgogne (France). H. Indicators based on climatic parameters permit to predict the flight period. obsoletus to the German viticulture. general stress responses and rapid senescence.: Incidence of BN in a small area of the Bekaa valley (Lebanon) on cvs Chardonnay and Alicante Bouschet. Pentastiridius sp. 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. Mescalchin and Mattedi: A description of the development of VK in Trentino (northern Italy). Kölber et al. Severity of symptoms vary from one year to the other.: Importance of outbreaks of BN in the province of Modena (Italy) and monitoring of potential insect vectors in addition to H.: The highest risk of phytoplasma contamination for grapevines coincides with the flight activity of adults of H.: Demonstration that H.: The situation of GY diseases in Croatia shows that only BN is present in the northern and eastern vineyards. Myrta et al. Same conclusions about tolerance and transient recovery of grapevines. The authors conclude that phytoplasma infection induces non specific. Laviña et al. obsoletus and Circulifer haematoceps were widespread and positive for stolbur or aster yellows phytoplasmas.: First detection in Albania of BN phytoplasma in grapevines of different age and varieties. Merlot and Pinot noir. every month for one year except in winter. respectively.: Observation of the occurrence of BN in Trentino in eight Chardonnay vineyards on more than 11.: Assessment of the best period for detection of BN in affected grapevines.: Anoder cixiid planthopper. Langer et al. obsoletus is the vector of LN in Italy. Gatineau et al. Neoaliturus fenestratus. Alma et al. obsoletus in Lebanon is recorded. Orenstein et al.: Comparison of infection risk by VK in organic and conventional vineyards in Germany.: Only BN identified in Switzerland on cvs Chardonnay.: Survey of GY from 1997 to 2002 on 21 cultivars in 12 Hungarian counties. obsoletus. Maixner et al. BN is the main disease and cvs Chardonnay and Zweigelt appear to be the more sensitive among whiteand red-berried varieties. Detection in December was the more constant. All affected grapevines and two naturally infected periwinkles contained the same stolbur isolate. the incidence of VK was significantly higher in conventional vineyards. Choueiri et al. which is widespread in the province.
The disease was identified in 1995. It is different from FD phytoplasma. Sabaté et al. obsoletus and frequence of C. Living insects checked for transmissibility of phytoplasma by feeding on artificial medium. Main symptoms: Symptoms are similar to those of other GY diseases and may affect the whole plant. obsoletus were positive for stolbur phytoplasma. 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.: Important expression of BN in cv Chardonnay in the Ukraine. O. Šeruga et al. 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). specific association of each isolate with a different weed is discussed. Control: No possibility of control because of erratic transmission by the vector. together with high populations of H. PGY phytoplasma is similar to the Alder yellows phytoplasma that is widespread in black alder in Germany.2003 2003 Petrovic et al. H. PALATINATE GRAPEVINE YELLOWS 1.: Important epidemics of "Schwarzholzkrankheit" (SHK = VK) in Franken (Bayern. -B and C) have been identified. dioica. vinifera seedlings that later developed GY symptoms. Trapping of insects on sticky traps or with D-Vac suction. Attempts to transmit PGY phytoplasma from grapevine to grapevine using reared S. Transmission to plants is not reported. Cicadellidae) trapped on alders and caged on V. 2003 2003 2004 2004 2004 C. Three isolates (PGY-A. using 4 fragments of phytoplasma DNA showed that all isolates are similar. Milkus et al. Langer and Maixner: Three different types of stolbur phytoplasma isolates found in grapevines. and rarely in groups. alni is highly monophagous and specimen do not survive long on grapevine. Several hemipters have been found to deliver stolbur phytoplasma to the medium. It is associated to phytoplasmas in the Elm yellows (16SrV) group.) Detection: Universal primers and RFLP analysis of amplicons can be used to identify a 16SrV phytoplasma. (a): Investigation on the variability of stolbur (16SrXII) phytoplasmas from extracts of different grapevines in Croatia. Other host plants: Alder trees (Alnus glutinosa L. occur most often at the border of plots. Agents: The agent is an EY (16SrV) group phytoplasma. Germany). Varietal susceptibility and sensitivity: PGY occurs in limited areas. 164 . was found carrying the alder phytoplasma at a higher rate than O. arvensis and U. affected plants are not numerous. titanus leafhoppers have failed.: Importance of a GY epidemics in the Drava region (Slovenia) in 2001 and 2002. obsoletus is rare. Description Palatinate grapevine yellows (PGY) is a disease occurring in Germany in Pfalz and Moselle regions (west of Germany) mainly on cv Scheurebe. alni but failed to transmit both to alder and grapevine. Another alder insect.: Search for vectors of stolbur phytoplasma in Cataluña (Spain) where H. Šeruga et al. obsoletus and weeds in different areas in Germany. (b): First identification of BN infection in the Macedonian republic. The latter results have not been published. Transmission: Transmission was obtained with the leafhopper Oncopsis alni Schrank (Hemiptera. mainly on cv Scheurebe. It is not epidemic and occurs most often in old vines in vineyards along creeks or rivers where alders (Alnus glutinosa L) are present. According to the data of field survey. Main synonyms: FD-Pfalz was a first name because the agent was shown to be related to the FD phytoplasma. However. Symptoms were recorded on 26% of 30-year old Scheurebe grapevines and 96% of tested individuals of H. Gilge et al. Alder is considered as the source of PGY infection. the psyllid Psylla alni.
obsoletus. Characterization of the three PGY strains showed that they were closer to FD than to strain EY1 from American elm. molecular characterisation and epidemiology of PGY. Same RFLP profiles of P1-P7 rDNA amplicons obtained from yellows affected grapevines. However. In the same period. using Heteroduplex mobility assay and sequencing of rDNA and FD9 fragment. respectively. Identification of the PGY-associated phytoplasma with phytoplasmas infecting alder in the vicinity of affected vineyards.: Further comparison between elm yellows group (16SrV) phytoplasmas. alni trapped on alders. In 1993. a phytoplasma isolate transmitted from alder to periwinkle in Italy. which was associated with two different phytoplasmas in group 16SrIII and 16SrI (aster yellows). The proportion of infective leafhoppers is lower with O. 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. Description Grapevine yellows (GY) were first reported in New York State (USA) in 1977. the latter findings were not confirmed. Main synonyms: Leaf curl and berry shrivel (LCBS). alni (Auchenorrhyncha. The strong adaptation of O. The differences in vector specificity between PGY and FD phytoplasmas are not supported by molecular data. (b): Transmission to V. 1997 1999 1999 2000 2000 2000 2001 2003 D. alni. Maixner et al. Darimont and Maixner: Comparison of the transmission efficiency of phytoplasma to their host plants and to grapevine by O. It is different from FD phytoplasma but molecular and serological data show a close relationship. Virginia grapevine yellows III (VGYIII) Main symptoms: North American Grapevine Yellows (NAGY) is a lethal disease causing yellowing of the leaves. vinifera seedlings of the PGY phytoplasma using specimen of O. a serious outbreak of GY occurred in Virginia. As much as 5 % of leafhoppers from Palatinate and 15 % from Moselle tested positive. Extensive survey of yellows-affected grapevines in several areas of Germany did not show the presence of other EY group phytoplasmas. 4 Italian and French strains of FD and elm and alder phytoplasmas. The risk of infection by PGY is lower than for VK because reservoir plants are fewer and outside vineyards. Oncopsis alni and Psylla alni specimen. NORTH AMERICAN GRAPEVINE YELLOWS 1. alder. American grapevine yellows. the vector of BN / VK.: Comparison between Elm yellows group (16SrV) phytoplasmas including PGY strains. Reinert: PhD thesis on detection. alni and H. The PGY strains are closest to the ALY phytoplasma from alder in Italy and to one strain of FD. In 1991. (b): Detection of a new EY-related phytoplasma in GY-affected cv Scheurebe in Germany. Historical review 1995 Maixner et al. both insect species trapped on alder. affected vines often do not survive winter frost because they lack reserves. 165 . Cicadellidae) (but not P. Variability analysis and sequencing of the FD9 DNA fragment and homology of the sequence to a part of the SecY gene. Maixner and Reinert: Demonstration that O. titanus living mostly on wild Vitis riparia around affected vineyards were found to carry an agent that was serologically related to that of FD. a phytoplasma belonging to the X-disease group (16SrIII) was detected in affected grapevines in New York. Angelini et al. The alder phytoplasma resembles ALY. alni to alder prevents a frequent inoculation to grapevine even if the infestation of the leafhopper population is high. Angelini et al. Main diseases: Virginia grapevine yellows I (VGYI). Reinert and Maixner: 1997. alni.2. In New York. populations of S. 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.
However. 2. Maixner and Pearson: First data on the study on S. Daire et al. vinifera. titanus in New York and transmission assays of a possible infectious agent. The latter observation suggests that another non-related MLO might have been present. tested positive for a MLO in the X-disease group with PCR-RFLP of 16S rDNA. the survey of vineyards and transmission assays to V. Sauvignon blanc and Cabernet franc. Symptoms. titanus. X-disease phytoplasma was detected in native Vitis sp. It appeared to be spreading and was perhaps insect borne. Chen et al. No transmission by vine planting material reported. it is very severe on Chardonnay vines but was observed on other varieties including Riesling.: 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). Detection: With molecular assays using universal or group-specific PCR primers. ELISA and immunfluorescence were positive from periwinkle. The X-disease phytoplasma occurring in New York was also detected with monoclonal antibodies and with probes for dot-blot hybridization. riparia to V. X-disease phytoplasma was detected in native Vitis sp. as well as direct PCR assays from insects.: Occurrence of FD-like symptoms on White Riesling grapevines in New York.Agents: The 16SrIII phytoplasma detected in NY has not been further characterized. observed in 1983. allowed the identification of a few candidate vectors of the aster yellows phytoplasma. show that the FDU (= FDI) strain from Italy and a MLO detected in a grapevine in Virginia (USA) are both related to X-disease. Historical review 1977 Uyemoto et al. and an aster yellows phytoplasma was detected in numerous weeds and shrubs within and around the vineyards. The FDI MLO was identified in a parallel work as a member of the X-disease group. very similar to those of LCBS. Maixner et al. Adults migrate from V. present in New York and its possible relationship with a GY disease. However. Symptoms may occur for several years in succession in the same vines. 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. The Virginia Grapevine Yellows (VGY) has been associated with the VGYIII phytoplasma (a strain of X-disease or 16SrIII. faba bean plants and feeding solutions. 1985 1991 1993 1993 1993 1993 166 . among which Agallia constricta. and aster yellows was detected from numerous weeds and shrubs within and around the vineyard.: Molecular comparisons between MLOs transmitted to periwinkle from grapevine or detected in a few naturally affected grapevines. Other host plants: Both phytoplasmas are widespread in the USA in weeds and cultivated species.: Two MLOs were transmitted from affected grapevines to periwinkles in Udine (Italy) (= FDI or FDU or GYU) and New York. subgroup III-B) and with the VGYI phytoplasma (a strain of aster yellows or 16SrI. 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. vinifera cuttings. the disease was first observed on cv De Chaunac. Prince et al. Varietal susceptibility and sensitivity: In New York. In Virginia. Larvae are more frequent on wild Vitis riparia at the border of the vineyards. subgroup I-A). then on White Riesling. Probes and primers yielded positive reactions using DNA extracted from grapevine. other grapevines with strong symptoms tested negative. Data showed that the Italian and American MLO were related. Transmission: No vector insects have been identified for VGY phytoplasmas. Pearson et al.: Monitoring of S. Control: There are no control methods since vectors and reservoirs of the associated phytoplasmas are still undetermined.
Beanland and Wolf: Search for possible insect vectors and reservoirs of NAGY phytoplasmas in Virginia. overlay one another like shingles and become leathery and brittle. The relatedness of the associated MLO to X-disease MLO is confirmed.5%) with the phytoplasmas associated to Papaya die back (PDB) in Australia and to Phormium yellow leaf (PYL) in New Zealand. The sequence similarity of the 16S rRNA gene shows that it is close to members of the aster yellows group (16SrI). Restricted Growth (RG). Mapping of diseased plants show that grapevines appear to be a terminal host of AGY. which is the more frequent and the Tomato big bud (TBB) phytoplasma. Agallia constricta is a suspected vector of VGYI (aster yellows phytoplasma). Shoots may display abortion of bunches or berry shrivel later in the season and death of terminal buds. followed by the progressive death of the shoots. The second phytoplasma. It belongs to the peanut witches' broom group (16SrII) and has a broad plant host range in Australia. leaves tend to fall early. suggesting that the source is from an alternative host. subgroup I-A. with a higher incidence in warmer inland districts of New South Wales and South Australia. clover phyllody phytoplasma (16SrI-C) being the closest . consistently associated with AGY symptoms and AGY phytoplasmas but their aetiology is not clear. BVGY phytoplasma has not been clustered in an established phytoplasma group. In addition. 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. a GY disease found in Victoria (Australia). Buckland Valley Grapevine Yellows (BVGY). is a member of the aster yellows (16SrI) group. waxy appearance and remain rubbery. It has been suggested that all three phytoplasmas are strains of the same phytoplasma species. Confirmation that a X-disease (16SrIII) group phytoplasma is involved and classification of the latter phytoplasma into a new subgroup. Conversely. Stems of affected shoots develop a blue. Some plants also display uneven bud development resulting in canes and cordons that are bare in places with little or no bunch development.: A GY disease observed in Virginia since 1983 affects large areas with a low incidence and a low spread rate. 1998 2003 E. The AGY phytoplasma has been designated as Candidatus species "Candidatus Phytoplasma australiense".1993 Wolf et al. The use of PCR has shown the association with AGY of three different phytoplasmas. Late Season Leaf Curl (LSLC). Late Season Leaf Curl (LSLC)-affected grapevines show symptoms in late summer with tightly downward rolled leaves that often remain green. two syndromes with particular symptoms (Restricted Growth and Late Season Leaf Curl) have been found in some cases. Other species tested positive and transmitted to feeding solutions. respectively. Restricted Growth (RG)-affected grapevines show retarded growth resulting in shortened shoots and smaller leaves. The association of AGY with phytoplasmas was confirmed as early as 1988. Main symptoms: All symptoms typical for GY can be found on AGY-diseased plants. Description Grapevine yellows were first reported from Australia in 1976 and called Australian grapevine yellows (AGY) in 1983. AUSTRALIAN GRAPEVINE YELLOWS 1. Early and recent surveys have shown that AGY are found in most viticultural regions of Australia. one node after the other. Both VGYI and VGYIII were found in several weeds and shrubs around and inside the vineyards. Davis et al. III-I. there are reported cases where they were not associated with AGY disease or phytoplasmas. riparia vines. also detected in wild V. It belongs to the stolbur group (16SrXII) but is distinct from the BN and VK phytoplasmas.: Two phytoplasmas are detected in yellows diseased grapevines in Virginia. The TBB phytoplasma has been designated as Candidatus species "Candidatus Phytoplasma australasia". Main diseases: AGY (sensu stricto). 167 . In this particular disease. The yellow leaf tissue can become necrotic. Agents : AGY disease (sensu stricto) is associated with two phytoplasmas: the AGY phytoplasma. The 16S rRNA gene has a high sequence similarity (more than 99.
suggesting that the disease is the result of an aerial transmission and that the phytoplasma was not present in the original planting material. and trunks in October is more reliable. Detection: Detection is obtained with PCR. However. AGY phytoplasmas are common in several crops in Australia. All generic "universal" primers may be used. 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. Control: There are no methods to control AGY diseases in the vineyard.and white-berried varieties.: The "Rhine Riesling problem" denoted "Australian Grapevine Yellows". Remission of AGY. branches. vector of TBB in tomato crops. Cixiidae) is known as the vector of PYL disease in New Zealand but the species has not been reported from Australia. suggesting that this contributes to the decline in phytoplasma numbers and to the remission of symptoms. Magarey and Wachtel: GY symptoms are found in most viticultural areas of Australia.: MLOs. Varietal susceptibility and sensitivity: Chardonnay and Riesling appear to be most often affected. once vines are infected by AGY. Several observations suggest that aerial transmission prevails. Magarey et al. Sanitation with hot water treatment is being developed in Australia. 140-510 nm in diameter. is found in high numbers together with two other potential MLO vectors in vineyards of Victoria (Australia).: Orosius argentatus. Magarey et al. Similarly. Specific primers have been designed on the 16S rRNA gene for detection of AGY phytoplasma (Ca. argentatus. The BVGY phytoplasma has no suspected insect vector. Reduction of symptom expression in vines injected with tetracycline. Phytoplasma australiense) either in single-step or nested PCR following a first amplification with universal primers. found in sieve tubes of diseased vines are associated with AGY symptoms. The incidence of disease may be temporarily high in some vineyards of Chardonnay. It is possible that AGY diseases have become prevalent in Australia in the last years with the increasing planting of Chardonnay. trunks and roots throughout the year and infection is persistent from year to year. 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. The AGY phytoplasma was detected in the common leafhopper Orosius argentatus (Evans) but transmission to grapevine has not been reported. 2. Magarey and Wachtel: Review of the AGY problem. Magarey: Comprehensive review of GY diseases in the world. The TBB phytoplasma is transmitted to other crops by O. RG and LSLC diseases has been observed. RG or LSLC. Magarey: The situation of GY in Australia. but phytoplasmas were also detected in other red. Sampling simultaneously from shoots. Phytoplasma australasia). Recent studies have shown that the latter phytoplasma can be acquired from infected grapevine by O. branches. However. 168 . Other host plants: As mentioned above. Detection can be from shoots. they are at greater risk of displaying symptoms again in the following years. such as Shiraz or Semillon. A "heat curing" effect of AGY disease by hot weather has been observed. Osmelak et al. indicating persistence of the disease. The planthopper Oliarus atkinsoni Myers (Hemiptera. Phloem fluorescence of diseased vines in the UV microscope. reservoirs must be important but the epidemiology is not fully understood. a specific primer in the intergenic 16S-23S spacer region has been designed for detection of TBB phytoplasma (Ca. argentatus and subsequently transmitted to faba bean but the ability to transmit to grapevine was not confirmed.Transmission: No vectors have been identified for any AGY disease. Hence. followed by RFLP analysis or sequencing.
insecticide sprays are useless for controlling the disease. Uneven distribution of phytoplasmas in infected plants. Beanland et al.: Assumption from field observations and monitoring that AGY.: BVGY phytoplasma is associated to GY symptoms that may show remission then reoccurrence but also appear on vines that were not affected previously.1995 1995 1996 Bonfiglioli et al. The problem of symptomlessly infected vines.: AGY phytoplasma detected in the body of Orosius argentatus leafhoppers.: Comparison of visual assessment and diagnostic assays.: Identification of a new phytoplasma in Chardonnay grapevines in Victoria (Australia).: Description of AGY symptoms. Constable et al.: No vector for AGY found. Detection of BVGY in another plot in the same area.: Close relationships between phytoplasmas associated with AGY. Constable et al. It is occasionnally detected in symptomless vines and in vines with LSLC or RG symptoms. Spring and summer are optimal for detection. and LSLC. Constable et al.: Further comparison of AGY phytoplasma with overseas GY phytoplasmas show that it is closely related to. Candidatus Phytoplasma australiense. Constable et al. The TBB phytoplasma is detected occasionnally in vines with AGY symptoms and also in LSLC-affected vines. (a and b): Positive detection of phytoplasma with PCR in AGY-affected grapevines and tentative survey of phytoplasmas in Australia. Wilson et al.: Detection of phytoplasmas associated with AGY.: The AGY phytoplasma is consistently present in the majority of symptomatic grapevines. Davis et al. Hence. 1997 1997 1997 1998 1998 1998 1999 1999 1999 2000 2000 2001 2001 2002 2002 2002 169 . but different from the stolbur phytoplasma associated with BN in France.: Use of hot water treatment in commercial nurseries of Australia. Padovan et al. RG and LSLC are associated.: TBB phytoplasmas can be acquired from diseased grapevine by Orosius argentatus and subsequently transmitted to faba bean. Constable and Symons: AGY phytoplasma is detected in most symptomatic grapevines but also TBB phytoplasma and a third uncharacterised phytoplasma. LSLC would be followed by AGY on the following year and AGY would be followed by RG in subsequent years. Kelly et al. Liefting et al. BVGY might form a new AY subgroup of AY (16Sr) group or even a new phytoplasma group. Habili et al. Padovan et al. Spain and Israel. Gibb et al. resulting in increasing cumulative incidence.: Description of the AGY phytoplasma and designation of a new taxon. (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). later called BVGY phytoplasma.: Study of the phylogenetic relationships of the BVGY phytoplasma with aster yellows and stolbur phytoplasmas. Papaya dieback and Phormium yellow leaf diseases. Bonfiglioli et al. Waite et al. established from a different source of plant material indicate that BVGY disease occurs as a result of aerial transmission. RG. Beanland et al. Constable: PhD thesis on the biology and epidemiology of AGY phytoplasmas.
detection is more reliable from samples taken from branches and trunks. Other varieties are cited in the different situations. TBB phytoplasma may have a role in the aetiology of LSLC. In Chardonnay. The second situation is the significant occurrence of X-disease group (16SrIII) phytoplasmas in countries other than the USA. The subgroup of aster yellows was not determined but primers specific for stolbur tested negative. Statistical analysis showed that RG and LSLC diseases were not always associated with AGY. RG and LSLC symptoms for 3 to 6 years in Chardonnay and Shiraz vineyards. The third situation is represented by poorly characterized GYs recorded from new countries. later designated 16SrXII. However. they seem to have some relevance in Israel and were recently reported from Tunisia. (a and b): Biology and epidemiology of AGYs. a 16SrI-B phytoplasma was found in mixed infection with a stolbur phytoplasma (at that time designated 16SrI-G). No variability was observed amongst BVGY phytoplasma isolates.: Detection of phytoplasma in the aster yellows group in GY affected plants in Slovenia and Croatia. The first is the detection in countries other than the USA. Varietal susceptibility and sensitivity: Chardonnay. Transmission: Aster yellows and. OTHER GRAPEVINE YELLOWS 1. which was the first designation for stolbur phytoplasma in some laboratories. 1997 170 . namely Israel. Constable et al. 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. Symptomatic shoots from AGY affected vines are reliable for detection in summer (January). They must not be confused with the 16SrI-G phytoplasma of earlier reports.: Survey of phytoplasmas infecting grapevine in northern Italy.2002 2003 Crocker et al. recurrence in other vines and new occurrence in previously unaffected vines. However. X-disease group phytoplasmas. a variety widely grown in all countries and very sensitive to all GY agents. In three instances. are very frequent in many host plant species and several insect species are known as vectors or potential vectors in different countries. It seems that aster yellows phytoplasmas have no important significance in European grapevine yellows. These aster yellows phytoplasmas belong to subgroups I-B or I-C (clover phyllody) which are ubiquitous in Europe. The AGY and the TBB phytoplasmas may persistently affect grapevines throughout the year and between years. 2. of phytoplasmas belonging to the aster yellows group (16SrI) in yellows-affected grapevines. Description Besides the cases reported above. three different situations must be described. There is no information of transmission of AGY by planting material. 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. The total cumulative incidence for AGY could be as high as 90%. sometimes as a second phytoplasma in co-infection with FD or BN phytoplasmas and sometimes also in symptomless grapevines. At other times of the year. 2004 2004 F. to a lesser extent. Saric et al.: Survey of AGY.: The management of the area for plant material in the nursery with the scope of hot water treatment. Historical review Europe 1996 Alma et al. Constable et al. is often reported in connection with these cases. Constable and Symons: Study with Heteroduplex mobility assay (HMA) of the elongation factor Tu (Tuf) gene of the AGY and the BVGY phytoplasmas. the diseases are characterized by remission in some plants.
H. including S. M'hirsi et al. Stolbur phytoplasma (16SrXII) was also detected. The spatial and temporal dispersion of the four species was analysed. Circulifer spp. and Anaceratagalia laevis were all found carrying an aster yellows phytoplasma. Tanne et al. 2001 2003 Tunisia 2003 Chabbouh et al. Trapping of hemipters and identification of numerous potential phytoplasma vectors. Detection with PCR show erratic identification of a 16SrI-B phytoplasma. titanus. 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. of phytoplasmas belonging to group 16SrI. Similar transmission of clover phyllody (16SrI-C) MLO by S.: Further survey of GY vectors in the Golan Heights. obsoletus was found in addition to the preceding 5 species. Macrosteles sexnotatus. Gajardo et al. some affected vines showed a sudden fall of leaves in summer. subgroups I-B and I-C and also to group 16SrVIII (Ash yellows) in mixed infection with I-B. a X-disease phytoplasma was detected in some Neoaliturus specimen. Megophthalmus scabripennis was positive for aster yellows phytoplasma. Transmission to periwinkle of yellows diseases using collected insects. Caudwell et al.: Identification in yellows-affected vines of varieties Petit Syrah. Orosius orientalis.: Survey over several years of potential vectors of phytoplasmas in vineyards of the Golan Heights showed that Neoaliturus fenestratus.: First results on the monitoring of GY diseases in Tunisia in imported table grape varieties. Merlot and Carmenere. titanus had been obtained in 1971 in France (cf.: 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). 1971b). Orenstein et al.: Formal identification of a 16SrI-B phytoplasma in GY-affected grapevines in Tunisia. Neoaliturus sp. D'Ascenzo et al. obsoletus and Circulifer haematoceps were all three abundant and positive for stolbur and aster yellows phytoplasmas. H.: Monitoring of potential phytoplasma vectors in vineyards in Israel and molecular detection of phytoplasmas in their body. called "Amarilliamento de Elqui".2001 Alma et al. In addition. 2003 171 . Klein et al..: Experimental transmission of the Chrysanthemum yellows phytoplasma (16SrI-B) to grapevine by four leafhopper species. 2003 South America 1980 Caudwell: Description of a disease of the yellows type in Chile. In addition to similarities of symptoms with FD.
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