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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.
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Boudon-Padieu 2006 .CIHEAM Centre International de Hautes Etudes Agronomiques Méditerranéennes Options méditerranéennes Directeur de la publication: Bertrand Hervieu Directory of Infectious Diseases of Grapevines and Viroses and Virus-like Diseases of the Grapevine: Bibliographic Report 1998-2004 Editors: G. Martelli. E.P.
Martelli. +39 080 542 45 87 e-mail: firstname.lastname@example.org Directory of Infectious Diseases of Grapevines and Viroses and Virus-like Diseases of the Grapevine: Bibliographic Report 1998-2004 Editors: G.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» . 55 ISSN : 1016-1228 ISBN : 2-85352-338-1 © CIHEAM.September 2006 tel.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. Série B : N. Italy. E. 279 Options Méditerranéennes. Boudon-Padieu Bari : CIHEAM (Centre International de Hautes Etudes Agronomiques Méditerranéennes) p.
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 .
by and large. ICVG has a long-lasting tradition in these endeavours. Furthemore. up to date as to the time of publication. to all those interested in viticultural matters. The present issue of Options Méditerrnéennes. 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. in the belief of rendering a good service to the scientific community and. whereas E. produced under the auspices of ICVG. 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. Martelli and E. 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.PREFACE Virus. except when virus-free vines are grown for comparison alongside with non-sanitized sister vines. Bovey The “Directory” is a work of great thoroughness and accuracy. Improving the sanitary conditions of the industry. and practitioners. being a recognized authority in this field. and to the quality and quantity of the crop. these diseases cause loss of vigour and often a decline of affected stocks. it is most unfortunate that dissemination of infectious diseases continues. or induce subtle debilitating effects which are difficult to percieve. hosts a new edition of the “Directory of Infectious Diseases of Grapevines” by G. grouping diseases and setting them in a well thought out order. Boudon-Padieu has a long lasting experience in the study of phytoplasmas. Boudon-Padieu and the latest Bibliographic Report (1998-2004) on “The Viroses and Virus-like Diseases of the Grapevine” by R. taking also into account the future role of the Mediterranean as a free-trade area. Bovey. they reduce graft compatibility of scions and rootstocks. Special care was taken in the organization of the subject matter. 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. is therefore a goal of utmost importance to which the Mediterranean Agronomic Institute of Bari (IAM-B). The “Bibliographic Report” is another most precious achievement by R. Sincere thanks are espressed to the authors of these contributions.P. As to the authors. Italy 7 . that fosters coperative studies in grapevine virology. G. promoting the publication of the Bibliografic Report 1985-1997 on virus and virus-like diseases of the grapevine. to which IAM-B was happy to contribute. whose publication IAM-B is happy to support. students.P. Notwithstanding these efforts and the promoting activity of the International Council for the Study of Virus and Virus-like Diseases of the Grapevine (ICVG). although the technology is now available for detection and elimination of most of the pathogens occurring in propagation material. Cosimo Lacirignola Director of IAM Bari. so as to produce a document which can readily be used by scientits. whose Secretatiat he has admirably conducted since its very establishment in 1962. technicians. is contributing with direct involvement in international research projects and through the activity the Mediterranean Network on Certification of Grapevines. As a whole. which reflect on the commercial value and productive life of the vineyards. a distinguished virologist of worldwide reputation and one of the father founders of ICVG. virus-like and phytoplasma diseases of grapevines constitute a major limiting factor to the development and well-being of the world viticultural industry.
Italy E. France 2006 .P. 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. Martelli University of Bari.
1986. which were compiled under the auspices of the International Council for the Study of Viruses and Virus Diseases of the Grapevine (ICVG). ICVG has been instrumental in fostering basic and applied research in grapevine virology. Caudwell A.. its 14th Conference having taken place in Italy in 2003 (Bovey and Gugerli. Hewitt and R.INTRODUCTION The grapevine (Vitis spp. and supporting initiatives for the establishment and implementation of clean stock and certification programmes. and R. A bibliographic report 1979-1984. phloem-and xylemlimited prokaryotes) play a major role. A bibliographic report 1998-2004. Bibliographie de 1965-1970. grapevines are exposed to the attacks of a variety of pests and pathogens among which infectious intracellular agents (viruses. shortening the productive life of vineyards. The importance of the grapevine industry and the magnitude of the problems caused by these pathogens has generated wide interest which. Bovey R. in turn.) undoubtedly represents one of the horticultural crops most widely grown in temperate climates. Thus. The papers up to 2004 are listed and commented upon in six bibliographic reports: Caudwell A. 1979. Hewitt W. Bovey R. ICVG has issued the recommendations that follow: The International Council for the Study of Virus and Virus-like Diseases of the Grapevine (ICVG). 76 pp. The viroses and virus-like diseases of the grapevine. recognizes that a number of the 60 or so infectious agents (viruses. Les virus de la vigne. A short history of ICVG. Vitis 11. 10-172. Office International de la Vigne et du Vin. 2006. viroids. as well as on the quality and quantity of the yield. nurserymen. The viroses and virus-like diseases of the grapevine. 2003.B. From the very beginning. Vitis 25. Vitis 18. Martelli. Since its foundation. among other things. 303-324. 2003). has produced an impressive series of papers which now number about 5. Bovey. and a highly valuable agricultural commodity. Gugerli.. and P. Bovey R. 29 (Series B. Extended Abstracts 14th Meeting of ICVG. that has been especially active at the international scale from the late 1950's onwards. and G. 227-275. and phytoplasmas) recorded from the grapevine can be highly detrimental to this crop.400. 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. Locorotondo 2003. all efforts should be made to improve its sanitary conditions.P. 11 . 205-279. 3rd part). growers. 3rd part). The increased attention paid to grapevine virological problems and the like. Paris. The viroses and virus-like diseases of the grapevine.. having a negative impact on the plant vigour and longevity. Options Méditerranéenes.B.. has fostered intensive research. 316-376. attracting the attention of scientists. and administrators on the detrimental effects of infectious diseases on the well-being of the industry. 1972.. Options Méditerranéenes. A bibliographic report 1985-1997. W. Bovey. As most of the vegetatively propagated crops. Bovey R. Infected propagating material is largely responsible for the spread of diseases among and within viticultural countries. and endangering the survival of affected vines. A bibliographic report 1971-1978. ICVG has met regularly every 3 to 4 years. 1-2. To this effect. causing heavy losses. 1999. viroids. xx (Series B. 1965. The viroses and virus-like diseases of the grapevine. Bibliographie des viroses de la vigne des origines à 1965.
(issued and approved in 1997 at the 12th ICVG Meeting. USA. Improvement of the sanitary level can be achieved through selection and sanitation.P. 1980. Their elimination from mother vines intended for propagation should therefore be pursued.F. Thus. and 3). Editions Payot. W. In order to preserve valuable grape clones and varieties. technology should make it possible to exclude additional disease-causing viruses from the certified stock. 93 pp. Uyemoto. and permit tracing the certified grapevines to the originally selected and tested plants. Set 1 (O. (a 2nd revised edition edited by W.The presence of diseases such as infectious degeneration. high standards are essential to ensure that no viticultural area is compromised by the introduction and spread of diseases. The certification process should specify conditions to prevent and detect subsequent infection of nursery plants by regulated pests. as Extended Abstracts. Infected propagation material is largely responsible for the spread of diseases among and within viticultural countries. (a revised edition by J. W. Tolin. all efforts should be made to improve its sanitary conditions.P. They represent a most valuable source of information. The American Phytopathological Society Press. we propose two sanitary classes. which are best performed in the framework of certification programmes encompassing also clonal selection. ensure clonal integrity. Rowhani will be published in 2005 in the series APS Plant Virus Image CD-ROM.B. Goheen and H. and fleck. (issued and approved in 2003 at the 14th ICVG Meeting. Grapevine (Vitis) virus and virus-like diseases. Uyemoto will be published in 2005). Class 1 should include only grape nursery stock which tests negative for the most damaging diseases/pathogens. Lisbon. It would move freely between regulatory boundaries. Inadequate certification standards have repeatedly resulted in disease problems for growers and nurserymen. Compendium of Grape Diseases. bois noir. 12 . Minnesota. In addition. 2. a moratorium will be established for these viruses. rugose wood. Locorotondo. Virus-like Diseases and Other Disorders).D. As efforts are made to harmonize grapevine certification protocols. Certified grapevines are derived from pathogen tested. Bovey R.K. the virological problems of grapevines have been extensively treated and illustrated in a number of books and major review articles: Uyemoto J. G. leafroll disease and associated closteroviruses (grapevine leafroll associated viruses 1. 29 pp.A. 100 slides. Hewitt. 1978. The agents that should be controlled by the Class 1 certification program are those associated with infectious degeneration and grapevine decline (nepoviruses). Goheen. having a negative impact on plant vigour and longevity.K. Plant Virus Slide Series. many of which can be highly detrimental to this crop. Gubler and J. Martelli and A. including the causal agents of fleck and rupestris stem pitting. lately.. W. Grape Section.K. Wilcox. Clemson. and phytoplasmas (flavescence dorée. viroids and phytoplasmas). regional viticultural conditions. recognises over 70 infectious agents affecting grapevine (viruses.C.C. Barnett and S. Martelli. as well as on the quality and quantity of the yield. Dias. No other stock should move outside regulatory regions. 1988.F. Virus and Virus-like Diseases of Grapevines. R. Portugal ) The International Council for the Study of Virus and Virus-like Diseases of the Grapevine (ICVG). Pearson R. G. and the need for preservation of heritage germplasm. In the future. Certification of grapevine nursery stock is a powerful and effective tool to control these agents. under the name of Grapevine Viruses. Lausanne. 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.W. Paul. A. G. Certified selections should be tested for specific pathogens. eds) Clemson University. Woodham. Gärtel. and other grapevine yellows). rugose wood (GVA. St. leafroll. Martelli and A.C. Until that time. GVB and GVD).G. is regarded as incompatible with an accepted sanitary status. Vuittenez. The regional certification standards for Class 2 stock should be created at a local level based on the rate of endemic infection. Italy) The Proceedings of all the international ICVG Conferences have been published in extenso or. 181 pp. However. valuable grape genetic resources exist which are infected with virus but are essential to the preservation of world viticultural heritage.P. clonally selected primary sources. and A. that enables vineyards to economically and sustainably maintain quality and productivity.
INRA Editions. sélection pomologique. Graft-transmissible Diseases of Grapevines. Italy.Université de Bourgogne Dijon. 192 pp. Hamze and Mr. Bulletin de l'OIV 69. Rome /International Board for Plant Genetic Resources. Walter B. M. n° 86. Martelli. 1993.P.P. Director of the Mediterrenean Agronomic Institute of Bari for supporting the publication of this work. France 13 .K. B. Koganezawa. the Steering Committee of ICGV decided to update the Directory and to entrust this task to G.P.Frison E. and B. Paris. 1997.P. 1996. Walter B. Ridé. Paul. R. Taylor. Scott.A. Martelli G. Boudon-Padieu. 1999. C. 1ere partie: Effets des viroses sur la culture de la vigne et ses produits. University of Bari. 263 pp. (ed. Giovanni P.H. They express their deep gratitude to Prof. 945-971. Martelli and published under the auspices of the Mediterranean Fruit Crop Improvement Council (MFCIC). 2ere partie: Sélection sanitaire. a body now estinguished. Walter. 261-276. respectively. In: Plant Virus Disease Control (A. M. Protocols for detection of viruses and virus-like diseases: Les Colloques. eds. Lacirignola. (ed). Martelli Professor of Plant Virology.A. 1997. Sanitary selection of the grapevine.R. Rome. and R. Bovey and G. Food and Agriculture Organization of the United Nations. Hadidi. Virus certification of grapevines. Bulletin de l'OIV 70. Elisabeth Boudon-Padieu Directeur de Recherches Head Biologie et Ecologie des Phytoplasmes Plante Microbe Environnement INRA . Collingwood. and G.P. Rezaian and R. Sélection sanitaire de la vigne: sélection sanitaire et sélection pomologique. Ikin. Hervieu. 1998.S. St. E.). Paris.P. Notwithstanding this wealth of published information a "Directory of Major Virus and Virus-like Diseases of Grapevines" was compiled in 1992 by R. of the Centre International des Hautes Etudes Agronomiques Méditerranéennes. Maladies à Virus.137 pp. Krake L. Influence des viroses et qualité. Detection and Diagnosis of Graft-transmissible Diseases of Grapevines. FAO/IBPGR Technical Guidelines for the Safe Movement of Grapevine Germplasm. bactéries et phytoplasmes de la Vigne. American Phytopathological Society Press. CSIRO Publishing. Bari.. Chairman of the Board of Directors and Secretary General. Walter B. Sélection sanitaire de la vigne: sélection sanitaire et sélection pomologique. Influence des viroses et qualité. 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.). Boudon-Padieu and M. H. Bordeaux. Walter B.CNRS . 5-23. and G. Martelli G. FAO Publication Division. Khetarpal. Editions Féret. At the 14th ICVG Meeting. Martelli. 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.. 54 pp. Martelli and E. 225 pp. 2000. 1991. and to Dr. Rome. N.
INFECTIOUS AGENTS OF GRAPEVINES More than 70 infectious agents among viruses (58). This represents the highest number of intracelluar pathogens ever found in a single crop. and insecttransmitted xylematic bacteria (1) have been recorded form grapevines. The viral scenario of Vitis: viruses and their taxonomic affiliation(a) FAMILY GENUS SPECIES A. phytoplasmas (8). 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 . viroids (5).
8Ith Report of the International Committee on Taxonomy of Viruses. M.A. Elsevier/Academic Press. Desselberger. 2005. (eds).A. C. Virus Taxonomy. (a) 16 . London. Taxonomically unassigned viruses Unnamed filamentous Grapevine Ajinashika virus (GAgV) Grapevine stunt virus (GSV) Grapevine labile rod-shaped virus (GLRSV) Raspberry bushy dwarf virus (RBDV) Strawberry latent ringspot virus (SLRSV) Sowbane mosaic virus (SoMV) Tobacco mosaic virus (TMV) Tomato mosaic virus (ToMV) Scientific names of definitive virus species are written in italics. L.. Mayo..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. The names of tentative species are written in Roman characters. Maniloff. 1258. pp.M. J. The updated taxonomy of all classified grapevine viruses can be found in: Fauquet. U.. Viruses belonging to unassigned genera Idaeovirus Sadwavirus Sobemovirus Tobamovirus C. and Ball.
INFECTIOUS DEGENERATION .
the yellowing fades rapidly and the canopy develops a normal green color. low yields and low fruit quality. GFLV and several of the other grapevine-infecting European nepoviruses have distorting and chromogenic strains and may occur in mixed infections. Their economic impact varies with the tolerance of the cultivar to the viruses.e. and xylem cells. Description Fanleaf is the oldest known and one of the most important and widespread virus disease of the grapevine. radial bars crossing the lumen of epidermal. These structures are readily visible by light microscopy in lignified shoots. Bunches are smaller and fewer in number. Infectious malformations are induced by virus strains causing distortions. Occasionally. and inflorescences). Foliar symptoms develop early in the spring and persist throughout the vegetative season becoming less distinct in summer.). 19 . puckered. Main synonyms: court-noué. vein banding symptoms were shown to be caused by a co-infection by Grapevine yellow speckle viroid and GFLV. deep lobes. fasciations. phloem. shoots. Fruit set is poor. especially in the basal internodes. and grapevine leaves with typical symptoms were found in herbaria established before the introduction of American rootstock hybrids. dégénérescence infectieuse (Fr. Shoots are also malformed. short internodes. are small-sized and set poorly. showing abnormal branching. Main symptoms: Two distinct syndromes caused by different strains of the causal agent characterize this disease. another disease sometimes occurring in vineyards affected by infectious degeneration. Often infected grapevines occur in patches. which exhibit widely open petiolar sinuses and abnormally gathered primary veins. mosaico giallo. shortened productive life. records of this disease date back some 150 years. The name of this virus comes from the peculiar malformation of infected leaves.). arricciamento. Tolerant cultivars produce fairly good crops whereas sensitive cultivars are severely affected. roncet. Symptomatic leaves show little malformation. but bunches are small and few. sometimes appearing as rings or lines to extensive mottling of the veins and/or interveinal areas to total yellowing . Gelbmosaik (Germ. and acute denticulations. With increased ambient temperatures during summer. FANLEAF 1. are diagnostic of grapevines infected by GFLV. Now the disease is known to occur worldwide. and decreased resistance to adverse climatic factors. low proportion of graft take. In the European literature.INFECTIOUS DEGENERATION (GRAPEVINE FANLEAF VIRUS) Several nepoviruses infect grapevines in Europe and the Mediterranean area. or indistinguishable from those of fanleaf. degenerazione infettiva (Ital. i. however. or endocellular cordons. More recently. Yellow mosaic is induced by chromogenic virus strains. reduced rooting ability of propagation material. asymmetrical.). The foliage develops bright chrome yellow discolorations early in the spring that may affect all vegetative parts (leaves. The consensus is that fanleaf degeneration may have existed in the Mediterranean Basin and the Near East since the earliest time of grape cultivation. chlorotic mottling may accompany foliar deformations. Leaves are variously and severely malformed. This disorder was first reported from California as a syndrome elicited by a specific strain of GFLV. urticado (Port. bunches are straggly. parenchyma.and zigzag growth.). may show open petiolar sinuses. and berries ripen irregularly. double nodes. that give the leaf the appearance of an open fan. panachure. Chromatic alterations of the leaves vary from a few scattered yellow spots. showing progressive decline. tendrils. causing degenerative diseases whose symptoms are similar to. a disorder induced by Grapevine fanleaf virus (GFLV). The characterizing symptoms of "Vein banding". The foliage and shoots show little if any malformation. 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. Reisigkrankheit (partly). Trabeculae. and the yield may be much reduced.
M. use of fumigants is more and more questioned for environmental reasons and is being progressively banned. GFLV was the first grapevine virus to be recovered by mechanical inoculation and to be thoroughly characterized physicochemically and molecularly. quinoa. In the laboratory. There are conflicting reports on seed transmission in grapevines.g. 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. 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. which are desirable as they cause no harm to human health. 20 . However. but is not a specific test. Detection of trabeculae can give information on the health of American rootstocks. Work is under way in different laboratories to create GFLV-resistant rootstocks or cultivar through traditional breeding methods or genetic transformation technology. the endosperm of grapevine seeds. rotundifolia hybrid rootstocks (e. Virus particles are often within tubular structures that accumulate in bundles in the cytoplasm or nucleus. although some like Vitis labrusca can be infected. Control: Use of virus-tested scionwood and rootstock material in the framework of clean stock or certification programmes. index (10 individuals) by ELISA and in single nematodes by RT-PCR and immunosorbent electron microscopy. RT-PCR and immunocapture RT-PCR are becoming increasingly popular. Gomphrena globosa). both required for infectivity. Natural GFLV infections have been detected in weeds in Hungary and Iran. but is not reliable. Specific transmission by X. Molecular assays using radioactive or digoxigenin-labelled probes. Resistance to X. For transformation.g. This resistance is controlled by two unlinked recessive genes. index is determined by the viral coat protein. vinifera x M. and is transmitted through seeds of C. vinifera. Observation of symptoms in the field is useful as a first step in selection. Varietal susceptibility: Almost all known Vitis vinifera L. mannose and xylose. reorganization. but show few symptoms. rotundifolia hybrids is thought to be controlled by a single dominant gene. amaranticolor. vuittenezi has been suspected but not proven. and redistribution and are thought to be sites of viral polyprotein processing and RNA replication. O36-16) show interesting levels of field resistance to GFLV. varieties are susceptible. GFLV can be transmitted by mechanical inoculation from infected grapevine tissues to various herbaceous hosts (e. and transmission by X. index feeding. GFLV has been detected in small groups of viruliferous X. In contaminated soils. consisting of two functional molecules 7342 nt (RNA-1) and 3774 nt (RNA-2) in size. Cytopathology: GFLV elicits the formation of intracellular cytopathic structures known as vesiculatevacuolate inclusion bodies which are often apposed to the nucleus. Detection: ELISA using polyclonal antisera and monoclonal antibodies is a quick. Virus elimination is readily achieved from vegetating shoot tips by heat treatment (38-40 °C for as little as four weeks).Agent: Grapevine fanleaf virus (GFLV) is a nepovirus with polyhedral particles of about 30 nm in diameter. C. the use of fumigants against nematode vectors gives only a temporary but economically valuable control of the disease. encapsidated in different particles. and soybean. cheap. index in V. amaranticolor. where they occur in a radial orientation. Some V. or by somatic embryogenesis. Indexing on Vitis indicators by grafting takes a lot of time and field or greenhouse space. Muscadinia (Vitis) rotundifolia and Vitis munsoniana are highly resistant to X. The virus occurs in the pollen of infected grapevines and herbaceous hosts. but it is still regarded as necessary for confirming freedom from virus infection. Transmission by Xiphinema italiae has not been consistently documented. a number of selectable marker genes toxic to non engineered vines are used. in a persistent manner by the longidorid nematode Xiphinema index feeding on the roots of grapevines and retaining the virus for several months. Nematode populations transmit local virus isolates with a higher efficiency than those from other geographical areas. A satellite RNA 1114 nt in size is associated with some virus isolates. rotundifolia can be infected by GFLV when graft inoculated. rupestris x M. with variable levels of sensitivity. Indexing on herbaceous hosts by mechanical inoculation requires climatized greenhouses and is less reliable than ELISA. Chenopodium quinoa. These inclusions derive from membrane proliferation. by in vitro meristem tip culture. Transmission: At a site. are toxic to many plants but not to V. and very sensitive method. Positive sense single-stranded RNA genome. RT-PCR is estimated to be four to sixfold more sensitive than ELISA. However. serologically rather uniform and occurring as a family of minor molecular variants. The best antigen sources for serological diagnosis are leaves collected in spring or cortical shavings from mature dormant canes. Dissemination over medium and long distances is through infected vegetatively propagated scionwood and rootstocks. but resists infection when the virus is transmitted by the nematode. However. American rootstocks are also susceptible and are generally very sensitive. C.
Montpellier. 871 pp. Healthy rooted cuttings or seedling of Rupestris du Lot were contaminated: 1. Historical review From the late 1800 to 1997. 2. Description of grapevine degeneration in Frontignan (France) Rathay: Description of fanleaf disease from Austria (Zwiewipflereben) Ruggeri : Description of fanleaf disease from Italy (Roncet) Cholin: Description of fanleaf disease from Germany (Reisigkrankheit) Baccarini: First suggestion that fanleaf may be due to a virus. Les maladies et les parasites de la vigne. Hypothesis about a possible role of phylloxera as a vector. Report on first experiments on heat treatment of grapevine in order to eliminate fanleaf. 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. 1977(b) 1865 1882 1895 1896 1902 1906 1910 1912 1912 1917 1918 Cazalis-Allut. but only circumstantial evidence. Tome 1: Les maladies dues à des végétaux (champignons. France. Petri: Grapevine "arricciamento" (fanleaf) has a viral origin Arnaud and Arnaud: Hypothesis of a viral origin for grapevine court-noué (fanleaf). For a comprehensive review on early observations. No conclusive results were obtained. (b) Galet P.2. (a) See references in the Introduction. see the book by Galet. Branas et al. No direct proof of transmission by this aphid. 21 . 1954 1958 Hewitt: Review on grapevine virus and virus-like diseases found in California. With roots of fanleaf-infected grapevines with phylloxera feeding on them. Imprimerie du "Paysan du Midi".. 1929 1931 1937 1937 1946 1950a. as well as on controversies about transmission by phylloxera. 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. bactéries. Branas et al. viroses et phanérogames).b Hewitt: Fanleaf and yellow mosaic recorded from California. Hypothesis that fanleaf is a fungal disease. 1977.: Experiments on the capacity of phylloxera to transmit fanleaf. With individual phylloxera (radicicolous or gallicolous) fed on infected vines. the ICVG Bibliographic Reports (a) have recorded more than 1000 papers dealing with fanleaf. Bovey: Review on grapevine virus and virus-like diseases. 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. research and hypotheses on fanleaf. Arnaud: Court-noué is considered as a soil-borne virus disease.: Hypothesis that court-noué (fanleaf) is caused by a virus transmitted by phylloxera. 3. With soil containing phylloxera.
whereas insecticide treatment has no effect. Control of the vector by soil fumigation. index. yellow mosaic and ArMV 1960a 1960b 1961 1961 1962 1962 1963 1963 1963a. 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.: Transmission of GFLV by Xiphinema italiae in Israel. Goheen and Hewitt: Description of vein banding as a GFLV-induced disease Dias: Host range and properties of fanleaf and yellow mosaic viruses.b Martelli and Hewitt: Comparative studies showed that Californian and Italian GFLV isolates are the same. index in France. Reproduction of fanleaf symptoms in mechanically inoculated grapevine seedlings is confirmed Galzy: Heat treatment of grapevine plantlets grown aseptically in vitro. Das and Raski: Studies on the relationships of GFLV with its vector X. Dias and Harrison: Relationships between the viruses causing fanleaf. Hewitt et al. index. Dichloropropane-dichloropropene (DD) at 1000 l/ha gave satisfactory results. Bercks: Research on the use of three serological methods for detecting plant viruses. Vuittenez: New observations on the effects of soil fumigants on fanleaf in contaminated soils. Hewitt et al. latex test and barium sulfate test. index occurred during the 6 year period of observation. Bercks and Querfurth: Use of latex-test for detecting GFLV and other nepoviruses in grapevine tissue extracts in Germany. Transmission of fanleaf virus by X. Yield was increased by 400 % in comparison with that of untreated controls. Reproduction of fanleaf symptoms in mechanically inoculated grapevine seedlings 1964 1964 1965 Taylor and Hewitt: Description and characterization of Australian isolates of GFLV.: Investigations on grapevine virus diseases in California. including fanleaf virus: bentonite flocculation test. Cohn et al. amaranticolor is confirmed. and no reinfestation by X.: Detection of GFLV particles in thin-sectioned grapevine root tissues.: Fanleaf virus is transmitted by the nematode Xiphinema index Cadman et al. 1965 1967 1968 1968 1969 1969 1970 22 .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.: Transmission of fanleaf virus from grapevine to herbaceous hosts by mechanical inoculation and preliminary characterization of the virus. Serological relationship with ArMV reported. Vuittenez: Mechanical transmission of fanleaf virus to Chenopodium quinoa and C. Goheen et al.: Description of the Davis method of heat therapy of grapevines. Discussion on the possible role of weeds in the epidemiology of the disease. Boubals and Dalmasso: Experiments on soil disinfection against X. Graniti and Russo: A light microscope and cytochemical study of endocellular cordons. Gerola et al. Description of the chipbudding method for indexing. Brückbauer and Rüdel: The virus (or viruses) of Reisigkrankheit (GFLV and/or other nepoviruses) are seed transmitted in some herbaceous indicator plants.
George for detecting GFLV. During the moult of the nematode.: Comparison of indexing by mechanical inoculation to Chenopodium quinoa and by graft-transmission to V. Raski et al. this lining is shed and ingested in the intestine.: Control of fanleaf by soil fumigation with 1. quinoa.3-dichloropropene or methyl bromide.: Detailed physico-chemical characterization of GFLV. Taylor: Review paper on grapevine vein banding. yellow mosaic and vein banding) are transmitted in the same way by X. Walter et al. rupestris St. Vineyards replanted in contaminated but treated soils remained healthy for at least 5 years. Mur et al. Raski et al. Both methods give satisfactory and similar results. The acquisition time threshold is less than 5 minutes.1970 1970 Hewitt et al. Raski and Schmitt: Progress in the control of the fanleaf-nematode complex by soil disinfection with 1. A dormant bud of the variety to be cured is grafted onto a healthy potted rootstock. Alfaro and Goheen: The different strains of fanleaf virus (fanleaf sensu stricto.: GFLV and marbrure (fleck) are not transmitted through the seeds of grapevine. 1970 1970 1970 1971 1971 1972 1972 1973 1973 1973a 1973b 1974 1974 1975 1976 1976 1977 1979 1979 23 . Dias: Review paper on grapevine yellow mosaic. quinoa. George. Querfurth and Paul: Protein A-coated latex-linked antiserum (PALLAS) method for detecting GFLV and other viruses. anterior oesophagus and oesophageal bulb of their nematode vectors. Bercks: Serological detection of grapevine viruses in West Germany. Uyemoto et al.: Use of green grafting as a quick and secure method for graft-indexing. Hévin et al. index. PALLAS is quicker and cheaper than ELISA. rupestris is more accurate than mechanical transmission to C. St. After bud take. Indexing by budding on V.3 dichloropropene or methyl bromide. but ELISA is more sensitive. Vuittenez: Review paper on grapevine fanleaf. especially with low titre antisera. 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. Goheen and Luhn: New method of heat therapy. Both tests are more sensitive than mechanical inoculation to C. The sensitivity of the latex test is increased. the plant is placed in a heat cabinet for treatment Hévin et al. index.: 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. 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. Van Velsen and Niejalke: Green budding for indexing grapevine with the indicator cvs.: GFLV particles observed in the lumen of the oesophagus of X.: Comparison between PALLAS latex test and ELISA for detecting GFLV in France.: Heat therapy of grape plantlets grown in vitro causes changes in some characteristics of the variety. Mission or LN 33.
: Experiments with systemic nematicides for controlling X. Bouquet: Muscadinia rotundifolia becomes infected by GFLV when the virus is transmitted by grafting but resists infection when transmission is by X.: Comparison of polyclonal and monoclonal antibodies for detecting fanleaf virus with ELISA in various grapevine tissues. index by ISEM Bovey et al. Morris-Krsinich et al. index by ELISA.3-dichloropropene failed to eradicate either nematodes or fanleaf virus from the soil but reduced the incidence of the disease to acceptable levels. 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. Savino et al. Hafez et al.: Detection of fanleaf virus and other sap-transmissible viruses in grapevine tissues by ISEM. a very common species in vineyards of Rheinhessen and Palatinate. A Middle Eastern V. Russo et al. Bouquet: Serological detection of GFLV in its vector X. index feeding.: Use of systemic nematicides for the control of X. That X.: Detection of fanleaf virus in grapevine tissues by ELISA and ISEM in different periods of the year. These are promising sources of germplasm for obtaining resistant rootstocks. as vector of GFLV.: Identification of several Vitis species and interspecific hybrids resistant to fanleaf virus. although it is not resistant to the virus itself. Forty days of treatment.: 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.3-D) or methyl bromide applied 75-90 cm deep with 90 cm spacing for 1. vinifera accession represent an excellent example of host plant resistance to GFLV. Rüdel: Discussion on the possible role of X. RNA-2 contains the cistron coding for the viral coat protein. the possibility that a few X.3-dichloropropene (1. vuittenezi. Carbon disulfide gave less satisfactory results. Bouquet: M. index and fanleaf in grapevines. especially in wood shavings of dormant canes during winter.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. Transmission trials gave a few positive results. Even in the cases where the virus was transmitted. Vuittenez: Review on serological methods of detection and identification of grapevine viruses. vuittenezi population used for the experiments could not be entirely ruled out.: 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. Krake and Woodham: Possibility that the agent of yellow speckle is involved together with GFLV in the etiology of vein banding. Walker et al. index larvae were present in the X. index.1979 1980 1980 Kalasian et al. Raski et al. index. The use of methyl bromide requires a continuous cover with polyethylene sheeting for some time after the treatment. index. Methyl bromide and 1. Raski et al. Efficiency of both methods is compared. 1980 1980 1980 1981 1981 1981 1981 1983a 1983b 1983 1983 1983 1985 1985 1986 1986 24 . Brown and Roberts: Detection of fanleaf virus in its vector X. in California.: Soil fumigation with 1. index.: Identification of a natural serological variant of GFLV from Tunisia Huss et al. vuittenezi might be a vector of GFLV is therefore uncertain. rotundifolia is resistant to fanleaf virus transmission by X. Lear et al. Monette: Heat therapy of GFLV and ArMV--infected grapevines with alternating temperatures.
1987 1987 1987 Huss et al. RpRV.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.: Improvement in the serological detection of GFLV and ArMV viruses using monoclonal antibodies. Serghini et al. rotundifolia showed good resistance to GFLV in California. Viruliferous nematodes were crushed in standard extraction buffer and tested in batches of 1-50 by means of DAS-ELISA. vinifera x V.: 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. and has strong homologies with the satellite RNA associated with ArMV.: Study of interactions between GFLV and ArMV isolates grown in C. cultivation of non-host plants and organic soil amendments are recommended. Etienne et al.: Detection of GFLV in the vector Xiphinema index by ELISA. the latter being especially damaging on the variety Kerner. Fuchs et al. quinoa and transmitted by heterografting to Vialla and Kober 5BB rootstocks.: Determination of the nucleotide sequence of the satellite RNA (RNA-3) of GFLV.: Production and use of monoclonal antibodies to GFLV. ArMV.: Use of green grafting technique for sensitive and quick GFLV detection under greenhouse conditions. Treated vines yielded more for over 4 years. Reliable results were obtained with samples of 20-50 nematodes. Lázár et al.: Determination of the complete sequence of GFLV RNA-2. Mild and severe strains were discriminated on the basis of field performance of infected Vitis. RNA-3 encodes a non structural protein. Rüdel: Severe restrictions set on the use of soil fumigants in West Germany for environmental reasons make control of "Reisigkrankheit" very difficult. Catalano et al. No eradication was obtained. Long term fallow (about 5 years). Walter et al. index and GFLV. Previous experience showed that 1. Walker and Meredith: Identification of two accessions of Vitis vinifera resistant to GFLV. RpRSV and ArMV are common. Walker et al. TBRV and leafroll from infected grapevines by in vitro meristem tip culture. Raski and Goheen: Comparison of 1.: Identification of a satellite RNA of GFLV.: Evidence of a differential efficiency of GFLV transmission by Xiphinema index populations from different geographical origins. Martelli and Taylor: Review article on nematode-transmitted viruses.: Two rootstock selections derived from crossings V. 1988 1988 1988 1989 1989 1989 1989 1989 1990 1990 1990 1990 1990 1991 1991 1991 25 . GFLV. Altmayer: Elimination of GFLV. The selection of resistant cultivars and rootstocks is considered of primary importance. Walter and Etienne: Detection of GFLV in wood shavings of dormant canes. Pinck et al.3-dichloropropene and methyl bromide for controlling X. Walter et al. Rüdel: Review on the most important virus diseases of grapevines in West Germany. Effect on yield and economic importance. SLRV. Resistance is controlled by two unliked recessive genes Walter et al. Catalano et al. Positive. but less consistent results were obtained with 1-10 nematodes. Mild strains were shown to confer protection towards severe challenging strains in Chenopodium and grapevine. Treatments with soil fumigants are no longer permitted in Germany.
: Use of modified GFLV coat protein genes for transformation of different Nicotiana species for inducing resistance. index.: Co-inoculation of C.: Development of cDNA probes to GFLV genomic and satellite RNAs and their use for virus detection directly in grapevine extracts.: Detection of GFLV in X. Satellite RNA appears to have a modulating effect on virus pathogenicity.1991a 1991b Fuchs et al. GFLV is eliminated from somatic embryos obtained from tissue cultures grown at 35 °C 1991 1991 1992 1992 1992a. which is also resistant to phylloxera. index by biotin-avidin ELISA Bardonnet 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. Fuchs et al. Ritzenthaler et al. quinoa with biologically active transcripts of GFLV F-13 satellite RNA and GFLV strains devoid of satellite. Esmenjaud et al. hybrids and breeding stocks after infection of the roots by means of viruliferous X. Horvath et al. Spielmann et al. Both became infected in the course of a 12-year trial but had no reduced crop yields. Esmenjaud et al.: Genomic RNA-1 of GFLV is completely sequenced and its genetic organization determined. identification of their replication determinants and evidence of replication in Chenopodium quinoa protoplasts 1993 1993 1993 1993 Martelli et al. Viry et al.: Production of biologically active transcripts from cloned cDNA of genomic RNAs of GFLV. index infested soils.:Detection of GFLV in single nematodes by RT-PCR. Staudt and Weischer: Vitis rotundifolia and Vitis munsoniana resist infection by GFLV transmitted by X. Walker et al. O39-16 in particular. 1993 1993 1993 1993 1993 1994 1994 1994 1994 26 . GFLV is a quasispecies occurring in the field as a series of minor molecular variants.b Hans et al. Goussard and Wiid: First application of somatic embryogenesis for sanitation of grapevines.: GFLV satellite RNA detected in 5 of 34 virus isolates from different geographical locations. index. This represents the first substantiated record of a natural GFLV infection in hosts other than Vitis.: In naturally GFLV-infected vineyards the hypovirulent ArMV A1 isolate induces delayed infection by GFLV Saldarelli et al. thus qualifying for use in X.: GFLV isolated in Hungary from naturally infected symptomatic plants of Aristolochia clematis and Lagenaria siceraria turbinata.: Production of GFLV satellite RNA transcripts.: Two Vitis vinifera x Muscadinia rotundifolia rootstock hybrids (O39-16 and O4343) grafted with Cabernet sauvignon showed a high level of tolerance to GFLV. Staudt: Study of the spread of GFLV in several Vitis species.: Evidence that transgenic tobacco plants expressing the coat protein of GFLV are protected from GFLV infection. Walter et al.
: Attempts to characterize molecularly X. including the two accessions reported as resistant by Walker and Meredith (1990).: Genetic transformation of American roostocks with the coat protein gene of GFLV for resistance induction. Belin et al. Pfeiffer et al.: Transformation of Nicotiana species and Vitis rupestris with different virusderived (coat protein. index feeding. Krastanova et al. Walker and Jin: V.: Genetic transformation of Vitis vinifera with the coat protein gene of GFLV for resistance induction. index. Ritzenthaler et al. Rowhani et al. 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. Of 531 accessions of European. Fuchs: Review article on genetic transformation of grapevines for resistance to GFLV and other pathogens. : Identification of RNA2-encoded proteins in the specific transmission of GFLV by X. 1997 1998 1998 1999 1999 2000 2000 2000 2000 2001 2002 2003 2003 2003 27 . Spielmann et al. Pfeiffer et al. Naraghi-Arani et al. American. RNase L) genes for inducing resistance to GFLV.: Variations observed following RT-PCR and RFLP analysis of the coat protein gene of nine GFLV isolates grown in different hosts confirm the quasispecies nature of this virus. rotundifolia hybrids show high resistance to X. Ritzenthaler et al. Pinck: A comprehensive review of the molecular aspects of GFLV genome and its replication strategy. except for four. Gaire et al.: Up to date account of GFLV replication strategy. rupestris x M.: Demostration that the movement protein of GFLV is located on the intracellular tubular structures containing rows of virus particles. This resistance is controlled by a single dominant gene.: Identification of membranes derived from the endoplasmic reticulum as sites of GFLV replication. Gölles et al. all were susceptible to the virus.: 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.: Demonstration that a 28 kDa protein coded by GFLV RNA-2 is involved in the replication of this RNA.: Identification of the molecular signal accounting for the systemic spread of GFLV in infected hosts. 5 oligoadenylate synthase. Mauro et al. The level of resistance obtained looks promising. Martelli and Walter: Review article on certification of grapevines. De Luca et al.: Development of a GFLV detection system based on PCR analysis of immobilized virions. Belin et al. replicase) and exogenous (2.: Successful transformation of somatic embryos of an European grape cultivar (Russalska 3) with the normal. and Asian Vitis species inoculated by green grafting with a GFLV source. Lahogue and Boulard: Search for genes of resistance in grapevines. index populations by PCR-RFLP and sequencing of the ITS region. truncated or nontranslatable coat protein gene of GFLV.
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Amsterdam. K. Several of these viruses have distorting and chromogenic strains and may occur in mixed infections with GFLV.F. eds). Lehto. 1978. 1996. Satellite nucleic acids. P. Piazzolla. In: Grapevine Viruses and Certification in EEC Countries: State of the Art. Martelli. Bulletin de l'OIV 69. Harrison B. Karasev..J. These viruses can readily be eliminated by heat therapy or in vitro meristem tip culture. 286 pp. Wellink. and A. Savino and P. G. 2005) Extensive reviews of the biological.. Sanfaçon. T. 1996. J. T.J. 1992. Kurstak. Family Comoviridae. New York.G. 1997) and their satellite RNAs (Mayo et al. Brown.. The Plant Viruses. Nematode Vectors of Plant Viruses. C. 1981. M. 1996. a nematode-borne virus originally classified as a tentative species in the genus Nepovirus. Taliansky. All have polyhedral particles about 30 nm in diameter and a positive sense. A. Sélection clonale de la vigne: sélection sanitaire et sélection pomologique. Mayo M. Fritsch. which are separately encapsidated.A. 3. Springer-Verlag. epidemiological.... 5. 2005).. A. causing diseases whose symptoms are similar to. Desselberger and L. In: Virus Taxonomy. ed.V. A tentative grouping of nepoviruses Phytopathologia Mediterranea 17.F. a non-taxonomic clustering. the Mediterranean and Middle East. subgroup C. Quaderno No. 1977) . M. London. K.H.V. Gallitelli. Mechanical transmission to herbaceous hosts or indexing on Vitis indicators can also be used. ed).. Iwanami. 2000) are available. 2000. Martelli. Palukaitis. Murant.e. Jones. G. Nepoviruses of grapevine and their nematode vectors in the EEC. 945-971. Rüdel M.P. 145-147. In: Virus Taxonomy. Simons and M. Vol. Eight Report of the International Committee on Taxonomy of Viruses (C. i.. Ball. are now classified in the genus Nepovirus. H. Scholtof. 1997. Murphy et al. eds) Elsevier/Academic Press. Murant (eds). 807-818. 362 pp. U.P. Harrison B. T.INFECTIOUS DEGENERATION (EUROPEAN AND MEDITERRANEAN NEPOVIRUSES) Besides GFLV several other nepoviruses can infect grapevine in Europe. Martelli and R. Academic Press. CAB International.B.. Milne.A. Taylor and Brown. Polyhedral Virions and Bipartite RNA Genomes. single-stranded RNA genome occurring as two functional species (RNA-1 and RNA-2). physico-chemical. Van Regenmortel et al. 1ere partie: Effects des viroses sur la culture des vignes et ses produits. 1977. D. J. Leibowitz. 23-39. Influence des viroses et qualité. (G. and A. In: Virus Taxonomy.. eds). subgroup A typified by Tobacco ringspot virus (TRSV). Maniloff.. San Diego. 2005. which were originally included in the Nepovirus group (Harrison and Murant. (E.P. Nepoviruses. Walter B. Many are transmitted by longidorid nematodes (Rüdel. and molecular characteristics of nepoviruses (Harrison and Murant. Mayo.). and G. References Goldbach R. Nepoviruses. 1955) and are subdivided into subgroups based on physico-chemical properties of member viruses. Quacquarelli. Bari. 1978. Serological (ELISA. Istituto Agronomico Mediterraneo.M. Seventh Report of the International Committee on Taxonomy of Viruses (M. Strawberry latent ringspot virus (SLRSV). Vienna. 10281032 Murant A. typified by Tomato ringspot virus (ToRSV) (Martelli et al. Family Comoviridae. Sixth Report of the International Committee on Taxonomy of Viruses (F. or indistinguishable from those of fanleaf. Le Gall et al.D.D. subgroup B. 1992). 1955. CMI/AAB Descriptions of Plant Viruses No. 185. Fauquet.A.A. Elsevier/Noth Holland. In: Handbook of Plant Virus Infections: Comparative Diagnosis. 341-347. RT-PCR) are routinely used for their detection in grapevine tissues (primarily cortical scrapings from dormant canes). Wetzel and N. Oxon. Le Gall O. family Comoviridae (Goldbach et al. Martelli G.F. Plenum Press.E and D. Their detrimental effects to grapevine culture and products have been summari zed by Walter and Martelli (1996).. V. 35 . Nepovirus Group. typified by Tomato black ring virus (TBRV). Yoshikawa.A.P. Murant 1981. 197-238. ISEM) and molecular assays (hybridization.F. Taylor C. has now been assigned to the newly established genus Sadwavirus (Le Gall et al.
SLRSV and TBRV found in grapevines with atypical Reisigkrankheit in West Germany. Bulgaria. AILV and GCMV. wt 2. Israel.2 x 106 (RNA-1) and 1. ArMV. USA (California). Quacquarelli et al. vinifera varieties. Gerola et al. consisting of two separately encapsidated molecules with Mol. Russo et al. symptoms are of the fanleaf type. Iran. respectively. In Germany. and. Coat protein has a single type of subunits with Mr 54 x 103. Cross-protection between ArMV and GFLV has been reported.: Interactions between nepoviruses in grapevine and herbaceous hosts. and sediment as three components (T. and B). have a angular outline. Jankulova and Kaitasova: ArMV found in grapevine in Bulgaria. Yugoslavia. ArMV and TBRV by ELISA in Israel. Its particles are about 30 nm in diameter. Dalmasso et al.1 x 106 (RNA-2). Transgenic plants expressing the coat protein gene of the virus have been produced.: ArMV detected in Japan in grapevines imported from Europe. losses up to 50 % have been recorded. Canada. Stellmach: Review paper on ArMV in grapevine. always in Germany the severe dieback disease of the cv. Turkey. 2. M. 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.4 x 106 and a size of 1104 nt. 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. Component T is made up of empy protein shells. Belli et al.: Ultrastructure of ArMV infections in plant tissues Martelli and Lehoczky: Detection of ArMV in grapevine in Hungary. and circular about 350 nt in size. Vuittenez et al. The genome is a positive sense ssRNA. Murant: ArMV description in the CMI/AAB Descriptions of Plant Viruses series. a typical nepovirus belonging in subgroup A of the genus Nepovirus. The virus supports the replication of two types of satellite RNAs. linear with Mol. Romania. This virus has also been found in grapevine in Switzerland. TBRV.Genus NEPOVIRUS ARABIS MOSAIC VIRUS (ArMV) 1. and Japan.: Isolation of ArMV from grapevine in Italy. and with other nepoviruses in the Reisigkrankheit complex of western Germany. Kobayashi et al. accounting for 27% (component M) and 41% (component B) of the particle weight. 36 .: Xiphinema diversicaudatum can transmit ArMV to grapevine.: Detection of ArMV by ISEM Tanne: Detection of GFLV. Rüdel: Transmission of ArMV to grape seedlings by Xiphinema diversicaudatum. Description ArMV. In other V. index. the vector of GFLV. is serologically related to GFLV.: Physico-chemical properties of GFLV.95-2. wt of 0. whereas components M and Bcontain RNA.: ArMV. Hungary. Brückbauer and Rüdel: Symptoms of atypical Reisigkrankheit in the vineyard are associated with ArMV in West Germany Bercks et al. It can infect many woody and herbaceous plants and is transmitted to grapevine by the nematode Xiphinema diversicaudatum but not by X. ArMV occurs often in mixed infections with GFLV in certain areas of France and Italy.
Of 407 accessions of 1986 1987 1988 1989 1989 1989 1989 1990 1990 1990 1991 1991 1992 1992 1993 1993 1994 1995 1995 1995 1996 1996 37 . Becker et al. : Comparison of coat proteins of ArMV and other nepoviruses.: Properties of ArMV small satellite RNA. whereas the rootstock remains infected. When a healthy scion is grafted onto an infected rootstock. Molecular assays are as good as ELISA for routine testing.: ArMV in Switzerland Lázár et al.Kerner to ArMV seems to be limited in time.: Cross-protection experiments in Chenopodium quinoa between ArMV and GFLV Gugerli et al.: ArMV is not seed-transmitted in grapevines Liu et al.1982 1984 1985 Brückbauer: Possibility of distinguishing GFLV. The virus cannot be recovered from leaves or buds of the Kerner scions.: The presence of ArMV satellite RNA can attenuate symptoms in certain hosts Bertioli et al. such as RpRSV or GFLV can be found in both rootstock and scion.: Transgenic Nicotiana plants transformed with the coat protien of ArMV produce empty viral shells.: ArMV recorded from Romania Huss et al. Faber.: Evidence that ArMV has two RNA-2 molecules and complete nucleotide sequence of both RNAs Etscheid et al. Ipach et al.: Association of ArMV-infected rootstocks with Kerner disease in West Germany. whereas other nepoviruses. Eppler et al. RRV. SLRV and TBRV Belli et al. Stellmach: Kerner disease is probably caused by ArMV. MacKenzie et al. ArMV. Kaper et al. Stellmach and Berres: The susceptibility of cv. Walter et al.: Properties of a strain of ArMV isolated from grapevine in Italy.: Nicotiana plants engineerd with ArMV coat protein gene show different degrees of tolerance to the virus Flak and Gangl: ArMV recorded from Austria Loudes et al.: Nucleotide sequenece of a small circular satelllite RNA Steinkellner et al.: A hypovirulent ArMV isolate delays GFLV infection in grapevines under field conditions. Steinkellner et al. Study of histological changes at the graft union level.: Nucleotide sequence of the ArMV satellite RNA Liu et al.: Survey for the presence of ArMV in Canadian vineyards Lahogue and Boulard: Search for genes of resistance in grapevines. Rüdel: In the Palatinate (West Germany) ArMV is transmitted by Xiphinema diversicaudatum and occurs often in mixed infections with GFLV in grapevine.: Transformation of grapevines with the coat protein gene of ArMV.: Use of cDNA probes for ArMV detection. the virus is recovered from the scion only during the first year.: Detection of ArMV by PCR in herbaceous hosts and grapevines Steinkellner et al. Hypothesis of a graft incompatibility when the rootstock is infected with ArMV. Yield losses may reach 77% in cv. Marc-Martin 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
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. Properties of a previously undescribed nepovirus fron south-east Anatolia.: First isolation by mechanical transmission of an unknown nepovirus from cv.95-2. wt 0. Bulletin OEPP/EPPO Bulletin 32. Biological. Historical review 1972 1972 1977 Ferreira and De Sequeira: Description and preliminary characterization of an unidentified virus denoted CM112.. Cigsar I. The virus can be detected directly in grapevine leaf extracts by immunodiffusion in agar plates..: GBLV description in the CMI/AAB Descriptions of Plant Viruses series. N. De Mendonça et al. Uyemoto et al. 2005. The scattered distribution of infected vines in the field suggests that the virus is spread primarily by infected propagative material.P. 1977 1978 41 . References Abou Ghanem-Sabanadzovic N. Digiaro. I. M.: A virus serologically related to GBLV isolated from Vitis labrusca cv.P. The genome is a positive sense ssRNA. is not seed borne and was reported only from south-eastern Turkey.: Isolation of virus CM112 from in vitro cultures of grapevine tissues. Two isolates of GBLV have been transmitted by mechanical inoculation to seedlings and rooted cuttings of several grapevine cultivars without inducing symptoms. Martelli et al.: Complete nucleotide sequence of GARSV RNA-2 3.1 x 106 (RNA-2). Abou Ghanem-Sabanadzovic. Description Grapevine Bulgarian latent virus (GBLV) owes it name to the fact that it was found for the first time in Bulgaria in 1971. 471-475 Gokalp K. Concord in New York State. physico-chemical and serological characterization of the virus and assignment to the Nepovirus group (now genus Nepovirus). M. 2. B1. Digiaro and G. S. isolated in 1970 in Portugal from symptomless vines. A strain of this virus had been found previously in Portugal and described as virus CM112. Martelli. 35-41. Boscia and G. D. The economic importance of the virus is minor. 2002. 2003. Historical review 2002 2003 2005 Cigsar et al. Kizlar tahtasi showing mild fanleaf-like symptoms Gokalp et al. M.: Description and thorough characterization of GARSV identified as a new species in the subgroup B of the genus Nepovirus Abou Ghanem-Sabanadzovic et al. Digiaro and G.P. wt of 2. Journal of Plant Pathology 85. The virus supports the replication of a satellite RNA with Mol. Complete nucleotide sequence of RNA-2 of two Turkish nepoviruses.: Description of GBLV. By contrast. De Stradis. whereas components B1 and B2 contain RNA. Virus Genes 30. Coat protein has a single type of subunits with Mr 54 x 103. Cigasr. Sanitary status of grapevine in south eastern and central Anatolia. Virus particles are about 30 nm in diameter and sediment as three components (T. GRAPEVINE BULGARIAN LATENT VIRUS (GBLV) 1. GBLV is a typical nepovirus belonging in subgroup C of this genus but its vector is not known.2 x 106 (RNA-1) and 1.5 x 106 (less than 1800 nt). 2. GBLV has also been recorded from Hungary and Yugoslavia. consisting of two separately encapsidated molecules with Mol. but different from GBLV. A. Component T is made up of empy protein shells. Martelli. where it is widespread and infects latently several grapevine varieties growing in widely separared areas. Martelli. Martelli et al. Sabanadzovic. The virus occurs as different closely related but serologically distinguishable strains. 335-340. and B2). accounting for 39% (componet B1) and 42% (component B2) of the particle weight.vector.
1992.P. Ferreira A. 186 Martelli G. Colmar. Quacquarelli and D. France..Sur l'isolement d'un virus à partir de cultures de tissus de vigne. A. 349-354. Annals of Applied Biology 85. Some properties of grapevine Bulgarian latent virus.. 269-272. Stace-Smith. Pocsai E. De Sequeira and A. and O. Gallitelli. Physical and chemical properties of the particles and ribonucleic acid of blueberry leaf mottle virus.. A. Pereira and V. Grapevine Bulgarian latent virus. D. Proceeding 4th Meeting of ICVG. Gallitelli. Niagara Falls 1980. 1972. Savino and A. Di Franco. V. Annales de Phytopathologie. Phytopathology 71.A. 468-472. De Sequeira O. Applicability of immunosorbent electron microscopy (ISEM) for the detection and identification of CM112 virus in grapevine. 217-222. M. Gallitelli et al. De Sequeira. Martelli G.. 21-24. Russo and V. P. Martelli et al. Varennes and De Sequeira: First application of ELISA for the detection of virus CM112. Colmar 1970. Pocsai: Occurence of GBLV in Hungary. Some properties of the new latent virus from grapevine rootstocks in Yugoslavia. Monografias INIA 18 . Garcia de Orta Estacao Agronomica 12. De Sequeira. Quacquarelli.. Savino and A. Proceeding 4th Meeting of ICVG. The ultrastructure of grapevine Bulgarina latent virus infections in natural and artificial hosts. 113-120. Savino and O. No. Occurence of grapevine Bulgarian latent virus in Hungary.: Ultrastructural study of GBLV infections in grapevine and C. 42 .. Numéro hors série. Krastanova S. 95-101. Simoes. A. 1980. 1980.A. 1980. 143-145 De Mendonça A. Martelli G. Numéro hors série.A. M. Yankulova. Properties of a distinctive strain of Grapevine Bulgarian latent virus. and R. Acta Phytopathologica Academiae Scientiarum Hungaricae 16. Russo et al. Gallitelli. Gallitelli D. 1972.1979 Martelli et al. 1985. Vasconcelos-Costa.N. Dimitrijevic B. Preliminary studies on an undescribed grapevine virus. Mota.A. 1978.. 1981. Rasteniev'dni Nauki 29. De Sequeira.: Improvement of ELISA protocol for GBLV detection the whole year round.. O. Krastanova et al. 1977. 1980 1980 1980 1980 1981 1981 1982 1983 1985 1992 3. and J.A. P. CMI/AAB Descriptions of Plant Viruses. V. Phytopathologia Mediterranea 22. Possibilities for the whole-year ELISA detection of viruses infecting grapevines. 1970. O.: Detection of GBLV in grapevine leaf extracts by ISEM.: Detection of virus CM112 in grapevine leaf extracts by ISEM. V. 1983. 51-58.. Abracheva. Proceedings 7th Meeting of ICVG. Ramsdell D. D. Ganeva and M. Savino.P. Niagara Falls 1980. References De Mendonça A. Niagara Falls 1980. Abracheva. Martelli G. Proceedings 6th Meeting of ICVG. M. quinoa. Ramsdell and Stace-Smith: The New York isolate of GBLV is a strain of BLMoV.: 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.. D. Quacquarelli. An immunoradiometric assay for the titration of a Portugese strain of grapevine Bulgarian latent virus (GBLV).C. Annales de Phytopathologie. Cordoba 1976. A preliminary report. The Portuguese strain supports the replication of a satellite RNA. 245-250. Dimitrijevic: GBLV found in Yugoslavia De Mendonça et al.: A comparative study of Bulgarian GBLV isolates and the Portuguese virus CM112 establishes that CM112 is a serologically close but distinguishable strain of GBLV.Proceedings 7th Meeting of ICVG. Proceedings 7th Meeting of ICVG. A manually transmissible latent virus of the grapevine from Bulgaria.A.P.P. 27-32. 135-141. 1979. De Sequeira and Vasconcelos-Costa: Use of an immunoradiometric assay for the titration of the Portuguese strain of GBLV.A. Jankulova. Ferreira. 1981.
Varennes A. Affected vines lack in vigour and may decline and die. and O. Some virus strains induce leaf deformity. Isolation and identification of a strain of grapevine Bulgarian latent virus in Concord grapevines in New York State. Plant Disease Reporter 61. 1977. wt of 1. sometimes together with X. wt of 2. Agronomia Lusitana 41. index. The virus is not serologically related with GFLV. Taschenberg and D. Jakó et al: HCMV affects pigment and sugar content of infected grapevine leaves. RNA-2 has Mol.6 x 106 . Tobacco pants and the rootstock 110R have been successfully transformed with the viral coat protein for induction of resistance.8 x 106 . 949-953. 1966 1968 1968 1969 1969 1971 1972a 1972b 1972 1975 1977 43 . a size of 7212 nt and accounts for 40% of the particle weight.. It has been recorded also from Czechoslovakia. pretty much like GFLV. 269-277.Uyemoto J. a symptom virtually indistiguishable from GFLV-induced yellow mosaic. Kenten: GCMV is distantly serologically related to Cacao necrosis virus. Mali et al. Lehoczky and Tasnady: A study of the effect of HCMV on yield and sugar content of infected grapevines. Martelli and Quacquarelli : Physico-chemical characterization of HCMV and comparison with TBRV. vuittenezi is very frequently found in vineyards with chrome mosaic patches. Martelli and Sarospataki: X. The coat protein has a single type of subunits of Mr 52 x 103. The genome is bipartite. No evidence that X. Croatia and Austria. The virus belongs in the same subgroup of TBRV (subgroup B) to which is distantly related serologically.: HCMV adversely affects photosynthetical carbon dioxide fixation. Pozsár et al. and was originally called Hungarian chrome mosaic virus. De Sequeira. The virus appears to be unrelated serologically to GFLV and is not transmitted by X. vuittenezi transmits GCMV or GFLV. a size of 4441 nt and accounts for 31% of the particle weight.A. Historical review 1966 Martelli et al. index as vector of the virus (unconfirmed results). Although GCMV particles have been detected by immunosorbent electron microscopy in Xiphinema index fed on infected hosts. GRAPEVINE CHROME MOSAIC VIRUS (GCMV) 1. transmitted to herbaceous hosts from Hungarian grapevines with symptoms similar to those of fanleaf and yellow mosaic. 2. early reports that this nematode could transmit the virus have not been confirmed.K. Description Grapevine chrome mosaic virus (GCMV) was first found in Hungary. symptomless infection may occur. Detection of CM122 latent grapevine virus by enzyme-linked immunosorbent assay (ELISA). E. Martelli et al. called Hungarian chrome mosaic virus.: Host range and properties of a spherical virus. Evaluation of short reaction times and re-use of a-globulin and conjugate. RNA-1 has Mol. 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. GCMV is transmitted through grapevine seeds. Saric and Hranuelli: GCMV recorded from Croatia. near Lake Balaton.: The isometric virus associated with Hungarian chrome mosaic is serologically distantly related to Tomato black ring virus (TBRV). Leaves of infected vines are partially or entirely bright yellow or whitish.F. index. Virus re-named Grapevine chrome mosaic virus. However. double nodes and short internodes. 1982.: GCMV recorded from Slovakia and report of X.K. Hummer.
Taylor and Brown: Results of GCMV transmission trials with X. L. T.: GCMV detected by ELISA in infected field-grown vines. Lanneau and J. Le Gall et al.M. Bretout C. Dodd and Robinson: GCMV and TBRV are molecularly related. R. 7809-7819. index extracts by ISEM. Brault. Roberts and Brown: Detection of GCMV in X. Le Gall et al. Laimer da Camara Machado and V. 258-262. Candresse. Delbos. Further demonstration that the two viruses are related. Ravelonandro and J. Lázár et al: Seed transmission of GCMV in grapevine. 44 . A. No assessment of resistance made.: Transformation of roostock 110R with the coat protein gene of GCMV. Candresse. Savino. 127-128.: GCMV recorded from Austria. T. 1989.. Vitis 34. 1995. Dunez. Lehoczky: Pinot noir and Jubileum 75 are good indicators for GCMV. Detection of nepoviruses in ligneous grapevine material using IC/PCR. Dunez. Le Gall. Russo et al.: GCMV and TBRV can recombine. O. Nucleotide sequence and genetic organization of Hungarian grapevine chrome mosaic nepovirus RNA2. Bretout et al. O. Doz et al. M.: Complete nucleotide sequence of GCMV RNA-1. Dunez. 89-97.. Brault et al.: Detection of GCMV in leaf dips by ISEM.: GCMV cross-protects Chenopodium quinoa from the severe apical necrosis induced by a TBRV strain.1979 1980 1980 1982 1984 1985 1985 1989 1989 1989 1990 1993 1993 1994 1994 1995 1995 1997 2000 Lehoczky et al.: Development of molecular probes for GCMV detection. Genetically engineered resistance against grapevine chrome mosaic virus. D'Onghia. 3.: Complete nucleotide sequence of GCMV RNA-2.: Up-to-date report on virus diseases of grapevines in Hungary and description of the clean stock programme implemented in the country. M.P. The virus vector is yet to be identified. Le Gall. 1993. M. index are inconclusive. Brault V. Kölber et al. Brault V. V. 1989. Brault et al. Le Gall et al. References Brandt S. Acta Horticulture 235. Lehoczky et al. O. This finding does not imply vectoring capacity by this nematode. 231-238. Himmler. and G.: Characterization of a GCMV strain and confirmation of its serological relationship with TBRV.: Tobacco plants genetically engineered with the coat protein gene of GCMV are resistant to infection. Occurrence of grapevine chrome mosaic nepovirus in Austria. T. Virus and RNAspecific molecular hydridization probes for two nepoviruses. Nucleic Acid Research 17. Journal of Phytopathology 142. Plant Molecular Biology 21. Dimou D.: Description of a certification scheme for the production of virus-free propagating material in Hungary. Hilbrand. Le Gall and J. Brandt and Himmler: Development of a IC-PCR protocol for GCMV detection in cortical scrapings from dormant grapevine canes. 1994. Dimou et al. Candresse. Lázár et al..
1966.J. Bojnansky. Phytopathologia Mediterranea 24. No.P. Acta Phytopathologica Academia Scientiarum Hungaricae 1. and G. Lázár J. Kiserletugyi-Kozlemenyek 64 (1-3). 1989. ArMV) in the seeds and seedlings of grapevine by ELISA. Lehoczky J. Detection of grapevine chrome mosaic virus in naturally infected vines by indexing. Extended Abstracts 13th Meeting of ICVG. Sarospataki and J. G. L. Kölber. Nucleotide sequence of Hungarian grapevine chrome mosaic nepovirus RNA1. Phytopathologia Mediterranea 24. 7795-7807. Host range and properties of a virus associated with Hungarian grapevines showing macroscopic symptoms of fanleaf and yellow mosaic. Martelli G. 129-134. 1-7. Lehoczky J. 119-126. 1970. 1993. O. Horváth. Jakó N. L. J. 1966. Candresse. University of California. The purification and some properties of cocoa necrosis virus. 49-64. Martelli G. Martelli G. 567-568..C. Candresse and A. A. 123-141. 1972a. Studies on the nitrogen metabolism of vines infected with yellow mosaic virus. Jakó N. Hungarian chrome mosaic of grapevine and tomato black ring: two similar but unrelated plant viruses. Hungarian chrome mosaic. J. Pozsár B. 1972. Vuittenez. Lehoczky .. Proceedings International Conference on Virus and Vector on Perennial Hosts. Transmission by nematodes of some grapevine viruses occurring in Czecholovakia and Hungary. 1995. 1979. Farkas. and D.Ser. 1285-1289. Division of Agricultural Sciences. Lehoczky J. Lehoczky J. Le Gall O. Kenten R.) University of California. NW Frazier (ed..Tasnady. 1969. Dunez.J. M.P. Hajdú. Numero hors série. Lehoczky and G. Martelli G. Szonyegi and M. Division of Agricultural Sciences. Doz B. Kölber M. Annales de Phytopathologie 11. Adelaide 2000. Y. Beczner.P. Lázár. Devergne. Agrobacterium-mediated genetic transformation of grapevine somatic embryos and regeneration of transgenic plants expressing the coat protein of grapevine chrome mosaic nepovirus (GCMV). Grapevine chrome mosaic virus. Candresse and J.M. with Special Reference to Vitis. Annales de Phytopathologie. a serotype ot tomato black ring virus. J.P. Nouvelle évidence d'une parenté sérologique éloignée entre ce virus et celui deas anneaux noirs de la tomate (tBRV). Detection of grapevine chrome mosaic virus in field-grown vines by ELISA. A. T. Pol'nohopodarska Veda . Kölber M.. E.. Martelli G. Nucleic Acid Research 17. Lehoczky. T. Montreux 1993. Proceedings International Conference on Virus and Vector on Perennial Hosts.. V. Kölber and J. Robinson. Delbos and J. S.P. 505. Quacquarelli. 29-44. 1990. The change of pigment and sugar content in the chrome mosaic virus infected leaves of grapevine. Les Colloques de l'INRA 11. G. GFV-YM. Mali V. Pacsa and J. Lehoczky. Luntz. R. Caractérization d'une souche du virus de la mosaïque jaune crome de la vigne (GCMV) isolée en Hongrie de vignes non panachées. 1969. Sarospataki. 45 . Lehoczky and A.. 172-173. T. Sarospataki. Bouquet. 2000. Lehoczky and A. Kuszala and A. J. with Special Reference to Vitis. and A. Davis 1965. G. M.R. 165-173. Le Gall O. 1984. Lehoczky. 1-130. Detection of some nepoviruses (GFV. Weinberg und Keller 15. Muranyi . GCMV.. Prémunition: une competition entre souches faibles et souches sévères pour un voie commune d'expression de symptômes. 1968.Dodd S. S. 1731-1740. Berkeley.. J.. 1982. Certification scheme for production of virus-free grape propagating material and its results in Hungary. Acta Phytopathologica Academia Scientiarum Hungaricae 3. Le Gall O. Torregrosa . University of California. G. Martelli G... L. 389-401. Journal of General Virology 65. Mikulás.. and G. 236-237. Journal of General Virology 76. 1966. Quacquarelli. Davis 1965. Preliminary report on purification and serology of a virus associated with Hungarian grapevines showing macroscopic symptoms of fanleaf and yellow mosaic. Quacquarelli and J.. Annals of Applied Biology 71.. Effect of the grape chrome mosaic and grape fanleaf-yellow mosaic virus infection on the photosynthetical carbon dioxide fixation in vine leaves. CMI/AAB Descriptions of Plant Viruses. Division of Agricultural Sciences. 185-192. Serologische Verwandschaft eines mit dem ungarischen "chrome mosaic" vergesellschafteten Virus mit einem Stamm des "tomato black ring virus". and S. and A. Danglot. 58-72. J. 169-170.. Vitis 8. 1994. J. 1972b. 171-170. J. Martelli G. Brault and J. 402-410. Dunez. Sarospataki. L... Kertgazdasag 22. 1985. Grapevine virus diseases and clean stock program in Hungary. 135-140. 1971. Dunez.P. The effect of fanleaf and chrome mosaic virus diseases on yield and the fruit sugar content of grapevine. 1985. 1968. 103. 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. S. Lehoczky and G. In: Virus Diseases of Small Fruits and Grapevines. Vanek and V. Sarospataki. Nucleotide sequence homologies among RNA species of strains of tomato black ring virus and other nepoviruses. Plant Science. 102.. Extended Abstracts 11th Meeting of ICVG. Sznyegi. Cardin.P.H. Phytopathologia Mediterranea 8. 206-210. A Handbook. Nematodes of the family Longidoridae (Thorne 1935) Meyl 1960 found in Hungarian vineyards and virus transmission trials with Xiphinema index Thorne et Allen. Quacquarelli. Quacquarelli. 3. 1975.. Lázár J.
and belongs in the subgroup C of the genus Nepovirus. Description Grapevine Tunisian ringspot virus (GTRSV). mol. 35-41.. The scattered distribution of infected vines in the field suggests that the virus is spread primarily by infected propagative material. identified as a new species in the subgroup A of the genus Nepovirus. wt of 2. Immunosorben electron microscopy for detecting saptransmissible viruses of grapevine. 2. Saric A.P. Proceedings of the 7th Meeting of ICVG.: Description and thorough characterization of GDefV. Martelli. Grapevine deformation virus. 251-257.4 x 106 daltons and apparent size of c. K.. showing leaf and cane deformations Cigsar et al. 335-340 Cigsar I. Coat protein subunits have a Mr 53 x 103. is not seed-borne and reported only from south-eastern Turkey. Boscia and G.P.. including all those known to infect grapevine. M. Particles are isometric c. 5. GDefV is readily detected by ELISA and PCR using primers designed on the coat protein sequence. Journal of Plant Pathology 85. G.). Abou Ghanem-Sabanadzovic. 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. Sanitary status of grapevine in south eastern and central Anatolia. 2003.: First isolation by mechanical transmission of an unknown nepovirus from cv. : Complete nucleotide sequence of GDefV RNA-2 3.800 nt) and B (particles containing a molecule of RNA-1 with Mol. 286 pp. CAB International. 6.P.The virus has no recognized vector. a novel nepovirus from Turkey.P. Niagara Falls 1980. Proceedings of the Conference on Excoriosis and Virus Diseases of Grapevine. 1977. and D. The genome is bipartite. Digiaro. A. The virus sediments as three components: T (empty shells). The virus belongs in the subgroup A of the genus Nepovirus. Hranuelli. References Abou Ghanem-Sabanadzovic N. 46 . M (particles contaning a molecule of RNA-2 with Mol. Complete nucleotide sequence of RNA-2 of two Turkish nepoviruses.. 1980. GTRSV is serologically unrelated to any of 19 nepoviruses tested. M. wt of 1. Martelli. is distantly related serologically to ArMV but not to GFLV. Historical review 1991 Ouertani et al. and T. wt of 2 x 106 daltons and apparent size of c. 471-475 Cigsar I. Investigations on grapevine viruses in Croatia. 2002. Oxon. Description Grapevine deformation virus (GDefV) was recovered from Turkish grapevines showing distinct fanleaf-like symptoms. N. Martelli. Digiaro and G. 30 nm in diameter and sediment as three components. Gokalp. 2.3 x 106 Da and a size of 3753 nt. Nematode Vectors of Plant Viruses.J. Digiaro and G. 149-151. Virus Genes 30. 1997. GRAPEVINE DEFORMATION VIRUS (GDefV) 1. Abou Ghanem-Sabanadzovic et al. Taylor C. Mostar 1977. Martelli and V.800 nt. Sabanadzovic.: A mechanically transmissible virus was recovered by sap inoculation from Tunisian grapevines showing mild fanleaf-like symptoms..F Brown. Historical review 2002 2003 2005 Cigsar et al.6 x 106 Da and RNA-2. De Stradis. GRAPEVINE TUNISIAN RINGSPOT VIRUS (GTRSV) 1. E. Bulletin OEPP/EPPO Bulletin 32. D. 2005. S. No vector is known and no information is available on the distribution and economic importance of the virus. was isolated from a Tunisian grapevine with mild fanleaflike symptoms. RNA-1 has a mol. Savino.Russo M. wt of 2. M. distantly serologically related with ArMV.
A. Scottish and English. 1992. and sediment as three components (T. M. G.P.. Description Raspberry ringspot virus (RpRSV) is a nepovirus belonging in subgroup A of this genus. 1992. 1978. C. accounting for 29% (component M) and 43% (component B) of the particle weight. and differs from the type strain for it often sediments as if it were a single centrifugal component.J.: Sequencing and molecular characterization of two German isolates of RpRSV from grapevine 2003 3. 198.M. D.6 x 106 (RNA-2). Boscia. Two strains of different virulence occur in the Palatinate. D. 283-300. 107-117.C. 2.: Nucleotide sequence of RpRSV RNA-2 Jones et al. G. Archives of Virology 126. Ebel R. Rüdel and B. Extended Abstracts 14th Meeting of ICVG. consisting of two separately encapsidated molecules with Mol.J.L. A. Locorotondo 2003. Savino. Properties of an unusual isolate of raspberry ringspot virus from grapevine in Germany and evidence of its possibile transmission by Paralogidorus maximus. The grapevine strain of this virus is serologically very distantly related to the two main serotypes. M. Wardell J. 2189-2194.. The type strain of RpRSV is transmitted by Longidorus macrosoma but the grapevine strain is transmitted by Paralongidorus maximus. McGavin. V. RASPBERRY RINGSPOT VIRUS (RpRSV) 1.6 x 106 (RNA-1) and 1. Murant: Description of RpBRSV in the CMI/AAB Plant Virus Description series Stellmach and Querfurth: Study of a strain of RpRSV isolated from cv. Brückbauer H. 2003. Martelli and N. Heat therapy of infected grapevines. The nucleotide sequence of RNA 2 of raspberry ringspot nepovirus. Greco. Murant A. G.A. Wetzel. Reustle. Krczal and T. Historical review 1970 1978 1978 1982 1992 1994 Vuittenez et al. Elbling in West Germany. Robinson. References Blok V. Crop losses can be higher than 30 %. A. Jones A. No. Edwards and M. CMI/AAB Descriptions of Plant Viruses. The virus has only been found in grapevine in western Germany. The coat protein has a single type of subunits with Mr 54 x 103. These differences strongly suggest that the grapevine-infecting RpRSV may be a different viral species. Raspberry ringspot virus. The viral genome is a bipartite positive sense ssRNA. 1994. Manoukian... Brown. Jolly. 1982.3. D. The virus is transmitted by Paralogidorus maximus Ebel et al. Symptoms are similar to those of fanleaf. Mögliche Beziehungen zwischen Virus und Symptomausprägung bei der Rebe. W. This strain differs considerably from the English type strain of the virus although is serologically closely related to it. Particles are about 30 nm in diameter. RNA-2 is 3928 nt in size and contains a single ORF encoding a polypeptide with Mr of 124 kDa. Minafra.F. M. wt of 2. References Ouertani R. Journal of General Virology 73. Altmayer. and B). have angular outline. Molecular characterization of two German Raspberry ringspot virus isolates infecting grapevines and construction of full length infectious clones.T.A. 88-118. M. Brückbauer: RpRSV can be distinguished from other nepoviruses on the basis of symptoms induced on Vitis idicator plants Blok et al. Castellano.J. Annals of Applied Biology 124.: Recovery of RpRSV from grapevines of Palatinate. FS4 is a good indicator.: Biological and physico-chemical characterization of the grapevine strain of RpRSV.F. 47 . A Schnabel. Die Wein-Wissenschaft 37. Properties of a previously undescribed grapevine nepovirus from Tunisia. Mayo. 16.
The vector to grapevine is Longidorus attenuatus. latex test and barium sulfate test. Joannes Seyve. and Ontario (Canada). Virus particles are isometric.5 x 106 daltons and a size of 1327 nt. Pathologie und Thermo-Labilität mehrerer Reben-Isolate des Himbeerringflecken-Virus (raspberry-ringspot. or directly in grapevine leaf extracts using bentonite flocculation test. and B). but they can be high. SLRV and TBRV found in grapevine in the Palatinate. Lehoczky and Burgyan: Occurrence of TBRV in grapevines in Hungary. Losses are not known precisely. respectively. Ontario as the cause of severe damage to cv. Querfurth. Greece. TBRV supports the replication of a satellite RNA with Mol. Turkey. Vuittenez et al. Annales de Phytopathologie 2.: Nucleotide sequence of a TBRV satellite RNA. Rüdel: Transmission of TBRV to grapevine by Longidorus attenuatus. Tanne: Detection of TBRV by ELISA in Israel. wt of 2. RpRV and TBRV detected serologically in grapevine in West Germany.65 x 106 (RNA-2) accounting for 44% and 37% of the particle weight. RNA-2 is 4662 nt in size encoding a polyprotein with Mr of 150 kDa. Stellmach and Bercks: Further investigations on TBRV in grapevine. 2. RRV and TBRV are not transmitted by contact of roots or foliage in the vineyard. Rüdel and H Brückbauer. Untersuchungen zur Serologie.: RRV. Stobbs and Van Schagen: First record of TBRV from Canada. rings and lines on the leaves of recently infected plants. Historical review 1963 1965 1966 Stellmach and Bercks: TBRV detected in rootstock Aramon x V. 1967 1970 1970b 1976 1977 1980 1984 1984 1986 48 .. Bercks and Stellmach: ArMV. is a strain of this virus. Bercks and Querfurth: GFLV. known to cause severe damage to the grapevine variety Joannes Seyve in Ontario. its own subgroup. either by agar gel diffusion with extracts of herbaceous hosts previously infected mechanically from grapevine. mottling of the older leaves. TBRV produces a reduction in growth and yield. The virus is a definitive nepovirus species classified in subgroup A of this genus. J. The latex test is considered as the most sensitive and the least time consuming method. Stellmach: Review paper on TBRV in grapevine. The genome is a bipartite positive sense single-stranded RNA occurring as two separately encapsidated functional molecules with Mol.7 x 106 (RNA-1) and 1. then in Yugoslavia. The virus was detected in grapevines in the Niagara Peninsula. Their coat protein consists of a single type of subunits with Mr of about 57 kDa. Bercks: Comparison of three serological tests for detecting several viruses. Weinberg und Keller 25. M. 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. M. including TBRV : bentonite flocculation test. Kuszala.virus). 279-327 TOMATO BLACK RING VIRUS (TBRV) 1. ArMV. and G.Stellmach G. wt of 0. Vuittenez A. 128-136. Description Tomato black ring virus (TBRV) was first found in grapevines in Germany. and an increase in graft failure. 1978. Meyer et al. Détection et étude selogique du virus latent des taches annulires du frasier (strawberry latent ringspot). riparia 143 A in West Germany. Joannes Seyve virus. RNA-1 is 7356 nt in size and contains a single open reading frame encoding a polypeptide with Mr of 254 kDa. chlorotic spots. 1970. Israel. about 30 nm in diameter have angular outline and sediment as three components (T.
and J. O. Nucleotide sequence of tomato black ring virus RNA-1. 1984. Erdiller. Turkiye. et du virus des taches annulaires du Framboisier (raspberry ringspot) chez des vignes du Palatinat. Journal of General Virology 69. Researches on grapevine virus diseases and determination of their incidence in Ankara. Hemmer and C. Virus particles are isometric.6 x 106 (RNA-2). Brückbauer.W. Occurrence of tomato black ring virus on grapevine in southern Ontario. wt 2. The nucleotide sequence of tomato black ring virus RNA-2. Rüdel and H. Mayo and C. Meyer M.1986 1988 1993 Meyer et al. 2. Complete nucleotide sequence of a satellite RNA of tomato black ring virus. Vuittenez A. It was also detected in imported vines in Turkey and Portugal. and G. M. and 1. Kuszala. Journal of Turkish Phytopathology 22. 1993. Kreiah et al. Their coat protein consists of two types of subunits with Mr 43 x 103 and 27 x 103. The genome is a bipartite positive sense singlestranded RNA occurring as two separately encapsidated functional molecules with Mol. Description Strawberry latent ringspot virus (SLRSV) has been isolated from grapevine in the Palatinate (Germany) and in northern Italy.G. du virus des anneaux noirs de la Tomate (tomato black ring). 55-61. M. Détection et étude sérologique du virus latent des taches annulaires du Fraisier (strawberry latent ringspot).A. The virus supports the replication of a satellite RNA 1118 nt in size. Babini and Bertaccini: Electron microscope study SLRSV infections in plant tissues.6 x 106 (RNA-1) accounting for 38% of the particle weight. 279-327. Annales de Phytopathologie 2. RNA-2 is 3824 nt in size and encodes a single ORF expressing a polypetide with Mr of about 99 kDa. Canadian Plant Disease Survey 64. References Akbas B.: Nucleotide sequence of SLRSV satellite RNA. Greif C.: Nucleotide sequence of TBRV RNA-2 Greif et al. Assignement of SLRSV to the new genus Sadwavirus. The virus is a definitive species in the genus Sadwavirus. Fritsch. O. Meyer M. The virus is transmitted by Xiphinema diversicaudatum. and B). Symptoms on affected European grapes are of the fanleaf type. Savino et al. Brückbauer: SLRSV can be distinguished from other nepoviruses on the basis of symptoms induced on Vitis idicator plants. O. 1257-1271 Stobbs L. Genus SADWAVIRUS STRAWBERRY LATENT RINGSPOT VIRUS (SLRSV) 1. 1575-1583.: Nucleotide sequence of TBRV RNA-1 Abkas and Erdiller: TBRV recorded from grapevines in Turkey 3. 3-5. 1517-1529. 1970. Journal of General Virology 65. 1988. 1986.: Nucleotide sequence of SLRSV RNA-2. Fritsch. 49 . J. respectively..: SLRSV and other nepoviruses isolated from grapevines in Germany Credi et al. Hemmer. Le Gall et al. Journal of Gerneal Virology 67. Historical review 1974 1977 1981 1982 1982 1987 1993 1994 2005 Murant: Description of SLRSV in the CMI/AAB Descriptions of Plant vrises series. 1984. Van Schagen. Hemmer and C. Fritsch. Bercks et al. about 30 nm in diameter have angular outline and sediment as three components (T. Kreiah et al.: SLRSV found in grapevine in Turkey.: SLRSV recorded from grapevine in Italy. M...
T.A. References Babini A. Brückbauer. Desselberger. Strunk and J.. Wellink. 1994.. J.. A. Maniloff. Journal of General Virology. Phytopathologia Mediterranea 20. 2527-2532. L. Mögliche Beziehungen zwischen Virus und Symptomausprägung bei der Rebe. A. K.F. Credi R. U. H.. Sequence analysis and location of capsid protein within RNA 2 of strawberry latent ringspot virus. Bercks R. Wetzel and N. 1977. Strawberry latent ringspot virus.. Yoshikawa. Gelli.. Kreiah S. 1163-1165. A. 1982. 1993. Savino V.. Untersuchungen überdie Viruskrankheited der rebe unter besonderer Berüchsichtigung “atypischer Formen” der Reisigkrankheit.A.. Bertaccini and C. (eds) 799-802 Elsevier/Academic Press. Die Wein-Wissenschaft 37. 1974.M. G. The nucleotide sequence of a satellite RNA associated with strawberry latent ringspot virus. Rüdel. Cooper and G.M. FAO Plant Protection Bulletin 35. Viral aggregates induced by a distinctive strain of strawberry latent ringspot virus from grapevine. D'Onghia and M. 133-180. 2005. H. G. L. Yilmaz. J.V. In: Virus Taxonomy. CMI/AAB Descriptions of Plant Viruses No. Turkey. 50 . Karasev.I. 74. Martelli.3. J. G. and A. Kreiah S. Sanfaçon.. Murant A. Eight Report of the International Committee on Taxonomy of Viruses (Fauquet. Bertaccini.. A. C. Le Gall O. Brückbauer H. Mayo. T.. 1981.R. Cooper. Iwanami. M.I. Genus Sadwavirus. 88-118. and Ball. Strawberry latent ringspot virus in grapevine. 56-63. 102-104.A. Phytopathologische Zeitschrift 104. Weinberg und Keller 24. T Jones. Strunk.. Betti. Querfurth and M.R. 1987. A distinctive isolate of Strawberry latent ringspot virus from grapevines in Italy. 126. 1982. Babini. London. 304-308. Journal of General Virology 75. Lehto.P.
and/or ring spots) and distorted leaves. ingiallimento nervale (Ital. ToRSV and BLMoV are also seed transmitted in grapes. rivesi transmit ToRSV type strain (decline). riparia. A number of rootstocks containing V. except for BLMoV which may have been introduced from Europe. single-stranded RNA occurring as two separately encapsidated functional molecules with Mol . grapevine decline. positive-sense. TRSV and PRMV. BLMoV. rupestris plasma show field resistance to the northern US strain of ToRSV and to TRSV and PRMV. California) yield but not vigour is affected. poor fruit setting. All roostocks and. X. In warmer climates (Maryland. V. whereas in V. This type of resistance is hypersensitivity. depérissement de la vigne (Fr. Description Main synonyms: Yellow vein. Infected vines decline slowly over time. Description Blueberry leaf mottle virus (BLMoV) is named after the disease induced in highbush blueberry (Vaccinium corymbosum). californicum transmits ToRSV yellow vein strain. straggly and shelled clusters. PRMV. ELISA and molecular tools are useful for testing field-infected material.35 x 106 (RNA-1) and 2. berlandieri or V. the infecting virus and the climatic conditions. labrusca is also resistant to TRSV. jaunissement des nervures. Virus particles are isometric. Concord it delays bud burst. Partial sequence of the 3' 51 . little grape (Eng. mottled (oak leaf pattern. Long distance spread takes place primarily through infected propagating material. V.) Main symptoms: Symptomatological responses of grapevines vary according to the species (i. elongatus. distortion of canes. In cold climates (e. Longidorus diadecturus and L. Peach rosette mosaic virus (PRMV) in V. labrusca cv. sedimenting as three components. americanum sensu lato and PRMV by X.e. Their coat protein consists of a single type of subunits with Mr of about 54 x 103 Da. most V. Adernvergilbung (Germ. malformation and mottling of the leaves.). Transmission: These viruses are all transmitted by grafting and mechanical inoculation. The genome is a bipartite. labrusca causes a severe disease characterized by delayed bud burst. TRSV is transmitted by X. and ToRSV separately or in combination. No vector is known for BLMoV. Varietal susceptibility: There are great variations in the susceptibility of Vitis species and cultivars. deperimento della vite. are endemic in North America and thought to be native of the region. labrusca.). interspecific hybrids). wt of 2. its main host. and poor fruit setting. Vitis vinifera. induces fanleaf-like symptoms on leaves and canes. All these viruses. americanum sensu stricto. 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. PRMV. Bunches are small and straggly (Maryland's grapevine little berry) and leaves may show chrome yellow flecking along the veins (California's yellow vein).GRAPEVINE DEGENERATION AND DECLINE (AMERICAN NEPOVIRUSES) 1. Blueberry leaf mottle virus (BLMoV) infects latently European grapes.). Nematicidal control of vectors is possible. All other viruses are transmitted by longidorid nematodes: Xiphinema americanum sensu stricto and X. BLUEBERRY LEAF MOTTLE VIRUS (BLMoV) 1. about 30 nm in diameter with angular outline. New York State and Ontario) own-rooted European grapes affected by Tomato ringspot virus (ToRSV) and Tobacco ringspot virus (TRSV) decline rapidly. exhibiting stunted growth. BLMoV is a definitive nepovirus species assigned to subgroup C. interestingly. Control: Use of virus-free propagating material and resistant rootstocks.g. which in blueberry is transmitted by pollen. but not conclusive.15 x 106 (RNA-2). and poor fruit setting Agent: The above mentioned four distinct nepoviruses. TRSV. Alternative weed hosts that have epidemiological significance are known for ToRSV. V. are involved in the aetiology of North American grapevine degeneration and decline.
K. 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. sedimenting as three components. References Bacher J. D. Warkentin.C. Their coat protein consists of a single type of subunits with Mr of about 57 x 103 Da. and has no economic importance. Uyemoto J. Hummer. Description Peach rosette mosaic virus (PRMV) is named after the disease induced in peach. 1994.Use of resistant roostock hybrids and of certified planting material is recommended. mottled and variously deformed leaves and delayed bud burst. Grapevines (Vitis labrusca) are infected in New York State (USA) by a serologically distinct strain of the virus. but not eradicate. 2. but in highbush blueberry the virus is pollen-borne and suspected to be pollen-transmitted .W. PRMV can also be introduced in a site by infected planting material and be spread by vectors to adjacent vines. 1977. which induces fanleaf-type symptoms and is distantly related to GBLV. Clusters are straggly. Physical and chemical properties of the particles and ribonucleic acid of blueberry leaf mottle virus. 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. Healthy grapevines become infected when planted in soils from diseased vineyards. Historical review 1977 Uyemoto et al: BLMoV isolated from New York 'Concord' vines showing fanleaf-like symptoms. The virus is a definitive nepovirus species assigned to subgroup C. Crop losses up to 60% and death of susceptible V.. where the disease occurs in more or less circular patches and spreads slowly. labrusca cultivars (Concord. Taschenberg and D. As the virus is endemic and seed-borne in some perennial weeds Taraxacum officinale (dandelion). about 28 nm in diameter with angular outline. Plant Disease Reporter 61. and with extensive shelling of the berries..F. 145-156 Ramsdell D.termini of both RNA molecules has been determined.F. PRMV is soil-borne. 949-953.Rasmdel and J. Occasional. Hancock. The virus is seed-transmitted in grapevines and C. The genome is a bipartite positive sense single-stranded RNA occurring as two separately encapsidated functional molecules with Mol. The virus is transmitted through seeds in grapevines and C. RNA1 is 8004 nt in size and contains a single open reading frame encoding a polypeptide with Mr of 240 kDa. 468-472. : Partial nucleotide sequence of BLMoV RNA-1 and RNA-2 1981 1994 3. Vines are stunted and show a progressive decline. Vectors are the Dorylaimoid nematodes Xiphinema americanum sensu lato and Longidorus diadecturus. Isolation and identification of a strain of grapevine Bulgarian latent virus in Concord grapevines in New York State. mostly to vines adjacent to previously infected plants.5% of seedlings from seeds taken from diseased vines proved to be infected. 1981. Virus particles are isometric.K. The vector is unknown. E. Stace-Smith. wt of 2. possibily non specific transmission by L.4 x 106 (RNA-1) and 2. vector populations . Virus Research 33. one of its crop plant hosts. elongatus has also been reported. Sequence analysis of the 3' temini of RNA 1 and RNA 2 of blueberry leaf mottle virus. when a vineyard is planted susceptible cultivars may become infected by nematode vectors.2 x 106 (RNA-2) accounting for 44% and 37% of the particle weight. PRMV is seed-borne in both naturally infected dandelion (4% of infected seeldlings) and in artificially infected C. especially) and a number of American-French hybrids have been recorded. Infected grapevines show shortened and crooked shoots. smaller and fewer than normal . 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. but identified as a strain of GBLV. Pollen grains of cv. and R. which may lead to their death. D. quinoa. 52 . Concord grapes are apparently virus-free but 9. Solanum carolinense (Carolina horse nettle) and Rumex crispus (curly dock). quinoa (90% of infected seedlings). respectively. PEACH ROSETTE MOSAIC VIRUS (PRMV) 1. Phytopathology 71.
Transmission of peach rosette mosaic virus to peach grape and cucumber by Longidorus diadecturus obtained from diseased orchards in Ontario.F.G Van Schagen and B. Historical review 1972a 1972b 1974 1976 Dias: Preliminary characterization of the grapevine isolate of PRMV. Transmission of raspberry ringspot. quinoa and has reproduced in part the field syndrome when inoculated mechanically to Concord grape seedlings. Allen et al. 1972b.. 53 . Numero hors sèrie.: Investigation on the susceptibility to PRMV infection of American and European grapevines and hybrid rootstocks.F. The characterization of a virus responsible for peach rosette mosaic and grape decline in Michigan.: Nucleotide sequence of RNA-1 of the grapevine strain of PRMV. Lammers et al. Dias H.R. 1982. Ramsdell: Review article on PRMV. 1984. 1228-1239.: Xiphinema americanum is an efficient vector of PRMV. Numero hors sèrie. S. Dias: Grapevine and peach strains of PRMV can be differentiated serologically. J.A Ebsary.. Annales de Phytopathologie. Annales de Phytopathologie.G Van Schagen and E. Ramsdell and Gillet: List of grapervine cultivars and roostocks showing differential susceptibility to PRMV.R.F. Dias H. Allen and Ebsary: Longidorus attenuatus transmits PRMV non specifically and with low efficiency. Cation. Ramsdell and Myers: Description of PRMV-induced grapevine degeneration and association of X. Canadian Journal of Botany 54.. 1-5 Allen W. Carolinense. americanum with the disease. Canadian Journal of Plant Pathology 6. tomato black ring and peach rosette mosaic viruses by an Ontario population of Longidorus elongatus. and D. and B.R. Canadian Journal of Plant Pathology 4. crispus) and transmission through grapevine seeds. Allen et al. 1978 1979 1980 1982 1983 1984 1985 1988 1988 1995 1998 1999 3. 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.A Ebsary. Dias and Cation: Biological characterization of the grapevine strain of PRMV. Purification and some characteristics of peach rosette mosaic virus (grape isolate). The virus is seedborne in C. Canadian Journal of Plant Pathology 10. 1988. Ramsdell and Myers: Field spread of PRMV is associated with the presence of infected weeds (T. Transmission of peach rosette mosaic virus by Ontario populations of Longidorus diadecturus and Xiphinema americanum (Nematoda: Longidoridae). 105-106. Strains of peach rosette mosaic virus differentiated by cross absorption and immunodiffusion tests.: Longidorus diadecturus transmits PRMV to grapevines. Dias and Allen: Physico-chemical characterization of PRMV.S Everleigh. J. officinale.2. Ramsdell and Gillet: Description of PRMV in the AAB Descriptions of Plant Viruses. Rasmdell et al.. 29-32 Allen W. References Allen W. 97-103. 16-18. 1976. Ramsdell et al: Use of ELISA for PRMV detection in grapevines. Dias H. 1972a. R.
Phytopathologia Mediterrenea 24. Andrews.M. 1978..H.C. Characterization of the single protein and the two nucleic acids of peach rosette mosaic virus. G. Plant Disease Reporter 63. Canadian Journal of Botany 58. 625-627.C.L. but was recorded from grapevines only in New York State and Pennsylvania. Description Tobacco ringspot virus (TRSV) is the type species of the genus Nepovirus and the prototype of subgroup A. Superimposed shallow and deep soil fumigation to control Xiphinema americanum and peach rosette mosaic virus reinfection a Concord vineyard. 447-450 Ramsdell D. 1988. TRSV is soil-borne and is transmitted by Xiphinema americanum sensu stricto.C. Paul. There is no evidence of seed trasmission in the grapevine. 1985. wt of 2. Bird GW. 54 . M.R. Preventive control measures are the use of resistant roostock hybrids and of certified planting material. French hybrids and European grape cultivars to infection by peach rosette mosaic virus. Pearson and A. 4143. AAB Descriptions of Plant Viruses. and J.3 x 106 (RNA-2). Ramsdell D. The genome is a bipartite positive sense single-stranded RNA occurring as two separately encapsidated functional molecules with Mol. Gillet. Uyemoto et al. and R. St. 57-73. Buzayan et al. R. Ramsdell D. R. Ramsdell D.7 x 106 (RNA-1) and 1. Gillet and L. Virus particles are isometric. Myers. Ramsdell D. J. Powell et al. respectively. Symptoms elicited by TRSV are the same as those of ToRSV in native cultivars. Cloning an sequencing of peach rosette mosaic virus RNA1. sedimenting as three components (T.). 154-157.: Nucleotide sequence of TRSV satellite RNA.W. TRSV has a relatively wide natural host range. 1995.M. 1983. Phytopathology 64. .M. Relative susceptibility of American.C. TOBACCO RINGSPOT VIRUS (TRSV) 1. J. In: Compendium of Grape Diseases (R. Allen.E Morris. Lammers A.: A review of viruses infecting grapevines in New York vineyards. Ramsdell D. Peach rosette mosaic virus decline. Gillet and C. Goheen eds.W. 2. Epidemiology of peach rosette mosaic virus in a Concord grape vineyard. Peach rosette mosaic virus.M. about 30 nm in diameter with angular outline. American Phytopathological Society Press. Myers.C..M..M. and B).Dias H.C. and W.. symptomatology and nematodes associated with grapevine 'degeneration' in Michigan. accounting for 44% and 28% of B and M particle weight. but in European grapes responses are similar to those elicited by GFLV. Ramsdell D. Susceptibility of American grapevine scion cultivars and French hybrid roostocks and scion cultivars to infection by peach rosette mosaic virus. Foster and Morris-Krsinich: In vitro translation of TRSV RNA-1 and TRSV RNA-2 yields major polypeptides with Mr of 225K and 116K. 364. Bird. Plant Disease 79. 51-52. 1974. A comparison between enzyme-linked immunosorbent assay (ELISA) and Chenopodium quinoa for detection of peach rosette mosaic virus in 'Concord' grapevines. 1998.C Ramsdell. Phytopathology 68. is endemic in Central and Eastern North America. and R. and J. Peach rosette mosaic virus.C. 1979. G. The virus supports the replication of a circular satellite RNA 359 nt in size.: TRSV agent of a new grapevine disease in New York State. Gillet. Gillet and G.C. Plant Disease 67. 74-78.C.L.: Survey of ToRSV and TRSV in Pennsylvanian vineyards. American Vitis species reported to be resistant to ToRSV and TRSV. respectively. RNA-1 is 7514 nt in size and contains a single open reading frame encoding a polypeptide with Mr of 225 kDa. Ramsdell D. Stace-Smith: Description of TRSV in the AAB Descriptions of Plant Viruses series.W. Rose. 1747-1754. 1174-1178. Historical review 1970 1977 1985 1985 1986 1990 Gilmer et al. 1980.F Allison and D. 1999.C. RNA-2 has been sequenced only in part. No. Virus Research 65. Coat protein consists of a single type of subunits with Mr of about 57 x 103 Da..C.
Nucleotide sequence and in vitro expression of the capsid protein gene of tobacco ringspot virus. Kelts. ToRSV is soil-borne. the infecting virus strain. Keese and A. which often leads to death.J. Vines are stunted and show a progressive rapid decline. Buzayan J.S.M.8 x 106 (RNA-1) and 2. and B. : Partial nucleotide sequence of TRSV RNA-2..K. Virology 151. 55 . 1996. RNA-1 is 8214 nt and RNA-2 is 7273 nt in size. Virus particles are isometric.S. mottled and distorted leaves and short internodes. Morris-Krsinich. B. Incidence of tomato rigspot virus and tobacco ringspot virus in grapevines in Pennsylvania.. In California ToRSV affects the yield rather than the vine's growth. Preventive control measures are the use of resistant roostock hybrids and of certified planting material. smaller and fewer than normal. A new grapevine disease induced by tobacco ringspot virus.L. Bruening. californicum in California The virus can be introduced in a site by infected planting material and be spread by vectors to adjacent vines.R. 619-627. Powell C. Disease named yellow vein. 186-199.A. Two serological ToRSV variants are known to infect grapevines. 1993. 335-349. American Journal of Enology and Viticulture 28. 1-8. Symptomatological responses vary according to the species (V. Uyemoto J.A. The genome is a bipartite positive sense single-stranded RNA occurring as two separately encapsidated functional molecules with Mol. Bruening. J.L. AAB Descriptions of Plant Viruses. Vines grow vigorously but bear little or no fruit. Foster R. Forer. Both RNAs contain a single open reading frame encoding polypeptides with Mr of 244 kDa (RNA-1) and 207 kDa (RNA-2).M. Stace-Smith R. where it occurs iin the region of the Great Lakes and in the Pacific seaboard from California to British Columbia.K. Zalloua P. Gooding and Hewitt: A mechanically transmissible virus found to be associated with yellow vein. Historical review 1954 1956 1962 Hewitt: Report of an "unfruitful vine" condition in California to which a yellow speckling of the leaves is associated. "yellow vein" being the characterizing syndrome of its infections. V. Longenecker and L. Description Tomato ringspot virus (ToRSV) is a definitive species in the genus Nepovirus and the prototype of subgroup C. W. more severely in colder than in warmer climates. Coat protein consists of a single type of subunits with Mr of about 58 x 103 Da. ToRSV has a relatively wide natural host range and is endemic in North America.. P. sedimenting as three components (T. 309.. wt of 2. Chemical cleavage of the 5'-linked protein of tobacco ringspot virus genomic RNAs and characterization of the protein-RNA linkage. 131-136. vinifera. Silva and S. Nucleotide sequence of satellite tobacco ringspot virus RNA and its relationship to multimeric forms. 516-519. Virus Research 30. Infected vines have small. Abawi. 1990. Vectors are the Dorylaimoid nematodes Xiphinema americanum sensu stricto and X. Singh. 1977.. 1985. 2.B. No. Cummins and G. J.R.M. respectively.4 x 106 (RNA-2) accounting for 44% and 41 % of the particle weight. contrary to the decline strain which is seed-transmitted. 1970. ToRSV-induced decline affects European cultivars. J. Buzayan and G. rivesi in north American States and Canada and X. Virology 144. and the climatic conditions. Tobacco ringspot virus. especially if self-rooted. References Buckley B.M. The yellow vein strain of the virus is pollen-borne but is not transmitted through seeds.. J.M. Clusters are straggly. 1986. Virology 219. Gerlach G.1993 1996 Buckley et al. about 30 nm in diameter with angular outline.A. Zallua et al. Hewitt: Successful graft transmission of unfruitful vine condition. Phytopathology 60. The virus has been occasionaly recorded from grapevines outside of North America. labrusca. 702-704. Gould. 1985. TOMATO RINGSPOT VIRUS (ToRSV) 1. Virus and virus-like diseases affecting grapevines in New York vineyards. Uyemoto and L. Plant Disease 74. and B).L. Synthesis and processing of the translation products of tobacco ringspot virus in rabbit reticulocyte lysates.: Nucleotide sequence of TRSV RNA-1 3. and with extensive shelling of the berries. interspecific hybrids). Gilmer R.
: List of grapevine roostocks and cultivars showing differential susceptibility to ToRSV in Canada.: ToRSV found in grapevines in Taiwan.: Complete nucleotide sequence of ToRSV RNA-2. Gilmer and Uyemoto: ToRSV agent of a decline of Baco noir in New York State.: Complete nucleotide sequence of ToRSV RNA-1. Rott et al. Rott et al. Allen et al. Martelli and Taylor: Review of nematode-borne viruses and their vectors.: A review of viruses infecting grapevines in New York State vineyards.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. Baumgartnerova and Subikova: ToRSV recorded form grapevine in Slovakia.: Confirmation that the grape yellow vein and the the grape decline strains of ToRSV are serological variants of the same virus. Herrera and Madariaga: ToRSV recorded from grapevine in Chile. Stace-Smith and Ramsdell: Review of nepoviruses of the Americas.: Xiphinema rivesi identified as the main vector of ToRSV in Ontario vineyards.: Survey of ToRSV and TRSV in Pennsylvanian vineyards. Martelli: Review of nematode-borne viruses of grapevines and their epidemiology. Bitterlin and Gonslaves: ToRSV retained and transmitted by viruliferous Xiphinema rivesi stored for two years at 1-3°C. Teliz et al. Gonsalves: ToRSV is irregularly distributed in infected vines but can be detected by ELISA. Cory and Hewitt: The yellow vein strain of ToRSV is not transmitted through seeds. Yang et al. Li et al: ToRSV identified in China in grapevine seedlings grown from seeds imported from France. Piazzolla et al. Allen and Dias: Physico-chemical characterization of ToRSV. Bays and Tolin: ToRSV found in grapevines in Virginia Powell et al. Rowhani et al: Description of sampling strategy for detection of ToRSV. Uyemoto and Gilmer: Spread of ToRSV through the soil of New York State vineyards recorded. Uyemoto: Seed transmission of the decline strain of ToRSV. 56 . Corbett and Podleckis: Ultrastructural study of ToRSV-infected grapevine tissues. californicum). 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. americanum (now X. American Vitis species reported to be resistant to ToRSV and TRSV.: Transmission of the yellow vein strain of ToRSV by X. Allen et al. Uyemoto et al. Dias: Record of ToRSV in the Niagara peninsula.
Bulletin of the California Department of Agriculture 43. Rott M.. Plant Disease Reporter 61. L. Téliz D. 1968. 861-863.E. Transmission of tomato ringspot.A. Tomato ringspot virus associated with little grape disease of Vidal 256 grapevines. 1977. Podleckis E.G. Works of the Institute of Experimental Phytopathology and Entomology .J. Tremaine and D'A. and B. Plant Disease Reporter 56.G and V. Ivanka pri Dunaji 4. Membrane-associated spherical particles in extracts and tissues of virus-infected grapevines.L.R. and M. Incidence of tomato rigspot virus and tobacco rigspot virus in grapevines in Pennsylvania.F. 408-411. Bitterlin M. Beijing 2004. M.. Nepoviruses of the Americas. Incidence of tomato ringspot virus in grape in Virginia.M. 465-473.B. A severe outbreak of virus-induced grapevine decline in Cascade grapevines in New York. Allen W. J. Identification of tomato rinsgspot virus in leafroll diseased grapevines. Gilchrist. J. M. peach yellow bud mosaic. 1990. V. Phytopathologia Mediterranea 24.M. 663 Martelli G. A. Spatial distribution of Xiphinema rivesi and persistence of tomato ringspot virus and its vector in soil.A. Rokni. 1954. 1989. 393-400 Hewitt W. 658-663. Presence and incidence of grapevine viruses in central Chile. 35-44. Journal of General Virology 72. CMI/AAB Descriptions of Plant Viruses.E. 31-34. 1982. M. Distribution of viruses and their nematode vectors.B. Cory L.. 1987.F. and the association of a mechanically transmissible virus with the disease. 1977. Proceedings 15th International Plant Protection Congress.R. Tomato ringspot virus.3.. A comparison of Grapevine yellow vein virus and a Grapevine isolate of Tomato ringspot virus.G. Phytopathology 53. Hewitt. Phytopathologia Mediterranea 24. 1028-1037. Savino. Canadian Journal of Plant Pathology 4.A. and E. and S. Corbett M. Piazzolla P. Rochon.. Dias and J. 196-203. Uyemoto. 1992.F.B. Nucleotide sequence of tomato ringspot virus RNA 1.W. 1982.G. 1972. Susceptibility of grape cultivars and rootstocks to an Ontario isolate of tomato ringspot virus. Proceedings 7th Meeting of IGVG. 1995. 1505-1514. Uyemoto J. 1962. Rott M. Xiang . Rowhani A.A. 290 Stace-Smith R. Ebsary.E. 131-166. Some virus and virus-like diseases of grapevines. and W. and W. Advances in Disease Vector Research 6. Van Schagen. 275-277. Nematode-borne viruses of grapevine. and grape yellow vein viruses by Xiphinema americanum.. 1985. J.V... 1980. Plant Disease 74. Zhang and Y.P. Podleckis. Association of Xiphinema species with soil types ang grapevines infected with tomato ringspot virus in Ontario. Current Topics in Vector Research 5. 1991. L. Y. their epidemiology and control.. Vitis 31. 1984. No. Gonsalves D.K. Phytopathology 56.K. Tomato ringspot virus in "Baco noir" grapevines in New York. and H.. Gilmer R.B. Chen. Subikova.H. Van Schagen and B.F.. Lownsberry.K. Canada. References Allen W. Allen W.C. Lee and D'A. Bays D. Gonsalves. Forer. 1978. Stace-Smith R.F.. 1988. 44-50. Powell C. Walker and S. Nematologia Mediterranea 6. 2004. Sampling strategies for the detection of grapevine fanleaf virus and the grapevine strain of tomato ringspot virus.. Gooding G. Canadian Journal of Botany 55. 24-28. Grape yellow vein: symptomatology.R. 475-480. American Journal of Enology and Viticulture 13.. 151-189. G.V. N. 702-704. 710. Plant Disease 72. 1987.A. Nucleotide sequence of tomato ringspot virus RNA-2. Phytopathology 79.F. Li. 2001. Castellano and D.P. Properties of the single protein and two nucleic acids of tomato ringspot virus. 1985.W.. Martelli G. Agricultura Tecnica 61. 157-164. Grogan and B. Longenecker and L. Journal of General Virology 76. R. H. 95-106. Some grapevine viruses in pollen and seed.C. Detection of tomato ringspot virus in grapevines: irregular distribution of virus. 133-135. 13161320 Dias H. Ramsdell. 1989. Li M. Tolin. Herrera M. 1993.V. 57 . and D. 47-64. Taylor.. Purification and serology of a virus associated with the grape yellow vein disease. Baumgartnerova H.S Whei. Incidence and geographic distribution of tomato ringspot virus in De Chaunac vineyards in the Niagara peninsula. 1169. Hewitt.V. Phytopathology 58. identification. Dias. Phytopathology 72. 1963. Musci. 1-27. 1975. Plant Disease Reporter 59. and V. Plant Disease 71.F. Corbett. and C. 1966. Niagara Falls 1980. Stobbs. and J. Rochon. Madariaga. 98-101. Detection and identification of plant viruses for quarantine in China.K. Gooding G.
R. Chen.M. Virus and virus-like diseases affecting grapevines in New York vineyards.. 1972. 58 .L. 1977. 1062-1064.S. Plant Disease Reporter 56. Yang I. T.C.J.K. Uyemoto J. and R. 1986. Journal of Agricultural Research China 35. Sap-transmissiible viruses associated with grapevine yellow mottle disease in Taiwan. J. 131-136. 504-510. Deng and M. American Journal of Enology and Viticulture 28.K.Uyemoto J. Spread of tomato ringspot virus in "Baco Noir" grapevines in New York. Abawi.. Gilmer. Cummins and G.
vinifera.5 nm. Virus particles are very flexuous filaments about 12 nm wide. exhibiting open structure and distinct cross banding with a pitch of about 3. 61 . brittle and rolls downwards. These spots enlarge with time and coalesce so that. infection of rootstocks is symptomless. differentiation of GLRaVs at the species level was by Roman numerals. called grapevine leafroll-associated viruses (GLRaV). GLRaV-8. and low in sugar. In the most severe cases. Leafroll decreases grapevine yield (by 15-20% in average) and affects negatively rooting ability. Careful observation of field symptoms in infected vines reveals that there are several types of leafroll. and GLRsV-9 in the newly established genus Ampelovirus. thus suggesting that there can be several causal agents. thus impairing carbohydrate translocation from foliar parenchymas. usually leaving a narrow green band along the primary and secondary veins.). Particle length varies from 1400 to 2200 nm according to individual viruses. Its occurrence in viticultural areas is worldwide. the whole leaf surface becomes deep purple. The leaf blade becomes thick. 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. GLRaV-5.).). GLRaV-2 in the genus Closterovirus.). accartocciamento. 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. accartocciamento fogliare (Ital. except for a variable decrease in vigour. enrollamiento de la hoja. enrollado (Sp. Description The first descriptions of grapevine leafroll date back to the mid 19th century. differing somewhat in aspect and in severity. especially the phloem. depending on the climate and geographic location. most of the leaf surface becomes reddish. enroulement (Fr. but the leaves become chlorotic to yellowish. GLRaV-1.).) Main symptoms: In red-berried cultivars of Vitis vinifera reddish spots develop in the lower leaves in late spring or summer. Rollkrankheit. whereas the Mr of all other viruses ranges between 35 and 44 kDa. whereas GLRaV-7 is presently classified as unassigned species to the family.e. Agents: To date. Enrolamento de la folha (Port. Main synonyms: White Emperor disease (Eng. Hence. nine different viruses with filamentous particles. i. instead of reddish. potassium depletion in the leaf blade and accumulation in the petioles. in autumn. the symptoms are similar. they are inferior in quantity and quality. respectively. the same as the size of coat protein (CP) subunits. GLRaV-2 CP has a Mr of 24 kDa. and is probably the most widespread virus disease of grapevine. Leafroll is no less important than fanleaf in economic importance. and with many cultivars. have been found in leafroll-infected vines. as estimated by polyacrylamide gel electrophoresis. the risk of disseminating the disease is great if untested rootstocks are used. The fruits often mature late and irregularly. Sizes deduced from the nucleotide sequence of the CP cistron are 22 kDa for GLRaV-2 . GLRaV-6. Blattrollkrankheit (Germ. Starch accumulates in degenerated chloroplasts causing increased thickness and brittleness of the leaf blades. reduction of protein content. and lowering of sugar content.GRAPEVINE LEAFROLL 1. These negative effects are reverted if the disease is eliminated by sanitation treatments. Until 1995. changes in the pattern of peroxidase and polyphenoloxidase isoenzymes. In white-berried cultivars of V. GLRaV-3. All GLRaVs belong in the family Closteroviridae. graft take and plant vigour. Also the composition and aromatic profile of the musts are modified. Sieve elements are obliterated and crusheds. These symptoms progress towards the top of the canes as the season advances. GLRaV-4. In most cases. A number of other physiological parameters are affected. Other such viruses may exist. Also plant anatomy is affected. which are differentiated from one another by a progressive number.
biologically. longispinus. and has structural organization differing from that of other sequenced closteroviruses. Indexing on red-fruited cultivars such as 'Cabernet sauvignon'. and some are commercially avaliable. vinfera and cortical shavings from mature dormant canes of V. with none of which they are serologically related. The genome is a monopartite. So far. American Vitis species and rootstocks are the best antigen sources for serological assays.528 nt in size. 'Cabernet Franc' 'Pinot noir'. grafted vines) which is largely responsible for its dissemination over medium and long distances. Symptom expression depends on the variety. and degenerate primers have been designed and successfully used for PCR detection of virtually all GLRaVs. Detection: In many cases. GLRaV-3. As to nucleic acid-based assays. Composite samples should be used to minimize false negative responses that may originate from the unven distribution of GLRaVs in chronically infected vines. 29. Its soft scale insects vectors are Pulvinaria vitis and Neopulvinaria innumerabilis. viburni and Ps. 'Merlot'. contains 13 ORFs. but it does not discriminate between GLRaVs and was reported to be less sensitive than ELISA. Pl. Other GLRaVs may exist in nature as suggested by reports. roostocks. Varietal susceptibility and sensitivity: No immune variety or rootstock is known. the type member of the genus. Ps. Ps. comstocki. has a number of minor biological variants which can be differentiated by the reaction of inoculated Nicotiana species. climate. vinifera varieties show symptoms most clearly because of the reddening of the leaves.919 nt in size. American rootstocks are usually symptomless carriers of GLRaVs. Natural field spread of leafroll disease has been reported from many countries in Europe and elsewhere. Pseudococcus longispinus. which is the type species of the novel genus Ampelovirus has a genome 17. contains nine open reading frames (ORF) and has structural organization identical to that of Beet yellow virus (BYV). leafroll can be detected by its symptoms in the field on red-fruited varieties. GLRaV-2. Leaf tissues or petioles from mature symptomatic leaves of V. All GLRaVs can be identified by serological and nucleic acid-based techniques.5 kDa for GLRaV-4 . citri. GLRaV-3. positive sense RNA molecule. Foliar tissues are not recommended for serological GLRaVs detection in American Vitis species and roostocks. Ps.647 nt in size and contains 10 major ORFs. The presence of high molecular weight double-stranded RNAs (dsRNA) in phloem tissue extracts can be used as infection marker. This has been ascertained experimentally for GLRaV-1. None of the GLRaVs is known to be seedborne. Spread at a site is mediated by mealybug and soft scale insect vectors. Mealybug vectors of GLRaV-3 are Planococcus ficus. affinis. Parthenolecanium corni. 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.35 kDa for both GLRaV-3 and GLRaV-1. number an types of infecting viruses. ultrastructurally. only vectors of GLRaV-1. Membranous vesicles can derive eitrher from peripheral vesiculation of mitochondria followed by disruption of the organelles (GLRaV-1. GLRaVs differ in various vays (molecularly. GLRaV-5 and GLRaV-9 are phylogenetically close to one another but are serologically unrelated. broadspectrum. calceolariae. soil condition and probably. The genome of GLRaV-2 is 15. GLRaV-1 is transmitted in nature by the pseudococcid mealybugs Heliococcus bohemicus and Phenacoccus aceris and the soft scale insects Pulvinaria vitis.Transmission is semipersistent and does not appear to be vector-specific. The genome of GLRaV-4 and GLRaV-9 are incompletely sequenced but show a structural organization compatible with that of the genus. GLRaV-5 and GLRaV-9 are both transmitted by Ps. 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. and Neopulvinaria innumerabilis. singlestranded. GLRaV-5) or from vesiculation of the endoplasmic reticulum (GLRaV-2 and GLRaV-7) Transmission: Leafroll is graft-transmissible and persists in propagative material (budwood. GLRaV-2 and GLRaV-3. Red-berried V. or the hybrid LN 33 is still the most popular method for identifying the disease. and epidemiologically) from most of the known closteroviruses. GLRaV-3. Polyclonal antisera and/or monoclonal antibodies have been raised to each single GLRaV . and some of them are used as indicators. GLRaV-5 and GLRaV-9 have been identified. GLRaVs show molecular variations which give rise to a population of strains. Disappearance of dsRNAs from vines submitted to 62 . vinifera. in agreement with the quasispecies nature of viruses. Ps. maritimus. The genome of GLRaV-1 is 17. Regardless of whether they belong to the genus Closterovirus or Amplelovirus. These reagents are routinely used for ISEM. the only member of the group to be mechanically transmissible. classical double antibody sandwich ELISA (Chromo-ELISA) or Lumino-ELISA. 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. or by molecular techniques.
2. These particles are probably closteroviruses associated with leafroll. Goheen et al. As no apparent spread of the disease was observed.sanitation treatments is regarded as evidence for successful virus elimination. dsRNAs cannot be utilized for virus identification. Bovey: Leafroll in Switzerland. Ravaz and Verge: Occurrence in France of “rougeau”. unless they are hybridized with virus-specific probes. Harmon and Snyder: The "White Emperor" disease is graft-transmissible and is regarded a virus disease. Dimitrijevic: Leafroll in Yugoslavia.: Leafroll in Italy. Vuittenez: Leafroll in France. Control: Production and use of clonally selected and sanitized propagation material is very effective and the only preventive method for leafroll control available. However. Introduction of transgenic resistance to GLRaV-2 and GLRaV-3 is being attempted by engineering different viral genes into rootstocks and European grape cultivars. Hypothesis of the viral origin of leafroll.: Transmission of a virus to herbaceous plants from a leafroll-infected vine in Israel. Chamberlain: Leafroll in New Zealand. Belli et al. Tanne and Nitzany: Leafroll in Israel Tanne et al.: Studies on the effects of leafroll on yield of grapevines in California. 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”. The incidence if the disease was less than 20% as compared with 80 to 100% in commercial vineyards. Scheu: Leafroll is widespread in German vineyards. Fraser: Leafroll in Australia. Lehoczky et al. 1971 1973 1974 1975 63 . roostocks are suggested as the major souces of leafroll dissemination. Later studies showed that the virus is an occasional contaminant Lider et al.: Leafroll virus can be inactivated in vivo by heat therapy. a grapevine disorder similar to leafroll. Luhn and Goheen: Leafroll found in the original grapevine stocks imported from Europe into California in 1890. Hewitt: Leafroll in California Goheen et al. Mendgen: Presence of filamentous particles in grapevines with symptoms of flavescence dorée in West Germany. Blattny et al. a grapevine disorder similar to leafroll Scheu: Demonstration of graft transmission of leafroll from diseased to healthy Vitis vinifera.: Leafroll in Czechoslovakia. vinifera and there is no published information on how to protect healthy stocks from vector-mediated reinfection in the field. Hoefert and Gifford: Study of the effects of leafroll infection on vine anatomy.: Leafroll in Hungary. No sources of resistance are known in V.: White Emperor and leafroll are identical diseases.
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. Sasahara et al. Kliever and Lider: Study of biochemical changes found in grapevine infected with leafroll in California. Absence of such particles in healthy grapevines. Rosciglione and Gugerli: GLRaV-1 and GLRaV-2 with particles of 2200 nm and 1800 nm respectively. Hu and Gonsalves: Monoclonal antibodies produced against GLRaV-3. Mossop et al. Abracheva: Leafroll in Bulgaria. Production of polyclonal antisera for use in ELISA. but not in similar praparations from healthy plants.: 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.: Closterovirus-like particles purified from leafroll-diseased grapevines in France. Hofmann: Symptoms of leafroll in affected clones of Pinot noir and performance in West Germany. Antiserum against a New York isolate also reacted with GLRaV-3 from Europe.: 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. Barlass at al. previously found in grapevines in Switzerland. Zimmermann et al.: First record of successful elimination of leafroll in grapevine by using meristem tip culture in Japan. A large dsRNA molecule is consistently isolated from leafroll-diseased grapes.: ELISA testing reveals that GLRaV-3 has an uneven distribution in grapevine tissues. but not in healthy controls. Corbett et al: Electron microscope observations by negative staining of leaf extracts from leafroll-diseased grapevines in South Africa showed the presence of closterovirus. Tanaka: Leafroll in Japan.: Aggregates of closterovirus-like particles observed in thin sections of phloem from leafroll-diseased grapevines. Faoro et al. Von der Brelie and Nienhaus: Light and electron microscope study of cytopathological changes induced by leafroll in grapevines. A third closterovirus type called GLRaV-3. Confirmation that GLRaV-3 and the New York closterovirus isolate cross react serologically. Namba et al. Martelli et al. Castellano et al. Suggestion that a closterovirus may be the agent of the disease. Teliz et al. 1981 1981 1982 1982 1983 1984 1984 1984 1985 1986 1986 1987 1987 1988 1988 1989 64 . Presence of virus-like particles in thin sections of leafroll-diseased vines. are also present in leafroll-affected grapevines from Italy. Purification and serology of associated closterovirus-like particles.: Ultrastructural study of leafroll-infected grapevine tissues.like particles.: Elimination of leafroll by in vitro meristm tip culture and apex fragmentation. Gugerli et al.: Review on the detrimental effects of viral infection on grapevine physiology. found in grapevines affected by leafroll.: Studies on the cytopathology of leafroll-diseased grapevines.: Closterovirus-like particles and specific dsRNA found in leafroll-diseased grapevines in New Zealand. Rosciglione and Gugerli: GLRaV-3 is transmitted by the mealybug Planococcus ficus. Production of rabbit and hen antibodies for ELISA and ISEM to GLRaV-1 and GLRaV-3. Zee et al.
: Transmission of GLRaV-3 from grapevine to grapevine by the mealybug Pseudococcus longispinus in Israel.: Immunocytological detection and localization of GLRaV-1 and GLRaV-3 by immunogold labelling in grapevine thin sections. There is evidence that other GLRaVs are involved in leafroll etiology. Credi and Santucci: GLRaV-1 and GLRaV-3 cannot be detected by direct ELISA in leaves of graft-inoculated American rootstock. later in the leaves. GLRaV-2. stem pitting and corky bark spread rapidly. With leafroll. Gugerli et al. especially those containing V. Agran et al. but not GLRaV-1. Savino et al. Boscia et al. indicating the presence of other leafroll-associated viruses. The virus was detected in root tissues.: 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. Téliz et al. 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.: Detection of leafroll-associated closterovirus in recently infected grapevines in New York.: Use of green grafting for detecting virus-like diseases of grapevine.: Characterization of leafroll-associated closterovirus-like particles from grapevine using also monoclonal antibodies. ficus fed on infected vines. Faoro et al. was detected in P. vinifera varieties used as inoculum source. In Mexico leafroll. 1989 1989 1989 1990 1990 1990a. Identification of GLRaV-5.: Production of monoclonal antibodies to GLRaV-1 and GLRaV-3.1989 1989 Tanne et al. rupestris plasma. Pseudococcus longispinus is present on weeds around diseased vineyards. symptoms are obtained within 20-70 days. ELISA is to be applied to cortical scrapings rather than leaf tissues. Heat therapy requires very long treatments and is only 20-30 % successful. Auger et al.: Production and characterization of monoclonal antibodies specific to GLRaV-3. Zimmermann et al. -2 and -3 are common whereas GLRaV-5 is rarely detected. whereas meristem tip culture yields up to 100 % sanitation Walter and Zimmermann: Further characterization of closteroviruses associated with leafroll in France.: Evidence of the irregular distribution of GLRaV-3 in American rootstocks.: Further characterization of GLRaV-1 and GLRaV-3 by monoclonal antibodies.: Leafroll and associated closteroviruses in China Engelbrecht and Kasdorf: Transmission of GLRaV-3 by Planococcus ficus from grapevine to grapevine in South Africa. 1990 1991 1991 1991 1991 1991 1991 1991 65 . For reliable testing. GLRaV-1. b Hu et al. but they are easily detected in inoculated LN33 vines and in V. Good results in summer with extracts of basal leaves and in autumn or winter with wood shavings macerated in buffer. Gugerli: Review of grapevine closteroviruses. Transmission of GLRaV-3 by the mealybug Planococcus ficus. GLRaV-1 and GLRaV-2 were not transmitted. Identification of GLRaV-4 1990 1990 1990 1990 Walter et al.: Comparison of heat therapy and meristem tip culture for eliminating GLRaV-3 from Italian grape varieties. Gugerli et al.
: Revision of the nomenclature of GLRaVs and use of Arabic numerals in the species names. Former GLRaV 2 is re-named GLRaV-6. Tzeng et al. Golino et al.: Comparison of different assay methods for detecting GLRaVs : ELISA. Flak and Gangl: Leafroll and associated closteroviruses in Austria. Segura et al. Castellano et al. Chasselas shows the prsence of two different GLRaV-2. Gugerli and Ramel: Analysis by monolconal antibodies of a Swiss source of cv.1% in 1988 to 93. Faoro and Carzaniga: Ustrastructural study of GLRaV-1 and GLRV-3 infections.1% in 1992 Ioannou: Leafroll and natural spread of associated closteroviruses in Cyprus. Habili et al.: Mechanical transmission of GLRaV-2 and ultrastructural study of infected tissues of Nicotiana benthamiana.: Transmission of GLRaV-3 by Pseudococcus affinis in California.: Leafroll and associated closteroviruses in Romania. Kassemeyer: Detection of GLRaVs in Germany. Boehm and Martins: Leafroll in Portugal. Katis et al: Leafroll and associated closteroviruses in Greece. Jordan: In a New Zealand commercial vineyard GLRaV-3 incidence increased from 9. Belli et al. Milkus et al. Martelli et al.b Saldarelli et al.: Leafroll in Taiwan.: Production of antisera to GLRaVs using electrophoretically separated coat protein subunits as antigens.1991 Hu et al. Minafra and Hadidi: Detection of GLRaV-3 in viruliferous mealybugs by PCR.: Leafroll and associated closteroviruses in Moldova. 1994 1994 1994 1994 1994 1995 1994 1995 1995 1995 1995 Merkuri et al.: Leafroll and associated closteroviruses in Ukraine. Observation of peripherically vesiculated mitochondria. denoted GLRaV 2a and GLRaV 2b. Leafroll and associated closteroviruses in Yemen. Bondarchuk et al.: Purification and physico-chemical characterization of grapevine corky bark associated virus. Boscia et al. ELISA is recommended for large screening. Namba et al. ISEM and dsRNA analysis.: Production of radioactive and non-radioactive molecular probes to GLRaV-3 from denatured dsRNA template and their use for virus identification.: Analysis for the presence of double-stranded RNAs can be used for assessing virus elimination following sanitation treatments. 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.: Leafroll and associated closteroviruses in Albania. Gozsczynski et al. later identified as GLRaV-2. Pop et al. 66 . whereas the other assays are more suitable for analyzing samples that gave inconclusive results with ELISA.: Leafroll and associated closteroviruses in Spain 1991 1991 1991 1991 1991 1991 1992 1993 1993 1993 1993 1993 1993 1994a.
Wilcox et al. Fortusini et al.: GLRaV-3 is transmitted by Planococcus citri in a semipersisten manner.: Use of degenerate primers for PCR detection of GLRaV-4 and GLRaV-5.: Leafroll and associated closteroviruses in Lebanon.: GLRaV-3 detected in native American vines in Western New York. Al-Tamimi et al. La Notte et al.: The sequenced genome of GLRaV-2 has the same structural organization of that of Beet yellows virus.: Serological characterization of GLRaV-6 and production of monoclonal antibodies. Lahogue and Boulard: Search for genes of resistance in grapevines. the type specied of the genus Closterovirus. and Asian Vitis species inoculated by green grafting with a GLRaV-1 and GLRaV-3 sources were resistant. Alkowni et al. Ling et al.GLRaV-5 induces mitochondrial vesiculation. Faoro: Comprehensive review of the ultrastructure of GLRaVs infections.: Association of GLRaV-2 in Italy and France with a graft incompatibility revealed by Kober 5BB. American. Haidar et al. Zhu et al.: Identification of two different mechanically transmissibile strains of GLRaV-2. Martelli et al.: Leafroll and associated closteroviruses in Jordan. Habili and Nutter: In an Australian commercial vineyard GLRaV-3 incidence increased from 23.: Transmission of GLRaV-1 by the soft scale insects Parthenolecanium corni and Neopuvinaria innumerabilis.1% in 1986 to 51. Cabaleiro et al. Ling et al.: Comprehensive review of the properties of GLRaVs.: Distribution and incidence of GLRaVs in Canadian viticultural districts.: Elimination of GLRaV-3 by heat therapy improves agronomic performances of a Nebbiolo clone and the quality of the must. MacKenzie et al.: Development of a spot-PCR technique for GLRaVs identification.: GLRaV-2 and GLRaV-3 genes engineered in grapevine rootstocks to induce resistance.9% in 1996. Gugerli et al.: Cloning an sequencing of the coat protein gene of GLRaV-3 and its expression in transgenic tobacco.: Identification of GLRaV-7 and production of a polyclonal antiserum.: Leafroll and associated closteroviruses in Palestine. Choueiri et al. GLRaV-3 appears to be a typical closterovirus. No vector was identified. Gozsczynski et al. Routh et al. Viruses are irregularly distributed in the vines. Guidoni et al.: Extensive sequencing of GLRaV-3 genome. None of 223 accessions of European. 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1998 1998 1998 1998 1998 1998 1998 67 . Petersen and Charles: Pseudococcus calceolariae acts as vector of GLRaV-3 in New Zealand. Rowhani and Uyemoto: Comparative trials between indexing and laboratory detection methods show that the latter are more sensitve for GLRaVs detection. Krastanova et al.1995 1996 1996 1996 1996 1996 Greif et al.
: Revision of the family Closteroviridae.: Mealybug species Pseudococcus longispinus. Abou Ghanem -Sabanadzovic et al. affinis and Planococcus citri transmit with various efficiency GLRaV-3 in California but not GLRaV-1. Fazeli and Rezaian: Partial sequencing of GLRaV-1 genome.: Evidence that GLRaV-1 and GLRaV-3 are serologically related based on the cross-reactivity of a monoclonal antibody raised to GLRaV-1. a new genus having GLRaV-3 as type species.: Leafroll and associated closteroviruses in South Korea. Demonstration that the membranous vesicles accumulating in the cytoplasm derive from proliferation of the endoplasmic reticulum. GLRaV-2 and GLRaV-4. 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.: Association of an unusual strain of GLRaV-2 with a graft incompatibility condition described from California as young vine decline.: Completion of the nucleotide sequence of GLRaV-3 genome and use of the HSP90 and coat protein genes for producing transgenic rootstocks.: New monoclonal antibodies to GLRaV-2. Castellano et al: Ultrastructural study of GLRaV-2 and GLRV-7 infections. Kim et al. Evidence of the quasispecies nature of the virus.: Identification of hypervariable regions in GLRaV-1 genome. Gonsalves: Review article on the molecular traits of GLRaVs. Mannini and Credi. viburni.: Evidence that vines sanitized from leafroll have superior qualitative and quantitative traits.: Intriguing association of GLRaV-6 with cv. Martelli et al. Little et al. Seddas et al.: New vectors of GLRaV-1 are the mealybugs Heliococcus bohemicus and Phenacoccus aceris and the soft scale insect Pulvinaria vitis. Establishment and description of Ampelovirus . Cardinal in Italy and Greece. Ling et al. Boscia et al. Monis: Identification of GLRaV-8 and production of monoclonal antibodies. Gugerli: Detection of GLRaVs by Lumino-ELISA. Proposal for the establishment of Vinvirus (Ampelovirus) . transmitted by mealybugs and soft scale insects. maritimus. Zhou et al. Ps. Rowhani et al. Digiaro et al.: Completion of the nucleotide sequence of GLRaV-2 genome. Karasev: Review article on closteroviruses. one of which is especially suited for ELISA testing.: 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. Ps. Golino et al.: Survey of GLRaVs in Mediterranean and Near East countries.1998 2000 2000 2000 Saldarelli et al. Meng et al.: Identification and partial characterization of a new strain of GLRaV-2. Sforza et al. Turturo et al. a genus with monopartite RNA species. Golino et al.: Use of degenerate primers for PCR detection of GLRaV-1 and GLRaV-7. : Partial sequencing of GLRaV-7 genome. Production of a new set of monoclonal antibodies. Ps.
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P. Use of degenerate primers for partial sequencing and RT-PCR-based assays of grapevine leafroll-associated viruses. and P.P. 1998. European Journal of Plant Pathology 104. Minafra. 135-141. 1993. C. Kiryat Anavim 1987. Kiryat Anavim 1987. Detection of grapevine leafroll-associated closterovirus III by molecular hybridization. D. Histological and cytological studies on the infectious leafroll disease of the grapevine.. 35-38.. 1989. 207211.P. Sforza R. D. Maladies de l'enroulement et du bois strié de la vigne: analyse microscopique et sérologique. Martelli. Sela and I. Arboriculture.E. Volos 1990. Die Rollkrankheit des Rebstockes. 68-69.. 2000. 1989 Detection of a grapevine leafroll-associated closterovirus in recently infected tissues in New York and spread of the disease in Mexico. 1924. Zhang. Gonsalves and F. Martelli. Tanne E. 356358. 1997. Walter. and J.K. P. 157-160. Von der Brelie D. Iri. Transmission of grapevine leafroll disease and an associated closterovirus to healthy grapevine by the mealybug Planococcus ficus Signoret. 345-346.A..P.P. Walter. Transmission of closterolike particles associated with grapevine leafroll by mealybugs (Pseudoccidae) in Israel. I. Gugerli. 2000. Adelaide 2000. Tanaka S.. Rowhani A. Establishment of a relationship between grapevine leafroll closteroviruses 1 and 3 by use of monoclonal antibodies. Scheu G. Seddas A.C. Turturo C. A. 17-18. Takezawa and M. Extended Abstracts 11th Meeting of ICVG. Uyemoto.. Yafeng. 1976. Partial molecular characterization and RT-PCR detection of grapevine leafroll-associated virus 7. Montreux 1993. Il rossore delle viti.. Tada. D. Savino V.J. Sannino F. New scale insect vectors of grapevine closteroviruses. Greif. M. 704-709. 1994a. Rowhani.P. Saldarelli. 14. D'Onghia and G. Nitzany.. Segura A. Turturo C. 1906. V. Boscia. Rivista di Patologia Vegetale 1. Extended Abstracts 13th Meeting of ICVG. K. Adelaide 2000. Minafra and M. M. Use of degenerate primers in a RT-PCR assay for the identification and analysis of some filamentous viruses. 11-17. Zhang . Tanne. Tanne E. Plant Pathology Bulletin 3. Rowhani A. Martelli and B. Zee. Routh G. Y.. Indexing grapes in Japan for viruses. Gugerli 1989.S. 1987.. Phytopathology 88. Saldarelli. Annals of the Phytopathological Society of Japan 42. G. P. Saldarelli P. Proceedings 9th Meeting of ICVG. Komar and C.A . 1973. Reichnährstand Verlag. Rowhani. 169175. W. M. 156-167. Y.. 80-85. 71-73. 110-113.. 67-69. E. 82. Preliminary investigations of genetic variability of Grapeviune leafroll associated virus 3 isolates. Teliz D. Regeneration of plantlets by meristem tip culture for virus-free grapevine.. Cloquemin and B. Cabaleiro. Martelli. 945-950.D.. Tzeng H.M. C. Der Deutsche Weinbau 14. Rosciglione B. 799-801. and J. Vitis 33.. Ben-Dov and B. Saldarelli and A. Sasahara H. Haidar. A. 1936. Greif. J. V. Proceedings 9th Meeting of ICVG. Saldarelli P. Scheu G. A. Transmission of grapevine leafroll virus to herbaceous plants. Rott.M. M. Minafra. A comparison between serological and biological assays in detecting grapevine leafroll-associated viruses. 222-223..L. with special reference to closteroand vitiviruses of the grapevine. Revue Suisse de Viticulture. 1238-1243. 1986. Progrès Agricole et Viticole 45. Extended Abstracts 13th Meeting of ICVG. Uyemoto. Plant Disease 81. Extended Abstracts 14th Meeting of ICVG. 1998.. Verge. Proceedings 10th Meeting of ICVG. Teliz D. Martelli.Ravaz L. M. Tazaki. A. 508-517. Vitis 12. Hu and D. Phytopathologische Zeitschrift 80. Nienhaus. and G. Plant Disease 71. Y. Golino..P. Jacquet. Gonzales and C. D. Horticulture 18. Effect of heat therapy and meristem tip culture on the elimination of grapevine leafroll-associated closterovirus type III. P. Virus diseses of grapevine in Israel. 192-196. 1994. 222-225 Tanne E. Saldarelli P.L. 1994b. Journal of the Japanese Society for Horticultural Science 50. 1974. Digiaro. Presence of grapevine leafroll in North West Spain. 1935. 74 . A. Zeitschrift für Pflanzenkrankheiten und Pflanzenschutz 89. Mein Winzerbuch. 1981. M. Plant Pathology 49. Le rugeau de la vigne. Harpaz. 162-163. 176-180. and F. Digiaro. and F. Extended Abstracts 13th Meeting of ICVG. Isolation and partial characterization of two new viruses from grapevine. H. Rosciglione B. G. Hummer. Adelaide 2000. Savino and G. 38.. 2000. 91-96.S. Chen and D.K. 1982a. Guglielmi Montano and G. 433-436. and P. D. Field serological detection of viral antigens associated with grapevine leafroll disease. Jelkmann and G. Raccah. Phytoparasitica 17. 125-126. Non radioactive molecular probes for the detection of three filamentous viruses of the grapevine. Locorotondo 2003. 8689. Plant Pathology 43.. Tzeng. The occurrence of grapevine leafroll disease among the main grapevine cultivars and breeding stocks in Taiwan. 2000.. Gonsalves.K. T. Berlin. 1991.
Phytopathology 77. Z. Lee. Adelaide 2000. Legin. Production and characterization of monoclonal antibodies specific to closterovirus-like particles associated with grapevine leafroll disease. D. R. Wilcox F. 313-316. McFerson and D. Goszczynski and M. N. Journal of Phytopathology 130. 75 . The use of a greengrafting technique for the detection of virus-like diseases of the grapevine. D. 1427-1434. D. Agronomie 8.Y.Vuittenez A. Vernoy. Gonsalves. Zimmermann D. Walter and M. Le Gall. Nucleotide sequence and genome organization of Grapevine leafroll-associated virus 2 are similar to Beet yellows virus. 2003. Boscia.E Goszczynski. 1062. A. Potere.. 1987. O.F. Potere. 1958. 1990. Pool and R.V. G.P.. Kim. P. Vesselle. Ling. Zimmermann.Y. A. Bass. and D. Zhu H. Extended Abstracts 13th Meeting of ICVG. Further characterization of closterovirus-like particles associated with the grapevine leafroll disease. 2000..R. Zimmermann D. Extended Abstracts 14th Meeting of ICVG. Castellano. Saldarelli. Jiang and D. Cytopathology of leafroll diseased grapevines and the purification and serology of associated closteroviruslike particles. C. R. Monoclonal antibodies for detection and characterization of Grapevine leafroll-associated virus 2. 1289-1298. Abou-Ghanem. Plant Disease 82. and G. Zhou Z. D. 1998 Leafroll virus is common in cultivated American grapevines in Western New York.. Collas and G..S. J. 1990. Martin. O. Van Regenmortel. Zee F.H. Volos 1990. C. Journal of Phytopathology 128. 731-741. K. Zhou Z. R. Production and characterization of monoclonal antibodies specific for Grapevine leafroll-associated virus 3 and epitope mapping of the coat protein. 130. Locorotondo 2003.A. Boscia. B.. Purification de particules virales associées à l'enroulement de la vigne et mise au point d'un protocole ELISA permettant leur détection.W. 1988. Goheen. 277-288. 137-145. 1991. B. 1998.E. Walter B. K. Comptes Rendues de l'Academie d'Agriculture de France 44. Gonsalves.S. Turturo.. Walter B. Sommermeyer. Gonsalves. 203. Proceedings 10th Meeting of ICVG. D. Transmission par greffage d'une virose type enroulement foliarie commune dans le vignobles de l'Est e du Centre-Est de la France. Martelli. 62-66. the closterovirus type member. P. Journal of General Virology 79. Walter and O..
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
In general. to a restricted range of herbaceous hosts (mostly Nicotiana species). Histological study of diseased vines. but not foveaviruses. the putative agent of rupestris stem pitting. experimental evidence has been obtained of the very close association of GRSPaV.: Corky bark is remarkably heat stable and difficult to eliminate by heat therapy. Graniti: Detailed description of rugose wood symptoms.were observed in self-rooted cultivars and ungrafted roostock stocks (V. though with difficulty. Using a Nicotiana benthamiana model system. The best antigen sources for serological diagnosis are cortical shavings from mature dormant canes. since symptomless infections make sanitary selection not totally reliable. with vein necrosis. GVA can be eliminated to a very hight rate (up to 97%) by the procedure used for cryopreservation of grapevine shoot tips. Recently. Thus. Name of the disease changed into corky bark. Thus. 2. rupestris. several resistant plant lines were obtained by transformation with the coat protein and the movement protein genes of GVA and GVB. Graniti and Martelli: Demonstration of the infectious nature of rugose wood. In addition. other assays include: single step or nested reverse trascription-polymerase chain reaction (RT-PCR).: First record of rugose wood outside of Italy. Suggestion that rugose wood may be a disease of combination requiring the contact of scion and rootstock for the development of symptoms. 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. 1965 1965 1967 81 . Immuno-capture RT-PCR is 1000-fold more sensitive than ELISA for virus detection in grapevines. "stem pitting" and "stem grooving". Faccioli: First histological study of corky bark-affected grapevines. Martelli et al. Graniti and Ciccarone: First record of rugose wood from southern Italy. Individual viruses can be identified by ELISA or dot immunobinding on nylon membranes using polyclonal antisera and/or monoclonal antibodies when available. Goheen et al. Kober 5BB and LN 33) is the only reliable method for detecting and sorting out the diseases of the complex. described from California. Hewitt et al. and that it may be a composite disease resulting from the interaction of different viruses among which GFLV. are synonymized with "rugose wood". meristem tip culture. all sources must be indexed and/or laboratory tested. Historical review Names like "legno riccio". Transgene expression was detected in these plants and in transformed grapevine explants. Suggestion that it may be caused by a virus. rupestris and Kober 5BB). possibly in relation with the scion/stock combination and climatic conditions. Control of mealybugs is difficult for they overwinter under the bark of grapevines and possess an unwettable waxy covering. Beukman and Goheen: Brief account of the histological modifications of corky bark-affected LN 33. immunocapture RT-PCR. Vitiviruses. Latent infection can occur also in grafted vines.: Graft transmission of rough bark to LN 33. The intensity of wood abnormalities (pitting and grooving) vary. However. 1954 1961 1962 1963 1963 1964 1965 Hewitt: Rough bark. rupestris. rugose wood agents can be eliminated with reasonable efficiency by heat therapy. or spot-RT-PCR using degenerate or virus-specific primers. 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. as shown by 110R. it is plausible to regard 110R as a specific indicator of rupestris stem pitting in addition to V. Detection: Indexing on indicators (V. if not otherwise associated with a specific syndrome. no strategy has yet been developed for the chemical control of vectors. or a combination of the two. a virus-like disease. are mechanically transmissible. Goidanich and Canova: First record of corky bark in Europe.
Rugose wood may not require a grafted plant for full symptom expression. Castillo et al. No nepoviruses implicated in their etiology. Boccardo and D'Aquilio: Physicochemical characterization of GSP-AV Abracheva: Survey of over 650 grapevine cultivars and hybrids for rugose wood reaction in Bulgaria.: Green grafting useful for corky bark indexing.: Heat therapy effective against rugose wood.b. Engelbrecht and Nel: Rugose wood and fanleaf are not related. from a rugose wood-infected vine. Goheen: Evidence that corky bark and leafroll. Mink and Parsons: Use of growth chambers for rapid symptom expression of corky bark in Vitis indicators. 1968 1969 1970 1970 1971 1971 1972 1973 1973 1975 1975 1977 1978 1979 1979 1980 1980 1981 1981 1982 1983 82 .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. First record of rugose wood symptoms outside of Europe (Israel). Teliz et al. Lehoczky: Destructive effects of rugose wood registered in Hungary in both self-rooted and grafted European grape varieties. Conti et al. Graniti and Martelli: Up-to-date review of rugose wood. is transmitted by the pseudoccid mealybug Pseudococcus longispinus. based on graft transmission tests. despite similarities in the s y m p t o m s o n t h e foliage are different diseases. Goheen and Luhn: Heat treatment of dormant buds grafted onto LN 33 is effective against corky bark. Rosciglione et al. Anonymous: A review of rugose wood in Italy. Legin et al. Goheen and Luhn: Suggestion that corky bark and rugose wood are the same disease.: First experimental evidence that a filamentous virus (GVA).: Observation of rugose wood symptoms in self-rooted vines. At 38 °C the minimum inactivation period for leafroll is 56 days and for corky bark 98 days.: Recovery by mechanical inoculation of a closterovirus with particles 800 nm long.: Rugose wood recorded from Australia.Virus provisionally called grapevine stem pitting-associated virus (GSP-AV). a. Bovey and Brugger: Further evidence that GFLV may not be implicated in the etiology of rugose wood in Switzerland. Hewitt: Successful graft transmission of Californian rugose wood. Beukman and Goheen: Up-to-date review of corky bark. Beukman and Gifford: Detailed account of adverse effects of corky bark on the anatomy of Vitis. Hewitt and Neja: First record of rugose wood in USA (California). Hewitt: Up-to-date review on grapevine virus and virus-like disease worldwide.1968 1968 Lehoczky et al. Sarooshi et al.
Monette and James: Detection of two biologically distinct but serologically indistinguishable isolates of GVA.3 and 4. Engelbrecht and Kasdorf: Natural field spread of corky bark in South Africa associated with the presence of P. an additional disease of the rugose wood complex. Monette et al. The first two disorders appear to be spreading in the vineyards. thesis describing rupestris stem pitting disease in comparison with corky bark. First report of Kober stem grooving. Rosciglione and Castellano: Demonstration that GVA is transmitted also by Planococcus citri and P.: Two distinct dsRNAs with a mol. ficus. but not consistently in grapevines from New York. Kuniyuki and Costa: Rugose wood recorded from Brasil Goheen: First published description of rupestris stem pitting. Corky bark and Rupestris stem pitting.ficus. ficus.: Evaluation of the effect of rugose wood on cv.: Heracleum latent virus and GVA are distantly serologically related. No closterovirus-like particles in samples with rupestris stem pitting. Corbett and Wiid: Closterovirus-like particles found in extracts from vines affected by corky bark and rugose wood in South Africa. Castrovilli and Gallitelli: Physicochemical comparison of two Italian isolates of GVA. Prudencio: M.: Assessment of crop losses induced by rugose wood to two different European grape varieties. Savino et al.: Experimental confirmation of the complex nature of rugose wood based on the differential reaction of woody indicators. denoted Grapevine virus B (GVB). Rugose wood and corky bark are not the same disease. Engelbrecht et al. Savino et al.: First record of rugose wood from China. Suggestion that the disease is not related to closteroviruses associated with GLRaV and corky bark. Italia propagated on six different rootstocks. Gallitelli et al.: Evidence that GSP-AV can occur in grapevines together with another similar but serologically unrelated virus with short closterovirus-like particles. Martelli: Rugose wood recorded in southern Mediterranean and Arab countries. Similar dsRNA species were detected.: Application of spot hybridization for the detection of GVA in grapevine sap.: First indication of the possible existence of LN 33 stem grooving.: Experimental confirmation that rugose wood may not express symptoms in grafted indicators.4 x 106 associated with rupestris stem pitting in grapevines from California and Canada.: Three types of wood disorders of the stem-grooving type observed in South African grapevines. GSP-AV re-named Grapevine virus A (GVA).: A low molecular weight dsRNA associated with rupestris stem pitting. Savino et al.Sc. Garau et al. Murant et al.1984 Milne et al. Garau et al. Azzam et al. Tanne et al. 1985 1985 1985 1985 1985a 1985b 1985 1985 1985 1987 1988 1989 1989 1989 1989 1989 1989 1990 1990 1991 1991 83 . wt of 5. similar to Kober stem grooving. Li et al.: Transmission of corky bark by the mealybug P.
: Rugose wood recorded from Albania. Namba et al. 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.: Purification and properties of GVB.: 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. Thorough comparative study of nine GVB isolates from different countries. Martelli et al.: Development of digoxigenin-labelled riboprobes for the detection of GVA and GVB in infected tissue extracts.: Production of monoclonal antibodies to GVA and their use for ELISA detection of the virus in infected vines. 1991 1991 1991 1991 1991 1991 1991 1991 1992 1992 1993 1993 1993 1993 1994 1994 1994 1994 1994 1994 1994 1994 1995 84 . Minafra and Hadidi: Detection of GVA and GVB in viruliferous mealybugs by PCR. Suggestion that GVA may be the causal agent of the disease.: A closterovirus with particles 1440-2000 nm long serologically unrelated to all other known grapevine closteroviruses found in corky bark-affected vines.: Rugose wood recorded from Ukraine Boscia et al.: Clear-cut connection of GVA and rugose wood.: Sequence of the 3' end of GVA and GVB genome. Virus named Grapevine virus C (GVC). Torbato in Italy. both associated with a stem pitting condition of grapevines rather than with leafroll.: Contemporary occurrence of Rupestris stem pitting and Kober stem grooving in symptomless scions of cv. Virus transmission by the mealybug Ps. Saldarelli et al.: Presence of two distinct serotypes of GVA. Merkuri et al. Suggestion that GVA is implicated in the aetiology of the disease. Boscia et al.: Synthesis of a cloned probe for GVA.: Rugose wood recorded from Malta Monette and Godkin: Recovery of a closterovirus-like virus by mechanical inoculation from a corky bark-affected vine. Garau et al.: Development and diagnostic use of a cloned probe to GVB. Both viruses qualify for the inclusion in the genus Trichovirus. Minafra et al.1991 1991 Gugerli et al. Boulila et al. Digiaro et al. Minafra et al. Martelli et al. Saric and Korosec-Koruza: Rugose wood recorded from Croatia and Slovenia Ioannou: Rugose wood recorded from Cyprus.:Rugose wood recorded from Tunisia Milkus et al.: GVA and Kober stem grooving are closely associated. Padilla: Rugose wood recorded from Spain Boscia et al. Garau et al. ficus induced corky bark symptoms in LN 33 Saldarelli et al. Chavez and Varon de Agudelo: Rugose wood recorded from Colombia.
: GVA and GVD are serologically distantly related. GRSPaV is assigned to this genus. though not consistently in vines showing a syndrome denoted "corky rugose wood". GVB.: GVB is consistently associated with corky bark and is present. Chevalier et al.: GVA and GVB are transmitted by Pseudococcus affinis. Further support of the cause-effect relationship between GVA and this disease. Minafra et al. Goszczynski et al. Zhang et al. this may be the first visualization of GRSPaV.: GVA is transmitted by by Ps. Guidoni et al.: Consistent detection of GVA in Kober stem grooving-infected grapevines by immunocapture-polymerase chain reaction.: Review of the properties of putative grapevine-infecting trichoviruses (GVA.: Estasblishment of the genus Vitivirus with GVA as type species. The virus is not seed-borne. GVA. Saldarelli et al.1995 Monette and Godkin: Detection of non mechanically transmissible capillovirus-like particles in a grapevine affected by rugose wood.: Sequencing of a Californian isolate of GRSPaV. Boscia et al. GVC.: Rugose wood recorded from Lebanon. and GVD removed from the genus Trichovirus and assigned to the new genus.: Description of Grapevine virus D (GVD). Martelli and Jelkmann: Establishment of the genus Foveavirus. GVC. longispinus in a semi-persistent manner. Bonavia et al. and GVD) later assigned to the genus Vitivirus. 1995 1995 1995 1996 1996 1996 1996 1996 1997 1997 1997 1997 1997a 1997b 1997 1997 1997 1997 1998 1998 1998 1998 85 .: A study of the spatial distribution pattern of corky bark in aThompson seedless vineyard in Israel. Suggestion that spreading is by a vector that transmits in a semipersistent manner. Garau et al. Boscia et al. Haidar et al. Meng et al.: Nucleotide sequence of GVA genome and taxonomic position of the virus.: Sequence and structrural organization of Grapevine rupestris stem pittingassociated virus genome (GRSPaV). Alkowni et al: Rugose wood in Palestine.: Rugose wood recorded from Jordan. Efficient detection method based on TAS-ELISA developed. GVB.: Elimination of GVA from cv Nebbiolo clones by heat therapy improves agronomic performance of the vines and quality of the must. Choueiri et al. La Notte et al. Tanne et al. Martelli et al. Since particle size (600-700 nm in length) is compatible with that of Grapevine rupestris stem pitting-associated virus (GRSPaV) particles identified in 2002.: 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.: GVA and GVB are serologically related. Rubinson et al.: Nucleotide sequence of GVB genome. La Notte et al.: Development of a PCR technique for the detection of GVA and GVB in nylon membrane-spotted sap.
: GVA transmission by Pseudococcus comstocki. Nucleotide sequence identity within groups is 91-99.: Movement proteins of GVA and GVB detected by gold immunolabelling in association with cell walls and plasmodemata of infected cells. Meng et al.: Rugose wood viruses in Egypt.: The function of GVA genes identified by mutation analysis of individual ORFs of a full-length infectious viral clone. Galiakparov et al. Buzkan et al. intermingled with virus particle aggregates. i. Kominek et al.: One-sided phenotyping mixing. GVB and Corky bark and GRSPaV and Rupestris stem pitting. GVA coat protein encapsidating GVB RNA.: GRSPaV particles.: GVA particles carry a highly structured epitope centered on a common peptide region of the coat protein sequence. Petrovic et al.: Rugose wood viruses in the Czeck Republic. Saldarelli et al.1999 1999 2000a Meng et al.3% among groups. Stewart and Nassuth: An improved extraction method allows RT-PCR detection of GRSPaV virtually thoughout the year in all grapevine tissues. Habili et al. 2000b 2000 2001 2001 2001 2002 2002 2003 2003 2003 2003 2003 2003 2003 2003 2003 2003 2003 86 . GVA movement protein is also present in great quantity in the cytoplasm. are filamentous and measure 723 nm in length. Goszczynski and Jooste: Use of single-strand conformation polymorphism reveals molecular heterogenity in GVA populations. The virus was not detected in 245 seedlings from infected cv.: Production of monoclonal antibodies to GVB. Further support to the cause-effect relationship of GRSPaV with this disease Galikparov et al.: Production of an infectious RNA transcript from a full-length cDNA clone of GVA. occurs in Nicotiana plants doubly infected with GVA and GVB. Boscia 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. 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. Confirmation of the cause-effect relationship between GVA and Kober stem grooving. Virus detected in bleached seeds suggesting that it is present inside the seeds. observed for the first time.: Consistent association of GRSPaV with vines indexing positive for Rupestris stem pitting. GCD) and foveavirus (GRSPaV) sequences in two steps. Samples made up of three buds from dormant canes are less laboriour to prepare than cane shavings and yield comparable results. Dovas and Katis: Improved RT-PCR method for the simultanous detection in grapevine extracts of vitivirus (GVA. and III) identified in South Africa based primarily on sequence homology of the 3' end of the viral genome. Dell'Orco et al.: Rugose wood viruses in Iran. Ahmed et al. Seyval seedlings.e. Goszczynski and Jooste: Thre groups of GVA strains (I. II. Saldarelli et al. Wang et al. Nakano et al. GVB.: Elimination of GVA by cryopreservation. Minafra et al.8% and 78-89.: Synthesis of full-length cDNA copies of GVA and GVB genomes.: Production of a polyclonal antiserum to a recombinant coat protein of GRSPaV and its use in dot immunobinding on polyvynil difluoride membranes for virus detection in grapevine tissue extracts.
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GRAFT INCOMPATIBILITY .
so as to represent veritable emerging diseases. Transmission: GLRaV-2.g. appears to be involved in California's young vine decline. 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. New Zealand. California. shoots are short. though not consistently and. but the yield is reduced. with margins more or less extensively rolled downwards. Australia and Chile (young vine decline). Infected propagative material is to be blamed for its dissemination. Chile. Other molecular variants of GLRaV-2 were reported from New Zealand (Alphie virus). The heat-labile graft-transmissible agent present in the hybrid 140R. only GLRaV-2 RG was identified. whereas varieties grafted on susceptible roostocks (e. and together with Grapevine virus B (GVB).). 5C. A transitory form of incompatibility was reported from Italy under the name of bushy stunt. associated with grapevine bushy stunt is still unidentified. Argentina and probably elsewhere. 101-14) exhibit a green canopy and perform rather well. 03916. Normal growth resumes with the second or third leaf. scions show a stunted and bushy vegetation due to the contemporary proliferation of apical and axillary buds. However. A virus originally detected in cv. Severely affected vines decline and may die within one or two years. which are usually not accompanied by wood abnormalities (pitting or grooving). In this case. decline and may die. up to five different grafttransmissible agents inducing incompatibility could be differentiated in California. the increased use of clonal material is disclosing unprecedented conditions of generalized decline that develop dramatically in certain scionrootstock combinations. 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. Kober 5BB. Freedom. Cases of graft union disorders have been observed in Europe (Kober 5BB incompatibility). Based on the differential responses of a panel of 18 rootstocks. This latter condition has been known for a long time and occurs also in rugose wood-affected vines. leaves are small-sized. is not transmitted by mealybugs and does not have a known vector. Detection: Indexing on Caberent sauvignon is a reliable method for detecting incompatibility conditions. A prominent swelling forms at the scion/rooststock junction and variously extended necrotic lesions may develop on the rootstock stem. 1. Agents: An ordinary strain of Grapevine leafroll-associated virus 2 (GLRaV-2) is consistently associated with Kober 5BB incompatibility (Europe). except for GRSPaV. Syrah decline is a severe disease occurring in France.g.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). 1103P. 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. 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). Of these. in Argentinian grapes. 3309) develop a discolored canopy . incompatibilità d'innesto (Ital. Harmony. and Australia in association with young vine decline conditions. although none of a number of known grapevine-infecting viruses has been found in affected vines. but the colour of the canopy remains green. a member of the genus Closterovirus.) Symptoms: Newly planted vines grow weakly. consistently. and again California (rootstock stem lesions). Other assays include nucleic acid-based tecniques such as single step or nested reverse transcription-polymerase chain reaction (RT-PCR) and 95 . 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. Salt creek . Description Main synonyms: Incompatibilité au greffage (Fr. The same virus was detected in diseased Chilean grapes. The nature of this disease has not been ascertained but one or more graft-transmissibile agents may be involved in its aetiology. European grape varieties grafted on tolerant rootstocks (e. GVB is mealybug-borne and can be spread at a site by these insects. The putative agent of bushy stunt was consistenly found in clones of the rootstock 140R in which it is latent. Varietal susceptibility: Appearance of graft union disorders depends more on the rootstock rather than the scion.
No conclusion are drawn. Gomez Talquenca et al: GLRaV-2 is consistently associated with declining Cabernet sauvingon vines grafted on different roostocks in Argentina. or a combination of the two. Historical review 1942 1950 1973. whenever feasible. 1977 1979 1986 1991 1995 2000 2000 2000 2001 Jacob: Description of graft incompatibility in different scion/stock combinations.: GLRaV-2 and GVB are consistently associated with young vine decline in California. Currently known graft incompatibility agents can be eliminated with reasonable efficiency by heat therapy. Uyemoto and Rowhani: Indexing on 18 different grape rootstocks reveals the existence of at least five different agents causing graft incompatibility. Bertazzon and Angelini: Comparison of several detection methods for the broad or specific identification of Grapevine leafroll-associated virus 2 variants. The problem is very complex and may involve several still unidentified factors. though not consistently. Greif et al.: GLRaV-2 is the cause of a graft incompatibility revealed by Kober 5BB. GRSLV re-named Redglobe strain of GLRaV-2.immunocapture RT-PCR. 2003 2003 2003 2003 2003 2003 2004 96 . Durquety et al. decline. utilization of tolerant roostocks is advisable. Boubals: Report of a national French study group investigating the aetiology of Syrah decline.: GLRaV-2 is associated.: Report of a new molecular variant of GLRaV-2 from New Zealand. 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. and death of the vines. Prodan et al. Golino et al. Suggestion that they are molecular variants of the same virus species.: Third paper of a series on incompatibility on Kober 5BB. the use of sensitive rootstocks is to be avoided and. If scionwood is infected. Strategies on how to protect healthy stocks from vector-mediated GVB reinfection in the field are yet to be developed.: Identification of an apparently new closterovirus denoted Grapevine rootstock stem lesion virus (GRSLV) causing stem necrosis of rootstocks. with a decline condition of young Thomposn seedless vines in Chile. 2. GRSLV has about 75% nucleotide homology with GLRaV-2. Boubals: Syrah decline occurs in Argentina Uyemoto et al. Renault Spilmont et al. Graft-transmission of the incompatibility factor. using degenerate or virus-specific primers.: Two papers describing incompatibility phenomena between clonal selections of different cultivars grafted on Kober 5BB Fallot et al. Bonfiglioli et al.: Description of bushy stunt and evidence that it is caused by a graft-transmissibile heat-sensitive agent carried by some clonal rootstocks. Martelli: GRSLV and GLRaV-2 are serologically related and are both recognized by a panel of 18 monoclonal antibodies. meristem tip culture. Boubals and Huglin: Report on graft incompatibility of certain varieties grafted on 57R.: Updated report on the state of the art of investigations carried out in France on Syrah decline. Control: Prevent introduction of infected vines in the vineyard by using certified grafted plants or virusfree scionwood and rootstocks. Savino et al.
Martelli G. Rowhani. 1973. Durquety P. Ruchaud. Renault Spilmont A.M. 167-173.. Nouvelles recherches sur l'incompatibilité clonale d'Abouriou greffé sur 5BB. Luvisi and R. 2003. 2003.E. Extended Abstracts 14th Meeting of ICVG. A young grafted vine decline syndrome in Argentina vineyards.K. The relationship of grapevine leafroll-associated virus 2 with a graft incompatibility condition of grapevines.3. S. Progrés Agricole et Viticole 90. Autres cas chez la vinge.. D.. Le clone et ses reactions au greffage. Proceeding of the American Society of Horticultural Science 41. Advances in the detection of Grapevine leafroll-associated virus 2 variants. Locorotondo 2003. Extended Abstracts 14th Meeting of ICVG. 1979. Progrés Agricole et Viticole 96. New closterovirs in 'Redglobe' grape causes decline of grafted plants. Boursiquot. C. Di Terlizzi. 144. Prodan S. References Bertazzon N. La trasmission de l'incompatibilité au greffage entre 5BB et Vitis vinifera.III. Fallot. 420-427 Fallot J. Etude de l'incompatibilité au graffege de certain cépages et du 57R. Bonfiglioli R. Rowhani. Sim and A. Fallot. Extended Abstracts 14th Meeting of ICVG. 2001.S. California Agriculture 55 (4). Boscia.M. 277.M. 85-86. Boubals D.. Savino V.P. Dauty. Prota. 85-86.. Extended Abstracts 14th Meeting of ICVG.M. Locorotondo 2003. A. 2000. Gazeau and J. Prota.K. Existence dans un cépage population de clones présentant divers degrés de compatibilité avec certains porte-greffes. Proceedings 10th Meeting of ICVG. 28-31. Di Silvio. 1991. Montalegre and N. 283-290. Le clone et ses reactions au greffage. Uyemoto J. 3-10. 2003. Extended Abstracts 14th Meeting of ICVG. 2000.S.P.. Grenan and J. Ruchaud. 1977. Huglin.. Examples of incompatibility between grape varieties and roostocks. Adelaide 2000. 201-203. M. 1950. 142-143. and B. 2004. 183-189. Volos 1990.. Gomez Talquenca G. 2003. Locorotondo 2003. Progrés Agricole et Viticole 117. C. Fiore. 137-141 Boubals D. I. and E. Phytopathologia Mediterranea 34. 211-216. S. Jacob H. Progrés Agricole et Viticole 103. Progrés Agricole et Viticole 117. Molecular studies on a graft incompatibility syndrome in New Zealand vineyards yields another probable variant of Grapevine leafroll-associated virus 2. and P.. Boubals D. 122-129 and 171-178.. Progrés Agricole et Viticole 67. O. Walter and U. Gracia. Krag. C. V. Walter. Locorotondo 2003. Syrah decline in French vineyards. 1995. J. Benassac and R. Aetiology of decline in Thompson seedless grafted table grape plants. Journal of Plant Pathology 86. Greif C. Ruchaud.A. Extended Abstracts 14th Meeting of ICVG. Garau. J. 2000. S. Le dépérissement de la Syrah. J. 141. 1986. Identification of the latent viruses associated with young vine decline in California. J. Rowhani. R. Fiori. B. Le dépérissement de la Syrah existe aussi en Argentine. P. D. Discovery of different grapevine sources with graft-transmissible agents causing union-incompatibility on sensitive rootstocks. and A.P. Progrés Agricole et Viticole 94. Compte-rendu de la réunion du Groupe de Travail National. Legin R. Rivieccio and F. 202-210 Uyemoto J. 2003. Locorotondo 2003. S.. F.. 1942. 97 . Locorotondo 2003. Presence in clonal rootstocks of a grafttransmissible factor that induces stunting and bushy growth in European grapevines. Grau. Extended Abstracts 13th Meeting of ICVG.. Etude de phénomènes d'incompatibilité au greffage chez la vigne.. Angelini. Edwards and A. Pantaleo..A. Durquety and J. Durquety P. Golino D. Bass. 2003. 139-140. B. P.II. Gazeau. Le clone et ses reactions au greffage. Grapevine virology highlights 2000-2003.P. 279-283. Garcia Lampasona and O.
FLECK COMPLEX .
6 x 106 ) is 7564 nt in size and contains four open reading frames (ORF) that 101 . 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. Fleck. Grapevine fleck: Marbrure (Fr. rupestris. maculatura infettiva. Sultanina).). Affected vines are often stunted. wt of 2. adverse influence on vigour. positive sense RNA with high cytosine content (c. Leaf symptoms usually become less severe in summer. c. GFkV. which are twisted and asymmetric. rupestris following graft inoculation. T made up of empty protein shells and B. The putative causal agent of the disease has only been found in California. and Grapevine rupestris vein feathering virus (GRVFV) are all phloem-limited and non mechanically transmissible. Marmorierung der Rebe (Germ. twisted and puckered along the veins. 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. Leaves with intense flecking are wrinkled. GAMaV. grapevine asteroid mosaic. 50%). capped. The disease is latent in European grapevine varieties and in most American rootstocks. sometimes with necrotic center. irregularly distributed over the leaf blade. vinifera in California. reported only from Japan. leaf petioles and veinlets. GFkV genomic RNA (Mol. Symptoms are expressed in Vitis rupestris and consist of clearing of the veins of third and fourth order.FLECK COMPLEX The fleck complex consists of several diseases (grapevine fleck. rooting ability of rootstocks and on graft take has been reported. GFkV particles sediment as two centrifugal components. Italy and California. Description Main synonyms: A.). twisted and may curl upward. Rupestris necrosis. but likely to occur elsewhere. In V. rupestris. Grapevine asteroid mosaic-associated virus (GAMaV).). 25 kDa. This disease. vinifera. In V. Grapevine red globe virus (GRGV) is a Grapevine fleck virus (GFkV)-like virus which apparently does not induce symptoms in European grapevine varieties (e. producing localized translucent spots. Asteroid mosaic symptoms have been observed in several varieties of V. screziatura (Ital. Red globe) nor in V. 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. Severe strains induce also varying degrees of stunting.g. Mild asteroid mosaic-like symptoms are shown by some European grapevine varieties (e. and GRVFV genomes are available.). which is used as indicator. GFkV genomic RNA constitutes about 35% of the particle weight. Symptoms: a. V. The coat protein (CP) of GFkV and GRGV particles is made up of a single protein species with Mr of c. Grapevine asteroid mosaic: Mosaïque étoilée (Fr. b. Asteroid mosaic.). mosaico stellare (Ital. Fleck is an ubiquitous disease reported from most viticultural countries in the world. Although the elusive nature of the complex hinders the assessment of its economic impact. which is a monopartite single-stranded. containing the genome. e. the disease elicits creamy-yellow bands developing along the major veins of the leaves. grapevine rupestris necrosis. Transient mild chlorotic discolourations of the primary and secondary leaf veins develop in V. Leaves are asymmetric. whereas the CP of GAMaV and GRVFV consists of a major protein of 21 kDa and a minor prtotein of 25 kDa. rupestris reacts with localized necrosis of the shoots. Rupestris vein feathering. leaf symptoms are characterized by star-shaped chlorotic spots. Recorded from California and Italy . Sternmosaik der Rebe (Germ. and produce little or no fruit. B.).g. Agents: All viruses of the complex. GRGV. d. is latent in European grapevine varieties. The putative causal agent of the disease so far has been found in Greece. 1. The complete sequence of GFkV and partial sequences of GRGV.
molecular and ultrastuctural studies disclosed sufficient similarities between maculaviruses. 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". whereas GAMaV induces peripheral vesiculation of chloroplasts.9 kDa (ORF 4) with unknown function. No information is available on individual susceptibility. GRGV. GFkV is not seed transmitted. the CP (ORF 2).George. 102 . Polyclonal antisera and monoclonal antibodies to GFkV heve been raised. of which it represents the type species. rupestris St. 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. Historical review 1954 1962 1966 1966 Hewitt: First description of asteroid mosaic in California. The same sanitation procedures are likely to operate successfully with the other viruses of the complex.4 polypeptide with the conserved motif of replication associated proteins (ORF 1). Further physico-chemical. but cannot be used for any of the other members of the complex due to the unvailability of antisera. whilst GAMaV and GRVFV were assigned to the genus Marafivirus. Although observations from Italy. GAMaV. 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. As the disease is rare and does not appear to be spreading. These deranged organelles are thought to be sites of virus replication. but no experimental data are available. Because of its molecular characteristics. a disease inducing symptoms similar to those of fleck in V.: "Marbrure". and two proline rich polyproteins of 31. Control: Because of the latency of symptoms sanitary selection of European grapevine cultivars and most American rootstock hybrids is unrealiable. Refatti: Review paper on asteroid mosaic. Comparison of symptoms with those of other mosaic diseases of grape.rupestris described in France. Varietal susceptibility: GFkV and possibly all the other viruses of the complex infect naturally a large number of varieties and Vitis species. Virus specific and degenerate primers have been designed for single or multiplex RT-PCR detection of GFkV. The 3' end of GRGV genome is structurally similar to that of GFkV except for the lack of ORF 4. its economic importance is low. 2. meristem tip or fragmented shoot apex culture. Detection: Indexing on V. primary dissemination of these and the other viruses of the complex is through infected propagative material. marafiviruses and members of the genus Tymovirus to warrant the establisment of the a new family denoted Tymoviridae. GFkV can be eliminated by heat therapy. Vuittenez et al. Transmission: No vector is known for any of the viruses of the fleck complex.encode a 215. 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. and GRVFV.: First record of fleck as an unidentified symptom different from fanleaf and transmissible from symptomless varieties to V. Hewitt et al. ELISA is currently employed for routine detection of GFkV. GFkV was identified as the representative of a new genus denoted Maculavirus.4 kDa (ORF 3) and 15. Therefore. Transmission through dodder of GFkV has been reported but it is of no epidemiological importance. Attempts to transmit the disease by mechanical inoculation to herbaceous test plants or by Xiphinema index were unsuccessful.
Confirmation that the virus can be eliminated by heat therapy and is not related to leafroll. Milkus: Suggestion of a prokaryotic etiology for fleck. The efficiency of heat treatment for disease elimination is unsatisfactory.: Purification of GPLIV from naturally diseased vines and production of a specific antiserum.: Report of widespread occurrence of fleck in visually selected grapevine clones in southern Italy. Castellano et al.: Successful elimination of fleck through heat therapy. Dismissal of the prokaryote etiology hypothesis. Triolo and Materazzi: Fleck has a detrimental effect on the quality V. Castellano et al. Report that a virus serologically similar to GPLIV is associated with the disease.: Identification of a dsRNA of about 7 Kb pairs in diseased vines.: Physicochemical characterization of GPLIV. Rives: Further demonstration that fleck is distinct from fanleaf based on differential responses to heat treatment. 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. Hévin et al. Woodham and Krake: Dodder transmission of fleck from vine to vine.: Observation of a non mechanically transmissible virus. Bovey: Identification of fleck in Switzerland as a latent disease of Chasselas transmissible to V. Verderevskaya et al. Barlass et al. 1982 1983 1983 1983 1984 1985 1985 1987 1989 1990 1990 1990 1991 103 . Savino et al. Ottenwaelter et al. in sieve tubes of field-grown vines with leafroll symptoms but likely to be affected by other diseases. Report of multivesiculate inclusion bodies probably connected with GPLIV infection.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. rupestris.: Fleck is not seed transmissible. Castellano and Martelli: Confirmation that GPLIV is associated with multivesiculate bodies and demonstration that these derive from deranged mitochondria. Goheen and Luhn: A novel heat therapy system based on virus inactivation in buds grafted onto healthy LN 33 rootstocks is effective against fleck.: Successful elimination of fleck through fragmented shoot apex culture in vitro. Triolo and Resta: Tetracycline treatments are ineffective against fleck.: Description of fleck as an independent graft-transmissible disease present in many European varieties and American rootstocks. leafroll and corky bark).: Observation of an isometric non mechanically transmissible virus in the phloem of diseased vines. Dolja et al. Engelbrecht and Kasdorf: Observation of natural field spread of fleck in South Africa. rupestris propagating wood. Hewitt et al. Rooting ability and graft take are adversely affected Yamakawa: Field spread of fleck in Japan Boulila et al. later called grapevine phloem-limited isometric virus (GPLIV).
Symptoms are severe and affected vines are almost fruitless. Adverse effect on Teleki 5A is negligible. Marsumoto and Ohki: A spherical virus resembling GFkV identified in thin sectioned cells of V. ELISA used successfully for virus detection in large scale survey.: Report of the close association with fleck symptoms in V. rupestris. GFkV-like multivesiculate bodies derived from deranged mitochondria are present in infected cells.: 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.: Natural field spread of GFkV observed in Northern Italy Schieber et al.: Additional monoclonal antibodies raised in France. Molecular properties of this virus further support the notion that it warrants classification in a genus of its own. Detection of sequences of an undentified virus from a Greek grapevine. Sabanadzovic et al. later named Grapevine rupestris vein feathering virus (GRVFV). GAMaV.: Molecular reagents (degenerate primers) developed for the specific identification of viruses of the fleck complex (GFkV. vein necrosis and vein mosaic has a detrimental effect on rootstock growth. Virus renamed Grapevine fleck virus (GFkV).rupestris of an isometric virus latent in V. Walter and Cornuet: Confirmation by ELISA of the consistent association of GFkV with fleck disease. Pruning wood is reduced by 51% in 420A and by 37% in Kober 5BB. Boscia et al.: Two GFkV-specific monoclonal antibodies raised in Italy can successfully be used in ELISA Kuniyuki and Costa: Three strains of GFkV reported from Brazil. This cytological feature recalls that later found in vines infected by Grapevine rupestris vein feathering virus. vinifera. Sabanadzovic et al. Virus named Grapevine asteroid mosaicassociated virus (GAMaV) Boscia et al. Kyriakopoulou: Description of a disease similar to asteroid mosaic observed in V.: Complete nucleotide sequence of GFkV genome. vinifera cv. rupestris with a necrotic disease. The disease seems to be spreading naturally. Boscia et al. June-July are the best months for ELISA detection of the virus in Alsace. Elbeaino et al.1991 1991a 1991b 1991 Gugerli et al. Virus named Grapevine redglobe virus.: GPLIV shown to be the agent of fleck. Fortusini et al. Sultanina in Greece. based on the differential reactions of indicators Credi and Babini: Infection by fleck. Namba et al. Boscia et al. Meristem tip culture effectively eliminates the virus. GRGV). 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 non mechanically transmissible isometric virus similar but unrelated to GFkV identified in asteroid mosaic-infected grapevines. 1991 1993 1994 1995 1995 1996 1996 1997 1997 1998 2000 2001 2001 104 .: Report of a highly consistent association of GPLIV with fleck in naturally infected and graft-inoculated V.: Use of degenerate primers designed on the methyl transferase and polymerase cistrons of members of Tymovirus and Marafivirus genera and of GFkV amplified a genome fragment of GFkV. GAMaV and of another virus with GFkV-like particles phylogenetically but not serologically related to GFkV present in a cv Red globe vine.
R. a new genus of plant viruses having GFkV as type species and GRGV as tentative species. fleck. 160-163. Castellano M. V. Martelli.G. Iran..P. Phytopathologia Mediterranea 24. and G. 1982. GAMaV and of a virus of Greek origin which induces vein feathering in V.A. Plant Pathology 44. D. Di Terlizzi. and G. V. Savino and M.G. Savino. 3134. Production of monoclonal antibodies to grapevine fleck virus. N. Vitis 22. Digiaro. GRVFV recorded from California and confirmation that GAMaV does not occur outside of California.P. Elicio. S. Savino. Abou Ghanem-Sabanadzovic. Virus-like particles and ultrastructural modifications in the phloem of leafroll-affected grapevines.V.A.: Sequencing of the 3' end of the genome of GRGV. Karsev.F.P. Boscia. Engelbrecht D. Castellano.D.P. Identification of the agent of grapevine fleck disease. 97-105. Greek virus recognized as a species in its own right denoted Grapevine rupestris vein feathering virus (GRVFV). Numéro hors série. 151-158. 23-39.E. Argentina.: Development of a multiplex RT-PCR protocol for the simultaneous detection of GFkV-like viruses using plant mRNA as an internal control. Virus Genes 27. and Japan) only the variant without insertion (GFkV353) was detected. 1984. Minafra.R.G. Savino. Savino and G. Molecular detection of Grapevine fleck virus-like viruses. N. Extended Abstracts 14th Meeting of ICVG. M. Bovey R.E.2002a 2002b 2003a Martelli et al. 1996.A. P. Savino. Boscia. Martelli et al. V. Martelli. Vitis 30. and A. 173-174. Martelli and M. Boulila M. S. V. Un virus latent dans le Chasselas. South Africa. Sequencing of the 3' end of three grapevine fleck virus-like viruses . respectively..P. G. Rowhani P...V.P. V. 195. Martelli. 95-98. 191. Boyko.: Description of the family Tymoviridae. Proceedings 8th Congress of the Mediterranean Phytopatological Union. Sabanadzovic .A. Castellano M. Skene.. A. In other countries (USA. Shi et al. 1990. Annales de Phytopathologie.P. G. Babini. Castellano. Sabanadzovic and G. Woodham and L. Barlass M. 2003a. Boscia D.. Kyriakopoulou and G. 1985. Sabanadzovic. 1995. G. 1991a. Martelli and V. S. 165-169. 2003b. Regeneration of virus-free grapevines using in vitro apical culture. Vitis 40: 65-68. Association of a phloem-limited non mechanically transmissible isometric virus with grapevine fleck disease. Sabanadzovic. 1990. References Abou Ghanem-Sabanadzovic. S. rupestris confirms the assignment of GRGV to the genus Maculavirus and of GAMaV and the Greek virus to the genus Marafivirus.P. Volos 1990. U.P.. M. 2003b 2003 3. Annals of Applied Biology 101. Abou Ghanem-Sabanadzovic et al. and G. Martelli 2001. Some properties of a phloem-limited non mechanically-transmissible grapevine virus. leafroll and Shiraz decline diseases and associated viruses in South African grapevines. N.M. Locorotondo 2003. 105 . V. Tomashevskaya. A non mechanically transmissible virus associated with asteroid mosaic of the grapevine. Phytophylactica 22. Krake. 291-295.P. Kyriakopoulou and G.. B.A. Martelli.J. A.. Journal of Phytopathology 129. Dolja V. O.. R.C. Journal of Ultrastructure Research 89. Martelli. A. Rowhani and G. Credi R. K. 11-16 Abou Ghanem-Sabanadzovic.. Boscia D. Proceedings 10th Meeting of ICVG. 1972. Boscia D. Kasdorf.: Description of Maculavirus. 1990. Advances in Horticultural Science 10. 1991b. Atabekov. Verderevskaya and J. 1983.. Double stranded RNA associated with fleck disease of grapevine.. Ultrastructure and nature of vesiculated bodies associated with isometric virus-like particles in diseased grapevines. 347-354.P.: A sequence variant of GFkV (GFkV416) with a 63 nucleotide insertion in the replicase gene identified in Australia and New Zealand. Progress in the study of the phloem-limited isometric virus-like particles associated with leafroll-diseased grapevines. Martelli. Effect of virus and virus-like infections on the growth of grapevine rootstocks. V.P. Vitis 33. 56-64. Castellano. T. Savino and D. Agadir 1990. G. 101-102 Boscia D. ElBeaino T. 1994. A. Field spread of corky bark. Abou Ghanem-Sabanadzovic et al. Multiplex RTPCR detection of Grapervine fleck virus-like viruses in grapevine with co-amplification of control plant mRNA. Martelli. Martelli. Castellano M. comprising the genera Maculavirus and Marafivirus that include GFkV/GRGV and GAMaV/GRVFV.A.
I.IV. (Ed. 567-571. Martelli. Jr. Lisbon 1997. 39-43. Purification and properties of spherical virus particles associated with grapevine Ajinashica disease. Saldarelli and G. Phytopathologia Mediterranea 24. a new genus of plant viruses. Davis 1965. Annals of Applied Biology 142. Ser. Hévin M. A possible new necrotic diseases of grapevine associated with small isometric particles and novel membrane-bound large particles. D. 31-32. S. Informatore Fitopatologico 46 (12). Abou Ghanem. 47-64. Gugerli P. affected by marbour. Heat inactivation of viruses in grapevines. Proceedings International Conference on Virus and Vector on Perennial Hosts. 1973. J. 1998. 43-47. and A. University of California. J. Digiaro and G. vinifera clones and in V.B. Archives of Virology 147. 2009-2015 Savino V. 1991.): Virus Diseases of Small Fruits and Grapevines (A Handbook).P. Division of Agricultural Sciences.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. C. Symons. Complete nucleotide sequence and genome organization of grapevine fleck virus. 1997. Luhn. Investigating the transmission of marbrure and fan-leaf through the seed in the grapevine.E. Numéro hors série. 1966. 1996. In Frazier N.P. J. Volos 1990. Grapevine fleck disease.P. 1954. S. The responses of the Grapevine fleck agent to tetracycline-HCl antibiotic and Dienes' stain. Milkus B. S. Seddas. Gooding.. Archives of Virology 147. M. Monoclonal antibodies for detection. D.. 281-285. S. 1973. Acta Phytopathologica Academiae Scientiarum Hungaricae 9. Proceedings 10th Meeting of ICVG. Annales de Phytopathologie. 1995. Parsons. Abou Ghanem-Sabanadzovic. Boscia. Rosciglione. 9.. Sabanadzovic. M. 1991. Ottenwaelter M. 106 .P.. Ottenwaelter. Walter. Grapevine fleck virus-like viruses in Vitis. is transmitted by graft inoculation.. Resta. Séparation de la marbrure et du court-noué (panachure) chez la vigne par thermothérapie. M. A.C Edwards and T. Costa.. 1985.. Martelli G. Bulletin of the California Department of Agriculture 43. Kuniyuki H. Martelli.B.C. T. M.. Namba S.M. Goheen. P. 9.V. 197-203. Refatti E.. 1997. Castellano. P. Nucleotide sequence variation in a small region of Grapevine fleck virus replicase provide evidence for two sequence variants of the virus. Yamashita. 287-289. Hévin. Plant Disease 75. Some virus and virus-like diseases of grapevines. Gugerli. 1977. Heat therapy eliminates the ability to transmit the causal agent of "marbrure" in several V. Gonsalves. Luhn. Annales de Phytopathologie. Raski and G. 2003. Proceedings 10th Meeting of ICVG. Cory and C. L. Rivista di Patologia Vegetale. Goheen. Matsumoto T.C. European Journal of Plant Pathology 103.. Refatti E. The family Tymoviridae. Shi B. Saldarelli. Symptoms of grapevine asteroid mosaic in Greece. Hewitt W. 560-564.-E. Extended Abstracts 12th Meeting of ICVG. 212-214. Habili and R.. 143-146. Sabanadzovic S. Rives. Asteroid mosaic of grapevine. 1847-1853.A. 5960. Journal of General Virology 82.S. Scattini. Dreher. and E.J. Phytopathologia Mediterranea 24. A. serological characterization and immunopurification of grapevine fleck virus. Annals of the Phytopathologial Society of Japan 64. 253-258. Ramel and F. IV.P. Rives. Brugger.. with Special Reference to Vitis. Fitopatologia Brasileira 20.T. Grapevine asteroid mosaic. and J. N. 57-83. Prati. 157-164.W. N. 1974. Plant Disease Reporter 61. 385-388. and C. Schieber O.C. Fortusini A. Doazan and M. N. 204-207.IV. Doazan and M. 1985. Martelli G.or chlamidia-like bodies in grape. 75-77. 2002a. M.. Studies on virus diseases of the grapevine in California. Rives M. George). 349-355. Hewitt W.Faoro F and P. A. 1837-1846. Leclair. latent in many varieties.B.. Maculavirus. N. Abou Ghanem-Sabanadzovic. Volos 1990. Kyriakopoulou P. 618-622.. Cinquanta and S.H. Procedures for rapid detection of virus and viruslike diseases of grapevine. 553-565. Sabanadzovic. Vitis 3. B. Occorrencia de mais um isolado do virus do mosaico da nervuras da videira que no causa sintomas no porta-enxerto Kober 5BB. rupestris "du Lot" (St. Mycoplasma. 9. Incidence of some graft-transmissible virus-like diseases of grapevine in visually selected and heat-treated stocks from Southern Italy. M. Further characterization of grapevine leafroll disease. Tsuchizaki and D. N.. 1972.M. 1991. Ser. Triolo E. Rivista di Patologia Vegetale. Mink G. Sabanadzovic S.P.P. Cytological alterations associated with an unidentified isometric grapevine virus (UIGV). 2000.. Hewitt W. Tremea.-J. 1970. 2002b. Goheen A. Belin and B.L. Abou Ghanem-Sabanadzovic and P. Berkeley. Rivista di Patologia Vegetale. and S. 1972. Bonnard. Martelli. 2001. G.. Archives of Virology 145. Ohki. D. 12491253. Numéro hors série. Boscia and G.. 767-774. 1973.. Ser. 1962. S.
Vuittenez A. Legin and J.G.D. Verderevskaja T. 1983. probablement indépendante du virus du Court-noué. Walter B. Semtschik. and P..C. Kuszala. 221-226. 193-198. 1966.S. Investigations on transmission of grapevine leafroll. Journal of the Japanese Society of Horticultural Science 58. Woodham R. Annales des Epiphyties 17. Krake.R. Agronomie 13. 1983. 1989. V. yellow speckle and fleck diseases by dodder. Ätiologie und Diagnose der Marmorierung der Weinrebe. 3209-324. 107 . R.. Archiv für Phytopathologie und Pflanzenschutz 19. La Recherche Agtonomique en Suisse 26. 67-73. Observations sur une mosaïque de la Vigne.. Marinesku and E. Virus reinfection of virus-free Cabernet sauvingon and Cabernet franc vines. George. Materazzi. Numéro hors série.. and A. 1993. 651-657. Cornuet. La maculatura infettiva della vite: influenza di isolati diversi sull'attitudine alla propagazione vegetativa di Vitis rupestris St.Triolo E. Phytopathologische Zeitschrift 106. and L. Yamakawa Y. ELISA detection of grapevine fleck virus (GFkV). 1987. 297-302.
with the following mol. with which specific viruses are associated and thought to be their possible causal agents. European diseases GRAPEVINE YELLOW MOTTLE 1. Jankulova: AMV infections recorded from Bulgaria. The spring growth shows more or less extensive yellowing of the leaf blades that does not extend to the veins. has differently shaped particles. Grapevine berry inner necrosis). 1. Description Main synonyms: None. 0. There may be differential susceptibility among cultivars. Plant vigour and yield do not seem appreciably affected.g. the type species of the genus Alfamovirus. Beczner and Lehoczky: AMV infections recorded from Hungary. RNA-3. Transmission: AMV is efficiently transmitted by aphids in a non persistent manner and can cause epidemic outbreaks in many of its natural hosts. is the putative causal agent. 30 to 57 nm in size. Hungary. from quasi isometric to bacilliform. with Mr 24x 103 Da. Detection: AMV is mechanically transmissible to herbaceous hosts and can also be identified by ELISA and moleculat techniques in infected vines. In grapevines. but some are of economic relevance locally (e. suggesting that the virus spreads primarily through infected planting material. Their overall importance is minor if compared with that of the major diseases dealt with in previous chapters. Faint yellow speckling. wts: RNA-1. Historical review 1973 1975 1976 1979 1978 1981 Bercks et al. however. rupestris and the hybrid Grézot 1 x 5C. RNA-2. rings and lines are typical summer responses of infected vines. A.73 x 106 (2593 nt). AMV. infections are scattered and occasional. 18% of the particle weight. Main symptoms: Various patterns of yellow discolouration characterize the disease. Agent: Alfalfa mosaic virus (AMV). Control: Use of healthy material obtained by heat treatment. 109 .: First record of AMV infections and description of symptoms in German grapevines. Bulgaria. Yellow mottle has been reported from Germany. Bovey and Cazelles: AMV particles visualized in thin sectioned grapevine leaves. Chardonnay and Veltliner rouge précoce identified as reliable indicators. a mechanically transmissible virus.MINOR VIRUS DISEASES Several graft-transmissible diseases are known. and a tripartite RNA genome accounting for c. Some of these diseases have been recorded only from Europe. Capsid proteins subunits are of one type. and Turkey.04 x 106 Da (3644 nt). Novak and Lanzova: AMV infections recorded from hop and grapevine in Czechoslovakia. former Czechoslovakia. Switzerland. Virus elimination by treating 37 days at 37-38°C. 2.62 x 106 (2037 nt). Bovey and Brugger: AMV recorded from Switzerland in grapevine and transmitted by grafting to V. others occur in Japan. 0. Varietal susceptibility: Little information available.
Jubileum 75. . 1993. Monografias INIA No. References Akbas B. 1976. Munich 1978. Acta Phytopathologica Academiae Scientiarum Hungaricae 16. Alfalfa mosaic virus on grapevine.. 63-65. This disease is known to occur only in Hungary. Journal of Turkish Phytopathology 22. ed. Control: No information. 1979. sativa DC. Main symptoms: Leaves show bright yellow discolourations that form marginal rings. Detection: GLPV is mechanically transmissible to herbaceous hosts. or the upper part of the blade. Cordoba 1976.. and J.1985 1993 1993 Francki: Comprehensive review of the properties of AMV and other viruses with tripartite genome. Detection and Diagnosis of Graft Transmissible Diseases of Grapevines. quasi spherical 25-30 nm in diameter to bacilliform 40 to 75 nm in length. Lehoczky. 119-128.). Several V. 3rd International Congress of Plant Pathology. Martelli: Yellow mottle suggested as the name for the disease caused by AMV in grapevines. Graft transmission to cv. Jankulova M. 1-18. 1981. Arboriculture. 263 pp. D. Francki. Vigour and yield are reduced.I. In: The Plant Viruses. and J. Le virus de la mosaïque de la luzerne sur la vigne. is seed-transmitted and spreads with diseased propagative materials. has differently shaped particles. New York .P. Novak J. GRAPEVINE LINE PATTERN 1. 152-154. Rome. The viruses and their taxonomy. and G. Über den Nachweis des Alfalfa mosaic virus in einer Weinrebe. Transmission: GLPV has no known vector. 55-63. 110 . 3.Hegi) plants in Czechoslovakia. 1978. FAO Publication Division. Horticulture 7. vinifera cultivars are susceptible. 1985. Martelli G. Varietal susceptibility: No information. Grapevine line pattern virus (GLPV) a possible member of the genus Ilarvirus. and O. Beczner L. 166-171. Erdiller. 1975. or maple leaf-like line patterns typically confined to the petiolar area. 39. Biologia Plantarum 18. Description Main synonyms: None.B. 1973. Polyhedral Virions with Tripartite Genomes (R. Investigation on certain viruses spread on grapevines in Bulgaria. Proceedings 6th Meeting of ICVG. 2. Bovey R. Phytopathologische Zeitschrift 76. Plenum Press.B. Brugger.B. Bovey R. 131-134.18 . Lesemann and G. Agent: The putative agent. Revue Suisse de Viticulture. 1993. Researches on grapevine virus diseases and determination of their incidence in Ankara Turkiye. (ed. Francki R. roughly following its contour. and J.I. Grapevine disease in Hungary caused by alfalfa mosaic virus infection. Querfurth. Cazelles.).) and grapevine (Vitis vinifera subsp. Bercks R. scattered spots or blotches.-J. Identification of alfalfa mosaic virus and tomato bushy stunt virus in hop (Humulus lupulus L. and a multipartite genome. Akbas and Erdiller: AMV infections recorded from Turkey. Historical rewiev 1985 Francki: Comprehensive review of the properties of AMV an other viruses with tripartite genome. Lanzova.
ed. J. Beczner and G. Lehoczky et al.P. 115-116. 1-18. 2.A. L. Martelli and J. 1985. CarMV is an isometric virus 30 nm in diameter. one of the putative agents of corky bark (rugose wood complex) has a very high association with diseased grapevines. Lehoczky et al. Martelli. Historical review 1989 1991 Rumbos and Avgelis: Roditis leaf discoloration described in Greece. G. Evidence of its grafttransmissibility. Kiryat Anavim. Francki. However.: Description of line pattern disease in Hungary. References Francki R. D. Control: No information.. The disease is graft-transmissible and spreads through infected propagating material. Mission. Transmission: No vector is known.I. 1987. Description Main synonyms: None. Burgyan. Bunches are reduced in numbers. Farkas. Castellano.1987 1989 1992 Lehoczky et al. 3. a novel virus disease of grapevine in Hungary. The disease has been recorded only from Greece. Polyhedral Virions with Tripartite Genomes (R. Israel. New York .). Detection: Graft-transmission to V. wt 1. Boscia. 1992. M. family Tombusviridae. according to more recent findings. Castellano.B. Farkas.B. 1989. Seed transmission of grapevine line pattern virus. By converse. Grapevine virus B (GVB). Agent: Symptomatic grapevines were reported to be doubly infected by GFLV and Carnation mottle virus (CarMV) the type species of the genus Carmovirus.4 x 106 (4003 nt in size) and coat protein subunits of Mr 38 x 103 Da. Line pattern. M. 23-30. Plenum Press. In: The Plant Viruses. Occurrence of the line pattern hitherto unknown virus disease of grapevine in Hungary. 1987. J. Varietal susceptibility: No information. L. with Mol. vinifera cv. size and have low sugar content. or variously extended sectors of the leaf blade. RODITIS LEAF DISCOLORATION 1..: Purification and characterization of GLPV and suggestion that it is the causal agent of the disease. Main symptoms: Symptoms are prominent in late summer and consist of yellow and/or reddish discolourations of the tissues along the veins. 111 . Phytopathologia Mediterranea 31.. has a monopartite RNA genome accounting for c. Boscia.: Evidence that GLPV is transmitted through grapevine seeds. Viruses associated with the disease are readily transmitted by sap inoculation and can be detected by ELISA and molecular techniques. Avgelis and Rumbos: Double infection of diseased vines by GFLV and CarMV reported. Evidence that a graft-and mechanically transmissible virus is associated with it. 18% of the particle weight.A.. D. GFLV may not be involved in the aetiology of the disease. Lehozcky J. Kertgazdasag 19. 61-79 Lehoczky J.P. especially near the petiole Leaves are deformed in correspondence of discoloured sectors. Lazar. The viruses and their taxonomy.I. G. Proceedings 9th Meeting of ICVG. Beczner and G. Burgyan. Lehoczky J. the interveinal areas.
Rumbos I. F. P. and ELISA. Whether this mechanism operates with grapevines has not been ascertained. C.I. the closest being Tobacco streak virus (TSV). and I. Bem.P. 2000. but differs from GLPV. thus is regarded as the agent of the disease. Transmission: GAMV is pollen-borne in herbaceous hosts and was able to infect pollinated plants. Control: No information 2. References Avgelis A. Proceedings 10th Meeting of ICVG. Avgelis. reproduced the field syndrome in mechanically inoculated grapevine seedlings.C. Infected grafting material is likely to be responsible for virus dissemination. Journal of Phytopathology 125. crinkling and deformation of the leaves. References Girgis S.: Evidence that GAMV is the agent of grapevine angular mosaic disease. Flowers abortion results in straggly bunches with small wrinkled berries bearing non viable seeds. 2003. The etiology of a new virus disease: grapevine angular mosaic. Baresana x Baresana. A. Girgis et al. A new ilarvirus isolated from grapevine in Greece. Historical review 2000 2003 Girgis et al.E. P. Laski Rizling from Slovenia exhibit a yellow line pattern syndrome resembling the grapevine line pattern disease described from Hungary. 3. C.: First record of GAMV. the only other ilarvirus reported from grapevine. Dovas. Varietal susceptibility: No information Detection: Indexing on cv. A.D. P. Sklavounos. Dovas.. Roditis leaf discoloration -.a new virus disease of grapevine: symptomatology and transmission to indicator plants. 19. decline gradually and some die.3. GAMV is molecularly related to a number of ilarviruses.. GRAPEVINE ANGULAR MOSAIC 1. S. YELLOW LINE PATTERN (Rasperry bushy dwarf virus) 1. Carnation mottle virus isolated from vines affected with "Roditis leaf discoloration". Extended Abstracts 14th Meeting of ICVG Locorotondo 2003. Avgelis. Girgis S.I. Katis. 1991. Plant Disease 84. Infected grapevines are stunted. Agent: Grapevine angular mosaic virus (GAMV). F.E. N. 112 . A. 437-443. Katis. Kyriakopoulou. Tsagris. and A. Bem. Description Main synonyms: None Main symptoms: Symptoms are chlorotic angular spots on the leaf blades.C. mechanical transmision to herbaceous hosts. Avgelis and N. 274-278. discoloration of tissues bordering the veins. Kyriakopoulou. Tzortzakakis and M. P. Rumbos.M. The disease has been recorded only from Greece. 1345. 1989. Volos 1990.D.M. Description Main synonyms: None Main symptoms: Grapevines of cv. a virus with a tripartite RNA genome and a 30 kDa coat protein.
Koshu. The vay of natural spreading in grapevine. a dimeter of about 33 nm. the 3' terminal region of which (2469 nts) has been sequenced. However. 20 Murant A. Almost all Japanese table grape cultivars derived from crosses with cv. a mechanically transmissible definitive member of the genus Trichovirus. representing the most important virus disease in Yamanashi Prefecture. The virus spreads naturally in the vineyards. and RT-PCR. Raspberry bushy dwarf virus.5 x 106 Da. CMI/AAB Description of Plant Viruses. M. Description Main synonyms: None Main symptoms: Infected grapevines have low vigor. and Pione whereas cvs. ELISA . Raspberry bushy dwarf virus infection of grapevine in Slovenia. being transmitted by the eryophid mite Colomerus vitis. 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. No. Campbell Early are suscetible as well as cvs Takao.2 kb in size). rings and line patterns are shown by leaf blades. wt of 2 x 106 Da (5. Grapevine berry inner necrosis has been reported only from Japan.5 Kb in size) and RNA-2 with Mol. Transmission: In raspberry the virus infects progeny seedlings and pollinated plants through pollen. Delaware. 113 . delayed bud break and young shoots with short internodes and internal browning. Transmission: GINV is transmitted by grafting to grapevines and by mechanical inoculation to herbaceous hosts. 1976. Ripening of bunches is delayed. Zezlina.8 x 106 Da (2.. Locorotondo 2003. berries are small and show external discolorations and internal necrosis. Virscek Marn and I. Varietal susceptibility: No information Detection: Mechanical transmission to herbaceous hosts. Historical review 1976 2003 Murant: Description of RBDV. Kyoho. References Mavric I. Extended Abstracts 14th Meeting of IGVG. Japanese diseases GRAPEVINE BERRY INNER NECROSIS 1. Control : No information 2. 165 B. Healthy vines of cvs Kyoho and Pione became naturally infected in the field within one year from planting. 3. and a bipartite singlestranded RNA genome accounting for c.Agent: Raspberry bushy dwarf virus (RBDV) was isolated from symptomatic vines.g. F. 30 x 103). Chlorotic mottling. Mavric et al. is unknow. wt 0. wt of 7. infected propagative material can disseminate the RBDV. Vitis riparia Gloire) are also susceptible. Agent: The disease agent is Grapevine berry inner necrosis virus (GINV). 24% of the particle weight and consisting of two functional species RNA-1 with Mol. 2003. Varietal susceptibility: Symptom severity varies with the cultivar.: First record of RBDV in grapevine.. if any. GINV has filamentous particles about 750 nm in length and a single-stranded RNA genome with Mol. Some rootstocks (e. RBDV. and Kaiji are infected latently.
47-56. 362.. Transmission of grapevine berry inner necrosis virus by the grape erineum mite Colomerus vitis in Yamanashi. Induction of berry necrosis in Kyoho back-inoculated with the virus isolate from grapevine mosaic diseased clones and renaming to grapevine berry inner necrosis. Historical review 1984 1985 1987 1992 Tanaka: Description of a mosaic disease in cv. Annals of the Phytopathological Society of Japan 55. Tanaka H. Studies on the grapevine berry innner necrosis virus disease. Kunugi. GINV is mechanically transmissible to herbaceous hosts and can be identified by ELISA and moleculat techniques in infected vines. Extended Abstracts 11th Meeting of ICVG. 77-78 Yanase H. 1985. Yanase H. 1992. Terai and A. 2000. 1987. Description Main synonyms: None. H. Yoshikawa N. Y. 1351-1363 GRAPEVINE STUNT 1. Terai. Kyoho back inoculated with the filamentous virus isolated from mosaic-diseased grapevines. Goto. Annals of the Phytopathological Society of Japan 51. 2. Grapevine berry inner necrosis. Magome. Kunigi Y. Bulletin of Yamanashi Fruit Tree Experimental Station 10.Annals of the Phytopathological Society of Japan 53. 536 Terai Y.: An account of the varietal susceptibility to the disease and natural field spread Kunigi et al. Disease re-named Grapevine berry inner necrosis Terai et al.. Y.. Nishijima T. Back-transmission of a grapevine filamentous virus to grapevine seedlings and induction of foliar and berry symptoms in grapevine. A new virus disease. Montreux 1993. H. 617.Annals of the Phytopathological Society of Japan 58. 57-63. Shinkai 2000. 2. Archives of Virology 142.. Mosaic symptoms on cv Kyoho. Takahashi and Y. and H. References Kunugi Y. and Terai Y. Asari. Terai. Iida. 1.Detection: Indexing on cvs Kyoho or Pione. 1993.: Partial sequencing of GINV genome and assignement of the virus in the genus Trichovirus Nishijima et al.: First account of grapevine berry inner necrosis disease in a non Japanese publication.. 1997. a new trichovirus: comparative studies with several known trichoviruses. and Yanase H. S. 1984. Symptoms on vines. S.: Experimental evidence that GINV is transmitted by the the grape erineum mite Colomerus vitis 1993 1997 2000 2000 3. Studies on the grapevine berry innner necrosis virus disease. Purification of a filamentous virus isolated from grapevine berry inner necrosis and foliar mosaic. Kyoho in Japan Yanase: Purification of a filamentous virus isolated from mosaic-diseased grapevines Yanase and Terai: Induction of mosaic symptoms in grapevines inoculated with the filamentous virus Terai and Yanase: Induction of berry internal necrosis in cv. Yanase. Yoshikawa et al. varietal susceptibility and natural spread. Terai and Y. Bulletin of Yamanashi Fruit Tree Experimental Station 10.. grapevine berry inner necrosis with natural spread in Japan. Control: Use of tolerant cultivars in areas where the disease spreads epidemically. 114 . Y. T.. 423..
Y. Hatamoto. Fujii. Y. Iwanami. Yamashita and Y.. Namba. Doi. 1-17. Yamashita. internodes are short. Hatamoto et al. GRAPEVINE AJINASHIKA DISEASE 1. 33. No. Yamashita and Y. 115 . 1982. sometimes. 396 Hatamoto M. 1987). Transmission of grapevine stunt disease by the grapevine leafhopper Arboridia apicalis Nawa. fruit setting is impaired and bunches are few and shelled. 1986. S. The disease is apparently restricted to the V. FFTC Book series. Taiwan (Reference 2951 in the Review of Plant Pathology 66. In Plant Virus Diseases of Horticultural Crops in the Tropics and Subtropics. The disease has only been reported from Japan. S. 137 Namba S. Fujii. Annals of the Phytopathological Society of Japan 47. Taipei. phloem-limited. Control: Use of disease-free material obtained through heat therapy. T. The same paper appears in Taiwan Food and Fertilizer Technology Center Technical Bulletin 92. Food and Fertilizer Technology Center for the Asian and Pacific Region. Annals of the Phytopathological Society of Japan 50.Main symptoms: Spring vegetation is delayed. Main symptoms: No appreciable symptoms are visible on the foliage of cv. 85 Namba S. Graft transmissibility of grapevine stunt disease. A small spherical virus associated with grapevine stunt disease.: Purification and characterization of the virus associated with stunt disease. S. Three phloem-limited viruses of grapevine: direct fluorescence detection. Varietal susceptibility: No information. 1981 1982 1984 Namba et al. leaves are small. are pale-coloured and have a low sugar content. Because of heat recovery. Detection: Grafting to Campbell Early and ELISA using extracts from infected vine tissues. Hatamoto et al.109-126.. Doi and M. S. The disease has only been reported from Japan.: Successful graft-transmission of stunt disease. 1984. Historical review. 1981.: Evidence that the disease is transmitted by the leafhopper Arboridia apicalis. Description Main synonyms: none. with scorched margins. which makes the crop unmarketable. Campbell Early. This condition gives the name to the disease which in Japanese means "unpalatable fruits with low sugar content". Evidence that it is not related to the presumed agent of ajinashika disease. vinifera cv. curled and. S. Inflorescences are undersized.: A small isometric virus associated with stunt disease in Japan.. 3. summer vegetation is apparently normal. Doi and K. Annals of the Phytopathological Society of Japan 48. Agent: An isometric. American rootstocks are infected without showing symptoms. Spread occurs also through infected propagative material. Transmission: The disease is transmitted by the leafhopper Arboridia apicalis. Koshu nor any apparent reduction of vigour and yield. S. M. This virus is serologically distinct from the putative agent of ajinashika disease. Yora. non mechanically transmissible virus about 25 nm in diameter is consistently associated with diseased vines and regarded as the possible causal agent. M. Yamashita. however. References Hatamoto M. 316. Doi. p. 1986 Namba et al. Namba.. The berries. 2.
1979 1980 1986 1991 1991 Namba et al. Ajinashika disease: a combined effect of grapevine leafroll and grapevine fleck viruses on sugar content in the Japanese grape cultivar Koshu. Namba et al. Transmission: No vector is known. Terai: Additional report on ajinashika disease as derived from the combined effect of leafroll and fleck. Boscia. Yora. Doi and K. Niagara Falls 1980.: First mention of ajinashika disease and report of the association with it of a non mechanically transmissible virus with isometric particles. Y. 1-17. Three phloem-limited viruses of grapevine: direct fluorescence detection. was suggested as the possible causal agent. 1980. Hatamoto. Y. 70-73. Proceedings 10th Meeting of ICVG. Ajinashika disease of the grapevine cultivar Koshu in Japan. Namba S. The disease seems to be restricted to V. Iwanami. Terai and Yano: Description of ajinashika disease and suggestion that it is caused by the concomitant infection of leafroll and fleck. Yamashita. 116 . consistently found in infected vines. 1986.. T. phloem-limited.Agent: The disease was reported to be caused by the concurrent infection of leafroll and fleck. Historical review.. T. 1979. and R. Yamashita. 1991. Varietal susceptibility: No information. 67-70. Koshu and ELISA using extracts from infected vine tissues. an isometric. Control: Use of disease-free material obtained through heat therapy. 15-19. References Namba S. non mechanically transmissible virus about 25 nm in diameter. 12491253. Terai Y. D. Gonsalves. Dissemination is through infected propagative material. No relationship found with fleck.. and D. Tsuchizaki. Volos 1990. Terai Y. S. Proceedings 7th Meeting of ICVG. 3.: Partial characterization of the isometric virus associated with the disease and its detection by ELISA in infected vines. vinifera cv. Annals of the Phytopathological Society of Japan 45. Purification and properties of spherical virus particles associated with grapevine ajinashika disease. S.: Further characterization of the isometric virus and claim that it is the putative agent of the disease. Detection: Graft transmission to cv. Plant Disease 75. Namba et al. A small spherical virus associated with the ajinashika disease of Koshu grapevine. 2. Namba S. 1991. Taiwan Food and Fertilizer Technology Center Technical Bulletin 92. Doi and M. Yano. Yamashita. S. However. Koshu.
VIRUS-LIKE DISEASES .
Historical review 1891 1937 1954 Buchenau: First description of enation disease of grapevine occurring in Germany. and often show longitudinal cracks between the nodes. Primus. Symptom expression varies year by year. Detection: By observing the symptoms on the leaves and by indexing on LN 33 hybrid. The disease has been reported from many European and extra-European countries Agent: The etiology of enation disease is not yet clear. The disease is perpetuated by vegetative propagation. Enations develop mostly on the underside of the leaves at the base of the shoots. apparently in relation with climatic conditions. The crop can be drastically reduced (up to about 50%. However. some of which have a clear-cut detrimental effect on the crop. Graft transmission suggests that it is a virus disease.). Varietal susceptibility and sensitivity: There is little information available. Latin America (Venezuela). vinifera varieties in Europe. which gives a bushy aspect to the plant. 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. Severely affected leaves drop prematurely. 2.VIRUS-LIKE DISEASES Several latent or semi-latent grapevine diseases are known.). Leaves developed later in the season are usually normal. Symptoms have been observed on many V. malattia delle enazioni. Australia and New Zealand. North (Tunisia) and South Africa. has now been dismissed Transmission: By vegetative propagation. The transmission by graft is rather erratic. Grenache and Tokay appear to be quite susceptible and develop severe symptoms when infected. Description Enation disease of grapevine is one of the oldest known disorders of European grapes. ENATION DISEASE 1. are often mis-shapen. whether they bear enations or not. with a fanlike aspect and abnormal indentation. as symptom expression is variable in successive years and graft transmission not 100 % successful. maladie des énations (Fr. 119 . They are often thicker than normal. Riesling. the absence of symptoms does not necessarily mean that the plant is healthy. Hewitt: Description of symptoms in California. according to the cultivar) and is of poor quality. They are outgrowths 2-3 mm high and 3-5 mm long or more. Basal leaves. irregular and mis-shapen. All persist in propagative material and are transmitted by grafting. with drawings of symptoms. but attempts to transmit it by graft or mechanical inoculation gave no results. Their agents are still unknown. The varieties Panse Precoce. growth tends to become normal again. Later in the year. with prominent veins. Gigante : Research on histological and cytological aspects of enations. Control: Use of healthy material. however. Italia. Main symptoms: Affected vines show a delayed opening of the buds and a slow growth of the shoots in the spring. omeoplasie crestiformi (Ital. The infectious agent of the disease perennates in propagating material. North America (California). There is no information on the possibility of curing this disease by heat treatment or meristem culture. its description dating back to the late 1800s Main synonyms: Enationenkrankheit der Rebe (Germ. which run more or less parallel to the main veins. This hypothesis. but some are heat-labile and can be eliminated by heat therapy.). The basal internodes are short. and is probably due to a virus-like agent.
attempts to transmit the disease by graft. Malvasia (10. only fanleaf virus was recovered. the hybrid LN 33 was found to be the most sensitive and reliable indicator variety. the proportion of diseased vines was highest in cv. The mean yield loss of diseased vines ranged from 17.: More data on the effects of enation on the yield of cv. . Beobachtungen und Untersuchungen über die Enationenkrankheit der Rebe. 120 . McGechan: Enation disease in Australia Tekinel et al. 1978. but report on observations made in Australia where no fanleaf virus could be recovered from enation-affected vines. Confirmation of graft transmissibility of the disease. In the vineyards under observation. Prota et al. Nieder: Enation disease in Austria Garau et al. Italia in Sardinia. Brückbauer: Description of symptoms of enation in Germany and confirmation of graft transmission of its agent. lowest in cv. and C. but diseased vines which had not shown enation symptoms for several years had almost normal yields. Enationaffected vines produced less than 50 % of the yield of healthy plants. Trebbiano romagnolo and reduces the yield from 13% to 23% according to the severità of symptom expression.: Enation disease in Czechoslovakia Avgelis and Xafis: Enation disease in Greece Prota and Garau: Enation disease found in Sardinia.5%).: Successful transmission of enation disease from diseased to healthy grapevine by graft strongly supports the hypothesis of a viral origin. The authors discuss the hypothesis that enation is caused by a strain of fanleaf virus. with negative results. Xafis. and possibly caused by a virus. Presence of enations in Razaki grapevine in Crete (Greece). Padilla et al. 195 Brückbauer H. Marinesku and Bondarchuk: Enation disease in Moldova Brückbauer: Influence of enation disease on growth and yield of grapevine in West Germany. although the rate of symptom expression does not exceed 30% Credi: Enation diseasae affects the vegetative vigour of cv. Graniti and Martelli: Review paper on enation. The authors conclude that the disease is probably of European origin. historical account. Phytopathologia Mediterranea 17.: Detailed description of macroscopic and microscopic symptoms of enation disease..: Enation disease in France Pozdena et al. Vernaccina (1. Weinberg und Keller 15.: In experiments on graft transmission of enation disease aimed at determining the best indicator.5%).: Enation disease in Turkey Hevin et al.4 to 48. 79-112. The possible role of fanleaf virus in the etiology of enation requires further investigation. 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. Mechanical transmission tests were also negative.1966 Graniti et al. Martelli et al. Refatti: Hypothesis of a correlation between fanleaf and enation disease.3% . 1968. References Avgelis A. There is some evidence that enation can be carried in the rootstocks.
9. 1976. Lisbon 1997. Berkeley. 207-217. Leclair and M. The research of the virus diseases of grapevine in CSSR. Ser. VEIN MOSAIC 1. Spain. Graniti A. A study of infectious degeneration (fanleaf) in vineyards of the Mediterranean region. Lamberti and A. 823-827. Graniti A.G. producing often a vein banding effect. Quacquarelli. California. Proceedings International Conference on Virus and Vector on Perennial Hosts. I. 326332..). Cordoba. 1971.. Gazeau. 48. Occurence of grapevine enations in the Moldavian SSR.). 1970.P. Enations of grapevine in Sardinia.P. Dolar. Hevin M. B. but when fanleaf virus transmission to herbaceous hosts became possible. Bericht der deutschen botanischen Gesellschaft 9. 1966. Graniti. F.. 1954. Savino. 225-246. 47. In the most sensitive varieties. with Special Reference to Vitis. This disease is widespread and probably worldwide. 179-189.. 17.). Main symptoms: Pale green mosaic affecting mostly the tissues adjacent to the main veins or the smaller ones. Enation symptoms found in France. 2.P. 1975. Die Eantionenkrankheit der rebe erstmal auch in Osterreich nachgewiesen. Deutsches Weinbau-Jahrbuch 31. small fruit and grapevine in Moldavia. 1970. 121 . 47-64. Vein mosaic appears to be of low economic importance. Division of Agricultural Sciences. Martelli G. 1997.D.1989. Refatti E. Bitki Koruma Buletin 11. 203-206.. Nas. Important virus diseases of grapevine in New South Wales. Studies on some variable characters of grapevines affected by enation disease in Sardinia. Description The symptoms of vein mosaic have been confused for some time with those of fanleaf/yellow mosaic. Israel.). Grapevine enation disease in Murcia (Spain).. Salih and Y. it was clear that vein mosaic was not caused by fanleaf virus. Rives.B.V. Bondarchuk. Phytopathologia Mediterranea 20. Extended Abstracts 12th Meeting of ICVG.. Cugusi. 1965. University of California. Bulletin of the California Department of Agriculture 43.. 1997. Garau R.. Tekinel N. Proceedings 6th Meeting of ICVG. Symptom expression seems to depend on climatic conditions. 1966. In: Virus Diseases of Small Fruits and Grapevines (A Handbook). Su una possibile correlazione fra il virus del complesso dell'arricciamento e la malattia delle enazioni della vite. Facultet Landbouwwetenschap (Gent) 40. Roma.) Virus and mycoplasma-like diseases of fruit trees. Chabbouh N and V. Salcan. Bolletino della Regia Stazione di Patologia Vegetale. 1983. Some virus and virus-like diseases of grapevines. Lamberti. 1979. Petria 6.. Garau. Phytopathologia Mediterranea 5. Rivista di Patologia Vegetale. Gigante R. 1973. Garau R. 145-152.. U. McGechan J. 1981. 1937. Prota U. Buchenau F. Main synonyms: Mosaïque des nervures (Fr.W. Ricerche istologiche sulle omeoplasie crestiformi (Enations) delle foglie di vite affette da rachitismo. 1966. Ita and F. A similar disease has been reported in Australia under the name of summer mottle.K.Brückbauer H. Prota U. 293-306. Proceedings 9th Meeting of ICVG. Padilla V. 1987.. 59-64. Pflanzenharzt 36. and R. 97-98. Martelli. And G. Rivista di Patologia Vegetale S.W. 241-243. Frazier N. and G. Vanek. Occurrence of enation disease in Tunisia. In: Verderevskaya T. 46-51. and M. G. Marinesku V. H. Adernmosaik (Germ. Enation disease of grapevine in Italy. Credi R. M. Mosaico delle nervature (Ital. 122-124. 7-12. Prota and M. Extended Abstracts 12th Meeting of ICVG. 251-252 Hewitt W.IV. 1983. G. Martelli and F. Mededel. Pozdena J. 1980.. n. O.. and V. Benayas. 1996. areas of the leaf blade may become necrotic. Einfluss von Virusinfektionen auf Wachstum und Ertrag der Rebe. but these necroses do not affect the veins as it is the case with vein necrosis. Z. (ed. (N.P.s. Kiryat Anavim. Davis. Agricultural Gazette of New South Wales 81. Lisbon 1997.S. Studies on reproduction of enation symptoms by grafting in Sardinia. Trasmissione per innesto della "malattia delle enazioni" della vite. 169-192. Garcia. Cugusi. Monografias INIA 18. A. 1891. Nieder G. IV. Abnorme Blattbildungen. Effetto della malattia delle enazioni della vite sulla produzione e sullo sviluppo vegetativo della cv Trebbiano romagnolo. 349-352. Moldavian Research Institute of Horticulture Kishinev. Enations.
Babini and A. Pop: Vein mosaic in Romania. Alphonse Lavallée. Ehrenfelser showing symptoms of vein mosaic. Vitis 17. riparia Gloire de Montpellier. Marinesku and Bondarchuk: Vein mosaic in Moldova. 266-270. 1979 1980 1982 1983 1985 1993 2004 3. but this hypothesis has not been confirmed. Krake and Woodham: Description in Australia of a systemic mottling syndrome which is expressed during summer on the leaves of some varieties.Agent: Unknown. The disease can be eliminated by heat therapy. Krake L. 17-23.. Varietal susceptibility and sensitivity: Vitis riparia Gloire de Montpellier is especially sensitive and is used as indicator. and Gamay apparently show little or no symptoms. Chasselas.A. Control: Use of indexed material. References Credi R. 1993. Canova. Bonfiglioli: Vein mosaic in New Zealand (unpublished). and R.: Vein mosaic in URSS.: Observation of a type of mosaic of grapevines which appears to be independent of fanleaf virus. Muscat de Hambourg. 122 . Mycoplasma-like organisms were supposed to be the cause of vein mosaic. Chardonnay. Potential interaction between rootstocks and grapevine latent viruses. Phytopathologia Mediterranea 24. Kuniyuki: Vein mosaic in Brazil. American Journal of Enology and Viticulture 44. vein mosaic and vein necrosis. Servant. Woodham. Occurrence of grapevine vein necrosis and grapevine vein mosaic in the Emilia-Romagna region (northern Italy). Golino D. Comparison of symptoms of fleck. 148-152. 2. vinifera cvs. Samonina et al.R. No claim is made that these putative viruses are connected with the disease. Transmission: By graft and vegetative propagation. Woodham and Krake: Comparison of summer mottle and vein mosaic. Pearl of Csaba). Grapevine summer mottle: a new graft-transmissible disease. in the absence of any detectable virus. Abracheva: Vein mosaic in Bulgaria. Vuittenez and Stocky: Electron microscope study of thin-sectioned tissues of leaves from Vitis riparia and Vitis vinifera cv. Symptoms are very similar to those of vein mosaic in Europe. show symptoms (Syrah.R. A. Milkus et al.. Pinot.: Vein mosaic in Ukraine. LN 33 is also sensitive. 1985. Viognier. Detection: Indexing with V. No vector known. Saric and Hranuelli: Vein mosaic in Croatia. Legin and Vuittenez: Description of vein mosaic. A number of cytological modifications primarily involving chloroplasts were observed along with the presence of bundles of filamentous structures resembling closterovirus particles. 1978. Golino: Vein mosaic in California. Several V.C. Historical review 1966 1973 1973 1976 1973 1973 1978 Vuittenez et al.
67-73. Krake.V. 1982. Rivista di Patologia Vegetale S IV 9. 1973.G. 1985. Investiation on grapevine viruses in the SR Croatia. de la marbrure de V. 1983. URSS.. IV . Cabernet sauvignon.N.. Vuittenez A. Symptoms show on vegetative growth that develops at temperatures in excess of 30 °C. La mosaique des nervures. Proceedings of the Conference on Excoriosis and Virus Diseases of Grapevine. 41-42. and V.N. Krylova. Niagara Falls 1980. Main symptoms: Pale green to yellowish dicolourations of the tissues adjacent to the main or secondary veins. Rivista di Patologia Vegetale. 9 . poorly developed and with small berries. and Hranuelli T. Vinogradr. Summa Phytopathologica 11. Agent: Unknown. Saric A. Detection: Graft transmission to a number of cvs. Numéro hors série. 191-204. Ser. A. 110 R.R. 1973. 2. Cabernet franc.V. maladie à virus de la vigne. Woodham R. rupestris and LN33 are infected symptomlessly.V. Comparaison des symptômes et transmission par greffage d'une mosaïque nervaire de Vitis vinifera . 68-72. Evidence that the disease is graft-transmissible. Vuittenez A. 243-250 Samonina I. Woodham and Krake: Comparative graft transmission trials demonstrate that summer mottle 123 . ou la “feuille rouge”. and G. 149-141. 1973. Proceedings 7th Meeting of IGCV.-Berl. e. 48-49 Marinesku V.: Elimination of the disease agent by culturing fragmented shoot apices. whereas the opposite occurs with summer mottle. Stocky. Vinodel. Kuszala. Bondarchuk. Moldavii 31. and L. These symptoms appear in summer and persist through the autumn. Historical review 1978 1982 1983 Krake and Woodham: Description of summer mottle in Australia. Mostar 1977. R. Milkus. V. Bunches of infected cvs Sideritis and Cabernet sauvignon are fewer. Pop I.Legin R. resembles in some respects the European vein mosaic and the Greek Roditis leaf discolouration (see under Minor virus diseases).V.. 9. Transmission: No vector is known. several European grape cultivars show symptoms. Vitis 22. Control: Use of disease-free propagating material obtained by culture of fragmented shoot apices. Vuittenez. Observations sur une Mosaïque de la Vigne. Grapevine vein mosaic.C. Barlass et al. suspected to be a virus or a viroid. Legin and J. 1976. Adverse effect of light and of high temperature on symptom expression of grapevine vein mosaic in Sao Paulo. producing a feathering or banding effect. A grapevine virus disease in the Primorye territory. IV. rupestris et d'une affection nécrotique des nervures de l'hybride Rup. Roditis leaf discolouration and summer mottle have similarities suggesting that they may be the same disease... 1977. Description Main synonyms: None. 57-63.g. 1. Rivista di Patologia Vegetale . B. Krylov and V. 1966. Ultrstructure de vignes infecteés par deux maladies de type viral: la mosaique des nervures. A comparison of grapevine summer mottle and vein mosaic diseases. Mission. 247-252. Mataro. However.. Annales des Epiphyties 17. SUMMER MOTTLE Summer mottle. S. Spread is through infected propagative material but is has also been observed between adjacent vines Varietal susceptibility: No grapevine tested was immune to infection. Symptoms of vein mosaic develop under mild weather conditions and fade during hot weather. probablement indépendante du virus du Court-noué. Kuniyuki H. an Australian disease. and A.
Grapevine summer mottle: a new graft-transmissible disease. rupestris x V. later on younger leaves as they develop. especially under greenhouse conditions. Detection: By grafting on 110 R. Woodham R. No vector known. VEIN NECROSIS Although vein necrosis is currently regarded as a disease in its own right.R. Krake L. Necrotic reactions are best seen on the lower face of the leaf blade. most likely GRSPaV. K. Graft-transmissible Diseases of Grapevines.). recently an association exceeding 95% has been experimentally observed between Grapevine rupestris stem pitting-associated virus (GRSPaV) and vines of V.R. Also the tendrils and many shoots can necrotize. Woodham and L. 70-74. Description Vein necrosis has probably a worldwide distribution. Transmission: By grafting and vegetative propagation. Agent: Suspected to be a virus. Krake. but their etiological relationship with the disease has not been proven. Krake L. So far. Krake. at first on the leaves at the base of the shoots. and the only Vitis species that is clearly affected is the rootstock 110 R. growth is much reduced and necrosis of the leaf veins appears. 247-252. Taylor. and some infected plants may die. 1978. Vitis 17.C. References Barlass M. 2. varieties or hybrids. 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.C. Australia. Possible viroidal etiology put forward. Main synonyms: Nécrose des nervures (Fr. A comparison of grapevine summer mottle and vein mosaic diseases. This strongly supports the hypothesis that vein necrosis is a specificic reaction of the rootstock 110 R to GRSPaV infection. 291-295. and L. Annals of Applied Biology 101. Little is known about sensitivity of other Vitis species. Skene.H.R. 1999. Many grapevine varieties and rootstocks are infected symptomlessly.. Scott. Historical review 1973 1978 1978 Legin and Vuittenez: Discovery and description of this virus-like disease while searching for indicators for fleck. and R. CSIRO Publishing. its economic importance has not been assessed.S... M. Mycoplasma-like organisms have been observed in the phloem of symptomatic vines. 1999 Krake et al.). Varietal susceptibility and sensitivity: The rootstock 110 R is most sensitive. Regeneration of virus-free grapevines using in vitro apical culture. Adernnekrose (Germ. 1983.: Vein necrosis in Italy and Bulgaria 124 . R.: Suggestion that summer mottle and Roditis leaf discolouration are the same disease 3. 266-270.A.C. necrosi delle nervature (Ital. 1. Vitis 22.differs from vein mosaic. Martelli et al.). Main symptoms: On the rootstock 110 R.M. Control: Use of indexed planting material. N. Rezaian and R. Woodham. Collingwood. berlandieri 110 Richter (110R) with vein necrosis symptoms.R. The agent of vein necrosis can be eliminated by heat therapy. 1982. Cause-effect relationships between MLOs and the disease has never been ascertained. Milkus and Kalashyan: Mycoplasma-like organisms found in phloem tissues of vines with vein necrosis.G.
Savino. rupestris et d'une affection nécrotique des nervures de l'hybride Rup. Boscia. Rumbos I C. 1985. V. G. References Bouyahia H. and L R. 148-152. Incidence of some graft-transmissible virus-like diseases of Grapevine in visually selected and heat-treated stocks from Southern Italy. Martelli. and A.. 59-65. 9. 1989. 1978. D. Suggestion than vein necrosis is a specificic reaction of 110R to GRAPaV. de la marbrure de V. i Chim Nauka 1.N.: In southern Italy. J. Vein necrosis: new virus-like disease in Turkish vineyards. Mycoplasma-like bodies in phloem tissue of grapevine affected by vein necrosis. V. 1994. Journal of Turkish Phytopathology 17. D. Abracheva and B. 1973. SSR. Bulletin OEPP/EPPO Bulletin 22. Vein necrosis a disease that is latent in most grapevine cultivars of the State of Rio grande do Sul.. 1988. Is Grapevine vein necrosis a reaction to Grapevine rupestris stem pitting-associated virus? Journal of Plant Pathology 86.Z. Biol. Necrosi delle nervature della vite in Italia e Bulgaria..P. 3-5 Martelli G. Krake. Kalashyan.A.. rupestris. La Notte. 43-45 Kuhn G. Vein necrosis. Martelli G. 1978. D. 61 Savino V. Woodham R. Krake: Vein necrosis in Australia Savino et al. 1993. Heat therapy reduced this proportion to 35. Potential interaction between rootstocks and grapevine latent viruses.: Vein necrosis in Hungary Gursoy: Vein necrosis in Turkey Rumbos: Vein necrosis in Greese Martelli et al. 125 . C. Boscia and V. Fitopatologia Brasileira 19. Journal of the Australian Institute of Agricultural Sciences 50. Savino. 79-83 Legin R. Ser. 1986. Golino D. 29-30.C. Lazar.1984 1985 Woodham R.. Castellano. Viruses of grapevine in Malta.. 204-207. 57-63. Rivista di Patologia Vegetale. M. Kergazdasag 18 (4). Detection of vein necrosis virus (GVNV) in the vines of cultivated grape varieties.: Vein necrosis in Malta Golino: Vein necrosis in California Khun: Vein necrosis in Brazil Bouyahia et al. Grapevine vein necrosis disease detected in rooststocks in Australia. Lehoczky et al. Informatore Fitopatologico 28 (10). Pirolo.. Farkas. 301.-Berl.: An association exceeding 95% observed between GRSPaV and 110R vines showing vein necrosis symptoms in indexing trials.R. No veing necrosis observed in 110R top grafted on GRSPaV-free V. Phytopathologia Mediterranea 24. 1986 1988 1989 1992 1993 1994 2004 3. P.. IV. Vuittenez. Boscia and G. Minafra and G. ser. Rosciglione. American Journal of Enology and Viticulture 44. and J. Martelli. A. 1984. 606-612. H. Savino. Gursoy Y.. Phytoparasitica 17. P.A. but did not eliminate the disease entirely.P. 58-60.C. the incidence of vein necrosis in visually selected stocks of table and wine grape varieties is on the average 71 %. and L.5 %. 1992. Milkus B. Izvestiya Akademii nauk Moldav. 110 R. fleck and leafroll in Vitis vinifera and grapevine rootstocks in central Greece. 2004.P.B. Galea Souchet. Lehozcky J.A.P. Comparaison des symptômes et transmission par greffage d'une mosaïque nervaire de Vitis vinifera.
VIROID (Yellow speckle) .
Neither of them is able to replicate in herbaceous hosts but both were succesfully inoculated to grapevine seedlings reproducing the yellow speckle syndrome. Europe. 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. Description Main synonyms: Moucheture jaune (Fr. or gathering along the main veins to give a vein banding pattern. Gelbsprenkelung der Rebe (Germ. 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. are subviral pathogerns endowed with autonomous replication in their hosts. Interestingly.). It was first detected in Japan and transmitted to cucumber and grapevine seedlings in which. climatic conditions. respectively and both belong in the genus Apscaviroid. Citrus exocortis viroid grapevine strain (CEVd-g). AGVd.). Five grapevine-infecting viroids are known.). and AGVd. Randles and J. Like viruses. CEVd-g. The symptomatology varies depending on the cultivar. and may have a worldwide distribution. GYSVd-1 and GYSVd -2 cause the disease individually or in combination. Agents: Two distinct viroids. HSVd-g has been recorded from Australia. presence of ribozymes. 129 . a member of the genus Apscaviroid.W. Viroids. Grapevine yellow speckle viroid 2 (GYSVd-2). HSVd-g. a disease characterized by chrome yellow flecks localized along the main veins of mature leaves and progressing into the interveinal areas.. Very often. R. and perhaps the type of infecting viroidal sequence variant.VIROIDS Viroids. 370 pp. are not associated to any specific symptomatology. plant age. and Tunisia. Main symptoms: Few to many minute chrome yellow spots or flecks scattered over the leaf surface. Semancik (eds. Vein banding. Sometimes. has a genome 369 nt in size. 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. infected vines are symptomless or show symptoms erratically. Australian grapevine viroid (AGVd). References Hadidi A. respectively from a cv. GYSVd-1 and GYSV-2 have a worldwide distribution The three additional viroids that have been detected in grapevines. GYSVd 1 and GYSVd 2 are made up of 366 and 363 nucleotides (nt). viroids are classified in families. Collingwood. the USA. however. however. the non coding genomes. Only GYSVd-1 and GYSVd-2 are pathogenic. north and south America. and plastidial replication (Avsunviroideae ). Flores. Cabernet franc and a cv. Both these viroids wer first isolated in Australia. HSVd-g. genera and species. 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. vein banding-like symptoms can be observed in vines infected only by yellow speckle viroids. These symptoms appear in the height of summer on a limited number of mature leaves and persist for the rest of the vegetative season. was demonstrated to be caused by a co-infection by yellow speckle viroids and GFLV. it did not induce symptoms. They are made up of a non encapsidated circular RNA of 246-375 nucleotides. 2003. a size much smaller than that the smallest viral genome. AGVd has been reported from Australia. has a genome 297 nt in size. J. Hop stunt viroid grapevine strain (HSVd-g). Kyoto vine with yellow speckle symptoms. picchiettatura gialla (Ital. all of which belong in the family Pospiviroideae: Grapevine yellow speckle viroid 1 (GYSVd-1). thought to be elicited by a specific strain of GFLV.). the type species of the genus Hostuviroid. CSIRO Publishing. YELLOW SPECKLE 1. inducing a disease called yellow speckle.S.
Krake and Woodham: Evidence that the agent of yellow speckle is implicated in the etiology of vein banding.: A disease of cv. if not all citrus-growing countries. 1984 1985 1985 1985 1987 1987 1988 1988 1988 130 . Flores et al. Transmission: No vector is known. a disease formerly thought to be caused by a chromogenic strain of GFLV. First identification of AGVd Koltunow and Rezaian: Identification and sequencing of grapevine yellow speckle viroid. and distribution of infected propagating material. yellow speckle and fleck through dodder. Sano et al.: The Japanese grapevine viroid identified as a strain of hop stunt viroid. It was first recoverd in Spain from symptomless grapevines. Semancik et al. CEVd-g and AGVd.: Successful mechanical transmission of viroids to grapevines. Shikata et al. besides Spain. a member of the genus Pospiviroid. Experimental transmission through dodder is possible. Mink and Parsons: Yellow speckle can be detected by growing vines for 2-3 weeks at 32 °C under continuous illumination. Woodham and Krake: Evidence of field spread of yellow speckle.: Two new viroids. Seed transmission has been demostrated for GYSVd-1.CEVd-g. Barlass et al. GYSVd-2.: A vein banding condition of cv. In the great majority of grapevine germplasm infection is latent. its grapevine strain so far has only been recorded from Australia and the USA. Prota et al. Rcatzitelli resembling yellow speckle reported from Bulgaria.: Yellow speckle eliminated by in vitro apical culture. the authors consider the results as inconclusive. This latter way of dissemination has been considered as more efficient and frequent than mechanical transmission.: Four viroids found in Australian grapevines. Woodham and Krake: Artificial transmission of grapevine leafroll. Detection: Some viroids can be transmitted mechanically to herbaceous hosts but this is not an efficient detection method. one of which identified as the agent of citrus exocortis. grafting. Three different viroids found in a number of accessions in a Californian varietal collection. For yellow speckle. Polyacrylamide gel electrophoresis has been used extensively before the advent of nucleic acid-based assays (hybridization and RT-PCR) which constitute far better detection and identification tools. has a genome 369 nt in size.: Reconstruction of the secondary structure of CEVd-g Szychowski et al. Cannonau not associated with the presence of GFLV reported from Italy. found in grapevine accessions from Europe and California. Although CEVd is present in most.: Evidence that viroids are widespread in grapevines. Varietal susceptibility: All Vitis species. Rezaian et al. Control: Use of viroid-free propagative material obtained by meristem tip culture. Natural dissemination takes place by mechanical inoculation through surface-contaminated cutting tools during management operations. Abracheva et al. 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). Garcia Arenal et al. hybrids and cultivars appear to be susceptible. 2.: First recovery of a viroid from grapevines in Japan. as the disease may have spread naturally.
Cordoba. A. Perreault and H.: First report of AGVd in the Mediterranean. Citrus exocortis viroid grapevine strain (CEVd-g). Journal of General Virology 66. Flores R. The occurence is reported of HSVd. Annals of Applied Biology 101. Martelli: Brief review of grapevine viroid situation supporting the idea that vein banding is primarily induced by viroidal rather than GFLV infection.: A survey of viroids of grapevine in Italy.1988 1989 1989 1989 1990 1990 Duran-Vila et al.: Review of viroids. Juarez and J. M. Little and Rezaian: Updated review of grapevine viroids. which require amplification in an alternate host. Molecular evidence that this viroid originated from recombination between five different viroids among which GYSVd-1 and GYSVd-2 1991a. according to an international agreement reached during the 10th Meeting of ICVG held in 1990 at Volos. Rezaian et al. N. 1991 1996 1997 1999 2001 2003 2003 3. Elleuch et al. 1982. 131 . 45-57. based on phylogenetical analysis of hop and grapevine isolates of this viroid. Grapevine yellow speckle viroid 2 (GYSVd 2) Australian grapevine viroid (AGVd). Proceedings of the 6th Meeting of ICVG. Skene. Production of viroid-free grapevines by shoot tip culture. 2095-2102. Regeneration of virus-free grapevines using in vitro apical culture.S. Quacquarelli and V.M... A possible virus disease of Rcatzitelli vines in Bulgaria. R. Minafra et al. Semancik.: Improvement of meristem tip culture technique for the production of viroid-free grapevines. Savino. Greece. These viroids.G. Wan and Symons: Transmission of GYSVd-1. 1978. (ii) enhanced viroids. Koltunow et al. References Abracheva P.: Extensive comparative analysis of grapevine accessions from California and Europe reveal a similar pattern of viroid distribution. Fakhfakh. G. Flores et al. 1991 Semancik and Szychowski: There are two classes of grapevine viroids: (i) apparent viroids.M.. Journal of Plant Pathology 85. Koltunow and Rezaian: Description and sequencing of grapevine viroid 1B (later renamed Grapevine yellow speckle viroid 2). Krake.b Szychowski et al. Wang et al. Woodham and L. Grapevine yellow speckle viroid 1 (GYSVd 1). V. 291-295. GYSVd-2. 127-130. CEVd-g and AGVd via grape seeds. 1976. J. Marrakchi.: Evidence that two related viroids (GYSVd 1 and GYSVd 2) can cause yellow speckle disease independently. Sano et al.P. 217-220. American Journal of Enology and Viticulture 39. 2003. Spain. Duran-Vila N. GYSVd-1 and GYSVd-2 Rezaian: Complete nucleotide sequencing of AGVd. which can readily be isolated directly from grapevines..P.: Structural analysis reveals that five distinct viroids infect commercial grapevine varieties. K. First report of Australian grapevine viroid from the Mediterranean region. Monografias INIA 18.C. are to be named as follows: Hop stunt viroid grapevine strain (HSVd-g). Duran-Vila.: Suggestion that the viroid causing stunting in hop (HSVd) originated from grapevines. 1985. Arregui.. Detection of viroid and viroid-like RNAs from grapevine.R. Elleuch A. J. Barlass M. Pallas and J. 1988. Martelli.: First record of grapevine viroids in China.
N.Woodham. 1990. 1989. In: Viroids (A. Zhang and X. Australian Journal of Agricultural Research 23. 25-36. Viroids of grapevines in Italy.S. Arab Journal of Plant Protection 7.a newly recognized grafttransmissible disease of Vitis. Krake.R. 53-59. Krake. Hadidi. Credi.. Nature. J.C. 1972.). 285-291 Wang G.A. Semancik. 173-182. Grapevine viroid 1B. Johnson and M.A. Rezaian M.L. A.P.. 869-872. Rezaian. Leclair. 1990.. Taylor R. Grapevine yellow speckle viroid: structural features of a new viroid group. 1985. detection. American Journal of Enology and Viticulture 38. Seminars in Virology 8. Grapevine viroids.R. Flores. Martelli and V.A.. Semancik.. American Journal of Enology and Viticulture 39. Rezaian. 24-28. Koltunow. 3411-3419. 1988. 2003. 1990. Krake L. a new member of the apple scar skin viroid group contains the left terminal region of tomato planta macho viroid. 1985.W. 35-40. Duran-Vila.M. Relationships among grapevine viroids from sources maintained in California and Europe. Woodham. 1987. China. P.M. Semancik J. Ohshima.. and J. Proceedings of the 10th Meeting of ICVG. Minafra.H.S.R. Phytopathologia Mediterranea 24. Skene. 1988. Rapid indexing procedures for detecting yellow speckle disease in grapevines. Di Serio and C. and M. Okado. L. Grapevine yellow speckle -. Nucleic Acids Research 16. Journal of General Virology 66. Hong. Z. Koltunow A. Duran-Vila. Journal of General Virology 70. Phylogenetic analysis of hop and grapevine isolates of hop stunt viroid supports a grapevine origin for hop stunt disease. Comparative properties of viroids of grapevine origin isolated from grapevines and alternate hosts.M. Australian grapevine viroid . Pallas and R. and J. 1991b... Wolpert and J. Szychowski J. Koltunow A. Isolation of three viroids and a circular RNA from grapevines.R. The field investigation of virus and viroid diseases of grapevine and stone fruit trees in Shandong and Liaoning province. J. Ohno and Y. A. yellow speckle and fleck diseases by dodder. 1989.A. S. Proceedings 10th Meeting of ICVG.Leclair. Uyeda. and M. Szychowski. 447-452.D. 65-73. Semancik J.A. Meshi. Semancik (eds. CSIRO Publishing. Garnier. 337-345. Investigations on transmission of grapevine leafroll. F. Virology 170. R. M. 2001. Doazan. R. 132 .. 1975. 39-41. Greece. Mimura and K.P. T.C. Garnier. 849-864.A. 195-206. An infectious low molecular weight RNA was detected in grapevines by molecular hybridization with hop stunt viroid cDNA. Proceedings of the 10th Meeting of ICVG. Little A. Wan C.R. A. Viroids: the non coding genomes. Sano T.evidence for extensive recombination between viroids. Plant Disease Reporter 59. R. N.A. Goheen and J.P. Proceedings of the Japanese Academy of Science 60(B). Grapevine yellow speckle agent implicated in the aetiology of vein banding disease. Koltunow A. China Fruits 4.S. Krake and K. 193-198.C. 1991. Woodham R. Sano T. Rezaian M. Phytopathologische Zeitschrift 106. 297. 1990. and L. Minafra. 210-219.A. Transmission of viroids via grape seeds. Greece.Flores R. T. Sano and I.A. Zhang. I. 270-278.. Minafra A. Shikata. Prota U. S. Garau. M. J. 1996. Virus Genes 22. Vitis 29.C. Greece. A viroid-like RNA isolated from grapevine has high sequence homology with hop stunt viroid. 287-288. Szychowski J.P. R. Goheen. 40-50. Journal of Phytopathology 147.S. R.I. 1983.M.A. 333338. Investigations on a vein banding disease of Grapevine in Sardinia. Uyeda. Grapevine yellow speckle disease: studies on natural spread observed in the field.A. 1988..Jiang. Widespread occurrence of viroid-like RNAs in grapevines. Randles and J. Semancik. Collingwood. 575-578.R. Hernadez.G. Martelli G..W.S. 1983. T. Parsons. and L. and R. 1987. Koltunow and L. sanitation and situation in the Arab countries. 1997. Volos.M.H. Symons. 370 pp. E. Doazan. Rezaian. Wolpert and J. 1982. Two related viroids cause grapevine yellow speckle disease independently.C. 413-422. Shikata E. Vitis 30. J. Infectious diseases of grapevines. Krake. Mechanical transmission and rootstock reservoirs as factors in the widespread distribution of viroids in grapevines. and R. P. Garcia Arenal F.A.. and R. Journal of General Virology 69. Flores. V. Dore. 5587-5598. 213-216. A. Volos. Rivera-Bustamante and A.C. and M.A. 1990. Vitis 22. Szychowski J.. Grapevine viroids. R. 1991a. The sequence of a viroid from grapevine closely related to to severe isolates of citrus exocortis viroid. N. Nucleic Acids Research 10. Credi. G.P. Relationship and patterns of distribution among grapevine viroids from California and Europe. 202-205. 1984. Mink G.S. 4203-4210. Cugusi and M. Woodham R. Rezaian. Volos.. Vitis 21. 1989. L. Krake.. J. A. Rezaian M. Nucleic Acids Research 15. 1991.. 1999. Savino. M.
Vergilbungskrankheit. The first GY to be recorded was Flavescence dorée in France in the 1950's. They are wall-less. Palatinate grapevine yellows. including roots. Phytoplasmas in the 16SrIII group (or X-disease group) are associated with grapevine yellows in North America and Israel. a phytoplasma belonging to group 16SrII (or peanut witches' 135 . 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. Quite often. shoots. vector species have not been identified for all GY diseases. Golden flavescence. Leaf blades are crispy and brittle. persistence of infection in dormant plant material.GRAPEVINE YELLOWS: GENERAL PROPERTIES 1. Nonetheless. The different GYs cannot be differentiated on the basis of symptoms. and transmission by specific leafhopper or planthopper vectors. inflorescence and berries. However. Stolbur phytoplasmas. Phytoplasma cells are vesicular rounded bodies. and serology. depending on the particular disease and other conditions such as variety. 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. autumn fall of leaves occurs later on diseased than on healthy vines. Nevertheless. in addition to other criteria such as symptomatology. Schwarzholzkrankheit. involving also the main veins. resulting in reduction of quantity and quality of the crop. associated to Bois noir (BN). GY agents have been identified in no less than 5 groups of phytoplasma clade. though some sieve tubes may appear crowded with phytoplasma cells in the electron microscope. Legno nero. Vergilbungskrankheit. Amarilliamento. Candidatus Phytoplasma australasia. Description Grapevine yellows (GYs) are a group of severe phytoplasma-induced diseases occurring worldwide on a number of Vitis vinifera cultivars. Phytoplasmas have the smallest genome reported for procaryotic organisms (560 2. 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. host range. Severely infected stocks decline rapidly. The main characteristics of GY diseases are important losses or damage to the yield. a severe decline of the vine. Downwards rolling of the leaves and sectorial discolorations of the blades. Australian grapevine yellows. The occurrence and spread of a particular GY mainly depend on the presence of efficient vectors in the vineyards or their close vicinity. 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). However. canes. Rubbery canes fall downwards with a typical "weeping" aspect. their movement is slow and their distribution in the plant is erratic. However. Bois noir. Flavescence dorée (FD) and Palatinate Grapevine Yellows (PGY) phytoplasmas belong to group 16SrV (or Elm yellows group). Phytoplasmas form a homogeneous phylogenetic clade subdivided into about 20 groups and subgroups. In 1997. but not seeds. buds. based mainly on sequence similarity of their rRNA genes and of a few other genes. phloem-restricted bacteria that belong to the class Mollicutes. Bunches wither in early summer or berries shrivel later in season. Main diseases: Flavescence dorée. Main synonyms: Flavescence dorée-like disease. North American grapevine yellows. Flavescenza dorata. climate and infection pressure. 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. their genome is poorly known because they cannot be cultivated. Flavescenza dorata. Their titre is usually very low in grapevine. the term phytoplasma has become the genus name of these plant pathogenic agents under the provisional taxonomic status Candidatus. develop on leaves. variable in shape and size (50-1000 nm in diameter) that circulate in phloem sieve tubes and can be seen in the electron microscope passing through pores of sieve plates.200 kbp). all organs of the plant may be infected. such as the ribosomal protein genes or the elongation factor Tu. after their aetiology had been understood and molecular methods for the characterization of associated phytoplasmas had been developed.
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. shoots. Double infections have been reported. Vitis riparia and American rootstock varieties and hybrids do not show typical symptoms of GY. These phenomena also depend on the disease complex (phytoplasma. Transmission: Phytoplasmas are vectored by hemipters. vector and abundance of reservoirs). Then. Individual symptoms can be confused with other diseases or disorders. Pinot noir. and bunches is strong evidence for phytoplasma infection. Other host plants: Most phytoplasmas are ubiquitous plant parasites that can be hosted by several sometimes tolerant plant species. Simultaneous presence of symptoms on leaves. Riesling and also numerous local varieties. but the latter technique has never been successfull on field-grown grapevines. 136 .broom group) is a second agent of Australian grapevine yellows. Phytoplasmas persist in their vectors but vertical transmission to the progeny of infected gravid females is not possible or scarce. The three ascertained vectors of GY diseases are univoltine leafhopper or planthopper species. Cabernet Sauvignon. on less conserved genes. Full recovery and transient remission of symptoms have been observed in low and medium sensitive cultivars. mainly hoppers (cicadellids or cixiids) or psyllids. more specific primers can be used. including the agents of FD and BN. canes. Propagation material may be the infection source for long distance dissemination of GY diseases. when grapevine is not a preferred host for the vector. Though the rate of transmission to plant material can be very low. Phytoplasmas can be detected by graft transmission to sensitive vines. When no information on the particular phytoplasma type is available. which allows restoring the sanitary status of affected vineyards when the vector activity is controlled. When the presence of a particular disease is suspected. using DAPI staining. Phytoplasmas in the 16SrI group (or aster yellows group) are associated with endemic GY diseases in several countries of Europe and in the USA. are critical. erratic transmission to grapevine may nevertheless take place. influence symptom expression and differential sensitivity of cultivars. These methods are not specific for any phytoplasma and are too laborious for disease monitoring. Others. Most of these PCR primers have been designed on conserved regions of the rRNA gene and are "universal" for all known phytoplasmas. Detection can also be made by transmission electron microscopy of thin sections or by scanning electron microscopy. However. It is probable that the feeding preference of insect vectors. are group specific. Varietal susceptibility and sensitivity: Numerous V. ELISA detection is possible from vascular tissue of symptomatic grapevines. Hence. The best antigen source are leaf veins and petioles. are available in the literature. are very sensitive to all GY diseases. designed on variable regions of the rDNA. The extraction from infected tissues of total DNA containing enough phytoplasma DNA of good quality and the elimination of inhibitors of the PCR reaction. as well as their feeding activity. Phytoplasmas also persist in vine stocks during winter. Outstanding progress in detection was achieved with DNA-based techniques. or on random selected non-ribosomal DNA fragments. They can also be observed in sieve tubes by light microscopy. Polyclonal and monoclonal antisera have been raised to a few phytoplasmas. vinifera cultivars have been reported to be sensitive or very sensitive to GYs in all countries where infection occurs. Random Fragment Length Polymorphism (RFLP) analysis of the amplicon can provide further characterization of the infecting phytoplasma. PCR amplification with universal primers is preferred. Some cultivars such as Chardonnay. The situation of cv Syrah is controversial. Detection: GY syndrome is characteristic. Particular procedures for extraction of phytoplasma antigens from infected vines must be used to enhance the sensitivity of detection. A range of primers that can be used for amplification with Polymerase Chain Reaction (PCR) of characteristic DNA fragments of phytoplasmas. such type of transport is risky when potential insect vector species occur on the site of planting. Infection of rootstocks has been detected only in the case of Flavescence dorée but it cannot be excluded for other GY agents.
BN is considered as a non epidemic form of FD. However. tolerance and potential for recovery of varieties. i. Control of GY diseases depends on the knowledge of vector insect species. into hot water (50 °C) for a sufficient length of time (minimum 30 mn). Generally. 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. of a disease occurring in the vineyards of the Mosel Valley and of the Rhine Valley.Other molecular methods are being developed. Schvester et al. an insect that has been introduced recently from America. The biology of the insect is described. Pruning or top-grafting are useless because roots and trunks are infected. 1956 1957 1961 1961 1961 1964 1965 137 . soaking of dormant material before or after grafting. weeds or other crops. In any case. under the name Flavescence dorée. Investigations are being conducted on sensitivity. The disease can be transmitted by grafting and is probably a virus disease. FD can be detected from leaves of non-symptomatic American roostocks. FD is transmitted by the leafhopper S. on physiological changes and defence mechanisms induced by infection. Control: There are no direct curing methods of infected plants. cultural practices. Symptoms (macroscopic and microscopic). Detection of phytoplasma DNA can be achieved from all plant organs or insect vectors.e. Caudwell: Characterization of a new type of flavescence. but does not rule out the possible involvement of a pathogen. phytoplasmas are unevenly distributed in GY-affected vines. evolution of the disease in space and time.. and on elicitors of defence reactions to these phloem-restricted bacterial agents. Control methods of vector populations are being experimented with natural insecticides. efforts should address prevention by planting healthy propagation material and limitation of vector populations and their inoculative activity. hypotheses on its nature. Production of sanitized propagation material is possible with hot water treatment (HWT). The author suggests this disease to be due to adverse climatic and soil conditions. on the possibility to limit or eradicate their population and prevent their migration or movement in vineyards. Vidano: S. littoralis Ball found in Italy in 1963. Caudwell (b): Description of recovery in grapevines affected with FD. The disease is thought to be caused by root damage. Real-time PCR has also been successfully used. 2. but is also possible using young leaves shortly before symptom expression or phloem scrapings from lignified canes. Detection in the upper parts of the vines is usually done from veins and petioles of the leaves of symptomatic shoots. Nonsymptomatic grapevines that show irregular symptoms from one year to the next may also test positive. Caudwell (a): Studies on Bois noir (BN) and on its relationships with FD. and natural enemies. Branas (a and b): Description of flavescence epidemics on Baco 22A and Chardonnay in France. Historical review 1955 Levadoux: Description of a severe outbreak of "Flavescence dorée" on Baco 22A in southwestern France. and on the identification of reservoirs of inoculum such as infected grapevines. using detection of PCR amplification products with DNA-DNA hybridization with a specific probe. denoted Flavescence dorée (FD) in agreement with the name proposed by Levadoux (1955) because of the golden yellow metallic aspect of leaves. Gärtel: Description. littoralis Ball.
The author assumes that both FD and its vector were introduced from North America with varieties of Vitis labrusca between 1927 and 1950. Caudwell et al. The disease has been observed for the first time in 1968. littoralis. Symptoms on V. 1967 1968 1970 1971 1971 1973 1973 1975 1977 1979 1980 1982 1982 1983 1984 1985 1985 138 . Caudwell et al. littoralis Ball is present in Ticino (southern Switzerland). An important FD-like disease is reported. (a): Mycoplasma-like organisms (MLOs) are observed in diseased grapevines and infective leafhoppers and in Vicia faba submitted to feeding inoculation with infective S. Regina. First record in 197576. Potential existence of several different diseases: leafhoppers (Euscelidius sp. (b): Evidence that FD and BN are two different diseases with similar symptoms.: Discovery of Mycoplasma or PLT group-like microorganisms in the phloem elements of plants showing dwarfing. the hypothesis is put forward that this nematode is the vector of the disease. Osler et al. Magarey and Wachtel: Description of a new disease of grapevine of the yellows type on the cv Rhine Riesling in South Australia. Recommendation that mother plants should be grown in areas far away from FD-infected regions. As similar organisms have been found in nematodes of the species Xiphinema index. Tanne and Nitzany: Occurrence of a yellows disease resembling FD in Israel. faba are different from those obtained with feeding inoculation with infective S.1966 Caudwell: Attempts to inhibit the agent of FD in vivo by heat treatment. and Euscelis sp. titanus (= littoralis) is not a vector. with special reference to grapevine.) caught in the vicinity of vineyards could transmit a yellows disease to Vicia faba plants. This phytoplasma was later on identified as a Clover phyllody phytoplasma. titanus fed on the latter V. Caudwell et al. Granata: Description of an epidemic yellows disease on cv Inzolia in Sicily. Caudwell: Symptoms of grapevine yellows (Amarilliamento) reported in Chile on cv Elqui. Bois noir. Caudwell and Larrue: Spread of FD in France is related to sanitary status of planting material. Inoculation of grapevine seedlings with S. Immersing cuttings taken on FD-affected grapevines in water at 30°C prior to planting reduced the proportion of infected plants by about 83%. littoralis. mainly on cv. littoralis is present in the same region of Oltrepò Pavese. Mosel and Rhine regions are considered as the causal pathogens of this disease. of which S. is probably of European origin. Conversely.: S. Doi et al. Belli et al. Rumbos et al. Belli et al. witches' broom and yellowing.: 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.: Rickettsia-like organisms observed in roots of grapevines with "Vergilbungskrankheit" in the Saar.: Presence of a disease similar to FD in the Oltrepò Pavese (northern Italy). faba plants produced typical GY symptoms. these findings have not been confirmed.: S. Rafaila and Costache: Yellows of grapevine with symptoms similar to those of FD found in 1967 for the first time in Romania. So far. Provisory name "Rhine riesling problem". Caudwell: Discussion on the origins of yellows diseases of plants. Baggiolini: S. Credi and Babini: Grapevine yellows are reported in Emilia Romagna (Italy). titanus found in 1984 in vineyards of northern Italy.
vector of FD.: Contribution to the knowledge of FD (or similar diseases) in the Veneto region. titanus. Vidano et al. In addition to S. Vidano et al. Recovery and crop loss are variable.: Observation of MLOs in phloem tissue of yellows-affected grapevines in Australia.: Survey of potential Auchenorryncha vectors of the pathogen agent of FD in Piemonte (Italy). Euscelidius variegatus and Euscelis incisus are taken into consideration. Borgo et al.: Occurrence of FD-like symptoms in the valley of Sarca in Trentino. vector of FD.: Presence and distribution of a FD-like disease in the region Friuli-Venezia Giulia in Italy. their etiology is unclear. Quaroni et al. northern Italy. Pinot nero) and comparison with other similar diseases. Seljak: Presence of Scaphoideus titanus in western Slovenia (Yugoslavia).: New data on the spread of FD in three vineyards of Oltrepò pavese (northern Italy) from 1985 to 1987. Hyalesthes obsoletus. Pinot bianco. titanus.or xylem-limited prokaryotes in Europe. Chardonnay is the more affected variety. Boudon-Padieu and Larrue: ELISA detection of FD pathogen on infected reared experimental and wild natural leafhopper vectors (E. control measures. in viticultural areas of northern Italy and of other Auchenorrhynchas susceptible to play a role in the transmission of yellows diseases in this region. whereas an unsprayed vineyard was more affected. using the scanning electron microscope. 1987 1987 1987 1988 1988 1988 1988 1988 1988 1989 1989 139 . Italy. Egger and Grasselli: Presence in Toscana. Magarey et al. variegatus and S. Susceptibility and sensitivity of affected varieties (Chardonnay. However. of a FD-like disease on Chardonnay.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. for the presence of a FD-like disease. of MLOs in phloem tissues of grapevines affected by "FD" in northern Italy.: Observations.: Description in Italy of the presence of FD or FD-like diseases. Mescalchin et al. Carraro et al. titanus. titanus. Rui et al.: Presence of a FD-like disease in Emilia-Romagna. Italy. disease distribution.: Study on the distribution of S. resulting in a reduced diffusion of the disease. Martelli: A review on the knowledge on grapevine diseases induced by phloem. Credi et al. The results suggest that the disease is brought into the vineyards from outside local sources. Credi and Callegari: Survey of vineyards in Emilia-Romagna. titanus is not always associated to the disease. Fortusini and Belli: Description of the development of epidemics of FD-like diseases in northern Itay. Fortusini et al. Survey for the presence of S. Conti: A review of intracellular procaryotic phytopathogenic agents. that are responsible for severe decline of vines. S. Two of these vineyards were sprayed with insecticides in 1986 and 1987. Italy. respectively). Italy. Borgo: General information on the presence of FD-type diseases in northern Italy. Credi: Description of a FD-like disease in Emilia Romagna and the reaction of 3 different cultivars.
Chardonnay develops in Tuscany. S. Deng and Hiruki: A method to amplify 16S rRNA gene from culturable and nonculturable mollicutes. Specific primers can be used for MLOs. Granata and Russo: An epidemic disease with FD-type symptoms develops in Sicily on cv. Symptoms also developed on plants obtained from buds taken on diseased Chardonnay and Sangiovese grafted on healthy Kober 5BB. Refatti et al. 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). Similar symptoms on Chardonnay could be obtained by insect inoculation (Euscelis incisus) and by cleft-grafting of tissues taken from yellows-infected elm. Restriction profiles after digestion with endonucleases permit the differenciation of MLOs associated to different plant diseases. Di Terlizzi et al.: Occurrence of grapevine yellows in Moldavian SSR. Caudwell et al.: The FD-like disease developing in Friuli-Venezia Giulia since the early 1980's was not decreased significantly in vineyards sprayed with insecticides though S. Natural infection with MLOs of bait plants (Catharanthus roseus and Vicia faba) placed in vineyards in spring and summer. Treatment prior to storage is more efficient. Vidano et al. Caudwell: Review on the epidemiology and characterization of FD and other GY diseases.: Bench grafting experiments of buds of indicator cvs. Caudwell and Kuszala: Detection with ELISA of FD-MLO in naturally FD-affected grapevines. Boidron and Grenan: Construction and testing in ENTAV. (a and b): Identification of numerous species of Auchenorrhynchas in the vineyard ecosystem in Piemonte (northen Italy) and of ampelophilous species. 1989 1990 1990 1990 1990 1991 1991 1991 1991 1991 1991 1991 1992 1992 1992 140 . titanus was not found in the area. Credi and Santucci: Development of "Flavescenza dorata" in grapevine in Emilia-Romagna and rate of infestation in different systems of cultivation. Inzolia in Sicily.: Grapevine graftwood shoots infected with FD can be disinfected by hot water treatment in the dormant stage.: Presence of yellows-like symptoms in Apulian grapevines (central Italy). Inzolia. Conti: A yellows-type disease of cv.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. The annual rate of symptomatic plants was never greater than 20 % over a 6-year period. Granata and Grimaldi: MLOs are observed with the electron microscope on affected grapevines of cv. The recommended temperature / time combination is 50°C / 3560 min. Credi et al. Several weeds with symptoms of phyllody were found to contain MLO in electron microscope studies. The aetiology is not fully ascertained though the presence of MLO is probable. 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". Le Grau du Roi (France) of a device for secure soaking of dormant buds and canes into hot water (50°C. Chardonnay and Baco 22A grafted on donor yellows-diseased plants of cvs Pinot blanc and Sangiovese. Only part of the plants were infected. Marinesku et al. titanus was present in all vineyards checked. This is a prospect for investigation on MLOs associated to plant yellows. Affected grapevines in the Rhône valley tested negative. 45mn). suggesting an unrelated agent.
Boubals (b): A severe epidemic of grapevine yellows occurs in Golan (Israel). Caveccia and Chardonnay vines to periwinkle by the dodder species Cuscuta campestris. but are different from known aster yellows cluster MLO strains. No spontaneous recovery. Arzone et al. Typical symptoms on several cvs. (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. Meignoz et al. IPVR is related to aster yellows MLO. Davis et al.: Description of MLOs and associated disorders in the phloem of FD-affected LN33 experimentally inoculated with S. Osler et al. titanus is not present. FD-MLO DNA can be distinguished from other MLO DNA with Dot-blot hybridization and Southern-blotting. titanus and other leafhopper species show that yellows disease present in northeastern Italy differs from flavescence dorée as it occurs in France. Arnò et al. 4 positive results (0.: FD can be transmitted by symptomless rootstocks. Senescent or degraded forms of MLOs identified inside degenerate sieve elements. MLOs were visible in sections of petioles of periwinkle showing symptoms and subsequent graft transmission from periwinkle to periwinkle was successful.: 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). (a and b): Observations supporting the existence of different types of grapevine yellows in northern Italy. It is more related to the agent of Elm yellows than to other yellows agents.: ELISA with FD-antibodies permit to detect related antigens in S. One MLO strain transmitted with Macrosteles quadripunctulatus appears similar or close to aster yellows MLO. genomic tests). Chen et al. titanus population in vineyards and the rate of spread of the disease. Bertaccini et al. Hot water treatments suppress the transmission to Chardonnay indicators. The MLO strains found in grapevine are related with aster yellows MLOs. Bianco et al.: The survey in Piemonte of Chardonnay vineyards affected with a FD-like disease shows that no correlation can be made between the importance of the S. Attempts to characterize MLO DNA extracted from insects with molecular methods. titanus leafhoppers. Caudwell et al. detection (ELISA. Daire et al. (a and b): Characterization of Italian periwinkle virescence (IPVR) MLO (obtained on bait periwinkle plants exposed in yellows-affected vineyards in Italy) is possible with DNA-DNA hybridization and PCR. S. However.6%) were obtained. Quacquarelli and Barba: Review of the distribution of flavescence dorée and other grapevine yellows in EEC viticultural countries.: Transmission experiments from grapevine to grapevine and to herbaceous hosts with S. results of research. Evolution of the disease and of its vector in the various viticultural regions of France. transmission by feeding to healthy grapevines succeeded only in 1 attempt out of 100.1992 Credi and Santucci: Of 628 attempts to transmit the agent of a yellows disease of grapevine from Sangiovese.: 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 . Alma et al. (b): Evidence of an association of MLOs with grapevine yellows in Emilia Romagna. titanus. Boubals (a): Report on a meeting of the French working group on FD. FD can be readily distinguished from Bois noir. The titre of MLO appears very low in grapevine. suggesting the presence of two different diseases at least in Veneto and Friuli-Venezia Giulia.
Credi (a and b): Observation of MLO in yellows-affected grapevines in northern Italy. Kuszala et al. S.called FDU. BN (stolbur MLO) detected in several regions of Italy and in Israel. titanus) and a second one. 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). Description of pathological changes in phloem of leaves and identification of senescent forms of MLO inside degenerate sieve elements. 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. c and d): Grapevine yellows (Vergilbungskrankheit) are common in the Moselle and Rhine valley. (a and b): Various attempts to transmit FD-like disease in different regions of northern Italy. Carraro et al. In northeastern Italy. International Committee of Systematic Bacteriology (ICSB). aster yellows and Xdisease. Detection in grapevines from Virginia (USA) of MLO related to X-disease. BN belongs to the stolbur group. (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. Only FD and BN have been identified in naturally affected grapevines. FD detected in leaves of rootstock 3309C and is also present in samples from Veneto (Italy). GYU was later on characterized by other authors as a phytoplasma belonging to the Xdisease (16SrIII) group. Osler et al. FD belongs to the elm yellows group but is different from elm phytoplasmas. Computer analysis for spatial patterns of diffusion show a non-random distribution and suggest transmission from other vines or from weeds. titanus is not present. Maixner (a. S. on the effects of pollarding affected vines and of chemical sprays against vectors. Positive assays obtained only on samples from France and Veneto. Classification of grapevine MLOs into three RFLP groups: elm yellows (FD from France).: FD-ELISA method was used on tissues of yellows-affected grapevines from diverse countries. then GYU phytoplasma) and comparison of specificity and sensitivity of the latter tools to detect phytoplasmas in grapevines in New York and Italy. healthy plants protected with plastic screens did not develop symptoms and diseased plants protected with screens showed transient recovery. Prince et al. So. 1993 Daire et al.: Six-year transmission trials of different types of yellows with S. using insects. probably transmitted by another insect. Experiments on the use of hot water treatment on planting material. MLO appear to be in very low titre. at least two different phytoplasmas are infecting grapevines in Veneto: FD sensu stricto (transmitted by S. FD is related to elm yellows and BN is related to stolbur. b. riparia in New York and failure of detection in some symptomatic grapevines in New-York and Italy. Daire: PhD thesis on detection and differentiation with DNA-based methods of the MLO agents of GY diseases in France. Transmission has been obtained only from vines of Veneto. Girolami and Egger: Importance of contamination of vineyards and diffusion of Grapevine yellows in northern Italy. Maixner: Demonstration that Hyalesthes obsoletus (Auchenorrhyncha: Cixiidae) is a vector of German grapevine yellows (Vergilbungskrankheit).: Diversity of MLOs associated with grapevine yellows and transmitted to periwinkle. titanus to test plants of cvs Perera and Chardonnay in Veneto and Friuli-Venezia Giulia (Italy). Electron microscopic observation of MLO in periwinkle and grapevine. Di Terlizzi et al. A MLO has been transmitted to periwinkle with dodder.: Important outbreak of a yellows disease on local varieties in Apulia (southeastern Italy) and infection of periwinkle as bait plants and with dodder transmission. Detection in non symptomatic V. titanus is present in the Friuli-Venezia Giulia region but no transmission assay has been successful. Specific probes cloned and specific primers designed for PCR amplification of phytoplasma DNA. 1993 1993 1993 1993 1993 1993 1994 1994 1994 1994 1994 142 . graft and dodder.
(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. Egger et al. Del Serrone et al. Ten weeds were found harboring MLOs with transmission electron microscopy. Italy). titanus which has not been found. (a and b): Grapevine yellows in Latium (southwestern Italy) are not associated with S. Albanese et al.: Four different phytoplasmas (FD. Boudon-Padieu (a and b): Reviews on knowledge and research on grapevine yellows. with the aim of identifying sources of tolerance or resistance to yellows diseases. Consideration on possible resistance and research on interaction of phytoplasma with nepovirus infection. Arzone et al. sometimes in mixed infection. in grapevines with yellows symptoms in Soave (Veneto. probably of Bois noir type.: Identification of BN (stolbur phytoplasma) in Spain. Molecular detection of aster yellowsrelated phytoplasma. Prince et al. (a): Specific detection with PCR of stolbur phytoplasma in grapevines affected with Vergilbungskrankheit. Fortusini et al.: The phytoplasma associated with Australian grapevine yellows is close to but different from stolbur phytoplasma causing BN. Haidar: First report of symptoms of yellows on grapevines in Lebanon.: Grapevines from Piemonte (northern Italy) contained several different phytoplasmas. Italy).: Presence and distribution of GY in Sicily.: Survey of yellows symptoms in an ampelographic collection of 1281 cultivars in Conegliano (Veneto.1994 1994 Parente et al. Padovan et al. The most frequent belonged to group 16SrI-G (later designated as 16SrXII or stolbur group). One grapevine out of 16 contained a 16SrV (= elm yellows) phytoplasma together with a 16SrI-G 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. obsoletus and in weeds growing in the vineyard in Germany.: Emphasis on the possible role of weeds as reservoir for MLO in vineyards. Alma et al. Double infection is reported. Koruza: Dispersal of grapevine yellows in Slovenia. aster yellows. Detection is also possible during winter on wood scrapings. Laviña et al. Maixner et al. Related MLOs were also detected in wild grapevines. Murari et al. Padovan et al. PCR detection of phytoplasma with universal primers. 1995 1995 1995 1995 1995 1995 1995 1995 1995 1996 1996 1996 1996 1996 1996 1996 1996 143 . Del Serrone and Barba (a and b): Monitoring of detection of phytoplasma in the organs of grapevine according to the vegetative state. Maixner et al (b): Detection of a new phytoplasma type related to FD in cv Scheurebe showing symptoms of yellows in Palatinate (Germany). in the vector H.: Presence of grapevine yellows in northern Portugal and electron microscopic visualization of MLOs in the phloem tissue of petioles of affected plants.: MLOs associated to grapevine yellows in Virginia belong to 2 different groups : X-disease group (16SrIII) and aster yellows group (16SrI). The importance of proper identification of disease type for control measures is emphasized. stolbur and apple proliferation) can be detected.
(b): Survey of possible auxiliaries for the biological control of leafhoppers and planthoppers in vineyards. Bosco et al. Borgo: Description and colour photographs of symptoms of grapevine yellows and knowledge on their aetiology and vectors. The term phytoplasma will become the genus name of plant pathogenic mycoplasmas under the provisional taxonomic status Candidatus. Tanne and Orenstein (a and b): Successful heterografting transmission of phytoplasma from symptomatic grapevine to periwinkle and easy subsequent identification of phytoplasma type. Del Serrone : A review in Italian on the knowledge on etiology. vectors and detection of the agents of Grapevine yellows. Murari et al. Among 32 species identified. Subclades are considered to represent the equivalent of distinct species. transmission. 10 are confirmed phytoplasma vectors. Belli et al. Subcommittee on the Taxonomy of Mollicutes: The taxonomy of phytoplasmas will refer to their molecular phylogeny. Maixner et al. Boudon-Padieu and Maixner: Updating in French and English of present knowledge on grapevine yellows: etiology. Phytoplasma belonging to the stolbur subgroup have been identified. Garau et al. AY and X-disease related phytoplasmas have been transmitted and detected.: Survey of hoppers in vineyards of the province of Piacenza (Emilia Romagna. Davis et al. (a): Only stolbur and FD phytoplasmas were detected in samples of yellows-affected grapevines from numerous regions of France and from Italy. Aster yellows and X-disease types were never detected.: Survey of leahoppers in vineyards of Piemonte. titanus is not present. The possibilities of control 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1998 1998 1998 144 . Italy).: Recurrent and severe yellows symptoms observed since the 1980's in Sardinia (Itlay) on several cultivars.: History and control of GY diseases in Italy. Batlle et al. titanus is not reported. (a): Description of available methods and strategies for detection of phytoplasmas in grapevine.: Identification of Flavescence dorée in Spain. Daire et al. 10 species were known vectors of phytoplasmas. International Committee of Systematic Bacteriology (ICSB). Spain and Israel. 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.1996 1997 1997 1997 Rüdel: Review on the history and diversity of Vergilbungskrankheit of grapevine in Germany. No double infection has been found to occur. (b): Prospects on detection and identification of grapevine phytoplasmas with new molecular tools and consequence on future epidemiology studies.: Symptoms of grapevine yellows observed in Hungary in the early 1970's. Maixner et al. Kölber et al. S. titanus reported for the first time in this region. 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). biology and control.: Use of Hot water treatment (HWT) to eliminate phytoplasmas (FD and AY) from 3 cultivars of grapevine. spread. northern Italy. Aldini et al. Presence of S. No epidemic outbreak has been observed. No positive detection with PCR on the HW-treated vegetative progeny of mature canes of symptomatic plants. S. Positive reaction with a DNA probe specific to aster yellows phytoplasmas.
1998 Lee et al. Lherminier et al. Batlle et al. epidemiology of these diseases. titanus. 1998 1998 1998 1998 1999 1999 1999 1999 2000 2000 2000 2000 2000 145 . titanus. BN is very frequent but H. FD-related phytoplasmas (elm yellows group) have been detected in grapevines from Treviso and Verona and transmitted from vine to vine using S. History. Carraro et al.: Summary of the complex situation of GY in northeastern Italy. Only FD and BN have been identified.: Evaluation on the efficiency and security of hot water treatment used to eradicate phytoplasmas from propagation material. Similar attemp with FD phytoplasma were not successful.: First report of the use of oligonucleotides for in situ hybridization with transmission electron microscopy to localize phytoplasma in plant cells.: Unsatisfactory results on the use of hot water treatments to eliminate phytoplasmas from grapevine wood.: A comprehensive review of the classification of phytoplasmas in the world. In the Veneto region only stolbur-group phytoplasma was recorded in some areas and both stolbur and elm yellows group phytoplasmas in other areas. Insecticide sprays against S. Sensitivity of assays must be good because of the low titre of phytoplasmas in affected plants. In addition a phytoplasma (FDU=GYU) transmitted from GYaffected vines to Catharanthus roseus by means of dodder in Friuli-Venezia Giulia. Seemüller et al. Maixner: A review of thermotherapy to cure phytoplasma infected material.: Phylogenetic studies of phytoplasmas associated to numerous plant diseases. FD is restricted to the north of Cataloña.: A review of the situation of GY in Spain. symptoms. Firrao et al. are compulsory in France.: A history of GYs in north-eastern Italy. obsoletus is rarely found in vineyards. epidemiology and etiology of GYs in the world. Guadagnini et al. Maixner and Reinert: Updating in German of the research made in different countries on grapevine yellows diseases. has been identified as a member of the X-disease group. Borgo et al.: Monitoring grapevine yellows in North-eastern Italy and attempts to rationalize routine laboratory testing for the differentiation between FD and stolbur phytoplasmas. Boudon-Padieu: Review on recent progress in the knowledge on grapevine yellows. though S. vector of FD. Phytoplasmas for which 16S rDNA sequence is available have been classified into 20 major groups and subgroups. Frausin et al. Search for alternative vectors.: Report of positive detection of aster-yellows phytoplasma in grapevine fed with Metcalfa pruinosa and in the body of the insects used for transmission. methods of detection of phytoplasmas in grapevine tissues and in insect vectors. with a particular reference to the work done in Germany. Refatti et al. Boudon-Padieu (b): Chapter in a Handbook on viral and bacterial diseases of the grapevine. On the basis of RFLP of PCR-amplified 16S rRNA gene. Reinert W. GY affect mainly the northern provinces. Survey with PCR of phytoplasmas infecting grapevines in Friuli-Venezia Giulia and Trento province (Italy) and in Slovenia. Probes were antisense to stolbur specific sequences in the 16S rDNA and used on stolbur-infected periwinkle. have shown that only stolbur group phytoplasma was consistently found. phytoplasmas can be classified into 14 groups and 38 subgroups. distribution in the world. according to the most recent studies on molecular structure of rDNA. Grapevine phytoplasmas classify into different groups. titanus is more widely distributed. nucleic acid hybridization and serological comparisons.depend on the knowledge of the biology of phytopathogenic agents and of their vectors. No negative effect was observed on scion buds but rootstock cuttings were more sensitive to negative effects of the treatment. and M.
: The presence of phytoplasmas in liquid medium on which insect vectors have been fed can account for potential vectorship of the species. A high rate of recovery is observed from one year to the next. although its vector S. 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.: 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. The latter was difficult to evaluate because of the non homogenous distribution of phytoplasmas in canes. Klein et al. HWT (50°C/40 mn) gave acceptable survival but none of the conditions experimented provided a total elimination of phytoplasmas. BN and aster yelllows in vineyards of southeastern Piemonte (Italy).: Use of immersion in hot water (HWT) or in chemicals to suppress phytoplasmas from grapevine propagation material in Italy. Plant variety and cutting diameter influence the rate of surviving cuttings. Phytoplasmas were detected in all five species.: Recommandations for the organisation of nursery in Australia and handling of hot water treatment to avoid dehydration of cuttings during treatment. Zahavi et al.: Development of a new technique with DNA chips to detect phytoplasmas in grapevine. Dual infection may occur (13 %). Bertaccini et al. Circulifer sp. which are all known as vector of phytoplasmas. Neoaliturus sp.: In Golan Heights (Israel) Hyalesthes obsoletus. confirmed the presence of three phytoplasmas associated to GY: stolbur (70%). Seljak and Petrovic: Overview on phytoplasma diseases of grapevine and fruit trees in Slovenia. sugar metabolism. nitrate and nitrite reductase. The phytoplasma agents are not specified. Quartau et al. 2000 2001 2001 2001 2001 2001 2001 2001 2001 2001 2001 2002 2002 2002 2002 2002 2003 146 . Marzachi et al. Crocker et al. Phytoplasmas seemed to disappear from the insect body after the transfer of insects to healthy plants. Seljak: A review of non-European hoppers introduced in Slovenia. Primers fitted for specific detection of these 3 phytoplasmas permit cheaper and quicker diagnosis. Clair et al. aster yellows (11 %) and X-disease groups phytoplasmas.: A 2-year survey in Golan Heights.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. Waite et al. Experimental evaluation of the method and application to field-trapped individuals of suspected vector species. Orenstein et al.: Use of hot water treatment in commercial nurseries in Australia to eradicate diverse pests and pathogen agents from propagation material. suggesting that they did not multiply in the body of the insects. Israel.: Survey of GY in Israel.: Scaphoideus titanus is present in Portugal. Hyalesthes obsoletus has been found but not in vineyards. Macrosteles quadripunctulatus and Orosius orientalis were found on weeds in vineyards or trapped on sticky traps. Influence of rootstock on the rate of GY symptoms in the same vineyard and same grapevine variety. FD has not been identified. Tanne et al. Boudon-Padieu: Updating of knowledge and research on grapevine yellows worldwide. Moretti et al. Osler and Refatti: The situation of GYs in northern Italy. (a): Presence of FD.: Report of experiments combining temperature and time of treatment with hot water on 5 grapevine cultivars to cure planting material from phytoplasmas. Frosini et al. titanus is widespread in western vineyards..
respectively.: Data on 15-year studies on efficiency and effect on propagation material of hot water treatment.. Chabbouh et al. (a and b): FD phytoplasma (16S rV-C) and S. Discussion on benefits and limits of artificial feeding medium to assess the transmission capability of phytoplasma by insects. Lessio et al. Ge and Maixner (a): Transmission to feeding medium is more efficient than transmission to natural host plants and to grapevine for Hyalesthes obsoletus and Oncopsis alni. Marzachi et al. 2003 2003 2003 2003 2003 2003 2003 2003 2003 2004 2004 2004 147 .: Compared efficiency. rapidity and sensitivity of methods for use in routine diagnosis of grapevine phytoplasmas.: Important presence of GY in Abruzzo (central Italy).: 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. Crocker et al. No important damage has been reported. stolbur phytoplasmas are suspected to occur because of long-lasting record of stolbur disease on many host plants in Serbia. BN (stolbur phytoplasma) has an incidence of about 30 %.: Identification of aster yellows (16SrI-B and C) and of ash yellows (16Sr-VII) related phytoplasmas associated to yellows of grapevine in Chile. in particular a clover phyllody type (16SrI-C) which is quite frequent.: Measurements of basal respiration rate of grapevine dormant wood as an indicator of the best period for application of hot water treatment.: Development of a sensitive PCR procedure for dual identification of FD and BN phytoplasmas. Milkus et al.2003 2003 2003 2003 2003 2003 Boudon-Padieu et al. M'hirsi et al.: Important outbreak of grapevine yellows on cv Chardonnay and identification of a stolbur phytoplasma in diseased plants. Tassart-Subirats et al. Gajardo et al. Osler et al.: Improvement of detection of FD and BN in field-grown grapevines with real-time PCR. Megophthalmus scabripennis positive for aster yellows phytoplasma. Hyalesthes obsoletus and Circulifer haematoceps abundant and positive for stolbur and aster yellows phytoplasmas. An aster yellows phytoplasma is suspected.: Flight activity of Scaphoideus titanus and Hyalesthes obsoletus and their infection status towards FD and BN (stolbur) phytoplasma. two nonpreferentially ampelophagous vectors of grapevine phytoplasmas. Clair et al. Leitner: The situation of Grapevine yellows in Austria is now surveyed by means of molecular detection assays. Recovery is a progressive phenonenon developing over 3 years in the case of BN. Orenstein et al. Other phytoplasmas are reported. Duduk et al. Myrta et al. Study of the spatial and temporal dispersion of the four species. fitted to routine detection. Kuzmanovic et al.: Report of stolbur phytoplasmas in grapevine affected with yellows in Albania.: Survey of potential vectors of phytoplasmas in vineyards of the Golan Heights. Neoaliturus fenestratus. Consequently roguing of affected vines should be avoided when possible and especially if vectors are controlled.: Report of aster yellows phytoplasmas in grapevine affected with yellows in Tunisia.: Grapevines affected with yellows are found free of phytoplasmas after recovery. Apart from FD phytoplasma reported by Duduk et al.: Identification of grapevine yellows symptoms in Tunisia and attempts to identify pathogen agent and vector. titanus found associated to grapevine yellows with a high incidence in vineyards of Rastina (South Serbia). D'Ascenzo et al.
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. Agents: FD was first regarded as a physiological disorder. Diseased vines have a patchy distribution in the plot. However. introduced into Europe at the beginning of the 20th century. Transmission occurs also by vegetative propagation and grafting. Moreover. Several isolates have been characterized with molecular criteria. were characterized in Italy and shown to be transmitted also by S. Symptoms develop on leaves and shoots in June and July to become oustanding in August and autumn. show a general asymmetric bent posture and necrosis of terminal bud. isolated infected vines have been identified in northern vineyards of Burgundy and Alsace. Two isolates denoted FD-D and FD-C. Lignification of the canes is usually incomplete. In addition. Leaves persist longer on affected plants. FD88. littoralis Ball) (Homoptera. 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. Most of the time. symptoms affect the whole plant. Infected rootstocks are important means of dissemination because they are symptomless. titanus is well established. Molecular comparisons have shown that all these isolates are more closely related among them than with other phytoplasma strains in the same group. 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. After the discovery of other GY diseases. littoralis Ball). north of Spain and Portugal. FD88. It is widespread in all northern provinces of Italy and an outbreak was recently reported from southern Serbia. Vineyards in these areas are under the threat of an FD outbreak that could occur if FDinfected planting material is introduced. Affected vines of most varieties may recover in the second year when they are protected against reinoculation with insecticide treatments. extremely sensitive varieties do not recover. These isolates could be distinguished by Western-blot analysis using FD antisera and monoclonal antibodies (Mabs). Crop losses may be very high. Infected rootstocks show little or no symptoms. Isolates F70. This leafhopper is a neartic species specialized on Vitis sp. Cicadellidae). indicating a vine-to-vine transmission. the term "FD sensu stricto" was applied to diseases and phytoplasmas that are transmitted by S. titanus. that has colonized a wide climatic area. southern Italy. It is also present in regions from which FD has not been reported in France. decline progressively and die. then as a virus disease. FD92 and FD-D could not be distinguished from one another. In France. If the plants are inoculated after recovery. Though the rate of transmission by bench grafting can be very low.) using S. However. Transmission: The FD agent is transmitted in the persistent mode by the leafhopper Scaphoideus titanus Ball (= S. FLAVESCENCE DORÉE 1. with an incidence that increases rapidly from one year to the next. transmission is possible from early June (about one month after the beginning of hatching) until the death of adults. and western Slovenia. titanus. Switzerland. It is highly epidemic and extremely dangerous because of the biology and ethology of its leafhopper vector. 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. FD92 and FD2000 were experimentally transmitted to broadbean (Vicia faba L. 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. Scaphoideus titanus Ball (= S.GRAPEVINE YELLOWS: INDIVIDUAL DISEASES A. symptoms may be limited to a few shoots. Eggs are laid in summer on two-year old wood and trunk. It was described in a limited area in north-eastern Cataluña (Spain). titanus individuals collected in infected vineyards of south-western France. First symptoms may 149 . Description Flavescence dorée (FD) was the first Grapevine yellows (GY) disease to be reported. Corsica. it occurs in all southern vine-growing regions. FD is endemic only in regions where S. this way of spreading is significant when infected grafted plants are planted in vineyards hosting the vector. S. rooted cuttings from infected canes of a few rootstock varieties can develop "vinifera-like" symptoms on the wood and leaves. and Savoie.
Italy and Spain are susceptible with various degrees of sensitivity. Simultaneous amplification in a multiplex procedure of both FD9 and Stol11. Nevertheless. such as real-time PCR or identification of PCR-amplified product with DNA-DNA hybridization using specific probes. In 2003 a phytoplasma resembling the FD-C isolate was found in wild Clematis in Veneto (Italy). Control: FD is a quarantine organism in the EC. Material from mother plants that have developed symptoms in the following growing season. Indirect control is obtained by insecticide treatments against S. Grenache. a DNA fragment specific to BN (stolbur) phytoplasma can be used for the survey and monitoring of these two GY diseases. sensitive amplification with nested PCR of the DNA fragment FD9. The presence of S. Alicante Bouschet. such as Merlot. 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. 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. The second treatment. has become the preferred detection method because it is easy to perform and was improved in reliability and sensitivity. which is specific for 16SrV-group phytoplasmas allowed the reliable detection of FD phytoplasma in grapevine. aims at killing newly hatched insects. Sequence or RFLP analysis of the FD9 DNA fragment permit to readily differentiate all the known FD isolates. The low rate of transmission by grafting and the long delay in symptom expression make the detection of infected vines in planting material difficult. Soaking of dormant material in hot water (HW) is recommended. Detection with laboratory methods is possible on insect vectors and infected vines. control measures are compulsory according to legal regulations. Two additional treatments are applied during summer. must be destroyed. In spite of the possibility of recovery. Bois noir (BN) is also frequent in regions inhabited by this leafhopper. Varietal susceptibility and sensitivity: All varieties grown in the various FD-affected regions in France. at the beginning of July. has been used as the official method for large scale survey of FD in France from 1993 to 2003. Natural insecticides can be used to preventively limit vector populations but their efficiency is limited in epidemic outbreaks. Molecular detection by PCR of selected phytoplasma DNA fragments. Antigen extraction from vascular tissue of symptomatic grapevines requires the use of high Tris molarity and strong detergents in the extraction buffer. An indirect DAS-ELISA using coating of wells with rabbit polyclonal antisera and detection with a cocktail of several Mabs. Monitoring natural enemies of S. roguing of diseased vines is advisable when the epidemic pressure is high. has allowed the identification of potential auxiliaries for biological control. Other host plants: No host plants other than Vitis sp. Other DNA-based methods. Vitis riparia can be infected but shows little symptoms. titanus in an affected vineyard is an important indication that may lead to suspect a FD infection. Polyclonal antisera and Mabs have been raised to FD phytoplasma. 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. Cabernet Sauvignon. have been found carrying FD phytoplasma until recently. Chardonnay. Roguing is compulsory in France. 150 . Standard conditions (50°C for 45 min) proved highly efficient for sanitizing infected rootstocks and grapevine cultivars. at the beginning of August. Nielluccio and Garganega are very sensitive varieties that usually do not recover after infection. is directed against winged adults migrating from nearby vineyards or wild vines. Detection: FD symptoms can be confused with those of other GYs and also with other diseases or disorders. and the risk of disease spreading important. whereas the third treatment. are currently under development. titanus. In France and Italy.appear on young vines as late as four years after grafting. The same is true for American rootstocks which can therefore be dangerous sources of infection and dissemination of the disease. although heavily infected vines can be observed. The first treatment is applied 30 days after first instar emergence at the beginning of the period of potential transmission. Sauvignon blanc. Planting of HW treated material is especially important in areas where the vector is present. appear more tolerant. titanus in New York State (USA). Ugni blanc (Trebbiano) and Prosecco are sensitive varieties that may recover when they are protected from new inoculations. Recently. Symptoms are rare in Syrah. Their rearing is still under development. its area of origin.
Caudwell et al. Vidano: Study of the ecology and biology of S. littoralis from Ticino (southern Switzerland). Study of the role in the aetiology of FD of mycoplasmalike organisms (MLO) observed in insect vectors and in inoculated grapevine and V. Branas (a and b): The new disease is thought to be caused by root damage.: First report of S. littoralis. S. No positive results of transmission trials with 15 species of Hemiptera (mainly Cicadellidae) other than S. First report that abandoned or wild vines may be a source of infection. littoralis in its area of origin in North America. (b): Biology of S. Caudwell: Study of the phenomenon of recovery of FD-affected vines. Caudwell: Attempts to inhibit the agent of FD in vivo by heat treatment.: Successful transmission of FD to herbaceous hosts by increasing the feeding access period of the vector S. Baggiolini et al. Use of 4-5 treatments with DDT or parathion and removal or destruction of pruning wood older that two years. Schvester et al. littoralis. Caudwell et al. Schvester et al. S. Schvester et al. Spontaneous recovery occurs after a 1-2 year crisis period. Description and study of recovery phenomenon and of localised symptoms. littoralis Ball. littoralis. Caudwell: The disease is denoted Flavescence dorée (FD) in agreement with the name proposed by Levadoux (1955) because of the golden yellow metallic aspect of the leaves. Caudwell and Poitou: Study of the interaction between fanleaf and FD. 1960 1960 1961 1961 1962 1962 1963 1964 1964 1966 1966 1967 1968 1969 1970 1971 1971 1972 151 . Boubals and Caudwell: FD epidemics observed in Corsica. littoralis is present.2. (a). Schvester et al. The aetiology is unknown. littoralis on plants other than the grapevine and transmission trials of FD to other host plants. Historical review 1955 1956 1957 Levadoux: Description of an outbreak of "Flavescence dorée" on Baco 22A in southwestern France. (a): Control of FD by insecticide treatments against the vector. (a): Possibility to limit the populations of S. littoralis recorded from in Italy in 1963. The disease can be transmitted by grafting and has probably a viral aetiology. littoralis. littoralis in Corsican vineyards with winter treatment of dormant eggs with oleoparathion.: Demonstration that FD is transmitted by the leafhopper S. Caudwell et al. biology and feeding damage and of the symptoms of FD in France on Baco 22A. Caudwell: PhD thesis on FD.: Studies on the survival of S. considered as a virus disease. faba plants. Carle and Moutous: Study on possible secondary or alternative vectors of FD or Bois noir (BN).: Field tests for insecticide control of S. Description of macroscopic and microscopic symptoms. an insect recently introduced from North America. Bonfils and Schvester: Relationships of leafhoppers with FD of grapevine in western France. Recovered vines may be infected again but the symptoms are lighter. Description of the insect. 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. littoralis. evolution of the disease in space and time. Caudwell et al. Vidano: S.
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
according to the severity of the FD epidemics. (a and b): Demonstration of curing effect of hot water treatment on FD-affected plant material. Mori et al. titanus laid in the bark were killed. Belli et al.: RFLP studies of 16SrV phytoplasmas. Posenato et al. The amplified FD9 fragment is specific and variable among 16SrV (elm yellows) group phytoplasmas. Control recommendations.: Important outbreak of FD in Lombardia (northern Italy). especially Prosecco in the period 1993-1996. titanus reared on healthy plants. (a and b): Situation of FD and S. titanus in north-eastern Italy where vines are trained in the pergola system. First outbreak in the 1980's on cv Perera. The biological significance of this finding is unknown. Italy). Serological relationships with elm yellows phytoplasma is confirmed Alma et al. Three different RFLP patterns at least are shown in FD isolates from France.: Development of specific PCR primers for the detection of FD or BN phytoplasmas. Vindimian et al. Daire et al.: A phytoplasma of the 16SrI (aster yellows) group is detected in eggs. Survey of hemipters in the vineyard and presence of other known phytoplasma vector species. Caudwell et al. nymphs and adults of S. the eggs of S. Numerous species identified. Martini et al. Spain and Italy.: Verification of hot water treatment conditions to suppress FD phytoplasma infection in dormant wood of grapevine. Bianco et al. Infection is rapidly spreading to the east. titanus in Switzerland. Pavan et al. Transmission by grafting is studied. 1996 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1999 1999 2000 2000 155 . Rousseau: A review of different ways to reduce the populations of S. Marcone et al. Clerc et al. The possible role of six species in the transmission of GY is discussed. titanus in the canton of Geneva (Switzerland). Jermini and Baillod: Combination of visual observation of instar larvae and trapping of adults on sticky traps to monitor the populations of S. Kuszala: Specific ELISA procedures to detect FD or BN phytoplasmas in extracts from diseased grapevines from the field. FD has not been identified. titanus in Trentino (Italy).: Demonstration that two isolates of FD sensu stricto (FD-D and FD-C) are spreading in Veneto (Italy).: First report of S. In the same conditions. (a and b): Evolution of FD in the Treviso province (Veneto.: In spite of the presence and rapid spread of S. Batlle et al. Bosco et al. Several different peptides from membrane proteins are identified by Western-blot. Seddas et al.1996 1996 1996 1996 1996 Borgo: The first occurrence of FD sensu stricto in the province of Treviso (northern Italy) was reported in the 1980's. titanus in biological viticulture. The possibility that direct inoculations have occurred in nursery is discussed. Lherminier and Boudon-Padieu: Use of transmission electron microscopy with in situ immunolabeling to detect FD phytoplasma in grapevine and herbaceous host plants. FD is not present in southern Italy.: Monoclonal antibodies raised to immunoaffinity purified FD phytoplasmas.: Insecticide control of S. Increase and spread to other cultivars. titanus in Italy.: First report of FD outbreak in northern Cataluña (Spain).: Monitoring of leafhoppers in vineyards in Italy.
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
Credi and Callegari: Survey of vineyards in Emilia-Romagna. and different susceptibility and sensitivity of grapevine cultivars. Cazelles et al. Recovery and remission of symptoms for several consecutive years are frequent in vines affected by BN. Description of symptoms of VK. Mendgen: 1971. for the presence of a FD-like disease. Leclant and Lacote: Study of the ability to transmit stolbur to herbaceous plants by H. Leclant: Presence of Hyalesthes obsoletus a cixiid planthopper in the south of France. suppression of affected branches may improve harvest quality and help in progressive recovery. For direct differentiation between FD and BN/VK phytoplasmas.: Rickettsia-like organisms from VK-affected grapevines are cultured in chick embryos. roguing should be avoided. Description of structures that were probably closteroviruses. littoralis.: 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. (b): Epidemiological observations of BN lead to the assumption that an aerial vector is involved. unknown at that time. Borgo: General information on the presence of FD-type diseases in northern Italy. cytological and electron microscope study of tissues of affected grapevine. Chemical sprays against H. (b): BN is distinct from FD based on non transmissibility by S.: Rickettsia-like organisms found in roots of VK-affected grapevines in the Saar. obsoletus in the south of France. These findings have never been confirmed. Findings never confirmed. Though pruning does not cure infected vines. titanus is not always associated with the disease. Caudwell et al. S. except for declining vines.DNA fragments. Hence. Rumbos et al. Italy. The disease occurs in the Moselle valley and the Rhine valley. Rumbos et al. The species has been described as the vector of stolbur "virus" by Suchov in USSR in 1948.: GY symptoms are present in Switzerland (Suisse romande and Ticino) but they are not associated with S. 2. the multiplex nested-PCR assay cited in the FD chapter can be used. absence of recovery. 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. Nienhaus et al. BN is considered as a non epidemic form of FD. Superficial soil cultivation in summer or winter ploughing are methods intended to damage the larval instars and reduce vector populations. because of the complex biological cycle and multiple host plants of the insect species. no direct control is possible. Control: As with other GY diseases. 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. history. while others seem more tolerant and do not show symptoms. obsoletus are not efficient. Some grapevines show symptoms repeatedly in successive years. Gärtel: Description of VK under the name of Flavescence dorée. Caudwell et al. Natural enemies are being investigated. 1972 1977 1978 1978 1988 1989 1992 159 . The results suggest that the disease is brought into the vineyards from outside local sources. Historical review 1961 1965 1968 1969 1971 1971 Caudwell: Study on BN and its relationships to FD. titanus.
Alma et al. Borgo: Presence of BN in mixed infections with FD in the province of Treviso (Veneto. (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. PCR-RFLP analysis confirms relationship to aster yellows phytoplasmas.: PCR-RFLP analysis of MLO-DNA detected in VK-affected grapevines in Germany show that this MLO is related to stolbur. Italy) where FD is prevalent. Minucci et al. c and d) Transmission of a yellows to periwinkle with dodder in Germany. Computer analysis for spatial patterns of spread shows a non-random distribution and suggests preferential transmission from other vines or from weeds. clustered by the authors into aster yellows group but shown later to belong to the stolbur group. Boudon-Padieu (b): History. 1993 1993 1993 1993 1993 1993 1993 1994 1994 1994 1995 1995 1995 1996 1996 1996 1996 160 . It is suggested that vectors with preference for other host plants act as vectors for VK. Bianco et al. Davis et al. Bertaccini et al. Bertaccini et al. Biology of the insect. Maixner et al.: First detection of BN (stolbur phytoplasma) in Spain.: Grapevines from Piemonte (northern Italy) contain several different phytoplasmas. the most frequent belonging to group 16SrI.: A monoclonal antibody raised to stolbur mycoplasma-like organism is used for ELISA detection of the agent in herbaceous host plants and insects. 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. The latter MLO was shown later to belong to the stolbur group. Maixner (b. aetiology and transmission of BN in France. (a): Development of stolbur-specific primers on phytoplasma rDNA and use for detection of stolbur phytoplasma in grapevines affected with VK and in the vector H. Maixner : Demonstration that H. Bianco et al. but are different from known strains of MLOs of the aster yellows cluster. Daire et al. Maixner et al. MLO associated to GY in Italy are related to aster yellows MLO. H. BN MLO is detected in grapevines from France. Laviña et al. (a and b): PCR amplification of MLO DNA shows that MLOs associated with GY in several regions of northern Italy are related to the Italian periwinkle virescence (IPVR) MLO. Italy. The MLO strains found in grapevine are related with aster yellows MLOs. obsoletus. Brescia and Pavia (northern Italy). Reservoirs and possible role of other vectors remain to be elucidated. obsoletus is a vector of VK in Germany. from several regions of Italy. H.: The agent of a GY disease in the Italian Riviera is detected with a probe specific for Eastern aster yellows. Several weeds are host plants of the stolbur phytoplasma. subgroup I-G (later renamed as 16SrXII or stolbur group). obsoletus is confirmed as a vector of stolbur disease plants in France.1992 Fos et al. obsoletus identified as the vector. and from Israel. (b): Presence of phytoplasmas in the group 16SrI. (b): Evidence of an association of MLOs with grapevine yellows in EmiliaRomagna.: One of the two phytoplasmas detected in yellows-diseased grapevines in Liguria is related to IPVR phytoplasma. subgroup I-G (later named group 16SrXII (stolbur) in the provinces of Vicenza. Davis and Prince: According to a different terminology. (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).
Weber et al. Characterization of a phytoplasma belonging to group 16SrI. Estimated crop damage ranges from 20 to 40 %. Maixner: The situation of VK in Germany: symptoms. as well as phytoplasmas transmitted by Credi to periwinkle in Emila Romagna and previously classified in the aster yellows group. a high proportion of vines in which symptoms re-occurred after one season or more. a strong annual increase. Kuszala: Detection in ELISA using a monoclonal antibody to stolbur phytoplasma of BNassociated antigens in diseased grapevines in France.: Prevalence of BN (stolbur phytoplasma) in Navarra and Cataluña (north eastern Spain). Daire et al.: Consistent association of stolbur phytoplasma with GY diseases in the Italian provinces of Fiuli-Venezia Giulia and Trento and in Slovenia. (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. Refatti et al. Osler et al. Laviña et al. obsoletus as a vector of BN. conversely. The maximum delay of symptom expression in young plants is 2 years. Albanese et al. biology of H. First detection of BN agent in diseased grapevines in Switzerland with the same procedure. vector transmission and possible control measures. obsoletus and its role as the vector of VK. titanus. Davis et al. a high proportion of previously infected vines that did not show symptoms for 2 years at least and.: Monitoring of field populations of H. using hot water treatment with conditions recommended in France. Ploughing in winter of soils infested by C. obsoletus and possible control measures. Maixner and Reinert (b): Spatio-temporal analysis of the distribution of VK in 4 vineyards in the Moselle an Rhine valleys over a 2-3 year period showed a high incidence.: Important spread of GY symptoms on several grapevine varieties in Umbria (central Italy). Kölber et al. (a and b): BN is poorly transmitted by bench-grafting. subgroup I-G (identified later as belonging to the stolbur group or 16SrXII). Search for variability among BN phytoplasmas with RFLP analysis of the specific non ribosomal DNA was unsuccessful. Vindimian: LN but not FD is present in Trentino (northern Italy) in spite of the presence and active spread of S. Marcone et al. Maixner and Reinert (a): Trials on elimination of VK infection from vine wood in Germany. arvensis is recommended to expose instar larvae and kill them with frost. Sforza and Boudon-Padieu: Description of H. 1996 1996 1996 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1998 1998 1998 1998 161 . Preferential spread along the rows.: Occurrence of GY diseases in Hungary and identification of stolbur phytoplasma in affected grapevines. Results were satisfactory. obsoletus in Germany. search for vectors. Maixner and Reinert: A review of the research and present knowledge on VK and other GY diseases. (c): Identification of stolbur phytoplasma (16SrXII) as agents of GYs in Greece and Israel.: Phytoplasmas associated to GY in Campania (southern Italy) are genetically uniform and belong to the stolbur group. Sforza: PhD thesis on the epidemiology of BN in France. Weber: PhD thesis on the biology of the cixiid H.1996 1996 Koruza: Survey of GY in three regions of Slovenia shows that the disease is of BN type.
Up to 34% of insects were infected with stolbur phytoplasma. Acquisition may occur at larval stages since emerging 5th instars were infective. Evidence secured for the occurrence of tolerance (no symptom at all for 15 years) and temporary remission (no symptoms for a few years) .1998 1998 Sforza and Bourgoin: Anatomy of reproductory organs and mating of H. obsoletus in vineyards is efficient with sticky traps placed at the soil level. the ability to recover of cvs Riesling and Pinot in spite of systemic symptoms. Weber and Maixner (b): Etholology of H. No other grapevine phytoplasma was detected. High rate of BN / LN infection in the different vine-growing regions. No other phytoplasma found in the South of the country.: Only stolbur phytoplasma detected in grapevines showing GY symptoms.: BN reported only in north-western and eastern vineyards of Croatia.: In a large survey of vineyards in 8 Hungarian counties conducted in 1998.: VK disease dramatically increased in incidence in the Moselle and Rhine valleys during the 1990's. PCR detection is reliable when testing together 25 insects among which only one is infected. obsoletus. The same pathogen detected in weeds and other crops. The study confirms a high rate of infected H. among which hoary cress. Identification of two infected symptomless weeds in the vineyard. Bourquin et al. Identification of main reservoir weeds.: Use of PCR to confirm the presence of stolbur phytoplasma in GY-affected grapevines in Switzerland. obsoletus can be high because of the presence of reservoir weeds in the vineyard. Transmission to natural host plants is higher than for grapevine with both insects. Up to 30 % of the insects were infected and the highest rate was in vineyards with high populations of Convolvulus arvensis. Confirmation of the role of H. regardless of the cultivar in Campania (southern Italy).: First detection of stolbur phytoplasma in grapevines in Croatia. the second GY disease in Germany.: Epidemiology of BN in France. and a high proportion of vines that escaped infection during the 3-5 year period of monitoring. Braccini and Pavan: Report of survey of Auchenorrynchas in vineyards of Toscana (central Italy). Škoric et al.: Widespread presence of LN in Toscana (central Italy). Marcone et al. Maixner et al. Darimont and Maixner: Comparison of the transmission efficiency of phytoplasma to their host plants and to grapevine by H. obsoletus. Sforza et al. Sforza et al. obsoletus as a vector to grapevine. The rate of infected H. different phytoplasmas were identified in a number of cultivars. Šeruga et al. Females are more often infected than males. obsoletus and Metcalfa pruinosa were found positive for BN phytoplasma (16SrXII-A). H. Seljak and Petrovic: A review of the Slovenian situation of GY diseases. Weber and Maixner (a): Procedures for the survey of infection status of populations of H. Phytoplasmas in the 16SrXII group (stolbur) were the most frequent. obsoletus and Oncopsis alni. the vector of Palatinate grapevine yellows (PGY). First launching of colonies of the insect under controlled conditions. Varga et al. obsoletus and possibility to control the insect in vineyards.: Several vineyards affected by BN in Burgundy (France) monitored for over 15 years and assessed for symptom severity. Braccini et al. Larrue et al. formerly reported in western Europe. 1998 1998 1998 1999 1999 2000 2000 2000 2000 2000 2000 2000 2000 2000 2000 162 . Maixner and Reinert: Collection of H. obsoletus.: Ethological onservations and morphological descriptions of adults and larvae of H. obsoletus related to VK infection.
Gatineau et al. chlorophyll fluorescence and photosynthetic activities suggest that Photo System II is affected. which is widespread in the province. Kölber et al. All affected grapevines and two naturally infected periwinkles contained the same stolbur isolate.2001 Bertamini and Nedunchezhian: Evaluation of effects of BN disease (stolbur phytoplasma) in field-grown grapevines. is a natural vector of stolbur phytoplasma to sugarbeet in Bourgogne (France). Neoaliturus fenestratus. Samples were taken on all organs of 10 affected grapevines. on pigments. obsoletus.000 plants over 12 years. obsoletus in Lebanon is recorded. In spite of a similar abundance of H.: Survey of GY from 1997 to 2002 on 21 cultivars in 12 Hungarian counties. Choueiri et al. Mescalchin and Mattedi: A description of the development of VK in Trentino (northern Italy). Alma et al.: The situation of GY diseases in Croatia shows that only BN is present in the northern and eastern vineyards.: Survey of potential vectors of phytoplasmas in vineyards of the Golan Heights. Study of the spatial and temporal dispersion of the four species. Morandell: Detection of VK in south Austria. respectively. obsoletus and the presence in organic vineyards of several weeds known as host plants carrying larvae on the roots. the incidence of VK was significantly higher in conventional vineyards.: Only BN identified in Switzerland on cvs Chardonnay. general stress responses and rapid senescence. BN is the main disease and cvs Chardonnay and Zweigelt appear to be the more sensitive among whiteand red-berried varieties. obsoletus. obsoletus and Circulifer haematoceps were widespread and positive for stolbur or aster yellows phytoplasmas. 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. Detection in December was the more constant. H. Pentastiridius sp. very similar to European isolates. while Megophthalmus scabripennis was positive for aster yellows phytoplasma. Myrta et al. Cicotti et al. Same conclusions about tolerance and transient recovery of grapevines. The authors conclude that phytoplasma infection induces non specific. Maixner et al. Merlot and Pinot noir. Gugerli et al. obsoletus to the German viticulture. obsoletus is the vector of LN in Italy. Orenstein et al. 2001 2001 2001 2002 2002 2002 2003 2003 2003 2003 2003 2003 2003 2003 2003 2003 163 .: Observation of the occurrence of BN in Trentino in eight Chardonnay vineyards on more than 11. Bertaccini et al. Darimont and Maixner: Importance of the presence and infectivity of H. every month for one year except in winter. Langer et al. dioica) as a reservoir of stolbur phytoplasma and as a host for the insect.: Demonstration that H. The presence of H. Attempt to bring up the larvae to the surface of the soil by ploughing so that they are killed by frost in winter is under evaluation.: First detection in Albania of BN phytoplasma in grapevines of different age and varieties. Curkovic Perica et al.: Anoder cixiid planthopper.: Incidence of BN in a small area of the Bekaa valley (Lebanon) on cvs Chardonnay and Alicante Bouschet. Role of stinging nettle (U.: Assessment of the best period for detection of BN in affected grapevines. reached by other authors. Indicators based on climatic parameters permit to predict the flight period. Laviña et al.: The highest risk of phytoplasma contamination for grapevines coincides with the flight activity of adults of H.: Comparison of infection risk by VK in organic and conventional vineyards in Germany.
arvensis and U. Germany). It is associated to phytoplasmas in the Elm yellows (16SrV) group. Varietal susceptibility and sensitivity: PGY occurs in limited areas. Description Palatinate grapevine yellows (PGY) is a disease occurring in Germany in Pfalz and Moselle regions (west of Germany) mainly on cv Scheurebe. Agents: The agent is an EY (16SrV) group phytoplasma. together with high populations of H. -B and C) have been identified.: Important expression of BN in cv Chardonnay in the Ukraine. specific association of each isolate with a different weed is discussed. titanus leafhoppers have failed. Control: No possibility of control because of erratic transmission by the vector. It is different from FD phytoplasma. obsoletus were positive for stolbur phytoplasma. Gilge et al. was found carrying the alder phytoplasma at a higher rate than O. Main symptoms: Symptoms are similar to those of other GY diseases and may affect the whole plant. Main synonyms: FD-Pfalz was a first name because the agent was shown to be related to the FD phytoplasma.: Search for vectors of stolbur phytoplasma in Cataluña (Spain) where H. obsoletus is rare. (a): Investigation on the variability of stolbur (16SrXII) phytoplasmas from extracts of different grapevines in Croatia. occur most often at the border of plots. Trapping of insects on sticky traps or with D-Vac suction. It is not epidemic and occurs most often in old vines in vineyards along creeks or rivers where alders (Alnus glutinosa L) are present.2003 2003 Petrovic et al. alni but failed to transmit both to alder and grapevine. H. Symptoms were recorded on 26% of 30-year old Scheurebe grapevines and 96% of tested individuals of H. mainly on cv Scheurebe. and rarely in groups. Another alder insect. Alder is considered as the source of PGY infection. 2003 2003 2004 2004 2004 C. Milkus et al. Living insects checked for transmissibility of phytoplasma by feeding on artificial medium. According to the data of field survey. Langer and Maixner: Three different types of stolbur phytoplasma isolates found in grapevines. (b): First identification of BN infection in the Macedonian republic. O. dioica. Other host plants: Alder trees (Alnus glutinosa L. obsoletus and frequence of C.: Importance of a GY epidemics in the Drava region (Slovenia) in 2001 and 2002. Transmission to plants is not reported. the psyllid Psylla alni. Šeruga et al. Transmission: Transmission was obtained with the leafhopper Oncopsis alni Schrank (Hemiptera. The disease was identified in 1995. Three isolates (PGY-A. alni is highly monophagous and specimen do not survive long on grapevine. Šeruga et al. PGY phytoplasma is similar to the Alder yellows phytoplasma that is widespread in black alder in Germany. 164 . PALATINATE GRAPEVINE YELLOWS 1. Sabaté et al. obsoletus and weeds in different areas in Germany. Attempts to transmit PGY phytoplasma from grapevine to grapevine using reared S. Cicadellidae) trapped on alders and caged on V.) Detection: Universal primers and RFLP analysis of amplicons can be used to identify a 16SrV phytoplasma. The latter results have not been published.: Important epidemics of "Schwarzholzkrankheit" (SHK = VK) in Franken (Bayern. using 4 fragments of phytoplasma DNA showed that all isolates are similar. Several hemipters have been found to deliver stolbur phytoplasma to the medium. affected plants are not numerous. 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). vinifera seedlings that later developed GY symptoms. However. 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.
Same RFLP profiles of P1-P7 rDNA amplicons obtained from yellows affected grapevines. As much as 5 % of leafhoppers from Palatinate and 15 % from Moselle tested positive. alni and H. Variability analysis and sequencing of the FD9 DNA fragment and homology of the sequence to a part of the SecY gene. alder. Extensive survey of yellows-affected grapevines in several areas of Germany did not show the presence of other EY group phytoplasmas. Virginia grapevine yellows III (VGYIII) Main symptoms: North American Grapevine Yellows (NAGY) is a lethal disease causing yellowing of the leaves. (b): Transmission to V. a phytoplasma isolate transmitted from alder to periwinkle in Italy. a serious outbreak of GY occurred in Virginia. In the same period. alni) is able to transmit the alder yellows phytoplasma to healthy seedlings of alder.: Further comparison between elm yellows group (16SrV) phytoplasmas. the latter findings were not confirmed.2. 1997 1999 1999 2000 2000 2000 2001 2003 D. Darimont and Maixner: Comparison of the transmission efficiency of phytoplasma to their host plants and to grapevine by 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. It is different from FD phytoplasma but molecular and serological data show a close relationship. Main synonyms: Leaf curl and berry shrivel (LCBS). Historical review 1995 Maixner et al. which was associated with two different phytoplasmas in group 16SrIII and 16SrI (aster yellows). obsoletus. using Heteroduplex mobility assay and sequencing of rDNA and FD9 fragment. Cicadellidae) (but not P. The strong adaptation of O. alni. In 1993. In 1991. NORTH AMERICAN GRAPEVINE YELLOWS 1. In New York. The differences in vector specificity between PGY and FD phytoplasmas are not supported by molecular data. The proportion of infective leafhoppers is lower with O. vinifera seedlings of the PGY phytoplasma using specimen of O. Characterization of the three PGY strains showed that they were closer to FD than to strain EY1 from American elm. alni (Auchenorrhyncha. Reinert: PhD thesis on detection. affected vines often do not survive winter frost because they lack reserves. a phytoplasma belonging to the X-disease group (16SrIII) was detected in affected grapevines in New York.: Comparison between Elm yellows group (16SrV) phytoplasmas including PGY strains. alni to alder prevents a frequent inoculation to grapevine even if the infestation of the leafhopper population is high. both insect species trapped on alder. The risk of infection by PGY is lower than for VK because reservoir plants are fewer and outside vineyards. Infected grapevines often die within months from the onset of symptoms and significant losses of vines have been observed. 4 Italian and French strains of FD and elm and alder phytoplasmas. Identification of the PGY-associated phytoplasma with phytoplasmas infecting alder in the vicinity of affected vineyards. populations of S. Reinert and Maixner: 1997. Description Grapevine yellows (GY) were first reported in New York State (USA) in 1977. American grapevine yellows. alni trapped on alders. However. Maixner et al. Main diseases: Virginia grapevine yellows I (VGYI). The PGY strains are closest to the ALY phytoplasma from alder in Italy and to one strain of FD. Maixner and Reinert: Demonstration that O. Angelini et al. (b): Detection of a new EY-related phytoplasma in GY-affected cv Scheurebe in Germany. 165 . die back of shoot tips and abortion of developing fruit. alni. Reinert and Maixner: All the 3 strains of PGY were detected both in alder trees and in specimen of O. Oncopsis alni and Psylla alni specimen. the vector of BN / VK. The alder phytoplasma resembles ALY. respectively. molecular characterisation and epidemiology of PGY. Angelini et al.
Detection: With molecular assays using universal or group-specific PCR primers. X-disease phytoplasma was detected in native Vitis sp. However. titanus. Pearson et al.: Molecular comparisons between MLOs transmitted to periwinkle from grapevine or detected in a few naturally affected grapevines. subgroup I-A).: Occurrence of FD-like symptoms on White Riesling grapevines in New York. The latter observation suggests that another non-related MLO might have been present. and an aster yellows phytoplasma was detected in numerous weeds and shrubs within and around the vineyards. However. the disease was first observed on cv De Chaunac. Historical review 1977 Uyemoto et al. Detection was positive in symptomless wild Vitis riparia growing near vineyards in New York and in some symptomatic grapevines. it is very severe on Chardonnay vines but was observed on other varieties including Riesling. 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. No transmission by vine planting material reported. Larvae are more frequent on wild Vitis riparia at the border of the vineyards. Maixner and Pearson: First data on the study on S. observed in 1983. very similar to those of LCBS. as well as direct PCR assays from insects. Symptoms. In Virginia. Varietal susceptibility and sensitivity: In New York. and aster yellows was detected from numerous weeds and shrubs within and around the vineyard. The Virginia Grapevine Yellows (VGY) has been associated with the VGYIII phytoplasma (a strain of X-disease or 16SrIII. Daire et al.Agents: The 16SrIII phytoplasma detected in NY has not been further characterized. riparia to V. the survey of vineyards and transmission assays to V. Other host plants: Both phytoplasmas are widespread in the USA in weeds and cultivated species.: Monitoring of S. Transmission: No vector insects have been identified for VGY phytoplasmas. tested positive for a MLO in the X-disease group with PCR-RFLP of 16S rDNA. Symptoms may occur for several years in succession in the same vines. 1985 1991 1993 1993 1993 1993 166 . show that the FDU (= FDI) strain from Italy and a MLO detected in a grapevine in Virginia (USA) are both related to X-disease. Control: There are no control methods since vectors and reservoirs of the associated phytoplasmas are still undetermined. Maixner et al.: Two MLOs were transmitted from affected grapevines to periwinkles in Udine (Italy) (= FDI or FDU or GYU) and New York. faba bean plants and feeding solutions. (a): A Riesling plant from New York and a periwinkle carrying the FDI MLO described above. The FDI MLO was identified in a parallel work as a member of the X-disease group. then on White Riesling. subgroup III-B) and with the VGYI phytoplasma (a strain of aster yellows or 16SrI. Sauvignon blanc and Cabernet franc. X-disease phytoplasma was detected in native Vitis sp. other grapevines with strong symptoms tested negative. vinifera cuttings. ELISA and immunfluorescence were positive from periwinkle. Adults migrate from V. Probes and primers yielded positive reactions using DNA extracted from grapevine. 2. among which Agallia constricta. It appeared to be spreading and was perhaps insect borne. Chen et al. Prince et al. titanus in New York and transmission assays of a possible infectious agent. present in New York and its possible relationship with a GY disease. vinifera. allowed the identification of a few candidate vectors of the aster yellows phytoplasma. The X-disease phytoplasma occurring in New York was also detected with monoclonal antibodies and with probes for dot-blot hybridization. Data showed that the Italian and American MLO were related.: 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).
Restricted Growth (RG).1993 Wolf et al. The TBB phytoplasma has been designated as Candidatus species "Candidatus Phytoplasma australasia". It belongs to the peanut witches' broom group (16SrII) and has a broad plant host range in Australia. Description Grapevine yellows were first reported from Australia in 1976 and called Australian grapevine yellows (AGY) in 1983. The relatedness of the associated MLO to X-disease MLO is confirmed. In addition. The AGY phytoplasma has been designated as Candidatus species "Candidatus Phytoplasma australiense". is a member of the aster yellows (16SrI) group. with a higher incidence in warmer inland districts of New South Wales and South Australia. Buckland Valley Grapevine Yellows (BVGY). The second phytoplasma. one node after the other. Stems of affected shoots develop a blue. also detected in wild V. riparia vines. overlay one another like shingles and become leathery and brittle. Early and recent surveys have shown that AGY are found in most viticultural regions of Australia. a GY disease found in Victoria (Australia). Mapping of diseased plants show that grapevines appear to be a terminal host of AGY. Restricted Growth (RG)-affected grapevines show retarded growth resulting in shortened shoots and smaller leaves.: A GY disease observed in Virginia since 1983 affects large areas with a low incidence and a low spread rate. Other species tested positive and transmitted to feeding solutions. followed by the progressive death of the shoots. It belongs to the stolbur group (16SrXII) but is distinct from the BN and VK phytoplasmas. Agents : AGY disease (sensu stricto) is associated with two phytoplasmas: the AGY phytoplasma. 1998 2003 E. Both VGYI and VGYIII were found in several weeds and shrubs around and inside the vineyards. Davis et al. 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. consistently associated with AGY symptoms and AGY phytoplasmas but their aetiology is not clear. Shoots may display abortion of bunches or berry shrivel later in the season and death of terminal buds. The sequence similarity of the 16S rRNA gene shows that it is close to members of the aster yellows group (16SrI). BVGY phytoplasma has not been clustered in an established phytoplasma group. In this particular disease. Conversely. Late Season Leaf Curl (LSLC). subgroup I-A. waxy appearance and remain rubbery. Main diseases: AGY (sensu stricto). there are reported cases where they were not associated with AGY disease or phytoplasmas. The use of PCR has shown the association with AGY of three different phytoplasmas. suggesting that the source is from an alternative host. The 16S rRNA gene has a high sequence similarity (more than 99. Beanland and Wolf: Search for possible insect vectors and reservoirs of NAGY phytoplasmas in Virginia. two syndromes with particular symptoms (Restricted Growth and Late Season Leaf Curl) have been found in some cases. clover phyllody phytoplasma (16SrI-C) being the closest . AUSTRALIAN GRAPEVINE YELLOWS 1.: Two phytoplasmas are detected in yellows diseased grapevines in Virginia. The association of AGY with phytoplasmas was confirmed as early as 1988.5%) with the phytoplasmas associated to Papaya die back (PDB) in Australia and to Phormium yellow leaf (PYL) in New Zealand. Confirmation that a X-disease (16SrIII) group phytoplasma is involved and classification of the latter phytoplasma into a new subgroup. which is the more frequent and the Tomato big bud (TBB) phytoplasma. Main symptoms: All symptoms typical for GY can be found on AGY-diseased plants. The yellow leaf tissue can become necrotic. Late Season Leaf Curl (LSLC)-affected grapevines show symptoms in late summer with tightly downward rolled leaves that often remain green. It has been suggested that all three phytoplasmas are strains of the same phytoplasma species. 167 . III-I. leaves tend to fall early. respectively. Some plants also display uneven bud development resulting in canes and cordons that are bare in places with little or no bunch development. Agallia constricta is a suspected vector of VGYI (aster yellows phytoplasma).
140-510 nm in diameter. Remission of AGY. argentatus and subsequently transmitted to faba bean but the ability to transmit to grapevine was not confirmed. Magarey et al. suggesting that this contributes to the decline in phytoplasma numbers and to the remission of symptoms. RG and LSLC diseases has been observed. vector of TBB in tomato crops. followed by RFLP analysis or sequencing. Similarly. suggesting that the disease is the result of an aerial transmission and that the phytoplasma was not present in the original planting material. However. Hence. The TBB phytoplasma is transmitted to other crops by O. Sanitation with hot water treatment is being developed in Australia. Varietal susceptibility and sensitivity: Chardonnay and Riesling appear to be most often affected. The BVGY phytoplasma has no suspected insect vector. Detection: Detection is obtained with PCR. indicating persistence of the disease. The planthopper Oliarus atkinsoni Myers (Hemiptera. Several observations suggest that aerial transmission prevails.: MLOs. Reduction of symptom expression in vines injected with tetracycline. once vines are infected by AGY. branches. Detection can be from shoots. 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. It is possible that AGY diseases have become prevalent in Australia in the last years with the increasing planting of Chardonnay. All generic "universal" primers may be used. Phytoplasma australiense) either in single-step or nested PCR following a first amplification with universal primers. A "heat curing" effect of AGY disease by hot weather has been observed. Magarey and Wachtel: GY symptoms are found in most viticultural areas of Australia. Phytoplasma australasia). they are at greater risk of displaying symptoms again in the following years. Recent studies have shown that the latter phytoplasma can be acquired from infected grapevine by O. but phytoplasmas were also detected in other red. reservoirs must be important but the epidemiology is not fully understood. trunks and roots throughout the year and infection is persistent from year to year.and white-berried varieties. Sampling simultaneously from shoots. Control: There are no methods to control AGY diseases in the vineyard. Magarey and Wachtel: Review of the AGY problem. Magarey: The situation of GY in Australia. found in sieve tubes of diseased vines are associated with AGY symptoms. branches. Magarey: Comprehensive review of GY diseases in the world.: The "Rhine Riesling problem" denoted "Australian Grapevine Yellows". Magarey et al. Osmelak et al.Transmission: No vectors have been identified for any AGY disease. 2. RG or LSLC. AGY phytoplasmas are common in several crops in Australia. is found in high numbers together with two other potential MLO vectors in vineyards of Victoria (Australia). such as Shiraz or Semillon. Other host plants: As mentioned above. argentatus. and trunks in October is more reliable. The incidence of disease may be temporarily high in some vineyards of Chardonnay. However. 168 . Phloem fluorescence of diseased vines in the UV microscope. a specific primer in the intergenic 16S-23S spacer region has been designed for detection of TBB phytoplasma (Ca. The AGY phytoplasma was detected in the common leafhopper Orosius argentatus (Evans) but transmission to grapevine has not been reported.: Orosius argentatus. Specific primers have been designed on the 16S rRNA gene for detection of AGY phytoplasma (Ca. Cixiidae) is known as the vector of PYL disease in New Zealand but the species has not been reported from Australia. 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.
: Description of the AGY phytoplasma and designation of a new taxon.: AGY phytoplasma detected in the body of Orosius argentatus leafhoppers. Constable et al.: Use of hot water treatment in commercial nurseries of Australia.: Assumption from field observations and monitoring that AGY. RG and LSLC are associated. Davis et al. Gibb et al. Wilson et al. later called BVGY phytoplasma. Hence. (a and b): Positive detection of phytoplasma with PCR in AGY-affected grapevines and tentative survey of phytoplasmas in Australia.: Identification of a new phytoplasma in Chardonnay grapevines in Victoria (Australia). Padovan et al. It is occasionnally detected in symptomless vines and in vines with LSLC or RG symptoms. Spring and summer are optimal for detection. Bonfiglioli et al. Constable et al.: Comparison of visual assessment and diagnostic assays. (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). BVGY might form a new AY subgroup of AY (16Sr) group or even a new phytoplasma group. RG. Waite et al.: Study of the phylogenetic relationships of the BVGY phytoplasma with aster yellows and stolbur phytoplasmas. Constable et al. The problem of symptomlessly infected vines. established from a different source of plant material indicate that BVGY disease occurs as a result of aerial transmission. Habili et al.: TBB phytoplasmas can be acquired from diseased grapevine by Orosius argentatus and subsequently transmitted to faba bean. Candidatus Phytoplasma australiense. Papaya dieback and Phormium yellow leaf diseases. The TBB phytoplasma is detected occasionnally in vines with AGY symptoms and also in LSLC-affected vines. Spain and Israel.: BVGY phytoplasma is associated to GY symptoms that may show remission then reoccurrence but also appear on vines that were not affected previously. Uneven distribution of phytoplasmas in infected plants. Padovan et al. Liefting et al.: No vector for AGY found.: The AGY phytoplasma is consistently present in the majority of symptomatic grapevines. Kelly et al. Constable et al.: Description of AGY symptoms. resulting in increasing cumulative incidence. Beanland et al. Detection of BVGY in another plot in the same area. Constable: PhD thesis on the biology and epidemiology of AGY phytoplasmas.: Detection of phytoplasmas associated with AGY.1995 1995 1996 Bonfiglioli et al. and LSLC. Constable and Symons: AGY phytoplasma is detected in most symptomatic grapevines but also TBB phytoplasma and a third uncharacterised phytoplasma.: Close relationships between phytoplasmas associated with AGY. 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. Beanland et al. LSLC would be followed by AGY on the following year and AGY would be followed by RG in subsequent years. insecticide sprays are useless for controlling the disease.: Further comparison of AGY phytoplasma with overseas GY phytoplasmas show that it is closely related to.
The subgroup of aster yellows was not determined but primers specific for stolbur tested negative. Constable et al. Transmission: Aster yellows and. In Chardonnay. are very frequent in many host plant species and several insect species are known as vectors or potential vectors in different countries. TBB phytoplasma may have a role in the aetiology of LSLC. they seem to have some relevance in Israel and were recently reported from Tunisia. a variety widely grown in all countries and very sensitive to all GY agents.: Survey of AGY. sometimes as a second phytoplasma in co-infection with FD or BN phytoplasmas and sometimes also in symptomless grapevines. The first is the detection in countries other than the USA. There is no information of transmission of AGY by planting material. Constable et al. to a lesser extent. Description Besides the cases reported above. namely Israel. 1997 170 . Historical review Europe 1996 Alma et al. The second situation is the significant occurrence of X-disease group (16SrIII) phytoplasmas in countries other than the USA.: Detection of phytoplasma in the aster yellows group in GY affected plants in Slovenia and Croatia. 2004 2004 F. later designated 16SrXII. X-disease group phytoplasmas. However.: Survey of phytoplasmas infecting grapevine in northern Italy. The third situation is represented by poorly characterized GYs recorded from new countries. OTHER GRAPEVINE YELLOWS 1. 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.2002 2003 Crocker et al. Statistical analysis showed that RG and LSLC diseases were not always associated with AGY. (a and b): Biology and epidemiology of AGYs. 2.: The management of the area for plant material in the nursery with the scope of hot water treatment. Other varieties are cited in the different situations. However. The total cumulative incidence for AGY could be as high as 90%. the diseases are characterized by remission in some plants. detection is more reliable from samples taken from branches and trunks. Saric et al. In three instances. 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. They must not be confused with the 16SrI-G phytoplasma of earlier reports. No variability was observed amongst BVGY phytoplasma isolates. a 16SrI-B phytoplasma was found in mixed infection with a stolbur phytoplasma (at that time designated 16SrI-G). recurrence in other vines and new occurrence in previously unaffected vines. It seems that aster yellows phytoplasmas have no important significance in European grapevine yellows. Symptomatic shoots from AGY affected vines are reliable for detection in summer (January). At other times of the year. These aster yellows phytoplasmas belong to subgroups I-B or I-C (clover phyllody) which are ubiquitous in Europe. which was the first designation for stolbur phytoplasma in some laboratories. RG and LSLC symptoms for 3 to 6 years in Chardonnay and Shiraz vineyards. is often reported in connection with these cases. of phytoplasmas belonging to the aster yellows group (16SrI) in yellows-affected grapevines. The AGY and the TBB phytoplasmas may persistently affect grapevines throughout the year and between years. three different situations must be described. Varietal susceptibility and sensitivity: Chardonnay. Constable and Symons: Study with Heteroduplex mobility assay (HMA) of the elongation factor Tu (Tuf) gene of the AGY and the BVGY phytoplasmas.
obsoletus was found in addition to the preceding 5 species. Merlot and Carmenere. subgroups I-B and I-C and also to group 16SrVIII (Ash yellows) in mixed infection with I-B. 1971b).: Survey over several years of potential vectors of phytoplasmas in vineyards of the Golan Heights showed that Neoaliturus fenestratus. H. and Anaceratagalia laevis were all found carrying an aster yellows phytoplasma. Orosius orientalis. H.: Monitoring of potential phytoplasma vectors in vineyards in Israel and molecular detection of phytoplasmas in their body. titanus had been obtained in 1971 in France (cf. Klein et al. 2001 2003 Tunisia 2003 Chabbouh et al. including S. Gajardo et al. Stolbur phytoplasma (16SrXII) was also detected.: First results on the monitoring of GY diseases in Tunisia in imported table grape varieties. obsoletus and Circulifer haematoceps were all three abundant and positive for stolbur and aster yellows phytoplasmas. Detection with PCR show erratic identification of a 16SrI-B phytoplasma.2001 Alma et al. Macrosteles sexnotatus. called "Amarilliamento de Elqui". In addition to similarities of symptoms with FD. The spatial and temporal dispersion of the four species was analysed. In addition. M'hirsi et al. Megophthalmus scabripennis was positive for aster yellows phytoplasma.: Identification in yellows-affected vines of varieties Petit Syrah. Tanne et al. Transmission to periwinkle of yellows diseases using collected insects. Similar transmission of clover phyllody (16SrI-C) MLO by S. of phytoplasmas belonging to group 16SrI.: Further survey of GY vectors in the Golan Heights. Orenstein et al. Caudwell et al. Neoaliturus sp. Trapping of hemipters and identification of numerous potential phytoplasma vectors. some affected vines showed a sudden fall of leaves in summer. titanus. 2003 171 . D'Ascenzo et al.: Formal identification of a 16SrI-B phytoplasma in GY-affected grapevines in Tunisia. a X-disease phytoplasma was detected in some Neoaliturus specimen.: 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). Circulifer spp. 2003 South America 1980 Caudwell: Description of a disease of the yellows type in Chile..: Experimental transmission of the Chrysanthemum yellows phytoplasma (16SrI-B) to grapevine by four leafhopper species. 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.
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