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Options Méditerranéennes Série B n.55 Directory of Infectious Diseases of Grapevines and Viroses and Virus-like Diseases of the Grapevine: Bibliographic Report 1998-2004
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Les opinions, les données et les faits exposés dans ce numéro sont sous la responsabilité des auteurs et n'engagent ni le CIHEAM, ni les pays membres.
P. E. Martelli.CIHEAM Centre International de Hautes Etudes Agronomiques Méditerranéennes Options méditerranéennes Directeur de la publication: Bertrand Hervieu Directory of Infectious Diseases of Grapevines and Viroses and Virus-like Diseases of the Grapevine: Bibliographic Report 1998-2004 Editors: G. Boudon-Padieu 2006 .
279 Options Méditerranéennes.This volume of Options Méditerranéennes Series B has been formatted and paged up by the IAM Bari Editing Board La maquette et la mise en page de ce volume de Options Méditerranéennes Series B ont été réalisées à l'Atelier d'Edition de l'IAM Bari Copy number : 500 Printed by IDEAPRINT -Bari.P. +39 080 542 45 87 e-mail: ideaprint@virgilio. Série B : N. 55 ISSN : 1016-1228 ISBN : 2-85352-338-1 © CIHEAM. Italy. E. Boudon-Padieu Bari : CIHEAM (Centre International de Hautes Etudes Agronomiques Méditerranéennes) p.it Directory of Infectious Diseases of Grapevines and Viroses and Virus-like Diseases of the Grapevine: Bibliographic Report 1998-2004 Editors: G.September 2006 tel. Martelli. 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» .
TABLE OF CONTENTENTS PREFACE INTRODUCTION INFECTIOUS AGENTS OF GRAPEVINES INFECTIOUS DEGENERATION (GRAPEVINE FANLEAF VIRUS) INFECTIOUS DEGENERATION (EUROPEAN AND MEDITERRANEAN NEPOVIRUSES) GRAPEVINE DEGENERATION AND DECLINE (AMERICAN NEPOVIRUSES) GRAPEVINE LEAFROLL RUGOSE WOOD COMPLEX GRAFT INCOMPATIBILITY FLECK COMPLEX MINOR VIRUS DISEASES VIRUS-LIKE DISEASES VIROIDS GRAPEVINE YELLOWS: GENERAL PROPERTIES GRAPEVINE YELLOWS: INDIVIDUAL DISEASES GRAPEVINE YELLOWS: REFERENCES PIERCE'S DISEASE THE VIROSES AND VIRUS-LIKE DISEASES OF THE GRAPEVINE: BIBLIOGRAPHIC REPORT .1998-2004 INTRODUCTION SUBJECT INDEX REFERENCES 7 11 15 17 33 49 59 76 93 99 107 117 127 133 149 173 195 203 205 207 219 5 .
Furthemore. being a recognized authority in this field. produced under the auspices of ICVG. except when virus-free vines are grown for comparison alongside with non-sanitized sister vines. whose Secretatiat he has admirably conducted since its very establishment in 1962. The “Bibliographic Report” is another most precious achievement by R. 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. Martelli and E. 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.P. Sincere thanks are espressed to the authors of these contributions. and practitioners. in the belief of rendering a good service to the scientific community and. grouping diseases and setting them in a well thought out order. so as to produce a document which can readily be used by scientits. and to the quality and quantity of the crop. ICVG has a long-lasting tradition in these endeavours. The present issue of Options Méditerrnéennes. or induce subtle debilitating effects which are difficult to percieve. whereas E. up to date as to the time of publication. these diseases cause loss of vigour and often a decline of affected stocks. promoting the publication of the Bibliografic Report 1985-1997 on virus and virus-like diseases of the grapevine. One way of achieving this goal is to produce and make available updated accounts of the state-of-theart of grapevine virology and comprehensive bibliographic reports. Boudon-Padieu has a long lasting experience in the study of phytoplasmas. Notwithstanding these efforts and the promoting activity of the International Council for the Study of Virus and Virus-like Diseases of the Grapevine (ICVG). G. a distinguished virologist of worldwide reputation and one of the father founders of ICVG. students.PREFACE Virus. although the technology is now available for detection and elimination of most of the pathogens occurring in propagation material. hosts a new edition of the “Directory of Infectious Diseases of Grapevines” by G. 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. Special care was taken in the organization of the subject matter. that fosters coperative studies in grapevine virology. Bovey The “Directory” is a work of great thoroughness and accuracy. virus-like and phytoplasma diseases of grapevines constitute a major limiting factor to the development and well-being of the world viticultural industry. is therefore a goal of utmost importance to which the Mediterranean Agronomic Institute of Bari (IAM-B). they reduce graft compatibility of scions and rootstocks. by and large. it is most unfortunate that dissemination of infectious diseases continues. to which IAM-B was happy to contribute.P. Cosimo Lacirignola Director of IAM Bari. Improving the sanitary conditions of the industry. As a whole. to all those interested in viticultural matters. Boudon-Padieu and the latest Bibliographic Report (1998-2004) on “The Viroses and Virus-like Diseases of the Grapevine” by R. As to the authors. which reflect on the commercial value and productive life of the vineyards. Bovey. technicians. taking also into account the future role of the Mediterranean as a free-trade area. whose publication IAM-B is happy to support. Italy 7 . is contributing with direct involvement in international research projects and through the activity the Mediterranean Network on Certification of Grapevines.
France 2006 . Martelli University of Bari. Boudon-Padieu INRA Dijon.P. Italy E.International Council for the Study of Virus and Virus-like Diseases of the Grapevine (ICVG) DIRECTORY OF INFECTIOUS DISEASES OF GRAPEVINES G.
A bibliographic report 1971-1978. 76 pp. Gugerli.INTRODUCTION The grapevine (Vitis spp. Vitis 25.P. ICVG has met regularly every 3 to 4 years. 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. 1986. shortening the productive life of vineyards. 2003. among other things. To this effect. 303-324. Office International de la Vigne et du Vin. has fostered intensive research. The viroses and virus-like diseases of the grapevine.. As most of the vegetatively propagated crops. 316-376. and endangering the survival of affected vines. Options Méditerranéenes. Bibliographie de 1965-1970. attracting the attention of scientists. The viroses and virus-like diseases of the grapevine. Vitis 18. Martelli. in turn.. Bovey R. 205-279. Extended Abstracts 14th Meeting of ICVG. Bovey R.. Thus. as well as on the quality and quantity of the yield. recognizes that a number of the 60 or so infectious agents (viruses. 227-275. viroids. From the very beginning. A short history of ICVG. 3rd part). 1999.B. The viroses and virus-like diseases of the grapevine. ICVG has been instrumental in fostering basic and applied research in grapevine virology. Paris. 2006. and phytoplasmas) recorded from the grapevine can be highly detrimental to this crop. Bovey R.B. Options Méditerranéenes. Caudwell A. 2003). and administrators on the detrimental effects of infectious diseases on the well-being of the industry. grapevines are exposed to the attacks of a variety of pests and pathogens among which infectious intracellular agents (viruses. Vitis 11. W. 29 (Series B. Bovey. Bibliographie des viroses de la vigne des origines à 1965. causing heavy losses. and P. and R.. phloem-and xylemlimited prokaryotes) play a major role. Bovey R. 1972. having a negative impact on the plant vigour and longevity. viroids. all efforts should be made to improve its sanitary conditions. and G. nurserymen. The importance of the grapevine industry and the magnitude of the problems caused by these pathogens has generated wide interest which. and a highly valuable agricultural commodity. The increased attention paid to grapevine virological problems and the like. and supporting initiatives for the establishment and implementation of clean stock and certification programmes. 1979.) undoubtedly represents one of the horticultural crops most widely grown in temperate climates. that has been especially active at the international scale from the late 1950's onwards. A bibliographic report 1998-2004. 11 . Infected propagating material is largely responsible for the spread of diseases among and within viticultural countries. The viroses and virus-like diseases of the grapevine. A bibliographic report 1985-1997. Bovey. growers. its 14th Conference having taken place in Italy in 2003 (Bovey and Gugerli.. Locorotondo 2003. Since its foundation. ICVG has issued the recommendations that follow: The International Council for the Study of Virus and Virus-like Diseases of the Grapevine (ICVG). Les virus de la vigne. which were compiled under the auspices of the International Council for the Study of Viruses and Virus Diseases of the Grapevine (ICVG).400. has produced an impressive series of papers which now number about 5. The papers up to 2004 are listed and commented upon in six bibliographic reports: Caudwell A. Hewitt W. 1965. A bibliographic report 1979-1984. 10-172. 3rd part). Hewitt and R. 1-2. xx (Series B.
1988. and permit tracing the certified grapevines to the originally selected and tested plants. They represent a most valuable source of information. W. The agents that should be controlled by the Class 1 certification program are those associated with infectious degeneration and grapevine decline (nepoviruses). Martelli. Uyemoto. 1980.K. 1978. high standards are essential to ensure that no viticultural area is compromised by the introduction and spread of diseases. Martelli and A. having a negative impact on plant vigour and longevity. Improvement of the sanitary level can be achieved through selection and sanitation.K. valuable grape genetic resources exist which are infected with virus but are essential to the preservation of world viticultural heritage. Editions Payot. Gubler and J. Martelli and A. Bovey R.D. Uyemoto will be published in 2005). Italy) The Proceedings of all the international ICVG Conferences have been published in extenso or. Rowhani will be published in 2005 in the series APS Plant Virus Image CD-ROM. As efforts are made to harmonize grapevine certification protocols.The presence of diseases such as infectious degeneration.A. leafroll. as Extended Abstracts. Woodham. Set 1 (O. It would move freely between regulatory boundaries. 29 pp. Compendium of Grape Diseases. Lausanne. is regarded as incompatible with an accepted sanitary status. Pearson R. (a revised edition by J.P. The regional certification standards for Class 2 stock should be created at a local level based on the rate of endemic infection.P. many of which can be highly detrimental to this crop.C. (a 2nd revised edition edited by W.C. viroids and phytoplasmas). G. R. W. and other grapevine yellows). a moratorium will be established for these viruses. Goheen. G.B. which are best performed in the framework of certification programmes encompassing also clonal selection.C. A. 181 pp. Paul. and fleck. The certification process should specify conditions to prevent and detect subsequent infection of nursery plants by regulated pests. ensure clonal integrity. Gärtel. recognises over 70 infectious agents affecting grapevine (viruses.. Vuittenez. Minnesota. In order to preserve valuable grape clones and varieties. No other stock should move outside regulatory regions. Wilcox. Virus and Virus-like Diseases of Grapevines. all efforts should be made to improve its sanitary conditions. Inadequate certification standards have repeatedly resulted in disease problems for growers and nurserymen. 93 pp. Hewitt. In addition. Class 1 should include only grape nursery stock which tests negative for the most damaging diseases/pathogens. rugose wood. as well as on the quality and quantity of the yield. technology should make it possible to exclude additional disease-causing viruses from the certified stock. Certified grapevines are derived from pathogen tested. 100 slides.P. Locorotondo. Portugal ) The International Council for the Study of Virus and Virus-like Diseases of the Grapevine (ICVG).K. USA. Grapevine (Vitis) virus and virus-like diseases. GVB and GVD). Until that time. The American Phytopathological Society Press. W. Certified selections should be tested for specific pathogens. Their elimination from mother vines intended for propagation should therefore be pursued. Barnett and S. 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. and 3). bois noir. leafroll disease and associated closteroviruses (grapevine leafroll associated viruses 1. Clemson. G. clonally selected primary sources. under the name of Grapevine Viruses. and the need for preservation of heritage germplasm. including the causal agents of fleck and rupestris stem pitting. regional viticultural conditions. Certification of grapevine nursery stock is a powerful and effective tool to control these agents. (issued and approved in 2003 at the 14th ICVG Meeting. Thus. that enables vineyards to economically and sustainably maintain quality and productivity.F. Virus-like Diseases and Other Disorders). and A. Plant Virus Slide Series. the virological problems of grapevines have been extensively treated and illustrated in a number of books and major review articles: Uyemoto J. However. In the future. rugose wood (GVA. we propose two sanitary classes.W. Lisbon. 2. St.G. and phytoplasmas (flavescence dorée. Infected propagation material is largely responsible for the spread of diseases among and within viticultural countries. Tolin. lately.F. eds) Clemson University. 12 . (issued and approved in 1997 at the 12th ICVG Meeting. Dias. Grape Section. Goheen and H.
Bulletin de l'OIV 70. (ed. Khetarpal. Koganezawa. 2000.. Krake L. 945-971. Sélection sanitaire de la vigne: sélection sanitaire et sélection pomologique. 1996. France 13 . Boudon-Padieu and M. C.K. and B. Sanitary selection of the grapevine. Lacirignola. Ridé. Virus certification of grapevines. and R. Martelli. Martelli and E. the Steering Committee of ICGV decided to update the Directory and to entrust this task to G. Martelli G. INRA Editions. Influence des viroses et qualité. Director of the Mediterrenean Agronomic Institute of Bari for supporting the publication of this work. Editions Féret. 54 pp. Notwithstanding this wealth of published information a "Directory of Major Virus and Virus-like Diseases of Grapevines" was compiled in 1992 by R. 1991. 5-23. Protocols for detection of viruses and virus-like diseases: Les Colloques. Bovey and G. St. Chairman of the Board of Directors and Secretary General. Bulletin de l'OIV 69. and G. E. Martelli and published under the auspices of the Mediterranean Fruit Crop Improvement Council (MFCIC).P. Maladies à Virus. 1993. Rome. Walter B.P. Paul. n° 86. Giovanni P.P.). They express their deep gratitude to Prof. N. FAO/IBPGR Technical Guidelines for the Safe Movement of Grapevine Germplasm. 192 pp. and G. eds.). University of Bari. a body now estinguished. Italy. Scott. Martelli G. Hervieu. Paris.H. Walter B.Frison E. Bordeaux.Université de Bourgogne Dijon. 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. Taylor. Rome /International Board for Plant Genetic Resources. CSIRO Publishing. Walter B.137 pp. Rezaian and R. At the 14th ICVG Meeting. Boudon-Padieu. Walter. FAO Publication Division.P. Hadidi. of the Centre International des Hautes Etudes Agronomiques Méditerranéennes. Sélection sanitaire de la vigne: sélection sanitaire et sélection pomologique. and to Dr. 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. (ed). 1999. Hamze and Mr. Collingwood. H.S. B. 263 pp. Paris. 225 pp. Ikin. Elisabeth Boudon-Padieu Directeur de Recherches Head Biologie et Ecologie des Phytoplasmes Plante Microbe Environnement INRA . Detection and Diagnosis of Graft-transmissible Diseases of Grapevines. 1997.CNRS . Graft-transmissible Diseases of Grapevines.A. American Phytopathological Society Press. Bari.R. respectively. 261-276. 1998. Walter B. Influence des viroses et qualité. 1997. Martelli Professor of Plant Virology. R. Food and Agriculture Organization of the United Nations.P. In: Plant Virus Disease Control (A. M. Martelli. 1ere partie: Effets des viroses sur la culture de la vigne et ses produits. bactéries et phytoplasmes de la Vigne. sélection pomologique. 2ere partie: Sélection sanitaire..A. Rome. M.P.
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. The viral scenario of Vitis: viruses and their taxonomic affiliation(a) FAMILY GENUS SPECIES A. phytoplasmas (8). viroids (5). and insecttransmitted xylematic bacteria (1) have been recorded form grapevines. Viruses belonging to genera included into families BROMOVIRIDAE Alfamovirus Cucumovirus Ilarvirus Alfalfa mosaic virus (AMV) Cucumber mosaic virus (CMV) Grapevine line pattern virus (GLPV) Grapevine angular mosaic virus (GAMoV) BUNYAVIRIDAE CLOSTEROVIRIDAE Tospovirus Closterovirus Ampelovirus Tomato spotted wilt virus (TSWV) Grapevine leafroll-associated virus 2 (GLRaV-2) Grapevine leafroll-associated virus 1 (GLRaV-1) Grapevine leafroll-associated virus 3 (GLRaV-3) Grapevine leafroll-associated virus 4 (GLRaV-4) Grapevine leafroll-associated virus 5 (GLRaV-5) Grapevine leafroll-associated virus 6 (GLRaV-6) Grapevine leafroll-associated virus 7 (GLRaV-7) Grapevine leafroll-associated virus 8 (GLRaV-8) Grapevine leafroll-associated virus 9 (GLRaV-9) Two new putative ampeloviruses are being characterized in France and the USA COMOVIRIDAE Fabavirus Nepovirus Broadbean wilt virus (BBWV) Artichoke italian latent virus (AILV) Arabis mosaic virus (ArMV) Blueberry leaf mottle virus (BBLMV) Cherry leafroll virus (CLRV) Grapevine Bulgarian latent virus (GBLV) Grapevine Anatolian ringspot virus (GARSV) Grapevine deformation virus (GDefV) Grapevine Grapevine fanleaf virus (GFLV) Grapevine Tunisian ringspot virus (GTRV) Peach rosette mosaic virus (PRMV) Raspberry ringspot virus (RpRV) Tobacco ringspot virus (TRSV) Tomato ringspot virus (ToRSV) Tomato blackring virus (TBRV) FLEXIVIRIDAE Foveavirus Grapevine rupestris stem pitting-associated virus (GRSPaV) chrome mosaic virus (GCMV) 15 .
Desselberger. The names of tentative species are written in Roman characters. U.. 2005. 8Ith Report of the International Committee on Taxonomy of Viruses. C. Mayo.. and Ball. pp. Maniloff. (a) 16 .M. Taxonomically unassigned viruses Unnamed filamentous Grapevine Ajinashika virus (GAgV) Grapevine stunt virus (GSV) Grapevine labile rod-shaped virus (GLRSV) Raspberry bushy dwarf virus (RBDV) Strawberry latent ringspot virus (SLRSV) Sowbane mosaic virus (SoMV) Tobacco mosaic virus (TMV) Tomato mosaic virus (ToMV) Scientific names of definitive virus species are written in italics.A.A.. J. L. 1258. London.Potexvirus (GINV) Potato virus X (PVX) Trichovirus Vitivirus Grapevine berry inner necrosis virus Grapevine virus A (GVA) Grapevine virus B (GVB) Grapevine virus C (GVC) Grapevine virus D (GVD) Carnation mottle virus (CarMV) Tobacco necrosis virus D (TNV-D) Grapevine Algerian latent virus (GALV) Petunia asteroid mosaic virus (PAMV) Grapevine asteroid mosaic-associated virus (GAMaV) Grapevine redglobe virus (GRGV) Grapevine fleck virus (GFkV) Grapevine rupestris vein feathering virrus (GRVFV) TOMBUSVIRIDAE Carmovirus Necrovirus Tombusvirus Marafivirus Maculavirus TYMOVIRIDAE A new putative marafivirus is being characterized in the USA POTYVIRIDAE Potyvirus(?) Unidentified potyvirus-like virus isolated in Japan from a Russian cultivar B. Virus Taxonomy. Viruses belonging to unassigned genera Idaeovirus Sadwavirus Sobemovirus Tobamovirus C. (eds). Elsevier/Academic Press. The updated taxonomy of all classified grapevine viruses can be found in: Fauquet. M.
INFECTIOUS DEGENERATION .
may show open petiolar sinuses. The consensus is that fanleaf degeneration may have existed in the Mediterranean Basin and the Near East since the earliest time of grape cultivation. 19 . tendrils. Chromatic alterations of the leaves vary from a few scattered yellow spots. urticado (Port.). Bunches are smaller and fewer in number. More recently. and xylem cells. Shoots are also malformed. bunches are straggly. 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. panachure. dégénérescence infectieuse (Fr. mosaico giallo. and decreased resistance to adverse climatic factors.INFECTIOUS DEGENERATION (GRAPEVINE FANLEAF VIRUS) Several nepoviruses infect grapevines in Europe and the Mediterranean area. Often infected grapevines occur in patches. and acute denticulations. which exhibit widely open petiolar sinuses and abnormally gathered primary veins. Main synonyms: court-noué. GFLV and several of the other grapevine-infecting European nepoviruses have distorting and chromogenic strains and may occur in mixed infections. With increased ambient temperatures during summer. and inflorescences). puckered. that give the leaf the appearance of an open fan. however. fasciations. but bunches are small and few. Infectious malformations are induced by virus strains causing distortions. deep lobes. Symptomatic leaves show little malformation. parenchyma.and zigzag growth. sometimes appearing as rings or lines to extensive mottling of the veins and/or interveinal areas to total yellowing . Gelbmosaik (Germ. chlorotic mottling may accompany foliar deformations. the yellowing fades rapidly and the canopy develops a normal green color. shortened productive life. especially in the basal internodes. radial bars crossing the lumen of epidermal. FANLEAF 1. This disorder was first reported from California as a syndrome elicited by a specific strain of GFLV. Tolerant cultivars produce fairly good crops whereas sensitive cultivars are severely affected. Foliar symptoms develop early in the spring and persist throughout the vegetative season becoming less distinct in summer.). shoots.e. These structures are readily visible by light microscopy in lignified shoots. In the European literature. arricciamento. Occasionally. Reisigkrankheit (partly). causing degenerative diseases whose symptoms are similar to. The name of this virus comes from the peculiar malformation of infected leaves. or indistinguishable from those of fanleaf. The foliage and shoots show little if any malformation.). short internodes. or endocellular cordons. reduced rooting ability of propagation material. are small-sized and set poorly. Leaves are variously and severely malformed. are diagnostic of grapevines infected by GFLV. low proportion of graft take. Fruit set is poor. Yellow mosaic is induced by chromogenic virus strains. vein banding symptoms were shown to be caused by a co-infection by Grapevine yellow speckle viroid and GFLV. and the yield may be much reduced. The foliage develops bright chrome yellow discolorations early in the spring that may affect all vegetative parts (leaves. double nodes.). The characterizing symptoms of "Vein banding". phloem. and grapevine leaves with typical symptoms were found in herbaria established before the introduction of American rootstock hybrids. Description Fanleaf is the oldest known and one of the most important and widespread virus disease of the grapevine. showing abnormal branching. a disorder induced by Grapevine fanleaf virus (GFLV). and berries ripen irregularly. showing progressive decline. asymmetrical. degenerazione infettiva (Ital. 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. Trabeculae. Their economic impact varies with the tolerance of the cultivar to the viruses. i. low yields and low fruit quality. roncet. Now the disease is known to occur worldwide. records of this disease date back some 150 years.
20 . Virus particles are often within tubular structures that accumulate in bundles in the cytoplasm or nucleus. Dissemination over medium and long distances is through infected vegetatively propagated scionwood and rootstocks. vinifera. Detection: ELISA using polyclonal antisera and monoclonal antibodies is a quick. In the laboratory. but is not a specific test. Chenopodium quinoa. GFLV has been detected in small groups of viruliferous X. However. reorganization. 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. but show few symptoms. encapsidated in different particles. by in vitro meristem tip culture. or by somatic embryogenesis. and very sensitive method. index (10 individuals) by ELISA and in single nematodes by RT-PCR and immunosorbent electron microscopy. C. where they occur in a radial orientation. Work is under way in different laboratories to create GFLV-resistant rootstocks or cultivar through traditional breeding methods or genetic transformation technology. and transmission by X. and soybean. but is not reliable. cheap. Indexing on Vitis indicators by grafting takes a lot of time and field or greenhouse space. although some like Vitis labrusca can be infected. Transmission: At a site. RT-PCR is estimated to be four to sixfold more sensitive than ELISA. The best antigen sources for serological diagnosis are leaves collected in spring or cortical shavings from mature dormant canes. Control: Use of virus-tested scionwood and rootstock material in the framework of clean stock or certification programmes. a number of selectable marker genes toxic to non engineered vines are used. index in V. However. Observation of symptoms in the field is useful as a first step in selection. A satellite RNA 1114 nt in size is associated with some virus isolates. O36-16) show interesting levels of field resistance to GFLV. American rootstocks are also susceptible and are generally very sensitive. In contaminated soils. serologically rather uniform and occurring as a family of minor molecular variants. in a persistent manner by the longidorid nematode Xiphinema index feeding on the roots of grapevines and retaining the virus for several months.g. Specific transmission by X. rotundifolia hybrids is thought to be controlled by a single dominant gene. rotundifolia hybrid rootstocks (e. Indexing on herbaceous hosts by mechanical inoculation requires climatized greenhouses and is less reliable than ELISA. Positive sense single-stranded RNA genome.g. C. There are conflicting reports on seed transmission in grapevines. GFLV can be transmitted by mechanical inoculation from infected grapevine tissues to various herbaceous hosts (e. Transmission by Xiphinema italiae has not been consistently documented. These inclusions derive from membrane proliferation. Resistance to X. 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. index is determined by the viral coat protein. For transformation. the use of fumigants against nematode vectors gives only a temporary but economically valuable control of the disease. rotundifolia can be infected by GFLV when graft inoculated. mannose and xylose. vinifera x M. Natural GFLV infections have been detected in weeds in Hungary and Iran. vuittenezi has been suspected but not proven. and is transmitted through seeds of C. rupestris x M. Muscadinia (Vitis) rotundifolia and Vitis munsoniana are highly resistant to X. Gomphrena globosa). M. the endosperm of grapevine seeds. This resistance is controlled by two unlinked recessive genes. 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. and redistribution and are thought to be sites of viral polyprotein processing and RNA replication. index feeding.Agent: Grapevine fanleaf virus (GFLV) is a nepovirus with polyhedral particles of about 30 nm in diameter. Varietal susceptibility: Almost all known Vitis vinifera L. amaranticolor. The virus occurs in the pollen of infected grapevines and herbaceous hosts. Virus elimination is readily achieved from vegetating shoot tips by heat treatment (38-40 °C for as little as four weeks). Some V. amaranticolor. Detection of trabeculae can give information on the health of American rootstocks. Nematode populations transmit local virus isolates with a higher efficiency than those from other geographical areas. with variable levels of sensitivity. quinoa. RT-PCR and immunocapture RT-PCR are becoming increasingly popular. However. Molecular assays using radioactive or digoxigenin-labelled probes. are toxic to many plants but not to V. GFLV was the first grapevine virus to be recovered by mechanical inoculation and to be thoroughly characterized physicochemically and molecularly. use of fumigants is more and more questioned for environmental reasons and is being progressively banned. but resists infection when the virus is transmitted by the nematode. but it is still regarded as necessary for confirming freedom from virus infection. which are desirable as they cause no harm to human health. both required for infectivity. varieties are susceptible.
Les maladies et les parasites de la vigne. 1977(b) 1865 1882 1895 1896 1902 1906 1910 1912 1912 1917 1918 Cazalis-Allut. No conclusive results were obtained. Historical review From the late 1800 to 1997. France. Bovey: Review on grapevine virus and virus-like diseases. the ICVG Bibliographic Reports (a) have recorded more than 1000 papers dealing with fanleaf. With roots of fanleaf-infected grapevines with phylloxera feeding on them. Montpellier. research and hypotheses on fanleaf. Branas et al.. Hypothesis that fanleaf is a fungal disease. 1929 1931 1937 1937 1946 1950a. With individual phylloxera (radicicolous or gallicolous) fed on infected vines. Tome 1: Les maladies dues à des végétaux (champignons. 2.: Hypothesis that court-noué (fanleaf) is caused by a virus transmitted by phylloxera. Hypothesis about a possible role of phylloxera as a vector. 21 . For a comprehensive review on early observations. bactéries. Healthy rooted cuttings or seedling of Rupestris du Lot were contaminated: 1. see the book by Galet. Arnaud: Court-noué is considered as a soil-borne virus disease. (a) See references in the Introduction. 871 pp. but only circumstantial evidence. 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. 1977. Branas et al. Imprimerie du "Paysan du Midi". 1954 1958 Hewitt: Review on grapevine virus and virus-like diseases found in California. 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. Report on first experiments on heat treatment of grapevine in order to eliminate fanleaf. No direct proof of transmission by this aphid. 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. With soil containing phylloxera. as well as on controversies about transmission by phylloxera. Schiff-Giorgini: Graft-transmission of fanleaf disease Pantanelli: Fanleaf disease can be trasmitted through the soil Pantanelli : Fanleaf disease has a patchy distribution in the field Petri: Association of trabeculae with fanleaf. 3. (b) Galet P. Petri: Grapevine "arricciamento" (fanleaf) has a viral origin Arnaud and Arnaud: Hypothesis of a viral origin for grapevine court-noué (fanleaf).2.: Experiments on the capacity of phylloxera to transmit fanleaf.b Hewitt: Fanleaf and yellow mosaic recorded from California. viroses et phanérogames).
: Detection of GFLV particles in thin-sectioned grapevine root tissues. Control of the vector by soil fumigation. Hewitt et al. including fanleaf virus: bentonite flocculation test. Serological relationship with ArMV reported. Reproduction of fanleaf symptoms in mechanically inoculated grapevine seedlings is confirmed Galzy: Heat treatment of grapevine plantlets grown aseptically in vitro.: Description of the Davis method of heat therapy of grapevines. 1965 1967 1968 1968 1969 1969 1970 22 . amaranticolor is confirmed.: Transmission of fanleaf virus from grapevine to herbaceous hosts by mechanical inoculation and preliminary characterization of the virus. Cohn et al. Vuittenez: New observations on the effects of soil fumigants on fanleaf in contaminated soils.b Martelli and Hewitt: Comparative studies showed that Californian and Italian GFLV isolates are the same. Gerola et al. yellow mosaic and ArMV 1960a 1960b 1961 1961 1962 1962 1963 1963 1963a.: Investigations on grapevine virus diseases in California. whereas insecticide treatment has no effect. index. index. and no reinfestation by X. latex test and barium sulfate test.: Transmission of GFLV by Xiphinema italiae in Israel. Bercks and Querfurth: Use of latex-test for detecting GFLV and other nepoviruses in grapevine tissue extracts in Germany. Vuittenez: Mechanical transmission of fanleaf virus to Chenopodium quinoa and C. index in France.: Fanleaf virus is transmitted by the nematode Xiphinema index Cadman et al. Goheen and Hewitt: Description of vein banding as a GFLV-induced disease Dias: Host range and properties of fanleaf and yellow mosaic viruses. Transmission of fanleaf virus by X. Goheen et al. Dias and Harrison: Relationships between the viruses causing fanleaf. Bercks: Research on the use of three serological methods for detecting plant viruses. index occurred during the 6 year period of observation.1958 1958 1960 Vuittenez: Fumigation of fanleaf-contaminated soil with nematicides prevents infection of healthy grapevines replanted immediately. Graniti and Russo: A light microscope and cytochemical study of endocellular cordons. Discussion on the possible role of weeds in the epidemiology of the disease. Yield was increased by 400 % in comparison with that of untreated controls. Das and Raski: Studies on the relationships of GFLV with its vector X. Gifford and Hewitt: Use of heat therapy and in vitro shoot tip culture to eliminate fanleaf virus from infected grapevines. Description of the chipbudding method for indexing. Reproduction of fanleaf symptoms in mechanically inoculated grapevine seedlings 1964 1964 1965 Taylor and Hewitt: Description and characterization of Australian isolates of GFLV. Dichloropropane-dichloropropene (DD) at 1000 l/ha gave satisfactory results. Brückbauer and Rüdel: The virus (or viruses) of Reisigkrankheit (GFLV and/or other nepoviruses) are seed transmitted in some herbaceous indicator plants. 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. Boubals and Dalmasso: Experiments on soil disinfection against X. Hewitt et al.
rupestris St.: Use of green grafting as a quick and secure method for graft-indexing. Raski et al. Bass and Vuittenez: Thermotherapy was improved by growing shoot apices of heat treated vines aseptically on nutritive media or by grafting them on aseptic grape seedlings in vitro. Raski et al. Both tests are more sensitive than mechanical inoculation to C. Alfaro and Goheen: The different strains of fanleaf virus (fanleaf sensu stricto. but ELISA is more sensitive.: GFLV particles observed in the lumen of the oesophagus of X. anterior oesophagus and oesophageal bulb of their nematode vectors. St.3-dichloropropene or methyl bromide.: Control of fanleaf by soil fumigation with 1. Indexing by budding on V. quinoa.3 dichloropropene or methyl bromide. index. Hévin et al. 1970 1970 1970 1971 1971 1972 1972 1973 1973 1973a 1973b 1974 1974 1975 1976 1976 1977 1979 1979 23 .: Detailed physico-chemical characterization of GFLV. Van Velsen and Niejalke: Green budding for indexing grapevine with the indicator cvs.: Comparison between PALLAS latex test and ELISA for detecting GFLV in France. this lining is shed and ingested in the intestine. A dormant bud of the variety to be cured is grafted onto a healthy potted rootstock. Raski and Schmitt: Progress in the control of the fanleaf-nematode complex by soil disinfection with 1.: Heat therapy of grape plantlets grown in vitro causes changes in some characteristics of the variety. The acquisition time threshold is less than 5 minutes. After bud take. rupestris is more accurate than mechanical transmission to C. Bercks: Serological detection of grapevine viruses in West Germany. George. index. Walter et al. Mur et al. Querfurth and Paul: Protein A-coated latex-linked antiserum (PALLAS) method for detecting GFLV and other viruses.: Comparison of indexing by mechanical inoculation to Chenopodium quinoa and by graft-transmission to V. the plant is placed in a heat cabinet for treatment Hévin et al. Dias: Review paper on grapevine yellow mosaic.: 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. The sensitivity of the latex test is increased. PALLAS is quicker and cheaper than ELISA. During the moult of the nematode. Mission or LN 33. George for detecting GFLV. 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. quinoa.1970 1970 Hewitt et al. Vineyards replanted in contaminated but treated soils remained healthy for at least 5 years. Uyemoto et al. Vuittenez: Review paper on grapevine fanleaf. Taylor: Review paper on grapevine vein banding.: GFLV and marbrure (fleck) are not transmitted through the seeds of grapevine. Both methods give satisfactory and similar results. yellow mosaic and vein banding) are transmitted in the same way by X. Goheen and Luhn: New method of heat therapy. especially with low titre antisera.
a very common species in vineyards of Rheinhessen and Palatinate. Rüdel: Discussion on the possible role of X. rotundifolia is resistant to fanleaf virus transmission by X. Walker et al. index. Carbon disulfide gave less satisfactory results. although it is not resistant to the virus itself. Russo et al.: Detection of fanleaf virus and other sap-transmissible viruses in grapevine tissues by ISEM.: Detection of fanleaf virus in grapevine tissues by ELISA and ISEM in different periods of the year.: Identification of several Vitis species and interspecific hybrids resistant to fanleaf virus. Even in the cases where the virus was transmitted. Brown and Roberts: Detection of fanleaf virus in its vector X. index and fanleaf in grapevines. 1980 1980 1980 1981 1981 1981 1981 1983a 1983b 1983 1983 1983 1985 1985 1986 1986 24 .: Experiments with systemic nematicides for controlling X. especially in wood shavings of dormant canes during winter. in California. Vuittenez: Review on serological methods of detection and identification of grapevine viruses. Hafez et al. the possibility that a few X.: 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. Lear et al. as vector of GFLV.: GFLV particles are arrayed in long parallel rows in thin-sectioned mesophyll cells of infected grapevines. A Middle Eastern V.: Use of systemic nematicides for the control of X. These are promising sources of germplasm for obtaining resistant rootstocks.: Comparison of polyclonal and monoclonal antibodies for detecting fanleaf virus with ELISA in various grapevine tissues.1979 1980 1980 Kalasian et al. Morris-Krsinich et al. index.3-D (1400 l/ha) and 50-75 cm deep with 165 cm spacing for methyl bromide (448 kg/ha) gave a good control of X. Raski et al. The use of methyl bromide requires a continuous cover with polyethylene sheeting for some time after the treatment. Bouquet: Serological detection of GFLV in its vector X. vuittenezi population used for the experiments could not be entirely ruled out. Efficiency of both methods is compared. RNA-2 contains the cistron coding for the viral coat protein. Savino et al. That X. index.: Study on the effectiveness of soil fumigation for the control of X.: Identification of a natural serological variant of GFLV from Tunisia Huss et al. Transmission trials gave a few positive results. vinifera accession represent an excellent example of host plant resistance to GFLV. 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. Methyl bromide and 1. Raski et al.3-D) or methyl bromide applied 75-90 cm deep with 90 cm spacing for 1. index by ISEM Bovey et al. index larvae were present in the X. index feeding. Monette: Heat therapy of GFLV and ArMV--infected grapevines with alternating temperatures. Bouquet: Muscadinia rotundifolia becomes infected by GFLV when the virus is transmitted by grafting but resists infection when transmission is by X. vuittenezi might be a vector of GFLV is therefore uncertain. Bouquet: M. vuittenezi. Krake and Woodham: Possibility that the agent of yellow speckle is involved together with GFLV in the etiology of vein banding. Forty days of treatment. index by ELISA. index.: Soil fumigation with 1.3-dichloropropene (1.3-dichloropropene failed to eradicate either nematodes or fanleaf virus from the soil but reduced the incidence of the disease to acceptable levels.
Reliable results were obtained with samples of 20-50 nematodes. TBRV and leafroll from infected grapevines by in vitro meristem tip culture. Walker and Meredith: Identification of two accessions of Vitis vinifera resistant to GFLV. the latter being especially damaging on the variety Kerner. Catalano et al. No eradication was obtained. index and GFLV. Mild strains were shown to confer protection towards severe challenging strains in Chenopodium and grapevine. GFLV. Mild and severe strains were discriminated on the basis of field performance of infected Vitis. rotundifolia showed good resistance to GFLV in California.3-dichloropropene and methyl bromide for controlling X.1987 1987 1987 Huss et al.: Use of green grafting technique for sensitive and quick GFLV detection under greenhouse conditions. vinifera x V.: Detection of GFLV in grapevine seeds and seedlings by ELISA. 1988 1988 1988 1989 1989 1989 1989 1989 1990 1990 1990 1990 1990 1991 1991 1991 25 .: Identification of a satellite RNA of GFLV. Walter and Etienne: Detection of GFLV in wood shavings of dormant canes. but less consistent results were obtained with 1-10 nematodes. Long term fallow (about 5 years). RNA-3 encodes a non structural protein. Catalano et al. Previous experience showed that 1. quinoa and transmitted by heterografting to Vialla and Kober 5BB rootstocks.: Detection of GFLV in the vector Xiphinema index by ELISA. Treatments with soil fumigants are no longer permitted in Germany.: Improvement in the serological detection of GFLV and ArMV viruses using monoclonal antibodies.: Study of interactions between GFLV and ArMV isolates grown in C. Effect on yield and economic importance. Martelli and Taylor: Review article on nematode-transmitted viruses. SLRV. The selection of resistant cultivars and rootstocks is considered of primary importance. Viruliferous nematodes were crushed in standard extraction buffer and tested in batches of 1-50 by means of DAS-ELISA.: Two rootstock selections derived from crossings V. RpRSV and ArMV are common. cultivation of non-host plants and organic soil amendments are recommended. ArMV. RpRV. Walter et al.3-dichloropropene or methyl bromide fumigation following one year fallow period can give a satisfactory control of the disease for at least 12-15 years. Lázár et al. Etienne et al. Pinck et al.: Possibility of detecting several nepoviruses or serotypes of nepoviruses in grapevine leaves or wood shavings in a single DAS-ELISA test using a mixture of different polyclonal antisera. Positive. Fuchs et al. Walter et al. Serghini et al. Resistance is controlled by two unliked recessive genes Walter et al. Rüdel: Severe restrictions set on the use of soil fumigants in West Germany for environmental reasons make control of "Reisigkrankheit" very difficult. Walker et al.: Determination of the complete sequence of GFLV RNA-2.: Production and use of monoclonal antibodies to GFLV.: Evidence of a differential efficiency of GFLV transmission by Xiphinema index populations from different geographical origins. Altmayer: Elimination of GFLV. Treated vines yielded more for over 4 years. Raski and Goheen: Comparison of 1. Rüdel: Review on the most important virus diseases of grapevines in West Germany. and has strong homologies with the satellite RNA associated with ArMV.: Determination of the nucleotide sequence of the satellite RNA (RNA-3) of GFLV.
hybrids and breeding stocks after infection of the roots by means of viruliferous X. index.: GFLV satellite RNA detected in 5 of 34 virus isolates from different geographical locations.: Detection of GFLV in X. Satellite RNA appears to have a modulating effect on virus pathogenicity. Goussard and Wiid: First application of somatic embryogenesis for sanitation of grapevines.: Production of GFLV satellite RNA transcripts. Viry et al. Esmenjaud et al.: Production of biologically active transcripts from cloned cDNA of genomic RNAs of GFLV. Staudt: Study of the spread of GFLV in several Vitis species. 1993 1993 1993 1993 1993 1994 1994 1994 1994 26 .: Evidence that transgenic tobacco plants expressing the coat protein of GFLV are protected from GFLV infection. Horvath et al. thus qualifying for use in X. quinoa with biologically active transcripts of GFLV F-13 satellite RNA and GFLV strains devoid of satellite. Walker et al. O39-16 in particular.: Development of cDNA probes to GFLV genomic and satellite RNAs and their use for virus detection directly in grapevine extracts. Spielmann et al.1991a 1991b Fuchs et al. index.: 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.: Two Vitis vinifera x Muscadinia rotundifolia rootstock hybrids (O39-16 and O4343) grafted with Cabernet sauvignon showed a high level of tolerance to GFLV. which is also resistant to phylloxera. Both became infected in the course of a 12-year trial but had no reduced crop yields. delays symptom expression by 1-2 days and adversely affects virus replication. index by biotin-avidin ELISA Bardonnet et al. This represents the first substantiated record of a natural GFLV infection in hosts other than Vitis. Walter et al. Ritzenthaler et al.: Genomic RNA-1 of GFLV is completely sequenced and its genetic organization determined. GFLV is eliminated from somatic embryos obtained from tissue cultures grown at 35 °C 1991 1991 1992 1992 1992a.: Use of modified GFLV coat protein genes for transformation of different Nicotiana species for inducing resistance. Fuchs et al.: Co-inoculation of C.b Hans et al. GFLV is a quasispecies occurring in the field as a series of minor molecular variants.: In naturally GFLV-infected vineyards the hypovirulent ArMV A1 isolate induces delayed infection by GFLV Saldarelli et al.: GFLV isolated in Hungary from naturally infected symptomatic plants of Aristolochia clematis and Lagenaria siceraria turbinata.:Detection of GFLV in single nematodes by RT-PCR. identification of their replication determinants and evidence of replication in Chenopodium quinoa protoplasts 1993 1993 1993 1993 Martelli et al. Esmenjaud et al. Staudt and Weischer: Vitis rotundifolia and Vitis munsoniana resist infection by GFLV transmitted by X. index infested soils.
Lahogue and Boulard: Search for genes of resistance in grapevines. De Luca et al. Naraghi-Arani et al. American. Walter and Martelli: Review article on detrimental effects of viruses on grape yields. including the two accessions reported as resistant by Walker and Meredith (1990).: Up to date account of GFLV replication strategy. Ritzenthaler et al.: Demonstration that a 28 kDa protein coded by GFLV RNA-2 is involved in the replication of this RNA. 5 oligoadenylate synthase.: 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. Krastanova et al. Gölles et al. Gaire et al.: Identification of membranes derived from the endoplasmic reticulum as sites of GFLV replication. 1997 1998 1998 1999 1999 2000 2000 2000 2000 2001 2002 2003 2003 2003 27 . index. Pinck: A comprehensive review of the molecular aspects of GFLV genome and its replication strategy. Belin et al. Of 531 accessions of European.: Transformation of Nicotiana species and Vitis rupestris with different virusderived (coat protein. Belin et al. Mauro et al. Martelli and Walter: Review article on certification of grapevines.: Identification of the molecular signal accounting for the systemic spread of GFLV in infected hosts. rotundifolia hybrids show high resistance to X. index feeding.: Genetic transformation of Vitis vinifera with the coat protein gene of GFLV for resistance induction. The level of resistance obtained looks promising. Pfeiffer et al. Pfeiffer et al. replicase) and exogenous (2. RNase L) genes for inducing resistance to GFLV. and Asian Vitis species inoculated by green grafting with a GFLV source. Walker and Jin: V.: Attempts to characterize molecularly X. Fuchs: Review article on genetic transformation of grapevines for resistance to GFLV and other pathogens.1995 1995 1995 1995 1995 1996 1996 Brandt and Himmler: Use of immunocapture RT-PCR for GFLV detection in host tissues. all were susceptible to the virus. Rowhani et al.: Genetic transformation of American roostocks with the coat protein gene of GFLV for resistance induction. index populations by PCR-RFLP and sequencing of the ITS region. : Identification of RNA2-encoded proteins in the specific transmission of GFLV by X.: The membranous structures appressed to the nucleus of infected cells known as vacuolate-vesiculate inclusion bodies are virus factories as they are the likely site of RNA replication and processing of viral polyproteins. rupestris x M.: Successful transformation of somatic embryos of an European grape cultivar (Russalska 3) with the normal. This resistance is controlled by a single dominant gene.: Demostration that the movement protein of GFLV is located on the intracellular tubular structures containing rows of virus particles.: Development of a GFLV detection system based on PCR analysis of immobilized virions. Ritzenthaler et al. Spielmann et al. truncated or nontranslatable coat protein gene of GFLV. except for four.
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A.P. 1981. Wellink. In: Virus Taxonomy.A. Taylor and Brown. eds).). Amsterdam. 286 pp.G. Martelli and R. Nepovirus Group. These viruses can readily be eliminated by heat therapy or in vitro meristem tip culture. 1978. G. 1992.. Murant (eds). Influence des viroses et qualité. and A. D.. Sélection clonale de la vigne: sélection sanitaire et sélection pomologique. M. Brown.V. Martelli. Piazzolla. subgroup B. the Mediterranean and Middle East. Quacquarelli. CMI/AAB Descriptions of Plant Viruses No. has now been assigned to the newly established genus Sadwavirus (Le Gall et al. Quaderno No. All have polyhedral particles about 30 nm in diameter and a positive sense.A. Plenum Press...J.. (G. 1996. 807-818.. Murant. Eight Report of the International Committee on Taxonomy of Viruses (C. 1996. Jones. Rüdel M. family Comoviridae (Goldbach et al. Scholtof. Sixth Report of the International Committee on Taxonomy of Viruses (F. Gallitelli. 1955) and are subdivided into subgroups based on physico-chemical properties of member viruses.A. Taliansky.P.. 2000) are available..F.B. V. K. H. J. 1992). Many are transmitted by longidorid nematodes (Rüdel. K.E and D. Fauquet. (E. J. Seventh Report of the International Committee on Taxonomy of Viruses (M. Lehto. 1996. 362 pp. G. and A. M. Le Gall O. CAB International. single-stranded RNA genome occurring as two functional species (RNA-1 and RNA-2). Oxon.D. Sanfaçon. a non-taxonomic clustering. Iwanami. epidemiological. New York. The Plant Viruses. In: Handbook of Plant Virus Infections: Comparative Diagnosis. 3. 23-39..e. Karasev. In: Virus Taxonomy.F. 35 . subgroup C. 1955. Leibowitz. U. which were originally included in the Nepovirus group (Harrison and Murant. Vol. ed. 2005. 10281032 Murant A. A. Milne. typified by Tomato ringspot virus (ToRSV) (Martelli et al. Strawberry latent ringspot virus (SLRSV). Mayo.D. Family Comoviridae. Bulletin de l'OIV 69. subgroup A typified by Tobacco ringspot virus (TRSV). Nepoviruses. Several of these viruses have distorting and chromogenic strains and may occur in mixed infections with GFLV. ed). Van Regenmortel et al. Vienna. Mechanical transmission to herbaceous hosts or indexing on Vitis indicators can also be used. Murphy et al. Desselberger and L.V. Bari.A.P. Walter B. 2005). Satellite nucleic acids. Ball. A tentative grouping of nepoviruses Phytopathologia Mediterranea 17. 1997) and their satellite RNAs (Mayo et al. Istituto Agronomico Mediterraneo. and molecular characteristics of nepoviruses (Harrison and Murant.. eds) Elsevier/Academic Press. Martelli. Murant 1981. 2005) Extensive reviews of the biological.. Nepoviruses. Harrison B.M. Nepoviruses of grapevine and their nematode vectors in the EEC. In: Virus Taxonomy. Kurstak. Harrison B. 2000. Family Comoviridae. Academic Press. References Goldbach R. 1997.H. 945-971. physico-chemical. Wetzel and N. 1978. which are separately encapsidated. 1977) . 145-147. ISEM) and molecular assays (hybridization. 197-238. Nematode Vectors of Plant Viruses. are now classified in the genus Nepovirus.. typified by Tomato black ring virus (TBRV). 1ere partie: Effects des viroses sur la culture des vignes et ses produits. Maniloff..P. or indistinguishable from those of fanleaf. 5.J. In: Grapevine Viruses and Certification in EEC Countries: State of the Art. T. Taylor C. 185. RT-PCR) are routinely used for their detection in grapevine tissues (primarily cortical scrapings from dormant canes). Their detrimental effects to grapevine culture and products have been summari zed by Walter and Martelli (1996). Martelli G.F. London. 341-347. Fritsch. 1977. T. C. Polyhedral Virions and Bipartite RNA Genomes. Serological (ELISA. Springer-Verlag. Palukaitis. San Diego. Yoshikawa. Elsevier/Noth Holland. Savino and P.F. eds). Mayo M.A. Simons and M. P. i. Le Gall et al. T.INFECTIOUS DEGENERATION (EUROPEAN AND MEDITERRANEAN NEPOVIRUSES) Besides GFLV several other nepoviruses can infect grapevine in Europe. a nematode-borne virus originally classified as a tentative species in the genus Nepovirus. and G. causing diseases whose symptoms are similar to.
Vuittenez et al. In other V.: Physico-chemical properties of GFLV.1 x 106 (RNA-2). 36 . Kerner appears to be caused by ArMV infection. USA (California). 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. always in Germany the severe dieback disease of the cv. Bulgaria. and Japan. Transgenic plants expressing the coat protein gene of the virus have been produced. 2.4 x 106 and a size of 1104 nt.: Interactions between nepoviruses in grapevine and herbaceous hosts. wt 2.: Isolation of ArMV from grapevine in Italy. Stellmach: Review paper on ArMV in grapevine.: Ultrastructure of ArMV infections in plant tissues Martelli and Lehoczky: Detection of ArMV in grapevine in Hungary. have a angular outline. Component T is made up of empy protein shells. Yugoslavia. Hungary. and with other nepoviruses in the Reisigkrankheit complex of western Germany. respectively. Cross-protection between ArMV and GFLV has been reported.95-2. Description ArMV. The virus supports the replication of two types of satellite RNAs. Rüdel: Transmission of ArMV to grape seedlings by Xiphinema diversicaudatum. Iran. accounting for 27% (component M) and 41% (component B) of the particle weight. wt of 0. TBRV.: ArMV. Its particles are about 30 nm in diameter. Russo et al. and circular about 350 nt in size. Turkey. M.Genus NEPOVIRUS ARABIS MOSAIC VIRUS (ArMV) 1. Quacquarelli et al. Dalmasso et al. The genome is a positive sense ssRNA. It can infect many woody and herbaceous plants and is transmitted to grapevine by the nematode Xiphinema diversicaudatum but not by X. consisting of two separately encapsidated molecules with Mol. 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. symptoms are of the fanleaf type. and.: Xiphinema diversicaudatum can transmit ArMV to grapevine. Murant: ArMV description in the CMI/AAB Descriptions of Plant Viruses series. vinifera varieties. Coat protein has a single type of subunits with Mr 54 x 103.: ArMV detected in Japan in grapevines imported from Europe. AILV and GCMV. is serologically related to GFLV. Romania. whereas components M and Bcontain RNA. Israel. ArMV and TBRV by ELISA in Israel. SLRSV and TBRV found in grapevines with atypical Reisigkrankheit in West Germany. ArMV occurs often in mixed infections with GFLV in certain areas of France and Italy. In Germany.2 x 106 (RNA-1) and 1. Jankulova and Kaitasova: ArMV found in grapevine in Bulgaria. and sediment as three components (T. and B). Gerola et al. losses up to 50 % have been recorded. Canada. linear with Mol. index. a typical nepovirus belonging in subgroup A of the genus Nepovirus.: Detection of ArMV by ISEM Tanne: Detection of GFLV. ArMV. This virus has also been found in grapevine in Switzerland. Kobayashi et al. Belli et al. Brückbauer and Rüdel: Symptoms of atypical Reisigkrankheit in the vineyard are associated with ArMV in West Germany Bercks et al. the vector of GFLV.
: 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. the virus is recovered from the scion only during the first year. Walter et al.: Properties of ArMV small satellite RNA.: Evidence that ArMV has two RNA-2 molecules and complete nucleotide sequence of both RNAs Etscheid et al. : Comparison of coat proteins of ArMV and other nepoviruses. When a healthy scion is grafted onto an infected rootstock. Rüdel: In the Palatinate (West Germany) ArMV is transmitted by Xiphinema diversicaudatum and occurs often in mixed infections with GFLV in grapevine. Becker et al. Steinkellner et al. Hypothesis of a graft incompatibility when the rootstock is infected with ArMV.: Detection of ArMV by PCR in herbaceous hosts and grapevines Steinkellner et al. Yield losses may reach 77% in cv.: Association of ArMV-infected rootstocks with Kerner disease in West Germany.: A hypovirulent ArMV isolate delays GFLV infection in grapevines under field conditions.: Nucleotide sequenece of a small circular satelllite RNA Steinkellner et al.: Cross-protection experiments in Chenopodium quinoa between ArMV and GFLV Gugerli et al.: Nucleotide sequence of the ArMV satellite RNA Liu et al. Stellmach: Kerner disease is probably caused by ArMV.: Use of cDNA probes for ArMV detection. Eppler et al. RRV.Kerner to ArMV seems to be limited in time.: Transgenic Nicotiana plants transformed with the coat protien of ArMV produce empty viral shells. whereas the rootstock remains infected. Kaper et al. Study of histological changes at the graft union level. The virus cannot be recovered from leaves or buds of the Kerner scions.1982 1984 1985 Brückbauer: Possibility of distinguishing GFLV. Ipach et al. such as RpRSV or GFLV can be found in both rootstock and scion. Faber. MacKenzie et al. ArMV.: Properties of a strain of ArMV isolated from grapevine in Italy.: ArMV recorded from Romania Huss et al. SLRV and TBRV Belli et al.: ArMV in Switzerland Lázár et al.: Transformation of grapevines with the coat protein gene of ArMV. whereas other nepoviruses.: ArMV is not seed-transmitted in grapevines Liu et al. Molecular assays are as good as ELISA for routine testing. Stellmach and Berres: The susceptibility of cv.: The presence of ArMV satellite RNA can attenuate symptoms in certain hosts Bertioli et al.: Survey for the presence of ArMV in Canadian vineyards Lahogue and Boulard: Search for genes of resistance in grapevines. Marc-Martin et al. Of 407 accessions of 1986 1987 1988 1989 1989 1989 1989 1990 1990 1990 1991 1991 1992 1992 1993 1993 1994 1995 1995 1995 1996 1996 37 .
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
Component T is made up of empy protein shells. wt of 2. GBLV has also been recorded from Hungary and Yugoslavia.: Isolation of virus CM112 from in vitro cultures of grapevine tissues. Martelli. Boscia and G. Two isolates of GBLV have been transmitted by mechanical inoculation to seedlings and rooted cuttings of several grapevine cultivars without inducing symptoms. physico-chemical and serological characterization of the virus and assignment to the Nepovirus group (now genus Nepovirus). Sabanadzovic. References Abou Ghanem-Sabanadzovic N. The scattered distribution of infected vines in the field suggests that the virus is spread primarily by infected propagative material..: Description and thorough characterization of GARSV identified as a new species in the subgroup B of the genus Nepovirus Abou Ghanem-Sabanadzovic et al.vector. whereas components B1 and B2 contain RNA.P.: GBLV description in the CMI/AAB Descriptions of Plant Viruses series. Uyemoto et al. Martelli et al. B1.: First isolation by mechanical transmission of an unknown nepovirus from cv. Martelli. Historical review 2002 2003 2005 Cigsar et al. I. 2.. The virus occurs as different closely related but serologically distinguishable strains. Digiaro and G.2 x 106 (RNA-1) and 1. 2002. Digiaro and G. De Stradis. The virus can be detected directly in grapevine leaf extracts by immunodiffusion in agar plates. Cigsar I.: Complete nucleotide sequence of GARSV RNA-2 3. Sanitary status of grapevine in south eastern and central Anatolia. GRAPEVINE BULGARIAN LATENT VIRUS (GBLV) 1.95-2. Journal of Plant Pathology 85. 2. 2003. Martelli. accounting for 39% (componet B1) and 42% (component B2) of the particle weight.: Description of GBLV. 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. Virus particles are about 30 nm in diameter and sediment as three components (T. Coat protein has a single type of subunits with Mr 54 x 103. D. GBLV is a typical nepovirus belonging in subgroup C of this genus but its vector is not known. Description Grapevine Bulgarian latent virus (GBLV) owes it name to the fact that it was found for the first time in Bulgaria in 1971. The economic importance of the virus is minor.: A virus serologically related to GBLV isolated from Vitis labrusca cv. Biological. M. N. 35-41. A strain of this virus had been found previously in Portugal and described as virus CM112. Properties of a previously undescribed nepovirus fron south-east Anatolia. Bulletin OEPP/EPPO Bulletin 32. Cigasr. is not seed borne and was reported only from south-eastern Turkey. De Mendonça et al. but different from GBLV.5 x 106 (less than 1800 nt). M. where it is widespread and infects latently several grapevine varieties growing in widely separared areas. The virus supports the replication of a satellite RNA with Mol. 1977 1978 41 . Concord in New York State.P. By contrast. 335-340. consisting of two separately encapsidated molecules with Mol. Abou Ghanem-Sabanadzovic. The genome is a positive sense ssRNA. wt 0. Complete nucleotide sequence of RNA-2 of two Turkish nepoviruses. A. Kizlar tahtasi showing mild fanleaf-like symptoms Gokalp et al. S. 471-475 Gokalp K.1 x 106 (RNA-2). Digiaro. and B2). Virus Genes 30. 2005. isolated in 1970 in Portugal from symptomless vines.P. Martelli et al. M. Historical review 1972 1972 1977 Ferreira and De Sequeira: Description and preliminary characterization of an unidentified virus denoted CM112.
Ferreira. Numéro hors série. France. Varennes and De Sequeira: First application of ELISA for the detection of virus CM112. and O. De Sequeira O. De Sequeira and A.P. Cordoba 1976. 95-101. Stace-Smith. 1972.A. P. A. Gallitelli.P. Niagara Falls 1980. Abracheva. No. De Sequeira and Vasconcelos-Costa: Use of an immunoradiometric assay for the titration of the Portuguese strain of GBLV. P. M. 27-32. Garcia de Orta Estacao Agronomica 12.. Quacquarelli. Properties of a distinctive strain of Grapevine Bulgarian latent virus. Rasteniev'dni Nauki 29. Di Franco. Gallitelli D. Some properties of the new latent virus from grapevine rootstocks in Yugoslavia. Pocsai E.A. 113-120.. Proceedings 7th Meeting of ICVG. Gallitelli et al.A. 42 . 143-145 De Mendonça A. Annales de Phytopathologie.C. 21-24. D. The ultrastructure of grapevine Bulgarina latent virus infections in natural and artificial hosts. 1980. Niagara Falls 1980. V.: Improvement of ELISA protocol for GBLV detection the whole year round. References De Mendonça A. 135-141.: Ultrastructural study of GBLV infections in grapevine and C. Savino and A. 1979. 1972. Pocsai: Occurence of GBLV in Hungary. Martelli G. Russo et al. Ramsdell D.. Numéro hors série.: Detection of virus CM112 in grapevine leaf extracts by ISEM. Ganeva and M. 1981. 1985. Grapevine Bulgarian latent virus. 1980. Pereira and V.A. and R. V. 1978. Quacquarelli. Simoes. O.1979 Martelli et al. Vasconcelos-Costa. The Portuguese strain supports the replication of a satellite RNA. Occurence of grapevine Bulgarian latent virus in Hungary. Proceedings 6th Meeting of ICVG. Niagara Falls 1980. O. 217-222. 1980. Savino and O. quinoa. D. Martelli et al.N. M. and J. De Sequeira. Martelli G. Savino and A. Phytopathology 71. Colmar 1970. Colmar.A. V. 468-472. 349-354. A preliminary report. Preliminary studies on an undescribed grapevine virus. Dimitrijevic B. Krastanova et al.. M. Dimitrijevic: GBLV found in Yugoslavia De Mendonça et al.P. Acta Phytopathologica Academiae Scientiarum Hungaricae 16. Ferreira A. Phytopathologia Mediterranea 22. D. CMI/AAB Descriptions of Plant Viruses. A. Gallitelli. De Sequeira. Some properties of grapevine Bulgarian latent virus..: 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. Possibilities for the whole-year ELISA detection of viruses infecting grapevines.P. Gallitelli.A.. A.Sur l'isolement d'un virus à partir de cultures de tissus de vigne. Krastanova S.. Abracheva..Proceedings 7th Meeting of ICVG. Jankulova.. Ramsdell and Stace-Smith: The New York isolate of GBLV is a strain of BLMoV. 245-250. Monografias INIA 18 . Proceeding 4th Meeting of ICVG.. 269-272. 186 Martelli G. Physical and chemical properties of the particles and ribonucleic acid of blueberry leaf mottle virus. 1983. 1977.A. 1970. Applicability of immunosorbent electron microscopy (ISEM) for the detection and identification of CM112 virus in grapevine. Yankulova. Russo and V. Martelli G. Quacquarelli and D.: 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. A manually transmissible latent virus of the grapevine from Bulgaria. Annales de Phytopathologie. De Sequeira. Savino. 1992. 51-58.: Detection of GBLV in grapevine leaf extracts by ISEM. 1981. Annals of Applied Biology 85. An immunoradiometric assay for the titration of a Portugese strain of grapevine Bulgarian latent virus (GBLV). Proceeding 4th Meeting of ICVG. 1980 1980 1980 1980 1981 1981 1982 1983 1985 1992 3. Mota.
RNA-2 has Mol. Lehoczky and Tasnady: A study of the effect of HCMV on yield and sugar content of infected grapevines. It has been recorded also from Czechoslovakia. Affected vines lack in vigour and may decline and die. and O. The virus belongs in the same subgroup of TBRV (subgroup B) to which is distantly related serologically. De Sequeira. Martelli and Quacquarelli : Physico-chemical characterization of HCMV and comparison with TBRV. Kenten: GCMV is distantly serologically related to Cacao necrosis virus. a size of 4441 nt and accounts for 31% of the particle weight. a symptom virtually indistiguishable from GFLV-induced yellow mosaic.K. Martelli and Quacquarelli: Description of HCMV in the CMI/AAB Descripitons of Plant Viruses series. Taschenberg and D. The genome is bipartite.. 269-277. Jakó et al: HCMV affects pigment and sugar content of infected grapevine leaves. E. Plant Disease Reporter 61. Tobacco pants and the rootstock 110R have been successfully transformed with the viral coat protein for induction of resistance. sometimes together with X. Hummer. RNA-1 has Mol. The virus appears to be unrelated serologically to GFLV and is not transmitted by X. vuittenezi is very frequently found in vineyards with chrome mosaic patches. Isolation and identification of a strain of grapevine Bulgarian latent virus in Concord grapevines in New York State. 1977.: GCMV recorded from Slovakia and report of X. GRAPEVINE CHROME MOSAIC VIRUS (GCMV) 1. a size of 7212 nt and accounts for 40% of the particle weight. 1966 1968 1968 1969 1969 1971 1972a 1972b 1972 1975 1977 43 .: 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. wt of 1. Mali et al.: HCMV adversely affects photosynthetical carbon dioxide fixation. The coat protein has a single type of subunits of Mr 52 x 103.K. Although GCMV particles have been detected by immunosorbent electron microscopy in Xiphinema index fed on infected hosts.6 x 106 . and was originally called Hungarian chrome mosaic virus. index. pretty much like GFLV. Croatia and Austria. transmitted to herbaceous hosts from Hungarian grapevines with symptoms similar to those of fanleaf and yellow mosaic.: Host range and properties of a spherical virus. near Lake Balaton. wt of 2. called Hungarian chrome mosaic virus.Uyemoto J. index as vector of the virus (unconfirmed results). Detection of CM122 latent grapevine virus by enzyme-linked immunosorbent assay (ELISA). Saric and Hranuelli: GCMV recorded from Croatia. 1982. Martelli and Sarospataki: X. double nodes and short internodes. However. GCMV is transmitted through grapevine seeds. 2. symptomless infection may occur. Some virus strains induce leaf deformity. Virus re-named Grapevine chrome mosaic virus. No evidence that X.F. The virus is not serologically related with GFLV. 949-953.A. Historical review 1966 Martelli et al. Martelli: Purification and serology of the virus isolated from Hungarian grapevines with fanleafand yellow mosaic-like symptoms. Varennes A. Agronomia Lusitana 41.8 x 106 . Martelli et al. index. early reports that this nematode could transmit the virus have not been confirmed. Pozsár et al. vuittenezi transmits GCMV or GFLV. Description Grapevine chrome mosaic virus (GCMV) was first found in Hungary. Evaluation of short reaction times and re-use of a-globulin and conjugate.
Genetically engineered resistance against grapevine chrome mosaic virus. 231-238.: Tobacco plants genetically engineered with the coat protein gene of GCMV are resistant to infection. index extracts by ISEM. Le Gall et al.: Detection of GCMV in leaf dips by ISEM. Journal of Phytopathology 142. Lehoczky: Pinot noir and Jubileum 75 are good indicators for GCMV. Ravelonandro and J.: GCMV and TBRV can recombine. Delbos. Lanneau and J.. 44 . 3.: Description of a certification scheme for the production of virus-free propagating material in Hungary.. index are inconclusive. Brault V. M. T. O. L.: Transformation of roostock 110R with the coat protein gene of GCMV. Acta Horticulture 235. References Brandt S. Brault et al. Further demonstration that the two viruses are related. Brault V. 1993. Taylor and Brown: Results of GCMV transmission trials with X. Lehoczky et al. Le Gall and J. Hilbrand. 258-262. Vitis 34. Lázár et al: Seed transmission of GCMV in grapevine. Dodd and Robinson: GCMV and TBRV are molecularly related. Dimou D. Brault.: Complete nucleotide sequence of GCMV RNA-2.. Bretout C. Occurrence of grapevine chrome mosaic nepovirus in Austria. Le Gall.: Characterization of a GCMV strain and confirmation of its serological relationship with TBRV. Nucleotide sequence and genetic organization of Hungarian grapevine chrome mosaic nepovirus RNA2. D'Onghia. Lázár et al. Himmler. Candresse. 127-128. Kölber et al. Savino.P. 1989. Dunez. This finding does not imply vectoring capacity by this nematode. and G. O. 1994. V.: GCMV detected by ELISA in infected field-grown vines. Le Gall. Candresse. Brandt and Himmler: Development of a IC-PCR protocol for GCMV detection in cortical scrapings from dormant grapevine canes. Laimer da Camara Machado and V. T. M. T. Dunez. R. Bretout et al. The virus vector is yet to be identified.: GCMV cross-protects Chenopodium quinoa from the severe apical necrosis induced by a TBRV strain.: GCMV recorded from Austria. 1989. A. 1995.: Development of molecular probes for GCMV detection. Dimou et al. Doz et al.1979 1980 1980 1982 1984 1985 1985 1989 1989 1989 1990 1993 1993 1994 1994 1995 1995 1997 2000 Lehoczky et al. Le Gall et al. 7809-7819.M. M. Virus and RNAspecific molecular hydridization probes for two nepoviruses. O.: Complete nucleotide sequence of GCMV RNA-1. No assessment of resistance made.: Up-to-date report on virus diseases of grapevines in Hungary and description of the clean stock programme implemented in the country. Plant Molecular Biology 21. 89-97. Detection of nepoviruses in ligneous grapevine material using IC/PCR. Candresse. Brault et al. Nucleic Acid Research 17. Roberts and Brown: Detection of GCMV in X. Dunez. Le Gall et al. Russo et al.
and A. Martelli G. Robinson.. Nucleotide sequence homologies among RNA species of strains of tomato black ring virus and other nepoviruses. J. Plant Science. V.. Martelli G. J. 3. Journal of General Virology 76. Lehoczky and G. Kertgazdasag 22. Certification scheme for production of virus-free grape propagating material and its results in Hungary. Kuszala and A. 1969. 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. 29-44. 1970. Lehoczky. Studies on the nitrogen metabolism of vines infected with yellow mosaic virus. Vitis 8. Martelli G. A. Le Gall O.. M. Horváth. 1-130. Annals of Applied Biology 71. Extended Abstracts 13th Meeting of ICVG. Devergne.. Torregrosa . 1731-1740. University of California. and S. Sznyegi. and G. E. J. O... 1989. Y. Sarospataki and J. Sarospataki. G. Quacquarelli. ArMV) in the seeds and seedlings of grapevine by ELISA. and D..P.P. University of California. 1984. a serotype ot tomato black ring virus. 402-410. Lázár J. Szonyegi and M. Martelli G. Caractérization d'une souche du virus de la mosaïque jaune crome de la vigne (GCMV) isolée en Hongrie de vignes non panachées. 135-140. Doz B. Dunez. and A. Hungarian chrome mosaic. Prémunition: une competition entre souches faibles et souches sévères pour un voie commune d'expression de symptômes. T.. Bouquet. Division of Agricultural Sciences. J. Extended Abstracts 11th Meeting of ICVG. 49-64. Mali V. Quacquarelli. 1968. Bojnansky.R. Preliminary report on purification and serology of a virus associated with Hungarian grapevines showing macroscopic symptoms of fanleaf and yellow mosaic. S.. Serologische Verwandschaft eines mit dem ungarischen "chrome mosaic" vergesellschafteten Virus mit einem Stamm des "tomato black ring virus". Hungarian chrome mosaic of grapevine and tomato black ring: two similar but unrelated plant viruses. Vanek and V. Delbos and J. Lehoczky J. Division of Agricultural Sciences. Muranyi .J. GFV-YM. Transmission by nematodes of some grapevine viruses occurring in Czecholovakia and Hungary. Acta Phytopathologica Academia Scientiarum Hungaricae 3. Kölber M. Sarospataki. 236-237. Kiserletugyi-Kozlemenyek 64 (1-3).. Hajdú. 58-72. 165-173.. 1972. 1995. Candresse. Agrobacterium-mediated genetic transformation of grapevine somatic embryos and regeneration of transgenic plants expressing the coat protein of grapevine chrome mosaic nepovirus (GCMV). A. Numero hors série. Effect of the grape chrome mosaic and grape fanleaf-yellow mosaic virus infection on the photosynthetical carbon dioxide fixation in vine leaves.P. No. Quacquarelli. Quacquarelli. Kölber M. T. Berkeley. Nucleotide sequence of Hungarian grapevine chrome mosaic nepovirus RNA1. Annales de Phytopathologie. Weinberg und Keller 15. J. Davis 1965. Grapevine virus diseases and clean stock program in Hungary. Lehoczky and A.. Candresse and A.P. Martelli G. 45 . Lehoczky and A. Montreux 1993. Nematodes of the family Longidoridae (Thorne 1935) Meyl 1960 found in Hungarian vineyards and virus transmission trials with Xiphinema index Thorne et Allen.Ser. 172-173. Lehoczky J. L. 185-192.. Annales de Phytopathologie 11. Detection of grapevine chrome mosaic virus in field-grown vines by ELISA. T. Dunez. S. Luntz. Phytopathologia Mediterranea 8. G. The change of pigment and sugar content in the chrome mosaic virus infected leaves of grapevine. G. GCMV. Farkas. 1990. 1972a. 119-126. 103.P. 567-568. Detection of grapevine chrome mosaic virus in naturally infected vines by indexing. A Handbook. with Special Reference to Vitis. Nouvelle évidence d'une parenté sérologique éloignée entre ce virus et celui deas anneaux noirs de la tomate (tBRV). 2000. Lázár J. In: Virus Diseases of Small Fruits and Grapevines. Division of Agricultural Sciences. Detection of some nepoviruses (GFV.. Adelaide 2000.. Martelli G. 505. 1971. with Special Reference to Vitis.C. Lehoczky. Lehoczky and G.. Lázár.. Le Gall O. 206-210. Les Colloques de l'INRA 11. 1-7.M. Brault and J. Proceedings International Conference on Virus and Vector on Perennial Hosts. and G. Phytopathologia Mediterranea 24. Acta Phytopathologica Academia Scientiarum Hungaricae 1. Pol'nohopodarska Veda . Proceedings International Conference on Virus and Vector on Perennial Hosts. CMI/AAB Descriptions of Plant Viruses. Jakó N. Journal of General Virology 65. J. L. 1982. Quacquarelli and J. L. The purification and some properties of cocoa necrosis virus. 1969. NW Frazier (ed. 1966. 1968. Kölber. 1979.H. Phytopathologia Mediterranea 24. Host range and properties of a virus associated with Hungarian grapevines showing macroscopic symptoms of fanleaf and yellow mosaic. 1985. Pacsa and J. J. Jakó N.) University of California... L.J. The effect of fanleaf and chrome mosaic virus diseases on yield and the fruit sugar content of grapevine. 171-170.Dodd S. Nucleic Acid Research 17. Kenten R. Danglot. J. G. 1966. 129-134. 389-401. R. Lehoczky J.P.P.. 7795-7807. Candresse and J. M. Grapevine chrome mosaic virus. Kölber and J. Lehoczky . Le Gall O. Martelli G. Beczner. 1285-1289. Vuittenez. Sarospataki. Mikulás. 123-141. Sarospataki. 169-170.Tasnady. Cardin. 1966. 1994. Lehoczky J. S. Davis 1965. 1975. 1985. 102. 1972b. Pozsár B. 1993. Dunez. Lehoczky.
wt of 2 x 106 daltons and apparent size of c. The scattered distribution of infected vines in the field suggests that the virus is spread primarily by infected propagative material. mol.The virus has no recognized vector. CAB International. Nematode Vectors of Plant Viruses.4 x 106 daltons and apparent size of c. Particles are isometric c. Martelli. GRAPEVINE DEFORMATION VIRUS (GDefV) 1.P. M. Martelli and V. D. GDefV is readily detected by ELISA and PCR using primers designed on the coat protein sequence. Gokalp. 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. Proceedings of the Conference on Excoriosis and Virus Diseases of Grapevine. 5. 149-151. S.. Immunosorben electron microscopy for detecting saptransmissible viruses of grapevine. wt of 1. distantly serologically related with ArMV. Mostar 1977. E. Description Grapevine deformation virus (GDefV) was recovered from Turkish grapevines showing distinct fanleaf-like symptoms. 2002. 1997. 286 pp.. was isolated from a Tunisian grapevine with mild fanleaflike symptoms. wt of 2. Niagara Falls 1980. M.6 x 106 Da and RNA-2. Journal of Plant Pathology 85. including all those known to infect grapevine. is distantly related serologically to ArMV but not to GFLV. References Abou Ghanem-Sabanadzovic N. The genome is bipartite.Russo M. 6. Historical review 1991 Ouertani et al. Digiaro and G. 30 nm in diameter and sediment as three components.. 2.J.. N. Coat protein subunits have a Mr 53 x 103.P. GRAPEVINE TUNISIAN RINGSPOT VIRUS (GTRSV) 1.. 471-475 Cigsar I. Grapevine deformation virus. M (particles contaning a molecule of RNA-2 with Mol.800 nt.P. Hranuelli. A.: A mechanically transmissible virus was recovered by sap inoculation from Tunisian grapevines showing mild fanleaf-like symptoms. De Stradis. The virus sediments as three components: T (empty shells).F Brown. Complete nucleotide sequence of RNA-2 of two Turkish nepoviruses.P. wt of 2.800 nt) and B (particles containing a molecule of RNA-1 with Mol. 335-340 Cigsar I.). Bulletin OEPP/EPPO Bulletin 32. Oxon. showing leaf and cane deformations Cigsar et al. and belongs in the subgroup C of the genus Nepovirus. K. 251-257.: Description and thorough characterization of GDefV. G.3 x 106 Da and a size of 3753 nt. 46 . RNA-1 has a mol. Abou Ghanem-Sabanadzovic et al. Saric A. Martelli. identified as a new species in the subgroup A of the genus Nepovirus. a novel nepovirus from Turkey. Abou Ghanem-Sabanadzovic. 35-41. Virus Genes 30. Proceedings of the 7th Meeting of ICVG. 2005. Description Grapevine Tunisian ringspot virus (GTRSV). Digiaro and G. Boscia and G. The virus belongs in the subgroup A of the genus Nepovirus. Sanitary status of grapevine in south eastern and central Anatolia. and D. GTRSV is serologically unrelated to any of 19 nepoviruses tested. 1980. Investigations on grapevine viruses in Croatia. is not seed-borne and reported only from south-eastern Turkey. : Complete nucleotide sequence of GDefV RNA-2 3. 2003. Savino. Digiaro. Historical review 2002 2003 2005 Cigsar et al.: First isolation by mechanical transmission of an unknown nepovirus from cv. 2. 1977. Sabanadzovic. and T. Martelli. No vector is known and no information is available on the distribution and economic importance of the virus. M. Taylor C.
1978. This strain differs considerably from the English type strain of the virus although is serologically closely related to it. Castellano. Rüdel and B. 88-118. G. Annals of Applied Biology 124. Wetzel.M. CMI/AAB Descriptions of Plant Viruses. Savino. 1992.T. FS4 is a good indicator. 198. 2003. Brückbauer H.3. Brown.6 x 106 (RNA-2). accounting for 29% (component M) and 43% (component B) of the particle weight. D. Extended Abstracts 14th Meeting of ICVG. Molecular characterization of two German Raspberry ringspot virus isolates infecting grapevines and construction of full length infectious clones. Murant: Description of RpBRSV in the CMI/AAB Plant Virus Description series Stellmach and Querfurth: Study of a strain of RpRSV isolated from cv. V. Journal of General Virology 73. The grapevine strain of this virus is serologically very distantly related to the two main serotypes. M. The type strain of RpRSV is transmitted by Longidorus macrosoma but the grapevine strain is transmitted by Paralongidorus maximus. Reustle. Particles are about 30 nm in diameter. 47 . Minafra. have angular outline. Edwards and M. Scottish and English. G.: Sequencing and molecular characterization of two German isolates of RpRSV from grapevine 2003 3.L. Heat therapy of infected grapevines. Altmayer. and differs from the type strain for it often sediments as if it were a single centrifugal component. 1994. No. 1982. Jolly.. Martelli and N. The nucleotide sequence of RNA 2 of raspberry ringspot nepovirus. RNA-2 is 3928 nt in size and contains a single ORF encoding a polypeptide with Mr of 124 kDa.C. Brückbauer: RpRSV can be distinguished from other nepoviruses on the basis of symptoms induced on Vitis idicator plants Blok et al. References Blok V. 2. Description Raspberry ringspot virus (RpRSV) is a nepovirus belonging in subgroup A of this genus. 283-300.6 x 106 (RNA-1) and 1. C. Mayo. D.A. RASPBERRY RINGSPOT VIRUS (RpRSV) 1.J. Two strains of different virulence occur in the Palatinate.: Nucleotide sequence of RpRSV RNA-2 Jones et al. Manoukian. The virus has only been found in grapevine in western Germany. Crop losses can be higher than 30 %. Die Wein-Wissenschaft 37..A.F. wt of 2. W. M. The viral genome is a bipartite positive sense ssRNA.: Recovery of RpRSV from grapevines of Palatinate..: Biological and physico-chemical characterization of the grapevine strain of RpRSV. Ebel R. consisting of two separately encapsidated molecules with Mol. Elbling in West Germany. Properties of an unusual isolate of raspberry ringspot virus from grapevine in Germany and evidence of its possibile transmission by Paralogidorus maximus. Greco. D. Historical review 1970 1978 1978 1982 1992 1994 Vuittenez et al. The coat protein has a single type of subunits with Mr 54 x 103. Murant A. Properties of a previously undescribed grapevine nepovirus from Tunisia. 16. Robinson. 1992. M. Mögliche Beziehungen zwischen Virus und Symptomausprägung bei der Rebe. and sediment as three components (T. Raspberry ringspot virus. A. Boscia. Locorotondo 2003. References Ouertani R. Krczal and T.P.A. G. Jones A. Wardell J. A. 2189-2194. The virus is transmitted by Paralogidorus maximus Ebel et al. M.J.F.J. A Schnabel. Symptoms are similar to those of fanleaf. These differences strongly suggest that the grapevine-infecting RpRSV may be a different viral species. Archives of Virology 126. and B). 107-117. McGavin..
including TBRV : bentonite flocculation test.Stellmach G. Vuittenez et al. Détection et étude selogique du virus latent des taches annulires du frasier (strawberry latent ringspot). ArMV. Weinberg und Keller 25. mottling of the older leaves. M. The virus was detected in grapevines in the Niagara Peninsula. The vector to grapevine is Longidorus attenuatus. SLRV and TBRV found in grapevine in the Palatinate. Joannes Seyve. The latex test is considered as the most sensitive and the least time consuming method. Turkey. The virus is a definitive nepovirus species classified in subgroup A of this genus. TBRV produces a reduction in growth and yield. RNA-1 is 7356 nt in size and contains a single open reading frame encoding a polypeptide with Mr of 254 kDa. RpRV and TBRV detected serologically in grapevine in West Germany. Kuszala. is a strain of this virus. Untersuchungen zur Serologie. Bercks and Querfurth: GFLV.. 1970. Pathologie und Thermo-Labilität mehrerer Reben-Isolate des Himbeerringflecken-Virus (raspberry-ringspot. 279-327 TOMATO BLACK RING VIRUS (TBRV) 1. RNA-2 is 4662 nt in size encoding a polyprotein with Mr of 150 kDa. and an increase in graft failure. Historical review 1963 1965 1966 Stellmach and Bercks: TBRV detected in rootstock Aramon x V. Stellmach and Bercks: Further investigations on TBRV in grapevine. 2. and Ontario (Canada). 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. rings and lines on the leaves of recently infected plants. Rüdel: Transmission of TBRV to grapevine by Longidorus attenuatus. known to cause severe damage to the grapevine variety Joannes Seyve in Ontario. its own subgroup. and G. Tanne: Detection of TBRV by ELISA in Israel. 128-136. wt of 2.7 x 106 (RNA-1) and 1. Lehoczky and Burgyan: Occurrence of TBRV in grapevines in Hungary. Bercks and Stellmach: ArMV. Stellmach: Review paper on TBRV in grapevine. about 30 nm in diameter have angular outline and sediment as three components (T. riparia 143 A in West Germany. Annales de Phytopathologie 2. Israel. J. Joannes Seyve virus. Bercks: Comparison of three serological tests for detecting several viruses. respectively. and B). The genome is a bipartite positive sense single-stranded RNA occurring as two separately encapsidated functional molecules with Mol. Querfurth. then in Yugoslavia.5 x 106 daltons and a size of 1327 nt. Ontario as the cause of severe damage to cv.65 x 106 (RNA-2) accounting for 44% and 37% of the particle weight. TBRV supports the replication of a satellite RNA with Mol. Their coat protein consists of a single type of subunits with Mr of about 57 kDa. wt of 0. Virus particles are isometric. Meyer et al. or directly in grapevine leaf extracts using bentonite flocculation test. M. but they can be high. Rüdel and H Brückbauer. Vuittenez A. Greece. latex test and barium sulfate test. RRV and TBRV are not transmitted by contact of roots or foliage in the vineyard. 1978. 1967 1970 1970b 1976 1977 1980 1984 1984 1986 48 . either by agar gel diffusion with extracts of herbaceous hosts previously infected mechanically from grapevine.: RRV. Description Tomato black ring virus (TBRV) was first found in grapevines in Germany. Losses are not known precisely. Stobbs and Van Schagen: First record of TBRV from Canada. chlorotic spots.virus).: Nucleotide sequence of a TBRV satellite RNA.
1575-1583. Rüdel and H.: SLRSV found in grapevine in Turkey. and G. M. Journal of General Virology 65. Journal of General Virology 69. Kreiah et al. Greif C. Babini and Bertaccini: Electron microscope study SLRSV infections in plant tissues. Nucleotide sequence of tomato black ring virus RNA-1. J. Meyer M.: Nucleotide sequence of SLRSV satellite RNA. Fritsch. O.A. The virus is a definitive species in the genus Sadwavirus.1986 1988 1993 Meyer et al. wt 2. 1988. Détection et étude sérologique du virus latent des taches annulaires du Fraisier (strawberry latent ringspot). RNA-2 is 3824 nt in size and encodes a single ORF expressing a polypetide with Mr of about 99 kDa. Turkiye. Complete nucleotide sequence of a satellite RNA of tomato black ring virus. 279-327. Brückbauer.: SLRSV and other nepoviruses isolated from grapevines in Germany Credi et al. The virus supports the replication of a satellite RNA 1118 nt in size. and B).6 x 106 (RNA-2). References Akbas B. Description Strawberry latent ringspot virus (SLRSV) has been isolated from grapevine in the Palatinate (Germany) and in northern Italy. Mayo and C.. du virus des anneaux noirs de la Tomate (tomato black ring). M. Erdiller. 1984... Occurrence of tomato black ring virus on grapevine in southern Ontario. 55-61. Assignement of SLRSV to the new genus Sadwavirus. and 1. M. 3-5.: SLRSV recorded from grapevine in Italy. Le Gall et al. 2. 1970. respectively. 1993.: Nucleotide sequence of TBRV RNA-1 Abkas and Erdiller: TBRV recorded from grapevines in Turkey 3.G. Hemmer and C. Their coat protein consists of two types of subunits with Mr 43 x 103 and 27 x 103. The nucleotide sequence of tomato black ring virus RNA-2. Hemmer and C. 1257-1271 Stobbs L. O. 1986. 1517-1529. Researches on grapevine virus diseases and determination of their incidence in Ankara.6 x 106 (RNA-1) accounting for 38% of the particle weight. O. Hemmer. Canadian Plant Disease Survey 64. Journal of Turkish Phytopathology 22. Brückbauer: SLRSV can be distinguished from other nepoviruses on the basis of symptoms induced on Vitis idicator plants. and J. Genus SADWAVIRUS STRAWBERRY LATENT RINGSPOT VIRUS (SLRSV) 1. Journal of Gerneal Virology 67. Savino et al. Annales de Phytopathologie 2. et du virus des taches annulaires du Framboisier (raspberry ringspot) chez des vignes du Palatinat. Virus particles are isometric. 49 . Van Schagen. Historical review 1974 1977 1981 1982 1982 1987 1993 1994 2005 Murant: Description of SLRSV in the CMI/AAB Descriptions of Plant vrises series.W. The virus is transmitted by Xiphinema diversicaudatum. Fritsch. Kreiah et al. Kuszala. Fritsch. Symptoms on affected European grapes are of the fanleaf type. 1984. Meyer M. The genome is a bipartite positive sense singlestranded RNA occurring as two separately encapsidated functional molecules with Mol.: Nucleotide sequence of TBRV RNA-2 Greif et al. It was also detected in imported vines in Turkey and Portugal. about 30 nm in diameter have angular outline and sediment as three components (T. Bercks et al. Vuittenez A.: Nucleotide sequence of SLRSV RNA-2.
A.. 1982. Weinberg und Keller 24. M. Brückbauer H.I. D'Onghia and M. U. 126.A. Journal of General Virology. and A.R. Yilmaz. A. L. FAO Plant Protection Bulletin 35... 304-308. 133-180. A. T. Phytopathologische Zeitschrift 104.I. Savino V. Lehto. Strawberry latent ringspot virus.R.M. 1987. Querfurth and M.. G. Le Gall O. H. 102-104. Sanfaçon. 2527-2532.. T. Wellink. Viral aggregates induced by a distinctive strain of strawberry latent ringspot virus from grapevine.A. 1974. References Babini A. L. Die Wein-Wissenschaft 37. Rüdel.. London. 1977. Phytopathologia Mediterranea 20. G.. Iwanami. G. H. 2005. Sequence analysis and location of capsid protein within RNA 2 of strawberry latent ringspot virus. Bertaccini and C. Untersuchungen überdie Viruskrankheited der rebe unter besonderer Berüchsichtigung “atypischer Formen” der Reisigkrankheit. Murant A. Mayo.. Turkey. Bertaccini. Journal of General Virology 75. Babini. Maniloff. J. 74. J. Cooper.V. Mögliche Beziehungen zwischen Virus und Symptomausprägung bei der Rebe.M. 1994. In: Virus Taxonomy.. and Ball. Karasev. Bercks R.P. 88-118. A. 1993. Strunk and J. Eight Report of the International Committee on Taxonomy of Viruses (Fauquet. A distinctive isolate of Strawberry latent ringspot virus from grapevines in Italy. CMI/AAB Descriptions of Plant Viruses No. C. 56-63. 50 . J. 1163-1165. 1982.. Kreiah S. T Jones. Strunk. Genus Sadwavirus. Kreiah S. (eds) 799-802 Elsevier/Academic Press. Strawberry latent ringspot virus in grapevine.F. Credi R.A. Brückbauer. Betti. K.3. Cooper and G. Wetzel and N. Gelli. The nucleotide sequence of a satellite RNA associated with strawberry latent ringspot virus. 1981. Martelli.. Desselberger. Yoshikawa..
All these viruses. V. little grape (Eng. Peach rosette mosaic virus (PRMV) in V. its main host. Partial sequence of the 3' 51 . labrusca. which in blueberry is transmitted by pollen. americanum sensu lato and PRMV by X. Bunches are small and straggly (Maryland's grapevine little berry) and leaves may show chrome yellow flecking along the veins (California's yellow vein). labrusca causes a severe disease characterized by delayed bud burst. Transmission: These viruses are all transmitted by grafting and mechanical inoculation.GRAPEVINE DEGENERATION AND DECLINE (AMERICAN NEPOVIRUSES) 1. americanum sensu stricto. This type of resistance is hypersensitivity. Concord it delays bud burst. PRMV. rupestris plasma show field resistance to the northern US strain of ToRSV and to TRSV and PRMV. labrusca cv. All other viruses are transmitted by longidorid nematodes: Xiphinema americanum sensu stricto and X. exhibiting stunted growth.). positive-sense. labrusca is also resistant to TRSV. mottled (oak leaf pattern. are involved in the aetiology of North American grapevine degeneration and decline. grapevine decline. poor fruit setting. PRMV. V. berlandieri or V. X. Alternative weed hosts that have epidemiological significance are known for ToRSV. In cold climates (e. 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. Description Blueberry leaf mottle virus (BLMoV) is named after the disease induced in highbush blueberry (Vaccinium corymbosum). BLUEBERRY LEAF MOTTLE VIRUS (BLMoV) 1. except for BLMoV which may have been introduced from Europe. depérissement de la vigne (Fr.) Main symptoms: Symptomatological responses of grapevines vary according to the species (i. and poor fruit setting. riparia.15 x 106 (RNA-2). but not conclusive. ToRSV and BLMoV are also seed transmitted in grapes. Blueberry leaf mottle virus (BLMoV) infects latently European grapes. Infected vines decline slowly over time. single-stranded RNA occurring as two separately encapsidated functional molecules with Mol . All roostocks and. TRSV. In warmer climates (Maryland. interspecific hybrids). The genome is a bipartite. elongatus. rivesi transmit ToRSV type strain (decline). induces fanleaf-like symptoms on leaves and canes. Description Main synonyms: Yellow vein. and ToRSV separately or in combination. Nematicidal control of vectors is possible. Varietal susceptibility: There are great variations in the susceptibility of Vitis species and cultivars.g. about 30 nm in diameter with angular outline. ELISA and molecular tools are useful for testing field-infected material. ingiallimento nervale (Ital. V. and poor fruit setting Agent: The above mentioned four distinct nepoviruses. California) yield but not vigour is affected. and/or ring spots) and distorted leaves. No vector is known for BLMoV. are endemic in North America and thought to be native of the region. BLMoV. malformation and mottling of the leaves. TRSV is transmitted by X. TRSV and PRMV. jaunissement des nervures. the infecting virus and the climatic conditions. Adernvergilbung (Germ. straggly and shelled clusters. Control: Use of virus-free propagating material and resistant rootstocks. interestingly. wt of 2. most V. New York State and Ontario) own-rooted European grapes affected by Tomato ringspot virus (ToRSV) and Tobacco ringspot virus (TRSV) decline rapidly.35 x 106 (RNA-1) and 2. BLMoV is a definitive nepovirus species assigned to subgroup C. californicum transmits ToRSV yellow vein strain.). A number of rootstocks containing V. distortion of canes. Vitis vinifera. Longidorus diadecturus and L. Virus particles are isometric.e.). deperimento della vite. Long distance spread takes place primarily through infected propagating material. whereas in V. sedimenting as three components. Their coat protein consists of a single type of subunits with Mr of about 54 x 103 Da.
52 . Isolation and identification of a strain of grapevine Bulgarian latent virus in Concord grapevines in New York State. As the virus is endemic and seed-borne in some perennial weeds Taraxacum officinale (dandelion). Physical and chemical properties of the particles and ribonucleic acid of blueberry leaf mottle virus. The genome is a bipartite positive sense single-stranded RNA occurring as two separately encapsidated functional molecules with Mol. Hancock. Crop losses up to 60% and death of susceptible V. Pollen grains of cv. respectively. Their coat protein consists of a single type of subunits with Mr of about 57 x 103 Da..K. PEACH ROSETTE MOSAIC VIRUS (PRMV) 1.Use of resistant roostock hybrids and of certified planting material is recommended.F.. The virus is seed-transmitted in grapevines and C. 1977. The vector is unknown. PRMV is seed-borne in both naturally infected dandelion (4% of infected seeldlings) and in artificially infected C. E. 1994. Occasional. RNA1 is 8004 nt in size and contains a single open reading frame encoding a polypeptide with Mr of 240 kDa. Grapevines (Vitis labrusca) are infected in New York State (USA) by a serologically distinct strain of the virus. Virus Research 33. Vectors are the Dorylaimoid nematodes Xiphinema americanum sensu lato and Longidorus diadecturus. Vines are stunted and show a progressive decline.5% of seedlings from seeds taken from diseased vines proved to be infected. The virus is transmitted through seeds in grapevines and C. Clusters are straggly. possibily non specific transmission by L. and has no economic importance. about 28 nm in diameter with angular outline.2 x 106 (RNA-2) accounting for 44% and 37% of the particle weight. Virus particles are isometric. 2.W. Infected grapevines show shortened and crooked shoots. Stace-Smith. mottled and variously deformed leaves and delayed bud burst. Plant Disease Reporter 61. 1981. 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. 145-156 Ramsdell D. D. elongatus has also been reported. 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. but in highbush blueberry the virus is pollen-borne and suspected to be pollen-transmitted . quinoa (90% of infected seedlings). 949-953. Phytopathology 71. which may lead to their death. but not eradicate. labrusca cultivars (Concord. sedimenting as three components. PRMV can also be introduced in a site by infected planting material and be spread by vectors to adjacent vines. and with extensive shelling of the berries. Uyemoto J.termini of both RNA molecules has been determined. 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. : Partial nucleotide sequence of BLMoV RNA-1 and RNA-2 1981 1994 3. where the disease occurs in more or less circular patches and spreads slowly.F. Hummer. Description Peach rosette mosaic virus (PRMV) is named after the disease induced in peach. Historical review 1977 Uyemoto et al: BLMoV isolated from New York 'Concord' vines showing fanleaf-like symptoms.K. Sequence analysis of the 3' temini of RNA 1 and RNA 2 of blueberry leaf mottle virus. wt of 2.C. Concord grapes are apparently virus-free but 9. and R. Warkentin. when a vineyard is planted susceptible cultivars may become infected by nematode vectors. D. Taschenberg and D. especially) and a number of American-French hybrids have been recorded.Rasmdel and J. Solanum carolinense (Carolina horse nettle) and Rumex crispus (curly dock). smaller and fewer than normal . Healthy grapevines become infected when planted in soils from diseased vineyards. 468-472. one of its crop plant hosts. References Bacher J. quinoa. PRMV is soil-borne.4 x 106 (RNA-1) and 2. which induces fanleaf-type symptoms and is distantly related to GBLV. mostly to vines adjacent to previously infected plants. vector populations . but identified as a strain of GBLV. The virus is a definitive nepovirus species assigned to subgroup C.
J.G Van Schagen and B. 1984. Historical review 1972a 1972b 1974 1976 Dias: Preliminary characterization of the grapevine isolate of PRMV. Ramsdell and Gillet: List of grapervine cultivars and roostocks showing differential susceptibility to PRMV. Transmission of raspberry ringspot. 1972a.R. Rasmdell et al. 53 .F. 1988. Canadian Journal of Botany 54. officinale. 1228-1239. Numero hors sèrie. J. crispus) and transmission through grapevine seeds. Lammers et al. americanum with the disease. Dias H. Dias and Cation: Biological characterization of the grapevine strain of PRMV.S Everleigh. 1976. The virus is seedborne in C. 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.F. Cation. Ramsdell and Myers: Description of PRMV-induced grapevine degeneration and association of X. Ramsdell: Review article on PRMV. Dias H.R. Numero hors sèrie.F. References Allen W. Canadian Journal of Plant Pathology 4. R. Transmission of peach rosette mosaic virus to peach grape and cucumber by Longidorus diadecturus obtained from diseased orchards in Ontario.. Strains of peach rosette mosaic virus differentiated by cross absorption and immunodiffusion tests. 1972b. 97-103. Canadian Journal of Plant Pathology 6. Purification and some characteristics of peach rosette mosaic virus (grape isolate).: Nucleotide sequence of RNA-1 of the grapevine strain of PRMV. 1982. Transmission of peach rosette mosaic virus by Ontario populations of Longidorus diadecturus and Xiphinema americanum (Nematoda: Longidoridae). quinoa and has reproduced in part the field syndrome when inoculated mechanically to Concord grape seedlings. Dias H. 29-32 Allen W. 105-106. and B. Carolinense.. tomato black ring and peach rosette mosaic viruses by an Ontario population of Longidorus elongatus. Annales de Phytopathologie.G Van Schagen and E..R. Ramsdell et al: Use of ELISA for PRMV detection in grapevines.: Investigation on the susceptibility to PRMV infection of American and European grapevines and hybrid rootstocks. Allen et al.A Ebsary. Annales de Phytopathologie. Allen and Ebsary: Longidorus attenuatus transmits PRMV non specifically and with low efficiency. Ramsdell and Gillet: Description of PRMV in the AAB Descriptions of Plant Viruses. Canadian Journal of Plant Pathology 10. 16-18. and D.. Ramsdell and Myers: Field spread of PRMV is associated with the presence of infected weeds (T. 1978 1979 1980 1982 1983 1984 1985 1988 1988 1995 1998 1999 3.: Xiphinema americanum is an efficient vector of PRMV. 1-5 Allen W. Dias and Allen: Physico-chemical characterization of PRMV.2. Allen et al. S. The characterization of a virus responsible for peach rosette mosaic and grape decline in Michigan.: Longidorus diadecturus transmits PRMV to grapevines.A Ebsary. Dias: Grapevine and peach strains of PRMV can be differentiated serologically.
L.M. Buzayan et al. 74-78.: Nucleotide sequence of TRSV satellite RNA. 1978. Peach rosette mosaic virus. Ramsdell D.C.L.C. TRSV has a relatively wide natural host range. and R. Powell et al. Myers. American Phytopathological Society Press.. Peach rosette mosaic virus.C Ramsdell. TRSV is soil-borne and is transmitted by Xiphinema americanum sensu stricto.M. Bird. Plant Disease 67. sedimenting as three components (T.C. In: Compendium of Grape Diseases (R. AAB Descriptions of Plant Viruses.F Allison and D.C. about 30 nm in diameter with angular outline. Goheen eds. .7 x 106 (RNA-1) and 1. Uyemoto et al. R. Characterization of the single protein and the two nucleic acids of peach rosette mosaic virus. J. and B). Bird GW. Allen. Canadian Journal of Botany 58.: A review of viruses infecting grapevines in New York vineyards. 1980. Peach rosette mosaic virus decline. Ramsdell D. Superimposed shallow and deep soil fumigation to control Xiphinema americanum and peach rosette mosaic virus reinfection a Concord vineyard. Plant Disease 79. and R.C.C.C.C. Virus particles are isometric. Gillet. G.C. Ramsdell D.C. Coat protein consists of a single type of subunits with Mr of about 57 x 103 Da. Lammers A. Gillet and L. Myers. The genome is a bipartite positive sense single-stranded RNA occurring as two separately encapsidated functional molecules with Mol. Ramsdell D.E Morris. R. Epidemiology of peach rosette mosaic virus in a Concord grape vineyard. Relative susceptibility of American. RNA-1 is 7514 nt in size and contains a single open reading frame encoding a polypeptide with Mr of 225 kDa.: Survey of ToRSV and TRSV in Pennsylvanian vineyards. Paul. 364. respectively. Andrews. 54 . American Vitis species reported to be resistant to ToRSV and TRSV.M.). Cloning an sequencing of peach rosette mosaic virus RNA1. Preventive control measures are the use of resistant roostock hybrids and of certified planting material.H.M.C. J.M. TOBACCO RINGSPOT VIRUS (TRSV) 1. Ramsdell D.W. 1988. No. 1174-1178. accounting for 44% and 28% of B and M particle weight. 51-52. Ramsdell D. Phytopathology 68. There is no evidence of seed trasmission in the grapevine. 57-73. Gillet.. M. 154-157. St.. 1747-1754.W. Stace-Smith: Description of TRSV in the AAB Descriptions of Plant Viruses series. symptomatology and nematodes associated with grapevine 'degeneration' in Michigan. and J. wt of 2. Phytopathology 64. Ramsdell D. and J.: TRSV agent of a new grapevine disease in New York State. 1979. respectively. Virus Research 65.M.3 x 106 (RNA-2). 2. French hybrids and European grape cultivars to infection by peach rosette mosaic virus. 4143. Susceptibility of American grapevine scion cultivars and French hybrid roostocks and scion cultivars to infection by peach rosette mosaic virus. Historical review 1970 1977 1985 1985 1986 1990 Gilmer et al. G. 1995.Dias H. Phytopathologia Mediterrenea 24. Pearson and A. but in European grapes responses are similar to those elicited by GFLV. and W. Plant Disease Reporter 63. Gillet and C. Foster and Morris-Krsinich: In vitro translation of TRSV RNA-1 and TRSV RNA-2 yields major polypeptides with Mr of 225K and 116K.W. A comparison between enzyme-linked immunosorbent assay (ELISA) and Chenopodium quinoa for detection of peach rosette mosaic virus in 'Concord' grapevines. 625-627. Description Tobacco ringspot virus (TRSV) is the type species of the genus Nepovirus and the prototype of subgroup A. The virus supports the replication of a circular satellite RNA 359 nt in size.. Rose. RNA-2 has been sequenced only in part. 1985. 1974. 1998. is endemic in Central and Eastern North America.. but was recorded from grapevines only in New York State and Pennsylvania. 447-450 Ramsdell D. 1999. 1983. Gillet and G. Symptoms elicited by TRSV are the same as those of ToRSV in native cultivars.R.
1993 1996 Buckley et al.M.M. Disease named yellow vein. sedimenting as three components (T.8 x 106 (RNA-1) and 2. Virology 151. Infected vines have small.K. 1996. Forer.A. Buzayan and G. Keese and A. ToRSV is soil-borne. about 30 nm in diameter with angular outline. 55 . References Buckley B.R. the infecting virus strain. Vines are stunted and show a progressive rapid decline. J. American Journal of Enology and Viticulture 28. B. Gerlach G. The genome is a bipartite positive sense single-stranded RNA occurring as two separately encapsidated functional molecules with Mol.A. Zalloua P. Bruening. Vines grow vigorously but bear little or no fruit. The virus has been occasionaly recorded from grapevines outside of North America. ToRSV-induced decline affects European cultivars. smaller and fewer than normal. 702-704. W. Zallua et al. Abawi. 1977. wt of 2. P. TOMATO RINGSPOT VIRUS (ToRSV) 1.L. 1990. Uyemoto J. Virus particles are isometric. Tobacco ringspot virus. Phytopathology 60. : Partial nucleotide sequence of TRSV RNA-2. Historical review 1954 1956 1962 Hewitt: Report of an "unfruitful vine" condition in California to which a yellow speckling of the leaves is associated.J. Morris-Krsinich. 619-627. Both RNAs contain a single open reading frame encoding polypeptides with Mr of 244 kDa (RNA-1) and 207 kDa (RNA-2). Coat protein consists of a single type of subunits with Mr of about 58 x 103 Da.4 x 106 (RNA-2) accounting for 44% and 41 % of the particle weight. Virus Research 30.M. The yellow vein strain of the virus is pollen-borne but is not transmitted through seeds. Two serological ToRSV variants are known to infect grapevines. rivesi in north American States and Canada and X. 1993. 1985. AAB Descriptions of Plant Viruses.. Nucleotide sequence of satellite tobacco ringspot virus RNA and its relationship to multimeric forms. Bruening. Virology 219. 1970. 516-519.M. Uyemoto and L. 2. especially if self-rooted. 131-136. Silva and S. Longenecker and L. respectively. Synthesis and processing of the translation products of tobacco ringspot virus in rabbit reticulocyte lysates. interspecific hybrids). Virology 144. V. 186-199.L. 1985. more severely in colder than in warmer climates. Virus and virus-like diseases affecting grapevines in New York vineyards.. and the climatic conditions. Cummins and G.A. "yellow vein" being the characterizing syndrome of its infections. RNA-1 is 8214 nt and RNA-2 is 7273 nt in size. Vectors are the Dorylaimoid nematodes Xiphinema americanum sensu stricto and X. which often leads to death. In California ToRSV affects the yield rather than the vine's growth. J. Nucleotide sequence and in vitro expression of the capsid protein gene of tobacco ringspot virus. Clusters are straggly.. J.S. 1986. A new grapevine disease induced by tobacco ringspot virus. 1-8. Plant Disease 74.M.B. Singh. Powell C.L. contrary to the decline strain which is seed-transmitted. No. Gilmer R. Buzayan J. where it occurs iin the region of the Great Lakes and in the Pacific seaboard from California to British Columbia. Gooding and Hewitt: A mechanically transmissible virus found to be associated with yellow vein.. Chemical cleavage of the 5'-linked protein of tobacco ringspot virus genomic RNAs and characterization of the protein-RNA linkage. vinifera.. californicum in California The virus can be introduced in a site by infected planting material and be spread by vectors to adjacent vines. 335-349. Stace-Smith R. Description Tomato ringspot virus (ToRSV) is a definitive species in the genus Nepovirus and the prototype of subgroup C. J. 309. Kelts. Symptomatological responses vary according to the species (V. mottled and distorted leaves and short internodes.K.: Nucleotide sequence of TRSV RNA-1 3. Foster R. and with extensive shelling of the berries. Incidence of tomato rigspot virus and tobacco ringspot virus in grapevines in Pennsylvania.S.R. and B.. and B). ToRSV has a relatively wide natural host range and is endemic in North America. Gould. labrusca. Hewitt: Successful graft transmission of unfruitful vine condition. Preventive control measures are the use of resistant roostock hybrids and of certified planting material.
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.: Xiphinema rivesi identified as the main vector of ToRSV in Ontario vineyards. Allen et al. Piazzolla et al. Li et al: ToRSV identified in China in grapevine seedlings grown from seeds imported from France. Uyemoto et al. californicum). Corbett and Podleckis: Ultrastructural study of ToRSV-infected grapevine tissues.: Complete nucleotide sequence of ToRSV RNA-2. Gilmer and Uyemoto: ToRSV agent of a decline of Baco noir in New York State. 56 . Dias: Record of ToRSV in the Niagara peninsula. Uyemoto and Gilmer: Spread of ToRSV through the soil of New York State vineyards recorded.: A review of viruses infecting grapevines in New York State vineyards. Bays and Tolin: ToRSV found in grapevines in Virginia Powell et al. Stace-Smith and Ramsdell: Review of nepoviruses of the Americas. Herrera and Madariaga: ToRSV recorded from grapevine in Chile. Stace-Smith: Description of ToRSV in the CMI/AAB Descriptions of Plant Viruses series. americanum (now X. American Vitis species reported to be resistant to ToRSV and TRSV. Uyemoto: Seed transmission of the decline strain of ToRSV. Rott et al. Teliz et al. Martelli and Taylor: Review of nematode-borne viruses and their vectors.: List of grapevine roostocks and cultivars showing differential susceptibility to ToRSV in Canada.: Survey of ToRSV and TRSV in Pennsylvanian vineyards. Allen et al. Bitterlin and Gonslaves: ToRSV retained and transmitted by viruliferous Xiphinema rivesi stored for two years at 1-3°C. Allen and Dias: Physico-chemical characterization of ToRSV. Rowhani et al: Description of sampling strategy for detection of ToRSV. Martelli: Review of nematode-borne viruses of grapevines and their epidemiology.: Confirmation that the grape yellow vein and the the grape decline strains of ToRSV are serological variants of the same virus. Yang et al.: Transmission of the yellow vein strain of ToRSV by X. Gonsalves: ToRSV is irregularly distributed in infected vines but can be detected by ELISA.: ToRSV found in grapevines in Taiwan. Rott et al. Cory and Hewitt: The yellow vein strain of ToRSV is not transmitted through seeds. Baumgartnerova and Subikova: ToRSV recorded form grapevine in Slovakia.: Complete nucleotide sequence of ToRSV RNA-1. Podlekis and Corbett: ToRSV is the agent of little grape disease in Maryland.
Gonsalves.G.G and V. Plant Disease 74. 24-28. 465-473.B.B. Podleckis E. 1992. Phytopathology 53. Van Schagen. Grape yellow vein: symptomatology.K. Canadian Journal of Plant Pathology 4. No. 1963. Transmission of tomato ringspot. 98-101. Association of Xiphinema species with soil types ang grapevines infected with tomato ringspot virus in Ontario. 1978. Plant Disease Reporter 56. Plant Disease 72.H. Bulletin of the California Department of Agriculture 43. and J. Tomato ringspot virus associated with little grape disease of Vidal 256 grapevines. and S. Forer. 1987.G. 1966.A.. 151-189. and E. Nucleotide sequence of tomato ringspot virus RNA 1. Nepoviruses of the Americas. 1169. 57 . Walker and S. 1028-1037.M. 1954.V. 1980. Some virus and virus-like diseases of grapevines. Phytopathology 72. Gooding G.A. Hewitt. Nucleotide sequence of tomato ringspot virus RNA-2. Stobbs. Piazzolla P. R. Tomato ringspot virus. J. Dias. Tolin.C. Phytopathologia Mediterranea 24. Y.. Detection and identification of plant viruses for quarantine in China. 408-411. Nematologia Mediterranea 6.W. Incidence and geographic distribution of tomato ringspot virus in De Chaunac vineyards in the Niagara peninsula. Bitterlin M. 1505-1514. A comparison of Grapevine yellow vein virus and a Grapevine isolate of Tomato ringspot virus.F. Rott M. Some grapevine viruses in pollen and seed.K. 710. 1977. 1972. 861-863. Tomato ringspot virus in "Baco noir" grapevines in New York.F. Canada. N... Plant Disease 71. Spatial distribution of Xiphinema rivesi and persistence of tomato ringspot virus and its vector in soil.. Current Topics in Vector Research 5.K. 196-203. G. Martelli G. A. and V.F.. Gooding G. Grogan and B. Longenecker and L.V. Taylor.. 1989. Stace-Smith R. Corbett M..M. Detection of tomato ringspot virus in grapevines: irregular distribution of virus. Agricultura Tecnica 61. Uyemoto. 1962. 1990. Ramsdell. 275-277. J. and H. Advances in Disease Vector Research 6.3. and W.F. Plant Disease Reporter 61..E. Rott M.K. M. Xiang .P. 658-663. Purification and serology of a virus associated with the grape yellow vein disease. Rochon.V. and M. Subikova. Van Schagen and B. 2004. Chen. Susceptibility of grape cultivars and rootstocks to an Ontario isolate of tomato ringspot virus. 1982.F. Presence and incidence of grapevine viruses in central Chile. peach yellow bud mosaic. Ivanka pri Dunaji 4.A. Herrera M. M. 1988.P. Castellano and D. A severe outbreak of virus-induced grapevine decline in Cascade grapevines in New York. Madariaga. 393-400 Hewitt W. 1-27.C.. 131-166. Sampling strategies for the detection of grapevine fanleaf virus and the grapevine strain of tomato ringspot virus. 1975. Proceedings 15th International Plant Protection Congress. H. Savino. 1995. Phytopathology 58. Phytopathology 56. 1989. American Journal of Enology and Viticulture 13. Phytopathologia Mediterranea 24. Proceedings 7th Meeting of IGVG.G. Li M. 1977.R. Journal of General Virology 76. Lee and D'A. Uyemoto J.W. and B. and D. L.R. 1984. and grape yellow vein viruses by Xiphinema americanum.. Cory L.F. Incidence of tomato ringspot virus in grape in Virginia.E. Tremaine and D'A. References Allen W.V. Rowhani A..B. Membrane-associated spherical particles in extracts and tissues of virus-infected grapevines. Niagara Falls 1980. Rokni.. Canadian Journal of Botany 55. Gonsalves D. and W.S Whei. their epidemiology and control. Vitis 31. Li. Incidence of tomato rigspot virus and tobacco rigspot virus in grapevines in Pennsylvania. 1993. Powell C. 290 Stace-Smith R. Musci. V.J. M. 1985. Nematode-borne viruses of grapevine. 31-34. 133-135. 1991.B. Plant Disease Reporter 59. identification.R. CMI/AAB Descriptions of Plant Viruses. Podleckis. Works of the Institute of Experimental Phytopathology and Entomology .F. Corbett.. J. Rochon. Zhang and Y. Allen W. Téliz D. 2001.L. 475-480. 44-50. Allen W. Phytopathology 79.A. 702-704. Lownsberry. Beijing 2004. 1987. Gilchrist. 663 Martelli G. 35-44. Properties of the single protein and two nucleic acids of tomato ringspot virus. Baumgartnerova H. 1968. Dias and J. L. 157-164. Hewitt. Bays D. Ebsary. Distribution of viruses and their nematode vectors. and C. 13161320 Dias H.A.. 47-64. 1985.E. 1982. and the association of a mechanically transmissible virus with the disease. 95-106. Gilmer R. Journal of General Virology 72.. Identification of tomato rinsgspot virus in leafroll diseased grapevines.
1986. Plant Disease Reporter 56. 504-510.K. Journal of Agricultural Research China 35. Spread of tomato ringspot virus in "Baco Noir" grapevines in New York. Chen. J.R.S.. Abawi. Uyemoto J. and R.Uyemoto J. 131-136. 1972. Deng and M. 1062-1064.C. Yang I. Cummins and G. T. Virus and virus-like diseases affecting grapevines in New York vineyards.L. 1977. American Journal of Enology and Viticulture 28. Sap-transmissiible viruses associated with grapevine yellow mottle disease in Taiwan.J.K.. Gilmer. 58 .M.
Leafroll decreases grapevine yield (by 15-20% in average) and affects negatively rooting ability.). the same as the size of coat protein (CP) subunits.). and GLRsV-9 in the newly established genus Ampelovirus. Agents: To date. Starch accumulates in degenerated chloroplasts causing increased thickness and brittleness of the leaf blades.) Main symptoms: In red-berried cultivars of Vitis vinifera reddish spots develop in the lower leaves in late spring or summer. differentiation of GLRaVs at the species level was by Roman numerals. Sizes deduced from the nucleotide sequence of the CP cistron are 22 kDa for GLRaV-2 . have been found in leafroll-infected vines. most of the leaf surface becomes reddish. depending on the climate and geographic location. whereas GLRaV-7 is presently classified as unassigned species to the family. GLRaV-5. reduction of protein content. potassium depletion in the leaf blade and accumulation in the petioles. Particle length varies from 1400 to 2200 nm according to individual viruses. instead of reddish. in autumn. GLRaV-8. the symptoms are similar. thus suggesting that there can be several causal agents. respectively. Careful observation of field symptoms in infected vines reveals that there are several types of leafroll. and low in sugar. accartocciamento fogliare (Ital.). Description The first descriptions of grapevine leafroll date back to the mid 19th century. the risk of disseminating the disease is great if untested rootstocks are used. The leaf blade becomes thick. called grapevine leafroll-associated viruses (GLRaV). Enrolamento de la folha (Port. GLRaV-4. and is probably the most widespread virus disease of grapevine. but the leaves become chlorotic to yellowish.GRAPEVINE LEAFROLL 1. There are reports of early reddening of grapevine leaves regarded as physiological disorders and referred to as “Rugeau” or “Rossore” in the French and Italian literature.). the whole leaf surface becomes deep purple. The fruits often mature late and irregularly. usually leaving a narrow green band along the primary and secondary veins. All GLRaVs belong in the family Closteroviridae. GLRaV-3. These spots enlarge with time and coalesce so that. whereas the Mr of all other viruses ranges between 35 and 44 kDa. Blattrollkrankheit (Germ. Until 1995. and with many cultivars. Also plant anatomy is affected. infection of rootstocks is symptomless. In the most severe cases. enroulement (Fr. GLRaV-1. thus impairing carbohydrate translocation from foliar parenchymas. Main synonyms: White Emperor disease (Eng. GLRaV-2 CP has a Mr of 24 kDa. These negative effects are reverted if the disease is eliminated by sanitation treatments.5 nm. they are inferior in quantity and quality. vinifera. In white-berried cultivars of V. especially the phloem. GLRaV-6. which are differentiated from one another by a progressive number. Leafroll is no less important than fanleaf in economic importance. These symptoms progress towards the top of the canes as the season advances. In most cases. Its occurrence in viticultural areas is worldwide. except for a variable decrease in vigour. 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. A number of other physiological parameters are affected. brittle and rolls downwards. GLRaV-2 in the genus Closterovirus. 61 . enrollamiento de la hoja. changes in the pattern of peroxidase and polyphenoloxidase isoenzymes. Hence. exhibiting open structure and distinct cross banding with a pitch of about 3. Other such viruses may exist. graft take and plant vigour. Sieve elements are obliterated and crusheds. nine different viruses with filamentous particles. Also the composition and aromatic profile of the musts are modified. accartocciamento.). and lowering of sugar content.e. enrollado (Sp. differing somewhat in aspect and in severity. Rollkrankheit. i. Virus particles are very flexuous filaments about 12 nm wide. as estimated by polyacrylamide gel electrophoresis.
contains 13 ORFs. longispinus. leafroll can be detected by its symptoms in the field on red-fruited varieties. grafted vines) which is largely responsible for its dissemination over medium and long distances. which is the type species of the novel genus Ampelovirus has a genome 17. Mealybug vectors of GLRaV-3 are Planococcus ficus. Membranous vesicles can derive eitrher from peripheral vesiculation of mitochondria followed by disruption of the organelles (GLRaV-1. GLRaVs show molecular variations which give rise to a population of strains. the only member of the group to be mechanically transmissible. 'Cabernet Franc' 'Pinot noir'. climate. calceolariae. broadspectrum. Leaf tissues or petioles from mature symptomatic leaves of V. GLRaVs differ in various vays (molecularly. singlestranded. American Vitis species and rootstocks are the best antigen sources for serological assays. and degenerate primers have been designed and successfully used for PCR detection of virtually all GLRaVs. GLRaV-3. affinis. As to nucleic acid-based assays. GLRaV-2. GLRaV-5 and GLRaV-9 are phylogenetically close to one another but are serologically unrelated. None of the GLRaVs is known to be seedborne. citri. maritimus. American rootstocks are usually symptomless carriers of GLRaVs. the type member of the genus. So far. Symptom expression depends on the variety.919 nt in size. with none of which they are serologically related. This has been ascertained experimentally for GLRaV-1. vinfera and cortical shavings from mature dormant canes of V. and Neopulvinaria innumerabilis. 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. Ps. Polyclonal antisera and/or monoclonal antibodies have been raised to each single GLRaV . Ps. Parthenolecanium corni.647 nt in size and contains 10 major ORFs. number an types of infecting viruses.35 kDa for both GLRaV-3 and GLRaV-1. and epidemiologically) from most of the known closteroviruses. Pseudococcus longispinus. vinifera. Its soft scale insects vectors are Pulvinaria vitis and Neopulvinaria innumerabilis. Indexing on red-fruited cultivars such as 'Cabernet sauvignon'. 'Merlot'. Spread at a site is mediated by mealybug and soft scale insect vectors. GLRaV-1 is transmitted in nature by the pseudococcid mealybugs Heliococcus bohemicus and Phenacoccus aceris and the soft scale insects Pulvinaria vitis. and has structural organization differing from that of other sequenced closteroviruses. soil condition and probably. Pl. in agreement with the quasispecies nature of viruses. These reagents are routinely used for ISEM. GLRaV-3. The presence of high molecular weight double-stranded RNAs (dsRNA) in phloem tissue extracts can be used as infection marker. but it does not discriminate between GLRaVs and was reported to be less sensitive than ELISA. Ps. roostocks. The genome of GLRaV-4 and GLRaV-9 are incompletely sequenced but show a structural organization compatible with that of the genus. comstocki. GLRaV-2 and GLRaV-3. GLRaV-5 and GLRaV-9 are both transmitted by Ps.Transmission is semipersistent and does not appear to be vector-specific. or the hybrid LN 33 is still the most popular method for identifying the disease. 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. classical double antibody sandwich ELISA (Chromo-ELISA) or Lumino-ELISA. Disappearance of dsRNAs from vines submitted to 62 . contains nine open reading frames (ORF) and has structural organization identical to that of Beet yellow virus (BYV). Foliar tissues are not recommended for serological GLRaVs detection in American Vitis species and roostocks. GLRaV-3. viburni and Ps. positive sense RNA molecule. or by molecular techniques. The genome is a monopartite. biologically. has a number of minor biological variants which can be differentiated by the reaction of inoculated Nicotiana species. 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. Regardless of whether they belong to the genus Closterovirus or Amplelovirus. Detection: In many cases. The genome of GLRaV-2 is 15. Ps. 29. vinifera varieties show symptoms most clearly because of the reddening of the leaves. and some of them are used as indicators. Red-berried V. Natural field spread of leafroll disease has been reported from many countries in Europe and elsewhere. and some are commercially avaliable. The genome of GLRaV-1 is 17. Cytopathology: A characterizing feature of all GLRaV infections is the presence of intracellular inclusions in phloem tissues made up of aggregates of virus particles intermingled with single or clustered mebranous vesicles containing finely stranded material thought to be viral RNA. All GLRaVs can be identified by serological and nucleic acid-based techniques. Composite samples should be used to minimize false negative responses that may originate from the unven distribution of GLRaVs in chronically infected vines. Other GLRaVs may exist in nature as suggested by reports.5 kDa for GLRaV-4 . Varietal susceptibility and sensitivity: No immune variety or rootstock is known.528 nt in size. ultrastructurally. only vectors of GLRaV-1. GLRaV-5 and GLRaV-9 have been identified.
No sources of resistance are known in V. Harmon and Snyder: The "White Emperor" disease is graft-transmissible and is regarded a virus disease. roostocks are suggested as the major souces of leafroll dissemination. a grapevine disorder similar to leafroll.: White Emperor and leafroll are identical diseases. Ravaz and Verge: Occurrence in France of “rougeau”. Tanne and Nitzany: Leafroll in Israel Tanne et al. Belli et al. These particles are probably closteroviruses associated with leafroll. Lehoczky et al. unless they are hybridized with virus-specific probes. vinifera and there is no published information on how to protect healthy stocks from vector-mediated reinfection in the field. Control: Production and use of clonally selected and sanitized propagation material is very effective and the only preventive method for leafroll control available.: Leafroll in Hungary. Vuittenez: Leafroll in France. Hewitt: Leafroll in California Goheen et al. a grapevine disorder similar to leafroll Scheu: Demonstration of graft transmission of leafroll from diseased to healthy Vitis vinifera. Introduction of transgenic resistance to GLRaV-2 and GLRaV-3 is being attempted by engineering different viral genes into rootstocks and European grape cultivars. 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.: Leafroll in Italy. Fraser: Leafroll in Australia. Bovey: Leafroll in Switzerland.: Leafroll virus can be inactivated in vivo by heat therapy. Luhn and Goheen: Leafroll found in the original grapevine stocks imported from Europe into California in 1890. Hoefert and Gifford: Study of the effects of leafroll infection on vine anatomy. 1971 1973 1974 1975 63 .: Leafroll in Czechoslovakia. Mendgen: Presence of filamentous particles in grapevines with symptoms of flavescence dorée in West Germany. Later studies showed that the virus is an occasional contaminant Lider et al. Dimitrijevic: Leafroll in Yugoslavia.: Studies on the effects of leafroll on yield of grapevines in California. Blattny et al.sanitation treatments is regarded as evidence for successful virus elimination.: Transmission of a virus to herbaceous plants from a leafroll-infected vine in Israel. Hypothesis of the viral origin of leafroll. 2. dsRNAs cannot be utilized for virus identification. As no apparent spread of the disease was observed. Goheen et al. Chamberlain: Leafroll in New Zealand. Scheu: Leafroll is widespread in German vineyards. However.
1981 1981 1982 1982 1983 1984 1984 1984 1985 1986 1986 1987 1987 1988 1988 1989 64 .: 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. Zimmermann et al. Gugerli et al. Confirmation that GLRaV-3 and the New York closterovirus isolate cross react serologically.: Aggregates of closterovirus-like particles observed in thin sections of phloem from leafroll-diseased grapevines.: First record of successful elimination of leafroll in grapevine by using meristem tip culture in Japan. Mossop et al.: Closterovirus-like particles with an estimated length of 1000 nm found in thin sections of phloem tissue and in leaf dip preparations of leafroll-diseased grapevines in Japan. Production of rabbit and hen antibodies for ELISA and ISEM to GLRaV-1 and GLRaV-3.: Closterovirus-like particles and specific dsRNA found in leafroll-diseased grapevines in New Zealand. Rosciglione and Gugerli: GLRaV-3 is transmitted by the mealybug Planococcus ficus. Suggestion that a closterovirus may be the agent of the disease. but not in similar praparations from healthy plants. are also present in leafroll-affected grapevines from Italy.: ELISA testing reveals that GLRaV-3 has an uneven distribution in grapevine tissues.: Elimination of leafroll by in vitro meristm tip culture and apex fragmentation. Castellano et al. Presence of virus-like particles in thin sections of leafroll-diseased vines.: Ultrastructural study of leafroll-infected grapevine tissues. Von der Brelie and Nienhaus: Light and electron microscope study of cytopathological changes induced by leafroll in grapevines. 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. Namba et al. A large dsRNA molecule is consistently isolated from leafroll-diseased grapes. Barlass at al. Faoro et al. found in grapevines affected by leafroll. Kliever and Lider: Study of biochemical changes found in grapevine infected with leafroll in California. Martelli et al. Tanaka: Leafroll in Japan.: Closterovirus-like particles purified from leafroll-diseased grapevines in France. Zee et al. A third closterovirus type called GLRaV-3. previously found in grapevines in Switzerland. Hu and Gonsalves: Monoclonal antibodies produced against GLRaV-3. Sasahara et al.: Studies on the cytopathology of leafroll-diseased grapevines.like particles. Purification and serology of associated closterovirus-like particles. Production of polyclonal antisera for use in ELISA.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. but not in healthy controls. Abracheva: Leafroll in Bulgaria. Teliz et al. Rosciglione and Gugerli: GLRaV-1 and GLRaV-2 with particles of 2200 nm and 1800 nm respectively. Corbett et al: Electron microscope observations by negative staining of leaf extracts from leafroll-diseased grapevines in South Africa showed the presence of closterovirus.: Review on the detrimental effects of viral infection on grapevine physiology. Absence of such particles in healthy grapevines.
: Use of green grafting for detecting virus-like diseases of grapevine. -2 and -3 are common whereas GLRaV-5 is rarely detected. In Mexico leafroll. was detected in P.: 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. With leafroll. Identification of GLRaV-5. 1990 1991 1991 1991 1991 1991 1991 1991 65 . GLRaV-1.1989 1989 Tanne et al. Some vines indexing positive for leafroll do not react positively with any ofthe antisera. For reliable testing. Agran et al. vinifera varieties used as inoculum source.: Production and characterization of monoclonal antibodies specific to GLRaV-3. There is evidence that other GLRaVs are involved in leafroll etiology. Good results in summer with extracts of basal leaves and in autumn or winter with wood shavings macerated in buffer. GLRaV-2. b Hu et al.: Production of monoclonal antibodies to GLRaV-1 and GLRaV-3. Pseudococcus longispinus is present on weeds around diseased vineyards.: Detection of leafroll-associated closterovirus in recently infected grapevines in New York.: Leafroll and associated closteroviruses in Chile Kuhn: Leafroll in Brasil Li et al. Téliz et al.: Immunocytological detection and localization of GLRaV-1 and GLRaV-3 by immunogold labelling in grapevine thin sections. but not GLRaV-1. Gugerli et al. Heat therapy requires very long treatments and is only 20-30 % successful. The virus was detected in root tissues. Credi and Santucci: GLRaV-1 and GLRaV-3 cannot be detected by direct ELISA in leaves of graft-inoculated American rootstock. Zimmermann et al. Auger et al. symptoms are obtained within 20-70 days.: Evidence of the irregular distribution of GLRaV-3 in American rootstocks. rupestris plasma. 1989 1989 1989 1990 1990 1990a. ELISA is to be applied to cortical scrapings rather than leaf tissues.: Characterization of leafroll-associated closterovirus-like particles from grapevine using also 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. Boscia et al. Faoro et al. stem pitting and corky bark spread rapidly. later in the leaves. Identification of GLRaV-4 1990 1990 1990 1990 Walter et al. GLRaV-1 and GLRaV-2 were not transmitted. ficus fed on infected vines. whereas meristem tip culture yields up to 100 % sanitation Walter and Zimmermann: Further characterization of closteroviruses associated with leafroll in France. indicating the presence of other leafroll-associated viruses. Gugerli et al.: Transmission of GLRaV-3 from grapevine to grapevine by the mealybug Pseudococcus longispinus in Israel.: Comparison of heat therapy and meristem tip culture for eliminating GLRaV-3 from Italian grape varieties. Savino et al. Transmission of GLRaV-3 by the mealybug Planococcus ficus. but they are easily detected in inoculated LN33 vines and in V. Gugerli: Review of grapevine closteroviruses. especially those containing V.: Further characterization of GLRaV-1 and GLRaV-3 by monoclonal antibodies.
Leafroll and associated closteroviruses in Yemen.: Production of antisera to GLRaVs using electrophoretically separated coat protein subunits as antigens.: Purification and physico-chemical characterization of grapevine corky bark associated virus. Golino et al. Castellano et al.: Leafroll in Taiwan. Observation of peripherically vesiculated mitochondria.: Transmission of GLRaV-3 by Pseudococcus affinis in California. Minafra and Hadidi: Detection of GLRaV-3 in viruliferous mealybugs by PCR.1991 Hu et al. Tzeng et al. Habili et al.: Analysis for the presence of double-stranded RNAs can be used for assessing virus elimination following sanitation treatments.b Saldarelli et al. ELISA is recommended for large screening.: Leafroll and associated closteroviruses in Albania. denoted GLRaV 2a and GLRaV 2b. Gozsczynski et al.: Leafroll and associated closteroviruses in Spain 1991 1991 1991 1991 1991 1991 1992 1993 1993 1993 1993 1993 1993 1994a.: Comparison of different assay methods for detecting GLRaVs : ELISA. Boehm and Martins: Leafroll in Portugal. ISEM and dsRNA analysis. Pop et al. Boscia et al. 1994 1994 1994 1994 1994 1995 1994 1995 1995 1995 1995 Merkuri et al. Milkus et al. Belli et al.: Mechanical transmission of GLRaV-2 and ultrastructural study of infected tissues of Nicotiana benthamiana. Kassemeyer: Detection of GLRaVs in Germany. Gugerli and Ramel: Analysis by monolconal antibodies of a Swiss source of cv. Segura et al.: Leafroll and associated closteroviruses in Romania. Martelli et al.: Production of radioactive and non-radioactive molecular probes to GLRaV-3 from denatured dsRNA template and their use for virus identification.1% in 1992 Ioannou: Leafroll and natural spread of associated closteroviruses in Cyprus.: Leafroll and associated closteroviruses in Ukraine. Bondarchuk et al. Flak and Gangl: Leafroll and associated closteroviruses in Austria. Former GLRaV 2 is re-named GLRaV-6.: Transmission of GLRaV-3 by the soft scale insect Pulvinaria vitis. Faoro and Carzaniga: Ustrastructural study of GLRaV-1 and GLRV-3 infections. whereas the other assays are more suitable for analyzing samples that gave inconclusive results with ELISA.: Leafroll and associated closteroviruses in Moldova.1% in 1988 to 93. Chasselas shows the prsence of two different GLRaV-2. Jordan: In a New Zealand commercial vineyard GLRaV-3 incidence increased from 9.: Revision of the nomenclature of GLRaVs and use of Arabic numerals in the species names. 66 . Namba et al. Katis et al: Leafroll and associated closteroviruses in Greece. later identified as GLRaV-2. Krake: Characterization of leafroll disease based on symptoms shown by field-infected vines and graft-transmission tests.
: Cloning an sequencing of the coat protein gene of GLRaV-3 and its expression in transgenic tobacco.: GLRaV-3 detected in native American vines in Western New York. Routh et al. Haidar et al. Krastanova et al. Lahogue and Boulard: Search for genes of resistance in grapevines.: Association of GLRaV-2 in Italy and France with a graft incompatibility revealed by Kober 5BB. Choueiri et al. Martelli et al. 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1998 1998 1998 1998 1998 1998 1998 67 .: GLRaV-3 is transmitted by Planococcus citri in a semipersisten manner. None of 223 accessions of European.: Leafroll and associated closteroviruses in Palestine. Cabaleiro et al.: Identification of GLRaV-7 and production of a polyclonal antiserum.1995 1996 1996 1996 1996 1996 Greif et al. Gugerli et al.9% in 1996.: Leafroll and associated closteroviruses in Lebanon.: Serological characterization of GLRaV-6 and production of monoclonal antibodies.: Elimination of GLRaV-3 by heat therapy improves agronomic performances of a Nebbiolo clone and the quality of the must. Alkowni et al. Guidoni et al.1% in 1986 to 51. Fortusini et al. Gozsczynski et al.: The sequenced genome of GLRaV-2 has the same structural organization of that of Beet yellows virus.GLRaV-5 induces mitochondrial vesiculation. Wilcox et al.: GLRaV-2 and GLRaV-3 genes engineered in grapevine rootstocks to induce resistance. Habili and Nutter: In an Australian commercial vineyard GLRaV-3 incidence increased from 23. Zhu et al.: Distribution and incidence of GLRaVs in Canadian viticultural districts. Faoro: Comprehensive review of the ultrastructure of GLRaVs infections.: Development of a spot-PCR technique for GLRaVs identification. GLRaV-3 appears to be a typical closterovirus.: Use of degenerate primers for PCR detection of GLRaV-4 and GLRaV-5. No vector was identified.: Identification of two different mechanically transmissibile strains of GLRaV-2. Petersen and Charles: Pseudococcus calceolariae acts as vector of GLRaV-3 in New Zealand. Al-Tamimi et al. MacKenzie et al. Rowhani and Uyemoto: Comparative trials between indexing and laboratory detection methods show that the latter are more sensitve for GLRaVs detection. American. La Notte et al.: Comprehensive review of the properties of GLRaVs. the type specied of the genus Closterovirus. Ling et al.: Extensive sequencing of GLRaV-3 genome. and Asian Vitis species inoculated by green grafting with a GLRaV-1 and GLRaV-3 sources were resistant.: Leafroll and associated closteroviruses in Jordan.: Transmission of GLRaV-1 by the soft scale insects Parthenolecanium corni and Neopuvinaria innumerabilis. Viruses are irregularly distributed in the vines. Ling et al.
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M. 1987. Sela and I. D. 356358. Komar and C.. H. Non radioactive molecular probes for the detection of three filamentous viruses of the grapevine. Phytopathologische Zeitschrift 80.E. Saldarelli. Savino and G. Martelli. 1982a. European Journal of Plant Pathology 104. Extended Abstracts 14th Meeting of ICVG. Le rugeau de la vigne. Haidar. 192-196. Presence of grapevine leafroll in North West Spain.. P. Gonsalves. Golino. Volos 1990. Jacquet. Savino V. Martelli and B. Kiryat Anavim 1987. 35-38. Y. 1998. M. Nitzany. K. Rowhani.. A..L. Partial molecular characterization and RT-PCR detection of grapevine leafroll-associated virus 7. Arboriculture. D'Onghia and G. C.A. Vitis 33. Martelli.. 1993..D. Digiaro.J. D.S. Vitis 12. with special reference to closteroand vitiviruses of the grapevine. Reichnährstand Verlag. 157-160. and P. 162-163. Tada. Extended Abstracts 13th Meeting of ICVG.Ravaz L. 2000. Zee. 345-346. Rosciglione B. Proceedings 10th Meeting of ICVG.. Tanne E. Cloquemin and B. Tzeng H.S. Zeitschrift für Pflanzenkrankheiten und Pflanzenschutz 89. Minafra. Plant Pathology 43. 704-709. Progrès Agricole et Viticole 45. Uyemoto. Annals of the Phytopathological Society of Japan 42. 2000. 207211. Sannino F. Saldarelli P. 1981.. Detection of grapevine leafroll-associated closterovirus III by molecular hybridization. Zhang. Greif. Gonzales and C. 1974. Saldarelli P. Proceedings 9th Meeting of ICVG. V. New scale insect vectors of grapevine closteroviruses. 1924. Scheu G. Zhang . Indexing grapes in Japan for viruses. 799-801. 17-18. Tanne E. and J. 1994a.. Saldarelli and A.. Ben-Dov and B. M. A comparison between serological and biological assays in detecting grapevine leafroll-associated viruses. 91-96. Il rossore delle viti. Rott. Chen and D. Tanaka S. W. Rowhani A. Phytopathology 88. Extended Abstracts 11th Meeting of ICVG. P. and J. Uyemoto. 110-113. A. Von der Brelie D.P. Sasahara H. 222-223. V. Use of degenerate primers in a RT-PCR assay for the identification and analysis of some filamentous viruses.P. Segura A. G. 176-180. Rosciglione B. 8689. Berlin. Kiryat Anavim 1987. D. 1976. Plant Disease 71.K. 1994. and F. Teliz D. A. Phytoparasitica 17. 508-517. I.P. Greif.. Der Deutsche Weinbau 14. 222-225 Tanne E. Y. 1994b. Journal of the Japanese Society for Horticultural Science 50. 82. Rowhani A. 1935. Histological and cytological studies on the infectious leafroll disease of the grapevine.P.. Y. 945-950. Tanne. 169175. M.M.. and F. Raccah. Rivista di Patologia Vegetale 1. Verge. and G. Guglielmi Montano and G. Minafra. 67-69. D. Tzeng. A. J. Preliminary investigations of genetic variability of Grapeviune leafroll associated virus 3 isolates.P. Cabaleiro. Sforza R. Iri. 68-69. Virus diseses of grapevine in Israel. 1991. Harpaz. Adelaide 2000.. M. Mein Winzerbuch. 1973. The occurrence of grapevine leafroll disease among the main grapevine cultivars and breeding stocks in Taiwan. 71-73.K. Boscia. 74 . G.C. M. 1989 Detection of a grapevine leafroll-associated closterovirus in recently infected tissues in New York and spread of the disease in Mexico. Tazaki. Field serological detection of viral antigens associated with grapevine leafroll disease. Rowhani. Transmission of grapevine leafroll virus to herbaceous plants. Turturo C. Walter. Minafra and M. 433-436. Routh G. Plant Disease 81. Martelli. Saldarelli P. and P. A. Martelli. M. Horticulture 18. 2000. Locorotondo 2003.P. 38. Effect of heat therapy and meristem tip culture on the elimination of grapevine leafroll-associated closterovirus type III. C. Seddas A. 2000..K. Gugerli. Extended Abstracts 13th Meeting of ICVG. Transmission of grapevine leafroll disease and an associated closterovirus to healthy grapevine by the mealybug Planococcus ficus Signoret.. 1936. Jelkmann and G. Scheu G. Walter. Adelaide 2000. 156-167. P. Regeneration of plantlets by meristem tip culture for virus-free grapevine.. Plant Pathology 49. 1906. Turturo C. Takezawa and M. Hummer. Teliz D.A . Proceedings 9th Meeting of ICVG. Use of degenerate primers for partial sequencing and RT-PCR-based assays of grapevine leafroll-associated viruses. Plant Pathology Bulletin 3. Adelaide 2000. Die Rollkrankheit des Rebstockes. D. Nienhaus. 1998. 1238-1243.. Revue Suisse de Viticulture. Digiaro. Yafeng. 1997. 14.L.. 135-141. 125-126.. E. Extended Abstracts 13th Meeting of ICVG. 1986. 1989. 80-85. Gonsalves and F. Isolation and partial characterization of two new viruses from grapevine. Transmission of closterolike particles associated with grapevine leafroll by mealybugs (Pseudoccidae) in Israel. Establishment of a relationship between grapevine leafroll closteroviruses 1 and 3 by use of monoclonal antibodies. Saldarelli. Maladies de l'enroulement et du bois strié de la vigne: analyse microscopique et sérologique. Gugerli 1989. Hu and D. Montreux 1993. T.P. 11-17..
2000.. 1998 Leafroll virus is common in cultivated American grapevines in Western New York.. and D. Gonsalves. 1987. 313-316. Zimmermann D. J. McFerson and D. Journal of Phytopathology 128. Adelaide 2000. Further characterization of closterovirus-like particles associated with the grapevine leafroll disease. A. Boscia. 130. 1990. Vesselle.. Wilcox F. 62-66. Lee. D. Bass. Monoclonal antibodies for detection and characterization of Grapevine leafroll-associated virus 2. The use of a greengrafting technique for the detection of virus-like diseases of the grapevine. P.P. Walter and M.S. Ling. N. R. Martelli. Locorotondo 2003.Y. 1990. Z. P. D. Martin. D.R.. Van Regenmortel. the closterovirus type member.V. D. Production and characterization of monoclonal antibodies specific for Grapevine leafroll-associated virus 3 and epitope mapping of the coat protein. 1958. Potere. Volos 1990..W. Saldarelli.. 1289-1298. Comptes Rendues de l'Academie d'Agriculture de France 44.. Proceedings 10th Meeting of ICVG. 1427-1434. B. Zhou Z. 137-145. Jiang and D. D. Extended Abstracts 13th Meeting of ICVG. Kim. R. 203. Goheen. Purification de particules virales associées à l'enroulement de la vigne et mise au point d'un protocole ELISA permettant leur détection. Gonsalves. C. Collas and G. 731-741. Production and characterization of monoclonal antibodies specific to closterovirus-like particles associated with grapevine leafroll disease. O. Plant Disease 82. Gonsalves. Journal of Phytopathology 130. K. Le Gall. Zimmermann. Abou-Ghanem. G. Turturo. 2003. R.E Goszczynski. 277-288. Sommermeyer. Walter B. C. 1062.A. Agronomie 8.Y. 75 . Pool and R. 1998. Extended Abstracts 14th Meeting of ICVG. and G. A. Boscia. Zimmermann D. O. 1988. Zee F. Cytopathology of leafroll diseased grapevines and the purification and serology of associated closteroviruslike particles. Zhou Z. Journal of General Virology 79.H. Walter B. Transmission par greffage d'une virose type enroulement foliarie commune dans le vignobles de l'Est e du Centre-Est de la France. B. 1991. Nucleotide sequence and genome organization of Grapevine leafroll-associated virus 2 are similar to Beet yellows virus. Walter and O. Phytopathology 77.Vuittenez A.F. Vernoy.E. Goszczynski and M. Zhu H..S. Potere. Legin. K. Castellano.
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
Individual viruses can be identified by ELISA or dot immunobinding on nylon membranes using polyclonal antisera and/or monoclonal antibodies when available. 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. The best antigen sources for serological diagnosis are cortical shavings from mature dormant canes. Latent infection can occur also in grafted vines. as shown by 110R. Beukman and Goheen: Brief account of the histological modifications of corky bark-affected LN 33. if not otherwise associated with a specific syndrome. GVA can be eliminated to a very hight rate (up to 97%) by the procedure used for cryopreservation of grapevine shoot tips. meristem tip culture.were observed in self-rooted cultivars and ungrafted roostock stocks (V. Historical review Names like "legno riccio". Goheen et al. Thus. or spot-RT-PCR using degenerate or virus-specific primers. experimental evidence has been obtained of the very close association of GRSPaV. Graniti and Ciccarone: First record of rugose wood from southern Italy. since symptomless infections make sanitary selection not totally reliable. immunocapture RT-PCR. Martelli et al.: Graft transmission of rough bark to LN 33. a virus-like disease. the putative agent of rupestris stem pitting. 1965 1965 1967 81 . Name of the disease changed into corky bark. with vein necrosis. rupestris. rupestris and Kober 5BB). Graniti and Martelli: Demonstration of the infectious nature of rugose wood. it is plausible to regard 110R as a specific indicator of rupestris stem pitting in addition to V. Immuno-capture RT-PCR is 1000-fold more sensitive than ELISA for virus detection in grapevines. The intensity of wood abnormalities (pitting and grooving) vary. possibly in relation with the scion/stock combination and climatic conditions. Detection: Indexing on indicators (V. Suggestion that rugose wood may be a disease of combination requiring the contact of scion and rootstock for the development of symptoms. Suggestion that it may be caused by a virus. Transgene expression was detected in these plants and in transformed grapevine explants. 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. no strategy has yet been developed for the chemical control of vectors. are synonymized with "rugose wood". but not foveaviruses. several resistant plant lines were obtained by transformation with the coat protein and the movement protein genes of GVA and GVB. Hewitt et al. In addition. Recently. though with difficulty. rupestris. In general. "stem pitting" and "stem grooving". Kober 5BB and LN 33) is the only reliable method for detecting and sorting out the diseases of the complex. Thus. Histological study of diseased vines. described from California. 2. Vitiviruses. However. all sources must be indexed and/or laboratory tested. to a restricted range of herbaceous hosts (mostly Nicotiana species).: Corky bark is remarkably heat stable and difficult to eliminate by heat therapy. other assays include: single step or nested reverse trascription-polymerase chain reaction (RT-PCR). 1954 1961 1962 1963 1963 1964 1965 Hewitt: Rough bark.: First record of rugose wood outside of Italy. Using a Nicotiana benthamiana model system. or a combination of the two. Goidanich and Canova: First record of corky bark in Europe. are mechanically transmissible. rugose wood agents can be eliminated with reasonable efficiency by heat therapy. and that it may be a composite disease resulting from the interaction of different viruses among which GFLV. Graniti: Detailed description of rugose wood symptoms. Faccioli: First histological study of corky bark-affected grapevines. Control of mealybugs is difficult for they overwinter under the bark of grapevines and possess an unwettable waxy covering.
: Observation of rugose wood symptoms in self-rooted vines. Hewitt: Up-to-date review on grapevine virus and virus-like disease worldwide. Goheen: Evidence that corky bark and leafroll. Hewitt: Successful graft transmission of Californian rugose wood.: Green grafting useful for corky bark indexing.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. Anonymous: A review of rugose wood in Italy. 1968 1969 1970 1970 1971 1971 1972 1973 1973 1975 1975 1977 1978 1979 1979 1980 1980 1981 1981 1982 1983 82 . Sarooshi et al. Conti et al.: First experimental evidence that a filamentous virus (GVA). Lehoczky: Destructive effects of rugose wood registered in Hungary in both self-rooted and grafted European grape varieties. No nepoviruses implicated in their etiology.Virus provisionally called grapevine stem pitting-associated virus (GSP-AV). despite similarities in the s y m p t o m s o n t h e foliage are different diseases. Rosciglione et al. a. based on graft transmission tests. Legin et al.: Recovery by mechanical inoculation of a closterovirus with particles 800 nm long. Hewitt and Neja: First record of rugose wood in USA (California). Graniti and Martelli: Up-to-date review of rugose wood.b. Mink and Parsons: Use of growth chambers for rapid symptom expression of corky bark in Vitis indicators. Bovey and Brugger: Further evidence that GFLV may not be implicated in the etiology of rugose wood in Switzerland. Rugose wood may not require a grafted plant for full symptom expression. is transmitted by the pseudoccid mealybug Pseudococcus longispinus. Engelbrecht and Nel: Rugose wood and fanleaf are not related. Goheen and Luhn: Heat treatment of dormant buds grafted onto LN 33 is effective against corky bark. At 38 °C the minimum inactivation period for leafroll is 56 days and for corky bark 98 days. Goheen and Luhn: Suggestion that corky bark and rugose wood are the same disease. First record of rugose wood symptoms outside of Europe (Israel). Castillo et al. Beukman and Goheen: Up-to-date review of corky bark.1968 1968 Lehoczky et al. from a rugose wood-infected vine. Beukman and Gifford: Detailed account of adverse effects of corky bark on the anatomy of Vitis.: Rugose wood recorded from Australia. Teliz et al. Boccardo and D'Aquilio: Physicochemical characterization of GSP-AV Abracheva: Survey of over 650 grapevine cultivars and hybrids for rugose wood reaction in Bulgaria.: Heat therapy effective against rugose wood.
: Two distinct dsRNAs with a mol.: Transmission of corky bark by the mealybug P. 1985 1985 1985 1985 1985a 1985b 1985 1985 1985 1987 1988 1989 1989 1989 1989 1989 1989 1990 1990 1991 1991 83 . Italia propagated on six different rootstocks.ficus. Corbett and Wiid: Closterovirus-like particles found in extracts from vines affected by corky bark and rugose wood in South Africa.: First record of rugose wood from China. Engelbrecht et al.3 and 4.: Experimental confirmation that rugose wood may not express symptoms in grafted indicators. No closterovirus-like particles in samples with rupestris stem pitting. an additional disease of the rugose wood complex. Savino et al. Kuniyuki and Costa: Rugose wood recorded from Brasil Goheen: First published description of rupestris stem pitting. Rugose wood and corky bark are not the same disease. Tanne et al.: Assessment of crop losses induced by rugose wood to two different European grape varieties.: Heracleum latent virus and GVA are distantly serologically related. Monette et al. Gallitelli et al. Murant et al. ficus. Monette and James: Detection of two biologically distinct but serologically indistinguishable isolates of GVA.: Three types of wood disorders of the stem-grooving type observed in South African grapevines. but not consistently in grapevines from New York. Li et al. Similar dsRNA species were detected. Garau et al. ficus.: First indication of the possible existence of LN 33 stem grooving.: A low molecular weight dsRNA associated with rupestris stem pitting.: Evidence that GSP-AV can occur in grapevines together with another similar but serologically unrelated virus with short closterovirus-like particles. Azzam et al. Corky bark and Rupestris stem pitting.4 x 106 associated with rupestris stem pitting in grapevines from California and Canada. thesis describing rupestris stem pitting disease in comparison with corky bark. Martelli: Rugose wood recorded in southern Mediterranean and Arab countries. Suggestion that the disease is not related to closteroviruses associated with GLRaV and corky bark. similar to Kober stem grooving. Savino et al. wt of 5. Castrovilli and Gallitelli: Physicochemical comparison of two Italian isolates of GVA.: Evaluation of the effect of rugose wood on cv. Rosciglione and Castellano: Demonstration that GVA is transmitted also by Planococcus citri and P. GSP-AV re-named Grapevine virus A (GVA). Savino et al.Sc. First report of Kober stem grooving. Prudencio: M. Garau et al.1984 Milne et al. The first two disorders appear to be spreading in the vineyards. Engelbrecht and Kasdorf: Natural field spread of corky bark in South Africa associated with the presence of P.: Application of spot hybridization for the detection of GVA in grapevine sap. denoted Grapevine virus B (GVB).: Experimental confirmation of the complex nature of rugose wood based on the differential reaction of woody indicators.
Virus named Grapevine virus C (GVC). Virus transmission by the mealybug Ps.: GVA and Kober stem grooving are closely associated. Garau et al. Minafra and Hadidi: Detection of GVA and GVB in viruliferous mealybugs by PCR. Padilla: Rugose wood recorded from Spain Boscia et al. Monette and James: A closterovirus with short particles (725 nm) isolated from a corky barkaffected vine induces necrotic local lesions and systemic symptoms in Nicotiana benthamiana.: A closterovirus with particles 1440-2000 nm long serologically unrelated to all other known grapevine closteroviruses found in corky bark-affected vines. Garau et al. Martelli et al.: Rugose wood recorded from Yemen. both associated with a stem pitting condition of grapevines rather than with leafroll. Thorough comparative study of nine GVB isolates from different countries.1991 1991 Gugerli et al. Merkuri et al. Suggestion that GVA may be the causal agent of the disease.: Clear-cut connection of GVA and rugose wood. 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. Digiaro et al.: Sequence of the 3' end of GVA and GVB genome. Saldarelli et al.: Contemporary occurrence of Rupestris stem pitting and Kober stem grooving in symptomless scions of cv. Minafra et al.: Presence of two distinct serotypes of GVA.: Development and diagnostic use of a cloned probe to GVB.: Synthesis of a cloned probe for GVA. Boulila et al. Namba et al. 1991 1991 1991 1991 1991 1991 1991 1991 1992 1992 1993 1993 1993 1993 1994 1994 1994 1994 1994 1994 1994 1994 1995 84 . Saric and Korosec-Koruza: Rugose wood recorded from Croatia and Slovenia Ioannou: Rugose wood recorded from Cyprus. Boscia et al. ficus induced corky bark symptoms in LN 33 Saldarelli et al.: Rugose wood recorded from Malta Monette and Godkin: Recovery of a closterovirus-like virus by mechanical inoculation from a corky bark-affected vine.: Development of digoxigenin-labelled riboprobes for the detection of GVA and GVB in infected tissue extracts. Torbato in Italy.:Rugose wood recorded from Tunisia Milkus et al. Minafra et al. Both viruses qualify for the inclusion in the genus Trichovirus. Suggestion that GVA is implicated in the aetiology of the disease. Martelli et al.: Rugose wood recorded from Ukraine Boscia et al.: Rugose wood recorded from Albania. Chavez and Varon de Agudelo: Rugose wood recorded from Colombia.: Purification and properties of GVB.: Production of monoclonal antibodies to GVA and their use for ELISA detection of the virus in infected vines.
: Sequence and structrural organization of Grapevine rupestris stem pittingassociated virus genome (GRSPaV). Saldarelli et al. Haidar et al.: Antiserum to the movement protein of GVA is useful for virus detection in ELISA. Boscia et al. Bonavia et al. Martelli et al. Further support of the cause-effect relationship between GVA and this disease. Faoro: Review of the cytopathology of grapevine trichovirus infections. GVC. Since particle size (600-700 nm in length) is compatible with that of Grapevine rupestris stem pitting-associated virus (GRSPaV) particles identified in 2002. longispinus in a semi-persistent manner. Garau et al.: GVB is consistently associated with corky bark and is present. though not consistently in vines showing a syndrome denoted "corky rugose wood".: GVA and GVB are serologically related.: A study of the spatial distribution pattern of corky bark in aThompson seedless vineyard in Israel. Boscia et al. La Notte et al.1995 Monette and Godkin: Detection of non mechanically transmissible capillovirus-like particles in a grapevine affected by rugose wood. The virus is not seed-borne. GVB.: GVA and GVB are transmitted by Pseudococcus affinis. GRSPaV is assigned to this genus.: Nucleotide sequence of GVB genome.: Rugose wood recorded from Lebanon. Alkowni et al: Rugose wood in Palestine. Goszczynski et al. and GVD) later assigned to the genus Vitivirus. Choueiri et al.: Sequencing of a Californian isolate of GRSPaV. and GVD removed from the genus Trichovirus and assigned to the new genus. Rubinson et al. Abou Ghanem et al. Guidoni et al.: Review of the properties of putative grapevine-infecting trichoviruses (GVA.: Consistent detection of GVA in Kober stem grooving-infected grapevines by immunocapture-polymerase chain reaction. GVA. Martelli and Jelkmann: Establishment of the genus Foveavirus.: Estasblishment of the genus Vitivirus with GVA as type species.: GVA and GVD are serologically distantly related. Efficient detection method based on TAS-ELISA developed.: Development of a PCR technique for the detection of GVA and GVB in nylon membrane-spotted sap.: Description of Grapevine virus D (GVD). Minafra et al.: GVA is transmitted by by Ps. Tanne et al. Suggestion that spreading is by a vector that transmits in a semipersistent manner. GVC.: Elimination of GVA from cv Nebbiolo clones by heat therapy improves agronomic performance of the vines and quality of the must. this may be the first visualization of GRSPaV. Meng et al. La Notte et al. 1995 1995 1995 1996 1996 1996 1996 1996 1997 1997 1997 1997 1997a 1997b 1997 1997 1997 1997 1998 1998 1998 1998 85 . Zhang et al.: Nucleotide sequence of GVA genome and taxonomic position of the virus. GVB. Chevalier et al.: Rugose wood recorded from Jordan.
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.: The function of GVA genes identified by mutation analysis of individual ORFs of a full-length infectious viral clone. GCD) and foveavirus (GRSPaV) sequences in two steps.8% and 78-89.: GVA particles carry a highly structured epitope centered on a common peptide region of the coat protein sequence. Dovas and Katis: Improved RT-PCR method for the simultanous detection in grapevine extracts of vitivirus (GVA. The virus was not detected in 245 seedlings from infected cv. Virus detected in bleached seeds suggesting that it is present inside the seeds. observed for the first time.: 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. GVB and Corky bark and GRSPaV and Rupestris stem pitting.e. are filamentous and measure 723 nm in length. GVB.1999 1999 2000a Meng et al. GVA coat protein encapsidating GVB RNA. Goszczynski and Jooste: Thre groups of GVA strains (I. Petrovic et al. Saldarelli et al. Galiakparov et al.: One-sided phenotyping mixing.: Consistent association of GRSPaV with vines indexing positive for Rupestris stem pitting. Further support to the cause-effect relationship of GRSPaV with this disease Galikparov et al.: Rugose wood viruses in Iran. Ahmed et al. Nakano et al.3% among groups.: Rugose wood viruses in Egypt.: Rugose wood viruses in the Czeck Republic. Wang et al.: GRSPaV particles. i. Buzkan et al. Confirmation of the cause-effect relationship between GVA and Kober stem grooving. occurs in Nicotiana plants doubly infected with GVA and GVB.: Movement proteins of GVA and GVB detected by gold immunolabelling in association with cell walls and plasmodemata of infected cells. GVA movement protein is also present in great quantity in the cytoplasm.: Elimination of GVA by cryopreservation. and III) identified in South Africa based primarily on sequence homology of the 3' end of the viral genome. Minafra et al. Goszczynski and Jooste: Use of single-strand conformation polymorphism reveals molecular heterogenity in GVA populations. Saldarelli et al. Kominek et al. Boscia et al. Nucleotide sequence identity within groups is 91-99. II. Samples made up of three buds from dormant canes are less laboriour to prepare than cane shavings and yield comparable results.: GVA transmission by Pseudococcus comstocki. 2000b 2000 2001 2001 2001 2002 2002 2003 2003 2003 2003 2003 2003 2003 2003 2003 2003 2003 86 . Seyval seedlings. Habili et al. intermingled with virus particle aggregates.: Production of monoclonal antibodies to GVB.: Synthesis of full-length cDNA copies of GVA and GVB genomes.: Production of an infectious RNA transcript from a full-length cDNA clone of GVA. Meng et al. Stewart and Nassuth: An improved extraction method allows RT-PCR detection of GRSPaV virtually thoughout the year in all grapevine tissues. Dell'Orco 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.
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GRAFT INCOMPATIBILITY .
Australia and Chile (young vine decline). Based on the differential responses of a panel of 18 rootstocks. the increased use of clonal material is disclosing unprecedented conditions of generalized decline that develop dramatically in certain scionrootstock combinations. Description Main synonyms: Incompatibilité au greffage (Fr. 1. and together with Grapevine virus B (GVB). although none of a number of known grapevine-infecting viruses has been found in affected vines. with margins more or less extensively rolled downwards. European grape varieties grafted on tolerant rootstocks (e. GVB is mealybug-borne and can be spread at a site by these insects. Other assays include nucleic acid-based tecniques such as single step or nested reverse transcription-polymerase chain reaction (RT-PCR) and 95 . only GLRaV-2 RG was identified. Other molecular variants of GLRaV-2 were reported from New Zealand (Alphie virus). and Australia in association with young vine decline conditions. 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. Normal growth resumes with the second or third leaf. This latter condition has been known for a long time and occurs also in rugose wood-affected vines. whereas varieties grafted on susceptible roostocks (e.g.) Symptoms: Newly planted vines grow weakly. 101-14) exhibit a green canopy and perform rather well. though not consistently and. 3309) develop a discolored canopy . The same virus was detected in diseased Chilean grapes. The heat-labile graft-transmissible agent present in the hybrid 140R.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). but the colour of the canopy remains green. 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. so as to represent veritable emerging diseases. Varietal susceptibility: Appearance of graft union disorders depends more on the rootstock rather than the scion. Salt creek . Infected propagative material is to be blamed for its dissemination. Of these. Freedom. appears to be involved in California's young vine decline. Severely affected vines decline and may die within one or two years. scions show a stunted and bushy vegetation due to the contemporary proliferation of apical and axillary buds. A prominent swelling forms at the scion/rooststock junction and variously extended necrotic lesions may develop on the rootstock stem. 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). Detection: Indexing on Caberent sauvignon is a reliable method for detecting incompatibility conditions. a member of the genus Closterovirus. shoots are short. The nature of this disease has not been ascertained but one or more graft-transmissibile agents may be involved in its aetiology. and again California (rootstock stem lesions). but the yield is reduced. 1103P. Agents: An ordinary strain of Grapevine leafroll-associated virus 2 (GLRaV-2) is consistently associated with Kober 5BB incompatibility (Europe). 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. The putative agent of bushy stunt was consistenly found in clones of the rootstock 140R in which it is latent. 03916. consistently. associated with grapevine bushy stunt is still unidentified. leaves are small-sized. 5C. up to five different grafttransmissible agents inducing incompatibility could be differentiated in California. which are usually not accompanied by wood abnormalities (pitting or grooving). A transitory form of incompatibility was reported from Italy under the name of bushy stunt. incompatibilità d'innesto (Ital.). Chile. Harmony. Argentina and probably elsewhere. Cases of graft union disorders have been observed in Europe (Kober 5BB incompatibility). Syrah decline is a severe disease occurring in France. However. California. is not transmitted by mealybugs and does not have a known vector. decline and may die. 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. in Argentinian grapes. New Zealand. A virus originally detected in cv. Transmission: GLRaV-2. In this case. except for GRSPaV. Kober 5BB.g.
utilization of tolerant roostocks is advisable.: Updated report on the state of the art of investigations carried out in France on Syrah decline. Graft-transmission of the incompatibility factor. Renault Spilmont et al. Gomez Talquenca et al: GLRaV-2 is consistently associated with declining Cabernet sauvingon vines grafted on different roostocks in Argentina. Savino et al. Martelli: GRSLV and GLRaV-2 are serologically related and are both recognized by a panel of 18 monoclonal antibodies. Currently known graft incompatibility agents can be eliminated with reasonable efficiency by heat therapy. whenever feasible. No conclusion are drawn.: GLRaV-2 is associated. Durquety et al. the use of sensitive rootstocks is to be avoided and. Boubals and Huglin: Report on graft incompatibility of certain varieties grafted on 57R. and death of the vines. Boubals: Report of a national French study group investigating the aetiology of Syrah decline. GRSLV has about 75% nucleotide homology with GLRaV-2. Strategies on how to protect healthy stocks from vector-mediated GVB reinfection in the field are yet to be developed. though not consistently. The problem is very complex and may involve several still unidentified factors. 2.: Description of bushy stunt and evidence that it is caused by a graft-transmissibile heat-sensitive agent carried by some clonal rootstocks. meristem tip culture. 2003 2003 2003 2003 2003 2003 2004 96 .: GLRaV-2 is the cause of a graft incompatibility revealed by Kober 5BB.: Two papers describing incompatibility phenomena between clonal selections of different cultivars grafted on Kober 5BB Fallot et al.: Third paper of a series on incompatibility on Kober 5BB. decline. If scionwood is infected. Uyemoto and Rowhani: Indexing on 18 different grape rootstocks reveals the existence of at least five different agents causing graft incompatibility. 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. Boubals: Syrah decline occurs in Argentina Uyemoto et al. Control: Prevent introduction of infected vines in the vineyard by using certified grafted plants or virusfree scionwood and rootstocks.: Identification of an apparently new closterovirus denoted Grapevine rootstock stem lesion virus (GRSLV) causing stem necrosis of rootstocks. or a combination of the two. 1977 1979 1986 1991 1995 2000 2000 2000 2001 Jacob: Description of graft incompatibility in different scion/stock combinations. Historical review 1942 1950 1973. Greif et al. GRSLV re-named Redglobe strain of GLRaV-2. with a decline condition of young Thomposn seedless vines in Chile.: GLRaV-2 and GVB are consistently associated with young vine decline in California. using degenerate or virus-specific primers. Bertazzon and Angelini: Comparison of several detection methods for the broad or specific identification of Grapevine leafroll-associated virus 2 variants.immunocapture RT-PCR. Bonfiglioli et al. Golino et al. Prodan et al.: Report of a new molecular variant of GLRaV-2 from New Zealand. Suggestion that they are molecular variants of the same virus species.
3. and E. Prota. Journal of Plant Pathology 86. Autres cas chez la vinge. 137-141 Boubals D. Extended Abstracts 14th Meeting of ICVG. Le dépérissement de la Syrah.S. 2003. Uyemoto J. Garcia Lampasona and O... and P. Ruchaud. Locorotondo 2003. Nouvelles recherches sur l'incompatibilité clonale d'Abouriou greffé sur 5BB. S. Extended Abstracts 14th Meeting of ICVG. 201-203. Extended Abstracts 14th Meeting of ICVG. A young grafted vine decline syndrome in Argentina vineyards. 2000. New closterovirs in 'Redglobe' grape causes decline of grafted plants.A.P. Discovery of different grapevine sources with graft-transmissible agents causing union-incompatibility on sensitive rootstocks.. J. Boubals D.K. 2003.. Sim and A. P. 2000. Di Silvio.M. R. Bonfiglioli R. 2003. Di Terlizzi. Greif C. 1977. Golino D. Rivieccio and F. 122-129 and 171-178. Extended Abstracts 13th Meeting of ICVG. Gazeau and J. Pantaleo. Ruchaud. Progrés Agricole et Viticole 94. Grenan and J. Progrés Agricole et Viticole 90. Compte-rendu de la réunion du Groupe de Travail National. Extended Abstracts 14th Meeting of ICVG. Fiori.A. Grapevine virology highlights 2000-2003. Garau. Dauty. Progrés Agricole et Viticole 103. 277. D. Prodan S. Huglin. 3-10. 2003.. Savino V. Etude de phénomènes d'incompatibilité au greffage chez la vigne. 2000. Renault Spilmont A. M. 1986. Jacob H. F. Martelli G. 142-143.. Gracia. Progrés Agricole et Viticole 117. Gazeau. Rowhani. Le clone et ses reactions au greffage. Existence dans un cépage population de clones présentant divers degrés de compatibilité avec certains porte-greffes. Extended Abstracts 14th Meeting of ICVG. 141. B. Examples of incompatibility between grape varieties and roostocks. Molecular studies on a graft incompatibility syndrome in New Zealand vineyards yields another probable variant of Grapevine leafroll-associated virus 2. 279-283. Etude de l'incompatibilité au graffege de certain cépages et du 57R. Le clone et ses reactions au greffage. P. I. Locorotondo 2003. Extended Abstracts 14th Meeting of ICVG. Syrah decline in French vineyards.S. J.M. Rowhani. Locorotondo 2003. J. 1979. S. 28-31. Progrés Agricole et Viticole 96. C.. 2004. 2001. Locorotondo 2003.. Montalegre and N. 1991. S. Boubals D. Fiore. La trasmission de l'incompatibilité au greffage entre 5BB et Vitis vinifera. S. 1950. and A. Edwards and A. 167-173. Proceeding of the American Society of Horticultural Science 41. 97 .P. Bass. C. References Bertazzon N.. Volos 1990. Locorotondo 2003.. J. 420-427 Fallot J. Phytopathologia Mediterranea 34. 211-216. C. Progrés Agricole et Viticole 117.. Durquety P.. Rowhani. 202-210 Uyemoto J. Krag. 1973. Angelini. Luvisi and R. O. 2003. 85-86. Benassac and R. Durquety and J. Gomez Talquenca G. 2003. Boscia.M... Prota. The relationship of grapevine leafroll-associated virus 2 with a graft incompatibility condition of grapevines. Progrés Agricole et Viticole 67. Fallot. Le clone et ses reactions au greffage. Presence in clonal rootstocks of a grafttransmissible factor that induces stunting and bushy growth in European grapevines. 139-140. Locorotondo 2003. 85-86.P. B. Adelaide 2000.P. Le dépérissement de la Syrah existe aussi en Argentine.II. Identification of the latent viruses associated with young vine decline in California.. Grau. Advances in the detection of Grapevine leafroll-associated virus 2 variants. Aetiology of decline in Thompson seedless grafted table grape plants. Ruchaud. Legin R. 1942.K. Walter. 144.M. Proceedings 10th Meeting of ICVG. V. D. Walter and U.E. 183-189. Boursiquot. 283-290. California Agriculture 55 (4). and B. Fallot. 1995. A. Durquety P.III.
FLECK COMPLEX .
GFkV genomic RNA constitutes about 35% of the particle weight. GAMaV. Grapevine red globe virus (GRGV) is a Grapevine fleck virus (GFkV)-like virus which apparently does not induce symptoms in European grapevine varieties (e. c. Description Main synonyms: A. which is used as indicator. 25 kDa. grapevine rupestris necrosis. The coat protein (CP) of GFkV and GRGV particles is made up of a single protein species with Mr of c. Red globe) nor in V. rupestris following graft inoculation. leaf symptoms are characterized by star-shaped chlorotic spots. 1. Rupestris necrosis. e. Symptoms are expressed in Vitis rupestris and consist of clearing of the veins of third and fourth order. Marmorierung der Rebe (Germ. rupestris. irregularly distributed over the leaf blade. vinifera in California. V. Sultanina). B. Italy and California. but likely to occur elsewhere. leaf petioles and veinlets. T made up of empty protein shells and B. Severe strains induce also varying degrees of stunting.FLECK COMPLEX The fleck complex consists of several diseases (grapevine fleck. Mild asteroid mosaic-like symptoms are shown by some European grapevine varieties (e. The putative causal agent of the disease has only been found in California. 50%). 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. adverse influence on vigour. positive sense RNA with high cytosine content (c. Fleck is an ubiquitous disease reported from most viticultural countries in the world. GFkV particles sediment as two centrifugal components. and GRVFV genomes are available. producing localized translucent spots. Recorded from California and Italy . GFkV. 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.). Sternmosaik der Rebe (Germ. and produce little or no fruit. Agents: All viruses of the complex. Leaves are asymmetric. capped. This disease. is latent in European grapevine varieties. twisted and may curl upward. b. mosaico stellare (Ital. Asteroid mosaic symptoms have been observed in several varieties of V. Fleck. rupestris. The putative causal agent of the disease so far has been found in Greece.g. 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. Affected vines are often stunted. GRGV. Grapevine asteroid mosaic-associated virus (GAMaV). Grapevine fleck: Marbrure (Fr. sometimes with necrotic center.). Leaves with intense flecking are wrinkled. Transient mild chlorotic discolourations of the primary and secondary leaf veins develop in V. screziatura (Ital. grapevine asteroid mosaic.).). In V. Although the elusive nature of the complex hinders the assessment of its economic impact. which are twisted and asymmetric. which is a monopartite single-stranded. Rupestris vein feathering. Leaf symptoms usually become less severe in summer. Asteroid mosaic. maculatura infettiva. reported only from Japan. wt of 2. d.6 x 106 ) is 7564 nt in size and contains four open reading frames (ORF) that 101 .). whereas the CP of GAMaV and GRVFV consists of a major protein of 21 kDa and a minor prtotein of 25 kDa. Grapevine asteroid mosaic: Mosaïque étoilée (Fr. twisted and puckered along the veins. vinifera.). In V. rooting ability of rootstocks and on graft take has been reported. The disease is latent in European grapevine varieties and in most American rootstocks.g. GFkV genomic RNA (Mol. and Grapevine rupestris vein feathering virus (GRVFV) are all phloem-limited and non mechanically transmissible. containing the genome. The complete sequence of GFkV and partial sequences of GRGV. the disease elicits creamy-yellow bands developing along the major veins of the leaves. rupestris reacts with localized necrosis of the shoots. Symptoms: a.
Vuittenez et al. Control: Because of the latency of symptoms sanitary selection of European grapevine cultivars and most American rootstock hybrids is unrealiable. GRGV. and GRVFV. Historical review 1954 1962 1966 1966 Hewitt: First description of asteroid mosaic in California. but no experimental data are available. Because of its molecular characteristics. GFkV can be eliminated by heat therapy. 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. 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. GFkV is not seed transmitted. a disease inducing symptoms similar to those of fleck in V.9 kDa (ORF 4) with unknown function. primary dissemination of these and the other viruses of the complex is through infected propagative material.: First record of fleck as an unidentified symptom different from fanleaf and transmissible from symptomless varieties to V. Further physico-chemical.4 kDa (ORF 3) and 15.4 polypeptide with the conserved motif of replication associated proteins (ORF 1). 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. ELISA is currently employed for routine detection of GFkV. The 3' end of GRGV genome is structurally similar to that of GFkV except for the lack of ORF 4. These deranged organelles are thought to be sites of virus replication. molecular and ultrastuctural studies disclosed sufficient similarities between maculaviruses. Comparison of symptoms with those of other mosaic diseases of grape. Transmission through dodder of GFkV has been reported but it is of no epidemiological importance. rupestris St.George. Detection: Indexing on V. Refatti: Review paper on asteroid mosaic. 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". Attempts to transmit the disease by mechanical inoculation to herbaceous test plants or by Xiphinema index were unsuccessful. Polyclonal antisera and monoclonal antibodies to GFkV heve been raised. 102 . Virus specific and degenerate primers have been designed for single or multiplex RT-PCR detection of GFkV. As the disease is rare and does not appear to be spreading.rupestris described in France. whilst GAMaV and GRVFV were assigned to the genus Marafivirus. meristem tip or fragmented shoot apex culture. Therefore. GAMaV. marafiviruses and members of the genus Tymovirus to warrant the establisment of the a new family denoted Tymoviridae. of which it represents the type species. No information is available on individual susceptibility. Varietal susceptibility: GFkV and possibly all the other viruses of the complex infect naturally a large number of varieties and Vitis species. Although observations from Italy. its economic importance is low. The same sanitation procedures are likely to operate successfully with the other viruses of the complex.encode a 215. Hewitt et al. the CP (ORF 2). GFkV was identified as the representative of a new genus denoted Maculavirus. 2.: "Marbrure". Transmission: No vector is known for any of the viruses of the fleck complex. and two proline rich polyproteins of 31. but cannot be used for any of the other members of the complex due to the unvailability of antisera. whereas GAMaV induces peripheral vesiculation of chloroplasts.
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. Goheen and Luhn: A novel heat therapy system based on virus inactivation in buds grafted onto healthy LN 33 rootstocks is effective against fleck. Triolo and Resta: Tetracycline treatments are ineffective against fleck. Castellano et al. Dolja et al.: Description of fleck as an independent graft-transmissible disease present in many European varieties and American rootstocks. Barlass et al. Savino et al. Ottenwaelter et al.: Successful elimination of fleck through fragmented shoot apex culture in vitro.: Identification of a dsRNA of about 7 Kb pairs in diseased vines. 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). Engelbrecht and Kasdorf: Observation of natural field spread of fleck in South Africa.: Physicochemical characterization of GPLIV.: Successful elimination of fleck through heat therapy. Triolo and Materazzi: Fleck has a detrimental effect on the quality V. Rives: Further demonstration that fleck is distinct from fanleaf based on differential responses to heat treatment.: Observation of a non mechanically transmissible virus. Woodham and Krake: Dodder transmission of fleck from vine to vine. rupestris propagating wood.: Observation of an isometric non mechanically transmissible virus in the phloem of diseased vines.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.: Fleck is not seed transmissible. Dismissal of the prokaryote etiology hypothesis. Castellano et al. Castellano and Martelli: Confirmation that GPLIV is associated with multivesiculate bodies and demonstration that these derive from deranged mitochondria. Report that a virus serologically similar to GPLIV is associated with the disease. Report of multivesiculate inclusion bodies probably connected with GPLIV infection. Confirmation that the virus can be eliminated by heat therapy and is not related to leafroll.: Purification of GPLIV from naturally diseased vines and production of a specific antiserum. leafroll and corky bark).: Report of widespread occurrence of fleck in visually selected grapevine clones in southern Italy. in sieve tubes of field-grown vines with leafroll symptoms but likely to be affected by other diseases. Verderevskaya et al. 1982 1983 1983 1983 1984 1985 1985 1987 1989 1990 1990 1990 1991 103 . rupestris. The efficiency of heat treatment for disease elimination is unsatisfactory. Hévin et al. Bovey: Identification of fleck in Switzerland as a latent disease of Chasselas transmissible to V. Hewitt et al. Milkus: Suggestion of a prokaryotic etiology for fleck.
rupestris of an isometric virus latent in V. ELISA used successfully for virus detection in large scale survey. Detection of sequences of an undentified virus from a Greek grapevine. later named Grapevine rupestris vein feathering virus (GRVFV).: 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. Kyriakopoulou: Description of a disease similar to asteroid mosaic observed in V.: Report of a highly consistent association of GPLIV with fleck in naturally infected and graft-inoculated V. based on the differential reactions of indicators Credi and Babini: Infection by fleck. Walter and Cornuet: Confirmation by ELISA of the consistent association of GFkV with fleck disease. Sultanina in Greece. Virus named Grapevine redglobe virus. Pruning wood is reduced by 51% in 420A and by 37% in Kober 5BB. Elbeaino et al. Sabanadzovic et al.1991 1991a 1991b 1991 Gugerli et al. vein necrosis and vein mosaic has a detrimental effect on rootstock growth. GFkV-like multivesiculate bodies derived from deranged mitochondria are present in infected cells. Boscia et al. rupestris with a necrotic disease. Sabanadzovic et al.: A spherical virus purified from berries of Ajinashica disease-affected vines is serologically related to GPLIV (= GFkV) and has physicochemical properties comparable to those of GFkV.: A non mechanically transmissible isometric virus similar but unrelated to GFkV identified in asteroid mosaic-infected grapevines. rupestris.: Natural field spread of GFkV observed in Northern Italy Schieber et al. Boscia et al.: GPLIV shown to be the agent of fleck. vinifera. 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.: Molecular reagents (degenerate primers) developed for the specific identification of viruses of the fleck complex (GFkV. The disease seems to be spreading naturally.: Report of the close association with fleck symptoms in V. Namba et al. Marsumoto and Ohki: A spherical virus resembling GFkV identified in thin sectioned cells of V. Meristem tip culture effectively eliminates the virus. This cytological feature recalls that later found in vines infected by Grapevine rupestris vein feathering virus. June-July are the best months for ELISA detection of the virus in Alsace. Virus named Grapevine asteroid mosaicassociated virus (GAMaV) Boscia et al. Fortusini 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. GAMaV and of another virus with GFkV-like particles phylogenetically but not serologically related to GFkV present in a cv Red globe vine. Adverse effect on Teleki 5A is negligible. GRGV).: Complete nucleotide sequence of GFkV genome. vinifera cv. Boscia et al. Molecular properties of this virus further support the notion that it warrants classification in a genus of its own. Virus renamed Grapevine fleck virus (GFkV). Symptoms are severe and affected vines are almost fruitless. 1991 1993 1994 1995 1995 1996 1996 1997 1997 1998 2000 2001 2001 104 . GAMaV.: Additional monoclonal antibodies raised in France.
165-169.. Journal of Ultrastructure Research 89.. Martelli. Sabanadzovic . Abou Ghanem-Sabanadzovic. A. Babini. 347-354. S.J. G.: Sequencing of the 3' end of the genome of GRGV. V. Martelli. 160-163. Sabanadzovic. 23-39.A.. a new genus of plant viruses having GFkV as type species and GRGV as tentative species.P. V.A.P. Proceedings 8th Congress of the Mediterranean Phytopatological Union. Barlass M.P. Kyriakopoulou and G. 2003a. Journal of Phytopathology 129. D.G. 95-98. V. S. Castellano. 1991a. References Abou Ghanem-Sabanadzovic. A. Martelli.P. G.2002a 2002b 2003a Martelli et al. Martelli.D. Virus-like particles and ultrastructural modifications in the phloem of leafroll-affected grapevines. Savino.. and A. Phytopathologia Mediterranea 24. Association of a phloem-limited non mechanically transmissible isometric virus with grapevine fleck disease. Savino and G.A. Castellano. Minafra. Martelli and M.M. N. Virus Genes 27. 1995. N. 2003b. A.P.P. Boscia. S. Argentina. South Africa. Multiplex RTPCR detection of Grapervine fleck virus-like viruses in grapevine with co-amplification of control plant mRNA.. Castellano M. Sequencing of the 3' end of three grapevine fleck virus-like viruses . Plant Pathology 44. Kyriakopoulou and G. 105 . Ultrastructure and nature of vesiculated bodies associated with isometric virus-like particles in diseased grapevines. Boscia.F.G. Progress in the study of the phloem-limited isometric virus-like particles associated with leafroll-diseased grapevines. Savino. Castellano. 56-64. In other countries (USA. Savino. 291-295. Un virus latent dans le Chasselas. Vitis 30.. Savino.P.P. 1990.A. Atabekov. Annales de Phytopathologie. 1972. GRVFV recorded from California and confirmation that GAMaV does not occur outside of California.. 2003b 2003 3. Boyko. Karsev.. A non mechanically transmissible virus associated with asteroid mosaic of the grapevine. Martelli. Martelli et al.A. Elicio. 3134. Field spread of corky bark. Proceedings 10th Meeting of ICVG. Abou Ghanem-Sabanadzovic et al. 173-174. 1990. Sabanadzovic. Skene. Martelli and V. leafroll and Shiraz decline diseases and associated viruses in South African grapevines. Volos 1990. Abou Ghanem-Sabanadzovic et al. Di Terlizzi.. Vitis 22.A. Credi R. 1996. rupestris confirms the assignment of GRGV to the genus Maculavirus and of GAMaV and the Greek virus to the genus Marafivirus. Double stranded RNA associated with fleck disease of grapevine.P.: Description of the family Tymoviridae. N.: Development of a multiplex RT-PCR protocol for the simultaneous detection of GFkV-like viruses using plant mRNA as an internal control. V. Boscia D. 195. 1994. Molecular detection of Grapevine fleck virus-like viruses. and G.E. Bovey R. G. V.R. 1991b. 191. M. fleck. 101-102 Boscia D.: Description of Maculavirus. Savino and D. Agadir 1990. Advances in Horticultural Science 10.. Boscia D. Phytophylactica 22. G. GAMaV and of a virus of Greek origin which induces vein feathering in V. Vitis 40: 65-68. Martelli. Vitis 33. Dolja V. Martelli. T. R. Tomashevskaya. Martelli. 1990. K. comprising the genera Maculavirus and Marafivirus that include GFkV/GRGV and GAMaV/GRVFV. U.E. Krake. respectively. Verderevskaya and J. Greek virus recognized as a species in its own right denoted Grapevine rupestris vein feathering virus (GRVFV). Sabanadzovic and G. Iran. Production of monoclonal antibodies to grapevine fleck virus..V.V. Regeneration of virus-free grapevines using in vitro apical culture.: A sequence variant of GFkV (GFkV416) with a 63 nucleotide insertion in the replicase gene identified in Australia and New Zealand. Boscia D. B.P. M.. Kasdorf. 1984.R.G. P. A. S. 1982. V. Martelli 2001. O. and Japan) only the variant without insertion (GFkV353) was detected.C.P. Annals of Applied Biology 101. Boulila M. Engelbrecht D. Extended Abstracts 14th Meeting of ICVG. 151-158. ElBeaino T. and G. Numéro hors série. Rowhani and G. Rowhani P. Digiaro.P. 1985. 97-105. 11-16 Abou Ghanem-Sabanadzovic. Castellano M. Shi et al. Effect of virus and virus-like infections on the growth of grapevine rootstocks. Locorotondo 2003. 1983. and G. Some properties of a phloem-limited non mechanically-transmissible grapevine virus. Woodham and L. Identification of the agent of grapevine fleck disease.. Castellano M. V. Savino and M.
Sabanadzovic. Tremea. Rives.. is transmitted by graft inoculation. Kuniyuki H. 143-146.. 1972. Proceedings 10th Meeting of ICVG. M. George). Plant Disease Reporter 61. with Special Reference to Vitis. Doazan and M. Castellano. M. S. Gugerli.L. Rives M. Hévin. Heat therapy eliminates the ability to transmit the causal agent of "marbrure" in several V. Grapevine fleck virus-like viruses in Vitis. 1991. 553-565. Complete nucleotide sequence and genome organization of grapevine fleck virus.E. Costa. 1972.IV. Abou Ghanem. 47-64. Goheen. 1962. Goheen. T. Saldarelli.P.M. 1985. Division of Agricultural Sciences. Berkeley. A possible new necrotic diseases of grapevine associated with small isometric particles and novel membrane-bound large particles.-E. 385-388. Martelli.J. D.. The family Tymoviridae. Investigating the transmission of marbrure and fan-leaf through the seed in the grapevine. Monoclonal antibodies for detection. Rivista di Patologia Vegetale. Ser. Sabanadzovic S. Prati. 2009-2015 Savino V. Triolo E. Digiaro and G. S.I. 2002b. Abou Ghanem-Sabanadzovic. Extended Abstracts 12th Meeting of ICVG. 75-77. N. Fortusini A. 157-164. Dreher. rupestris "du Lot" (St.Faoro F and P. Informatore Fitopatologico 46 (12).C.H. 567-571. Volos 1990. S. P.. Archives of Virology 147. 1973. Martelli. A. The responses of the Grapevine fleck agent to tetracycline-HCl antibiotic and Dienes' stain. Annales de Phytopathologie. In Frazier N. 1973. Proceedings International Conference on Virus and Vector on Perennial Hosts. Martelli. 9. Ohki. 560-564. Gooding.W. Resta. 1998. vinifera clones and in V. serological characterization and immunopurification of grapevine fleck virus. Annals of the Phytopathologial Society of Japan 64. S... 9. Ramel and F. Saldarelli and G. Milkus B. 2002a.. A.. Proceedings 10th Meeting of ICVG. Symptoms of grapevine asteroid mosaic in Greece. Boscia and G. G. 1985. N. Gugerli P. M. Luhn. Numéro hors série. A. University of California. 1996. D.B. 204-207. 5960. Nucleotide sequence variation in a small region of Grapevine fleck virus replicase provide evidence for two sequence variants of the virus.. and E. Heat inactivation of viruses in grapevines. Procedures for rapid detection of virus and viruslike diseases of grapevine.P. 1966. Brugger. Annals of Applied Biology 142. 57-83. Raski and G. Martelli G.T. Namba S. B.. Rivista di Patologia Vegetale. Ser. N. Davis 1965. 349-355. 2001.M.A. M.P. Ser.P. Martelli G. Incidence of some graft-transmissible virus-like diseases of grapevine in visually selected and heat-treated stocks from Southern Italy. Phytopathologia Mediterranea 24. Journal of General Virology 82. Mycoplasma. D. 1973. Vitis 3. 767-774. Goheen A. Rives. Parsons. Cory and C.. 197-203. Sabanadzovic S. and J. 1837-1846. 281-285.): Virus Diseases of Small Fruits and Grapevines (A Handbook). Hévin M. 1995. S. 253-258.B. Tsuchizaki and D. C. Asteroid mosaic of grapevine. Matsumoto T. Volos 1990. Archives of Virology 145. Rivista di Patologia Vegetale. 618-622. 1997. Hewitt W. Schieber O. Some virus and virus-like diseases of grapevines. Shi B. Maculavirus. Jr. Plant Disease 75. Séparation de la marbrure et du court-noué (panachure) chez la vigne par thermothérapie. 106 . Belin and B. Acta Phytopathologica Academiae Scientiarum Hungaricae 9. (Ed.IV. Phytopathologia Mediterranea 24. Yamashita. 9. Hewitt W. Grapevine fleck disease. Cytological alterations associated with an unidentified isometric grapevine virus (UIGV). 287-289. Boscia. Abou Ghanem-Sabanadzovic and P. IV.C.V.. Annales de Phytopathologie. 2003. and A. Further characterization of grapevine leafroll disease. Symons. M. 1847-1853. J. Occorrencia de mais um isolado do virus do mosaico da nervuras da videira que no causa sintomas no porta-enxerto Kober 5BB.C Edwards and T. Cinquanta and S. Doazan and M. Grapevine asteroid mosaic.P. Luhn.or chlamidia-like bodies in grape. Sabanadzovic. Purification and properties of spherical virus particles associated with grapevine Ajinashica disease. a new genus of plant viruses.P.. 1991.C. 39-43. Hewitt W. Numéro hors série. 1991. Leclair. 212-214. 1954. Scattini. 12491253. Ottenwaelter M. Walter. 1970. European Journal of Plant Pathology 103. and S. affected by marbour. J. Kyriakopoulou P. Rosciglione.-J. M.P. N. Abou Ghanem-Sabanadzovic.. P. Archives of Virology 147..J. Refatti E.. 1997.S. 1974. Bulletin of the California Department of Agriculture 43.B.. Studies on virus diseases of the grapevine in California. Seddas. J.. Habili and R. L. Mink G. Refatti E. 2000. 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. and C. latent in many varieties. 1977. 31-32. 43-47.. Lisbon 1997. Fitopatologia Brasileira 20. Bonnard. Ottenwaelter. Gonsalves.. N.
Yamakawa Y. Krake. 1966. La Recherche Agtonomique en Suisse 26. Phytopathologische Zeitschrift 106.S. 1983. R. Marinesku and E. Observations sur une mosaïque de la Vigne.R. Agronomie 13. 107 . and L.C. and A.Triolo E. 651-657. 67-73... V. Journal of the Japanese Society of Horticultural Science 58. and P. 1987. 1983. Verderevskaja T.D.. probablement indépendante du virus du Court-noué. Numéro hors série. 193-198. Kuszala. Vuittenez A. Ätiologie und Diagnose der Marmorierung der Weinrebe. George. Investigations on transmission of grapevine leafroll. ELISA detection of grapevine fleck virus (GFkV). Semtschik. Archiv für Phytopathologie und Pflanzenschutz 19.G. Woodham R. 1989. 297-302. 1993. Annales des Epiphyties 17. Walter B. 221-226. Materazzi. Cornuet. Legin and J. 3209-324. Virus reinfection of virus-free Cabernet sauvingon and Cabernet franc vines. yellow speckle and fleck diseases by dodder.. La maculatura infettiva della vite: influenza di isolati diversi sull'attitudine alla propagazione vegetativa di Vitis rupestris St.
18% of the particle weight. 0. with Mr 24x 103 Da. Chardonnay and Veltliner rouge précoce identified as reliable indicators.73 x 106 (2593 nt). from quasi isometric to bacilliform. Bulgaria. Faint yellow speckling. has differently shaped particles. Transmission: AMV is efficiently transmitted by aphids in a non persistent manner and can cause epidemic outbreaks in many of its natural hosts. AMV.g. Switzerland. 2.62 x 106 (2037 nt). the type species of the genus Alfamovirus. wts: RNA-1.MINOR VIRUS DISEASES Several graft-transmissible diseases are known.: First record of AMV infections and description of symptoms in German grapevines. infections are scattered and occasional. others occur in Japan. Jankulova: AMV infections recorded from Bulgaria. Virus elimination by treating 37 days at 37-38°C. rupestris and the hybrid Grézot 1 x 5C. suggesting that the virus spreads primarily through infected planting material. is the putative causal agent. Grapevine berry inner necrosis). Historical review 1973 1975 1976 1979 1978 1981 Bercks et al. Varietal susceptibility: Little information available. Yellow mottle has been reported from Germany. with which specific viruses are associated and thought to be their possible causal agents. RNA-3. A. but some are of economic relevance locally (e. In grapevines. Bovey and Cazelles: AMV particles visualized in thin sectioned grapevine leaves. Capsid proteins subunits are of one type. Their overall importance is minor if compared with that of the major diseases dealt with in previous chapters. 109 . a mechanically transmissible virus. 30 to 57 nm in size. 1. 0.04 x 106 Da (3644 nt). rings and lines are typical summer responses of infected vines. Description Main synonyms: None. The spring growth shows more or less extensive yellowing of the leaf blades that does not extend to the veins. and Turkey. RNA-2. There may be differential susceptibility among cultivars. European diseases GRAPEVINE YELLOW MOTTLE 1. Plant vigour and yield do not seem appreciably affected. Control: Use of healthy material obtained by heat treatment. and a tripartite RNA genome accounting for c. Hungary. Some of these diseases have been recorded only from Europe. with the following mol. Detection: AMV is mechanically transmissible to herbaceous hosts and can also be identified by ELISA and moleculat techniques in infected vines. Main symptoms: Various patterns of yellow discolouration characterize the disease. Beczner and Lehoczky: AMV infections recorded from Hungary. former Czechoslovakia. Bovey and Brugger: AMV recorded from Switzerland in grapevine and transmitted by grafting to V. Novak and Lanzova: AMV infections recorded from hop and grapevine in Czechoslovakia. Agent: Alfalfa mosaic virus (AMV). however.
B. Novak J. Brugger. Rome. References Akbas B. 63-65. 1993. Alfalfa mosaic virus on grapevine. Grapevine disease in Hungary caused by alfalfa mosaic virus infection. 3. Proceedings 6th Meeting of ICVG. 119-128. Detection and Diagnosis of Graft Transmissible Diseases of Grapevines. Varietal susceptibility: No information. Graft transmission to cv. Francki R. Jubileum 75. Control: No information. has differently shaped particles. D. Cazelles. 1976. sativa DC. . is seed-transmitted and spreads with diseased propagative materials. 1985. Horticulture 7.. Identification of alfalfa mosaic virus and tomato bushy stunt virus in hop (Humulus lupulus L.1985 1993 1993 Francki: Comprehensive review of the properties of AMV and other viruses with tripartite genome. 1979. 39. 1973. Plenum Press. In: The Plant Viruses.).Hegi) plants in Czechoslovakia.. 3rd International Congress of Plant Pathology. Revue Suisse de Viticulture. or the upper part of the blade.P. or maple leaf-like line patterns typically confined to the petiolar area. and J. and G. Munich 1978. vinifera cultivars are susceptible. Martelli G. 1975. 166-171. Martelli: Yellow mottle suggested as the name for the disease caused by AMV in grapevines.B.B. Jankulova M. Polyhedral Virions with Tripartite Genomes (R. ed. Bovey R. Bercks R. Grapevine line pattern virus (GLPV) a possible member of the genus Ilarvirus. Detection: GLPV is mechanically transmissible to herbaceous hosts. FAO Publication Division. Monografias INIA No. Researches on grapevine virus diseases and determination of their incidence in Ankara Turkiye. This disease is known to occur only in Hungary. Journal of Turkish Phytopathology 22. roughly following its contour. The viruses and their taxonomy. 110 . (ed. Phytopathologische Zeitschrift 76. quasi spherical 25-30 nm in diameter to bacilliform 40 to 75 nm in length. New York . Arboriculture.I. Erdiller. Francki. Main symptoms: Leaves show bright yellow discolourations that form marginal rings. 1993. Historical rewiev 1985 Francki: Comprehensive review of the properties of AMV an other viruses with tripartite genome. Acta Phytopathologica Academiae Scientiarum Hungaricae 16. 1978.). Transmission: GLPV has no known vector. Über den Nachweis des Alfalfa mosaic virus in einer Weinrebe. and a multipartite genome. Lesemann and G.) and grapevine (Vitis vinifera subsp. and J. Several V.I.18 . Querfurth. 1981. Beczner L. Description Main synonyms: None. 1-18. scattered spots or blotches. 55-63.-J. 263 pp. Investigation on certain viruses spread on grapevines in Bulgaria. Bovey R. Le virus de la mosaïque de la luzerne sur la vigne. and O. 152-154. Biologia Plantarum 18. and J. 2. Akbas and Erdiller: AMV infections recorded from Turkey. Lehoczky. Lanzova. 131-134. Agent: The putative agent. GRAPEVINE LINE PATTERN 1. Vigour and yield are reduced. Cordoba 1976.
Plenum Press. L. Transmission: No vector is known. CarMV is an isometric virus 30 nm in diameter. Occurrence of the line pattern hitherto unknown virus disease of grapevine in Hungary. Burgyan.I. Polyhedral Virions with Tripartite Genomes (R. a novel virus disease of grapevine in Hungary. Francki. Main symptoms: Symptoms are prominent in late summer and consist of yellow and/or reddish discolourations of the tissues along the veins. Lehoczky et al.A.. wt 1. Control: No information. D. Description Main synonyms: None. Line pattern. 1985. Mission. Kiryat Anavim. By converse.B. Farkas.P. Evidence that a graft-and mechanically transmissible virus is associated with it. ed. 1989. Bunches are reduced in numbers. J.B. 1987. Seed transmission of grapevine line pattern virus. family Tombusviridae. New York . according to more recent findings. Martelli.4 x 106 (4003 nt in size) and coat protein subunits of Mr 38 x 103 Da. or variously extended sectors of the leaf blade. Farkas. G. Lehoczky et al. The disease is graft-transmissible and spreads through infected propagating material. 115-116.). D. Castellano. Boscia. one of the putative agents of corky bark (rugose wood complex) has a very high association with diseased grapevines. The disease has been recorded only from Greece. the interveinal areas. Agent: Symptomatic grapevines were reported to be doubly infected by GFLV and Carnation mottle virus (CarMV) the type species of the genus Carmovirus. M. References Francki R. 2. G. J. size and have low sugar content.1987 1989 1992 Lehoczky et al. L.. 1-18. In: The Plant Viruses. 1992. Beczner and G. M. Grapevine virus B (GVB). Martelli and J. GFLV may not be involved in the aetiology of the disease. Lazar.A. Boscia. Varietal susceptibility: No information. The viruses and their taxonomy.: Description of line pattern disease in Hungary. has a monopartite RNA genome accounting for c. 23-30. 111 . Viruses associated with the disease are readily transmitted by sap inoculation and can be detected by ELISA and molecular techniques. with Mol. 3. Burgyan. Proceedings 9th Meeting of ICVG. 61-79 Lehoczky J. Historical review 1989 1991 Rumbos and Avgelis: Roditis leaf discoloration described in Greece. Beczner and G. 1987... However. Lehozcky J. Evidence of its grafttransmissibility. Lehoczky J.: Purification and characterization of GLPV and suggestion that it is the causal agent of the disease. Castellano. Avgelis and Rumbos: Double infection of diseased vines by GFLV and CarMV reported. especially near the petiole Leaves are deformed in correspondence of discoloured sectors. Detection: Graft-transmission to V. RODITIS LEAF DISCOLORATION 1.: Evidence that GLPV is transmitted through grapevine seeds. Israel.P. 18% of the particle weight. Phytopathologia Mediterranea 31. Kertgazdasag 19. vinifera cv.I.
C. discoloration of tissues bordering the veins. 437-443. Tsagris. Proceedings 10th Meeting of ICVG. 3. decline gradually and some die. Girgis et al. a virus with a tripartite RNA genome and a 30 kDa coat protein..E. 274-278. Description Main synonyms: None Main symptoms: Symptoms are chlorotic angular spots on the leaf blades. crinkling and deformation of the leaves. Volos 1990. N. Infected grafting material is likely to be responsible for virus dissemination. Roditis leaf discoloration -. The etiology of a new virus disease: grapevine angular mosaic. Sklavounos. C. A new ilarvirus isolated from grapevine in Greece.D. A. but differs from GLPV. Bem. A.D. thus is regarded as the agent of the disease. and A. Avgelis. P.M. Plant Disease 84. and ELISA.I. Journal of Phytopathology 125.C. GRAPEVINE ANGULAR MOSAIC 1. Laski Rizling from Slovenia exhibit a yellow line pattern syndrome resembling the grapevine line pattern disease described from Hungary. Katis. P.a new virus disease of grapevine: symptomatology and transmission to indicator plants. P. 19. and I. reproduced the field syndrome in mechanically inoculated grapevine seedlings.E. Transmission: GAMV is pollen-borne in herbaceous hosts and was able to infect pollinated plants. Baresana x Baresana. 1989. Whether this mechanism operates with grapevines has not been ascertained. Varietal susceptibility: No information Detection: Indexing on cv. 1991.P. Flowers abortion results in straggly bunches with small wrinkled berries bearing non viable seeds. Extended Abstracts 14th Meeting of ICVG Locorotondo 2003. the only other ilarvirus reported from grapevine. Kyriakopoulou.3. Bem. Dovas.: Evidence that GAMV is the agent of grapevine angular mosaic disease.: First record of GAMV. Katis. Carnation mottle virus isolated from vines affected with "Roditis leaf discoloration". Rumbos I. F. Kyriakopoulou. The disease has been recorded only from Greece. Agent: Grapevine angular mosaic virus (GAMV). Tzortzakakis and M. GAMV is molecularly related to a number of ilarviruses. 112 . References Avgelis A. A.I.M. Girgis S. Avgelis and N. YELLOW LINE PATTERN (Rasperry bushy dwarf virus) 1. 2000. mechanical transmision to herbaceous hosts. Infected grapevines are stunted. S. the closest being Tobacco streak virus (TSV). References Girgis S. Historical review 2000 2003 Girgis et al. Description Main synonyms: None Main symptoms: Grapevines of cv.. Avgelis. F. Dovas. C. 2003. 1345. Control: No information 2. Rumbos. P.
M. The virus spreads naturally in the vineyards. 113 . 20 Murant A.2 kb in size). CMI/AAB Description of Plant Viruses. ELISA . and RT-PCR. Varietal susceptibility: No information Detection: Mechanical transmission to herbaceous hosts. infected propagative material can disseminate the RBDV. Raspberry bushy dwarf virus infection of grapevine in Slovenia. delayed bud break and young shoots with short internodes and internal browning. 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. Japanese diseases GRAPEVINE BERRY INNER NECROSIS 1. Delaware. representing the most important virus disease in Yamanashi Prefecture. Koshu. Chlorotic mottling. and Kaiji are infected latently. Extended Abstracts 14th Meeting of IGVG. Mavric et al. if any.g. Transmission: In raspberry the virus infects progeny seedlings and pollinated plants through pollen. The vay of natural spreading in grapevine.8 x 106 Da (2. Kyoho. Campbell Early are suscetible as well as cvs Takao. being transmitted by the eryophid mite Colomerus vitis. the 3' terminal region of which (2469 nts) has been sequenced. berries are small and show external discolorations and internal necrosis.. However. Almost all Japanese table grape cultivars derived from crosses with cv.Agent: Raspberry bushy dwarf virus (RBDV) was isolated from symptomatic vines. Vitis riparia Gloire) are also susceptible. No. wt of 2 x 106 Da (5.5 Kb in size) and RNA-2 with Mol. Varietal susceptibility: Symptom severity varies with the cultivar. 1976. References Mavric I. 2003. Historical review 1976 2003 Murant: Description of RBDV. Virscek Marn and I. 3. Locorotondo 2003. and a bipartite singlestranded RNA genome accounting for c.5 x 106 Da. Ripening of bunches is delayed.. wt of 7. Some rootstocks (e. Agent: The disease agent is Grapevine berry inner necrosis virus (GINV). Healthy vines of cvs Kyoho and Pione became naturally infected in the field within one year from planting. Transmission: GINV is transmitted by grafting to grapevines and by mechanical inoculation to herbaceous hosts. Grapevine berry inner necrosis has been reported only from Japan. 24% of the particle weight and consisting of two functional species RNA-1 with Mol. RBDV. is unknow.: First record of RBDV in grapevine. GINV has filamentous particles about 750 nm in length and a single-stranded RNA genome with Mol. Zezlina. a dimeter of about 33 nm. wt 0. rings and line patterns are shown by leaf blades. 30 x 103). a mechanically transmissible definitive member of the genus Trichovirus. 165 B. Control : No information 2. Raspberry bushy dwarf virus. and Pione whereas cvs. F. Description Main synonyms: None Main symptoms: Infected grapevines have low vigor.
Y.. Bulletin of Yamanashi Fruit Tree Experimental Station 10. Kyoho back inoculated with the filamentous virus isolated from mosaic-diseased grapevines. S.Detection: Indexing on cvs Kyoho or Pione.: First account of grapevine berry inner necrosis disease in a non Japanese publication. Takahashi and Y. 1997. Purification of a filamentous virus isolated from grapevine berry inner necrosis and foliar mosaic. Magome. Yanase. Symptoms on vines.. 77-78 Yanase H... 1985.. Archives of Virology 142. H.: Experimental evidence that GINV is transmitted by the the grape erineum mite Colomerus vitis 1993 1997 2000 2000 3. Grapevine berry inner necrosis. Goto. Transmission of grapevine berry inner necrosis virus by the grape erineum mite Colomerus vitis in Yamanashi. 1. 2. Kunigi Y. Iida. T. Disease re-named Grapevine berry inner necrosis Terai et al. S. Mosaic symptoms on cv Kyoho. References Kunugi Y. Kunugi. Bulletin of Yamanashi Fruit Tree Experimental Station 10. 423. 57-63. Y. 1993. Terai and Y. varietal susceptibility and natural spread. and Terai Y. Terai. Yoshikawa et al. Terai and A. 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. H. GINV is mechanically transmissible to herbaceous hosts and can be identified by ELISA and moleculat techniques in infected vines. 536 Terai Y. and H. Extended Abstracts 11th Meeting of ICVG. Studies on the grapevine berry innner necrosis virus disease. 1992. Montreux 1993. Yanase H. Induction of berry necrosis in Kyoho back-inoculated with the virus isolate from grapevine mosaic diseased clones and renaming to grapevine berry inner necrosis. 47-56. Description Main synonyms: None. A new virus disease.: An account of the varietal susceptibility to the disease and natural field spread Kunigi et al. 1987. Studies on the grapevine berry innner necrosis virus disease. grapevine berry inner necrosis with natural spread in Japan. Control: Use of tolerant cultivars in areas where the disease spreads epidemically.. Annals of the Phytopathological Society of Japan 51. 617. Nishijima T. a new trichovirus: comparative studies with several known trichoviruses. 362. 2. 1351-1363 GRAPEVINE STUNT 1.. Back-transmission of a grapevine filamentous virus to grapevine seedlings and induction of foliar and berry symptoms in grapevine. Tanaka H. and Yanase H. Terai.Annals of the Phytopathological Society of Japan 53.Annals of the Phytopathological Society of Japan 58. 1984. 2000.: Partial sequencing of GINV genome and assignement of the virus in the genus Trichovirus Nishijima et al. Yoshikawa N. Annals of the Phytopathological Society of Japan 55.. Historical review 1984 1985 1987 1992 Tanaka: Description of a mosaic disease in cv. 114 . Y. Shinkai 2000. Asari.
S. Yora. This condition gives the name to the disease which in Japanese means "unpalatable fruits with low sugar content".Main symptoms: Spring vegetation is delayed. Yamashita and Y. S. Campbell Early. References Hatamoto M. S. 1981. 33. Hatamoto et al. curled and. S. which makes the crop unmarketable. Taipei. internodes are short. Taiwan (Reference 2951 in the Review of Plant Pathology 66. 1984. p. FFTC Book series. Annals of the Phytopathological Society of Japan 48.: Successful graft-transmission of stunt disease. Transmission of grapevine stunt disease by the grapevine leafhopper Arboridia apicalis Nawa. In Plant Virus Diseases of Horticultural Crops in the Tropics and Subtropics. The berries. Hatamoto et al. Because of heat recovery. 1986. Description Main synonyms: none. Agent: An isometric.. The disease has only been reported from Japan. 3. vinifera cv. S. The disease is apparently restricted to the V. Historical review. 1-17. Spread occurs also through infected propagative material. Annals of the Phytopathological Society of Japan 50. M. 1986 Namba et al. Iwanami. Inflorescences are undersized. Main symptoms: No appreciable symptoms are visible on the foliage of cv. Namba. Fujii. leaves are small. Doi. Yamashita and Y. Doi and K. The same paper appears in Taiwan Food and Fertilizer Technology Center Technical Bulletin 92. Doi. fruit setting is impaired and bunches are few and shelled. however.: Evidence that the disease is transmitted by the leafhopper Arboridia apicalis. 316. 1987). 2. 1982. Three phloem-limited viruses of grapevine: direct fluorescence detection. Doi and M. Fujii. Control: Use of disease-free material obtained through heat therapy. S. 396 Hatamoto M.: A small isometric virus associated with stunt disease in Japan. A small spherical virus associated with grapevine stunt disease.: Purification and characterization of the virus associated with stunt disease. 85 Namba S. sometimes. Koshu nor any apparent reduction of vigour and yield. American rootstocks are infected without showing symptoms. This virus is serologically distinct from the putative agent of ajinashika disease. GRAPEVINE AJINASHIKA DISEASE 1. Y. phloem-limited. No. Annals of the Phytopathological Society of Japan 47.. 1981 1982 1984 Namba et al. Food and Fertilizer Technology Center for the Asian and Pacific Region. summer vegetation is apparently normal.. Namba. M. non mechanically transmissible virus about 25 nm in diameter is consistently associated with diseased vines and regarded as the possible causal agent.. with scorched margins.109-126. Graft transmissibility of grapevine stunt disease. T. 137 Namba S. Transmission: The disease is transmitted by the leafhopper Arboridia apicalis. The disease has only been reported from Japan. are pale-coloured and have a low sugar content. Detection: Grafting to Campbell Early and ELISA using extracts from infected vine tissues. Hatamoto. Evidence that it is not related to the presumed agent of ajinashika disease. Yamashita. Y. Varietal susceptibility: No information. 115 . Yamashita.
Boscia. Namba et al. Yora. Three phloem-limited viruses of grapevine: direct fluorescence detection. an isometric. consistently found in infected vines. 1991.. Hatamoto. Terai: Additional report on ajinashika disease as derived from the combined effect of leafroll and fleck. 12491253. Yano. References Namba S. No relationship found with fleck. 1986. Control: Use of disease-free material obtained through heat therapy. Purification and properties of spherical virus particles associated with grapevine ajinashika disease. Annals of the Phytopathological Society of Japan 45. Terai Y. Taiwan Food and Fertilizer Technology Center Technical Bulletin 92. and R.: Further characterization of the isometric virus and claim that it is the putative agent of the disease. Y. Namba et al. 70-73. Yamashita. phloem-limited. Detection: Graft transmission to cv. 1-17. D. However. Namba S. Koshu and ELISA using extracts from infected vine tissues. Dissemination is through infected propagative material. Transmission: No vector is known. Y. vinifera cv.Agent: The disease was reported to be caused by the concurrent infection of leafroll and fleck. Niagara Falls 1980.. Iwanami. 1979. 1980. Yamashita. The disease seems to be restricted to V. 1991. 116 . S. Terai Y. Koshu. 3. S. Ajinashika disease: a combined effect of grapevine leafroll and grapevine fleck viruses on sugar content in the Japanese grape cultivar Koshu. 15-19.: First mention of ajinashika disease and report of the association with it of a non mechanically transmissible virus with isometric particles.. was suggested as the possible causal agent. 1979 1980 1986 1991 1991 Namba et al. and D. Historical review. Doi and M. Volos 1990. Doi and K. Proceedings 10th Meeting of ICVG. Tsuchizaki.: Partial characterization of the isometric virus associated with the disease and its detection by ELISA in infected vines. non mechanically transmissible virus about 25 nm in diameter. Ajinashika disease of the grapevine cultivar Koshu in Japan. 67-70. S. Plant Disease 75. Terai and Yano: Description of ajinashika disease and suggestion that it is caused by the concomitant infection of leafroll and fleck. Proceedings 7th Meeting of ICVG. Namba S. Varietal susceptibility: No information. T. Yamashita. Gonsalves. T. A small spherical virus associated with the ajinashika disease of Koshu grapevine. 2.
VIRUS-LIKE DISEASES .
They are often thicker than normal. whether they bear enations or not. omeoplasie crestiformi (Ital. which run more or less parallel to the main veins. as symptom expression is variable in successive years and graft transmission not 100 % successful. The crop can be drastically reduced (up to about 50%. but attempts to transmit it by graft or mechanical inoculation gave no results. according to the cultivar) and is of poor quality. apparently in relation with climatic conditions. maladie des énations (Fr. which gives a bushy aspect to the plant. Graft transmission suggests that it is a virus disease. the absence of symptoms does not necessarily mean that the plant is healthy. and is probably due to a virus-like agent. Enations develop mostly on the underside of the leaves at the base of the shoots. North America (California). Their agents are still unknown. some of which have a clear-cut detrimental effect on the crop. with prominent veins. Later in the year. The varieties Panse Precoce. The infectious agent of the disease perennates in propagating material. Italia. irregular and mis-shapen. with drawings of symptoms. Latin America (Venezuela). and often show longitudinal cracks between the nodes. 2. 119 . has now been dismissed Transmission: By vegetative propagation. The basal internodes are short. Symptom expression varies year by year. Hewitt: Description of symptoms in California. The disease has been reported from many European and extra-European countries Agent: The etiology of enation disease is not yet clear. Description Enation disease of grapevine is one of the oldest known disorders of European grapes. Detection: By observing the symptoms on the leaves and by indexing on LN 33 hybrid. Australia and New Zealand. Symptoms have been observed on many V. They are outgrowths 2-3 mm high and 3-5 mm long or more. with a fanlike aspect and abnormal indentation.). Primus. Gigante : Research on histological and cytological aspects of enations. All persist in propagative material and are transmitted by grafting. Main symptoms: Affected vines show a delayed opening of the buds and a slow growth of the shoots in the spring. Riesling.VIRUS-LIKE DISEASES Several latent or semi-latent grapevine diseases are known. However. There is no information on the possibility of curing this disease by heat treatment or meristem culture. This hypothesis. The disease is perpetuated by vegetative propagation. but some are heat-labile and can be eliminated by heat therapy.). North (Tunisia) and South Africa. The transmission by graft is rather erratic. ENATION DISEASE 1. its description dating back to the late 1800s Main synonyms: Enationenkrankheit der Rebe (Germ. Varietal susceptibility and sensitivity: There is little information available. are often mis-shapen. growth tends to become normal again. Control: Use of healthy material.). malattia delle enazioni. however. Severely affected leaves drop prematurely. vinifera varieties in Europe. Basal leaves. Historical review 1891 1937 1954 Buchenau: First description of enation disease of grapevine occurring in Germany. 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. Leaves developed later in the season are usually normal. Grenache and Tokay appear to be quite susceptible and develop severe symptoms when infected.
only fanleaf virus was recovered. 79-112. The mean yield loss of diseased vines ranged from 17. the proportion of diseased vines was highest in cv. McGechan: Enation disease in Australia Tekinel et al. Padilla et al. References Avgelis A. Malvasia (10. with negative results. Marinesku and Bondarchuk: Enation disease in Moldova Brückbauer: Influence of enation disease on growth and yield of grapevine in West Germany.: Enation disease in Turkey Hevin et al. Italia in Sardinia. 195 Brückbauer H. Vernaccina (1. The possible role of fanleaf virus in the etiology of enation requires further investigation.5%). Mechanical transmission tests were also negative.. Refatti: Hypothesis of a correlation between fanleaf and enation disease. Brückbauer: Description of symptoms of enation in Germany and confirmation of graft transmission of its agent. but report on observations made in Australia where no fanleaf virus could be recovered from enation-affected vines. The authors discuss the hypothesis that enation is caused by a strain of fanleaf virus.3% . 120 .: More data on the effects of enation on the yield of cv. attempts to transmit the disease by graft. and possibly caused by a virus.: In experiments on graft transmission of enation disease aimed at determining the best indicator. Presence of enations in Razaki grapevine in Crete (Greece). Confirmation of graft transmissibility of the disease. 1978. and C. Enationaffected vines produced less than 50 % of the yield of healthy plants. Nieder: Enation disease in Austria Garau et al. The authors conclude that the disease is probably of European origin.5%). historical account. In the vineyards under observation.1966 Graniti et al.: Enation disease in France Pozdena et al. Trebbiano romagnolo and reduces the yield from 13% to 23% according to the severità of symptom expression. Xafis. Martelli et al. but diseased vines which had not shown enation symptoms for several years had almost normal yields. Beobachtungen und Untersuchungen über die Enationenkrankheit der Rebe. although the rate of symptom expression does not exceed 30% Credi: Enation diseasae affects the vegetative vigour of cv.: Detailed description of macroscopic and microscopic symptoms of enation disease.: Enation disease in Czechoslovakia Avgelis and Xafis: Enation disease in Greece Prota and Garau: Enation disease found in Sardinia. 1968. Phytopathologia Mediterranea 17. the hybrid LN 33 was found to be the most sensitive and reliable indicator variety. Prota et al.4 to 48.: Successful transmission of enation disease from diseased to healthy grapevine by graft strongly supports the hypothesis of a viral origin. lowest in cv. Weinberg und Keller 15. There is some evidence that enation can be carried in the rootstocks. . Graniti and Martelli: Review paper on enation. 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.
225-246. Quacquarelli. Enations. and V. Spain. 7-12. 1954. M. U. Roma. Cugusi. 17. Lisbon 1997. Bolletino della Regia Stazione di Patologia Vegetale.G. 179-189. Bondarchuk. and M.. Rivista di Patologia Vegetale. Bitki Koruma Buletin 11. 251-252 Hewitt W. The research of the virus diseases of grapevine in CSSR. and R. Rivista di Patologia Vegetale S. Buchenau F.. Trasmissione per innesto della "malattia delle enazioni" della vite. Extended Abstracts 12th Meeting of ICVG. Chabbouh N and V. F. 1979. 1987. G. Grapevine enation disease in Murcia (Spain)..V.. Prota and M. Studies on some variable characters of grapevines affected by enation disease in Sardinia.IV.). Garau R. Ricerche istologiche sulle omeoplasie crestiformi (Enations) delle foglie di vite affette da rachitismo. 1966. Hevin M. Bericht der deutschen botanischen Gesellschaft 9. Proceedings International Conference on Virus and Vector on Perennial Hosts.. Mededel. Martelli and F. Petria 6. Gazeau. In: Virus Diseases of Small Fruits and Grapevines (A Handbook). Effetto della malattia delle enazioni della vite sulla produzione e sullo sviluppo vegetativo della cv Trebbiano romagnolo. Martelli G. Monografias INIA 18. (N. Ser. University of California. 59-64. Lisbon 1997.P. Lamberti and A. Vanek. Graniti A. Abnorme Blattbildungen. 48. Savino. Nas. 1980.K.Brückbauer H. 1976. it was clear that vein mosaic was not caused by fanleaf virus.1989. Gigante R.. Rives. Salcan. 349-352. Studies on reproduction of enation symptoms by grafting in Sardinia.S. IV. 1966. In: Verderevskaya T. Einfluss von Virusinfektionen auf Wachstum und Ertrag der Rebe. Agricultural Gazette of New South Wales 81. 1975. 47.P. 1965. 46-51. 1981.. A similar disease has been reported in Australia under the name of summer mottle. 1997. areas of the leaf blade may become necrotic. 203-206. 1971. B. 1997.P. VEIN MOSAIC 1. Deutsches Weinbau-Jahrbuch 31. 47-64. Adernmosaik (Germ. Moldavian Research Institute of Horticulture Kishinev. small fruit and grapevine in Moldavia. 1966. Cordoba. This disease is widespread and probably worldwide.. Die Eantionenkrankheit der rebe erstmal auch in Osterreich nachgewiesen. Pflanzenharzt 36. Mosaico delle nervature (Ital. 169-192. Prota U. 1973. Marinesku V. Extended Abstracts 12th Meeting of ICVG. 1970. Important virus diseases of grapevine in New South Wales. Tekinel N.. H. A study of infectious degeneration (fanleaf) in vineyards of the Mediterranean region. 1937.).W. Occurence of grapevine enations in the Moldavian SSR.. producing often a vein banding effect. And G.P.s.. Phytopathologia Mediterranea 5. Facultet Landbouwwetenschap (Gent) 40. Leclair and M. Refatti E. 326332. Description The symptoms of vein mosaic have been confused for some time with those of fanleaf/yellow mosaic. Enations of grapevine in Sardinia. G. 122-124. In the most sensitive varieties. 121 . Benayas. Garau. Garau R. Cugusi. with Special Reference to Vitis. Berkeley. Symptom expression seems to depend on climatic conditions. 1891. Bulletin of the California Department of Agriculture 43. 241-243.B. and G. 2. Credi R. 1996. A. Garcia. 823-827. Lamberti. Graniti A. Dolar.). (ed. Main synonyms: Mosaïque des nervures (Fr. Phytopathologia Mediterranea 20..D. 1983. O.. Padilla V. Kiryat Anavim. I. 1970. Vein mosaic appears to be of low economic importance. Israel.W. Nieder G. Proceedings 9th Meeting of ICVG. Davis. Division of Agricultural Sciences. 1983. 9. but these necroses do not affect the veins as it is the case with vein necrosis. Occurrence of enation disease in Tunisia. Frazier N. Enation disease of grapevine in Italy. but when fanleaf virus transmission to herbaceous hosts became possible. California. Some virus and virus-like diseases of grapevines.).) Virus and mycoplasma-like diseases of fruit trees. Enation symptoms found in France. 207-217. Main symptoms: Pale green mosaic affecting mostly the tissues adjacent to the main veins or the smaller ones. 145-152. McGechan J. Martelli. n. Su una possibile correlazione fra il virus del complesso dell'arricciamento e la malattia delle enazioni della vite. 97-98. Prota U. Ita and F. Pozdena J. Salih and Y. 293-306. Z. Proceedings 6th Meeting of ICVG. Graniti..
Milkus et al. Pinot. Babini and A. 2. 1978.R. Mycoplasma-like organisms were supposed to be the cause of vein mosaic. Marinesku and Bondarchuk: Vein mosaic in Moldova. Alphonse Lavallée. 1993. Kuniyuki: Vein mosaic in Brazil.. The disease can be eliminated by heat therapy. and Gamay apparently show little or no symptoms. Detection: Indexing with V.A. Samonina et al. Woodham and Krake: Comparison of summer mottle and vein mosaic. 266-270. Abracheva: Vein mosaic in Bulgaria.Agent: Unknown.R. Comparison of symptoms of fleck. 122 . but this hypothesis has not been confirmed. No claim is made that these putative viruses are connected with the disease.: Vein mosaic in URSS. No vector known. Legin and Vuittenez: Description of vein mosaic. 1985. Pop: Vein mosaic in Romania. show symptoms (Syrah.. 148-152. Woodham. Muscat de Hambourg. Grapevine summer mottle: a new graft-transmissible disease. Saric and Hranuelli: Vein mosaic in Croatia. Historical review 1966 1973 1973 1976 1973 1973 1978 Vuittenez et al. Chasselas. Several V. Krake and Woodham: Description in Australia of a systemic mottling syndrome which is expressed during summer on the leaves of some varieties. Krake L.: Vein mosaic in Ukraine. Vitis 17. in the absence of any detectable virus. Servant. References Credi R. Golino: Vein mosaic in California. LN 33 is also sensitive. Symptoms are very similar to those of vein mosaic in Europe.: Observation of a type of mosaic of grapevines which appears to be independent of fanleaf virus. vein mosaic and vein necrosis. vinifera cvs. Ehrenfelser showing symptoms of vein mosaic. Viognier. Transmission: By graft and vegetative propagation. Canova. Potential interaction between rootstocks and grapevine latent viruses. 1979 1980 1982 1983 1985 1993 2004 3. A number of cytological modifications primarily involving chloroplasts were observed along with the presence of bundles of filamentous structures resembling closterovirus particles. Pearl of Csaba). 17-23. Occurrence of grapevine vein necrosis and grapevine vein mosaic in the Emilia-Romagna region (northern Italy). riparia Gloire de Montpellier. A.C. and R. Control: Use of indexed material. Phytopathologia Mediterranea 24. Chardonnay. Vuittenez and Stocky: Electron microscope study of thin-sectioned tissues of leaves from Vitis riparia and Vitis vinifera cv. American Journal of Enology and Viticulture 44. Bonfiglioli: Vein mosaic in New Zealand (unpublished). Varietal susceptibility and sensitivity: Vitis riparia Gloire de Montpellier is especially sensitive and is used as indicator. Golino D.
Woodham and Krake: Comparative graft transmission trials demonstrate that summer mottle 123 . Agent: Unknown. Vuittenez A. Mataro. 2. A comparison of grapevine summer mottle and vein mosaic diseases. Kuszala. Kuniyuki H. Observations sur une Mosaïque de la Vigne. Niagara Falls 1980. Evidence that the disease is graft-transmissible.. 1983. Vinodel. 1973. Cabernet franc. Symptoms show on vegetative growth that develops at temperatures in excess of 30 °C. suspected to be a virus or a viroid. Numéro hors série. Comparaison des symptômes et transmission par greffage d'une mosaïque nervaire de Vitis vinifera .: Elimination of the disease agent by culturing fragmented shoot apices. 243-250 Samonina I. 1985. ou la “feuille rouge”. URSS. 1977. 1976. 9 . Annales des Epiphyties 17.. Mostar 1977. 57-63.V. 48-49 Marinesku V. Transmission: No vector is known. 68-72. and A. e. Main symptoms: Pale green to yellowish dicolourations of the tissues adjacent to the main or secondary veins. A. resembles in some respects the European vein mosaic and the Greek Roditis leaf discolouration (see under Minor virus diseases). R. Milkus. Ser. Spread is through infected propagative material but is has also been observed between adjacent vines Varietal susceptibility: No grapevine tested was immune to infection. S. Symptoms of vein mosaic develop under mild weather conditions and fade during hot weather. an Australian disease. several European grape cultivars show symptoms.V. Bondarchuk. La mosaique des nervures. Legin and J. Bunches of infected cvs Sideritis and Cabernet sauvignon are fewer. Vuittenez. probablement indépendante du virus du Court-noué.Legin R. 149-141. Rivista di Patologia Vegetale.. Historical review 1978 1982 1983 Krake and Woodham: Description of summer mottle in Australia. Cabernet sauvignon. Vuittenez A. Pop I. These symptoms appear in summer and persist through the autumn..-Berl.R. Rivista di Patologia Vegetale S IV 9. Krake. and V. 110 R. B. Stocky. IV . Saric A. and G. Woodham R.g. 191-204. Mission. Barlass et al. Krylov and V. Proceedings of the Conference on Excoriosis and Virus Diseases of Grapevine.. A grapevine virus disease in the Primorye territory. 1973. whereas the opposite occurs with summer mottle. Rivista di Patologia Vegetale . 1982. Control: Use of disease-free propagating material obtained by culture of fragmented shoot apices.C. However. poorly developed and with small berries. IV. Detection: Graft transmission to a number of cvs. 41-42. SUMMER MOTTLE Summer mottle. Proceedings 7th Meeting of IGCV. and Hranuelli T. de la marbrure de V. 247-252. producing a feathering or banding effect. 1973. Moldavii 31. 1966.V. Krylova. rupestris and LN33 are infected symptomlessly. Investiation on grapevine viruses in the SR Croatia.N.G. V.N. Adverse effect of light and of high temperature on symptom expression of grapevine vein mosaic in Sao Paulo. Description Main synonyms: None. rupestris et d'une affection nécrotique des nervures de l'hybride Rup. 1.. Roditis leaf discolouration and summer mottle have similarities suggesting that they may be the same disease. 67-73. Ultrstructure de vignes infecteés par deux maladies de type viral: la mosaique des nervures. Vinogradr. Summa Phytopathologica 11.V. Grapevine vein mosaic. Vitis 22. and L. 9. maladie à virus de la vigne.
Vitis 17. So far. Varietal susceptibility and sensitivity: The rootstock 110 R is most sensitive. 1. This strongly supports the hypothesis that vein necrosis is a specificic reaction of the rootstock 110 R to GRSPaV infection. Krake L.differs from vein mosaic. 2. Many grapevine varieties and rootstocks are infected symptomlessly. K. Transmission: By grafting and vegetative propagation.R. 1999. especially under greenhouse conditions. Mycoplasma-like organisms have been observed in the phloem of symptomatic vines. Historical review 1973 1978 1978 Legin and Vuittenez: Discovery and description of this virus-like disease while searching for indicators for fleck. and L. Detection: By grafting on 110 R. 291-295. Rezaian and R. berlandieri 110 Richter (110R) with vein necrosis symptoms. Annals of Applied Biology 101. Description Vein necrosis has probably a worldwide distribution. Collingwood. Necrotic reactions are best seen on the lower face of the leaf blade. 70-74. Woodham. Krake.: Suggestion that summer mottle and Roditis leaf discolouration are the same disease 3.. recently an association exceeding 95% has been experimentally observed between Grapevine rupestris stem pitting-associated virus (GRSPaV) and vines of V. 1983. R. 1978. Woodham R. Main symptoms: On the rootstock 110 R.A. Scott. M.H. 266-270. Graft-transmissible Diseases of Grapevines.). Grapevine summer mottle: a new graft-transmissible disease. Krake L. at first on the leaves at the base of the shoots. Taylor.. CSIRO Publishing. The agent of vein necrosis can be eliminated by heat therapy. Woodham and L.C. Vitis 22. RT-PCR with virus specific primers and Western blot with an antiserum to recombinant coat protein of GRSPaV allow sensitive and reliable detection of this virus in symptomatic 110 R plants. Martelli et al.C.C. varieties or hybrids. Australia. Skene. Control: Use of indexed planting material. Krake. 247-252. and R.. Possible viroidal etiology put forward. Milkus and Kalashyan: Mycoplasma-like organisms found in phloem tissues of vines with vein necrosis. but their etiological relationship with the disease has not been proven. its economic importance has not been assessed. VEIN NECROSIS Although vein necrosis is currently regarded as a disease in its own right. growth is much reduced and necrosis of the leaf veins appears. necrosi delle nervature (Ital.: Vein necrosis in Italy and Bulgaria 124 .M.R. N. Little is known about sensitivity of other Vitis species. later on younger leaves as they develop. and the only Vitis species that is clearly affected is the rootstock 110 R. most likely GRSPaV. Main synonyms: Nécrose des nervures (Fr. Also the tendrils and many shoots can necrotize.).S. and some infected plants may die. No vector known. Regeneration of virus-free grapevines using in vitro apical culture. Agent: Suspected to be a virus.G.R.).R. rupestris x V. 1982. A comparison of grapevine summer mottle and vein mosaic diseases. 1999 Krake et al. Cause-effect relationships between MLOs and the disease has never been ascertained. Adernnekrose (Germ. References Barlass M.
Journal of Turkish Phytopathology 17. 1988. Necrosi delle nervature della vite in Italia e Bulgaria. Savino.. 148-152. Lehoczky et al. Incidence of some graft-transmissible virus-like diseases of Grapevine in visually selected and heat-treated stocks from Southern Italy. D. Vein necrosis: new virus-like disease in Turkish vineyards. Is Grapevine vein necrosis a reaction to Grapevine rupestris stem pitting-associated virus? Journal of Plant Pathology 86.Z. the incidence of vein necrosis in visually selected stocks of table and wine grape varieties is on the average 71 %. H. Pirolo. 1986.. 1992. Savino.: Vein necrosis in Hungary Gursoy: Vein necrosis in Turkey Rumbos: Vein necrosis in Greese Martelli et al. 1973. 79-83 Legin R. P. References Bouyahia H. A.R. 29-30.5 %. Boscia and V. Vein necrosis a disease that is latent in most grapevine cultivars of the State of Rio grande do Sul.-Berl.. SSR. and J. Suggestion than vein necrosis is a specificic reaction of 110R to GRAPaV. 9.A.. 1978. Rosciglione.. Grapevine vein necrosis disease detected in rooststocks in Australia. Informatore Fitopatologico 28 (10). D. Martelli G. Boscia.P. Martelli. 204-207. 1993.A. and L R. Comparaison des symptômes et transmission par greffage d'une mosaïque nervaire de Vitis vinifera. Krake. ser. V.C. and L. Abracheva and B. No veing necrosis observed in 110R top grafted on GRSPaV-free V. Biol.P. 58-60.: Vein necrosis in Malta Golino: Vein necrosis in California Khun: Vein necrosis in Brazil Bouyahia et al. Mycoplasma-like bodies in phloem tissue of grapevine affected by vein necrosis. de la marbrure de V. Lazar. Izvestiya Akademii nauk Moldav. Fitopatologia Brasileira 19. Minafra and G. V. Journal of the Australian Institute of Agricultural Sciences 50. 1984. Woodham R. D. 2004. Farkas. rupestris.P. Phytoparasitica 17. Phytopathologia Mediterranea 24. but did not eliminate the disease entirely. 57-63. 43-45 Kuhn G. J. 1985. Galea Souchet. La Notte. G. Potential interaction between rootstocks and grapevine latent viruses. rupestris et d'une affection nécrotique des nervures de l'hybride Rup. Milkus B. Gursoy Y. Castellano.1984 1985 Woodham R.P. 125 . Rumbos I C. 59-65. 1994.B.A. Lehozcky J. Vein necrosis. Boscia and G. 1978. Kalashyan. IV. Savino. 1989. Bulletin OEPP/EPPO Bulletin 22. and A. fleck and leafroll in Vitis vinifera and grapevine rootstocks in central Greece. 1986 1988 1989 1992 1993 1994 2004 3. 606-612.: In southern Italy. Golino D.. i Chim Nauka 1.: An association exceeding 95% observed between GRSPaV and 110R vines showing vein necrosis symptoms in indexing trials... 110 R. 3-5 Martelli G. Detection of vein necrosis virus (GVNV) in the vines of cultivated grape varieties. Vuittenez.. M. Rivista di Patologia Vegetale. 61 Savino V. Kergazdasag 18 (4). Krake: Vein necrosis in Australia Savino et al.C.N. Viruses of grapevine in Malta. C. P. Martelli. Heat therapy reduced this proportion to 35. American Journal of Enology and Viticulture 44. Ser. 301.
VIROID (Yellow speckle) .
They are made up of a non encapsidated circular RNA of 246-375 nucleotides. thought to be elicited by a specific strain of GFLV. are subviral pathogerns endowed with autonomous replication in their hosts. 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. the type species of the genus Hostuviroid. Viroids. Flores. it did not induce symptoms. presence of ribozymes.). AGVd. all of which belong in the family Pospiviroideae: Grapevine yellow speckle viroid 1 (GYSVd-1). Gelbsprenkelung der Rebe (Germ. HSVd-g. respectively and both belong in the genus Apscaviroid. Collingwood. plant age. 129 . 2003. Vein banding. the USA. Two families are known. Citrus exocortis viroid grapevine strain (CEVd-g). infected vines are symptomless or show symptoms erratically. and may have a worldwide distribution. CEVd-g. Europe. It was first detected in Japan and transmitted to cucumber and grapevine seedlings in which. Main symptoms: Few to many minute chrome yellow spots or flecks scattered over the leaf surface. north and south America. AGVd has been reported from Australia. Agents: Two distinct viroids. genera and species. and plastidial replication (Avsunviroideae ). YELLOW SPECKLE 1. Description Main synonyms: Moucheture jaune (Fr. a size much smaller than that the smallest viral genome. Australian grapevine viroid (AGVd). Neither of them is able to replicate in herbaceous hosts but both were succesfully inoculated to grapevine seedlings reproducing the yellow speckle syndrome.S. viroids are classified in families. a disease characterized by chrome yellow flecks localized along the main veins of mature leaves and progressing into the interveinal areas.. GYSVd 1 and GYSVd 2 are made up of 366 and 363 nucleotides (nt). was demonstrated to be caused by a co-infection by yellow speckle viroids and GFLV. GYSVd-1 and GYSV-2 have a worldwide distribution The three additional viroids that have been detected in grapevines. and Tunisia. HSVd-g has been recorded from Australia.). 370 pp. or gathering along the main veins to give a vein banding pattern. Five grapevine-infecting viroids are known. respectively from a cv. GYSVd-1 and GYSVd -2 cause the disease individually or in combination. has a genome 297 nt in size. The symptomatology varies depending on the cultivar. Sometimes. CSIRO Publishing. are not associated to any specific symptomatology. References Hadidi A.). climatic conditions. Only GYSVd-1 and GYSVd-2 are pathogenic. the non coding genomes. picchiettatura gialla (Ital. phylogenetic analysis of hop and grapevine isolates of HSVd has provided evidence that the viroid that causes hop stunt disease in Japan is a variant of HSVd-g. The suggestion is that HSVd moved from grapevine to hop probably 50-60 ago in the Nagano and/or Fukushima prefectures in which it is not uncommon to find hop gardens next to vineyards. however. Semancik (eds. These symptoms appear in the height of summer on a limited number of mature leaves and persist for the rest of the vegetative season. Cabernet franc and a cv.). inducing a disease called yellow speckle.W. vein banding-like symptoms can be observed in vines infected only by yellow speckle viroids. Kyoto vine with yellow speckle symptoms. Hop stunt viroid grapevine strain (HSVd-g). and perhaps the type of infecting viroidal sequence variant. has a genome 369 nt in size.VIROIDS Viroids. 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. Interestingly. R. J. Like viruses. however. Grapevine yellow speckle viroid 2 (GYSVd-2). a member of the genus Apscaviroid. Randles and J. Both these viroids wer first isolated in Australia. Very often. and AGVd. HSVd-g.
First identification of AGVd Koltunow and Rezaian: Identification and sequencing of grapevine yellow speckle viroid. Cannonau not associated with the presence of GFLV reported from Italy. 1984 1985 1985 1985 1987 1987 1988 1988 1988 130 . Varietal susceptibility: All Vitis species. found in grapevine accessions from Europe and California. Abracheva et al. Transmission: No vector is known. Prota et al. a member of the genus Pospiviroid. In the great majority of grapevine germplasm infection is latent. Seed transmission has been demostrated for GYSVd-1. 2. Woodham and Krake: Artificial transmission of grapevine leafroll.: The Japanese grapevine viroid identified as a strain of hop stunt viroid. Natural dissemination takes place by mechanical inoculation through surface-contaminated cutting tools during management operations. besides Spain.: Yellow speckle eliminated by in vitro apical culture.: Reconstruction of the secondary structure of CEVd-g Szychowski et al. grafting.: Evidence that viroids are widespread in grapevines. Rcatzitelli resembling yellow speckle reported from Bulgaria. It was first recoverd in Spain from symptomless grapevines. Detection: Some viroids can be transmitted mechanically to herbaceous hosts but this is not an efficient detection method. CEVd-g and AGVd. Experimental transmission through dodder is possible. and distribution of infected propagating material. Although CEVd is present in most. a disease formerly thought to be caused by a chromogenic strain of GFLV. has a genome 369 nt in size.: Four viroids found in Australian grapevines. 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. yellow speckle and fleck through dodder.CEVd-g. Sano et al. one of which identified as the agent of citrus exocortis. if not all citrus-growing countries. hybrids and cultivars appear to be susceptible. GYSVd-2. Semancik et al. For yellow speckle.: First recovery of a viroid from grapevines in Japan. Krake and Woodham: Evidence that the agent of yellow speckle is implicated in the etiology of vein banding. Flores et al. Control: Use of viroid-free propagative material obtained by meristem tip culture. Three different viroids found in a number of accessions in a Californian varietal collection.: Two new viroids. Garcia Arenal et al. Woodham and Krake: Evidence of field spread of yellow speckle. Historical review 1972 1975 1978 1982 1982 1983 1983 Taylor and Woodham: First description of yellow speckle as a graft transmissible disease separate from chromogenic disorders induced by grapevine fanleaf virus (GFLV). Mink and Parsons: Yellow speckle can be detected by growing vines for 2-3 weeks at 32 °C under continuous illumination. its grapevine strain so far has only been recorded from Australia and the USA. the authors consider the results as inconclusive.: A vein banding condition of cv. Shikata et al. This latter way of dissemination has been considered as more efficient and frequent than mechanical transmission.: Successful mechanical transmission of viroids to grapevines. Rezaian et al. Barlass et al.: A disease of cv. as the disease may have spread naturally.
Molecular evidence that this viroid originated from recombination between five different viroids among which GYSVd-1 and GYSVd-2 1991a. Grapevine yellow speckle viroid 2 (GYSVd 2) Australian grapevine viroid (AGVd). Citrus exocortis viroid grapevine strain (CEVd-g). 291-295. A. V. Koltunow et al. 127-130.: Review of viroids. Juarez and J.. based on phylogenetical analysis of hop and grapevine isolates of this viroid. G. (ii) enhanced viroids. Skene. Martelli: Brief review of grapevine viroid situation supporting the idea that vein banding is primarily induced by viroidal rather than GFLV infection. The occurence is reported of HSVd.M. Little and Rezaian: Updated review of grapevine viroids. 2095-2102.: Evidence that two related viroids (GYSVd 1 and GYSVd 2) can cause yellow speckle disease independently.M. Spain. 1988. Sano et al. Monografias INIA 18. R. Wang et al. Pallas and J. K. Proceedings of the 6th Meeting of ICVG. Production of viroid-free grapevines by shoot tip culture. Flores et al. GYSVd-1 and GYSVd-2 Rezaian: Complete nucleotide sequencing of AGVd. Marrakchi.: First record of grapevine viroids in China. Semancik. Journal of General Virology 66. American Journal of Enology and Viticulture 39. Elleuch A.: A survey of viroids of grapevine in Italy.. Journal of Plant Pathology 85. J.S.1988 1989 1989 1989 1990 1990 Duran-Vila et al. Koltunow and Rezaian: Description and sequencing of grapevine viroid 1B (later renamed Grapevine yellow speckle viroid 2).: Improvement of meristem tip culture technique for the production of viroid-free grapevines. Duran-Vila N. M. 1985. First report of Australian grapevine viroid from the Mediterranean region. 1991 Semancik and Szychowski: There are two classes of grapevine viroids: (i) apparent viroids. 45-57.P. These viroids. Flores R. Greece. Woodham and L. Detection of viroid and viroid-like RNAs from grapevine.P. Cordoba. 2003.R. Martelli.. Rezaian et al. GYSVd-2.b Szychowski et al. Wan and Symons: Transmission of GYSVd-1. Quacquarelli and V. CEVd-g and AGVd via grape seeds. which can readily be isolated directly from grapevines. which require amplification in an alternate host.C.: First report of AGVd in the Mediterranean. Duran-Vila. Grapevine yellow speckle viroid 1 (GYSVd 1). 1991 1996 1997 1999 2001 2003 2003 3. 217-220. Krake. Regeneration of virus-free grapevines using in vitro apical culture. are to be named as follows: Hop stunt viroid grapevine strain (HSVd-g). according to an international agreement reached during the 10th Meeting of ICVG held in 1990 at Volos. Minafra et al. 131 . 1976. Elleuch et al. Fakhfakh.G. Savino. J.. Arregui. N.: Suggestion that the viroid causing stunting in hop (HSVd) originated from grapevines. A possible virus disease of Rcatzitelli vines in Bulgaria. Barlass M.: Extensive comparative analysis of grapevine accessions from California and Europe reveal a similar pattern of viroid distribution. References Abracheva P.: Structural analysis reveals that five distinct viroids infect commercial grapevine varieties.. 1982. 1978. Annals of Applied Biology 101. Perreault and H.
Flores. Mimura and K.A. Rezaian. Rezaian M. Arab Journal of Plant Protection 7. 1972. S. and L.A. Sano T. Koltunow A. Krake.R. 5587-5598. Hong.. Krake L. V.A. Two related viroids cause grapevine yellow speckle disease independently. 285-291 Wang G. and M.. 2001. 370 pp.A. Investigations on a vein banding disease of Grapevine in Sardinia. 1975. 849-864. Vitis 22. Semancik. 297. Greece.A. 1990. J. 270-278. 1991.A. Cugusi and M. A. and R. 1985. 575-578. Rezaian M. 1985.P. yellow speckle and fleck diseases by dodder. Symons. Garcia Arenal F. Goheen and J. Minafra. A. 869-872. and J. Relationships among grapevine viroids from sources maintained in California and Europe.W.S. Koltunow and L.. Koltunow.S. Uyeda. Viroids: the non coding genomes. Ohshima. Grapevine yellow speckle viroid: structural features of a new viroid group. J. Collingwood. Semancik J. R. 53-59. Vitis 29. Savino. 1983.R.. 3411-3419.C. The sequence of a viroid from grapevine closely related to to severe isolates of citrus exocortis viroid.A. A viroid-like RNA isolated from grapevine has high sequence homology with hop stunt viroid. Journal of General Virology 66. Woodham R. Volos. Doazan. Nucleic Acids Research 10. 173-182. Shikata E.P. Relationship and patterns of distribution among grapevine viroids from California and Europe. 1999.L. Grapevine viroids. 1990. Nucleic Acids Research 15. Journal of General Virology 69..A. American Journal of Enology and Viticulture 39. 24-28. N. Sano T. Garau. Koltunow A. Investigations on transmission of grapevine leafroll. Semancik (eds. Duran-Vila. Journal of Phytopathology 147. P. Szychowski J. Phytopathologische Zeitschrift 106. and L. Shikata.I. Proceedings of the 10th Meeting of ICVG. Minafra..A.A.C. China. Woodham R. I. Widespread occurrence of viroid-like RNAs in grapevines. Nucleic Acids Research 16. 193-198. Skene. 1990. Krake and K. L. R. Grapevine yellow speckle disease: studies on natural spread observed in the field..P. L. Little A. 1982. Duran-Vila. Wolpert and J. sanitation and situation in the Arab countries.. Semancik J. 202-205. 1987. 35-40. Garnier. Semancik. Greece. Szychowski J.M.R. Infectious diseases of grapevines. Vitis 30.M. 1997. Rezaian. 132 . 195-206. 40-50.P. Mink G. Martelli G.C. Phylogenetic analysis of hop and grapevine isolates of hop stunt viroid supports a grapevine origin for hop stunt disease. Volos. E. Grapevine yellow speckle agent implicated in the aetiology of vein banding disease. Leclair. J. M. 1984.G. Vitis 21. N. T.a newly recognized grafttransmissible disease of Vitis. In: Viroids (A. Credi. 1990. Zhang. CSIRO Publishing. Proceedings 10th Meeting of ICVG. 1991b. Australian grapevine viroid . Randles and J. Rezaian M. Wan C. P. Rezaian. A. 25-36.D.R. M.P. Grapevine viroids. 1990. Ohno and Y.. Prota U.M. R. Dore. A. Virus Genes 22. An infectious low molecular weight RNA was detected in grapevines by molecular hybridization with hop stunt viroid cDNA. American Journal of Enology and Viticulture 38. J. S. detection.S. R. Phytopathologia Mediterranea 24.. Sano and I. Nature.A.A. Flores.S. J. Comparative properties of viroids of grapevine origin isolated from grapevines and alternate hosts. Hernadez. Semancik. Grapevine yellow speckle -. 1988. Krake. China Fruits 4. Woodham. Krake.Jiang. 1988. T. 413-422.R. Szychowski J.Leclair.C. 1987. Koltunow A. 1988. Australian Journal of Agricultural Research 23.. 213-216. 2003. and J. Okado. Zhang and X. Krake. Taylor R.H.evidence for extensive recombination between viroids. Viroids of grapevines in Italy. 65-73. Wolpert and J. Pallas and R. Journal of General Virology 70. and R.C.R. 39-41.. Goheen.H. T.. A. Isolation of three viroids and a circular RNA from grapevines. Meshi. Transmission of viroids via grape seeds. and M.. Seminars in Virology 8. Hadidi..S. R. G. Uyeda. 1989.Woodham. Doazan. Volos. Minafra A. 287-288. 1989.A.Flores R. Johnson and M. and M. 1983. Z. Rivera-Bustamante and A. F. 210-219. Plant Disease Reporter 59. Di Serio and C. 1996.). 337-345. 333338. Greece. R. 1989. 1991a. M. 447-452. Szychowski. N.C. 1991.. Credi. and R. 4203-4210.. Parsons. Rezaian.S. Mechanical transmission and rootstock reservoirs as factors in the widespread distribution of viroids in grapevines. a new member of the apple scar skin viroid group contains the left terminal region of tomato planta macho viroid. Rapid indexing procedures for detecting yellow speckle disease in grapevines. Garnier. Virology 170. Grapevine viroid 1B.M. Proceedings of the Japanese Academy of Science 60(B).W. The field investigation of virus and viroid diseases of grapevine and stone fruit trees in Shandong and Liaoning province. Martelli and V.M. Proceedings of the 10th Meeting of ICVG.
though some sieve tubes may appear crowded with phytoplasma cells in the electron microscope. after their aetiology had been understood and molecular methods for the characterization of associated phytoplasmas had been developed. host range. a phytoplasma belonging to group 16SrII (or peanut witches' 135 . develop on leaves. Quite often. shoots. Leaf blades are crispy and brittle. Vergilbungskrankheit. 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. However. associated to Bois noir (BN). inflorescence and berries. Bunches wither in early summer or berries shrivel later in season. The occurrence and spread of a particular GY mainly depend on the presence of efficient vectors in the vineyards or their close vicinity. Their titre is usually very low in grapevine. Phytoplasmas form a homogeneous phylogenetic clade subdivided into about 20 groups and subgroups. in addition to other criteria such as symptomatology. Schwarzholzkrankheit. Severely infected stocks decline rapidly. Flavescenza dorata. Phytoplasmas have the smallest genome reported for procaryotic organisms (560 2. Golden flavescence. The first GY to be recorded was Flavescence dorée in France in the 1950's. Several Candidatus phytoplasma species have been recently described. Australian grapevine yellows. Legno nero. Red-berried varieties show reddish to purple discolorations while white-berried varieties show golden to chlorotic discolorations. Main symptoms: Young shoots of affected Vitis vinifera are weak with short internodes and frequent necrosis of terminal buds. involving also the main veins. However. However. Flavescenza dorata. They are wall-less. canes.200 kbp). Stolbur phytoplasmas. The main characteristics of GY diseases are important losses or damage to the yield. Main diseases: Flavescence dorée. such as the ribosomal protein genes or the elongation factor Tu. The different GYs cannot be differentiated on the basis of symptoms. depending on the particular disease and other conditions such as variety. resulting in reduction of quantity and quality of the crop. Bois noir. Nevertheless. Phytoplasma cells are vesicular rounded bodies. buds. but not seeds. Description Grapevine yellows (GYs) are a group of severe phytoplasma-induced diseases occurring worldwide on a number of Vitis vinifera cultivars. Palatinate grapevine yellows. Flavescence dorée (FD) and Palatinate Grapevine Yellows (PGY) phytoplasmas belong to group 16SrV (or Elm yellows group). In 1997.GRAPEVINE YELLOWS: GENERAL PROPERTIES 1. based mainly on sequence similarity of their rRNA genes and of a few other genes. Amarilliamento. including roots. persistence of infection in dormant plant material. vector species have not been identified for all GY diseases. 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. and transmission by specific leafhopper or planthopper vectors. 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). Nonetheless. Vergilbungskrankheit. Candidatus Phytoplasma australasia. phloem-restricted bacteria that belong to the class Mollicutes. Downwards rolling of the leaves and sectorial discolorations of the blades. North American grapevine yellows. Phytoplasmas in the 16SrIII group (or X-disease group) are associated with grapevine yellows in North America and Israel. GY agents have been identified in no less than 5 groups of phytoplasma clade. 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. Rubbery canes fall downwards with a typical "weeping" aspect. their movement is slow and their distribution in the plant is erratic. their genome is poorly known because they cannot be cultivated. it was mainly in the 1990's that GYs could be differentiated. autumn fall of leaves occurs later on diseased than on healthy vines. and serology. climate and infection pressure. Main synonyms: Flavescence dorée-like disease. a severe decline of the vine. the term phytoplasma has become the genus name of these plant pathogenic agents under the provisional taxonomic status Candidatus. all organs of the plant may be infected.
more specific primers can be used. Detection can also be made by transmission electron microscopy of thin sections or by scanning electron microscopy. When no information on the particular phytoplasma type is available. are available in the literature. Outstanding progress in detection was achieved with DNA-based techniques. Phytoplasmas persist in their vectors but vertical transmission to the progeny of infected gravid females is not possible or scarce. Though the rate of transmission to plant material can be very low. vinifera cultivars have been reported to be sensitive or very sensitive to GYs in all countries where infection occurs. 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. using DAPI staining. including the agents of FD and BN. When the presence of a particular disease is suspected. Most of these PCR primers have been designed on conserved regions of the rRNA gene and are "universal" for all known phytoplasmas. Hence. designed on variable regions of the rDNA. Pinot noir. Simultaneous presence of symptoms on leaves. The three ascertained vectors of GY diseases are univoltine leafhopper or planthopper species. influence symptom expression and differential sensitivity of cultivars. They can also be observed in sieve tubes by light microscopy. The best antigen source are leaf veins and petioles. ELISA detection is possible from vascular tissue of symptomatic grapevines. or on random selected non-ribosomal DNA fragments. are critical. Individual symptoms can be confused with other diseases or disorders. Vitis riparia and American rootstock varieties and hybrids do not show typical symptoms of GY. However. The situation of cv Syrah is controversial. Particular procedures for extraction of phytoplasma antigens from infected vines must be used to enhance the sensitivity of detection. It is probable that the feeding preference of insect vectors. Cabernet Sauvignon.broom group) is a second agent of Australian grapevine yellows. These phenomena also depend on the disease complex (phytoplasma. are very sensitive to all GY diseases. are group specific. such type of transport is risky when potential insect vector species occur on the site of planting. 136 . Varietal susceptibility and sensitivity: Numerous V. Some cultivars such as Chardonnay. The extraction from infected tissues of total DNA containing enough phytoplasma DNA of good quality and the elimination of inhibitors of the PCR reaction. Then. Infection of rootstocks has been detected only in the case of Flavescence dorée but it cannot be excluded for other GY agents. shoots. Detection: GY syndrome is characteristic. vector and abundance of reservoirs). erratic transmission to grapevine may nevertheless take place. Full recovery and transient remission of symptoms have been observed in low and medium sensitive cultivars. A range of primers that can be used for amplification with Polymerase Chain Reaction (PCR) of characteristic DNA fragments of phytoplasmas. Transmission: Phytoplasmas are vectored by hemipters. Phytoplasmas also persist in vine stocks during winter. These methods are not specific for any phytoplasma and are too laborious for disease monitoring. but the latter technique has never been successfull on field-grown grapevines. mainly hoppers (cicadellids or cixiids) or psyllids. 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. Others. Riesling and also numerous local varieties. on less conserved genes. Phytoplasmas can be detected by graft transmission to sensitive vines. Phytoplasmas in the 16SrI group (or aster yellows group) are associated with endemic GY diseases in several countries of Europe and in the USA. when grapevine is not a preferred host for the vector. which allows restoring the sanitary status of affected vineyards when the vector activity is controlled. Random Fragment Length Polymorphism (RFLP) analysis of the amplicon can provide further characterization of the infecting phytoplasma. PCR amplification with universal primers is preferred. Double infections have been reported. Polyclonal and monoclonal antisera have been raised to a few phytoplasmas. as well as their feeding activity. canes. Propagation material may be the infection source for long distance dissemination of GY diseases.
and on the identification of reservoirs of inoculum such as infected grapevines. efforts should address prevention by planting healthy propagation material and limitation of vector populations and their inoculative activity. and on elicitors of defence reactions to these phloem-restricted bacterial agents. Caudwell: Characterization of a new type of flavescence. and natural enemies. but is also possible using young leaves shortly before symptom expression or phloem scrapings from lignified canes. Control methods of vector populations are being experimented with natural insecticides. Caudwell (b): Description of recovery in grapevines affected with FD. BN is considered as a non epidemic form of FD. under the name Flavescence dorée. The disease can be transmitted by grafting and is probably a virus disease. Generally. Historical review 1955 Levadoux: Description of a severe outbreak of "Flavescence dorée" on Baco 22A in southwestern France. of a disease occurring in the vineyards of the Mosel Valley and of the Rhine Valley. However. Vidano: S. evolution of the disease in space and time. using detection of PCR amplification products with DNA-DNA hybridization with a specific probe. Investigations are being conducted on sensitivity. Nonsymptomatic grapevines that show irregular symptoms from one year to the next may also test positive. Production of sanitized propagation material is possible with hot water treatment (HWT). littoralis Ball found in Italy in 1963. i. Control: There are no direct curing methods of infected plants. into hot water (50 °C) for a sufficient length of time (minimum 30 mn). Branas (a and b): Description of flavescence epidemics on Baco 22A and Chardonnay in France. 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. on physiological changes and defence mechanisms induced by infection. hypotheses on its nature. Detection in the upper parts of the vines is usually done from veins and petioles of the leaves of symptomatic shoots. cultural practices. 1956 1957 1961 1961 1961 1964 1965 137 . tolerance and potential for recovery of varieties.. Detection of phytoplasma DNA can be achieved from all plant organs or insect vectors. 2. denoted Flavescence dorée (FD) in agreement with the name proposed by Levadoux (1955) because of the golden yellow metallic aspect of leaves. soaking of dormant material before or after grafting. on the possibility to limit or eradicate their population and prevent their migration or movement in vineyards. The disease is thought to be caused by root damage. In any case. Control of GY diseases depends on the knowledge of vector insect species. The author suggests this disease to be due to adverse climatic and soil conditions. FD is transmitted by the leafhopper S. littoralis Ball. Pruning or top-grafting are useless because roots and trunks are infected.Other molecular methods are being developed. Symptoms (macroscopic and microscopic). Schvester et al. Caudwell (a): Studies on Bois noir (BN) and on its relationships with FD. phytoplasmas are unevenly distributed in GY-affected vines.e. an insect that has been introduced recently from America. but does not rule out the possible involvement of a pathogen. Gärtel: Description. FD can be detected from leaves of non-symptomatic American roostocks. weeds or other crops. The biology of the insect is described. Real-time PCR has also been successfully used.
Bois noir. titanus found in 1984 in vineyards of northern Italy. An important FD-like disease is reported. and Euscelis sp. First record in 197576. Credi and Babini: Grapevine yellows are reported in Emilia Romagna (Italy). faba plants produced typical GY symptoms. Belli et al.1966 Caudwell: Attempts to inhibit the agent of FD in vivo by heat treatment. mainly on cv. The author assumes that both FD and its vector were introduced from North America with varieties of Vitis labrusca between 1927 and 1950. with special reference to grapevine. Caudwell and Larrue: Spread of FD in France is related to sanitary status of planting material. This phytoplasma was later on identified as a Clover phyllody phytoplasma. Inoculation of grapevine seedlings with S. the hypothesis is put forward that this nematode is the vector of the disease. Caudwell et al. littoralis. Conversely. titanus (= littoralis) is not a vector. titanus fed on the latter V. Granata: Description of an epidemic yellows disease on cv Inzolia in Sicily.: S. of which S. Caudwell et al. Caudwell: Discussion on the origins of yellows diseases of plants. Regina.: Discovery of Mycoplasma or PLT group-like microorganisms in the phloem elements of plants showing dwarfing.: Rickettsia-like organisms observed in roots of grapevines with "Vergilbungskrankheit" in the Saar. faba are different from those obtained with feeding inoculation with infective S. these findings have not been confirmed. Tanne and Nitzany: Occurrence of a yellows disease resembling FD in Israel. Baggiolini: S. Belli et al.: 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. Rumbos et al. Mosel and Rhine regions are considered as the causal pathogens of this disease. littoralis.) caught in the vicinity of vineyards could transmit a yellows disease to Vicia faba plants. The disease has been observed for the first time in 1968. Osler et al. littoralis Ball is present in Ticino (southern Switzerland). Immersing cuttings taken on FD-affected grapevines in water at 30°C prior to planting reduced the proportion of infected plants by about 83%. Doi et al. Recommendation that mother plants should be grown in areas far away from FD-infected regions. littoralis is present in the same region of Oltrepò Pavese. is probably of European origin. Caudwell: Symptoms of grapevine yellows (Amarilliamento) reported in Chile on cv Elqui. Caudwell et al. As similar organisms have been found in nematodes of the species Xiphinema index. Magarey and Wachtel: Description of a new disease of grapevine of the yellows type on the cv Rhine Riesling in South Australia. witches' broom and yellowing.: S. 1967 1968 1970 1971 1971 1973 1973 1975 1977 1979 1980 1982 1982 1983 1984 1985 1985 138 . Potential existence of several different diseases: leafhoppers (Euscelidius sp. Symptoms on V. (b): Evidence that FD and BN are two different diseases with similar symptoms. (a): Mycoplasma-like organisms (MLOs) are observed in diseased grapevines and infective leafhoppers and in Vicia faba submitted to feeding inoculation with infective S. Provisory name "Rhine riesling problem".: Presence of a disease similar to FD in the Oltrepò Pavese (northern Italy). Rafaila and Costache: Yellows of grapevine with symptoms similar to those of FD found in 1967 for the first time in Romania. So far.
northern Italy. Quaroni et al.: Survey of potential Auchenorryncha vectors of the pathogen agent of FD in Piemonte (Italy). Italy. Hyalesthes obsoletus. Italy. whereas an unsprayed vineyard was more affected. titanus is not always associated to the disease. Mescalchin et al. Pinot nero) and comparison with other similar diseases. titanus. that are responsible for severe decline of vines. Credi and Callegari: Survey of vineyards in Emilia-Romagna. of a FD-like disease on Chardonnay. for the presence of a FD-like disease. Credi: Description of a FD-like disease in Emilia Romagna and the reaction of 3 different cultivars.or xylem-limited prokaryotes in Europe. However. Recovery and crop loss are variable.: Observation of MLOs in phloem tissue of yellows-affected grapevines in Australia. in viticultural areas of northern Italy and of other Auchenorrhynchas susceptible to play a role in the transmission of yellows diseases in this region.: Description in Italy of the presence of FD or FD-like diseases.: New data on the spread of FD in three vineyards of Oltrepò pavese (northern Italy) from 1985 to 1987. Two of these vineyards were sprayed with insecticides in 1986 and 1987. The results suggest that the disease is brought into the vineyards from outside local sources. control measures. Carraro et al. Fortusini and Belli: Description of the development of epidemics of FD-like diseases in northern Itay. titanus.: Presence and distribution of a FD-like disease in the region Friuli-Venezia Giulia in Italy. vector of FD. Vidano et al. Egger and Grasselli: Presence in Toscana.: Study on the distribution of S. titanus. Conti: A review of intracellular procaryotic phytopathogenic agents. Fortusini et al.: Presence of a FD-like disease in Emilia-Romagna. 1987 1987 1987 1988 1988 1988 1988 1988 1988 1989 1989 139 . Survey for the presence of S. variegatus and S. Chardonnay is the more affected variety. their etiology is unclear. Pinot bianco. Borgo et al. vector of FD. of MLOs in phloem tissues of grapevines affected by "FD" in northern Italy. using the scanning electron microscope. S. Susceptibility and sensitivity of affected varieties (Chardonnay. Rui et al. Italy. Euscelidius variegatus and Euscelis incisus are taken into consideration.: Observations. Credi et al. titanus. resulting in a reduced diffusion of the disease. Boudon-Padieu and Larrue: ELISA detection of FD pathogen on infected reared experimental and wild natural leafhopper vectors (E.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. In addition to S. Italy. Magarey et al. Martelli: A review on the knowledge on grapevine diseases induced by phloem. Borgo: General information on the presence of FD-type diseases in northern Italy. respectively). Seljak: Presence of Scaphoideus titanus in western Slovenia (Yugoslavia). disease distribution. Vidano et al.: Occurrence of FD-like symptoms in the valley of Sarca in Trentino.: Contribution to the knowledge of FD (or similar diseases) in the Veneto region.
Several weeds with symptoms of phyllody were found to contain MLO in electron microscope studies. Affected grapevines in the Rhône valley tested negative. Le Grau du Roi (France) of a device for secure soaking of dormant buds and canes into hot water (50°C. Credi and Santucci: Development of "Flavescenza dorata" in grapevine in Emilia-Romagna and rate of infestation in different systems of cultivation. Caudwell et al. Caudwell: Review on the epidemiology and characterization of FD and other GY diseases. Marinesku et al. Granata and Grimaldi: MLOs are observed with the electron microscope on affected grapevines of cv. Boidron and Grenan: Construction and testing in ENTAV. Conti: A yellows-type disease of cv. Inzolia. suggesting an unrelated agent. The aetiology is not fully ascertained though the presence of MLO is probable. The annual rate of symptomatic plants was never greater than 20 % over a 6-year period. Inzolia in Sicily.: Presence of yellows-like symptoms in Apulian grapevines (central Italy).: 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. Restriction profiles after digestion with endonucleases permit the differenciation of MLOs associated to different plant diseases.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. titanus was not found in the area. Specific primers can be used for MLOs. Di Terlizzi et al. Chardonnay develops in Tuscany. Deng and Hiruki: A method to amplify 16S rRNA gene from culturable and nonculturable mollicutes. 45mn). Granata and Russo: An epidemic disease with FD-type symptoms develops in Sicily on cv.: Bench grafting experiments of buds of indicator cvs. This is a prospect for investigation on MLOs associated to plant yellows. Natural infection with MLOs of bait plants (Catharanthus roseus and Vicia faba) placed in vineyards in spring and summer. Chardonnay and Baco 22A grafted on donor yellows-diseased plants of cvs Pinot blanc and Sangiovese. S. 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". (a and b): Identification of numerous species of Auchenorrhynchas in the vineyard ecosystem in Piemonte (northen Italy) and of ampelophilous species. Credi et al. The recommended temperature / time combination is 50°C / 3560 min. 1989 1990 1990 1990 1990 1991 1991 1991 1991 1991 1991 1991 1992 1992 1992 140 . Treatment prior to storage is more efficient.: Grapevine graftwood shoots infected with FD can be disinfected by hot water treatment in the dormant stage. Refatti et al. titanus was present in all vineyards checked. Only part of the plants were infected. 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). Vidano et al. Symptoms also developed on plants obtained from buds taken on diseased Chardonnay and Sangiovese grafted on healthy Kober 5BB. Similar symptoms on Chardonnay could be obtained by insect inoculation (Euscelis incisus) and by cleft-grafting of tissues taken from yellows-infected elm. Caudwell and Kuszala: Detection with ELISA of FD-MLO in naturally FD-affected grapevines.: Occurrence of grapevine yellows in Moldavian SSR.
(a and b): Observations supporting the existence of different types of grapevine yellows in northern Italy. Caveccia and Chardonnay vines to periwinkle by the dodder species Cuscuta campestris. Alma et al. Attempts to characterize MLO DNA extracted from insects with molecular methods.: 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 . Bianco et al.: First detection of FD-MLO DNA in grapevine extracts by DNA-DNA hybridization with DNA probes cloned from FD-MLO partially purified from experimentally infected plants (Vicia faba). However. Evolution of the disease and of its vector in the various viticultural regions of France. Bertaccini et al. IPVR is related to aster yellows MLO.1992 Credi and Santucci: Of 628 attempts to transmit the agent of a yellows disease of grapevine from Sangiovese. titanus population in vineyards and the rate of spread of the disease.: ELISA with FD-antibodies permit to detect related antigens in S.: 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. MLOs were visible in sections of petioles of periwinkle showing symptoms and subsequent graft transmission from periwinkle to periwinkle was successful.6%) were obtained. No spontaneous recovery. titanus is not present.: Transmission experiments from grapevine to grapevine and to herbaceous hosts with S. Boubals (b): A severe epidemic of grapevine yellows occurs in Golan (Israel).: FD can be transmitted by symptomless rootstocks. Quacquarelli and Barba: Review of the distribution of flavescence dorée and other grapevine yellows in EEC viticultural countries. S. Hot water treatments suppress the transmission to Chardonnay indicators. 4 positive results (0. Meignoz et al. detection (ELISA. (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. titanus. results of research. It is more related to the agent of Elm yellows than to other yellows agents. The titre of MLO appears very low in grapevine. titanus and other leafhopper species show that yellows disease present in northeastern Italy differs from flavescence dorée as it occurs in France. (a): Differentiation with PCR-RFLP analysis between a MLO related to aster yellows in GY-diseased plants from Lombardia and a MLO related to elm yellows in an affected grapevine in Friuli-Venezia Giulia.: Description of MLOs and associated disorders in the phloem of FD-affected LN33 experimentally inoculated with S. but are different from known aster yellows cluster MLO strains. Davis et al. Arzone et al. FD-MLO DNA can be distinguished from other MLO DNA with Dot-blot hybridization and Southern-blotting. genomic tests). Arnò et al. Typical symptoms on several cvs. suggesting the presence of two different diseases at least in Veneto and Friuli-Venezia Giulia. Boubals (a): Report on a meeting of the French working group on FD. Caudwell et al. (b): Evidence of an association of MLOs with grapevine yellows in Emilia Romagna. One MLO strain transmitted with Macrosteles quadripunctulatus appears similar or close to aster yellows MLO. Daire et al. transmission by feeding to healthy grapevines succeeded only in 1 attempt out of 100. The MLO strains found in grapevine are related with aster yellows MLOs. Osler et al. titanus leafhoppers. Chen et al. FD can be readily distinguished from Bois noir. Senescent or degraded forms of MLOs identified inside degenerate sieve elements.
Maixner (a. MLO appear to be in very low titre. b. at least two different phytoplasmas are infecting grapevines in Veneto: FD sensu stricto (transmitted by S. riparia in New York and failure of detection in some symptomatic grapevines in New-York and Italy. using insects. 1993 Daire et al. Maixner: Demonstration that Hyalesthes obsoletus (Auchenorrhyncha: Cixiidae) is a vector of German grapevine yellows (Vergilbungskrankheit).: Important outbreak of a yellows disease on local varieties in Apulia (southeastern Italy) and infection of periwinkle as bait plants and with dodder transmission. International Committee of Systematic Bacteriology (ICSB). A MLO has been transmitted to periwinkle with dodder.: FD-ELISA method was used on tissues of yellows-affected grapevines from diverse countries. on the effects of pollarding affected vines and of chemical sprays against vectors. 1993 1993 1993 1993 1993 1993 1994 1994 1994 1994 1994 142 . titanus is not present. BN belongs to the stolbur group. Transmission has been obtained only from vines of Veneto. then GYU phytoplasma) and comparison of specificity and sensitivity of the latter tools to detect phytoplasmas in grapevines in New York and Italy. Classification of grapevine MLOs into three RFLP groups: elm yellows (FD from France). So. Prince et al. In northeastern Italy. Description of pathological changes in phloem of leaves and identification of senescent forms of MLO inside degenerate sieve elements. aster yellows and Xdisease. FD detected in leaves of rootstock 3309C and is also present in samples from Veneto (Italy). Credi (a and b): Observation of MLO in yellows-affected grapevines in northern Italy. Electron microscopic observation of MLO in periwinkle and grapevine. Positive assays obtained only on samples from France and Veneto. Kuszala et al.called FDU. FD belongs to the elm yellows group but is different from elm phytoplasmas. titanus to test plants of cvs Perera and Chardonnay in Veneto and Friuli-Venezia Giulia (Italy). FD is related to elm yellows and BN is related to stolbur. Osler et al. S. Computer analysis for spatial patterns of diffusion show a non-random distribution and suggest transmission from other vines or from weeds. graft and dodder. probably transmitted by another insect. Daire: PhD thesis on detection and differentiation with DNA-based methods of the MLO agents of GY diseases in France. Only FD and BN have been identified in naturally affected grapevines. (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. Detection in non symptomatic V. Specific probes cloned and specific primers designed for PCR amplification of phytoplasma DNA. healthy plants protected with plastic screens did not develop symptoms and diseased plants protected with screens showed transient recovery. 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). titanus) and a second one. c and d): Grapevine yellows (Vergilbungskrankheit) are common in the Moselle and Rhine valley. titanus is present in the Friuli-Venezia Giulia region but no transmission assay has been successful.: Six-year transmission trials of different types of yellows with S. S. Di Terlizzi et al. Girolami and Egger: Importance of contamination of vineyards and diffusion of Grapevine yellows in northern Italy. 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. BN (stolbur MLO) detected in several regions of Italy and in Israel.: Diversity of MLOs associated with grapevine yellows and transmitted to periwinkle. GYU was later on characterized by other authors as a phytoplasma belonging to the Xdisease (16SrIII) group. Detection in grapevines from Virginia (USA) of MLO related to X-disease. Carraro et al. (a and b): Various attempts to transmit FD-like disease in different regions of northern Italy. Experiments on the use of hot water treatment on planting material.
Padovan et al. Alma et al. titanus which has not been found. Detection is also possible during winter on wood scrapings.: MLOs associated to grapevine yellows in Virginia belong to 2 different groups : X-disease group (16SrIII) and aster yellows group (16SrI). Ten weeds were found harboring MLOs with transmission electron microscopy. Prince et al. PCR detection of phytoplasma with universal primers. (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. The importance of proper identification of disease type for control measures is emphasized.: The phytoplasma associated with Australian grapevine yellows is close to but different from stolbur phytoplasma causing BN. The most frequent belonged to group 16SrI-G (later designated as 16SrXII or stolbur group). 1995 1995 1995 1995 1995 1995 1995 1995 1995 1996 1996 1996 1996 1996 1996 1996 1996 143 .1994 1994 Parente et al. (a and b): Grapevine yellows in Latium (southwestern Italy) are not associated with S. Molecular detection of aster yellowsrelated phytoplasma. Italy). Koruza: Dispersal of grapevine yellows in Slovenia. stolbur and apple proliferation) can be detected.: Six-year survey of evolution of yellows symptoms in a 6-12 year-old vineyard of cv Chardonnay that was never sprayed with insecticides. Boudon-Padieu (a and b): Reviews on knowledge and research on grapevine yellows. in grapevines with yellows symptoms in Soave (Veneto. Del Serrone et al. Maixner et al.: Survey of yellows symptoms in an ampelographic collection of 1281 cultivars in Conegliano (Veneto.: Emphasis on the possible role of weeds as reservoir for MLO in vineyards. probably of Bois noir type. One grapevine out of 16 contained a 16SrV (= elm yellows) phytoplasma together with a 16SrI-G phytoplasma.: Presence and distribution of GY in Sicily. Related MLOs were also detected in wild grapevines. aster yellows. Arzone et al. Murari et al. Egger et al. with the aim of identifying sources of tolerance or resistance to yellows diseases. obsoletus and in weeds growing in the vineyard in Germany. Del Serrone and Barba (a and b): Monitoring of detection of phytoplasma in the organs of grapevine according to the vegetative state. Haidar: First report of symptoms of yellows on grapevines in Lebanon. Double infection is reported. sometimes in mixed infection.: Four different phytoplasmas (FD. Consideration on possible resistance and research on interaction of phytoplasma with nepovirus infection. (a): Specific detection with PCR of stolbur phytoplasma in grapevines affected with Vergilbungskrankheit. Italy). Albanese et al. Maixner et al (b): Detection of a new phytoplasma type related to FD in cv Scheurebe showing symptoms of yellows in Palatinate (Germany).: Grapevines from Piemonte (northern Italy) contained several different phytoplasmas. Padovan et al.: Presence of grapevine yellows in northern Portugal and electron microscopic visualization of MLOs in the phloem tissue of petioles of affected plants.: Identification of BN (stolbur phytoplasma) in Spain. in the vector H. Fortusini et al. Laviña et al.
Phytoplasma belonging to the stolbur subgroup have been identified. Kölber et al. Daire et al. Belli et al. titanus reported for the first time in this region.: Recurrent and severe yellows symptoms observed since the 1980's in Sardinia (Itlay) on several cultivars. spread. titanus is not present. Del Serrone : A review in Italian on the knowledge on etiology. Maixner et al. 10 are confirmed phytoplasma vectors. Positive reaction with a DNA probe specific to aster yellows phytoplasmas. Garau et al.: Identification of Flavescence dorée in Spain.: Survey of leahoppers in vineyards of Piemonte. Aldini et al. International Committee of Systematic Bacteriology (ICSB). Aster yellows and X-disease types were never detected.1996 1997 1997 1997 Rüdel: Review on the history and diversity of Vergilbungskrankheit of grapevine in Germany. The possibilities of control 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1998 1998 1998 144 .: Symptoms of grapevine yellows observed in Hungary in the early 1970's. Presence of S. No double infection has been found to occur. biology and control.: Use of Hot water treatment (HWT) to eliminate phytoplasmas (FD and AY) from 3 cultivars of grapevine. (a): Description of available methods and strategies for detection of phytoplasmas in grapevine. No positive detection with PCR on the HW-treated vegetative progeny of mature canes of symptomatic plants. (a): Only stolbur and FD phytoplasmas were detected in samples of yellows-affected grapevines from numerous regions of France and from Italy. Bosco et al. The term phytoplasma will become the genus name of plant pathogenic mycoplasmas under the provisional taxonomic status Candidatus. Subclades are considered to represent the equivalent of distinct species. (b): Prospects on detection and identification of grapevine phytoplasmas with new molecular tools and consequence on future epidemiology studies. Italy). Murari et al. Maixner and Reinert (a): Hot water treatment (HWT) of 50°C for 60 min applied to dormant mature canes eliminated Vergilbungskrankheit phytoplasma with a small loss in survival of the cuttings.: History and control of GY diseases in Italy. AY and X-disease related phytoplasmas have been transmitted and detected. transmission. S. Among 32 species identified. Davis et al. S. Batlle et al. 10 species were known vectors of phytoplasmas. (b): Survey of possible auxiliaries for the biological control of leafhoppers and planthoppers in vineyards. vectors and detection of the agents of Grapevine yellows. Maixner et al. No epidemic outbreak has been observed. Tanne and Orenstein (a and b): Successful heterografting transmission of phytoplasma from symptomatic grapevine to periwinkle and easy subsequent identification of phytoplasma type. Subcommittee on the Taxonomy of Mollicutes: The taxonomy of phytoplasmas will refer to their molecular phylogeny. Borgo: Description and colour photographs of symptoms of grapevine yellows and knowledge on their aetiology and vectors.: Survey of hoppers in vineyards of the province of Piacenza (Emilia Romagna. 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). Boudon-Padieu and Maixner: Updating in French and English of present knowledge on grapevine yellows: etiology. Spain and Israel. titanus is not reported. northern Italy.
distribution in the world. Frausin et al. with a particular reference to the work done in Germany. methods of detection of phytoplasmas in grapevine tissues and in insect vectors. Only FD and BN have been identified. Guadagnini et al. Reinert W. nucleic acid hybridization and serological comparisons. titanus. Probes were antisense to stolbur specific sequences in the 16S rDNA and used on stolbur-infected periwinkle. In addition a phytoplasma (FDU=GYU) transmitted from GYaffected vines to Catharanthus roseus by means of dodder in Friuli-Venezia Giulia. Similar attemp with FD phytoplasma were not successful. In the Veneto region only stolbur-group phytoplasma was recorded in some areas and both stolbur and elm yellows group phytoplasmas in other areas. FD-related phytoplasmas (elm yellows group) have been detected in grapevines from Treviso and Verona and transmitted from vine to vine using S. History. Maixner: A review of thermotherapy to cure phytoplasma infected material.: Monitoring grapevine yellows in North-eastern Italy and attempts to rationalize routine laboratory testing for the differentiation between FD and stolbur phytoplasmas. obsoletus is rarely found in vineyards.depend on the knowledge of the biology of phytopathogenic agents and of their vectors. FD is restricted to the north of Cataloña. has been identified as a member of the X-disease group. 1998 Lee et al. Search for alternative vectors. Grapevine phytoplasmas classify into different groups. BN is very frequent but H. Borgo et al. according to the most recent studies on molecular structure of rDNA. Phytoplasmas for which 16S rDNA sequence is available have been classified into 20 major groups and subgroups. Boudon-Padieu: Review on recent progress in the knowledge on grapevine yellows. epidemiology of these diseases.: A history of GYs in north-eastern Italy. though S. Carraro et al.: Unsatisfactory results on the use of hot water treatments to eliminate phytoplasmas from grapevine wood. Seemüller et al.: First report of the use of oligonucleotides for in situ hybridization with transmission electron microscopy to localize phytoplasma in plant cells. and M. Insecticide sprays against S. 1998 1998 1998 1998 1999 1999 1999 1999 2000 2000 2000 2000 2000 145 .: A comprehensive review of the classification of phytoplasmas in the world. symptoms. vector of FD. Maixner and Reinert: Updating in German of the research made in different countries on grapevine yellows diseases. are compulsory in France. Refatti et al.: Summary of the complex situation of GY in northeastern Italy. phytoplasmas can be classified into 14 groups and 38 subgroups. epidemiology and etiology of GYs in the world. have shown that only stolbur group phytoplasma was consistently found.: Evaluation on the efficiency and security of hot water treatment used to eradicate phytoplasmas from propagation material.: Phylogenetic studies of phytoplasmas associated to numerous plant diseases. GY affect mainly the northern provinces. titanus.: Report of positive detection of aster-yellows phytoplasma in grapevine fed with Metcalfa pruinosa and in the body of the insects used for transmission. On the basis of RFLP of PCR-amplified 16S rRNA gene. Boudon-Padieu (b): Chapter in a Handbook on viral and bacterial diseases of the grapevine. Survey with PCR of phytoplasmas infecting grapevines in Friuli-Venezia Giulia and Trento province (Italy) and in Slovenia. No negative effect was observed on scion buds but rootstock cuttings were more sensitive to negative effects of the treatment. titanus is more widely distributed. Lherminier et al. Batlle et al. Sensitivity of assays must be good because of the low titre of phytoplasmas in affected plants.: A review of the situation of GY in Spain. Firrao et al.
HWT (50°C/40 mn) gave acceptable survival but none of the conditions experimented provided a total elimination of phytoplasmas. A high rate of recovery is observed from one year to the next. Bertaccini et al. Phytoplasmas seemed to disappear from the insect body after the transfer of insects to healthy plants. suggesting that they did not multiply in the body of the insects. The latter was difficult to evaluate because of the non homogenous distribution of phytoplasmas in canes. Seljak and Petrovic: Overview on phytoplasma diseases of grapevine and fruit trees in Slovenia. Tanne et al. Waite et al. 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. FD has not been identified.: The presence of phytoplasmas in liquid medium on which insect vectors have been fed can account for potential vectorship of the species. Zahavi et al.: Use of hot water treatment in commercial nurseries in Australia to eradicate diverse pests and pathogen agents from propagation material. The phytoplasma agents are not specified. Neoaliturus sp. Primers fitted for specific detection of these 3 phytoplasmas permit cheaper and quicker diagnosis. Influence of rootstock on the rate of GY symptoms in the same vineyard and same grapevine variety. Osler and Refatti: The situation of GYs in northern Italy.: Scaphoideus titanus is present in Portugal.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. Crocker et al. Boudon-Padieu: Updating of knowledge and research on grapevine yellows worldwide. Macrosteles quadripunctulatus and Orosius orientalis were found on weeds in vineyards or trapped on sticky traps. Phytoplasmas were detected in all five species. Seljak: A review of non-European hoppers introduced in Slovenia. Quartau et al. Israel. Frosini et al.: Report of experiments combining temperature and time of treatment with hot water on 5 grapevine cultivars to cure planting material from phytoplasmas. nitrate and nitrite reductase. Orenstein et al.: A 2-year survey in Golan Heights.: Survey of GY in Israel. Moretti et al. (a): Presence of FD. Dual infection may occur (13 %).: Recommandations for the organisation of nursery in Australia and handling of hot water treatment to avoid dehydration of cuttings during treatment. Circulifer sp.: Development of a new technique with DNA chips to detect phytoplasmas in grapevine. which are all known as vector of phytoplasmas. Experimental evaluation of the method and application to field-trapped individuals of suspected vector species.: In Golan Heights (Israel) Hyalesthes obsoletus. Marzachi et al.: Metcalfa pruinosa specimen reared in the laboratory may acquire FD and clover phyllody (aster yellows group) phytoplasmas by feeding on infected Vicia faba but feeding transmission to several plant species has not been obtained. sugar metabolism. Plant variety and cutting diameter influence the rate of surviving cuttings. Hyalesthes obsoletus has been found but not in vineyards. titanus is widespread in western vineyards.. Klein et al. BN and aster yelllows in vineyards of southeastern Piemonte (Italy). confirmed the presence of three phytoplasmas associated to GY: stolbur (70%).: Use of immersion in hot water (HWT) or in chemicals to suppress phytoplasmas from grapevine propagation material in Italy. aster yellows (11 %) and X-disease groups phytoplasmas. 2000 2001 2001 2001 2001 2001 2001 2001 2001 2001 2001 2002 2002 2002 2002 2002 2003 146 . Clair et al. although its vector S.
Myrta et al. stolbur phytoplasmas are suspected to occur because of long-lasting record of stolbur disease on many host plants in Serbia. Study of the spatial and temporal dispersion of the four species.: Flight activity of Scaphoideus titanus and Hyalesthes obsoletus and their infection status towards FD and BN (stolbur) phytoplasma. Consequently roguing of affected vines should be avoided when possible and especially if vectors are controlled.: Development of a sensitive PCR procedure for dual identification of FD and BN phytoplasmas. Crocker et al. rapidity and sensitivity of methods for use in routine diagnosis of grapevine phytoplasmas. D'Ascenzo et al. (a and b): FD phytoplasma (16S rV-C) and S. Recovery is a progressive phenonenon developing over 3 years in the case of BN. No important damage has been reported. Kuzmanovic et al. Leitner: The situation of Grapevine yellows in Austria is now surveyed by means of molecular detection assays. Lessio et al.: Report of severe symptoms of yellows on numerous grapevine varieties in Serbia and observation of organisms resembling phytoplasmas in sieve tubes of affected shoots collected in the Zupa area. Discussion on benefits and limits of artificial feeding medium to assess the transmission capability of phytoplasma by insects.2003 2003 2003 2003 2003 2003 Boudon-Padieu et al. M'hirsi et al. fitted to routine detection. respectively. titanus found associated to grapevine yellows with a high incidence in vineyards of Rastina (South Serbia).: Identification of aster yellows (16SrI-B and C) and of ash yellows (16Sr-VII) related phytoplasmas associated to yellows of grapevine in Chile.: Report of aster yellows phytoplasmas in grapevine affected with yellows in Tunisia.: Important outbreak of grapevine yellows on cv Chardonnay and identification of a stolbur phytoplasma in diseased plants. Megophthalmus scabripennis positive for aster yellows phytoplasma. Hyalesthes obsoletus and Circulifer haematoceps abundant and positive for stolbur and aster yellows phytoplasmas.: Compared efficiency. An aster yellows phytoplasma is suspected. Orenstein et al.. Milkus et al. Chabbouh et al. BN (stolbur phytoplasma) has an incidence of about 30 %. two nonpreferentially ampelophagous vectors of grapevine phytoplasmas. Marzachi et al.: Identification of grapevine yellows symptoms in Tunisia and attempts to identify pathogen agent and vector.: Report of stolbur phytoplasmas in grapevine affected with yellows in Albania. 2003 2003 2003 2003 2003 2003 2003 2003 2003 2004 2004 2004 147 . Neoaliturus fenestratus.: Survey of potential vectors of phytoplasmas in vineyards of the Golan Heights. Clair et al. Apart from FD phytoplasma reported by Duduk et al. Tassart-Subirats et al.: Important presence of GY in Abruzzo (central Italy). Osler et al.: Data on 15-year studies on efficiency and effect on propagation material of hot water treatment.: Measurements of basal respiration rate of grapevine dormant wood as an indicator of the best period for application of hot water treatment. Duduk et al. Gajardo et al.: Grapevines affected with yellows are found free of phytoplasmas after recovery. in particular a clover phyllody type (16SrI-C) which is quite frequent. Other phytoplasmas are reported.: Improvement of detection of FD and BN in field-grown grapevines with real-time PCR. 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.
However. titanus. symptoms affect the whole plant. FLAVESCENCE DORÉE 1. Infected rootstocks show little or no symptoms. However. After the discovery of other GY diseases. with an incidence that increases rapidly from one year to the next. Scaphoideus titanus Ball (= S. Several isolates have been characterized with molecular criteria. decline progressively and die. isolated infected vines have been identified in northern vineyards of Burgundy and Alsace. 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. it occurs in all southern vine-growing regions. 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. FD88. S. the term "FD sensu stricto" was applied to diseases and phytoplasmas that are transmitted by S. Transmission: The FD agent is transmitted in the persistent mode by the leafhopper Scaphoideus titanus Ball (= S. If the plants are inoculated after recovery. First symptoms may 149 . Crop losses may be very high. Hence. These isolates could be distinguished by Western-blot analysis using FD antisera and monoclonal antibodies (Mabs).) using S. Switzerland. In addition. indicating a vine-to-vine transmission. FD92 and FD-D could not be distinguished from one another. Molecular comparisons have shown that all these isolates are more closely related among them than with other phytoplasma strains in the same group. Cicadellidae). Moreover. 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). Though the rate of transmission by bench grafting can be very low. extremely sensitive varieties do not recover. Two isolates denoted FD-D and FD-C. Corsica. Infected rootstocks are important means of dissemination because they are symptomless. littoralis Ball) (Homoptera. Affected vines of most varieties may recover in the second year when they are protected against reinoculation with insecticide treatments. introduced into Europe at the beginning of the 20th century. Transmission occurs also by vegetative propagation and grafting. Description Flavescence dorée (FD) was the first Grapevine yellows (GY) disease to be reported. FD88. were characterized in Italy and shown to be transmitted also by S. titanus. symptoms may be limited to a few shoots. Feeding transmission starts after a 4-week latency in the body of the vector. that has colonized a wide climatic area. show a general asymmetric bent posture and necrosis of terminal bud. It is widespread in all northern provinces of Italy and an outbreak was recently reported from southern Serbia. Eggs are laid in summer on two-year old wood and trunk. titanus is well established. Most of the time. Leaves persist longer on affected plants. then as a virus disease. littoralis Ball). FD is endemic only in regions where S. This leafhopper is a neartic species specialized on Vitis sp. southern Italy. It is also present in regions from which FD has not been reported in France. Isolates F70. Agents: FD was first regarded as a physiological disorder. 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. Diseased vines have a patchy distribution in the plot. and western Slovenia. It is highly epidemic and extremely dangerous because of the biology and ethology of its leafhopper vector. this way of spreading is significant when infected grafted plants are planted in vineyards hosting the vector. FD92 and FD2000 were experimentally transmitted to broadbean (Vicia faba L. In France. and Savoie. Lignification of the canes is usually incomplete. transmission is possible from early June (about one month after the beginning of hatching) until the death of adults. It was described in a limited area in north-eastern Cataluña (Spain). north of Spain and Portugal. Symptoms develop on leaves and shoots in June and July to become oustanding in August and autumn. titanus individuals collected in infected vineyards of south-western France. rooted cuttings from infected canes of a few rootstock varieties can develop "vinifera-like" symptoms on the wood and leaves. 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.GRAPEVINE YELLOWS: INDIVIDUAL DISEASES A. Vineyards in these areas are under the threat of an FD outbreak that could occur if FDinfected planting material is introduced.
Vitis riparia can be infected but shows little symptoms. have been found carrying FD phytoplasma until recently. titanus in an affected vineyard is an important indication that may lead to suspect a FD infection.appear on young vines as late as four years after grafting. Two additional treatments are applied during summer. Bois noir (BN) is also frequent in regions inhabited by this leafhopper. a DNA fragment specific to BN (stolbur) phytoplasma can be used for the survey and monitoring of these two GY diseases. must be destroyed. Under these circumstances only chemical insecticides are efficient for eradication of the disease. and the risk of disease spreading important. appear more tolerant. The second treatment. Alicante Bouschet. has been used as the official method for large scale survey of FD in France from 1993 to 2003. Other DNA-based methods. whereas the third treatment. In 2003 a phytoplasma resembling the FD-C isolate was found in wild Clematis in Veneto (Italy). has allowed the identification of potential auxiliaries for biological control. Recently. at the beginning of August. Polyclonal antisera and Mabs have been raised to FD phytoplasma. which is specific for 16SrV-group phytoplasmas allowed the reliable detection of FD phytoplasma in grapevine. Sequence or RFLP analysis of the FD9 DNA fragment permit to readily differentiate all the known FD isolates. its area of origin. Material from mother plants that have developed symptoms in the following growing season. Ugni blanc (Trebbiano) and Prosecco are sensitive varieties that may recover when they are protected from new inoculations. Antigen extraction from vascular tissue of symptomatic grapevines requires the use of high Tris molarity and strong detergents in the extraction buffer. Varietal susceptibility and sensitivity: All varieties grown in the various FD-affected regions in France. An indirect DAS-ELISA using coating of wells with rabbit polyclonal antisera and detection with a cocktail of several Mabs. Nielluccio and Garganega are very sensitive varieties that usually do not recover after infection. In spite of the possibility of recovery. roguing of diseased vines is advisable when the epidemic pressure is high. aims at killing newly hatched insects. 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. Sauvignon blanc. The presence of S. although heavily infected vines can be observed. is directed against winged adults migrating from nearby vineyards or wild vines. Nevertheless. titanus in New York State (USA). Grenache. control measures are compulsory according to legal regulations. Control: FD is a quarantine organism in the EC. Their rearing is still under development. The first treatment is applied 30 days after first instar emergence at the beginning of the period of potential transmission. such as real-time PCR or identification of PCR-amplified product with DNA-DNA hybridization using specific probes. Planting of HW treated material is especially important in areas where the vector is present. Soaking of dormant material in hot water (HW) is recommended. The low rate of transmission by grafting and the long delay in symptom expression make the detection of infected vines in planting material difficult. Detection: FD symptoms can be confused with those of other GYs and also with other diseases or disorders. 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. Detection with laboratory methods is possible on insect vectors and infected vines. 150 . Monitoring natural enemies of S. Standard conditions (50°C for 45 min) proved highly efficient for sanitizing infected rootstocks and grapevine cultivars. Indirect control is obtained by insecticide treatments against S. titanus. sensitive amplification with nested PCR of the DNA fragment FD9. Planting material must not be collected from mother vines in areas or vineyards affected by the disease. Cabernet Sauvignon. Natural insecticides can be used to preventively limit vector populations but their efficiency is limited in epidemic outbreaks. Italy and Spain are susceptible with various degrees of sensitivity. has become the preferred detection method because it is easy to perform and was improved in reliability and sensitivity. Chardonnay. Other host plants: No host plants other than Vitis sp. such as Merlot. at the beginning of July. Simultaneous amplification in a multiplex procedure of both FD9 and Stol11. Other varieties. are currently under development. Roguing is compulsory in France. The same is true for American rootstocks which can therefore be dangerous sources of infection and dissemination of the disease. Symptoms are rare in Syrah. In France and Italy. Molecular detection by PCR of selected phytoplasma DNA fragments.
Schvester et al. littoralis. Caudwell: Study of the phenomenon of recovery of FD-affected vines. (b): Biology of S. S. Caudwell and Poitou: Study of the interaction between fanleaf and FD. littoralis is present. 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 recorded from in Italy in 1963. Description and study of recovery phenomenon and of localised symptoms.2. Schvester et al. Recovered vines may be infected again but the symptoms are lighter. 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. littoralis. (a): Control of FD by insecticide treatments against the vector.: First report of S. littoralis. Description of macroscopic and microscopic symptoms. 1960 1960 1961 1961 1962 1962 1963 1964 1964 1966 1966 1967 1968 1969 1970 1971 1971 1972 151 . littoralis from Ticino (southern Switzerland). Historical review 1955 1956 1957 Levadoux: Description of an outbreak of "Flavescence dorée" on Baco 22A in southwestern France. Schvester et al. Vidano: S. Study of the role in the aetiology of FD of mycoplasmalike organisms (MLO) observed in insect vectors and in inoculated grapevine and V. S.: Successful transmission of FD to herbaceous hosts by increasing the feeding access period of the vector S. The disease can be transmitted by grafting and has probably a viral aetiology. Caudwell et al.: Studies on the survival of S. Description of the insect. littoralis Ball. First report that abandoned or wild vines may be a source of infection. littoralis in its area of origin in North America. Caudwell et al. an insect recently introduced from North America. Schvester et al. biology and feeding damage and of the symptoms of FD in France on Baco 22A. Caudwell et al. Baggiolini et al. (a): Possibility to limit the populations of S. littoralis. faba plants.: Field tests for insecticide control of S. Vidano: Study of the ecology and biology of S. Bonfils and Schvester: Relationships of leafhoppers with FD of grapevine in western France. Use of 4-5 treatments with DDT or parathion and removal or destruction of pruning wood older that two years. (a). Caudwell et al. Boubals and Caudwell: FD epidemics observed in Corsica.: Demonstration that FD is transmitted by the leafhopper S. Caudwell: PhD thesis on FD. Branas (a and b): The new disease is thought to be caused by root damage. considered as a virus disease. littoralis. Spontaneous recovery occurs after a 1-2 year crisis period. The aetiology is unknown. Carle and Moutous: Study on possible secondary or alternative vectors of FD or Bois noir (BN). littoralis in Corsican vineyards with winter treatment of dormant eggs with oleoparathion. No positive results of transmission trials with 15 species of Hemiptera (mainly Cicadellidae) other than S. evolution of the disease in space and time. Caudwell: Attempts to inhibit the agent of FD in vivo by heat treatment. littoralis on plants other than the grapevine and transmission trials of FD to other host plants.
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
Spain and Italy. The biological significance of this finding is unknown. Infection is rapidly spreading to the east.: Monitoring of leafhoppers in vineyards in Italy. Several different peptides from membrane proteins are identified by Western-blot. Increase and spread to other cultivars.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. FD is not present in southern Italy. Clerc et al.: Development of specific PCR primers for the detection of FD or BN phytoplasmas.: First report of S. Kuszala: Specific ELISA procedures to detect FD or BN phytoplasmas in extracts from diseased grapevines from the field. First outbreak in the 1980's on cv Perera. Transmission by grafting is studied.: RFLP studies of 16SrV phytoplasmas. titanus in biological viticulture. Batlle et al. (a and b): Demonstration of curing effect of hot water treatment on FD-affected plant material. Jermini and Baillod: Combination of visual observation of instar larvae and trapping of adults on sticky traps to monitor the populations of S. Posenato et al. Caudwell et al. Vindimian et al. titanus in Italy. FD has not been identified. Belli et al.: Demonstration that two isolates of FD sensu stricto (FD-D and FD-C) are spreading in Veneto (Italy).: Important outbreak of FD in Lombardia (northern Italy).: Verification of hot water treatment conditions to suppress FD phytoplasma infection in dormant wood of grapevine. the eggs of S. Control recommendations. (a and b): Situation of FD and S. Bosco et al. The possibility that direct inoculations have occurred in nursery is discussed. titanus in Trentino (Italy). especially Prosecco in the period 1993-1996. 1996 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1999 1999 2000 2000 155 . Mori et al. titanus in Switzerland. In the same conditions. Italy). Daire et al. titanus laid in the bark were killed. Martini et al. according to the severity of the FD epidemics. titanus in north-eastern Italy where vines are trained in the pergola system. titanus in the canton of Geneva (Switzerland). Rousseau: A review of different ways to reduce the populations of S. Seddas et al.: First report of FD outbreak in northern Cataluña (Spain). The possible role of six species in the transmission of GY is discussed. nymphs and adults of S. Serological relationships with elm yellows phytoplasma is confirmed Alma et al. Bianco et al. Survey of hemipters in the vineyard and presence of other known phytoplasma vector species.: In spite of the presence and rapid spread of S. (a and b): Evolution of FD in the Treviso province (Veneto. Three different RFLP patterns at least are shown in FD isolates from France.: A phytoplasma of the 16SrI (aster yellows) group is detected in eggs.: Insecticide control of S. Marcone et al. The amplified FD9 fragment is specific and variable among 16SrV (elm yellows) group phytoplasmas. Pavan et al. titanus reared on healthy plants.: Monoclonal antibodies raised to immunoaffinity purified FD phytoplasmas. Lherminier and Boudon-Padieu: Use of transmission electron microscopy with in situ immunolabeling to detect FD phytoplasma in grapevine and herbaceous host plants. Numerous species identified.
Boudon-Padieu (a): Description, biology, spread, FD-transmission and control of S. titanus. Clair et al.: Design of internal nucleotides on the specific DNA fragment FD9 to be used as primers in nested-PCR for improving sensitivity of detection of elm yellows (16SrV) phytoplasmas in FD-affected grapevine or in elms affected with yellows in France. Cravedi and Aldini: Biology of S. titanus in Oltrepò pavese (Italy), a region affected by a severe outbreak of FD since 1999. Morone et al.: Occurrence of FD in Piemonte (northern Italy) since 1998. Roure: The situation of FD in France. Compulsory control (insecticide sprays and roguing of affected vines) on 300 000 ha. Scattini et al.: Important spread of FD in Lombardia (Italy) in cv Sangiovese. Angelini et al.: Phylogenetic comparison of FD isolates from France and Italy among them and other 16SrV (elm yellows) strains. One isolate from France (FD92 = FD88) and one from Italy (FD-D) are widespread and identical. Bianco et al.: S. titanus in northern Italy can transmit different grapevine isolates of 16SrV (elm yellows) phytoplasmas. Credi et al.: Detection of FD in grapevine samples from the provinces of Piacenza, Parma, Reggio Emilia and Modena in Emilia Romagna (Italy). Davis and Dally: A new 16S rDNA sequence of FD is produced and it is proposed that FD phytoplasmas are placed in two distinct subgroups. However, further work has shown that the 1994 original sequence was erroneous (see Angelini et al., 2003a). Harrison et al.: An elm yellows (16SrV) group phytoplasma close to the Italian isolate FD-C is detected in Virginia creeper plants (Parthenocissus quinquefolia) in southern Florida (USA). It is the first time that a phytoplasma resembling a FD phytoplasma is found in a plant species other than Vitis sp. Marzachi et al. (b): Development of a PCR strategy for large scale detection of FD phytoplasma in plants and insects. Pasquini et al.: Harmonization of procedures for diagnosis of FD in Italy. Quartau et al.: Presence of S. titanus in Portugal. Alma: A review of the distribution and biology of S. titanus in Italy. Bianco et al.: Development of a specific TaqMan® assay for detection of FD phytoplasmas. Borgo and Angelini: Spread of FD in Italy and the situation of mother vines and cultural practice. Boudon-Padieu: Present knowledge on the epidemiology, aetiology and diagnosis of FD. Credi et al. (b): Presence of FD in the Marche region (central Italy). Martini et al.: Further comparisons between FD phytoplasma isolates from France and Italy. Mori et al.: Positive transmission of the two Italian FD isolates (FD-D and FD-C) by S. titanus. Posenato et al.: Efficiency of various insecticides to destroy nymphs of S. titanus and Metcalfa pruinosa. Viggiani: Presence of S. titanus in the Basilicata region (southern Italy).
2000 2000 2000 2000 2001
2001 2001 2001
2001 2001 2001 2002 2002 2002 2002 2002 2002 2002 2002 2002
Angelini et al.: Use of Heteroduplex Mobility Assay and full sequencing on two genomic DNA fragments (ribosomal and non ribosomal) to further study the relationships between FD isolates and related 16SrV (elm yellows) group phytoplasmas. Bianco et al. (b): Presence of FD-D and FD-C isolates of FD phytoplasma in Lombardia (northern Italy). Botti and Bertaccini: Molecular variability between FD phytoplasma isolates using the 16S rRNA gene, the ribosomal protein operon and the FD9 DNA fragment. Distinction between epidemic and non epidemic isolates. Bressan et al.: An attempt to identify a pattern of seasonal transmission of FD, taking into account the transmission efficiency of S. titanus, the pattern of symptom expression by infected grapevines, the emergence of nymphs of S. titanus and the importance of vector population and diseased grapevines. Cavallini et al.: Presence of FD together with BN in the area of Modena (north central Italy). Clair et al.: A multiplex nested-PCR procedure for easy simultaneous diagnosis of FD and BN. The method is now the official method registered in the Official Journal of the French Republic to be used by laboratories in the frame of compulsory control of FD in vineyards. Constable and Boudon-Padieu: Isolation of FD phytoplasma chromosome in Pulse Field Gel Electrophoresis and first physical map for two FD isolates. De Sousa et al.: Identification of FD-D phytoplasma in S. titanus specimen trapped in Portugal. No description of FD-affected grapevines. Duduk et al. (a and b). Identification of FD in southern Serbia and presence of S. titanus. The phytoplasma isolate is similar to FD-C. Malausa et al.: Search for natural enemies of S. titanus in New York with the scope of their introduction in France as biological control auxiliaries. Nusillard et al. Results of a 2-year study in North America of the natural enemies of S. titanus. Santinelli et al.: Presence of S. titanus in Umbria (central Italy). Torres et al. The situation of FD in Spain. Confirmation that only FD-D is present. Angelini et al.: A phytoplasma identical to FD-C isolate is found in wild Clematis in the vicinity of a FD-C affected vineyard in Veneto (Italy). Lessio and Alma: Study of the distribution of S. titanus within a vineyard and use of chromatic sticky traps. Abundance is greater in normal- than in low-density planted vineyards. S. titanus is monophagous. Leafhoppers are not able to spread significantly outside a vineyard. Females are less likely to fly far away than males.
2003 2003 2003 2003 2003 2003 2003 2004 2004
B. BOIS NOIR (VERGILBUNGSKRANKHEIT, LEGNO NERO) 1. Description Bois noir (BN) and related diseases are Grapevine yellows (GY) known in Europe and Asia Minor. The first observations in Eastern France date back to the early 1940's. BN was formally described in 1961 in comparison with Flavescence dorée (FD), because symptoms were similar. It was assumed that the two diseases were different because BN occurred in regions where the vector of FD was not present and experimental transmission with S. titanus was not successful. Because the vector insect, Hyalesthes obsoletus, does not live on vines, affected plants are usually not grouped into patches as with FD, except for situations where a high percentage of affected plants is observed. Nevertheless, distribution is non-random and spread follows a main direction or is preferentially along the border of the plot. BN has been identified in all wine-growing regions of western and eastern Europe, in Israel, Lebanon and recently in the Ukraine.
Main synonyms: The syndrome named Vergilbungskrankheit (VK) (1971) or Schwarzholzkrankheit (SHK) (2003) in Germany, Legno nero (LN) in Italy, Bois noir in Spain, has been occasionally described under the name of Flavescence dorée-like disease. Moreover, it has been sometimes confused with FD in the 1970's and 1980's. It is only since the 1990's that the aetiology was established and that it was shown that the agents of these diseases in the different countries were similar or closely related. Furthermore, the same vector insect species, H. obsoletus, has been identified in all countries, except for Spain where it appears to be very rare. Main symptoms: All the main symptoms typical of GY can be found in BN / VK / LN - infected grapevines. In addition to leaf discoloration, growth abnormalities can be observed on canes and roots. Usually, only a few canes per plant show typical symptoms with either an uneven or a total lack of lignification. On whiteberried varieties, yellow banding develops along the veins which may undergo necrosis. On red-berried varieties, sectorial reddening of blades develops in summer and progressively invades the whole leaf. When berries develop they remain immature, greenish and eventually fall down. Remaining berries are tasteless and sour. Internodes often show longitudinal rows of brown pustules along the green bark of unripe wood. Severely affected vines decline and may die after a few years. Some vines may recover or express irregular symptoms in successive years. Agents: The phytoplasmas associated to these diseases belong to the stolbur group (16SrXII), a group that was for some time considered by some to be the I-G subgroup of the larger Aster yellows group (16SrI). Although stolbur phytoplasmas are ubiquitous pathogens, known to infect solanaceous crops as well as many weeds and bushes, only a limited number of strains have been identified. Hence the different names of BN disease appear to be more geographically than genetically significant. Transmission: Vergilbungskrankheit (= Schwarzholzkrankheit) was the first disease for which the vector was identified. The known vector insect of stolbur of tomato, H. obsoletus (Homoptera, Cixiidae) can transmit stolbur phytoplasma to grapevine, but this occurs during feed probing as the insect does not keep feeding on grapevine. Hence, vine-to-vine transmission does not occur. The role of H. obsoletus has been confirmed in France, in Italy and Israel. In Spain, where the species is rare, its role in BN epidemiology must be verified. Alternative vectors have been suspected but not demonstrated. H. obsoletus is a univoltine polyphagous species that overwinters as larval instars that live on the roots of host plants. Nymph emerge in spring and transmission occurs during adult mating flights. Overall, the presence of insects on vines may last a couple of months. Eggs are laid during summer on the ground at the base of the stem of host plants. Main host plants of the insect vector are common weeds such as Convolvulus arvensis, Ranunculus sp., Urtica dioica, Cardaria draba and Calystegia sepium. However, only C. arvensis and U. dioica were found consistently infected with stolbur phytoplasma, distinct isolates of which were identified in these hosts. From 10 to 80% of trapped populations of the vector may test positive for the presence of stolbur phytoplasma. Transmission by grafting is possible but only at a low rate. Since acquisition by the vector from infected vines does not seem possible, the significance of graft transmission on the spread of the disease is probably very low. Varietal susceptibility and sensitivity: In the different affected regions and countries, numerous cultivars of V. vinifera were found susceptible to the stolbur phytoplasma, cv Chardonnay being the most sensitive. The increasing incidence of BN has been related to the increase of surfaces planted with Chardonnay in the last decade in all countries. Infection of rootstock varieties has not been reported. In all varieties there is a proportion of affected grapevines that undergo transitory recovery or remission, while other plants in the same plot are tolerant or immune to infection. Other host plants: Stolbur phytoplasma is widespread, and has a wide host range. In addition to C. arvensis, U. dioica, Ranunculus sp., C. draba and C. sepium, shrubs and trees of Prunus spp., solanaceaous crops (tomato, tobacco, pepper, egg plant) and lavender (Lavandula spp.) may be significant reservoirs. Detection: The use of monoclonal antibodies raised to stolbur phytoplasma did not show enough sensitivity for routine ELISA detection in field-grown infected grapevines. Detection is achieved mainly with PCR assays or PCR-RFLP analyses of DNA from infected host plants and insect vectors. Most often, nested PCR is necessary for sensitive detection from infected grapevines. Universal primers were first used to detect both BN and FD in areas where the two diseases may coexist. Subsequent RFLP analyses of amplified phytoplasma rDNA permit the characterization of pathogens. Stolbur specific primers have also been designed, either on variable regions of the rRNA gene or on selected specific non-ribosomal
1972 1977 1978 1978 1988 1989 1992 159 . Cazelles et al. Historical review 1961 1965 1968 1969 1971 1971 Caudwell: Study on BN and its relationships to FD. for the presence of a FD-like disease. Recovery and remission of symptoms for several consecutive years are frequent in vines affected by BN. (b): BN is distinct from FD based on non transmissibility by S. obsoletus are not efficient. 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. The results suggest that the disease is brought into the vineyards from outside local sources. Rumbos et al. (b): Epidemiological observations of BN lead to the assumption that an aerial vector is involved. Leclant and Lacote: Study of the ability to transmit stolbur to herbaceous plants by H.: 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. Chemical sprays against H. Natural enemies are being investigated. Gärtel: Description of VK under the name of Flavescence dorée.: Rickettsia-like organisms from VK-affected grapevines are cultured in chick embryos. Leclant: Presence of Hyalesthes obsoletus a cixiid planthopper in the south of France.: Rickettsia-like organisms found in roots of VK-affected grapevines in the Saar. suppression of affected branches may improve harvest quality and help in progressive recovery. obsoletus in the south of France. Credi and Callegari: Survey of vineyards in Emilia-Romagna. and different susceptibility and sensitivity of grapevine cultivars. except for declining vines. 2. while others seem more tolerant and do not show symptoms. Though pruning does not cure infected vines. Some grapevines show symptoms repeatedly in successive years. titanus is not always associated with the disease. roguing should be avoided. Hence. For direct differentiation between FD and BN/VK phytoplasmas. the multiplex nested-PCR assay cited in the FD chapter can be used. titanus. Rumbos et al. The disease occurs in the Moselle valley and the Rhine valley. S. Nienhaus et al. cytological and electron microscope study of tissues of affected grapevine. Findings never confirmed. Caudwell et al. Superficial soil cultivation in summer or winter ploughing are methods intended to damage the larval instars and reduce vector populations. history. Control: As with other GY diseases. littoralis. unknown at that time. Mosel and Rhine valleys are considered as the causal agents of the disease and it is hypothesized that the nematode Xiphinema index is the vector of the disease. Description of symptoms of VK. no direct control is possible. Caudwell et al. The species has been described as the vector of stolbur "virus" by Suchov in USSR in 1948. Italy.DNA fragments.: GY symptoms are present in Switzerland (Suisse romande and Ticino) but they are not associated with S. These findings have never been confirmed. because of the complex biological cycle and multiple host plants of the insect species. Borgo: General information on the presence of FD-type diseases in northern Italy. Description of structures that were probably closteroviruses. Mendgen: 1971. BN is considered as a non epidemic form of FD.
(b): Presence of phytoplasmas in the group 16SrI. Borgo: Presence of BN in mixed infections with FD in the province of Treviso (Veneto. H. BN MLO is detected in grapevines from France. Bianco et al.: One of the two phytoplasmas detected in yellows-diseased grapevines in Liguria is related to IPVR phytoplasma. Italy) where FD is prevalent. subgroup I-G (later renamed as 16SrXII or stolbur group). c and d) Transmission of a yellows to periwinkle with dodder in Germany. (b): Evidence of an association of MLOs with grapevine yellows in EmiliaRomagna. Maixner 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. Maixner et al. Davis et al.: Grapevines from Piemonte (northern Italy) contain several different phytoplasmas. Biology of the insect. Bianco et al. Computer analysis for spatial patterns of spread shows a non-random distribution and suggests preferential transmission from other vines or from weeds. H. Reservoirs and possible role of other vectors remain to be elucidated. Maixner (b. obsoletus is a vector of VK in Germany. from several regions of Italy. Laviña et al. Several weeds are host plants of the stolbur phytoplasma. 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. Bertaccini et al. obsoletus. Daire et al. Maixner : Demonstration that H. (a): Development of stolbur-specific primers on phytoplasma rDNA and use for detection of stolbur phytoplasma in grapevines affected with VK and in the vector H. (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. (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). but are different from known strains of MLOs of the aster yellows cluster. The latter MLO was shown later to belong to the stolbur group.: PCR-RFLP analysis of MLO-DNA detected in VK-affected grapevines in Germany show that this MLO is related to stolbur. aetiology and transmission of BN in France. 1993 1993 1993 1993 1993 1993 1993 1994 1994 1994 1995 1995 1995 1996 1996 1996 1996 160 . the most frequent belonging to group 16SrI. Minucci et al.: The agent of a GY disease in the Italian Riviera is detected with a probe specific for Eastern aster yellows. obsoletus is confirmed as a vector of stolbur disease plants in France. MLO associated to GY in Italy are related to aster yellows MLO. It is suggested that vectors with preference for other host plants act as vectors for VK.: First detection of BN (stolbur phytoplasma) in Spain. subgroup I-G (later named group 16SrXII (stolbur) in the provinces of Vicenza. Davis and Prince: According to a different terminology. Bertaccini et al. obsoletus identified as the vector. Boudon-Padieu (b): History. PCR-RFLP analysis confirms relationship to aster yellows phytoplasmas.: A monoclonal antibody raised to stolbur mycoplasma-like organism is used for ELISA detection of the agent in herbaceous host plants and insects. Brescia and Pavia (northern Italy). The MLO strains found in grapevine are related with aster yellows MLOs.1992 Fos et al. clustered by the authors into aster yellows group but shown later to belong to the stolbur group. Alma et al. Italy. and from Israel.
a strong annual increase.: Prevalence of BN (stolbur phytoplasma) in Navarra and Cataluña (north eastern Spain). Weber et al.1996 1996 Koruza: Survey of GY in three regions of Slovenia shows that the disease is of BN type.: Phytoplasmas associated to GY in Campania (southern Italy) are genetically uniform and belong to the stolbur group. obsoletus as a vector of BN. Osler et al. (a and b): BN is poorly transmitted by bench-grafting. obsoletus and possible control measures. Ploughing in winter of soils infested by C. Laviña et al. (c): Identification of stolbur phytoplasma (16SrXII) as agents of GYs in Greece and Israel. biology of H. vector transmission and possible control measures. Albanese et al. search for vectors. Marcone et al. a high proportion of vines in which symptoms re-occurred after one season or more. titanus. Results were satisfactory. Maixner and Reinert: A review of the research and present knowledge on VK and other GY diseases.: Important spread of GY symptoms on several grapevine varieties in Umbria (central Italy). using hot water treatment with conditions recommended in France. Kölber et al. Davis et al.: Occurrence of GY diseases in Hungary and identification of stolbur phytoplasma in affected grapevines. Maixner: The situation of VK in Germany: symptoms. a high proportion of previously infected vines that did not show symptoms for 2 years at least and. 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. Sforza: PhD thesis on the epidemiology of BN in France. Estimated crop damage ranges from 20 to 40 %. Refatti et al. Search for variability among BN phytoplasmas with RFLP analysis of the specific non ribosomal DNA was unsuccessful. Characterization of a phytoplasma belonging to group 16SrI. 1996 1996 1996 1997 1997 1997 1997 1997 1997 1997 1997 1997 1997 1998 1998 1998 1998 161 . Weber: PhD thesis on the biology of the cixiid H. Preferential spread along the rows. obsoletus in Germany. The maximum delay of symptom expression in young plants is 2 years. obsoletus and its role as the vector of VK. Daire et al. as well as phytoplasmas transmitted by Credi to periwinkle in Emila Romagna and previously classified in the aster yellows group. arvensis is recommended to expose instar larvae and kill them with frost. Kuszala: Detection in ELISA using a monoclonal antibody to stolbur phytoplasma of BNassociated antigens in diseased grapevines in France. (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. Sforza and Boudon-Padieu: Description of H.: Monitoring of field populations of H. subgroup I-G (identified later as belonging to the stolbur group or 16SrXII). First detection of BN agent in diseased grapevines in Switzerland with the same procedure. conversely.: Consistent association of stolbur phytoplasma with GY diseases in the Italian provinces of Fiuli-Venezia Giulia and Trento and in Slovenia. Maixner and Reinert (a): Trials on elimination of VK infection from vine wood in Germany. Vindimian: LN but not FD is present in Trentino (northern Italy) in spite of the presence and active spread of S.
obsoletus. Maixner et al. the ability to recover of cvs Riesling and Pinot in spite of systemic symptoms. Confirmation of the role of H. and a high proportion of vines that escaped infection during the 3-5 year period of monitoring.: Ethological onservations and morphological descriptions of adults and larvae of H.: Only stolbur phytoplasma detected in grapevines showing GY symptoms. Up to 34% of insects were infected with stolbur phytoplasma. Weber and Maixner (b): Etholology of H. the second GY disease in Germany. Darimont and Maixner: Comparison of the transmission efficiency of phytoplasma to their host plants and to grapevine by H.: Several vineyards affected by BN in Burgundy (France) monitored for over 15 years and assessed for symptom severity. Sforza et al. the vector of Palatinate grapevine yellows (PGY). The same pathogen detected in weeds and other crops. Braccini and Pavan: Report of survey of Auchenorrynchas in vineyards of Toscana (central Italy). Seljak and Petrovic: A review of the Slovenian situation of GY diseases. formerly reported in western Europe. among which hoary cress. obsoletus and Oncopsis alni.: VK disease dramatically increased in incidence in the Moselle and Rhine valleys during the 1990's. obsoletus and Metcalfa pruinosa were found positive for BN phytoplasma (16SrXII-A). H. different phytoplasmas were identified in a number of cultivars. Marcone et al.: Epidemiology of BN in France. First launching of colonies of the insect under controlled conditions. Acquisition may occur at larval stages since emerging 5th instars were infective. obsoletus in vineyards is efficient with sticky traps placed at the soil level.1998 1998 Sforza and Bourgoin: Anatomy of reproductory organs and mating of H. Identification of two infected symptomless weeds in the vineyard. Braccini et al. obsoletus. Sforza et al. No other phytoplasma found in the South of the country. No other grapevine phytoplasma was detected.: In a large survey of vineyards in 8 Hungarian counties conducted in 1998. Evidence secured for the occurrence of tolerance (no symptom at all for 15 years) and temporary remission (no symptoms for a few years) . Up to 30 % of the insects were infected and the highest rate was in vineyards with high populations of Convolvulus arvensis. regardless of the cultivar in Campania (southern Italy). Identification of main reservoir weeds. obsoletus related to VK infection.: Use of PCR to confirm the presence of stolbur phytoplasma in GY-affected grapevines in Switzerland.: Widespread presence of LN in Toscana (central Italy). The rate of infected H. obsoletus and possibility to control the insect in vineyards. Transmission to natural host plants is higher than for grapevine with both insects. Females are more often infected than males. Weber and Maixner (a): Procedures for the survey of infection status of populations of H. The study confirms a high rate of infected H. 1998 1998 1998 1999 1999 2000 2000 2000 2000 2000 2000 2000 2000 2000 2000 162 . Bourquin et al. Phytoplasmas in the 16SrXII group (stolbur) were the most frequent. Maixner and Reinert: Collection of H.: First detection of stolbur phytoplasma in grapevines in Croatia. Larrue et al. Škoric et al. obsoletus.: BN reported only in north-western and eastern vineyards of Croatia. Varga et al. obsoletus as a vector to grapevine. obsoletus can be high because of the presence of reservoir weeds in the vineyard. High rate of BN / LN infection in the different vine-growing regions. Šeruga et al. PCR detection is reliable when testing together 25 insects among which only one is infected.
Choueiri et al. 2001 2001 2001 2002 2002 2002 2003 2003 2003 2003 2003 2003 2003 2003 2003 2003 163 . Darimont and Maixner: Importance of the presence and infectivity of H. the incidence of VK was significantly higher in conventional vineyards. Study of the spatial and temporal dispersion of the four species. Orenstein et al. Role of stinging nettle (U. Merlot and Pinot noir. Morandell: Detection of VK in south Austria. Mescalchin and Mattedi: A description of the development of VK in Trentino (northern Italy).: Comparison of infection risk by VK in organic and conventional vineyards in Germany. Detection in December was the more constant.: Anoder cixiid planthopper. Gatineau et al. reached by other authors. 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. Indicators based on climatic parameters permit to predict the flight period. Langer et al. Laviña et al. Same conclusions about tolerance and transient recovery of grapevines. very similar to European isolates. which is widespread in the province.: Demonstration that H. BN is the main disease and cvs Chardonnay and Zweigelt appear to be the more sensitive among whiteand red-berried varieties.: Importance of outbreaks of BN in the province of Modena (Italy) and monitoring of potential insect vectors in addition to H. dioica) as a reservoir of stolbur phytoplasma and as a host for the insect. In spite of a similar abundance of H.000 plants over 12 years. is a natural vector of stolbur phytoplasma to sugarbeet in Bourgogne (France). H. The presence of H.: Survey of GY from 1997 to 2002 on 21 cultivars in 12 Hungarian counties. obsoletus.: Only BN identified in Switzerland on cvs Chardonnay. Severity of symptoms vary from one year to the other.: First detection in Albania of BN phytoplasma in grapevines of different age and varieties. obsoletus and Circulifer haematoceps were widespread and positive for stolbur or aster yellows phytoplasmas.: Assessment of the best period for detection of BN in affected grapevines. respectively. Curkovic Perica et al.2001 Bertamini and Nedunchezhian: Evaluation of effects of BN disease (stolbur phytoplasma) in field-grown grapevines.: Survey of potential vectors of phytoplasmas in vineyards of the Golan Heights. Bertaccini et al. All affected grapevines and two naturally infected periwinkles contained the same stolbur isolate. Kölber et al. obsoletus is the vector of LN in Italy. Gugerli et al. Pentastiridius sp.: Observation of the occurrence of BN in Trentino in eight Chardonnay vineyards on more than 11. obsoletus to the German viticulture. Maixner et al. Samples were taken on all organs of 10 affected grapevines. Myrta et al. obsoletus and the presence in organic vineyards of several weeds known as host plants carrying larvae on the roots. on pigments.: The situation of GY diseases in Croatia shows that only BN is present in the northern and eastern vineyards. obsoletus. every month for one year except in winter. Cicotti et al.: Incidence of BN in a small area of the Bekaa valley (Lebanon) on cvs Chardonnay and Alicante Bouschet. The authors conclude that phytoplasma infection induces non specific. while Megophthalmus scabripennis was positive for aster yellows phytoplasma. Alma et al. general stress responses and rapid senescence. Neoaliturus fenestratus.: The highest risk of phytoplasma contamination for grapevines coincides with the flight activity of adults of H. obsoletus in Lebanon is recorded. chlorophyll fluorescence and photosynthetic activities suggest that Photo System II is affected.
PGY phytoplasma is similar to the Alder yellows phytoplasma that is widespread in black alder in Germany. Šeruga et al. was found carrying the alder phytoplasma at a higher rate than O. Control: No possibility of control because of erratic transmission by the vector. 2003 2003 2004 2004 2004 C. obsoletus and frequence of C. using 4 fragments of phytoplasma DNA showed that all isolates are similar.: Important expression of BN in cv Chardonnay in the Ukraine. According to the data of field survey. Description Palatinate grapevine yellows (PGY) is a disease occurring in Germany in Pfalz and Moselle regions (west of Germany) mainly on cv Scheurebe. Germany). The disease was identified in 1995. Gilge et al. It is associated to phytoplasmas in the Elm yellows (16SrV) group. alni but failed to transmit both to alder and grapevine. Alder is considered as the source of PGY infection. Cicadellidae) trapped on alders and caged on V. Main symptoms: Symptoms are similar to those of other GY diseases and may affect the whole plant. Langer and Maixner: Three different types of stolbur phytoplasma isolates found in grapevines.) Detection: Universal primers and RFLP analysis of amplicons can be used to identify a 16SrV phytoplasma. 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). and rarely in groups. mainly on cv Scheurebe. -B and C) have been identified. obsoletus is rare. 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. the psyllid Psylla alni. Another alder insect. However. arvensis and U. titanus leafhoppers have failed. Varietal susceptibility and sensitivity: PGY occurs in limited areas. affected plants are not numerous. Three isolates (PGY-A. Other host plants: Alder trees (Alnus glutinosa L. H. Šeruga et al. specific association of each isolate with a different weed is discussed. obsoletus were positive for stolbur phytoplasma. (a): Investigation on the variability of stolbur (16SrXII) phytoplasmas from extracts of different grapevines in Croatia. Transmission to plants is not reported. dioica. obsoletus and weeds in different areas in Germany. occur most often at the border of plots. 164 . Milkus et al.: Importance of a GY epidemics in the Drava region (Slovenia) in 2001 and 2002. Attempts to transmit PGY phytoplasma from grapevine to grapevine using reared S. Symptoms were recorded on 26% of 30-year old Scheurebe grapevines and 96% of tested individuals of H. It is different from FD phytoplasma. More specific characterization can be obtained with EY-group specific primers fAY/rEY that amplify a 16S rDNA fragment or with FD9 primers that amplifiy the non ribosomal FD9 DNA fragment. vinifera seedlings that later developed GY symptoms. It is not epidemic and occurs most often in old vines in vineyards along creeks or rivers where alders (Alnus glutinosa L) are present. Sabaté et al. Agents: The agent is an EY (16SrV) group phytoplasma. Trapping of insects on sticky traps or with D-Vac suction. alni is highly monophagous and specimen do not survive long on grapevine. PALATINATE GRAPEVINE YELLOWS 1. Transmission: Transmission was obtained with the leafhopper Oncopsis alni Schrank (Hemiptera. Living insects checked for transmissibility of phytoplasma by feeding on artificial medium. Several hemipters have been found to deliver stolbur phytoplasma to the medium. The latter results have not been published. together with high populations of H. (b): First identification of BN infection in the Macedonian republic.: Important epidemics of "Schwarzholzkrankheit" (SHK = VK) in Franken (Bayern.2003 2003 Petrovic et al. O.
Extensive survey of yellows-affected grapevines in several areas of Germany did not show the presence of other EY group phytoplasmas. The strong adaptation of O. Darimont and Maixner: Comparison of the transmission efficiency of phytoplasma to their host plants and to grapevine by O. both insect species trapped on alder. As much as 5 % of leafhoppers from Palatinate and 15 % from Moselle tested positive. The PGY strains are closest to the ALY phytoplasma from alder in Italy and to one strain of FD. However. affected vines often do not survive winter frost because they lack reserves. (b): Detection of a new EY-related phytoplasma in GY-affected cv Scheurebe in Germany. Reinert: PhD thesis on detection. 165 . Description Grapevine yellows (GY) were first reported in New York State (USA) in 1977. a phytoplasma isolate transmitted from alder to periwinkle in Italy. alni. respectively. alni to alder prevents a frequent inoculation to grapevine even if the infestation of the leafhopper population is high. The risk of infection by PGY is lower than for VK because reservoir plants are fewer and outside vineyards. Virginia grapevine yellows III (VGYIII) Main symptoms: North American Grapevine Yellows (NAGY) is a lethal disease causing yellowing of the leaves. Main synonyms: Leaf curl and berry shrivel (LCBS). vinifera seedlings of the PGY phytoplasma using specimen of O. In 1991. In 1993. Variability analysis and sequencing of the FD9 DNA fragment and homology of the sequence to a part of the SecY gene. alni. using Heteroduplex mobility assay and sequencing of rDNA and FD9 fragment. American grapevine yellows. In the same period.2. which was associated with two different phytoplasmas in group 16SrIII and 16SrI (aster yellows). Characterization of the three PGY strains showed that they were closer to FD than to strain EY1 from American elm. alni trapped on alders. obsoletus. a phytoplasma belonging to the X-disease group (16SrIII) was detected in affected grapevines in New York. Identification of the PGY-associated phytoplasma with phytoplasmas infecting alder in the vicinity of affected vineyards. titanus living mostly on wild Vitis riparia around affected vineyards were found to carry an agent that was serologically related to that of FD. alder. alni) is able to transmit the alder yellows phytoplasma to healthy seedlings of alder. Oncopsis alni and Psylla alni specimen. alni (Auchenorrhyncha. populations of S. (b): Transmission to V. 4 Italian and French strains of FD and elm and alder phytoplasmas. Reinert and Maixner: All the 3 strains of PGY were detected both in alder trees and in specimen of O. The differences in vector specificity between PGY and FD phytoplasmas are not supported by molecular data. Historical review 1995 Maixner et al. NORTH AMERICAN GRAPEVINE YELLOWS 1. Angelini et al. Angelini et al. In New York. Main diseases: Virginia grapevine yellows I (VGYI). Reinert and Maixner: 1997. 1997 1999 1999 2000 2000 2000 2001 2003 D. Maixner et al. the latter findings were not confirmed. Infected grapevines often die within months from the onset of symptoms and significant losses of vines have been observed. alni and H. The proportion of infective leafhoppers is lower with O.: Further comparison between elm yellows group (16SrV) phytoplasmas. It is different from FD phytoplasma but molecular and serological data show a close relationship. Same RFLP profiles of P1-P7 rDNA amplicons obtained from yellows affected grapevines. a serious outbreak of GY occurred in Virginia.: Comparison between Elm yellows group (16SrV) phytoplasmas including PGY strains. die back of shoot tips and abortion of developing fruit. Maixner and Reinert: Demonstration that O. the vector of BN / VK. molecular characterisation and epidemiology of PGY. Cicadellidae) (but not P. The alder phytoplasma resembles ALY.
and aster yellows was detected from numerous weeds and shrubs within and around the vineyard. 2. Historical review 1977 Uyemoto et al. Symptoms.: Occurrence of FD-like symptoms on White Riesling grapevines in New York.: Monitoring of S. very similar to those of LCBS. titanus. However. Varietal susceptibility and sensitivity: In New York. vinifera. Maixner et al. allowed the identification of a few candidate vectors of the aster yellows phytoplasma.: Two MLOs were transmitted from affected grapevines to periwinkles in Udine (Italy) (= FDI or FDU or GYU) and New York. and an aster yellows phytoplasma was detected in numerous weeds and shrubs within and around the vineyards. riparia to V. Chen et al. Data showed that the Italian and American MLO were related. The X-disease phytoplasma occurring in New York was also detected with monoclonal antibodies and with probes for dot-blot hybridization. titanus in New York and transmission assays of a possible infectious agent. among which Agallia constricta. 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. X-disease phytoplasma was detected in native Vitis sp. then on White Riesling.: Molecular comparisons between MLOs transmitted to periwinkle from grapevine or detected in a few naturally affected grapevines. Larvae are more frequent on wild Vitis riparia at the border of the vineyards. the survey of vineyards and transmission assays to V. Maixner and Pearson: First data on the study on S. (a): A Riesling plant from New York and a periwinkle carrying the FDI MLO described above. Prince et al. faba bean plants and feeding solutions. The latter observation suggests that another non-related MLO might have been present. Pearson et al. Symptoms may occur for several years in succession in the same vines. the disease was first observed on cv De Chaunac. However. The Virginia Grapevine Yellows (VGY) has been associated with the VGYIII phytoplasma (a strain of X-disease or 16SrIII. subgroup I-A). present in New York and its possible relationship with a GY disease. as well as direct PCR assays from insects. observed in 1983. Control: There are no control methods since vectors and reservoirs of the associated phytoplasmas are still undetermined. other grapevines with strong symptoms tested negative. show that the FDU (= FDI) strain from Italy and a MLO detected in a grapevine in Virginia (USA) are both related to X-disease. Adults migrate from V. Probes and primers yielded positive reactions using DNA extracted from grapevine. In Virginia. Sauvignon blanc and Cabernet franc.: Description of a disease of the yellows type affecting the variety De Chaunac in vineyards of New York State denoted Leaf curl and berry shrivel (LCBS). Detection: With molecular assays using universal or group-specific PCR primers. No transmission by vine planting material reported. it is very severe on Chardonnay vines but was observed on other varieties including Riesling. Daire et al. vinifera cuttings. tested positive for a MLO in the X-disease group with PCR-RFLP of 16S rDNA. subgroup III-B) and with the VGYI phytoplasma (a strain of aster yellows or 16SrI. X-disease phytoplasma was detected in native Vitis sp. ELISA and immunfluorescence were positive from periwinkle. Transmission: No vector insects have been identified for VGY phytoplasmas. It appeared to be spreading and was perhaps insect borne. The FDI MLO was identified in a parallel work as a member of the X-disease group. 1985 1991 1993 1993 1993 1993 166 .Agents: The 16SrIII phytoplasma detected in NY has not been further characterized. Other host plants: Both phytoplasmas are widespread in the USA in weeds and cultivated species. Detection was positive in symptomless wild Vitis riparia growing near vineyards in New York and in some symptomatic grapevines.
Main diseases: AGY (sensu stricto). Mapping of diseased plants show that grapevines appear to be a terminal host of AGY. Late Season Leaf Curl (LSLC)-affected grapevines show symptoms in late summer with tightly downward rolled leaves that often remain green. suggesting that the source is from an alternative host. a GY disease found in Victoria (Australia). which is the more frequent and the Tomato big bud (TBB) phytoplasma. It belongs to the stolbur group (16SrXII) but is distinct from the BN and VK phytoplasmas. subgroup I-A. BVGY phytoplasma has not been clustered in an established phytoplasma group. Both VGYI and VGYIII were found in several weeds and shrubs around and inside the vineyards. The yellow leaf tissue can become necrotic. Buckland Valley Grapevine Yellows (BVGY).: Two phytoplasmas are detected in yellows diseased grapevines in Virginia. Davis et al. It has been suggested that all three phytoplasmas are strains of the same phytoplasma species. Other species tested positive and transmitted to feeding solutions. with a higher incidence in warmer inland districts of New South Wales and South Australia. III-I. 1998 2003 E. Some plants also display uneven bud development resulting in canes and cordons that are bare in places with little or no bunch development. also detected in wild V. Confirmation that a X-disease (16SrIII) group phytoplasma is involved and classification of the latter phytoplasma into a new subgroup. 167 . The TBB phytoplasma has been designated as Candidatus species "Candidatus Phytoplasma australasia". The 16S rRNA gene has a high sequence similarity (more than 99. there are reported cases where they were not associated with AGY disease or phytoplasmas. Conversely. AUSTRALIAN GRAPEVINE YELLOWS 1. Description Grapevine yellows were first reported from Australia in 1976 and called Australian grapevine yellows (AGY) in 1983. It belongs to the peanut witches' broom group (16SrII) and has a broad plant host range in Australia. Shoots may display abortion of bunches or berry shrivel later in the season and death of terminal buds. overlay one another like shingles and become leathery and brittle. leaves tend to fall early. The sequence similarity of the 16S rRNA gene shows that it is close to members of the aster yellows group (16SrI). waxy appearance and remain rubbery. Affected vines carry one or more shoots with irregular veinal of interveinal yellowing and downward rolling of the leaves that overlay one another in a shingled fashion. two syndromes with particular symptoms (Restricted Growth and Late Season Leaf Curl) have been found in some cases. Early and recent surveys have shown that AGY are found in most viticultural regions of Australia. In addition.: A GY disease observed in Virginia since 1983 affects large areas with a low incidence and a low spread rate. In this particular disease. respectively. The use of PCR has shown the association with AGY of three different phytoplasmas. Late Season Leaf Curl (LSLC). is a member of the aster yellows (16SrI) group. Restricted Growth (RG)-affected grapevines show retarded growth resulting in shortened shoots and smaller leaves. riparia vines.5%) with the phytoplasmas associated to Papaya die back (PDB) in Australia and to Phormium yellow leaf (PYL) in New Zealand. clover phyllody phytoplasma (16SrI-C) being the closest . The relatedness of the associated MLO to X-disease MLO is confirmed. Stems of affected shoots develop a blue. one node after the other. followed by the progressive death of the shoots. Agents : AGY disease (sensu stricto) is associated with two phytoplasmas: the AGY phytoplasma. Restricted Growth (RG). consistently associated with AGY symptoms and AGY phytoplasmas but their aetiology is not clear. Beanland and Wolf: Search for possible insect vectors and reservoirs of NAGY phytoplasmas in Virginia. Main symptoms: All symptoms typical for GY can be found on AGY-diseased plants. The AGY phytoplasma has been designated as Candidatus species "Candidatus Phytoplasma australiense". The association of AGY with phytoplasmas was confirmed as early as 1988. Agallia constricta is a suspected vector of VGYI (aster yellows phytoplasma).1993 Wolf et al. The second phytoplasma.
Control: There are no methods to control AGY diseases in the vineyard. once vines are infected by AGY. Magarey: The situation of GY in Australia. Sanitation with hot water treatment is being developed in Australia. Reduction of symptom expression in vines injected with tetracycline. 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. Similarly. Magarey: Comprehensive review of GY diseases in the world. Magarey and Wachtel: Review of the AGY problem.: MLOs. The BVGY phytoplasma has no suspected insect vector. Varietal susceptibility and sensitivity: Chardonnay and Riesling appear to be most often affected. Detection: Detection is obtained with PCR. Phloem fluorescence of diseased vines in the UV microscope. All generic "universal" primers may be used. and trunks in October is more reliable. branches. argentatus and subsequently transmitted to faba bean but the ability to transmit to grapevine was not confirmed. 140-510 nm in diameter. trunks and roots throughout the year and infection is persistent from year to year.: The "Rhine Riesling problem" denoted "Australian Grapevine Yellows". followed by RFLP analysis or sequencing. Several observations suggest that aerial transmission prevails. found in sieve tubes of diseased vines are associated with AGY symptoms. AGY phytoplasmas are common in several crops in Australia. they are at greater risk of displaying symptoms again in the following years. The planthopper Oliarus atkinsoni Myers (Hemiptera.and white-berried varieties. Other host plants: As mentioned above.: Orosius argentatus. Magarey and Wachtel: GY symptoms are found in most viticultural areas of Australia. Phytoplasma australiense) either in single-step or nested PCR following a first amplification with universal primers. RG or LSLC. Phytoplasma australasia). The TBB phytoplasma is transmitted to other crops by O. The incidence of disease may be temporarily high in some vineyards of Chardonnay. The AGY phytoplasma was detected in the common leafhopper Orosius argentatus (Evans) but transmission to grapevine has not been reported. 168 . Remission of AGY. It is possible that AGY diseases have become prevalent in Australia in the last years with the increasing planting of Chardonnay. argentatus. such as Shiraz or Semillon. Magarey et al. a specific primer in the intergenic 16S-23S spacer region has been designed for detection of TBB phytoplasma (Ca. reservoirs must be important but the epidemiology is not fully understood. Hence. RG and LSLC diseases has been observed. suggesting that the disease is the result of an aerial transmission and that the phytoplasma was not present in the original planting material. vector of TBB in tomato crops. However. but phytoplasmas were also detected in other red. 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. However. Magarey et al. Specific primers have been designed on the 16S rRNA gene for detection of AGY phytoplasma (Ca. Detection can be from shoots. is found in high numbers together with two other potential MLO vectors in vineyards of Victoria (Australia). 2. Recent studies have shown that the latter phytoplasma can be acquired from infected grapevine by O. Sampling simultaneously from shoots. Cixiidae) is known as the vector of PYL disease in New Zealand but the species has not been reported from Australia. suggesting that this contributes to the decline in phytoplasma numbers and to the remission of symptoms. indicating persistence of the disease.Transmission: No vectors have been identified for any AGY disease. A "heat curing" effect of AGY disease by hot weather has been observed. Osmelak et al. branches.
: Description of AGY symptoms. Beanland et al. Padovan et al. Wilson et al.: Study of the phylogenetic relationships of the BVGY phytoplasma with aster yellows and stolbur phytoplasmas. (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). The problem of symptomlessly infected vines.: AGY phytoplasma detected in the body of Orosius argentatus leafhoppers. Candidatus Phytoplasma australiense. Liefting et al. Davis et al. Gibb et al. Beanland et al. (a and b): Positive detection of phytoplasma with PCR in AGY-affected grapevines and tentative survey of phytoplasmas in Australia. RG and LSLC are associated. LSLC would be followed by AGY on the following year and AGY would be followed by RG in subsequent years. Constable et al. 1997 1997 1997 1998 1998 1998 1999 1999 1999 2000 2000 2001 2001 2002 2002 2002 169 . Constable and Symons: AGY phytoplasma is detected in most symptomatic grapevines but also TBB phytoplasma and a third uncharacterised phytoplasma. Hence.: Description of the AGY phytoplasma and designation of a new taxon. It is occasionnally detected in symptomless vines and in vines with LSLC or RG symptoms.: Detection of phytoplasmas associated with AGY. Bonfiglioli et al. resulting in increasing cumulative incidence. later called BVGY phytoplasma. Constable et al. Habili et al. The TBB phytoplasma is detected occasionnally in vines with AGY symptoms and also in LSLC-affected vines. Waite et al.: Assumption from field observations and monitoring that AGY. BVGY might form a new AY subgroup of AY (16Sr) group or even a new phytoplasma group.: The AGY phytoplasma is consistently present in the majority of symptomatic grapevines. and LSLC. Uneven distribution of phytoplasmas in infected plants. Constable et al.: Use of hot water treatment in commercial nurseries of Australia. Kelly et al. Papaya dieback and Phormium yellow leaf diseases.: Comparison of visual assessment and diagnostic assays. Spring and summer are optimal for detection.1995 1995 1996 Bonfiglioli et al.: BVGY phytoplasma is associated to GY symptoms that may show remission then reoccurrence but also appear on vines that were not affected previously.: TBB phytoplasmas can be acquired from diseased grapevine by Orosius argentatus and subsequently transmitted to faba bean.: Further comparison of AGY phytoplasma with overseas GY phytoplasmas show that it is closely related to.: No vector for AGY found. RG. Padovan et al. but different from the stolbur phytoplasma associated with BN in France. Constable et al.: Close relationships between phytoplasmas associated with AGY. established from a different source of plant material indicate that BVGY disease occurs as a result of aerial transmission.: Identification of a new phytoplasma in Chardonnay grapevines in Victoria (Australia). Constable: PhD thesis on the biology and epidemiology of AGY phytoplasmas. Detection of BVGY in another plot in the same area. Spain and Israel. insecticide sprays are useless for controlling the disease.
Historical review Europe 1996 Alma et al. a 16SrI-B phytoplasma was found in mixed infection with a stolbur phytoplasma (at that time designated 16SrI-G). The first is the detection in countries other than the USA. In three instances. It seems that aster yellows phytoplasmas have no important significance in European grapevine yellows.: Detection of phytoplasma in the aster yellows group in GY affected plants in Slovenia and Croatia. Statistical analysis showed that RG and LSLC diseases were not always associated with AGY. Saric et al. three different situations must be described. No variability was observed amongst BVGY phytoplasma isolates. The total cumulative incidence for AGY could be as high as 90%. 2. These aster yellows phytoplasmas belong to subgroups I-B or I-C (clover phyllody) which are ubiquitous in Europe. 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. 1997 170 . Transmission: Aster yellows and. The third situation is represented by poorly characterized GYs recorded from new countries. of phytoplasmas belonging to the aster yellows group (16SrI) in yellows-affected grapevines.: 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. Constable and Symons: Study with Heteroduplex mobility assay (HMA) of the elongation factor Tu (Tuf) gene of the AGY and the BVGY phytoplasmas. a variety widely grown in all countries and very sensitive to all GY agents. Description Besides the cases reported above. recurrence in other vines and new occurrence in previously unaffected vines. However. The second situation is the significant occurrence of X-disease group (16SrIII) phytoplasmas in countries other than the USA.: Survey of phytoplasmas infecting grapevine in northern Italy. (a and b): Biology and epidemiology of AGYs. Constable et al. They must not be confused with the 16SrI-G phytoplasma of earlier reports.2002 2003 Crocker et al.: Survey of AGY. RG and LSLC symptoms for 3 to 6 years in Chardonnay and Shiraz vineyards. which was the first designation for stolbur phytoplasma in some laboratories. There is no information of transmission of AGY by planting material. 2004 2004 F. The AGY and the TBB phytoplasmas may persistently affect grapevines throughout the year and between years. is often reported in connection with these cases. Symptomatic shoots from AGY affected vines are reliable for detection in summer (January). detection is more reliable from samples taken from branches and trunks. At other times of the year. are very frequent in many host plant species and several insect species are known as vectors or potential vectors in different countries. to a lesser extent. 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. TBB phytoplasma may have a role in the aetiology of LSLC. OTHER GRAPEVINE YELLOWS 1. The subgroup of aster yellows was not determined but primers specific for stolbur tested negative. later designated 16SrXII. namely Israel. the diseases are characterized by remission in some plants. Constable et al. In Chardonnay. sometimes as a second phytoplasma in co-infection with FD or BN phytoplasmas and sometimes also in symptomless grapevines. Varietal susceptibility and sensitivity: Chardonnay. they seem to have some relevance in Israel and were recently reported from Tunisia. However. X-disease group phytoplasmas.
In addition. some affected vines showed a sudden fall of leaves in summer. Tanne et al. a X-disease phytoplasma was detected in some Neoaliturus specimen. of phytoplasmas belonging to group 16SrI. 2001 2003 Tunisia 2003 Chabbouh et al. 2003 171 . Stolbur phytoplasma (16SrXII) was also detected.. In addition to similarities of symptoms with FD. The spatial and temporal dispersion of the four species was analysed. Merlot and Carmenere.: Survey over several years of potential vectors of phytoplasmas in vineyards of the Golan Heights showed that Neoaliturus fenestratus. 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. Transmission to periwinkle of yellows diseases using collected insects. D'Ascenzo et al. Klein et al.: Formal identification of a 16SrI-B phytoplasma in GY-affected grapevines in Tunisia. including S. Detection with PCR show erratic identification of a 16SrI-B phytoplasma. and Anaceratagalia laevis were all found carrying an aster yellows phytoplasma. Megophthalmus scabripennis was positive for aster yellows phytoplasma.: Identification in yellows-affected vines of varieties Petit Syrah. Circulifer spp.2001 Alma et al. Caudwell et al. M'hirsi et al. 2003 South America 1980 Caudwell: Description of a disease of the yellows type in Chile. Orenstein et al. obsoletus was found in addition to the preceding 5 species. called "Amarilliamento de Elqui". Similar transmission of clover phyllody (16SrI-C) MLO by S. Macrosteles sexnotatus.: First results on the monitoring of GY diseases in Tunisia in imported table grape varieties. Neoaliturus sp.: Experimental transmission of the Chrysanthemum yellows phytoplasma (16SrI-B) to grapevine by four leafhopper species. 1971b). titanus.: Further survey of GY vectors in the Golan Heights.: Monitoring of potential phytoplasma vectors in vineyards in Israel and molecular detection of phytoplasmas in their body. H. subgroups I-B and I-C and also to group 16SrVIII (Ash yellows) in mixed infection with I-B. Orosius orientalis. H. Gajardo et al. obsoletus and Circulifer haematoceps were all three abundant and positive for stolbur and aster yellows phytoplasmas.: 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). titanus had been obtained in 1971 in France (cf. Trapping of hemipters and identification of numerous potential phytoplasma vectors.
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