Arabis mosaic and Prunus necrotic ringspot viruses in hop (Humulus lupulus L.)

Purified virus preparations made from nettlehead-diseased hop plants, or from Chenopodium quinoa, to which the virus was transmitted by inoculation of sap, contained polyhedral virus particles of 30 mμ diameter which were identified serologically as arabis mosaic virus (AMV). There were serological differences between AMV isolates from hop and from strawberry, and also differences in host range and in symptoms caused in C. quinoa and C. amaranticolor. AMV was always associated with nettlehead disease. The nematode Xiphinema diversicaudatum occurred in small numbers in most hop gardens, but was numerous where nettlehead disease was spreading rapidly. Preparations from nettlehead-affected hops also contained a second virus, serologically related to Prunus necrotic ringspot virus (NRSV), in mild and virulent forms which infected the same range of test plants but showed some serological differences. Mild isolates did not protect C. quinoa plants against infection by virulent isolates. Hop seedlings inoculated with virulent isolates of NRSV developed symptoms indistinguishable from those of split leaf blotch disease. Latent infection with NRSV was prevalent in symptomless hop plants. Nettlehead disease is apparently associated with dual infection of AMV and virulent isolates of NRSV. An unnamed virus with rod-shaped particles 650 mμ long was common in hop and was transmitted by inoculation of sap to herbaceous plants. Cucumber mosaic virus was obtained from a single plant of Humulus scandens Merr.     

The etiology of hop chlorotic disease

Hop chlorotic disease was first described in England in 1930, but it has since been seldom seen and its etiology has remained unknown. In 1983 a patch of plants with the disease occurred in a large area of hops (Humulus lupulus) cv. Bramling Cross planted at Yalding, Kent in 1967. All plants in a rectangular area enclosing the disease outbreak were infected with hop mosaic, hop latent and prunus necrotic ringspot viruses; the diseased plants were additionally infected with arabis mosaic virus (AMV). The disease was also associated with seed-transmitted AMV, and was induced in hop seedlings inoculated with partially purified preparations of AMV originating from chlorotic disease-affected hops prepared from Chenopodium quinoa. The disease appears to be caused by AMV, but AMV isolates from hops with chlorotic disease were serologically indistinguishable from AMV isolates from hops with symptoms of bare-bine and/or nettlehead and showed similar pathogenicity in diagnostic hosts. The basis of the difference between isolates in their pathogenicity in hop remains unknown.     

Strains of Prunus necrotic ringspot virus in hop (Humulus lupulus L.)

Purified preparations of Prunus necrotic ringspot virus (NRSV) from hop plants formed two light-scattering zones when centrifuged in sucrose density gradients; the upper and lower zones contained particles 25 mμ and 31 mμ in diameter respectively whose sedimentation coefficients were 79 S and 107 S. NSRV isolates from hop were of two distinct serological types: ‘A’ strains, serologically very closely related to NRSV isolates from apple; and ‘C’ strains more nearly related to NRSV from cherry. The variety Fuggle is tolerant to hop mosaic (not related to NRSV) and different selections of apparently healthy female plants usually contained A strains; but C strains were usually isolated from nettlehead-diseased plants. Either A or C strains occurred in male plants grown with the hop-mosaic tolerant varieties. In mosaic-sensitive varieties (Goldings and Bramlings) apparently healthy female plants tested were usually infected with C strains; either A or C types occurred in mosaic-sensitive male plants. NRSV was not detected in the seventy-four hop seedlings obtained from virus-infected plants. Some varieties developed nettlehead when infected with NRSV (A) or (C) + the hop form of arabis mosaic virus, but not with NRSV (A) or (C) alone. Others developed nettlehead when infected with arabis mosaic virus + NRSV (C) but not with arabis mosaic + NRSV (A). A and C strains can multiply together in the same hop plant. There is evidence of partial antagonism, however, and the fluctuating behaviour of the nettlehead syndrome probably reflects changes in the relative concentration of the two serotypes.     

A satellite-like nucleic acid of arabis mosaic virus associated with hop nettlehead disease

An isolate of arabis mosaic virus (AMV) from a hop plant with symptoms of nettlehead disease induced unusually severe symptoms when transmitted to Chenopodium quinoa. This isolate, called AMV-Ta, yielded particle preparations in which up to 80% of the nucleic acids consisted of a species of low molecular weight (SNA), estimated to be about 75 000 daltons by polyacrylamide gel (PAG) electrophoresis. An isolate free of detectable S-NA (AMV-To) was derived from AMV-Ta by inoculating plants with the two high molecular weight genomic RNA species of AMV (2–8 times 10⁶ and 1–3 times 10⁶ daltons; Murant, 1981) separated from S-NA by PAG electrophoresis. This isolate induced much milder symptoms in C. quinoa. Hop seedlings inoculated with AMV-Ta, either mechanically or by nematodes, developed characteristic nettlehead symptoms. Hop seedlings similarly inoculated with AMV-To remained free of nettlehead symptoms. Two species of S-NA associated with hop nettlehead isolates of AMV were detected at two sites in Kent, and two West Midlands sites. At both sites in Kent and at one of the West Midlands sites, the occurrence of the S-NA species was closely correlated with the incidence of nettlehead symptoms. At the other site in the West Midlands, the occurrence of nettlehead symptoms was too erratic to allow positive correlation of SNA with symptom development. Our results show that S-NA plays an active part in symptom production in experimentally inoculated plants of both hops and C. quinoa. In addition, the close correlation between the occurrence of AMV with additional nucleic acid species, and the incidence of nettlehead symptoms in commercially grown hops, suggests a role for S-NA in the aetiology of this disease.     

Host status of some plants for Xiphinema diversicaudatum (Micol.) and their susceptibility to viruses transmitted by this species

Forty plant species were grown in pots containing viruliferous Xiphinema diversicaudatum (Micol.) for 15 wk to assess the host range of the nematode in relation to infection with arabis mosaic (AMV) and strawberry latent ring-spot (SLRV) viruses. Host status for the nematode was determined mainly from changes in total populations, but the presence of eggs in the uteri of females and changes in the numbers of adults provided additional criteria. The nematode multiplied on relatively more woody perennials than on herbaceous crop plants or weeds. Chrysanthemum coronarium was the only plant on which numbers declined significantly below those on the controls. Most plant species became infected with either AMV or SLRV. Neither virus was detected in eight out of thirteen species of trees and shrubs although four were good hosts for the nematode. Galling or distortion of the terminal region of fine feeder roots, associated with X. diversicaudatum feeding, was seen on many of the experimental plants.     

The transmission of strawberry latent ringspot virus by Xiphinema diversicaudatutn (Nematoda)

Strawberry latent ringspot virus (SLRV) was found in diseased rose bushes growing in a glasshouse where the soil contained Xiphinema diversicaudatum (Micol). Adult female, adult male and juvenile X. diversicaudatum all transmitted the virus to cucumber seedlings, and nematodes kept without plants for 32 days after acquiring SLRV later transmitted it. When transferred to fresh plants every 2–4 days for 3 weeks, single nematodes transmitted up to three times; one nematode did not transmit until 19 days after the transfers began. One adult X. diversicaudatum, out of 141 tested, transmitted both SLRV and arabis mosaic virus. In all respects X. diversicaudatum behaved as a vector of SLRV as it does as a vector of arabis mosaic virus.     

Incidence and Control of Necrotic Leaf Mosaic Caused by Arabis Mosaic Virus in Lilium tigrinum splendens in the Netherlands

Necrotic mosaic on leaves and ring spots on bulb scales of Lilium tigrinum splendens, can be caused by arabis mosaic virus (AMV). Primarily infected bulbs can show spongy roots and large necrotic areas on creamy coloured bulb scales. Consecutive series of plants replanted for monthly periods in infested soil were mostly infected by AMV at a high rate (70%) throughout the year. Very low or undetectable numbers of Xiphinema diversicandatum, nematodes in soil dilution experiments infected lilies very efficiently (50–70 %). In general, soil disinfestation with dichloropropene, dazomet, methylbromide, and other disinfectants were variably fairly effective, particularly when yellow crocus among which couch was abundant, was previously grown for two years. The influx of AMV infected material into the soil was assumed to increase the number of AMV-carrying nematodes, and may be one cause of the failure of soil disinfestation. A survey of AMV infested soil in lily-growing regions in The Netherlands indicated its occurrence in a few fields only. Complex control measures applicable under growers' conditions, in addition to the variably effective soil disinfestation, are indicated.     

Hop line-pattern virus in relation to the etiology and distribution of nettlehead disease

Hop line-pattern virus (HLPV) was transmissible by mechanical inoculation to hop plants; it induced characteristic severe symptoms in Humulus lupulus L. var. neo-mexicanus Nels. & Cockerell and the commercial derivatives College Cluster and Keyworth's Midseason, but none in the traditional English varieties of H. lupulus (e.g. Fuggle).
Mechanical transmission of hop nettlehead virus (HNV) was facilitated by the presence of HLPV in the test plants; hop seedlings and clonal plants escaped infection by sap inoculum that infected plants of two varieties already infected with HLPV. HNV was also transferred by stem contact and by knife cuts to plants carrying HLPV.
Infection with HLPV was latent in twelve nettlehead-diseased Fuggle plants from different fields, and in diseased and symptomless plants in a nettlehead outbreak in W.G.V., a variety that previously had escaped infection. It is suggested either that HLPV predisposes hop plants to infection with HNV or that nettlehead disease is caused by dual infection with both viruses.
Localized and scattered patterns of nettlehead spread were observed in hop plantations; these two types are usually attributed to different modes of spread which would be compatible with a complex etiology of the disease.     

A damaging outbreak of arabis mosaic nepovirus in blackcurrant, the occurrence of other nepoviruses in Ribes species, and the demonstration that alfalfa mosaic virus is the cause of interveinal white mosaic in blackcurrant

In a crop of blackcurrant (Ribes nigrum), cv. Baldwin in Eire, chlorotic mottling and ringspot symptoms in leaves on plants and severe crop loss was associated with infection with arabis mosaic nepovirus (ArMV) and the presence in the soil of its nematode vector, Xiphinema diversicaudatum. This is only the second report of ArMV damaging a crop of blackcurrant. Tomato black ring (TBRV) and raspberry ringspot nepoviruses were detected in single plants of redcurrant (R. rubrum) in England and flowering currant (R. sanguineum) in Scotland respectively; each of these infected plants showed foliar chlorotic line-pattern symptoms. This is the first record of TBRV in redcurrant. A single blackcurrant plant in New Zealand showing symptoms typical of those described for interveinal white mosaic disease, contained alfalfa mosaic virus (AMV). When AMV particles were purified and concentrated from herbaceous test plants and mechanically inoculated to young blackcurrant plants, several became infected with AMV and most infected plants developed systemic symptoms typical of the original disease. This provides the strongest evidence to date that AMV is the causal agent of interveinal white mosaic disease.     

Seed-transmission of nematode-borne viruses

Transmission through seed of crop and weed plants seems to be characteristic of nematode-borne viruses. It occurred with tomato black ring virus (TBRV) in nineteen species (thirteen botanical families), with arabis mosaic virus (AMV) in thirteen species (eleven families), with raspberry ringspot virus (RRV) in six species (five families), and also, in more limited tests, with tomato ringspot, cherry leaf roll and tobacco rattle viruses. A remarkable feature was that infected seedlings, except those containing tobacco rattle virus, often appeared healthy. The occurrence and extent of seed-transmission depended on both the virus and the host plant. In many progenies more than 10%, and in some 100%, of seedlings were infected. The viruses were transmitted through at least two or three generations of seed of those host species tested. After 6 years' storage, TBRV- and RRV-containing seed of Capsella bursa-pastoris and Stellaria media germinated to give infected seedlings. In controlled crossing experiments with strawberry and raspberry, virus was transmitted to seed from both male and female parents but, at least in raspberry, the presence of competing virus-free pollen much decreased the ability of pollen from infected plants to set seed. There was no evidence that healthy mother plants became infected when their flowers were pollinated with infected pollen.     

Sites of virus retention in the alimentary tract of the nematode vectors, Xiphinema diversicaudatum (Micol.) and X. index (Thorne and Allen)

Electron microscopy of thin sections of Xiphinema diversicaudatum and X. index fed on plants infected respectively with arabis mosaic and grapevine fanleaf viruses showed that the viruses are retained as a monolayer of particles adsorbed on to the cuticle lining the lumina of the odontophore (stylet extension), anterior oesophagus and oesophageal bulb. During the moult of the nematode the cuticular lining is shed and together with the detached virus particles is ingested into the intestine through the oesophago-intestinal valve; this supports the limited experimental evidence that viruses transmitted by X. diversicaudatum and X. index are not retained through the moult.     

The isolation and identification of rhubarb viruses occurring in Britain

Virus-like symptoms were common in British crops of rhubarb. All plants tested of the three main varieties, ‘Timperley Early’, ‘Prince Albert’ and ‘Victoria’, were virus-infected. Turnip mosaic virus and a severe isolate of arabis mosaic virus (AMV) were obtained from ‘Timperley Early’; and ‘Prince Albert’ contained turnip mosaic virus, cherry leaf roll virus (CLRV), a mild isolate of AMV and, infrequently, cucumber mosaic virus (CMV). The main commercial variety ‘Victoria’ contained turnip mosaic virus, CLRV, a mild isolate of AMV and, infrequently, strawberry latent ringspot virus (SLRV). All the viruses were identified serologically. The rhubarb isolates did not differ markedly from other isolates of these viruses in herbaceous host reactions, properties in vitro or particle size and shape. A rhubarb isolate of CLRV was distinguished serologically from a cherry isolate of the virus. Turnip mosaic virus, CLRV and SLRV, were transmitted with difficulty, but AMV isolates were readily transmitted by mechanical inoculation. Turnip mosaic virus was also transmitted to rhubarb by Myzus persicae and Aphis fabae. CLRV was transmitted in 6–8% of the seed of infected ‘Prince Albert’ and ‘Victoria’ rhubarb and in 72% of the seed of infected Chenopodium amaranticolor. Mild isolates of AMV were also transmitted in 10–24% of the seed of infected ‘Prince Albert’ and ‘Victoria’ plants.     

The association of split-leaf blotch virus with nettlehead disease of hops

The distribution of two virus diseases, split-leaf blotch and nettlehead, in hop plantations suggested that infection with split-leaf blotch virus (SLBV) predisposed plants to nettlehead. Also in three field trials, the incidence of nettlehead disease was higher in plants (var. Fuggle) infected with SLBV than in healthy plants.
The symptoms shown by Fuggle plants infected with SLBV were unaffected by additional infection with hop line-pattern virus (HLPV), but plants of the varieties College Cluster and Keyworth's Midseason infected with both viruses developed symptoms resembling nettlehead disease in some respects. Such affected plants did not, however, induce nettlehead in plants of the indicator variety Early Prolific to which they were grafted, and therefore nettlehead disease is not the result of dual infection with SLBV and HLPV.
Despite the field association of split-leaf blotch and nettlehead symptoms, experimental data indicate that SLBV is not a component of the virus complex causing nettlehead disease.     

The association of Xiphinema diversicaudatum (Micoletsky) with strawberry latent ringspot and arabis mosaic viruses in a raspberry plantation

An outbreak of strawberry latent ringspot virus (SLRV) in a plantation of Mailing Jewel raspberry coincided with the greatest abundance of the nematode vector, Xiphinema diversicaudatum. Arabis mosaic virus (AMV) was not detected in the crop but was, together with SLRV, in many weed species present. AMV was transmitted through the seed of Poa annua, Capsella bursa-pastoris and Senecio vulgaris and SLRV through the seed of Mentha arvensis. X. diversicaudatum were more numerous within the rows than between them and vertical sampling showed that most occurred between 4 and 12 in depth in both locations. Monthly sampling showed that egg laying occurred from April to July; populations increased to a peak in late autumn but declined during the winter, resulting in about a twofold annual increase in numbers. Females, males and juveniles transmitted AMV and SLRV to cucumber seedlings, and in the absence of plants the nematode retained AMV for 112 days and SLRV for 84 days.     

Infectibility of some new raspberry cultivars with arabismosaic and raspberry ringspot viruses and further evidence for variation in British isolates of these two nepoviruses

In field trials at sites of an outbreak of arabis mosaic nepovirus (AMV) in England and of raspberry ringspot nepovirus (RRV) in Scotland, the results of exposure of some new raspberry cultivars to natural infection with these viruses showed discrepancies from those obtained in graft inoculation tests using AMV-Lib and RRV-S, the Scottish type isolates. In particular, cv. Glen Prosen, which is immune to AMV-Lib and RRV-S, was infected with AMV and RRV in the field trials. Studies on these and other field isolates of AMV and RRV showed that they differed from the type isolates in Rubus host range and in symptomatology in herbaceous hosts. However, whereas four isolates of RRV found infecting Rubus were distinguishable by spur formation in gel double-diffusion serological tests, six AMV isolates were indistinguishable by this method. Immunoelectrophoresis of virus particles did not distinguish the six AMV isolates, but isolates RRV-MX and RRV-T were distinguishable from RRV-S and the English type isolate, RRV-E. Like the two RRV type isolates, RRV-MX contained a single electrophoretic component, but it migrated must faster whereas RRV-T contained two components, one with a migration rate similar to that of RRV-MX and the other similar to that of the type isolates. Polyacrylamide gel electrophoresis of protein preparations from highly purified virus particles of RRV isolates E, S and MX detected a single polypeptide of estimated mol. wt 54 × 10³, 54 × 10³ and 50 × 10³ respectively but that of isolate T contained two polypeptides of estimated mol. wt 54 × 10³ and 50 × 10³. These data suggest that RRV-T is a mixture of two isolates. In laboratory tests the nematode Xiphinema diversicaudatum transmitted four isolates of AMV efficiently whereas two populations of the nematode Longidorus elongatus were less efficient vectors of four RRV isolates. Neither vector species transmitted virus to any of nine raspberry cultivars. The results are discussed in relation to the control of nepoviruses in raspberry and to the biology of these viruses.     

Seed-transmission in the ecology of nematode-borne viruses

Virus-free populations of vector nematodes can acquire tomato black ring (TBRV), raspberry ringspot (RRV) and arabis mosaic (AMV) viruses from weed seedlings grown from virus-carrying seed. When soils from fields where nematode-borne viruses occurred naturally were air-dried to kill vector nematodes and then moistened, TBRV and RRV occurred commonly in the weed seedlings that grew, but AMV occurred only rarely. Similar tests did not detect tobacco ringspot, grapevine fanleaf or tobacco rattle viruses in weed seeds in the single soil studied in each instance, although these three viruses are also seed-borne in some of their hosts. Many weed species, when infected experimentally, readily transmit TBRV and RRV to their seed, but the viruses were much commoner in naturally occurring seed of some of these species than of others. These discrepancies between the frequency of seed-transmission of viruses from experimentally infected plants and the extent of natural occurrence of infected seed seem largely to reflect the host preferences of the vectors. Infective Longidorus elongatus kept in fallow soil retained TBRV and RRV only up to 9 weeks. When weed seeds in the soil were then allowed to germinate, the nematodes reacquired virus from the infected seedlings. Some weed species were better than others as sources of virus. Persistence of these viruses in fields through periods of fallow or fasting of the vector therefore depends on a continuing supply of infected seedlings produced by virus-containing weed seeds. This is probably less true of viruses like AMV and grapevine fanleaf, which persist for 8 months or more in their vectors (Xiphinema spp.). A few seeds containing TBRV and RRV were found in soils free of vector nematodes, suggesting that the viruses are disseminated in weed seed. This probably explains how TBRV and RRV have reached a large proportion of L. elongatus populations in eastern Scotland.     

The detection by serological methods of viruses infecting the rose

Homogenates of herbaceous test plants infected with arabis mosaic virus (AMV), prunus necrotic ringspot virus (PNRSV), or strawberry latent ringspot virus (SLRV), and purified virus preparations were used to assess the sensitivities of four serological methods (the enzyme-linked immunosorbent assay - ELISA, immunodiffusion in gels, the latex flocculation assay, and serologically specific electron microscopy -SSEM) for the detection of these viruses. The latex test was up to 250 times more sensitive than gel immunodiffusion, but SSEM and ELISA were respectively up to 1000 and 200 times more sensitive than the latex test. Gel immunodiffusion and latex tests failed to detect any of the viruses in infected roses. Although ELISA reliably detected PNRSV and SLRV when leaves from infected roses were homogenised in a leaf: buffer ratio of 1 g:10 ml, AMV was occasionally undetected. However, when a modified ELISA technique, which reduced non-specific reactions, was used some PNRSV-infected roses were also not detected. Detection by SSEM was c. twice as sensitive as ELISA for all three viruses in rose extracts. The relative advantages of ELISA and SSEM for the detection of plant viruses are discussed.     

Further studies on a British form of pea early-browning virus

An isolate of pea early-browning virus from Britain (PEBV (B)) has tubular particles most of which are either about 103 or 212 mμ long with sedimentation coefficients of 210 and 286 S respectively. Both types show cross-banding at intervals of 2.5 mμ. Virus preparations containing only the shorter particles were not infective. PEBV (B) was transmitted to pea seedlings by both adult and juvenile Trichodorus primitivus (de Man) (Nematoda) and persisted for 32 days in T. primitivus kept without plants. In two experiments T. primitivus failed to transmit a Dutch isolate (PEBV (D)), which is distantly related serologically to PEBV (B). PEBV (B) was transmitted by nematodes to cucumber roots more readily in soil at 20d? than at 24d? C., and more readily at 24d? than at 29d? C. When transmitted by inoculation of sap, PEBV (B) and PEBV (D) caused similar symptoms in some pea varieties but differed in virulence towards others. Thirty-one varieties resistant to natural infection with PEBV in The Netherlands were susceptible to PEBV (B) when manually inoculated with sap or when grown in naturally infested soil from one site; twenty-six of these varieties did not become infected in soil from a second site, in which several other varieties that are susceptible in The Netherlands were infected. Varieties should therefore be tested for resistance by growing them on many infested fields. All but one of the pea varieties resistant to PEBV in The Netherlands became infected with the English form of tomato black ring virus when grown in soil containing infective Longidorus attenuatus Hooper.     

Epidemiology of three viruses infecting the rose in the United Kingdom

Studies on the epidemiology of arabis mosaic (AMV), prunus necrotic ringspot (PNRSV) and strawberry latent ringspot (SLRV) viruses were made in relation to commercial production of standard and bush roses. AMV or SLRV apparently induced either symptomless infection in rose cultivars and Rosa spp., or leaf symptoms ranging from small chlorotic flecks to severe chlorotic mosaic and, occasionally, plant death. Infection of R. canina ‘inermis’ or R. corymbifera by an isolate of SLRV from R. corymbifera also severely depressed flowering and hip formation. In addition, whereas this isolate could be graft-transmitted to all Rosa spp. tested, isolates from R. rugosa and R. multiflora failed to be graft-transmitted to R. canina ‘inermis’ or R. corymbifera. No difference was detected in graft-transmission tests of Rosa spp. with several isolates of AMV or PNRSV. In plantings of up to 7 yr none of the viruses was transmitted through pollen to healthy roses grown in nematode-free soil, and only SLRV was readily seed-transmitted, particularly in R. rugosa. Nevertheless, in soil containing viruliferous nematodes, AMV and/or SLRV were transmitted to c. 80% of healthy plants. AMV and particularly SLRV were each damaging to field-grown maiden rose bushes cv. Fragrant Cloud. SLRV delayed the onset of flowering, and reduced the number and size of blooms. Diseased bushes were less vigorous, and half or none of the AMV- or SLRV- infected bushes respectively, conformed to the British Standards Institution specifications for maiden bush roses. These results are discussed in relation to the commercial production of field-grown roses in the UK.     

Viral twig necrosis of sweet cherry. Modes of transmission and spread of petunia asteroid mosaic virus (PeAMV)

Grafting symptomless scions, derived from petunia asteroid mosaic virus (PeAMV)-infected trees, to healthy rootstocks resulted in only 3.3% infection in the resulting trees. Up to 90% of seeds from infected sweet cherries contained high quantities of PeAMV, but the virus was not transmitted to the seedlings apparently because of low virus content in the embryo and loss of infectivity during seed maturation and storage. Replanting healthy cherry trees cv. Sam, grafted to different rootstocks, into contaminated soils resulted in new infections. Eight of 13 trees on rootstocks derived from Prunus avium (F 12/1 and cv. Sam on its own roots) were infected with PeAMV within a period of four years but only one of 16 trees on Weiroot-rootstocks (selections from Prunus cerasus) became infected. The detection of PeAMV in naturally contaminated soil samples by the bait plant procedure, using Nicotiana clevelandii, was superior to testing soil eluates by enzyme-linked immunosorbent assay (ELISA) and immuno electron microscopy (IEM). Wild plants may contribute to virus propagation and maintenance of virus contamination of the soil as 25 of 310 samples from 712 herbaceous plants growing in the vicinity of infected trees contained PeAMV; the contaminated samples represented 12 species. The perpetuation of PeAMV by infected scion wood is probably of minor significance, and infection via the soil probably represents the most important means of spread of viral twig necrosis in northern Bavaria.