Decreased severity of ryegrass mosaic virus after passage through resistant perennial ryegrass

After exposure to infection in the field, the proportion of plants showing distinct symptoms of ryegrass mosaic virus (RMV) was less in perennial than in Italian ryegrass. The perennial ryegrass cv. Mascot had a smaller proportion of plants with symptoms than cv. S.23. Far milder symptoms were induced in test plants by RMV from naturally infected perennial ryegrass plots than from Italian ryegrass plots. Within perennial ryegrass, RMV from cv. Mascot caused milder symptoms than that from cv. S.23. Severe RMV isolated from Italian ryegrass cv. Trident (RMVT) became milder after one passage through cv. Mascot, although not as mild as RMV obtained from field plots of cv. Mascot (RMVM). Families from two highly resistant perennial ryegrass clones and two randomly selected clones of cv. S.23 crossed in all possible combinations varied in symptom severity when inoculated with RMVT but not when inoculated with RMVM. Families inoculated with RMVT also yielded virus which varied in the severity of symptoms induced in test plants, families with severe symptoms yielding severer virus. Thus, much of the variation in the resistance of these clones could be attributed to infection with RMV of differing severity. Resistance was controlled by several genes which were mainly additive in their effect.     

Selection of perennial and Italian ryegrass plants resistant to ryegrass mosaic virus

Perennial ryegrass plants collected from fields and Italian ryegrass plants grown from seed were selected for resistance to infection by ryegrass mosaic virus (RMV) by repeated manual inoculation. Two of 108 perennial ryegrass plants and one of 150 Italian ryegrass plants were symptomless after seven and nine inoculations respectively. These three plants were propagated vegetatively. Plants of the two perennial ryegrass clones showed no symptoms after further manual inoculations with the initial isolate of RMV, or with an inoculum from infected plants collected from several fields, or after inoculation by viruliferous mites. Electron microscopy and back tests indicated that the plants were virus free. Some plants of the selected Italian ryegrass clone became infected after a further inoculation with mites or sap, but fewer than similarly inoculated unselected plants.     

Serological detection of ryegrass mosaic virus and ryegrass seed-borne virus

Ryegrass mosaic virus (RMV) was reliably detected in both perennial (S24) and Italian (S22) ryegrass, by enzyme-linked immunosorbent assay (ELISA) when plants had been infected for 8 wk. ELISA detected more infections in field-grown perennial ryegrass cv. Premo than either visual assessment or electron microscopy. However, with plants of Italian ryegrass cultivars only recently infected with RMV, positive reactions were more difficult to separate from the reactions of RMV-free plants, which varied considerably with cultivar, some giving high absorbance values. Immunosorbent electron microscopy showed that the RMV antiserum also contained antibodies to ryegrass seed-borne virus (RGSV), suggesting that these high values were caused by RGSV infection in the material tested.     

Effect of cocksfoot mottle virus on resistant and susceptible cocksfoot cultivars grown alone or in combination with healthy or ryegrass mosaic virus-infected Italian ryegrass

Cocksfoot mottle virus (CfMV) was readily transmitted on cutting implements. However, after six harvests, only a quarter as many plants of the resistant cv. Cambria as of the susceptible cv. S.37 became infected. In addition to resistance to becoming infected, cv. Cambria also possessed some resistance once infected, and suffered less mortality than S.37. More CfMV-infected plants died when competing with healthy plants than when competing with other infected plants. Both CfMV and ryegrass mosaic virus (RMV) caused greater reductions in the yield of their respective hosts in mixed stands of cocksfoot and Italian ryegrass than in single species stands. CfMV reduced the proportion of cocksfoot in the mixture and RMV the proportion of ryegrass. However, CfMV-infected cv. Cambria competed better than CfMV-infected cv. S.37 with healthy or with RMV-infected ryegrass. It is concluded that grass cultivars selected for resistance to virus infection will slow down the rate of disease spread within the sward, and selecting for postinfection resistance will help maintain the aggressiveness and competitive ability of plants once they become infected.     

Interaction between crown rust (Puccinia coronata) and two viruses of ryegrass

In Lemtal Italian and S.24 perennial ryegrass plants, two isolates of ryegrass mosaic virus (RMV) suppressed the amount of crown rust emerging on leaves inoculated with Puccinia coronata uredospores by up to 75% compared with the amount on virus-free plants. Severity of rust infection on barley yellow dwarf virus (BYDV) infected plants generally did not differ significantly from that on virus-free plants. When both RMV and BYDV were present, rust was restricted in Lemtal plants to a level intermediate between those occurring on plants infected by either virus alone, and in S.24 plants to a level below that obtained with either virus alone. The mean water soluble carbohydrate (WSC) content of Lemtal plants was reduced more than 20% by RMV, but was not significantly altered by BYDV. In S.24 plants the WSC content was increased by 10% by RMV and by 60% by BYDV. Rust reduced the WSC content of healthy and virus-infected plants, the reduction being positively correlated with the level of rust on the sampled leaves. In plants of Lemtal, but not of S.24, the degree of rust infection was positively correlated with the WSC content of leaves from rust-free control plants.     

Tolerance to ryegrass mosaic virus, its assessment and effect on yield

Symptom severity of eighteen populations of Italian ryegrass infected artificially in the glasshouse and naturally in the field with ryegrass mosaic virus (RMV) was strongly correlated. A smaller proportion of plants of the more tolerant populations showed symptoms in the field, but this was probably due to an association of tolerance with increasing incidence of symptomless infection rather than with resistance to infection. Under sward conditions, the yield of a sensitive genotype was reduced by 27% and that of two more tolerant ones by 15 and 13 %. The percentage dry matter yield loss of the most sensitive genotype was similar in all cuts, despite the appearance of extensive necrosis at the time of one cut. With the more tolerant genotype the yield loss varied from 7 to 25 % according to cut. Over the period of the experiment RMV infection did not increase plant mortality.     

The transmission and effect on yield of ryegrass mosaic virus in a filtered air environment

The eriophyid mite vectors of ryegrass mosaic virus (RMV) were excluded from ryegrass by covering plots with a polyethylene film house ventilated by filtered air. Initially plots were manually inoculated to give different levels of RMV. During two seasons no mites were found within the house so that all transmission of virus was by sap. Rolling and frequent cutting of uninfected plots decreased yields, but surprisingly, increasing virus infection did not decrease yields significantly. Cutting transmitted virus within infected plots but not from infected to healthy plots, whereas combined rolling and cutting did transfer virus to healthy plots.     

Tolerance to ryegrass mosaic virus and its inheritance

Plant yield within and between four cultivars of perennial ryegrass infected with ryegrass mosaic virus (RMV) was closely related to symptom severity. Distribution of symptom severity was continuous in four perennial ryegrass and four Italian ryegrass cultivars infected with a severe RMV isolate, and also in another perennial ryegrass cultivar infected with a severe isolate of the virus, a mild one and one of intermediate severity. Symptom expression was polygenically inherited in both Italian (cv. RvP) and perennial (cv. S.24) ryegrass. Both additive and non-additive genetic variation was present in RvP, but the variation in S.24 was additive only. No significant maternal inheritance was present in either species.     

The elimination of ryegrass mosaic virus from Lolium multiflorum by meristem tip culture

Meristem tips were cultured from Lolium multiflorum plants infected with ryegrass mosaic virus (RMV). Meristem tips within the size range O'2-fi mm long were cultured on media with or without 2 ,4-D at 1 mg/1. The plants that regenerated in culture showed no symptoms over a 10 month period and no RMV particles were observed by electron microscopy. It was concluded that RMV had been eliminated. The method should prove useful in eliminating the virus from desirable genotypes used as parent material for seed production.     

The effect of some isolates of ryegrass mosaic virus on different genotypes of Lolium multiflorum

Some isolates of ryegrass mosaic virus produced necrosis in Lolium multiflorum much more frequently than others. This frequency could be increased or decreased by selecting plants with or without necrosis, respectively, as sources of inoculum. Infection with the milder, mosaic-inducing isolates which rarely caused necrosis did not protect the plants against subsequent infection with the more severe necrosis-inducing isolates. Oat plants were infected much more readily by isolates which frequently caused necrosis than by the less necrotic isolates, but passage through oat appeared to have no effect on the frequency with which an isolate produced necrosis in ryegrass.
Genotypes of L. multiflorum varied greatly in their tolerance of infection. Their level of tolerance was closely correlated with symptom expression. All genotypes tested were tolerant to a mild, mosaic-inducing, isolate. Two genotypes were tolerant to a severe, necrosis-inducing, isolate and to an original Rothamsted isolate; three genotypes were equally susceptible to both isolates, and two were more susceptible to the necrotic isolate than to the Rothamsted isolate.     

The adaptation of tomato mosaic virus to resistant tomato plants

Tomato mosaic virus derived from susceptible tomato plants (the standard virus) was cultured in resistant plants. Sap from non-inoculated leaves of resistant tomato plants infected with virus from the resistant host was more infective and contained more virus particles than leaf sap of resistant plants infected with the standard virus. Leaves of resistant tomatoes infected with virus from the resistant host also showed more obvious symptoms. Susceptible plants infected with virus from resistant plants not only showed fewer symptoms than when infected with standard virus, but samples were less infective and contained less virus up to 26 weeks, when values for infectivity were similar. This modification in activity was not reversible and was obtained with two lines of tomato having different types of resistance. Passage of virus from resistant plants through susceptible plants did not impair its ability to infect resistant plants.     

The effect of ryegrass mosaic virus on the yield of perennial ryegrass swards

The influence of ryegrass mosaic virus (RMV) on yield of perennial ryegrass swards was examined under two harvesting frequencies and at two levels of nitrogen fertilizer, in a field trial planted with a clone of ryegrass, cv. S 321, using small plots established with proportions of healthy and infected plants. RMV reduced the height and the number of tillers of plants grown in pots and reduced the height also of plants in the field. The swards were planted in August and at a harvest 8 wk later RMV reduced the yield from 2–12 to 1–52 t dry matter (d.m.)/ha. In the first full harvest year, RMV caused only a small reduction in yield where no N fertilizer was applied and the application of a very small amount of fertilizer would have recouped the loss. On the other hand RMV severely restricted the increase in yield resulting from heavy dressings of the fertilizer. Thus where 400 kg N/ha was applied, RMV reduced the total annual yield from 18-6 to 13-8 t d.m./ha. The effect of the virus was especially great both in the Spring, the period of maximum herbage production, and when the level of fertilizer was high.     

A field study of the pattern of local spread of ryegrass mosaic virus in mown grassland

In a field experiment at Hurley, small areas of old ryegrass sward, known to be infected with ryegrass mosaic virus (RMV) were partially surrounded by newly-sown Italian ryegrass. The experiment was harvested by mowing on five occasions. Samples of Italian ryegrass taken when it was 11 months old showed a very marked increase in the levels of RMV close to the areas of old sward. Thus, 84% of the Italian ryegrass plants bordering the old sward to the north contained RMV, compared with 63% of plants only 2.7 m away. The corresponding values to the south were 68% and 39%. The much higher values encountered to the north are presumed to be caused by the movement of the mite vector of RMV (Abacarus hystrix) in relation to the prevailing wind. Two different mowers were used, a rotary mower and a reciprocating knife mower. We could demonstrate no increase in the level of RMV as a result of mowing from the infected sward and on to the new sowing and could demonstrate no difference in the infection levels as a result of using the different mowers. We conclude that in mown swards migration by the mite is responsible for all or nearly all of very local spread of RMV.     

Transfer of polygenic resistance to ryegrass mosaic virus from perennial to Italian ryegrass by backcrossing

Polygenic resistance to ryegrass mosaic virus (RMV) was transferred from perennial to Italian ryegrass cv. Tribune by repeated cycles of backcrossing, polycrossing and selection. The resulting synthetic variety (Bb 2113) was highly resistant to infection by RMV whereas the Italian ryegrass cultivars Tribune, RvP, Bartolini, Augusta, Wilo and Dalita all were highly susceptible. Leaf width and length of Bb 2113 was similar to that of cv. Tribune and, like Tribune, Bb 2113 had awned florets. In a field trial, Bb 2113 had a similar persistency rating and number of inflorescences in the regrowth to cv. Tribune but produced 11% less dry matter. The results emphasise the need for a more efficient means of transferring small numbers of genes from one species of Lolium to another than backcrossing provides.     

The effects of mixing Italian ryegrass (Lolium multiflorum) with perennial ryegrass (L. perenne) or red clover (Trifolium pratense) on the incidence of viruses

Mixing the ryegrass mosaic virus (RMV) resistant perennial ryegrass (Lolium perenne) cv. Endura with the susceptible Italian ryegrass (L. multiflorum) cv. RvP decreased infection of RvP wth RMV from 37% when grown alone to 22% when mixed. However, Endura yielded less than RvP and there was no yield benefit from mixing the two cultivars. Mixing red clover (Trifolium pratense) cv. Hungaropoly with RvP had no detectable effect on RMV incidence in RvP but did decrease the incidence of red clover necrotic mosaic virus in Hungaropoly from 9% to 1% and of white clover mosaic virus from 53-5% to 41%. The yield of the mixture was equal to that of RvP grown alone but given nitrogen fertiliser. The numbers of eriophyid mites, including Abacarus hystrix the vector of RMV, on ryegrass leaves were similar in pure and mixed swards. It is concluded that with herbage crops, the common practice of sowing mixtures of species may help control virus diseases.     

Endophytic fungus decreases plant virus infections in meadow ryegrass (Lolium pratense)

We studied the effects of fungal endophyte infection of meadow ryegrass (Lolium pratense=Festuca pratensis) on the frequency of the barley yellow dwarf virus (BYDV). The virus is transferred by aphids, which may be deterred by endophyte-origin alkaloids within the plant. In our experiment, we released viruliferous aphid vectors on endophyte-infected and endophyte-free plants in a common garden. The number of aphids and the percentage of BYDV infections were lower in endophyte-infected plants compared to endophyte-free plants, indicating that endophyte infection may protect meadow ryegrass from BYDV infections.     

THE PATTERN OF FIELD INFECTION OF POTATO BY THE BEET RINGSPOT STRAIN OF TOMATO BLACK-RING, A SOIL-BORNE VIRUS

The potential importance of the beet ringspot strain of tomato black-ring, a soil-borne virus, was assessed by growing stocks of Kerr's Pink potato for 1 year on infested land and subsequently on uninfested land. The incidence of infection in two stocks was 39 and 8% in the first year on uninfested land, and 29 and 5% after 2 years.
The virus was usually restricted to the roots of plants in the first year of infection, but a few plants showed black rings and spots in their leaves. In the second year, 20–55% of the plants grown from tubers set by symptomless, but infected, mother plants were infected: many of these showed leaf necrosis, others had stunted shoots, and cupped and distorted leaves; some were symptomless although systemically infected. In the third and fourth years, most of the progeny from plants which had symptoms or which were symptomless but systemically infected, contained the virus: nearly all such infected plants were stunted and distorted or were symptomless. Infection decreased the weight of tubers produced by plants with severe necrotic spotting but not the yield of plants with less necrosis. The number and weight of tubers per plant were decreased by 15 and 20% respectively, in symptomless systemically infected plants, and by 20 and 30% in stunted plants.     

Variation in reaction to barley yellow dwarf virus in ryegrass and its inheritance

Genotypes of Italian and perennial ryegrass differed greatly in their reaction to infection with barley yellow dwarf virus (BYDV). One genotype of perennial ryegrass appeared unaffected, whilst the yield and height of other genotypes were reduced. Progeny of the Italian ryegrass genotypes showed a similar degree of variation. Tolerance, assessed by the effect of the virus on yield, was inherited in a wholly additive manner. Symptoms were rare and their severity was under both additive and non-additive genetic control and strongly correlated with increasing severity of reaction. The utilization of the variation in BYDV reaction in breeding programmes may prove difficult because plants with the highest performance tended to suffer most from BYDV infection, and half-sib progeny ranked in a different order of tolerance in the glasshouse from that of their respective parent genotypes in the field. It is suggested that the observed variation in BYDV response results from a readjustment in the balance of genotypically controlled and environmentally conditioned variation in plant growth.     

Multiple components of the resistance of potatoes to potato leafroll virus

In glasshouse experiments the ranking of potato genotypes for resistance to infection with potato leafroll virus (PLRV) using three concentrations of aphid-borne inoculum was the same as their field resistance ratings. In field-grown plants this resistance to infection increased in all genotypes as the plants aged but its rate of increase differed between genotypes. In tests on field-grown plants infected by aphid- or graft-inoculation, the proportion of virus-free progeny tubers increased the later the date of inoculation but was greater in resistant than in susceptible genotypes. This trend was most pronounced in the resistant clone G7445(1), in which the virus failed to move from the foliage to the tubers of some plants infected in glasshouse tests. The spread of PLRV will thus be minimised in crops of resistant compared with susceptible genotypes for three reasons: plants have greater resistance to infection, systemic spread of virus from their foliage to tubers is less likely and, as shown previously, the low concentration of virus particles in leaf tissue makes infected plants less potent sources of inoculum for aphids.