THE INFECTION OF PLANTS BY VIRUSES THROUGH ROOTS

Roots of young tomato plants became infected when inoculated with tomato bushy stunt, tobacco mosaic, and potato X viruses. Root infections also occurred when these viruses were added to soil or culture solutions in which plants were growing.
The viruses were sometimes localized around their initial entry points in roots; sometimes they invaded the root system but not the shoots, and sometimes they produced full systemic infection of roots and shoots. In some experiments, but not all, systemic infections were more frequent when the upper tap root or superficial roots were inoculated than when fibrous roots were inoculated.
In both tomato and potato, virus X spread from diseased to healthy plants sharing the same culture solution, if their roots were in contact, but not otherwise. Infection of the roots of potato plants by inoculation, produced only one plant with virus-infected haulms, although several had infected tubers.     

Types of change in potato virus X infections

In tobacco plants infected with mild strains of virus X , severe strains may arise as mutations which multiply locally.
Several strains of virus X gradually lost infectivity for potato on continued culture in other hosts such as tobacco.
Strains that are poorly infective or invasive for potato may take considerable time to move into the growing shoot from the tuber in detectable amounts.
Pronounced and apparently spontaneous increases in the severity of the strain that dominates in potatoes may occur in field crops.     

The relationships of some viruses causing necrotic diseases of the potato

Potato virus B , and some other viruses with reactions in potato varieties different from any previously described, are strains of virus X . All produce intracellular inclusions which vary with different hosts and virus strains. Except with virus B, the inclusions are larger and more frequent in potato than in tobacco or tomato. All give systemic infection when inoculated to tobacco, tomato and potato varieties in which they are carried or cause mosaic symptoms; some give systemic infection when inoculated to varieties in which they cause top-necrosis, whereas others give only local lesions.
Potato virus C is a strain of virus Y: in tobacco and a few potato varieties both produce similar symptoms, but in those varieties in which Y causes leaf-drop streak, C causes top-necrosis. C causes systemic infection when inoculated to tobacco and to potato varieties in which it causes mosaic symptoms, but not when inoculated to potato varieties in which it causes top-necrosis. Virus C was not transmitted by M . persicae. Viruses C and Y produce a few small intracellular inclusions in potato and tobacco.
Virus A is not related to Y or X : no inclusions were found in plants infected with A alone.     

SOME FACTORS AFFECTING THE TRANSMISSION OF LEAF-ROLL VIRUS BY APHIDS

Datura tatula is a more suitable host than potato for studying the factors influencing the transmission of potato leaf-roll virus by Myzus persicae ; it is more easily infected, provides a better source of virus for feeding aphids, produces symptoms more quickly and over a longer period of the year.
Loughnane's (1943) claim that leaf-roll virus is transmitted by starved aphids that feed for only 5 min. on infected potato plants was not confirmed. The shortest infection-feeding time in which M. persicae aphids became infective was 2 hr.; such aphids did not infect healthy plants in the first 2 days and, when transferred to a series of healthy plants at intervals, infected only few. The ability to cause infections was increased by increasing the length of infection feeding. Aphids fed for many days on infected plants could infect healthy plants in the first 15 min. of test feeding, and they continued to cause infections for long periods.
Aphids became infective more readily when feeding on recently infected Datura tatula , showing only slight symptoms, than on older plants with pronounced chlorosis; similarly, young potato sprouts showing no symptoms were better sources of virus for aphids than older plants showing severe leaf roll.
The differences in severity of symptoms shown by potato plants with leaf roll in the field mainly occur because of differences in virulence of accompanying strains of potato virus X , but isolates of leaf-roll virus were found that also varied in virulence.     

HEAT-THERAPY OF VIRUS-INFECTED PLANTS

Virus-free plants were produced from parents systemically infected with the following five viruses: tomato bushy stunt, carnation ring spot, cucumber mosaic, tomato aspermy and Abutilon variegation. The leaves formed while the infected plants were kept at 36°C. were free from symptoms, and test plants inoculated from these remained uninfected. When cuttings were taken from the infected plants at the end of the treatment most grew into healthy plants. The treated plants themselves usually developed symptoms after varying lengths of time at 20°C, but some that before treatment were infected with tomato aspermy, cucumber mosaic or Abutilon variegation viruses, remained permanently healthy.
The same method failed to cure plants infected with tomato spotted wilt, potato virus X and tobacco mosaic virus, although it decreased their virus content. Heat-therapy seems not to be correlated with the thermal inactivation end point of the virus in vitro.     

Study on potato virus M (PVM) occurrence in potato fields in Iran

57 native potato tuber samples collected from different potato growing region of Iran, planted on single rows in Karaj College experimental station. Plant samples of each single row plus 9.25 Fresh foliage samples collected from fields under new introduced cultivars were tested for potato virus (PVM) infection during growing season. Also 78 weeds and field crops belonging to Solonacae and Leguminosae from or neighboring to potato field were tested. Results indicated that PVM was not found on any plant other than potatoes. PVM was detected on 16 samples of 57 old vars, Virus was not seen in any samples collected from fields under new varieties. Results show that PVM is limiting in this crop. PVM detecting is difficult using assay hosts. Best test plants were French bean var Red kidney, Showing pinpoint necrotic LL, also Datura metel and Nicotiana debneyi are useful for virus detection showing chlorotic local lesion. Also microprecipition and gel diffusion test can be used for virus detection but Elisa was the best method. PVM infected plant showed 11-19.5 percent yield decrease in 3 cultivars tested.     

Evaluation of hairy nightshade as an inoculum source for aphid-mediated transmission of potato leafroll virus

Potato leafroll virus (PLRV) causes one of the most serious aphid-transmitted diseases affecting yield and quality of potatoes, Solanum tuberosum (L.), grown in the United States. The green peach aphid, Myzus persicae (Sulzer), is considered to be by far the most efficient vector of this virus. Even the most strict aphid control strategy may not prevent the spread of PLRV unless measures also are taken to keep virus source plants within and outside the crop at a minimum. Hairy nightshade, Solanum sarrachoides (Sendtner), is one of the preferred weed hosts for green peach aphid. The potential of this weed as an aphid reservoir and virus source and its spread or perpetuation were investigated. With the use of double antibody sandwich enzyme-linked immunosorbent assay, it was confirmed that green peach aphid can transmit PLRV to hairy nightshade and that aphids can become viruliferous after feeding on infected hairy nightshade plants. Transmission from hairy nightshade to potato is 4 times the rate of potato to potato or potato to hairy nightshade. The green peach aphid preferred hairy nightshade over potato plants and reproduced at a higher rate on hairy nightshade than on potato. Therefore, a low level of PLRV-hairy nightshade infection could enhance the disease spread in the field.     

Studies on the spread of certain plant viruses in the field

Studies on the spread of potato viruses X and Y and cucumber mosaic virus in the field are described: tobacco was used as the experimental plant. The plants were set out in the form of a cross, one series with the leaves in contact and one with the leaves not touching. No spread of potato virus X was observed, but there was extremely rapid permeation of virus Y throughout the plots. The spread of cucumber mosaic virus was much slower than that of virus Y.     

Volatiles from potato plants infected with potato leafroll virus attract and arrest the virus vector, Myzus persicae (Homoptera: Aphididae)

The influence of viral disease symptoms on the behaviour of virus vectors has implications for disease epidemiology. Here we show that previously reported preferential colonization of potatoes infected by potato leafroll virus (genus Polerovirus) (luteovirus) (PLRV) by alatae of Myzus persicae, the principal aphid vector of PLRV, is influenced by volatile emissions from PLRV-infected plants. First, in our bioassays both differential immigration and emigration were involved in preferential colonization by aphids of PLRV-infected plants. Second, M. persicae apterae aggregated preferentially, on screening above leaflets of PLRV-infected potatoes as compared with leaflets from uninfected plants, or from plants infected with potato virus X (PVX) or potato virus Y (PVY). Third, the aphids aggregated preferentially on screening over leaflet models treated with volatiles collected from PLRV-infected plants as compared with those collected from uninfected plants. The specific cues eliciting the aphid responses were not determined, but differences between headspace volatiles of infected and uninfected plants suggest possible ones.     

The appearance and spread of mosaic infection in the tomato crop and the relation to seed transmission of the virus

Appearance and spread of infection with mosaic-inducing viruses were studied for three seasons in tomato crops under glass. Comparison was made between the reactions of plants raised from virus-free seed and those of plants raised from virus-infected seed, on plots distributed at random in a house in which no precautions against entry and spread of virus were taken. Freedom from mosaic infection was maintained longest in plants raised from virus-free seed. An experiment was carried out after steam sterilization of the soil and under exceptionally favourable weather conditions. Appearance of mosaic symptoms occurred later in the life of the plants in this season and plants raised from virus-free seed did not react differently from other plants.
The location of plants first showing mosaic symptoms was related to the depth and texture of soil beneath those plants.
Tests were made of the apparent virus content of infected tomato seed during germination and differences were found in the persistence of virus during germination in seeds of differing origin.
Apparent, 'delayed' seed transmission of mosaic-inducing viruses occurs in the tomato crop, but as yet, this condition can only be interpreted in terms of differences in the resistance of plants raised from seed of differing origin to the multiplication and systemic spread of those viruses. The use of virus-free seed taken from well-nourished vigorous plants is essential to the production of a virus-free tomato crop under commercial conditions.     

ROGUING POTATO CROPS FOR VIRUS DISEASES

Removing virus-infected plants from plots of Majestic potatoes at Rothamsted on 2 July 1947 did not reduce the spread of leaf roll but reduced rugose mosaic (potato virus Y) to about one-fifth of that in plots rogued on 21 July or left unrogued. Roguing Arran Pilot potatoes on 16 June or 2 July reduced leaf roll to five-sixths of that in unrogued plots; roguing on 16 June reduced rugose mosaic to about half that in plots rogued on 2 July, and about a quarter of that in unrogued plots. Lifting Arran Pilot potatoes in mid-August reduced virus diseases to about two-thirds.
Roguing flattened the gradient (decrease in percentage plants diseased with increasing distance from the source of infection) with rugose mosaic, but had little effect with leaf roll. Evidently any plants prevented by roguing from contracting virus Y were near the initially infected plants.
In 1948, Majestic and King Edward potatoes at three places were rogued during 22–24 June and tubers were dug during 28–30 July and again at the end of the season. Leaf roll spread more in Majestic than in King Edward, and rugose mosaic spread more in King Edward. Roguing reduced the spread of both by about one-fifth at Rothamsted, but had no effect at Sutton Bonington. At Bretton, in the Derbyshire hills, roguing had no effect on leaf roll, but prevented the spread of rugose mosaic.
The small benefit occasionally achieved by roguing in the ware-growing districts of England does not make the practice economically worth while.     

Infection of potato mesophyll protoplasts with five plant viruses

Methods are described for preparing potato mesophyll protoplasts that are suitable for infection with inocula of virus nucleoprotein or RNA. The protoplasts could be infected with four sap-transmissible viruses (tobacco mosaic, tobacco rattle, tobacco ringspot and tomato black ring viruses) and with potato leafroll virus, which is not saptransmissible. No differences were observed in ability to infect protoplasts with potato leafroll virus strains differing either in virulence in intact plants or in aphid transmissibility.     

EXPERIMENTS ON THE COLONIZATION OF POTATO PLANTS BY APTEROUS AND BY ALATE APHIDS IN RELATION TO THE SPREAD OF VIRUS DISEASES

Batches of potato plants in pots were placed in the field for limited periods among plants infected with potato virus Y and leaf roll virus. Some of the potted plants were surrounded by sticky boards which prevented apterous aphids from reaching them. Almost as many plants within the boards as without became infected, indicating that most of the spread of virus was by winged aphids.
Apterae were probably responsible for spreading the viruses throughout a hill after one or more stems were infected. They may carry infection to neighbouring plants, but most of these will have been infected already by alatae.
The number of plants contracting infection was unaffected by watering.     

The epidemiology of tomato mosaic: Sources of TMV in commercial tomato crops under glass

Of the several possible sources of tomato mosaic virus, seeds and root debris in the soil are considered to be of greatest importance. A survey of 374,000 seedlings on ten commercial holdings found 0.05% of them infected, and although these were removed virus had been spread to other young plants which did not show infection when transplanted into the growing houses, seven of twenty-two of which contained a few infected plants when sampled shortly after planting. Virus overwintering on clothing, and debris on structures, are thought to be of minor importance, and smoking tobacco is seldom a source of infection for the tomato crop. A further survey of seventy-eight samples from tomato crops in Britain confirmed the 1960-61 survey: all were infected with tomato strains of TMV, none with tobacco strains, but one of the 187 infected seedlings referred to above was carrying a tobacco strain. Petunia was not as satisfactory as a special cultivar of White Burley tobacco for distinguishing between the tobacco and tomato TMV isolates. Observations and tests on a commercial holding showed that TMV was readily carried from plants in infected glasshouses into clean ones by workers, and once introduced, spread rapidly within the crop.     

Application of enzyme-linked immunosorbent assay to the detection of potato leafroll virus in potato tubers

Factors affecting the detection of potato leafroll virus (PLRV) by enzyme-linked immunosorbent assay (ELISA) in tubers of field-grown potato plants with primary or secondary infection were studied. The reactions of extracts of virus-free potato tubers were minimised by pre-incubating the extracts at room temperature and by careful choice of the dilution of enzyme-conjugated globulin. PLRV was reliably detected in tubers produced by secondarily infected plants of all six cultivars tested. PLRV concentration was greater in heel-end than in rose-end vascular tissue of recently harvested tubers but increased in rose-end tissue when tubers stored at 4°C for at least 5 months were placed at 15–24°C for 2 wk. PLRV occurred at greater concentration in tubers from plants of cv. Maris Piper with natural or experimentally induced primary infection than in tubers from secondarily infected plants; again PLRV concentration was greater in heel-end than in rose-end vascular tissue. Plants whose shoots were infected earliest in the growing season were invaded systemically and produced the greatest proportion of infected tubers; plants infected late in the season also produced infected tubers but PLRV was not detected in their shoot tops. PLRV concentration in tubers from the earliest-infected plants was less than in tubers from later-infected plants. PLRV was detected reliably by ELISA in tubers from progenies that were totally infected but was not detected in all infected tubers from partially infected progenies. ELISA is suitable as a routine method of indexing tubers for PLRV, although the virus will not be detected in all infected tubers produced by plants to which it is transmitted late in the growing season.     

THE SPREAD OF VIRUS DISEASES TO SINGLE POTATO PLANTS BY WINGED APHIDS

Young potato plants in pots exposed in the open near plots of potatoes for limited periods at intervals during the summer, became infested with large numbers of winged aphids only during warm, calm and dry weather. Although visited by aphids during May and June, when much of the spread of viruses occurred in nearby potato crops, few of the potted plants became infected. Most potted plants became infected in July when alate aphids were leaving neighbouring potato crops. Widely different proportions of the exposed plants became infected in different years; in two of the three years, many more plants were infected with virus Y than with leaf roll virus.     

Tests for transmission of four potato viruses through potato true seed

The Andean potato calico strain of tobacco ringspot virus (TRSV-Ca) was detected in 2–9% of potato seedlings grown from true seed from plants of cv. Cara and clone G5998(6) infected with TRSV-Ca. Similarly, a potato isolate of the oca strain of arracacha virus B (AVB-O) was detected in 4–12% of progeny seedlings of cv. Cara and clone D42/8 infected with AVB-O. Potato virus T (PVT) passed through 33–59% of seed from PVT-infected cv. Cara, but only 0–2% infection was detected in seedlings from seed of PVT-infected clone D42/8. By contrast, no infection was detected in seedlings grown from seed from plants of G5998(6), D42/8 or cv. Cara infected with Andean potato latent virus strains Hu (APLV-Hu) or Caj (APLV-Caj), although both strains passed through seed of Nicotiana clevelandii. AVB-O, PVT and TRSV-Ca were detected in all tests of pollen from flowers of infected potato plants, but APLV-Hu and APLV-Caj were detected less frequently. AVB-O and PVT were transmitted through 2% and 8% respectively, of seed from healthy potato plants pollinated with pollen from infected plants. However, no transmission through seed was detected when pollen from TRSV-Ca infected plants was used. None of the four viruses were transmitted to healthy potato plants pollinated with pollen from infected plants. APLV-Hu caused exceptionally severe symptoms in the cv. Cara plants used for seed production, but the Bolivian strain of PVT induced only mild symptoms rather than the severe systemic necrosis previously reported for the type of strain of PVT in this cultivar. No symptoms developed in potato seedlings infected with TRSV-Ca, AVB-O or PVT through the seed.     

FACTORS AFFECTING THE SPREAD OF APHID-BORNE VIRUSES IN POTATO IN EASTERN SCOTLAND

Experiments at Invergowrie, south-east Perthshire, showed that the extent of spread of potato leaf-roll and Y viruses varied from year to year and that virus Y consistently spread more than leaf roll. Most spread of Virus Y occurred before the end of June and of leaf-roll virus before the end of July. Both viruses spread slightly more in late- than in early-planted crops. When plants with leaf roll and already colonized by Myzus persicae were placed in a healthy crop of Majestic potatoes at intervals during the season, the amount of virus spread decreased rapidly with increasing age of the crop. Spread of leaf roll occurred in all of twenty-five 'seed' crops in different districts of eastern Scotland in 1955 but in only twenty out of thirty-six similar crops in 1956. Annual and regional differences in virus spread appear to reflect differences in the time at which migrant aphids reach potato crops in early summer and the rate at which infestation builds up in the crops.     

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.     

TMV-Infection syndromes in potato cultivars Erika and Krasava

Tre potato cultivars Erika and Krasava are unable to reproduce two strains of TMV systemically except when in graft symbiosis under normal greenhouse conditions with tomato which has been infected with the virus. The escape resistance of potato to TMV infection due to slight hypersensitivity was not changed either by long-lasting non-infectious symbiosis with TMV-sensitive tomato or in a constant environment (32±2 °C, 9 700 lx) which was adequate to change the hypersensitive reaction of some other TMV-hosts. On the contrary, enhanced temperature provoked a still more severe local and necrotic response in the potato varieties. This led to systemic necrosis and the quick death of potato components systemically infected in graft symbiosis with tomato under normal greenhouse conditions when transferred into an environment of enhanced temperature.