Some effects of spraying with thiabendazole on the susceptibility of sugar beet to yellowing viruses and on their vector, Myzus persicae (Sulz.)

In a field experiment fewer sugar-beet plants became infected with aphid-transmitted yellowing viruses in plots that had been sprayed with solutions of thiabendazole lactate than in water-sprayed plots, after exposure to natural infestation with aphids. Subsequent glasshouse tests showed that foliar sprays of o·o1 % thiabendazole lactate in water significantly reduced the proportion of inoculated sugar-beet plants which became infected with beet yellows virus (BYV) or beet mild yellowing virus (BMYV) after inoculation with viruliferous Myzus persicae (Sulz.). This effect on virus transmission was not apparently due to a direct insecticidal action of thiabendazole, because adult aphids usually survived equally well on sprayed and unsprayed plants. Treatment of test plants with thiabendazole did not affect the transmission of beet mosaic virus to them by M. persicae. The fecundity of M. persicae was greatly reduced by transferring them to plants which had been sprayed with thiabendazole or by spraying them with thiabendazole before transfer to unsprayed plants. The fertility of adult Aphis fabae Scop, was also reduced by spraying with thiabendazole. The mechanisms whereby thiabendazole affected fecundity of aphids and transmission of viruses are not understood.     

Studies on beet western yellows virus in oilseed rape (Brassica napus ssp. oleifera) and sugar beet (Beta vulgaris)

Oilseed rape (Brassica napus L. ssp. oleifera) was studied as a potential overwintering host for the sugar-beet yellowing viruses, beet yellows virus (BYV) and beet mild yellowing virus (BMYV), and their principal vector, Myzus persicae. In spring 1982, plants infected with a virus which reacted positively in enzyme-linked immunosorbent assay (ELISA) with BMYV antibody globulin were found in oilseed-rape crops; none of the plants contained virus which reacted with BYV antibody globulin. This virus was subsequently identified as beet western yellows virus (BWYV). No leaf symptoms could be consistently associated with infection of oilseed rape, but the virus was reliably detected by sampling any leaf on an infected oilseed-rape plant. Some isolates from oilseed rape did infect sugar beet in glasshouse tests, but the proportions of inoculated plants which became infected were low. Apparently there is therefore little danger of much direct transmission of BWYV by M. persicae from oilseed rape to sugar beet in spring. BWYV was introduced to and spread within oilseed-rape crops in autumn by M. persicae, and autumn-sown oilseed rape proved to be a potentially important overwintering host for M. persicae. In a survey of 80 autumn-sown crops of oilseed rape in East Anglia, northern England and Scotland in spring 1983, 78 were shown to be extensively infected with BWYV. Experimental plots of oilseed rape with 100% BWYV-infection yielded approximately 13.4% less oil than plots with 18% virus infection, the result of a decrease in both seed yield and oil content.     

Effects of sucrose sprays and darkness on aphid colonization of sugar beet and on aphid transmission of yellowing viruses

In the glasshouse, adult, apterous Myzus persicae (Sulz.) and Aphis fabae Scop, settled better and deposited more larvae on sucrose-sprayed sugar-beet plants than on water-sprayed plants. M. persicae settled badly and deposited few larvae on plants that were kept in the dark before or after infestation. The effects of darkness on aphids were reduced by spraying the host plants with 10% solutions of sucrose before infestation. Viruliferous M. persicae transmitted beet yellows virus (BYV) and beet mild yellowing virus (BMYV) less efficiently to dark-treated plants than to those grown in normal daylight. Spraying sugar beet with sucrose before inoculation with viruliferous M. persicae increased the proportion of successful BYV transmissions but only when the plants were dark-treated. The effects of sucrose and darkness on settling and larviposition of aphids and on virus transmission may be related to changes in the concentration of carbohydrates, particularly sugars, in the leaves.     

Measuring and modelling the effects of inoculation date and aphid flights on the secondary spread of Beet mosaic virus in sugar beet

The effect of the inoculation date on the spread of Beet mosaic virus (BtMV) in sugar beet field plots was studied. Two plants in the centre of each plot were inoculated with BtMV using Myzus persicae. The spread of the infection around these sources was monitored by inspecting the plants on two diagonal transects through the centre of the plot. Early inoculations resulted in a greater spread than late inoculations, but any inoculation before the onset of the aphid migration resulted in a similar‐sized spread. The spread was concentrated in patches around the inoculated plants, and its rate was explained by vector pressure, as shown by regression analysis and a mechanistic simulation model. This vector pressure was quantified using data obtained by catching aphids in a green water trap in the crop, catching aphids in a 12 m high suction trap at a distant location, and infection of bait plants from adjacent virus source plants. The daily total aphid catches obtained by a suction trap provided the best statistical explanation for the spread of this virus. The parameter r, describing the relationship between vector pressure and the rate of disease progress, was remarkably robust. This parameter varied less than 10% between treatments (infection date) within a single experiment, and less than a factor two between four experiments performed at different sites in two years. The robustness of this parameter suggests that the spread of a potyvirus may be predicted on the basis of the initial infection date and vector abundance.     

Aphid transmission of Beet Yellows Virus affected by homologous antiserum

A thin layer of homologous antiserum (against the beet yellows virus - BYV) between the leaf surface and a Parafilm membrane totally inhibited the acquisition of BYV by aphidsMyzus persicae (Sulz.), but it did not affect the inoculation of BYV by infective aphids. BYV transmission decreased with aphids picking up the virus from leaves coated with a normal rabbit serum. Aphids sucking on purified BYV suspension through the Parafilm membrane as well as aphids allowed to probe into leaves of healthy plants spread with an infectious purified BYV suspension failed to transmit BYV. No BYV particles could be detected in eluates from stylets and labia cut off from aphids which had probed on BYV infected plants by electron microscopic examination. The acquisition seems to be the most important phase for the aphid transmission of BYV which is apparently carried on the stylet surface.     

Purification and Some Properties of an Isolate of Beet Yellows Virus from Ukraine

A purification procedure, which yielded up to 15–30 mg of beet yellows virus (BYV) per 100 g of infected Tetragonia expansa leaves, has been developed. The procedure included sap clarification with Triton X-100, and two cycles of ultracentrifugation through sucrose cushion, which contained PEG-6000 and NaCl. A specific antiserum was prepared, and BYV infection was successfully detected by the double-antibody sandwich (DAS) ELISA in infected sugar beet leaves and roots diluted up to 1 × 10⁵ and 1 × 10⁴, respectively. The virus concentration was demonstrated to decrease in infected sugar beet roots slowly during 7 months, thus allowing successful diagnosis of planting material in winter storage. BYV presence in Myzus persicae aphids was also reliably detectable using the DAS-ELISA. In a competitive DAS-ELISA test, the Ukraine and the British BYV isolates were found serologically indistinguishable.     

The effect of plant age and infection with virus yellows on the survival of Myzus persicae on sugar beet

Survival of Myzus persicae confined in clip-cages on mature leaves of sugar beet declined as the plants aged. Death of aphids was often preceded by the appearance of a black deposit in the aphids' stomachs, which may have been the cause of death. Both the rate of death and the proportion of aphids dying with black deposits was significantly less when plants were infected with beet yellows virus or beet mild yellowing virus, by comparison with healthy plants. The implication of these phenomena on the onset of mature plant resistance is discussed.     

Effects of cereal borders, admixture with cereals and plant density on the spread of bean yellow mosaic potyvirus into narrow-leafed lupins (Lupinus angustifolius)

In studies of virus control measures, field experiments in 1987–1991 investigated the effects of cereal and fallow borders, admixture with cereals and plant density on spread of bean yellow mosaic potyvirus (BYMV) from pastures dominated by subterranean clover (Trifolium subterraneum) into plots of narrow-leafed lupins (Lupinus angustifolius). Virus spread was mainly monocyclic because BYMV killed infected lupin plants and between systemic movement and death there was only a brief period for BYMV acquisition and transmission to other plants by vector aphids. In plots with cereal borders, the rate and extent of BYMV spread into the lupins was decreased; at final assessment the numbers of infected plants were 43–60% less than in plots with fallow borders. Admixture with cereals also decreased the rate and extent of BYMV spread into lupin plots, numbers of infected plants being decreased by 76–96% at the time of final assessment. When lupins were sown at different seeding rates to generate a range of plant densities and weeds were removed, high densities decreased BYMV infection. The higher incidences of BYMV infection in sparse stands were attributed partly to smaller plant numbers and partly to incoming viruliferous vector aphids being more attracted to plants with bare earth around them, than to a plant canopy. BYMV infection decreased grain yield of samples from infected lupin plants by 94–100%. In plots with 34% infection and sparse stands, grain yield was decreased by about one third. Plotted progress curves for the accumulated numbers of alate aphids of the BYMV vector species Acyrthosiphon kondoi and Myzus persicae resembled those for numbers of BYMV infected plants in 1990, but in 1991 only the curve plotted for M. persicae did so. There was a 2 week delay between the curves for aphid numbers and virus counts which reflected the time taken for obvious systemic necrotic symptoms to develop in lupins.     

Breeding for resistance to infection with yellowing viruses in sugar beet

Differences in resistance to infection with beet yellows virus (BYV) and beet mild yellowing virus (BMYV) have been observed in virus-tolerant sugar-beet breeding material. The results of glasshouse virus-susceptibility tests usually agreed well with those of field experiments in which plants were exposed to artificial, or natural, infestation with viruliferous aphids. Breeding lines and varieties, which showed resistance to BYV when Myzus persicae Sulz, was used as vector, generally showed a similar resistance to this virus when Aphis fabae Scop. was used. Varieties which were resistant to infection with one virus were not necessarily resistant to the other, although some showed resistance to both BYV and BMYV. Preliminary results suggest that resistance to infection may be controlled by recessive genes which occur widely in sugar-beet cultivars. The mechanism of this form of resistance is not understood, but it does not appear to be closely associated with resistance to the aphid vectors of the viruses. The observed differences in resistance to infection demonstrate the possibility of breeding a sugar-beet variety in which two forms of resistance to virus yellows, tolerance and resistance to infection, are combined.     

A controlled environment study of the effect of leaf physiological age on the movement of apterous Myzus persicae on sugar-beet plants

Apterous Myzus persicae were found to move frequently from leaf to leaf on sugar-beet plants in controlled environment conditions. It is suggested that aphid movement can be related to changes in the rate and content of translocate flow during leaf development. These changes make newly-emerged leaves nutritionally favourable to colonising aphids and make expanding leaves slowly wane in favourability during the process of ‘sink to source’ conversion leading to aphid dispersal from the leaf. Variation in temperature was not found to alter the rate of aphid movement or the period (measured in thermal time) that aphids spent on particular leaves. However, the lower temperature was found to increase the rate of aphid development, aphid size and fecundity; these effects could also be due to nutritional factors. This dispersal behaviour may be a tactic to maximise food intake by a polyphagous aphid and increase the probability that nymphs are deposited on nutritionally-favourable leaves. The implications of the interleaf dispersal of apterous M. persicae for within- and between-plant spread of beet yellows virus (BYV) and beet mild yellowing virus (BMYV) are discussed.     

Insecticidal control of aphids and the spread of potato leafroll virus in potato crops in eastern Scotland

Field experiments were carried out in eastern Scotland in 1976-78 to test the ability of granular insecticides, applied to soil at planting, and of insecticide sprays applied to the foliage, to control aphids and spread of potato leafroll virus (PLRV) in potatoes. The three years provided contrasting opportunities for virus spread. In 1976, the main vector of PLRV, Myzus persicae, arrived in early June and multiplied rapidly in untreated plots, and PLRV spread extensively. In 1977, M. persicae arrived 4–6 wk later than in 1976 and most spread of PLRV, which was less than in 1976, occurred after the end of July. In 1978, few M. persicae were recorded but the potato aphid, Macrosiphum euphorbiae, arrived early and very large populations developed in untreated plots. However, little spread of PLRV occurred in 1978, supporting other evidence that M. euphorbiae is an inefficient vector of PLRV in field conditions. In each year, granular insecticides decreased PLRV spread to a quarter or less of that in control plots. Thiofanox gave somewhat better and longer-lasting control of aphid populations than disulfoton, especially of M. persicae, but did not give greater control of PLRV spread. Application of three (1976) or five (1977) sprays of demeton-S-methyl to plots treated with granular insecticides further improved the control of M. euphorbiae but had less or no effect on M. persicae, especially where organophosphorus resistant aphids (R1 strain) were found. These supplementary sprays of insecticide did not further improve the control of PLRV but, in 1978, four sprays of demephion or pirimicarb to plots not treated with granular insecticide decreased PLRV spread. These data, together with previous findings, indicate that the amount of virus spread depends on the date of arrival and rate of multiplication of M. persicae in relation to the timing and effectiveness of removal of PLRV sources in crops. It is concluded that in Scotland insecticide granules should be used routinely only in crops of the highest grade of seed potato. Their use for other grades need be considered only in years following mild winters, when aphids can be expected to enter crops earlier and in larger numbers.     

Sublethal effects of aldicarb on the behaviour of Aphis fabae and two clones of Myzus persicae and on the transmission of beet mosaic virus by these aphids

In laboratory experiments treatment of sugar-beet plants with aldicarb stimulated the mobility of Aphis fabae and two clones of Myzus persicae which were susceptible (S) and resistant (R) to carbamate-based insecticides, respectively. On the other hand, the number of aphids probing and the total number of probes made was reduced, and hence the transmission of beet mosaic virus (BMV) was restricted. In outdoor experiments the spread of BMV from aldicarb-treated plants by naturally infesting aphids was also restricted. The number of infected plants decreased with increasing distance from the sources of infection.

---

Résumé Des plantes de betterave traitées au laboratoire avec de l'aldicarbe ont stimulé la mobilité d'Aphis fabae et de deux clones de Myzus persicae, l'un sensible et l'autre résistant à des insecticides contenant des carbamates. Par ailleurs, le nombre de pucerons en train de sonder les feuilles ainsi que le nombre total de sondages ont été réduits et ainsi la transmission du virus de la mosaïque de la betterave (BMV) a été limitée. Dans des expériences à l'extérieur, la vitesse de propagation de BMV par des pucerons sur des plantes traitées à l'aldicarbe a été aussi plus limitée. Le nombre de plantes contaminées diminuait avec la distance de la source de contamination.

     

The effects of yellowing viruses on yield of sugar beet in field trials, 1985 and 1987

The separate effects of beet yellows virus (BYV) and beet mild yellowing virus (BMYV) on yield of sugar-beet cultivars inoculated at different growth stages were assessed in field trials in 1985 and 1987. Early or mid-season inoculation decreased sugar yield by up to 47% for BYV, and up to 29% for BMYV. Infections after the end of July had no significant effect on yield. Both viruses caused significant increases in the juice impurities sodium, potassium and amino-nitrogen after infecting plants early in the season. Yield losses associated with infection were determined by the causative virus, the time of infection, and susceptibility of the sugar-beet cultivars.     

The use of monoclonal antibodies to detect beet mild yellowing virus and beet western yellows virus in aphids

Information on infectivity of the aphids which invade sugar beet root crops each Spring is required for forecasting incidence and providing advice on control of virus yellows. Monoclonal antibodies, produced in the USA to barley yellow dwarf virus (BYDV) and in Canada to beet western yellows virus (BWYV), were used to distinguish between sugar-beet-infecting strains of the luteovirus beet mild yellowing virus (BMYV), and the non-beet-infecting strains of the closely-related BWYV in plant and aphid tissue. Totals of 773 immigrant winged Myzuspersicae and 124 Macrosiphum euphorbiae were caught in water traps in a crop of sugar beet between 25 April and 5 August 1990. Using the monoclonal antibodies and an amplified ELISA, 67%M. persicae and 19%M. euphorbiae were shown to contain BWYV; 8%M. persicae and 7%M. euphorbiae contained BMYV. In studies with live winged aphids collected from the same sugar beet field during May, 25 of 60 M. persicae and two of 13 M. euphorbiae transmitted BWYV to the indicator host plant Montia perfoliata; two M. persicae and two M. euphorbiae transmitted BMYV. In another study three of 65 M. persicae and one of three M. euphorbiae in which only BWYV was detected, transmitted this virus to sugar beet.     

Early-season predation impacts the establishment of aphids and spread of beet yellows virus in sugar beet

The potential of predators to impact the establishment of aphid vectors and the spread of beet yellows virus in sugar beet was examined. Myzus persicae carrying beet yellows virus (BYV) were released on six interior sites and six edge sites in each of four fields at the end of May. Aphids established at low densities and BYV was spread in circular patches around the infested plants at all sites. The number of diseased plants per patch at the end of September ranged from a field-average of 130 to 210 in the four fields. There was a weak tendency towards better aphid establishment and greater virus spread in fields in less complex landscapes. Edge sites had less virus spread than interior sites in one field, more virus spread in two other fields, and there was no statistically significant difference in the fourth field. In the field where virus spread was lowest at edge sites, we used predator exclosure and direct observation to manipulate and quantify the effects of early season predation. On a warm day in early June, 81% ofAphis fabae exposed to predators on young beet plants disappeared during a 24 h period, compared to 10% of aphids protected by clipcages. Intermediate levels of predator exclusion, allowing aphids to walk away but restricting predator access, showed that predation was responsible for aphid disappearance.Cantharis lateralis L. (Coleoptera: Cantharidae) was the most frequently observed foliar predator (>90%). It was found eating aphids on several occasions. The incidence of predators was 1.8 per plant per h in the field interior and 3.8 per plant per h. near the edge. In the same field, aphids and virus were released in six edge and six interior sites, that were surrounded by 0.5 m high plastic open-top barriers (‘exclosures’). Pitfall trapping inside the barriers reduced potential soil predator densities to ca. one-tenth of the open field level and arrivals of flying predators were reduced. Inside the exclosures, aphid establishment was enhanced, and virus spread at exclosure sites was increased by about 50% compared to open sites. Foliar and pitfall sampling yielded the following predators:Cantharis lateralis, C. rufa L. (Coleoptera: Cantharidae),Coccinella septempunctata L.,C. undecimpunctata L. (Coleoptera: Coccinellidae),Pterostichus cupreus (L.),Harpalus rufipes (de Geer),Patrobus atrorufus (Strom),Trechus quadristriatus (Schrk.),Bembidion lampros (Herbst) (Coleoptera: Carabidae). In a laboratory no-choice trial (with 10M. persicae /day offered), each of these species ate aphids with consumption rates varying from 1.7 to 9.2 aphids/day. The results show that early predation substantially impacted aphid establishment in one field, and resulted in reduced virus spread. Results in the other fields show that these results cannot be easily generalized.     

Distribution and incidence of yellowing viruses in sugar beet crops in Spain from 1990 to 1993

Surveys of the principal yellowing viruses of sugar beet, beet yellows virus (BYV) and beet mild yellowing virus (BMYV) in Spain were carried out from 1990–1993. Beet yellowing viruses were detected in all provinces, although the mean percentages of plants infected with BYV and BMYV were practically zero in the southern zone. Within the northern zone high variations from one province to another could be observed. The mean percentages of plants infected with BYV were higher in the Ebro Valley than in the Duero Valley. Areas infected with BYV were very restricted, while BMYV could be found to a variable extent all over Spain, although the infection levels were lower. The incidence and distribution of these viruses in the Spanish sugar beet crop makes the study and application of control measures for beet yellowing viruses necessary.     

Efficient dsRNA-mediated transgenic resistance to Beet necrotic yellow vein virus in sugar beets is not affected by other soilborne and aphid-transmitted viruses

Rhizomania caused by Beet necrotic yellow vein virus (BNYVV) is one of the most devastating sugar beet diseases. Sugar beet plants engineered to express a 0.4 kb inverted repeat construct based on the BNYVV replicase gene accumulated the transgene mRNA to similar levels in leaves and roots, whereas accumulation of the transgene-homologous siRNA was more pronounced in roots. The roots expressed high levels of resistance to BNYVV transmitted by the vector, Polymyxa betae. Resistance to BNYVV was not decreased following co-infection of the plants with Beet soil borne virus and Beet virus Q that share the same vector with BNYVV. Similarly, co-infection with the aphid-transmitted Beet mild yellowing virus, Beet yellows virus (BYV), or with all of the aforementioned viruses did not affect the resistance to BNYVV, while they accumulated in roots. These viruses are common in most of the sugar beet growing areas in Europe and world wide. However, there was a competitive interaction between BYV and BMYV in sugar beet leaves, as infection with BYV decreased the titres of BMYV. Other interactions between the viruses studied were not observed. The results suggest that the engineered resistance to BNYVV expressed in the sugar beets of this study is efficient in roots and not readily compromised following infection of the plants with heterologous viruses.     

Infection of potato plants with potato leafroll virus changes attraction and feeding behaviour of Myzus persicae

Potato leafroll virus (PLRV; genus Polerovirus, family Luteoviridae) is a persistently transmitted circulative virus that depends on aphids for spreading. The primary vector of PLRV is the aphid Myzus persicae (Sulzer) (Homoptera: Aphididae). Solanum tuberosum L. potato cv. Kardal (Solanaceae) has a certain degree of resistance to M. persicae: young leaves seem to be resistant, whereas senescent leaves are susceptible. In this study, we investigated whether PLRV‐infection of potato plants affected aphid behaviour. We found that M. persicae's ability to differentiate headspace volatiles emitted from PLRV‐infected and non‐infected potato plants depends on the age of the leaf. In young apical leaves, no difference in aphid attraction was found between PLRV‐infected and non‐infected leaves. In fact, hardly any aphids were attracted. On the contrary, in mature leaves, headspace volatiles from virus infected leaves attracted the aphids. We also studied the effect of PLRV‐infection on probing and feeding behaviour (plant penetration) of M. persicae using the electrical penetration graph technique (DC system). Several differences were observed between plant penetration in PLRV‐infected and non‐infected plants, but only after infected plants showed visual symptoms of PLRV infection. The effects of PLRV‐infection in plants on the behaviour of M. persicae, the vector of the virus, and the implications of these effects on the transmission of the virus are thoroughly discussed.     

Inherited resistance of sugar beet to aphid colonization

Results of glasshouse experiments have confirmed that inbred lines of sugar beet differ in each of three types of resistance to Myzus persicae Sulz. and Aphis fabae Scop., namely: resistance to settling, resistance to multiplication, and tolerance. Resistance to multiplication was not invariably associated with resistance to settling, although plants of some lines showed both forms of resistance. Plants that were resistant to settling of alatae were not always resistant to apterae of the same species, and there was not a close relationship between resistance to M. persicae and to A. fabae. The mechanisms involved in resistance to aphids in sugar beet are not understood. Progenies of plants, selected for resistance to aphids from inbred lines, were often more resistant than progenies of unselected plants. Inheritance of each type of resistance is probably polygenic. The potential value of the different kinds of resistance, in reducing direct feeding damage and controlling the spread of virus yellows in the field, is discussed. The ultimate breeding objective is to produce commercial varieties in which appropriate kinds of resistance to aphids are combined with resistance to virus yellows. The use of such varieties would reduce the need to control aphids in the field by applications of chemicals.     

The control of Alternaria species on leaves of sugar beet infected with yellowing viruses

Leaves of virus-free sugar-beet plants rarely became infected with Alternaria spp. in two field experiments at Cambridge in 1965. Infection with beet yellows virus (BYV) increased susceptibility of plants to Alternaria only slightly but infection with beet mild yellowing virus (BMYV) increased it greatly. There was a close association between the severity of Alternaria symptoms, shown by different breeding lines and varieties of sugar beet, and the losses of sugar yield which they sustained after infection with BYV and BMYV. Many lines and varieties were resistant to Alternaria even when infected with BMYV and their resistance seemed to be inherited as a dominant character. Individual plants of any one line or variety differed greatly in resistance to Alternaria, suggesting that selection should improve the present level of resistance. Spraying the foliage of Alternaria-susceptible varieties with fungicides had little effect on the severity of Alternaria symptoms or on sugar yield. This was probably because the wet summer of 1965 was ideal for the spread of Alternaria and because rain washed the fungicide deposits from the sprayed leaves.