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.     

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.     

The effects of the repellents dodecanoic acid and polygodial on the acquisition of non-, semi- and persistent plant viruses by the aphid Myzus persicae

Few Myzus persicae settled on polygodial (1 g litre^-1) or dodecanoic acid (5 g litre^-1)-treated leaves and few nymphs were deposited. Polygodial decreased acquisition of the semi-persistent beet yellows virus and the non-persistent potato virus Y and dodecanoic acid that of the persistent potato leafroll and beet mild yellowing viruses, of beet yellows virus and of the bimodally-transmitted cauliflower mosaic virus. However dodecanoic acid increased the acquisition of potato virus Y.     

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.     

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.     

Determining the relative roles of different aphid species as vectors of cucumber mosaic and bean yellow mosaic viruses in lupins

Glasshouse and field studies were done to determine the relative roles of different colonising and non-colonising aphid species as vectors of two non-persistently transmitted viruses, cucumber mosaic cucumovirus (CMV) and bean yellow mosaic potyvirus (BYMV) in narrow-leafed lupin (Lupinus angustifolius) crops in Australia. The abilities of nine different aphid species in transmitting CMV from infected to healthy lupins and BYMV from infected subterranean clover to healthy lupins were compared in the glasshouse using 5–10 min acquisition access feeds. The percentage transmission efficiencies found with lupin-colonising aphid species were (CMV/BYMV): Acyrthosiphon kondoi (6/15), Aphis craccivora (10/14) and Myzus persicae (11/77). With non-colonising species the respective efficiencies were: Brachycaudus rumexicolens (0.9/0), Lipaphis erysimi (4/8), Rhopalosiphum maidis (9/6), R. padi (5/5), Sitobion miscanthi (2/11) and Therioaphis trifolii (4/5). When flying aphids were trapped in the field in four successive years (1993–1996) on vertical nets downwind of virus-infected lupins, 13 different species were caught at a “wheatbelt” site and 18 at an urban irrigated site. Of 2833 aphids caught at the “wheatbelt” site, 64 transmitted CMV to lupin test plants. At the irrigated site, numbers of aphids transmitting CMV/numbers caught were 12/186 while the corresponding numbers for BYMV were 11/727. M. persicae, A. kondoi and R. padi transmitted both viruses, while additional vectors of CMV found were A. craccivora, Acyrthosiphon pisum, B. rumexicolens, L erysimi, R. insertum, T. trifolii and Toxoptera citricidus. Averaged over four years, A. kondoi accounted for 50% of CMV transmissions at the “wheatbelt” site, M. persicae for 16% and R. padi for 22%, and these three species were caught in the greatest numbers, comprising 28%, 13% and 37% respectively of the total catch. At the irrigated site R. padi accounted for half the CMV transmissions, while R. padi and A. kondoi together accounted for most of the BYMV transmissions. R. padi, A. kondoi, M. persicae and T. citridus were the most common aphid species at this site. These findings suggest that M. persicae, A. kondoi and R. padi are the aphid species likely to be most important as vectors of CMV and BYMV in narrow-leafed lupins grown in mediterranean-type climatic zones of southern Australia.     

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.     

The acquisition of a caulimovirus by different aphid species: comparison with a potyvirus

The acquisition and transmission of cauliflower mosaic virus (CaMV) by six aphid species and three clones of aphids was studied and compared with that of turnip mosaic virus (TuMV) with Myzus persicae. Two clones of Aphis fabae were unable to transmit CaMV, but the other species, Acyrthosiphon pisum, Brevicoryne brassicae, Megoura viciae, M. persicae and Rhopalosiphum padi transmitted in a bior multi-phasic manner. There was no statistical evidence of a bimodal transmission pattern. R. padi is recorded as a vector of CaMV for the first time. The transmission efficiency of CaMV varied with time of acquisition and suggested that accumulation of the virus occurred with two peaks of efficiency within the anterior region of the insect gut. The time at which these two peaks occurred varied between the species, but the basic pattern was common to all transmitting aphid species in this study. This pattern contrasted with that of TuMV. The transmission data are discussed in terms of bimodal transmission, the influence of feeding behaviour, the role of a helper protein associated with both TuMV and CaMV and the evidence for site specific attachment of CaMV.     

The influence of primary infection date and establishment of vector populations on the spread of yellowing viruses in sugar beet

In three field experiments in 1985 and 1986, we studied the effect of the date of primary infection on the spread of beet yellows closterovirus (BYV) and beet mild yellowing luteovirus (BMW) from artificially inoculated sugar beet plants. Laboratory-reared vector aphids, Myzus persicae, were placed on these sources of virus. There was no substantial natural immigration of vectors or viruses. In two experiments, one with BMYV in 1985 and the other in BYV in 1986, populations of vector aphids remained low and there was little virus spread, i.e. c. 50 infected plants from one primarily infected source. The cause of this small amount of spread was the low number of vector aphids. In the third experiment, with BYV in 1986, large populations of M. persicae developed and there was substantial virus spread: c. 2000 infected plants in the plots which were inoculated before canopy closure. In later-inoculated plots in the same experiment, there was much less spread: c. 100 infected plants per virus source plant. Differences between fields in predator impact are implicated as the most probable factor causing differences in vector establishment and virus spread between these three experiments. Virus spread decreased with later inoculation in all three experiments. A mathematical model of virus spread incorporating results from our work has been used to calculate how the initial proportion of infected plants in a crop affects the final virus incidence. This model takes into account the effect of predation on the development of the aphid populations. The processes underlying the spread and its timing are discussed.     

Factors affecting the relative abundance of two coexisting aphid species on sugar beet

1 The two most common species of aphid colonizing sugar beet Beta vulgaris L. are Myzus persicae (Sulzer) (Hemiptera: Aphididae) and Aphis fabae Scopoli (Hemiptera: Aphididae). 2 M. persicae colonizes sugar beet earlier than A. fabae but the population of the former also declines earlier. Despite similar numbers of each species migrating at the time of colonization, M. persicae is usually less abundant on the crop than A. fabae, suggesting differences between the species in their selection of, and performance on, sugar beet. 3 The intrinsic rate of increase of both species declines as sugar beet matures, however, at any given plant age the intrinsic rate of increase of A. fabae is one and a half times greater than that of M. persicae. This results in more rapid population growth and a later decline of the population. 4 Intraspecific competition appears to result in M. persicae becoming very restless, but there is no evidence for interspecific competition between the two species on this host. 5 A population growth model which takes account of the decline in host quality of sugar beet shows that the M. persicae population peaks 30 days before that of A. fabae, and, excluding differences in emigration rate, the maximum A. fabae population is 14 times greater than the maximum M. persicae population. These results are compared to field data.     

The combined use of mineral oils and pyrethroids to control plant viruses transmitted non- and semi-persistently by Myzus persicae

A mixture of the pyrethroid WL85871 (an enriched form of cypermethrin) and the mineral oil SC811 intoxicated adult apterae of an insecticide-susceptible clone of the peach-potato aphid Myzus persicae at a similar rate to a treatment containing only WL85871, but the mixture killed more. Mixtures of WL85871 andSC811 also gave better control of both acquisition and inoculation by M. persicae of the non-persistent potato virus Y (PVY) than either component alone. A mixture of the pyrethroids deltamethrin, cypermethrin and PP321 with SC811, or a mixture of WL85871 with the mineral oil Bayol 52 also decreased acquisition of PVY, and a mixture of WL85871 with SC811 decreased acquisition of another non-persistently transmitted virus, beet mosaic virus. Control with mixtures was generally better than that provided by each component applied separately. When testing acquisition or inoculation of the semi-persistent beet yellows virus, fewest plants were infected in treatments incorporating both WL85871 and SC811.     

Effects on Myzus persicae (Sulz.) and transmission of beet yellows virus of applying certain trace elements to sugar beet

Applications of lithium chloride (LiCl), zinc sulphate (ZnSO₄) or nickel sulphate (NiSO₄) to the roots of sugar-beet plants in the glasshouse encouraged settling on the leaves of adult apterae from a clone of Myzus persicae (Sulz.); conversely, treatment with boric acid (H₂B₂O₇) inhibited aphid settling. Larviposition of M. persicae was increased by NiSO₄ and tin chloride (SnCl₂). Viruliferous M. persicae transmitted beet yellows virus (BYV) more efficiently to plants treated with LiCl or H₂B₂O₇ than to those treated with copper sulphate (CuSO₄), ZnSO₄ or SnCl₂. The sulphate and chloride anions of the applied chemicals appeared to have little effect on M. persicae and virus transmission. It is suggested that applications of trace elements to sugar beet affected M. persicae and virus transmission by changing the concentrations of trace elements in the aphids' diet and by altering the metabolism of the leaf tissues in the host plant.     

Effect of dodecanoic acid on the colonisation of sugar beet by aphids and the secondary spread of virus yellows

The effect of dodecanoic acid, a behaviour-controlling chemical which alters aphid feeding behaviour and virus transmission efficiency, was studied under field conditions. In a first experiment, dodecanoic acid reduced the level of natural colonisation of sugar beet by Aphis fabae. A second experiment revealed no significant influence of dodecanoic acid on the secondary spread of the semi-persistent beet yellows virus and the persistent beet mild yellowing virus.     

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 aphid alarm pheromone derivatives and related compounds on non- and semi-persistent plant virus transmission by Myzus persicae

A derivative, prepared from the aphid alarm pheromone (E)-β-farnesene and the saturated straight 14-carbon chain dialkyl ester of acetylene dicarboxylic acid, was the most active compound tested for inhibiting acquisition by the aphid Myzus persicae of the non-persistently transmitted potato virus Y (PVY). Derivatives lacking, or with shorter, or branched or partially-unsaturated carbon side-chains were less active. The one derivative tested also inhibited inoculation of PVY, and acquisition of beet mosaic virus (also non-persistent) and the semi-persistent beet yellows virus. However, it had no obvious effect on aphid probing behaviour; related compounds lacking the (E)-β-farnesene moiety also inhibited acquisition of PVY.     

Serological Detection of Beet Western Yellows Luteovirus in the Non-vector,Brevicoryne brassicae (Homoptera: Aphididae)

The interaction between beet western yellows luteovirus (BWYV) and the aphid species Brevicoryne brassicae was investigated using virus transmission and serological detection experiments. This species failed to transmit a BWYV isolate from infected to healthy oilseed rape plants, although virus was readily detected by double-antibody sandwich enzyme-linked immunosorbent assay (DAS-ELISA) in single B. brassicae adults. When virus-carrying adults were tested by ELISA after different inoculation access periods, the number of virus-positive individuals decreased after 5 days, whereas with the efficient vector Myzus persicae, virus-positive individuals were found even after 10 days. This confirms the inability of B. brassicae to transmit BWYV, even though it may acquire the virus. It is suggested that B. brassicae, as compared with the efficient vector M. persicae, may serve as an experimental model for studying the mechanisms of the luteovirus-vector specificity     

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.     

Beet mosaic virus: its Vector and Host Relationships

In order to understand the various factors which affect Beet mosaic virus (BtMV) epidemics, different aspects of the relationships between this virus, its vectors and sugar beet were studied. The latency and incubation periods, determined under growth chamber and field conditions, responded inversely to the temperature and leaf growth rate. Field-infected plants could function as virus sources during the whole growing season. The virus was transmitted by Acyrthosiphon pisum, Aphis fabae, Macrosiphum euphorbiae, Metopolophium dirhodum, Myzus persicae and Ropalosiphum padi. Myzus persicae retained the virus for at least 16 h. Alatae and apterae of M. persicae transmitted the virus with the same efficiency, and in at least two consecutive probes. The proportion of infected plants increased as a logarithmic function of the number of alatae of six aphid species used in the arena tests.