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.     

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.     

Effects of position within wheat field and adjacent habitats on the density and diversity of cereal aphids and their natural enemies

The spatial structure of agricultural landscapes can have a strong impact on the distribution and diversity of insects. Here we studied the effects of within-field position (edge or center) as well as adjacent habitats on the community structure of the natural enemies of cereal aphids. Twelve agricultural sites were included in the study, with two spring wheat fields selected for each site (one adjacent to an alfalfa field, the other adjacent to a corn field). We sampled two rows per field (1 and 20 m from the edge) using pitfall trapping for ground-dwelling predators, sweep netting for leaf-dwelling predators and hand collecting of aphid mummies for parasitoids. Adjacent alfalfa areas, as opposed to corn fields, can significantly increase the abundance and diversity of leaf-dwelling predators and parasitoids near the field edges. Abundance and diversity were found significantly higher near the edges than in the centers of fields adjacent to alfalfa areas. In contrast, no significant differences were found between edges and centers of fields adjacent to corn fields. Of the fifteen most abundant species, Aphidius avenae (Haliday), A. gifuensis (Ashmead), Hippodamia variegata (Goeze) and Chrysopa sinica (Tjeder) were significantly more abundant near the edge than in the center. Being adjacent to alfalfa habitats could enhance parasitism and predator/prey ratios of leaf-dwelling predators at the edges, but has no effects on ground-dwelling predators. In conclusion, the effect of within-field position and adjacent habitats on natural enemies of agricultural pests was species specific. This should be considered for designing efficient plans of biological control.     

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.     

Intraguild predation by aphidophagous predators on parasitised aphids: the use of multiple video cameras

A video technique that allows simultaneous behavioural observations of several experimental replicates under field and laboratory conditions is described. The technique was used to analyse predation risk of parasitised aphids in a sugar beet field. The images of 16 black and white video cameras were recorded by a video multiplexer in combination with a time-lapse video recorder. Each camera was weather protected and equipped with a single infrared diode to allow observations during night times. Single leaves carrying aphid mummies only or mummies and unparasitised aphids were monitored. All colonies were exposed to predation and parasitation by the community of natural enemies in the field. Colonies with mummies and unparasitised aphids were visited significantly more often by predators than those without additional aphids. Predators also stayed significantly longer in patches with unparasitised aphids. Although an equal proportion of aphid mummies were destroyed in both treatments, the video analysis showed differences in predator species spectrum between treatments. In patches with aphids, coccinellid and hemipteran predators preyed on mummies, while in patches with only mummies, chrysopids accounted for most of the damage. The decrease in parasitoid survival could be attributed to the increasing number of predator visits in aphid patches and to a lesser extent to the decreasing number of unparasitised aphids (alternative prey). Parasitoid survival in colonies without alternative prey was correlated with the number of predator visits and the time predators spent on a leaf.Continuous video observations gave additional behavioural information for the interpretation of field data. Other prospective research fields for the use of the multi video camera technique are outlined and general advantages and disadvantages are discussed.     

Survey of two beet viruses in South Kazakhstan and Central Asia

During two surveys of beet root crops in South Kazakhstan and Central Asia conducted in 1988 and 1989, 465 of 990 samples were found to contain beet mosaic virus (BMV) by double-antibody sandwich (DAS) ELISA. BMV infection was widely scattered in the area surveyed, and its incidence varied considerably, reaching 100% in some fields adjacent to beet seed crops. BMV isolates from Kazakhstan, Kirgizia, Uzbekistan and Ukraine were found to be serologically closely related in DAS-ELISA test. Beet yellows virus (BYV) was not detected in any location surveyed in South Kazakhstan and Central Asia. BYV spread into the area is probably prevented by its geographical isolation.     

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.     

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.     

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.     

The potential of predators in natural control of aphids in wheat: Results of a ten-year field study in two German landscapes

In the present study, we investigated the natural control of aphids by predators in wheat fields in a low (L) and high-input cropping region (H) of Germany during a 10-year period. Data for the statistical analyses were obtained from weekly after the start of aphid emergence. The mean annual aphid indices, calculated as the sum of Sitobion avenae (Fabr.), Rhopalosiphum padi (L.), Metopolophium dirhodum (Walk.)(Homoptera: Aphididae), were 30.4 and 81.5 × 103 aphid days per m2, for L and H, respectively. Nine predator fractions were analysed: Coccinella septempunctata L., adults (1) and larvae (2), Propylea quatuordecimpunctata (L.) (Coleoptera: Coccinellidae) adults (3) and larvae (4), syrphid larvae (mostly Episyrphus balteatus [De Geer] (Diptera: Syrphidae)) (5), Chrysoperla carnea Steph. (Neuroptera: Chrysopidae) larvae (6), and adult carabids (7), staphylinids (8) and spiders (9). The two sites were comparable in terms of the mean size of the overall predator community, expressed in predator units (PU): 4.9 PU/m2 (L) vs. 5.4 PU/m2 (H). Most predator fractions responded numerically to increasing aphid densities. The numerical response was strongest in syrphid larvae, scarcely detectable in adult coccinellids, and virtually non-existent in epigeic arthropods. Multiple regression models revealed indirect relationships between the weekly overall predator community densities (PU/m2) and individual predator fractions (individuals/m2) and absolute rates of aphid density increase (individuals/m2) one or two weeks after baseline. A site-independent reduction of the aphid density increase to nil (y = 0) was observed at 3.9 to 4.2 PU/m2. Consequently, the 2.7 times higher aphid density at H cannot be attributed to the presence of fewer predators or lower effects of the overall predator community or of any individual predator fraction.     

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.     

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.     

Role of aphid predator guild in controlling spirea aphid populations on apple in West Virginia,USA

Spirea aphid populations and their predators were studied on apple to identify predators of importance in controlling aphid populations. Methods included random and non-random sampling from apple orchards in West Virginia, USA, sentinel aphid colonies, laboratory feeding studies, and predator exclusion studies. Aphidoletes aphidimyza (Diptera: Cecidomyiidae), chrysopids (Neuroptera: Chrysopidae), Harmonia axyridis (Coleoptera: Coccinellidae), and Orius insidiosus (Hemiptera: Anthocoridae) were the most abundant predators associated with spirea aphid colonies on apple. Parasitoids were all but absent in the study. Abundance of all predators was density dependent with greater responses to aphid populations at the orchard scale than to tree or individual colony scales. A. aphidimyza, O. insidiosus, chrysopids, and syrphids (Diptera) had the greatest degree of density dependence on aphid populations, and spiders showed inverse density dependence. Exclusion of predators with both cages and insecticides produced significantly higher aphid populations. Because of high abundance, good synchrony with aphid populations, and high impact per individual, H. axyridis adults were the most important spirea aphid predator on apple.     

Identification of Top-Down Forces Regulating Cotton Aphid Population Growth in Transgenic Bt Cotton in Central China

The cotton aphid Aphis gossypii Glover is the main aphid pest in cotton fields in the Yangtze River Valley Cotton-planting Zone (YRZ) in central China. Various natural enemies may attack the cotton aphid in Bt cotton fields but no studies have identified potential specific top-down forces that could help manage this pest in the YRZ in China. In order to identify possibilities for managing the cotton aphid, we monitored cotton aphid population dynamics and identified the effect of natural enemies on cotton aphid population growth using various exclusion cages in transgenic Cry1Ac (Bt)+CpTI (Cowpea trypsin inhibitor) cotton field in 2011. The aphid population growth in the open field (control) was significantly lower than those protected or restricted from exposure to natural enemies in the various exclusion cage types tested. The ladybird predator Propylaea japonica Thunberg represented 65% of Coccinellidae predators, and other predators consisted mainly of syrphids (2.1%) and spiders (1.5%). The aphid parasitoids Aphidiines represented 76.7% of the total count of the natural enemy guild (mainly Lysiphlebia japonica Ashmead and Binodoxys indicus Subba Rao & Sharma). Our results showed that P. japonica can effectively delay the establishment and subsequent population growth of aphids during the cotton growing season. Aphidiines could also reduce aphid density although their impact may be shadowed by the presence of coccinellids in the open field (likely both owing to resource competition and intraguild predation). The implications of these results are discussed in a framework of the compatibility of transgenic crops and top-down forces exerted by natural enemy guild.     

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.     

Aphids from mangold clamps and their importance as vectors of beet viruses

Mangold clamps are over-wintering sources of the aphid-transmitted beet mosaic, beet yellows and beet mild yellowing viruses, and of several species of aphid, three of the most common in clamps being Myzus persicae, Rhopalosiphoninus staphyleae tulipaellus and R. latysiphon. This study attempted to assess the relative importance of the different species in spreading viruses from clamps. Compared with M. persicae, R. s. tulipaellus and R. latysiphon are seldom trapped in flight, except near large infestations. Alatae of M. persicae and R. s. tulipaellus become common in clamps in April, but few fly below 15d? C., a temperature seldom reached in eastern England in early spring. Flight muscle autolysis, which occurs later in R. s. tulipaellus and R. latysiphon than in some aphid species, also probably prevents many alatae in clamps from flying. We confirmed the importance of clamps as sources of beet viruses, the percentage of infected plants decreasing with increasing distance from infested clamps. M. persicae is shown to be a better vector of beet viruses than the other clamp aphids, and is probably responsible for most virus spread from clamps. R. s. tulipaellus did not transmit beet mosaic virus, but it is a fairly efficient vector of beet yellows and beet mild yellowing viruses, and, although we did not find this species on sugar beet in the field, it probably spreads these viruses from clamps. R. latysiphon did not transmit any of the viruses, and the role of Macrosiphum euphorbiae, Aulacorthum solani and Myxus ascalonicus is probably small.     

Intraspecific variation of Myzus persicae on sugar beet (Beta vulgaris)

Differences in inherited resistance among seven sugar-beet stocks had similar effects on Myzus persicae clones representing the range of variation in aphid response to resistant and susceptible sugar beet observed in fifty-eight clones collected between 1969 and 1971. Three sugar-beet stocks were consistently resistant. Statistically significant interactions between beet stocks and aphid clones did not indicate the existence of biotypes with specific abilities to overcome resistance. M. persicae clones differed in their vigour of colonizing sugar beet, irrespective of the differences between beet stocks. The readiness of adult aphids to settle determined the size of aphid population produced and included a component related to the response of the aphid clone to sugar beet as a host, and a component related to the resistance ranking of the beet stock. Breeding sugar beet with resistance to aphids will be simplified, as the results indicate that, at present, differences between aphid biotypes need not be considered a problem.     

Predator-induced morphological shift in the pea aphid

Aphids exhibit a polymorphism whereby individual aphids are either winged or unwinged. The winged dispersal morph is mainly responsible for the colonization of new plants and, in many species, is produced in response to adverse environmental conditions. Aphids are attacked by a wide range of specialized predators and predation has been shown to strongly influence the growth and persistence of aphid colonies. In two experiments, we reared two clones of pea aphid (Acyrthosiphon pisum) in the presence and absence of predatory ladybirds (Coccinella septempunctata or Adalia bipunctata). In both experiments, the presence of a predator enhanced the proportion of winged morphs among the offspring produced by the aphids. The aphid clones differed in their reaction to the presence of a ladybird, suggesting the presence of genetic variation for this trait. A treatment that simulated disturbance caused by predators did not enhance winged offspring production. The experiments indicate that aphids respond to the presence of a predator by producing the dispersal morph which can escape by flight to colonize other plants. In contrast to previous examples of predator-induced defence this shift in prey morphology does not lead to better protection against predator attack, but enables aphids to leave plants when mortality risks are high.     

Functional response and reproductive attributes of the aphidophagous ladybird beetle,Harmonia dimidiata (Fabricius) in oak trees of sericultural importance

Harmonia dimidiata (Fabricius) (Coleoptera: Coccinellidae) is the dominant predator of the aphid species Cervaphis quercus Takahashi. This aphid is a serious pest of oak trees in several parts of north-east India. Young leaves of oak trees are used in sericulture by rural people and by industry in several parts of north-east India. The effect of different aphid densities on food consumption and fecundity of H. dimidiata was studied in the laboratory. Female beetles were maintained from the time of eclosion till death at a fixed density of 25, 50, 75,100 or 125 aphids. Both the functional response and the reproductive numerical response showed the upper asymptote at 100 adult aphids/female. At this density, females matured earlier and produced more eggs over a longer reproductive period. At lower prey densities, females matured late and they either did not produce eggs or produced fewer eggs. At the higher prey densities, females did not produce more eggs than the asymptote. Results suggested that H. dimidiata are an effective predator of C. quercus aphids on oak trees and could be exploited as a biological control agent in the rising phase of aphid population growth.     

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.