Bionomics of aphids reared on cereals and some Gramineae

In controlled temperature, light and relative humidity, Metopolophium dirhodum and Sitobion avenae multiplied more on young Proctor barley than on Blenda oats, and less on Cappelle wheat. Rhopalosiphum padi increased in number fastest on barley and slowest on oats. More survived, and generation lengths seemed shorter, on barley for M. dirhodum and S. avenae and on wheat for R. padi. Tests with young cereals outdoors generally agreed with those in controlled conditions. On mature plants, there were more M. dirhodum on barley, more R. padi on wheat and more S. avenae on oats than on the other cereals. Given a free choice in large cages outdoors, most aphids were found on barley. When allowed to choose between grasses, more M. dirhodum were on Dactylis glomerata, Poa pratensis and Festuca pratensis, more R. padi on Lolium perenne and F. pratensis, and more S. avenae on D. glomerata and L. perenne. Most aphids of all species combined were on F. pratensis, Lolium and Phleum, and fewest on Festuca rubra and Holcus mollis.     

Distribution of aphids in cereal crops

Cereal crops were examined weekly for aphids during 1969. Plants in twenty samples of row 0.3 m long were examined in a sheltered perimeter of a crop and along a transect 36.6 m into the crop. Aphids were usually first found within 1–4 weeks of the first alatae caught in a suction trap operating 12.2 m above ground. When first alatae caught in a suction trap operating 12.2 m above ground. When the first found from 10 to 27% of the 0.3 m lengths sampled contained aphids. Rhopalosiphum padi, first found late in May, were scarce (< 0.53/0.3 m) throughout June and July. Sitobium spp. and Metapolophium dirhodum, which appeared in mid-June, were more numerous than R. padi; most occurred during the second half of July, and populations decreased just before harvest in early August. Sitobium avenae was more abundant (max. 19.3/sample) than either S. fragariae (0.91) or M. dirhodum (2.51). More aphids occurred in oats (max. 52/0.3 m) during July than in wheat (45), and barley had fewer (6.8). S. avanae was more abundant than M. dirhodum in sheltered areas of barley and wheat, and in exposed areas of the same crop M. dirhodum was commonest. Along sheltered perimeters, the ratio of S. avenae to M. dirhodum was largest in barley (11:1), intermediate in oats (6:1) and smallest in wheat (3.7:1). Sitobium spp. were most numerous on the ears, when most M. dirhodum were on the leaves. Regression analyses of log. S² on log. m suggested that S. avenae was more evenly distributed within (36.6 m) the field (b = 1.056 + 0.109) than along the sheltered perimeter (b = 1.432 + 0.132), though it seemed similarly distributed along perimeters of barley, oats and wheat. The distributions of M. dirhodum and Sitobium spp. along sheltered perimeters of all crops were apparently similar.     

Interaction between barley yellow dwarf virus and rust in wheat, barley and oats, and the effects on grain yield and quality

In three separate experiments, the upper leaf surface of the fifth formed leaf of wheat cv. Highbury, the fourth and fifth leaves of barley cv. Julia and the third and fourth leaves of oat cv. Mostyn were inoculated in a spore settling tower with wheat brown rust (Puccinia recondita f. sp. tritici), barley brown rust (P. hordei) or oat crown rust (P. coronata f. sp. avenae), respectively. Fewer pustules developed on distal portions of leaves of plants infected with barley yellow dwarf virus (BYDV) than on similar portions of leaves from virus-free plants. There were no significant differences in the number of pustules on proximal leaf portions. In barley and oats, the number of pustules on distal leaf portions was negatively correlated with the amount of yellowing of the leaf areas scored. In wheat, symptoms of BYDV were mild and leaves were little affected by yellowing. The latent period of rust on wheat and oats was not affected by BYDV. In barley, BYDV reduced the latent period of rust on leaf 5, but not on leaf 4, and reduced it on proximal, but not distal, leaf portions. In other experiments, BYDV reduced the yield of wheat and oats by 44% and 66%, respectively, while BYDV-infected barley was almost sterile. The appropriate rust reduced the yield of wheat, barley and oats by 33%, 13% and 86%, respectively. When infected with both BYDV and rust, yield of wheat and oats was reduced by 63% and 91%, respectively. Neither BYDV nor rust affected the percentage crude protein content of wheat grain, nor did rust affect that of barley. In oats, BYDV and rust each significantly increased crude protein of grain, but rust infection of BYDV-infected plants tended to reduce it.     

The effect of barley yellow dwarf virus on honeydew production by the cereal aphids, Sitobion avenae and Metopolophium dirhodum

Individual S. avenae and M. dirhodum excreted significantly fewer droplets of honeydew on plants infected with BYDV than on healthy plants. S. avenae excreted less honeydew on the ears than on the leaves of wheat. M. dirhodum excreted less than S. avenae on the leaves. The size of honeydew droplets increased with the age of aphids but was not affected by BYDV infection. Possible reasons for the observed effects of BYDV on honeydew excretion are discussed.     

Spread of barley yellow dwarf virus and relative importance of local aphid vectors in central Washington

Data from bioassays of field collected aphids, barley indicator plants exposed to natural conditions, and various types of aphid traps were used to describe the spread of barley yellow dwarf virus (BYDV) in wheat and barley near Prosser, Washington. Bioassays were also used to assess the relative importance of local vector species. Of alate aphids collected from grain in the 1982 and 1983 fall migration seasons, 3.4–14–5% transmitted BYDV. Data from concurrent and post-migration assays of resident aphids (apterae and nymphs) reflected an increase in the proportion of infected plants in the field. Maximum increase in the percentage of viruliferous aphids occurred in late November and December of 1982 and November of 1983. The 1982 increase occurred after aphid flights had ceased for the year, suggesting active secondary spread. Collections in pitfall traps and infected trap plants from November to February confirmed aphid activity and virus spread. Rhopalosiphum padi was the most important vector in central Washington in 1982 and 1983 because of its abundance and relative BYDV transmission efficiency. Metopolophium dirhodum was more winter-hardy than R. padi and equal to R. padi in its efficiency as a vector; however, it was not as abundant as R. padi except during the mild winter of 1982–83, when it was a major contributor to secondary spread. Sitobion avenae may be important in years when it is abundant, but it was only a quarter as efficient as R. padi. Rhopalosiphum maidis was a much less efficient vector than R. padi and it only reached high populations in late autumn barley.     

Barley yellow dwarf virus in aphids caught in suction traps, 1969-73

Suction traps operating at low level (1 5 m) were used to catch live alate Rhopalosiphum padi, Macrosiphum (Sitobion) avenae and Metopolophium dirhodum which were tested for transmission of barley yellow dwarf virus (BYDV). The first species caught and infective was R. padi, followed by M. (S.) avenae infective some 2–3 wk later and M. dirhodum 3–4 wk later still. Never more than 11-5% of the annual catch of any species transmitted BYDV and the proportion fluctuated from week to week and between seasons in different years. The relative abundance of infective vectors of ths three species varied; annual numbers of infective M. (S.) avenae and M. dirhodum varied inversely with infective R. padi, the latter also usually transmitted severer virus. The results of the infectivity tests have been compared with the catches of these aphids by the Rothamsted Insect Survey and show that numbers of alate aphids do not necessarily indicate the likely incidence of BYDV.     

Serological and biological characterisation of isolates of barley yellow dwarf virus in Sweden in 1983

In 1983, cereal plants showing symptoms of barley yellow dwarf virus (BYDV), collected from 15 localities in Sweden, were tested for BYDV using enzyme-linked immunosorbent assay (ELISA). Antisera against two Swedish isolates of BYDV were used, a mild isolate (27/77) transmitted specifically by Sitobion avenae and a severe one (39/78) transmitted mainly by Rhopalosiphum padi. No virus was detected in 57 of 607 plants of oats and barley tested. Of the 550 plants in which virus was detected, 366 were infected with viruses similar to isolate 27/77, 116 with viruses similar to 39/78 and the remaining 68 reacted strongly with both antisera. When tested, the latter isolates were shown to be mixtures. Thirty-nine selected samples were also tested with antisera against the USA isolates RPV, RMV, MAV and PAV, and for transmission by S. avenae and R. padi. Twenty-six of these samples were transmitted specifically by S. avenae, one was transmitted only by R. padi and the remaining 12 samples were shown to be infected with a mixture of an S. avenae-specific isolate and one transmitted mainly by R. padi. Antisera against PAV and MAV each detected all isolates tested and the results were very similar to those with the antisera to the 39/78 and 27/77 isolates, respectively. None of the field isolates reacted with antisera against RMV or RPV. It was concluded that 1983 was an epidemic year for BYDV in Sweden and that isolates specifically transmitted by S. avenae predominated. Symptoms of infection by these isolates on oat plants ranged from mild to severe.     

Infection of cereal aphids by the fungus Entomophthora

Three species of Entomophthora killed many Metopolophium dirhodum, M. festucae and Sitobion avenae on wheat at Harpenden, Hertfordshire, in 1971. Rainfall was low and aphid numbers were small in May and there was no Entomophthora infection. E. planchoniana first infected M. dirhodum early in June and the percentage of each aphid species infected increased during and after heavier rain in the first 2 weeks of June. M. dirhodum and S. avenae were most often infected by E. planchoniana, and the less common M. festucae mostly by E. aphidis and E. thaxteriana. Relative frequencies of E. aphidis and E. thaxteriana were largest before, and of E. planchoniana after, mid-July. The largest percentages infected were 53% of M. dirhodum and 30% of S. avenae during the second half of July and 60% of M. festucae in late June. The percentages of old nymphs and apterous adults that were infected were similar and only about half those of infected alatae.     

Comparison of the potential rate of population increase of brown and green color morphs of Sitobion avenae (Homoptera: Aphididae) on barley infected and uninfected with Barley yellow dwarf virus

Life tables of brown and green color morphs of the English grain aphid, Sitobion avenae (Fabricius) reared on barley under laboratory conditions at 20 ± 1°C, 65% ± 5% relative humidity and a photoperiod of 16 : 8 h (L : D) were compared. The plants were either: (i) infected with the Barley yellow dwarf virus (BYDV); (ii) not infected with virus but previously infested with aphids; or (iii) healthy barley plants, which were not previously infested with aphids. Generally, both color morphs of S. avenae performed significantly better when fed on BYDV‐infected plants than on plants that were virus free but had either not been or had been previously infested with aphids. Furthermore, when fed on BYDV‐infected plants, green S. avenae developed significantly faster and had a significantly shorter reproductive period than the brown color morph. There were no significant differences in this respect between the two color morphs of S. avenae when they were reared on virus‐free plants that either had been or not been previously infested with aphids. These results indicate that barley infected with BYDV is a more favorable host plant than uninfected barley for both the color morphs of S. avenae tested, particularly the green color morph.     

The overwintering and abundance of cereal aphids

The longevity of Sitobion avenae varied inversely with temperature from 25 to over 100 days when reared outdoors on barley under lantern jars, being longest between December and April–May. This was due mainly to changes in the length of the pre- and post-reproductive periods rather than the length of the period of reproduction. Fecundity varied directly with temperature, from eight to sixty-seven nymphs per female. When reared on winter wheat and sheltered from wind, rain and snow, S. avenae survived best, Metopolophium dirhodum survived less well and Rhopalosiphum padi worst; when exposed in the open none of the three species survived. Close relationships were not demonstrable between the numbers of alate M. dirhodum, S. avenae and Rhopalosiphum spp. trapped in any year, between the abundance of these aphids and weather conditions between January and April, or between the numbers of these alatae and the size of crop infestations. This was probably because insufficient information was available about their biology, both inside and outside the crop.     

Infection of cereals and grasses by isolates of Polymyxa graminis (Plasmodiophorales)

The host range of isolates of Polymyxa was tested in mono-fungal sand cultures. Fourteen isolates of P. graminis, obtained from barley, wheat, oats or Poa annua and from several different countries, all infected barley and all but one infected wheat. Rye was also a good host, whereas oats (nine cultivars), Lolium multiflorum and Poa pratensis became only slightly infected. Wheat cultivars differed in susceptibility, with Galahad much more resistant than Avalon. Several common weed and pasture grasses were not infected by the two isolates tested. A range of wild Hordeum spp. were mostly susceptible to P. graminis and/or barley mild mosaic virus, which it transmits. An isolate of P. betae, used for comparison, caused slight infection on oats but not on other cereals. The variation within and between species of Polymyxa needs more detailed investigation.     

Identification, Distribution and Vector Population Dynamics of Barley Yellow Dwarf Virus in Three Cereal-Producing Areas of Spain

ELISA-based surveys during 1985–87 in three major cereal-growing areas of Spain confirmed the presence of barley yellow dwarf virus (BYDV). Samples of small grain cereals and grasses with and without BYDV-like symptoms were collected in the central, southwestern, and northeastern Spain. Infections were found in all cereal species sampled and in some grasses. About 37 % of the samples collected in 1985 were infacted with isolates of the PAV serotype. Isolates of the RPV serotype were less common, and were detected only in samples from the central region at El Encin, Madrid. Only a single sample, collected from El Encin in 1987, was unequivocally diagnosed as containing an isolate of the MAV serotype. Aphid vector population dynamics was monitored during fall and winter of 1984–87 in the central region. Rhopalosiphum padi L. appeared to be the most abundant species during fall and winter months, infesting grasses and volunteer wheat. Other species present were Sitobion avenae (F.), Metopolophium dirhodum (Walker) and Rhopalosiphum maidis (Fitch). Both R. padi and S. avenae seem to be anholocyclic in the central region of Spain, and are able to remain and reproduce on wheat volunteers and grasses until the beginning of spring. S, avenae populations increase quickly on wheat volunteers in April, while populations of R. padi remain low. Therefore, spread of S. avenae-transmitted BYDV types to neighbouring cereal fields seem more likely to occur than spread of other types. Other possible virus reservoirs, such as maize, also need investigation for a better understanding of BYDV epidemiology in the central and other cercal-growing areas of Spain.     

Survival and duration of development of Oscinella spp. (Diptera: Chloropidae) on different Gramineae in the laboratory

The duration of development and survival of Oscinella frit, O. nitidissima and O. nigerrima at different growth stages of several plant species was investigated in the laboratory. The effect of the infestation on the growth of plants was also quantified. Survival of O. frit larvae was significantly lower (P < 0.01) on barley than on either oats or wheat, between which there was no significant difference. Survival was usually highest when plants were infested at the two-leaf stage and lowest when plants were infested at the five-leaf stage. The duration of development of both male and female O. frit was longer on barley than on either oats or wheat and generally increased when older plants were infested. The proportion of male flies was greatest when older plants were infested. Duration of development and survival of O. frit was similar on oats and Lolium multiflorum, but larvae did not survive on Dactylis glomerata. Of the other Oscinella species, O. nitidissima larvae did not survive on oats, wheat and barley and O. nigerrima larvae did not survive on either oats or barley. The main shoots of cereal plants infested at the two-, three- or four-leaf stages were rapidly killed by O. frit larvae whereas many of those infested at the five-leaf stage continued to grow slowly. In response to attack proportionately more tillers were produced by oat plants than by either wheat or barley plants. When cereals were infested with O. nigerrima larvae the central shoots were damaged, but most recovered and continued to grow. Damage symptoms were most obvious when cereals were infested at the two-leaf stage, oats being more severely affected than either wheat or barley.     

Barley yellow dwarf virus, wheat, and Sitobion avenae: a case of trilateral interactions

We analysed interactions in the system of two Barley Yellow Dwarf Virus (BYDV) strains (MAV and PAV), and wheat (cv. Tinos) as host plant for the virus, and the cereal aphid Sitobion avenae (F.) as vector, in particular whether or not infection by the virus might alter host plant suitability in favour of vector development. By measuring the amino acid and sugar content in the phloem sap of infected and non‐infected wheat plants we found a significant reduction in the concentration of the total amount of amino acids on BYDV‐infected plants. Qualitative and quantitative analysis of honeydew and honeydew excretion indicated a lower efficiency of phloem sap utilisation by S. avenae on infected plants. In addition, S. avenae excreted less honeydew on infected plants. Both BYDV strains significantly affected aphid development by a reduction in the intrinsic rate of natural increase. Hence, infection by the virus reduced the host suitability in terms of aphid population growth potential on BYDV‐infected plants. However, more alate morphs developed on virus‐infected plants. These findings are discussed in relation to the population dynamics of S. avenae, and, as a consequence, the spread of BYDV.     

Entomophthora infecting the cereal aphids Metapolophium dirhodum and Sitobion avenae

Entomophthora aphidis, E. planchoniana, and E. thaxteriana killed many Metapalophium dirhodum and Sitobion avenae in barley during 1970. E. planchoniana first infected M. dirhodum late in June, after rain ended a long dry period, but few aphids were infected until after July 7, when heavy rain killed 65–80% of aphids. E. planchoniana was the most frequent of the 3 species until July 27, when E. aphidis and E. thaxteriana each became more abundant. S. avenae was more often infected by E. thaxteriana than the other species of Entomophthora. During the second half of July, 40–76% of adult M. dirhodum and 34–80% of S. avenae were infected. The 3 fungus species were equally common in M. dirhodum in sheltered fields, but E. thaxteriana was less common in an exposed field. In a sheltered field, E. thaxteriana was less frequent than the other species along the perimeter, and E. planchoniana was most common and E. aphidis the least common about 37 m into the crop.     

The invasion of root tips of cereals by the cereal cyst nematode Heterodera avenue

Newly germinated seedlings of susceptible cultivars of oats, wheat, barley and rye were inoculated with second-stage juveniles of Heterodera avenae in pots of sand. Subsequent examination showed oat root tips to be more commonly invaded, and by a greater range of nematode numbers than the other cereals. A comparison of oats and barley showed that lower nematode numbers in barley were not due to a higher emigration from barley; invasion, establishment and emigration by nematodes all being greater in oats. Second-stage juveniles were more likely to migrate prior to establishment in barley than in oats.     

The invasion and development of the cereal cyst-nematode, Heterodera avenae in the roots of autumn-and spring-sown cereals

Observations were made at 2 or 4 wk intervals from December to harvest on all stages of Heterodera avenae in winter oats growing on infested land. Second-stage larvae were present in all soil samples except on 5 and 20 July. Invasion and development of larvae was slow during winter. The nodal and seminal roots of winter oats were both heavily invaded by the nematode; larvae which invaded seminal roots tended to become male whereas those in nodal roots tended to become female. There was a small second invasion in August. Females were first observed on the roots of winter oats on 17 May, 214 days after the crop was sown and 62 days after the first fourth-stage larva was observed. The nodal roots of spring barley contained few H. avenae larvae whereas these roots were heavily invaded in winter wheat and oats. In spring barley the nodal roots were developing in June and July when few second-stage larvae were in the soil whereas in winter oats and wheat the nodal roots were growing rapidly in April when larvae were most numerous, and so were heavily invaded.     

The overwintering of cereal aphids on Gramineae in southern England, 1977–1980

Cereal aphids were sampled monthly on graminaceous host plants on farmland in Hampshire, UK from September 1977 to April 1980. Sitobion avenae overwintered anholocyclically on most types of Gramineae, particularly seedling grass crops and small-grain cereals. S. fragariae was probably mainly holocyclic on Rubus spp. Large summer populations of Metopolophium dirhodum were probably derived from holocyclic colonies on roses. M. festucae (subsp. cerealium) appeared to overwinter only anholocyclically, and was found mainly on mature grass crops and grasses in hedgerows. Rhopalosiphum padi was the most numerous species in autumn and overwintered anholocyclically in all three winters studied. R. insertum and R. maidis were rare and did not appear to survive the winter anholocyclically on graminaceous hosts.     

The effects of sowing date and choice of insecticide on cereal aphids and barley yellow dwarf virus epidemiology in northern England

During the mid-1980s, Sitobion avenae became recognised as an important vector of barley yellow dwarf virus (BYDV) in the Vale of York. A field trial at the University of Leeds Farm, North Yorkshire, was carried out during the autumn/winter of 1984-85 to evaluate different control procedures against S. avenae-transmitted BYDV and to investigate its epidemiology. Winter barley was sown on three dates in September, and plots were sprayed with either deltamethrin, demeton-S-methyl or pirimicarb on one of three dates between mid-October and mid-November, making a factorial design. Rhopalosiphum padi, the main vector of BYDV in southern England, were rarely found during the experiment, but the numbers of S. avenae were much higher, reaching a peak of 21% of plants infested in the unsprayed plots of the first sowing date. Single applications of each insecticide reduced populations of S. avenae to zero. Some treatments, particularly in the early sown plots and those treated with pirimicarb, however, did allow some recolonisation, and thus led to increased virus incidence and decreased yields. Sprays applied before the end of the migration of S. avenae were more efficient at controlling BYDV if the insecticide was persistent, otherwise a spray after this period, in November, was more effective. Virus incidence, although reduced by sprays, was generally low in plots sown on 18 and 27 September. In contrast, about 11% of plants were infected in unsprayed plots sown on 6 September and a small yield benefit was obtained with insecticidal treatments. Enzyme-linked immunosorbent assay (ELISA) of plants taken from the plots indicated that MAV- and PAV-like strains were present, and were most likely to have been transmitted by S. avenae.     

Preference of cereal aphids for different varieties of winter wheat

This paper reports on the development of a simple and robust preference meter (developed in-house) to score the host choice behavior of apterous aphids. With this tool, the preferences of two important cereal aphids Sitobion avenae (Fab.) and Metopolophium dirhodum (Walker) were investigated against four different varieties of winter wheat (Triticum aestivum L.) with a different susceptibility for Fusarium head blight (FHB). Differences in the choice behavior of both aphid species were observed for different wheat varieties. The preferred wheat variety of S. avenae and M. dirhodum was not the same. Also, both aphid species clearly had a differential preference for seedlings and ears. Using seedlings, M. dirhodum was about 1.8 times more rapid in making its choice than S. avenae. In separate experiments with ears, S. avenae was 4.5 times faster than in the experiments with seedlings. In the present study, we aim to highlight differences in preference behavior in relation to potential mechanisms for host selection.