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.     

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.     

Observations on the morphs of Macrosiphum avenae and Metopolophium dirhodum on cereals during the summer and autumn

From June to early August 1970, populations of Macrosiphum avenae and Metopolophium dirhodum on marked tillers of field barley were compared with the numbers of alatae trapped at crop height and at 12.2 m. There were always more M. dirhodum than M. avenae on the tillers. Only apterae were produced until mid-June when both alatae and apterae occurred; after mid-July only alate M. avenae were found. Until mid-June most of the flying alatae were caught at 12.2 m as they migrated from spring hosts to cereals. Thereafter, more alate M. avenae were trapped at 12.2 m than at crop level, whereas numbers of alate M. dirhodum were usually comparable at both heights. Although crop and flying populations occasionally showed temporal similarities, insufficient is known about their field distribution and the factors affecting their alate production and flight activity to interpret this relationship. In the autumn, two consecutive reproductive phases of M. dirhodum occurred on winter wheat grown in pots outdoors. Initially, apterous virginoparae and alatae, probably sexuparae, were produced, whereas only alate males appeared during the second phase. In contrast, M. avenae deposited mainly apterous virginoparae although some oviparae developed in October to lay scattered, probably infertile eggs on the tillers.     

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.     

Field assessments of resistance to the aphids Sitobion avenae and Metopolophium dirhodum in old and modern spring-sown wheats

Three ancient varieties of wheat and two modern spring-sown cultivars were investigated in the field to assess their resistance to the cereal aphids Sitobion avenae and Metopolophium dirhodum; the monitoring of natural populations and detailed observations using clip-cages were both undertaken. The ancient variety Einkorn showed both antixenotic and antibiotic resistance to S. avenae, and antibiotic resistance to M. dirhodum. The resistance to S. avenae was greater than that to M. dirhodum. The value of Einkorn as a component of a resistance-breeding programme is discussed with reference to its agronomic and genetic characteristics and with reference to the pest status of cereal aphids.     

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 effect of temperature and wind on the flight activity of cereal aphids

The lower temperature threshold for take-off in Sitobion avenae obtained from an analysis of daily 12·2 m suction trap catches was 16°C. In the laboratory, the take-off threshold for S. avenae was 17·5°C in increasing temperatures, but 19°C when aphids were kept at constant temperatures for a short period of time. The equivalent thresholds were both 20·5°C for Metopolophium dirhodum. Over a period of 16 h no S. avenae took-off from plants at 10°C, 70% at 15°C and all within 16 h at 20°C. It was concluded that suction trap catches can be used to compare the temperature thresholds for take-off of both different species and morphs of a species of aphid. Emigrants of Rhopalosiphum padi, but not of M. dirhodum, took-off at a higher temperature than the alate exules. High winds were found to delay but not inhibit take-off both in the field, and in the laboratory, using both artificial and plant substrates. All aphids eventually flew, even from favourable hosts. It was concluded that adverse weather conditions will delay but not prevent cereal aphid migration in early summer.     

Reproductive strategy of winged and wingless morphs of the aphids Sitobion avenae and Metopolophium dirhodum

The reproduction of apterous and alate morphs of the aphids Sitobion avenae and Metopolophium dirhodum is compared on the basis of fecundity in 5- and 10-day periods of adult life. Apterae of both species are consistently more fecund than alatae of comparable weight, producing about three more nymphs on average in any 5-day period. The reproductive differences are related to the number and quality of embryos at eclosion and to ovulation rates, both of which in turn appear to be linked to wing-muscle maintenance. These relationships between weight, embryos and reproduction may be used to predict a newly moulted adult aphid's fecundity, a method which may facilitate the assessment of resistance to aphids in new cereal varieties, by obviating lengthy recording of reproduction. The strategies by which alatae of these and other aphid species minimize the difference between their fecundity and that of apterae are discussed.     

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.     

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.     

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.     

Geographic and microgeographic genetic differentiation in two aphid species over southern England using the multilocus (GATA)4 probe

Samples of the grain aphid Sitobion avenue (F.) and the rose-grain aphid Metopolophium dirhodum (Walker) were collected in late March from wheat fields and adjacent road-side grasses at a number of locations in southern England. Unparasitized aphids were DNA fingerprinted using the multilocus (GATA)₄ probe. Over all locations, the fingerprints of individual S. avenue caught in wheat had lower overall average distances of band migration (ADBM) and shared a higher proportion of bands, than fingerprints of individuals caught in adjacent road-side grasses. The ADBM of fingerprints of S. avenue collected on road-side grasses altered significantly with geographical location, while the ADBM of fingerprints of S. avenue caught on wheat did not. A comparison of the fingerprints of individual M. dirhodum caught in wheat and neighbouring road-side grasses did not reveal any genetic differentiation. Fingerprints of M. dirhodum that were caught in the same host type did however, show significant variation in ADBM between different locations. With both S. avenue and M. dirhodum, spatial autocorrelation revealed that locations that were close together were no more likely to have individuals with similar ADBM than locations mat were far apart Our results suggest that (i) particular clones of S. avenue prefer to colonize wheat; and/or that (ii) particular clones of S. avenae perform better on wheat man other clones. It is unclear why M. dirhodum did not show any genetic structuring according to host type, but this species appears to engage in sexual reproduction much more frequently than S. avenae in southern England. M. dirhodum is likely to have displayed genetic heterogeneity between locations either because of founder effects, or because of genetic drift.     

Effects of vernalisation and aphid culture history on the relative susceptibilities of wheat cultivars to aphids

The weight and embryo number of adults of the cereal aphid Metopolophium dirhodum were compared on vernalised and unvernalised seedlings of six winter wheat cultivars. Aphids were smaller when reared on vernalised plants and contained fewer embryos than those reared on unvernalised material. Resistance rankings of the cultivars also changed markedly; cultivars at the extremes of the resistance sequence when vernalised were not separable statistically when unvernalised. When Sitobion avenae was tested on five of the six unvernalised cultivars, results agreed with earlier work on this species but the ranking differed from that for M. dirhodum. The plant species on which the test aphid's parent had been reared markedly affected the level of resistance and this and the vernalisation effects are discussed in relation to earlier work on aphid resistance in cereals.     

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.     

The influence of nitrogen fertiliser on the population development of the cereal aphids Sitobion avenae (F.) and Metopolophium dirhodum (Wlk.) on field grown winter wheat

The effect of nitrogen fertiliser on populations of the cereal aphids Sitobion avenae and Metopolophium dirhodum on winter wheat was investigated in a three year field experiment. Naturally occurring aphid populations were monitored on three nitrogen treatments; none, nitrogen application using Canopy Management guidelines (130–210 kg ha^-1) and conventional practice (190 kg ha^-1). Inoculations of laboratory reared S. avenae were used to enhance field populations on half the plots. Natural populations of M. dirhodum remained below the current UK spray threshold level of two-thirds of shoots infested at the start of flowering, or five aphids per shoot in all years, whilst populations of S. avenae exceeded the threshold in all years. The response of the two species to nitrogen differed. Significantly higher populations of M. dirhodum were recorded in both treatments which received nitrogen in all years, whilst the response of S. avenae varied between years. In 1994 and 1995 when environmental conditions favoured aphid development, higher populations were recorded in the two treatments which received nitrogen. In 1993 when high rainfall created unfavourable conditions, higher populations were recorded in the plots receiving no nitrogen. Differences in peak density and cumulative aphid index of S. avenae resulted from differences in the rate of population increase between ear emergence and peak density on the different treatments. Populations prior to ear emergence were higher in the plots which received nitrogen but the differences were not statistically significant. There was no evidence of a difference in the timing of population decline in the different treatments. In 1993 higher levels of infection by entomopathogenic fungi were observed in all treatments. Significantly higher levels of infection were recorded in the treatments receiving nitrogen, which may have accounted for the lower S. avenae populations recorded. It is possible that the larger canopies recorded in these treatments produced conditions which favoured infection by fungi, thereby limiting aphid population growth. The results indicate that application of nitrogen increases natural populations of M. dirhodum, and under favourable conditions, populations of S. avenae. However, in suboptimal climatic conditions, the application of nitrogen fertiliser can lead to lower populations of 5. avenae. The data also suggest that there is no consistent difference between a conventional and Canopy Managed approach to nitrogen fertiliser use in terms of the risk of infestation by cereal aphids.     

Preference of the Aphid Parasitoid Monoctonus paulensis (Hymenoptera: Braconidae,Aphidiinae) for Different Aphid Species: Female Choice and Offspring Survival

Host choice by Monoctonus paulensis (Ashmead) (Hymenoptera: Braconidae, Aphidiinae) was examined in the laboratory. In dichotomous choice tests, females were provided with four kinds of hosts that were similar in size and equally available. They preferred second instars of Acyrthosiphon pisum (Harris) over third instars of Sitobion avenae (F.) and both over second instars of Macrosiphum creelii Davis and Metopolophium dirhodum (Walker) (Hemiptera: Aphidoidea: Aphididae). Host choice was influenced by differences in aphid size. First instars of M. creelii and second instars of A. pisum were accepted at the same rate; first instars are smaller, have fewer defenses, and are easier to handle than second instars. Overall, A. pisum was the “best” host in terms of successful oviposition (76% of encountered aphids), eclosion from mummies (91%), and offspring sex ratio (72% females among offspring). Nearly 50% of encountered M. creelii escaped or were released by females without stinging, and 26% of encountered M. dirhodum survived stinging because either eggs were not laid or did not develop. Values for S. avenae were intermediate, with over 30% of encountered individuals escaping. Attacks on M. dirhodum and S. avenae, with or without oviposition, incurred lower time costs (17 s) than attacks on A. pisum and M. creelii (23 s). Relative to A. pisum (value = 1.0), a female's fitness returns in terms of offspring produced per encountered aphid were 0.75 for S. avenae, 0.36 for M. creelii, and 0.33 for M. dirhodum; her fitness costs in terms of time invested per offspring produced were 0.99 for S. avenae, 0.44 for M. dirhodum, and 0.35 for M. creelii. The ranked order of aphid species as reflected in host choice by females of M. paulensis was qualitatively the same as host order based on estimated fitness returns.     

The effect of barley yellow dwarf virus on field populations of the cereal aphids, Sitobion avenae and Metopolophium dirhodum

The numbers of cereal aphids, especially Metopolophium dirhodum in 1979, and Sitobion avenae in 1980, were significantly increased on BYDV infected wheat and oats in 1979, and wheat, barley and oats in 1980. The differences were probably caused by attraction of alates of each species to virus infected plants which had changed colour as a result of their infection. Significantly more alates of M. dirhodum were found on virus infected oats in 1979, and of S. avenae on oats and barley in 1980, although not on wheat in either year. probably because the colour contrast in wheat was less intense than in the other crops. Flight chamber experiments with alates of both species confirmed their visual attraction to virus-infected leaves. The interaction between virus, vector and host plants is discussed with reference to the ecology of virus spread.     

Some factors affecting the build-up of cereal aphid infestations in winter wheat

Aphid populations were measured at 14-day intervals during May to July 1974 in 17 fields of wheat at 10 sites along an 85 km transect running east from Hertfordshire to the Essex coast. Metopolophium dirhodum was the most numerous species initially, but was later superseded by Sitobion avenae. Rhopalosiphum padi was third in order of abundance. Several characteristics of each site and crop were examined in relation to the development of aphid infestations: two were associated with inter-site differences in aphid infestation. Infestations started earlier at the eastern than at the western sites due to the earlier arrival of alate migrants, and this resulted in higher peak populations densities of R. padi but not of M. dirhodum or S. avenae. Greater aphid densities were associated with crops having low nitrate concentrations in plant sap in the early stages of growth.     

The spatial distribution of canopy-resident and ground-resident cereal aphids (Sitobion avenae and Metopolophium dirhodum) in winter wheat

We investigated, within two cereal fields in Southern England, the within-canopy spatial distribution of the aphids Sitobion avenae and Metopolophium dirhodum in relation to crop yield and plant nitrogen. We extended the study to investigate the spatial distribution of aphids that fell to, or returned from, the ground in order to estimate availability of the within-canopy aphid population to ground-active predators. We revealed that crop canopy aphid spatial pattern was associated with nitrogen or yield. Differences were evident between species: S. avenae was generally negatively associated with yield or plant nitrogen, whilst M. dirhodum exhibited positive association. For both aphid species, we observed strong spatial pattern for aphids falling to the ground and conclude that this could, in part, mediate the effectiveness of ground-active predators as pest control agents.     

The value of three cereal aphid species as food for a generalist predator

The value of the cereal aphid species Metopolophium dirhodum (Wlk.), Sitobion avenae (F.) and Rhopalosiphum padi (L.) as prey for the linyphiid spider Erigone atra (Bl.) was assessed. Fecundity of females was determined for spiders fed on eight experimental diets: three single‐species aphid diets, a mixed diet of all three aphid species, three mixed diets with each aphid species in combination with fruit flies Drosophila melanogaster (Meig.), and pure D. melanogaster as a high quality comparison diet. The development and survival of first‐instar juveniles fed on three diets of single aphid species, and on a diet of Collembola were compared with those subjected to starvation. Prey value for adult females was assessed by egg production, hatching success and offspring size. In pure diets all three aphid species were of low value to the spiders, causing a rapid decline in egg production and supporting no growth of significance of first‐instar juveniles. No difference in value of aphid species of single‐species aphid diets was found in the fecundity experiment, while a ranking of aphid species of M. dirhodum > R. padi > S. avenae was revealed in the survivorship experiment. A mixed‐aphid diet was not found to be advantageous compared with single‐species aphid diets, and no advantage of including aphids in mixed diets with fruit flies was found. Metopolophium dirhodum and R. padi were neutral in mixed diets, while a diet of S. avenae and fruit flies caused reduced egg production compared with the pure diet of fruit flies, revealing a toxic effect of S. avenae on the spider. The value‐ranking of aphid species in mixed diets was similar to that of single‐species diets. A similar ranking of aphid species was found for different fitness parameters (fecundity of adult females and development of juveniles). A ranking of aphids by offspring size of mothers on aphid‐only diets was S. avenae > M. dirhodum > R. padi. All aphid‐fruit fly diets resulted in larger offspring than a diet of only D. melanogaster, with the overall largest offspring being produced on the diet of M. dirhodum and fruit flies.