Influence of crop background on aphids and other phytophagous insects on Brussels sprouts

More Brevicoryne brassicae and other alate aphids were caught in yellow water-traps in a weed-free crop of Brussels sprouts than in a crop with a weedy background. More B. brassicae colonized Brussels sprout plants in bare soil than in weeds; very few alatae were attracted to cruciferous weeds in the crop. Results in 1 yr suggest that initially larger populations on the weed-free sprouts became smaller than populations on the weedy sprouts because the larger aphid population attracted more natural enemies. Aleyrodes brassicae and certain Lepidoptera were also more abundant on sprout plants in bare soil than on sprouts surrounded by weeds; more adult A. brassicae were caught in water traps over the bare soil. More A. brassicae were present on sprout plants surrounded by a green than by a brown cloth background but the differences were not significant (P < 0–05). Numbers of B. brassicae on sprout plants with green and brown backgrounds varied greatly and did not differ significantly. In field cages, B. brassicae alatae were more attracted to potted sprout plants surrounded by bare soil than to ones surrounded by rings of living or cut grass or by artificial green rings. This effect was greater with small than with large sprout plants surrounded by grass rings. The maintenance of a limited weed cover is considered potentially useful in integrated control of some brassica pests.     

Seasonal changes in aphid populations on Brussels sprouts

Populations of Brevicoryne brassicae (L.) and Myzus persicae (Sulz.) were assessed weekly on Brussels sprouts at Wellesbourne, from July onwards in each of 3 years. A peak of B. brassicae numbers occurred every year about the end of July, and in 1968 and 1970 this was larger than the September peak. Some alates were produced in the generation born to the June immigrants and the dispersal of those produced at the end of July consistently brought a decline in B. brassicae populations in early August. In 1968 and 1970, however, outbreaks of entomogenous fungi which followed rain, added considerably to this decline. Records of predators and parasites indicated that these natural enemies had only a minor effect on aphid numbers. M. persicae colonized sprouts slightly earlier than B. brassicae; in 1969 and 1970 its numbers in July were low and some persisted after the first week of August in 1968 only. In each year the first eggs of B. brassicae were laid at the beginning of October, but viviparae were still numerous for a further 7 to 10 weeks.     

Influence of crop background on natural enemies of aphids on Brussels sprouts

The most abundant natural enemies of aphids on Brussels sprout crops were Syrphidae, different species being attracted differentially to weedy or weed-free plots according to whether they were more host-plant-orientated (e.g. Melanostoma spp., Platycheirus spp.) and thus affected directly by the background, or more aphid-orientated (e.g. Syrphus balteatus) and so less affected by background than by aphid numbers. Oviposition by Melanostoma spp. was usually much greater in weedy sprout crops than on sprouts in bare soil, and their eggs were also very abundant on weeds. Eggs of other syrphid species were scarcer on weeds. In contrast to Melanostoma, Platycheirus spp. usually oviposted preferentially on sprouts growing in bare soil. Oviposition by S. balteatus was in response to aphid abundance and thus tended to become greater on sprouts in bare soil. Notably more adults of non-aphido-phagous Syrphidae were caught over weedy than over non-weedy Brussels sprout plants. Anthocoris nemorum nymphs and adults were very common on sprout plants and weeds in the weedy crop but were scarce on sprouts in bare soil; A. nemorum oviposited on white and yellow charlock (Raphanus raphanistrum and Sinapis arvensis) occasionally. Parasitism of Brevicoryne brassicae by Diaeretiella rapae appeared to be related to aphid numbers and was only indirectly influenced by the crop background. Field experiments with green and brown cloth backgrounds showed that some syrphids were attracted to green; A. nemorum was relatively scarce over both artificial backgrounds. It is concluded that decreased natural enemy action is partly responsible for the initially greater abundance of B. brassicae in a weed-free crop of Brussels sprouts than in a weedy crop.     

The effect of leaf excision on the performance of Myzus persicae and Brevicoryne brassicae in relation to the nutrient treatment of the plants

Two groups of brussels sprout plants of one variety given different fertilizer treatments proved ‘resistant’ and ‘susceptible’ to both Myzus persicae and Breicoryrie brassicae, as measured by aphid mean relative growth rate. The hypothesis that leaf disc would fail to reflect differences in ‘nutritionallybased’ plant resistance was confirmed with B.brussicue but not M.persicae. Both aphids grew more slowly on discs than leaves of ‘susceptible’ plants; the reverse tendency showed on ‘resistant’ material. Total:(, amino nitrogen proved poorly correlated with aphid performance. Previously published (van Emden & Bashford, 1971) equations relating individual amino acids to aphid performance gave a better fit to the results, but still failed to account for the reduced performance of the aphids on ‘susceptible’ discs.     

Varietal differences in the susceptibility of Brussels sprouts to lepidopterous pests

The attack by cabbage caterpillars on open-pollinated cultivars of Brussels sprouts at Wellesbourne was recorded approximately weekly from July to October in 1973 and 1974. The red-foliaged cv. Rubine was much less attacked by Pieris rapae than any of the twenty-five green-foliaged cultivars tested, the least attacked of which were The Aristocrat, Continuity, Evergreen and Pride of the Market. Rubine also showed some resistance to attack by Mamestra brassicae and Evergestis forficialis, as did The Aristocrat. Continuity, though as resistant as any of the cultivars to E. forficialis, was not resistant to M. brassicae. No evidence of antibiosis was found and resistance, based on non-preference, is considered to result from ovipositing P. rapae adults discriminating against the red colour of Rubine and against the leaf composition (relative volatile content) of the green-foliaged resistants. This could also apply with M. brassicae and E. forficialis.     

Resistance of rape (Brassica napus) to attack by the cabbage aphid (Brevicoryne brassicae L.)

Resistance to Brevicoryne brassicae L. attacks in a New Zealand variety of forage rape resulted from a combination of host non–preference and antibiosis. Using clonal plant material obtained from cuttings, only half the numbers of immigrant alate cabbage aphids that settled to reproduce on the susceptible rape were to be found on the resistant rape. The reproduction rate of these alates was about 12 % slower on the resistant plants than on the susceptibles and the young took about 13% longer to mature. Antibiosis then shortened the reproductive life of the apterae by one-third, reduced their fecundity by nearly 50 % and caused 40% mortality in their progeny. The over-all effect of this was considerable and under conditions specified in the text could result in the population on the resistant plants being about one-eighth that on the susceptible plants in under 1 month. The resistant plants lost their resistance on flowering and became normally susceptible to cabbage aphid attack. With maturity, however, the B. brassicae-resistant plants became slightly resistant to Myzus persicae.     

Study on inheritance of resistance to root aphid, Pemphigus bursarius, in lettuce

The inheritance of resistance to lettuce root aphid, Pemphigus bursarius, was studied in lettuce using the Wellesbourne cultivars Avondefiance and Avoncrisp as resistant parents and Borough Wonder and Webb's Wonderful as aphid-susceptible parents. All four cultivars were crossed in all possible combinations including reciprocals and the response to root aphid of plants in the P₁F₁F₂ and BC generations was assessed using apterae of P. bursarius from the lettuce cv. Iceberg. Resistance to attack was clearly inherited and the parents appeared to be homozygous for their resistance or susceptibility. In the F₁ generation, however, in all crosses between resistant and susceptible parents, segregation into susceptible, resistant and some slightly less resistant plants occurred. This and the highly significant differences in segregation between pairs of reciprocal crosses in the F₁ and other generations indicate that the inheritance of resistance to root aphid is controlled by extra-nuclear factors. Modifying genes might also be involved but there appears to be no linkage of root aphid resistance with resistance to downy mildew, for which the Wellesbourne lettuces were bred.     

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.     

Falling aphids enhance impact of biological control by parasitoids on partially aphid-resistant plant varieties

Cage experiments, in which the population increase of Metopolophium dirhodum was measured either in the absence or presence of the parasitoid Aphidius rhopalosiphi, showed reductions of peak populations in the presence of the parasitoid of 30% on the susceptible wheat cultivar Armada, but of 57% on the partially resistant Rapier. On both cultivars, aphid population curves in the presence and absence of the parasitoid diverged before the first mummies were formed. The number of aphids leaving plants was measured, and was found to increase in the presence of the parasitoid, particularly with Rapier (almost double). Most aphids (about 75%) which left plants fell, and a smaller proportion regained plants when parasitoids were present than when they were absent. A field experiment with Brevicoryne brassicae and natural predation on Brussels sprouts also showed an increase in aphids leaving a less susceptible compared with a more susceptible cultivar.     

Variation to cause host injury between Russian wheat aphid (Homoptera: Aphididae) clones virulent to Dn4 wheat

Biotypes are infraspecific classifications based on biological rather than morphological characteristics. Cereal aphids are managed primarily by host plant resistance, and they often develop biotypes that injure or kill previously resistant plants. Although molecular genetic variation within aphid biotypes has been well documented, little is known about phenotypic variation, especially virulence or the biotype's ability to cause injury to cultivars with specific resistance genes. Five clones (single maternal lineages) of Russian wheat aphid, Diuraphis noxia (Kurdjumov) (Homoptera: Aphididae), determined to be injurious to wheat, Triticum aestivum L., with the Dn4 gene, were evaluated on resistant and susceptible wheat and barley, Hordeum vulgare L., for their ability to cause chlorosis, reduction in plant height, and reduction in shoot dry weight. Variation to cause injury on resistant 'Halt' wheat, susceptible 'Jagger' wheat, and resistant 'STARS-9301B' barley was found among the Dn4 virulent clones. One clone caused up to 30.0 and 59.5% more reduction in plant height and shoot dry weight, respectively, on resistant Halt than other clones. It also caused up to 29.9 and 55.5% more reduction in plant height and shoot dry weight, respectively, on susceptible Jagger wheat. Although STARS-9301B barley exhibited an equal resistant response to feeding by all five clones based on chlorosis, two clones caused approximately 20% more reduction in plant height and shoot dry weight than three other clones. The most injurious clones on wheat were not the most injurious clones on barley. This is the first report of variation to cause varying degrees of plant damage within an aphid biotype virulent to a single host resistance gene. A single aphid clone may not accurately represent the true virulent nature of a biotype population in the field.     

Population studies ofBrevicoryne brassicae (L.), its parasites and hyperparasites in England

Studies on populations of Brevicoryne brassicae (L.), its parasites and hyperparasites were carried out by actual counting in the sprouts field and by sticky and water traps. B. brassicae was found to be attacked by one primary parasite, Diaretiella rapae (McIntosh ), which in turn is parasitized by Alloxysta brassicae (Ashm. ),Asaphes vulgarisWalker ,A. suspensus (Nees ),Pachyneuron minutissimum (Förster ) and Dendrocerus carpenterii (Curtis ). The aphid population in the field was started by immigrant alates which were found flying too early to be synchronized with the sprouts plants. Similarly D. rapae was not synchronized with the aphids although many individuals could have been carried into a plot through parasitized immigrant alates, of which less than 30% were found parasitized. Because of high hyperparasitism (especially by A. brassicae)D. rapae was not able to maintain a high rate of parasitism to curb the aphid population growth. The maximum percentage mummies being 27.8%, while the maximum, percentage parasitism being 56.6% recorded only during the early 1974 season (mean=12.9%). The decline of aphid population from September onwards was largely due to the cold weather, Syrphid predation and occasionally fungal attack. The high rate of hyperparasitism by A. brassicae is attributed to its better synchronization with D. rapae. The mean percentage of parasite that emerged from mummies collected during 1973–74 were D. rapae 31.3%,A. brassicae 64.3%,A. vulgaris and A. suspensus 4.3%,D. carpenterii 0.2% and P. minutissimum 0.1%.     

Associational resistance or susceptibility: the indirect interaction between chemically‐defended and non‐defended herbivore prey via a shared predator

Many organisms possess chemical defences against their natural enemies, which render them unpalatable or toxic when attacked or consumed. These chemically‐defended organisms commonly occur in communities with non‐ or less‐defended prey, leading to indirect interactions between prey species, mediated by natural enemies. Although the importance of enemy‐mediated indirect interactions have been well documented (e.g. apparent competition), how the presence of prey chemical defences may affect predation of non‐defended prey in terrestrial communities remains unclear. Here, an experimental approach was used to study the predator‐mediated indirect interaction between a chemically‐defended and non‐defended pest aphid species. Using laboratory‐based mesocosms, aphid community composition was manipulated to include chemically‐defended (CD) aphids Brevicoryne brassicae, non‐defended (ND) aphids Myzus persicae or a mixed assemblage of both species, on Brassica oleracea cabbage plants, in the presence or absence of a shared predator (Chrysoperla carnea larvae). Aphid population growth rates, aphid distributions on host plants and predator growth rates were measured. In single‐species treatments, C. carnea reduced M. persicae population growth rate, but had no significant impact on B. brassicae population growth rate, suggesting B. brassicae chemical defences are effective against C. carnea. Chrysoperla carnea had no significant impact on either aphid species population growth rate in mixed‐species treatments. Myzus persicae (ND) therefore experienced reduced predation in the presence of B. brassicae (CD) through a predator‐mediated indirect effect. Moreover, predator growth rates were significantly higher in the M. persicae‐only treatments than in either the B. brassicae‐only or mixed‐species treatments, suggesting predation was impaired in the presence of B. brassicae (CD). A trait‐mediated indirect interaction is proposed, consistent with associational resistance, in which the predator, upon incidental consumption of chemically‐defended aphids is deterred from feeding, releasing non‐defended aphids from predatory control.     

An investigation of the resistance to cabbage aphid in brassica species

Fifteen Brassica species and subspecies, as well as accessions of Arabidopsis thaliana and Eruca sativa, were evaluated for their resistance to the cabbage aphid, Brevicoryne brassicae, in the field and laboratory at Horticulture Research International (HRI) Wellesbourne in 1992. In the laboratory, aphid performance was measured in terms of pre-reproductive period, reproductive period, population increase and insect survival. Using these parameters it was possible to identify brassicas possessing antibiosis resistance. In the field B. brassicae attack was assessed in terms of numbers of insects and it was possible to determine aphid preferences for alighting on different species. High levels of antixenosis and antibiosis resistance were discovered in accessions of Brassica fruticulosa and B. spinescens, in a Brassica juncea breeding line and in Eruca sativa. Partial resistance was found in several other Brassica species. The potential of these various sources as donors of resistance genes to be bred into cultivated brassicas is discussed.     

Feeding behavior of Brevicoryne brassicae in resistant and susceptible collard greens genotypes: interactions among morphological and chemical factors

The cabbage aphid, Brevicoryne brassicae (L.) (Hemiptera: Aphididae), is distributed throughout the tropical and subtropical areas of the world. The main crops attacked by B. brassicae are cabbage, collard greens, broccoli, Brussels sprouts, and cauliflower. To survive the attack of pest insects, plants have evolved various resistance mechanisms that may affect pest feeding behavior. The use of electronic monitoring through EPG (electrical penetration graph) can help characterize and distinguish the resistance mechanisms involved. This study evaluated the feeding behavior of B. brassicae in eight genotypes of collard greens, Brassica oleraceae L. var. acephala (Brassicaceae), exhibiting antixenosis and/or antibiosis resistance to this insect. Possible correlations were established between the glucosinolate levels, the hardness, and the epicuticular wax on the leaves vs. aphid feeding behavior. On the genotypes 22V, 5E, and 27VA, for which many ‘potential drop’ waves were performed, aphid development was slower, indicating antixenosis as resistance type. Aphids on the genotypes 22V and 24X required more time until accessing the phloem, also suggesting antixenosis as resistance category. Genotypes 22V and PE had hard leaves, which also points at antixenosis. Genotypes 20T and HS had higher total wax and wax mg⁻¹. Feeding parameters on ARI and 24X were similar to those observed on HS; antibiosis is likely to be the predominant resistance category of this germplasm. Because HS was considered as a susceptible standard genotype in this study, a higher gluconapin amount indicates that this compound does not influence cabbage aphid feeding behavior. The present study confirms that analysis of the physical and chemical aspects of collard greens genotypes by the EPG technique can provide a useful approach for the study of plant resistance to cabbage aphids.     

Population growth rate and relative virulence of the two South African biotypes of Russian wheat aphid,Diuraphis noxia,and bird cherry‐oat aphid,Rhopalosiphum padi,on resistant and non‐resistant barley

In South Africa a new biotype of the Russian wheat aphid (RWA), Diuraphis noxia (Kurdjumov) (Hemiptera: Aphididae), RWASA2, has appeared which exhibits an improved performance compared to the original biotype (RWASA1) on wheat containing the Dn1 resistance gene. We examined population growth rates as well as damage caused by RWASA1 and RWASA2, in addition to a different aphid species, the bird cherry‐oat aphid (BCA), Rhopalosiphum padi L. (Hemiptera: Aphididae), on three RWA‐resistant barley [Hordeum vulgare L. (Poaceae)] lines (STARS‐9577B, STARS‐0502B, and STARS‐9301B) and one susceptible control (PUMA). RWASA2 had a higher reproductive rate than RWASA1 on all barley lines tested, which is consistent with previous results on wheat. Two of the RWA‐resistant lines (STARS‐0502B and STARS‐9301B) also exhibited a similar resistance phenotype against BCA. In our experiments, severe chlorosis and leaf roll appeared earlier on the control PUMA barley variety as a result of RWASA2 feeding than was the case with RWASA1, probably due to the differences in reproductive rate. Although chlorosis appeared earlier on resistant plants after RWASA2 feeding, this symptom developed much faster during RWASA1 feeding on all three resistant lines tested. As chlorosis did not correlate well with aphid population numbers, we surmise that the differential chlorosis effects may be related to differences in the amount of saliva introduced by the two aphid clones during feeding. Our results indicate that the difference between RWASA2 and RWASA1 are broader than a ‘gene for gene’ interaction with the Dn1 resistance (R) gene in wheat, and that these biotypes also differ in important aspects of their biology.     

The influence of genetic background on resistance to the cabbage aphid (Brevicoryne brassicae) in Kale (Brassica oleracea var. acephala)

Resistance to Brevicoryne brassicae has been identified in the progeny of two selected kale (B. oleracea var. acephala) plants, one from the F1 hybrid cultivar ‘Arsis RS’ and one from the landrace ‘Butzo’. These plants were crossed with susceptible B. oleracea morphotypes that have different periods to flowering. The type of susceptible plant line used had an effect on the resistance phenotypc of the progeny. Tested F2 populations derived from these crosses show that resistance is not under simple genetic control. This, in addition to variation in aphid numbers within accessions, suggests that separation of genetic components of control from environmental ‘noise’ for any accession may only be possible by the production of double haploid plant lines.     

Genetic mapping revealed two loci for soybean aphid resistance in PI 567301B

The soybean aphid (Aphis glycines Matsumura) is the most damaging insect pest of soybean [Glycine max (L.) Merr.] in North America. New soybean aphid biotypes have been evolving quickly and at least three confirmed biotypes have been reported in USA. These biotypes are capable of defeating most known aphid resistant soybean genes indicating the need for identification of new genes. Plant Introduction (PI) 567301B was earlier identified to have antixenosis resistance against biotype 1 and 2 of the soybean aphid. Two hundred and three F_7:9 recombinant inbred lines (RILs) developed from a cross of soybean aphid susceptible cultivar Wyandot and resistant PI 567301B were used for mapping aphid resistance genes using the quantitative trait loci (QTL) mapping approach. A subset of 94 RILs and 516 polymorphic SNP makers were used to construct a genome-wide molecular linkage map. Two candidate QTL regions for aphid resistance were identified on this linkage map. Fine mapping of the QTL regions was conducted with SSR markers using all 203 RILs. A major gene on chromosome 13 was mapped near the previously identified Rag2 gene. However, an earlier study revealed that the detached leaves of PI 567301B had no resistance against the soybean aphids while the detached leaves of PI 243540 (source of Rag2) maintained aphid resistance. These results and the earlier finding that PI 243540 showed antibiosis resistance and PI 567301B showed antixenosis type resistance, indicating that the aphid resistances in the two PIs are not controlled by the same gene. Thus, we have mapped a new gene near the Rag2 locus for soybean aphid resistance that should be useful in breeding for new aphid-resistant soybean cultivars. Molecular markers closely linked to this gene are available for marker-assisted breeding. Also, the minor locus found on chromosome 8 represents the first reported soybean aphid-resistant locus on this chromosome.     

Maternal host plant effects on aphid performance: contrasts between a generalist and a specialist species on Brussels sprout cultivars

• 1 The performance of the second generation (G₂) of alates and apterae of a generalist, Myzus persicae, and a specialist, Brevicoryne brassicae, aphid species reared on Chinese cabbage or cabbage was evaluated on five cultivars of Brussels sprout.
• 2 Aphid performance was influenced both by the type of host on which the parent aphid had been reared and by the host on which it was feeding when reproducing.
• 3 The fecundity of the G₂ of alates of both aphid species reared on Chinese cabbage differed significantly between all the cultivars of Brussels sprout and, on average, was 25% higher than those reared on cabbage. These differences were also apparent for the intrinsic rate of increase of B. brassicae but not for M. persicae.
• 4 There was a trend for the G₂ of alates from Chinese cabbage to have greater fecundity compared with aphids from cabbage. These differences were significant for the fecundity of the G₂ of alates of both aphid species on Brussels sprout cultivars Fillbasket (30% higher), Red Delicious (35% higher) and Winter Harvest (25% higher) than those reared on cabbage.
• 5 The intrinsic rate of increase for the G₂ of alates of B. brassicae from Chinese cabbage was significantly different on all Brussels sprout cultivars tested. The intrinsic rate of increase differed significantly between aphids reared on either Chinese cabbage or cabbage on cultivars Oliver and Darkmar‐21 (M. persicae) and Red Delicious and Winter Harvest (B. brassicae). The cv. Oliver appeared to be the most consistently good host; Red Delicious was the poorest host overall.

     

Effectiveness of resistance genes to the large raspberry aphid, Amphorophora idaei Börner, in different raspberry (Rubus idaeus L.) genotypes and under different environmental conditions

The introduction into commerce of raspberry cultivars with major gene resistance to the large raspberry aphid, Amphorophora idaei, an important pest and virus vector on red raspberry in Europe, has been very effective both in decreasing pest numbers and greatly restricting infection with the viruses it transmits. However, biotypes of the aphid able to overcome these genes have developed in the field in recent years. Additionally, in field and laboratory tests, the response to aphid biotypes and recognised aphid strains of certain raspberry cultivars, such as Glen Prosen and Delight, differ markedly despite the fact that they are reputed to contain the same A. idaei‐resistance gene, A₁. In attempts to understand the reasons for this difference in response, analysis was made of the segregation of progeny seedlings from crosses between A. idaei‐resistant and ‐susceptible cultivars to two recognised strains of the aphid. These studies showed that, as expected, cv. Autumn Bliss contained the A. idaei‐resistance gene, A₁₀, and cvs Delight and Glen Prosen each contained the A. idaei‐resistance gene, A₁. When progeny seedlings were assayed in a heated glasshouse as young plants and in an unheated Tygan house as 1 m tall plants, the segregation ratios for resistance and susceptibility to A. idaei were largely unchanged. However, when the resistance of individual progeny plants was assessed, c. 37% of the putative gene A₁‐containing progeny and 9–23% of the putative gene A₁₀‐containing progeny, behaved differently in these two environments. Experiments involving an A. idaei‐resistant and ‐susceptible parent cultivar showed that shading plants increased their susceptibility to A. idaei colonisation. Whilst this shading effect has implications for experimentally detecting A. idaei‐resistant progeny in segregating raspberry seedlings, it does not explain the difference in field resistance to A. idaei of cvs Delight and Glen Prosen. Such differences in the field seem best explained by the presence in these cultivars of ‘minor’ genes for A. idaei resistance and/or susceptibility that influences the effectiveness of gene A₁.     

VARIETAL RESISTANCE OF LETTUCE TO ATTACK BY THE LETTUCE ROOT APHID, PEMPHIGUS BURSARIUS (L.)

Striking varietal differences in susceptibility to attack by the lettuce root aphid, Pemphigus bursarius L., were first found in lettuces grown at Wellesbourne in 1955. Subsequent work has confirmed that White Favourite and Imperials E-4 and 19551-M are highly resistant; Imperial 45634-M, Continuity and Iceberg are markedly resistant. Midas, Salad Bowl and Imperial 4164 each appear to be a mixture of susceptible and resistant plants.
Immigrant winged forms of P. bursarius showed no preference for colonizing any particular variety of lettuce, and it seems that resistance to attack results from antibiosis.
It is suggested that varietal differences in the composition of root sap may account for the differences in susceptibility to lettuce root aphid, but analyses of water and alcohol extracts of root sap from resistant and susceptible lettuce varieties have not shown consistent differences.