Aphid clonal resistance to a parasitoid fails under heat stress

Parasitoid virulence and host resistance are complex interactions depending on metabolic rate and cellular activity, which in aphids additionally implicate heritable secondary symbionts among the Enterobacteriaceae. As performance of the parasitoid, the aphid host and its symbionts may differentially respond to temperature, the success or failure of aphid parasitism is difficult to predict when temperature varies. We tested the hypothesis that resistance of the pea aphid Acyrthosiphon pisum to the parasitoid Aphidius ervi, which is linked to aphid secondary symbionts, may depend on temperature in several resistant and non-resistant aphid clonal lineages of different geographic origin and of known bacterial symbiosis, using experiments in controlled environments. Complete immunity to A. ervi at 20 degrees C in three different aphid clones whose symbiosis is characterized by the possession of Hamiltonella defensa reversed to high susceptibility at 25 degrees C and especially 30 degrees C, suggesting that the aphid's immune responses to the establishment and early development of the parasitoid is strongly reduced at moderately high temperatures. There was no evidence that a pea aphid control genotype that was susceptible to A. ervi at 20 degrees C could become more resistant as temperature increases, as has been suggested for insect fungal pathogens. By contrast, our results suggest that aphid clonal resistance to A. ervi and related parasitoids is characteristic of cool temperature conditions, similar to various other fitness attributes of aphids. Based on evidence that H. defensa symbionts characterized all three A. ervi resistant pea aphid clones studied, but was absent in control aphids that remained susceptible at all temperatures, we suggest that secondary symbiosis plays a key role in the heat sensitivity of aphid clonal resistance. Our study may also indicate that aphid natural control of variably susceptible host populations by aphid parasitoids is more likely at moderate to high temperatures.     

Impact of Rag1 aphid resistant soybeans on Binodoxys communis (Hymenoptera: Braconidae), a parasitoid of soybean aphid (Hemiptera: Aphididae)

Multiple strategies are being developed for pest management of the soybean aphid, Aphis glycines Matsumura; however, there has been little published research thus far to determine how such strategies may influence each other, thereby complicating their potential effectiveness. A susceptible soybean (Glycine max L.) variety without the Rag1 gene and a near isogenic resistant soybean variety with the Rag1 gene were evaluated in the laboratory for their effects on the fitness of the soybean aphid parasitoid, Binodoxys communis (Gahan). The presence or absence of the Rag1 gene was verified by quantifying soybean aphid growth. To test for fitness effects, parasitoids were allowed to attack soybean aphids on either a susceptible or resistant plant for 24 h and then aphids were kept on the same plant throughout parasitoid development. Parasitoid fitness was measured by mummy and adult parasitoid production, adult parasitoid emergence, development time, and adult size. Parasitoids that attacked soybean aphids on susceptible plants produced more mummies, more adult parasitoids, and had a higher emergence rate compared with those on resistant plants. Adult parasitoids that emerged from resistant plants took 1 d longer and were smaller compared with those from susceptible plants. This study suggests that biological control by B. communis may be compromised when host plant resistance is widely used for pest management of soybean aphids.     

Strong parasitoid-mediated selection in experimental populations of aphids

Clonal diversity in asexual populations may be maintained if different clones are favoured under different environmental conditions. For aphids, parasitoids are an important variable of the biotic environment. To test whether parasitoids can mediate selection among host clones, we used experimental populations consisting of 10 clones of the peach-potato aphid, Myzus persicae, and allowed them to evolve for several generations either without parasitoids or in the presence of two species of parasitoid wasps. In the absence of parasitoids, strong shifts in clonal frequencies occurred, mostly in favour of clones with high rates of increase. The parasitoid Diaeretiella rapae hardly affected aphid densities but changed the outcome of competition by favouring one entirely resistant clone and disfavouring a highly susceptible clone. Aphidius colemani, the more infective parasitoid, strongly reduced aphid densities and dramatically changed host clonal frequencies. The most resistant clone, not a successful clone without parasitoids, became totally dominant. These results highlight the potential of temporal or spatial variation in parasitoid densities to maintain clonal diversity in their aphid hosts.     

Combined effects of host-plant resistance and intraguild predation on the soybean aphid parasitoid Binodoxys communis in the field

Soybean varieties that exhibit resistance to the soybean aphid Aphis glycines have been developed for use in North America. In principle, host-plant resistance to soybean aphid can influence the interactions between the soybean aphid and its natural enemies. Resistance could change the quality of soybean aphids as a food source, the availability of soybean aphids, or resistance traits could directly affect aphid predators and parasitoids. Here, we focus on the effect of soybean aphid resistance on the interactions between soybean aphids, the parasitoid Binodoxys communis (Hymenoptera: Braconidae), and predators of these two species. We determined whether host-plant resistance affected within-season persistence of B. communis by releasing parasitoids into resistant and susceptible soybean plots. We observed higher B. communis densities in susceptible soybean plots than in resistant plots. There were also higher overall levels of intraguild predation of B. communis in susceptible plots, although the per-capita risk of intraguild predation of B. communis was affected neither by plant genotype nor by aphid density. We discuss these effects and whether they were caused by direct effects of the resistant plants on B. communis or indirect effects through soybean aphid or predators.     

Effect of cabbage cultivars with varying levels of resistance to aphids on the performance of the parasitoid,Aphidius colemani (Hymenoptera: Braconidae)

The effect of aphid resistance in plant cultivars on parasitism of Myzus persicae (Sulzer) and Brevicoryne brassicae (Linnaeus) by Aphidius colemani Viereck was investigated under laboratory conditions using three cultivars of common cabbage, Brassica oleracea var. capitata. Significantly greater aphid populations were found on cv. Derby Day (green-leaved, susceptible to both aphid species), regardless of the presence or absence of parasitoids, compared with cv. Minicole (green-leaved, partially resistant with antibiosis factors for B. brassicae) or cv. Ruby Ball (red-leaved, partially resistant with antixenosis factors for B. brassicae). Minicole had the greatest proportion of parasitized aphids and Derby Day the least. A significantly lower percentage of emerged parasitoids were recorded on Minicole for both aphid species compared with Derby Day and Ruby Ball. The implication of the results are discussed in relation to the integrated control of aphids.     

Interactions of Russian wheat aphid, a hymenopterous parasitoid and resistant and susceptible slender wheatgrasses

Interactions among three trophic levels of resistant and susceptible slenderwheat grasses, Elymus trachycaulum (Link) Goule ex Shinners ex. H.F. Lewis, Russian wheat aphid, Diuraphis noxia (Mordvilko), and a hymenopterous parasitoid were studied in the laboratory and greenhouse. These relationships were compared with a commercial susceptible wheat Triticum aestivum L. variety. Aphids reared on the resistant entries showed significantly lower weights and numbers. Significant reduction of parasitoid mummy weight and adult size was positively correlated with the effects on the aphids. Resistant entries also induced a longer prereproductive period for both the aphids and parasitoids. Numbers of aphids and aphid damage were significantly modified by the addition of parasitoids. Parasitism was higher on plants that did not have leaf rolling. These findings may indicate that antibiosis resistance studied here is not the most desirable because it decreases natural enemy vitality.     

The impact of Meu1-mediated resistance in tomato on longevity, fecundity and behavior of the potato aphid, Macrosiphum euphorbiae

The effect of the tomato resistance gene, Meu1, on feeding, longevity, fecundity and developmental rate of the pink biotype of the potato aphid, Macrosiphum euphorbiae (Thomas) (Hemiptera, Aphididae), was determined using nearly isogenic tomato (Lycopersicon esculentum Mill, Solanaceae) lines. Aphid mortality was significantly higher on resistant plants, with 60% of the aphids dying by the 4th day of exposure. By the 10th day, all the aphids on the resistant plants were dead whereas 100% of the aphids on susceptible plants were alive. Meu1-mediated resistance resulted in significantly decreased fecundity with a ten-fold decrease in the net fertility rate (4.5 and 45.7 progeny per aphid on resistant and susceptible tomato, respectively). A qualitative analysis showed that honeydew was produced by aphids on resistant and susceptible plants, suggesting that aphids initiate feeding on both plant types. However, significantly lower quantities of honeydew were present when aphids were caged on resistant plants. There were also significant differences in aphid location on resistant and susceptible leaves. Experiments evaluating behavior in less than 24 h showed that aphids left resistant leaves after relatively short exposure (3–6 h). Aphids transferred from resistant to susceptible tomato at intervals between 3 h and 24 h resumed feeding as evidenced by presence of honeydew. Although the mechanism by which Meu1-mediated resistance operates is not yet known, our data suggest that resistance factors act rapidly after initiation of feeding and that lower fecundity and longevity are related to reduction in aphid feeding.     

Responses of Schizaphis graminum (Homoptera: Aphididae) to leaf excision in resistant and susceptible sorghum

Greenbugs, Schizaphis graminum (Rondani), were reared on intact and excised leaves of varieties of sorghum which differed in their suitability as hosts for this aphid. Aphids grew poorly on intact leaves of three resistant varieties, but grew well on excised leaves of the same varieties. Leaf excision did not affect aphid growth on three susceptible varieties. By electronically monitoring the feeding behaviour of aphids on two resistant and one susceptible variety, significant differences were found in many parameters between aphids assayed on excised vs. intact leaves of only the resistant varieties. Aphids on excised leaves of the resistant varieties, and on excised or intact leaves of the susceptible variety, made fewer probes to the phloem, spending more time ingesting from phloem during each probe, compared to aphids on intact resistant plants. There was a higher level of free amino acids in excised leaves of all varieties, but aphid growth and feeding behaviour improved as a result of excision only on resistant varieties. This observation, coupled with the fact that intact plants of all varieties have similar amino acid levels, indicates that these nutrients are not of primary importance in sorghum suitability to the greenbug. Other explanations for the aphids' responses to excised leaves are discussed.     

Effects of pea aphid secondary endosymbionts on aphid resistance and development of the aphid parasitoid Aphidius ervi: a correlative study

In order to reduce parasite‐induced mortality, hosts may be involved in mutualistic interactions in which the partner contributes to resistance against the parasite. The pea aphid, Acyrthosiphon pisum Harris (Hemiptera: Aphididae), harbours secondary bacterial endosymbionts, some of which have been reported to confer resistance against aphid parasitoids. Although this resistance often results in death of the developing parasitoid larvae, some parasitoid individuals succeed in developing into adults. Whether these individuals suffer from fitness reduction compared to parasitoids developing in pea aphid clones without symbionts has not been tested so far. Using 30 pea aphid clones that differed in their endosymbiont complement, we studied the effects of these endosymbionts on aphid resistance against the parasitoid Aphidius ervi Haliday (Hymenoptera: Braconidae: Aphidiinae), host–parasitoid physiological interactions, and fitness of emerging adult parasitoids. The number of symbiont species in an aphid clone was positively correlated with a number of resistance measurements but there were also clear symbiont‐specific effects on the host–parasitoid interaction. As in previous studies, pea aphid clones infected with Hamiltonella defensa Moran et al. showed resistance against the parasitoid. In addition, pea aphid clones infected with Regiella insecticola Moran et al. and co‐infections of H. defensa–Spiroplasma, R. insecticola–Spiroplasma, and R. insecticola–H. defensa showed reduced levels of parasitism and mummification. Parasitoids emerging from symbiont‐infected aphid clones often had a longer developmental time and reduced mass. The number of teratocytes was generally lower when parasitoids oviposited in aphid clones with a symbiont complement. Interestingly, unparasitized aphids infected with Serratia symbiotica Moran et al. and R. insecticola had a higher fecundity than unparasitized aphids of uninfected pea aphid clones. We conclude that in addition to conferring resistance, pea aphid symbionts also negatively affect parasitoids that successfully hatch from aphid mummies. Because of the link between aphid resistance and the number of teratocytes, the mechanism underlying resistance by symbiont infection may involve interference with teratocyte development.     

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.     

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.     

EVOLUTION OF AN APHID-PARASITOID INTERACTION: VARIATION IN RESISTANCE TO PARASITISM AMONG APHID POPULATIONS SPECIALIZED ON DIFFERENT PLANTS

The evolution of associations between herbivorous insects and their parasitoids is likely to be influenced by the relationship between the herbivore and its host plants. If populations of specialized herbivorous insects are structured by their host plants such that populations on different hosts are genetically differentiated, then the traits affecting insect-parasitoid interactions may exhibit an associated structure. The pea aphid (Acyrthosiphon pisum) is a herbivorous insect species comprised of genetically distinct groups that are specialized on different host plants (Via 1991a, 1994). Here, we examine how the genetic differentiation of pea aphid populations on different host plants affects their interaction with a parasitoid wasp, Aphidius ervi. We performed four experiments. (1) By exposing pea aphids from both alfalfa and clover to parasitoids from both crops, we demonstrate that pea aphid populations that are specialized on alfalfa are successfully parasitized less often than are populations specialized on clover. This difference in parasitism rate does not depend upon whether the wasps were collected from alfalfa or clover fields. (2) When we controlled for potential differences in aphid and parasitoid behavior between the two host plants and ensured that aphids were attacked, we found that pea aphids from alfalfa were still parasitized less often than pea aphids from clover. Thus, the difference in parasitism rates is not due to behavior of either aphids or wasps, but appears to be a physiologically based difference in resistance to parasitism. (3) Replicates of pea aphid clones reared on their own host plant and on a common host plant, fava bean, exhibited the same pattern of resistance as above. Thus, there do not appear to be nutritional or secondary chemical effects on the level of physiological resistance in the aphids due to feeding on clover or alfalfa, and therefore the difference in resistance on the two crops appears to be genetically based. (4) We assayed for genetic variation in resistance among individual pea aphid clones collected from clover fields and found no detectable genetic variation for resistance to parasitism within two populations sampled from clover. This is in contrast to Henter and Via's (1995) report of abundant genetic variation in resistance to this parasitoid within a pea aphid population on alfalfa. Low levels of genetic variation may be one factor that constrains the evolution of resistance to parasitism in the populations of pea aphids from clover, leading them to remain more susceptible than populations of the same species from alfalfa.     

Impact of environmental stress on aphid clonal resistance to parasitoids: Role of Hamiltonella defensa bacterial symbiosis in association with a new facultative symbiont of the pea aphid

Resistance to endoparasitoids in aphids involves complex interactions between insect and microbial players. It is now generally accepted that the facultative bacterial symbiont Hamiltonella defensa of the pea aphid Acyrthosiphon pisum is implicated in its resistance to the parasitoid Aphidius ervi. It has also been shown that heat negatively affects pea aphid resistance, suggesting the thermosensitivity of its defensive symbiosis. Here we examined the effects of heat and UV-B on the resistance of A. pisum to A. ervi and we relate its stability under heat stress to different facultative bacterial symbionts hosted by the aphid. For six A. pisum clones harboring four different facultative symbiont associations, the impact of heat and UV-B was measured on their ability to resist A. ervi parasitism under controlled conditions. The results revealed that temperature strongly affected resistance, while UV-B did not. As previously shown, highly resistant A. pisum clones singly infected with H. defensa became more susceptible to parasitism after exposure to heat. Interestingly, clones that were superinfected with H. defensa in association with a newly discovered facultative symbiont, referred to as PAXS (pea aphid X-type symbiont), not only remained highly resistant under heat stress, but also expressed previously unknown, very precocious resistance to A. ervi compared to clones with H. defensa alone. The prevalence of dual symbiosis involving PAXS and H. defensa in local aphid populations suggests its importance in protecting aphid immunity to parasitoids under abiotic stress.     

Resistance to Aphis glycines (Hemiptera: Aphididae) in various soybean lines under controlled laboratory conditions

The soybean aphid, Aphis glycines Matsumura (Hemiptera: Aphididae), a pest of soybean, Glycine max (L.) Merr., native to Asia, has recently become a principal pest of this crop in many areas of North America. Insecticides are currently used to manage A. glycines, but host plant resistance is a potential alternative management tool. Tests were conducted to determine resistance to A. glycines among soybean lines. 'Cobb,' 'Tie-feng 8,' and 'Jackson' were resistant to population growth of A. glycines compared with 'Cook' and '91B91,' a susceptible control. Antibiosis was evident in Cobb, Jackson, and Tie-feng 8 from lowered survival of first generation A. glycines, and in Cobb, Jackson, Tie-feng 8, and 'Braxton' from diminished reproduction by first generation aphids. Antixenosis was apparent in Cobb and Jackson during initial infestation of aphid population growth tests, because A. glycines were unsettled and dispersed readily from placement points on unifoliolate leaves. Decreased nymphiposition by A. glycines occurred on Cobb and Jackson, and it may have been caused by antibiotic chemicals in these lines, failure of aphids to settle, or both. Differences in distribution of A. glycines between unifoliolate leaves and other shoot structures suggest that unifoliolate leaves were acceptable feeding sites on 91B91 and Cook, whereas unifoliolate leaves and other shoot structures were roughly equally acceptable feeding sites on Braxton, Tie-feng 8, Jackson, and Cobb. However, Jackson and Cobb had relatively low counts of A. glycines on shoots that may have been due to abandonment of plants by aphids, decreased aphid survival, or both. Results confirm earlier findings that Jackson is a strong source of resistance to A. glycines, and they suggest that Tie-feng 8, Braxton, and especially Cobb are potentially useful sources of resistance.     

Tritrophic interactions among Macrosiphum euphorbiae aphids, their host plants and endosymbionts: Investigation by a proteomic approach

The Mi-1.2 gene in tomato confers resistance against certain clones of the potato aphid (Macrosiphum euphorbiae). This study used 2D-DIGE coupled with protein identification by MALDI-TOF-MS to compare the proteome patterns of avirulent and semivirulent potato aphids and their bacterial endosymbionts on resistant (Mi-1.2+) and susceptible (Mi-1.2−) tomato lines. Avirulent aphids had low survival on resistant plants, whereas the semivirulent clone could colonize these plants. Eighty-two protein spots showed significant quantitative differences among the four treatment groups, and of these, 48 could be assigned putative identities. Numerous structural proteins and enzymes associated with primary metabolism were more abundant in the semivirulent than in the avirulent aphid clone. Several proteins were also up-regulated in semivirulent aphids when they were transferred from susceptible to resistant plants. Nearly 25% of the differentially regulated proteins originated from aphid endosymbionts and not the aphid itself. Six were assigned to the primary endosymbiont Buchnera aphidicola, and 5 appeared to be derived from a Rickettsia-like secondary symbiont. These results indicate that symbiont expression patterns differ between aphid clones with differing levels of virulence, and are influenced by the aphids’ host plant. Potentially, symbionts may contribute to differential adaptation of aphids to host plant resistance.     

Aphelinus albipodus(Hymenoptera: Aphelinidae) andDiaeretiella rapae(Hymenoptera: Braconidae) Parasitism onDiuraphis noxia(Homoptera: Aphididae) Infesting Barley Plants Differing in Plant Resistance to Aphids

Russian wheat aphid,Diuraphis noxia(Mordvilko), as a pest of small grains, has prompted research into biological control and host plant resistance. In the presence of Russian wheat aphid, leaves of a susceptible barley (Morex) are curled and chlorotic and sustain large densities of this aphid, while leaves of a resistant barley (STARS-9301B) remain flat and green and sustain fewer aphids. Might parasitism of Russian wheat aphid byAphelinus albipodusHayat & Fatima andDiaeretiella rapaeMcIntosh be affected differently by these plant types? When presented the plants separately and based on parasitism rate relative to aphid density, the largerD. rapaewas more effective in parasitizing relatively high densities of aphids within curled leaves of Morex than relatively low densities of aphids on uncurled leaves of STARS-9301B. Parasitism byA. albipodusdid not significantly differ among the plants. When given a choice of plants, approximately equal rates of parasitism occurred on the two plant lines for both parasitoid species, and parasitism byD. rapaewas greater thanA. albipodus.These data indicate that using parasitoid size as an indicator of success in a physically restricted environment may be misleading, when considered in a plant environment responsive in several manners to aphids (chlorosis, curling, and ability to sustain Russian wheat aphid). We expect that use of resistant barley will result in decreased parasitoid abundance as aphid densities decrease. However, parasitism rates are expected to be approximately equal on resistant and susceptible barley. In this system, plant resistance and biocontrol are compatible management strategies.     

Assessment of cotton aphids, Aphis gossypii, and their natural enemies on aphid-resistant and aphid-susceptible wheat varieties in a wheat–cotton relay intercropping system

The cotton aphid, Aphis gossypii Glover (Homoptera: Aphididae), is an important cotton pest in northern China, especially in the seedling stage of cotton. After large scale commercial use of transgenic Bt cotton, cotton aphids became one of the most important cotton pests. A 2‐year study was conducted to evaluate the role of four winter wheat varieties that were resistant or susceptible to wheat aphid, Sitobion avenae Fabricius (Homoptera: Aphididae), in conserving arthropod natural enemies and suppressing cotton aphids in a wheat–cotton relay intercropping system in northern China. The results indicated that wheat–cotton intercropping preserved and augmented natural enemies more than a monoculture of cotton. The density of natural enemies in cotton was significantly different among relay‐intercropping fields with different wheat varieties. The highest density of natural enemies and low cotton aphid populations were found in the treatment of cotton in relay intercropped with the wheat variety Lovrin10, which is susceptible to wheat aphid. The lowest density of predators and parasitoids associated with high cotton aphid populations were found with the wheat variety KOK1679, which is resistant to wheat aphid. The results showed that wheat varieties that are susceptible or moderately resistant to wheat aphid might reduce cotton aphids more effectively than an aphid‐resistant variety in the intercropping system by enhancing predators to suppress cotton aphids during the cotton seedling stage.     

棉花抗蚜性与苯丙氨酸解氨酶活性的关系

为探讨苯丙氨酸解氨酶活性与棉花抗蚜性的关系,在室内人工接蚜危害和田间自然蚜群2种条件下,测定不同抗蚜性棉花品种受棉蚜Aphis gossypii Glover危害后叶片的PAL活性。结果表明:未受棉蚜危害时,抗、感品种PAL活性无明显差异;棉蚜危害胁迫棉花PAL活性升高,抗蚜品种受到棉蚜危害诱导所产生的PAL活性远比感蚜品种高。田间有蚜株率与棉花叶片苯丙氨酸解氨酶活性的呈显著的负相关关系。研究说明PAL酶活性对棉花的抗蚜性有一定的影响。     

Feeding behavior and performance of Nasonovia ribisnigri on grafts,detached leaves,and leaf disks of resistant and susceptible lettuce

Aphids are dependent on the phloem sap of plants as their only source of nutrients. Host‐plant resistance in lettuce, Lactuca sativa L. (Asteraceae), mediated by the Nr gene is used to control the lettuce aphid Nasonovia ribisnigri (Mosely) (Hemiptera: Aphididae). The resistance is located in the phloem; however, the exact mechanism of resistance is unknown. In this study, we investigated whether the resistance factor (or factors) is synthesized in the root or in the shoot. The feeding behavior and performance of avirulent N. ribisnigri were studied on grafts of resistant and susceptible lettuce. In addition, the persistence of resistance in excised lettuce tissue was measured, by studying the feeding behavior and performance of N. ribisnigri on detached leaves and leaf disks of resistant lettuce. It appears that the resistance factor encoded by the Nr gene is produced in the shoots: aphid feeding was reduced on resistant shoots grafted on susceptible roots, whereas aphids were able to feed on grafts of susceptible shoots on resistant roots. Partial loss of resistance was observed after detachment of leaves and excision of leaf disks from resistant plants. Aphids fed longer on excised resistant plant tissue compared with intact resistant plants; however, compared with excised plant tissue of the susceptible cultivar, the time spent on feeding was shorter, indicating resistance was not completely lost. Our findings caution against the use of excised leaf material for aphid resistance bioassays.     

A genetic basis for the manipulation of sink–source relationships by the galling aphid <Emphasis Type="Italic">Pemphigus batae</Emphasis>

We examined how the galling aphid Pemphigus batae manipulates resource translocation patterns of resistant and susceptible narrowleaf cottonwood Populus angustifolia. Using carbon-14 (¹⁴C)-labeling experiments in common garden trials, five patterns emerged. First, although aphid galls on resistant and susceptible genotypes did not differ in their capacity to intercept assimilates exported from the leaf they occupied, aphids sequestered 5.8-fold more assimilates from surrounding leaves on susceptible tree genotypes compared to resistant genotypes. Second, gall sinks on the same side of a shoot as a labeled leaf were 3.4-fold stronger than gall sinks on the opposite side of a shoot, which agrees with patterns of vascular connections among leaves of the same shoot (orthostichy). Third, plant genetic-based traits accounted for 26% of the variation in sink strength of gall sinks and 41% of the variation in sink strength of a plant’s own bud sinks. Fourth, tree susceptibility to aphid gall formation accounted for 63% of the variation in ¹⁴C import, suggesting strong genetic control of sink–source relationships. Fifth, competition between two galls was observed on a susceptible but not a resistant tree. On the susceptible tree distal aphids intercepted 1.5-fold more ¹⁴C from the occupied leaf than did basal aphids, but basal aphids compensated for the presence of a distal competitor by almost doubling import to the gall from surrounding leaves. These findings and others, aimed at identifying candidate genes for resistance, argue the importance of including plant genetics in future studies of the manipulation of translocation patterns by phytophageous insects.