麦长管蚜和禾谷缢管蚜对小麦植株挥发物及蚜害诱导挥发物的行为反应

用四臂嗅觉计测定了麦长管蚜Macrosiphum avenae和禾谷缢管蚜Rhopalosiphum padi对小麦植株挥发物及麦蚜取食诱导挥发物的行为反应,揭示了2种麦蚜的嗅觉及小麦植株的诱导防御反应特点.在所选的13种小麦植株挥发物及蚜害诱导挥发物组分中,6-甲基-5-庚烯-2-酮、6-甲基-5-庚烯-2-醇和水杨酸甲酯对这2种蚜虫表现出强的驱拒作用;反-2-己烯醛对麦长管蚜的有翅和无翅蚜的吸引作用最强;反-2-己烯醇对禾谷缢管蚜的无翅蚜吸引作用最强,反-3-己酰醋酸酯对禾谷缢管蚜有翅蚜的吸引作用最强.说明麦蚜取食能诱导小麦植株的防御反应,麦长管蚜和禾谷缢管蚜及其不同蚜型间嗅觉反应的特点不同.     

Effects of variable maternal temperature on offspring development and reproduction of Rhopalosiphum padi,a serious global pest of wheat

1. Global warming is occurring at an unprecedented rate. Information about how variable temperature affects insect life-history traits is still scarce. 2. The current study first evaluated the effects of two variable-temperature treatments [high-temperature-variation (HT) treatments and mild-temperature-variation (MT) treatments] on the life-history traits of a maternal generation (F₀) of Rhopalosiphum padi, a serious global pest, using a constant normal-temperature (NT) treatment as a control. Following this, the life-history traits of the offspring generation (F₁) under the NT, MT and HT scenarios were analysed. 3. The total developmental duration of the aphid F₀ generation was significantly shortened by MT treatments, while it was significantly increased by HT treatments. Adult longevity and the fecundity of F₀ were significantly decreased by HT treatments compared with those in the MT and NT treatments, whereas no significant difference was found between the latter two treatments. The HT treatments applied to the F₀ generation significantly prolonged the total developmental duration of the F₁ generation. The offspring adult longevity was not affected by the mothers' temperature experience. The offspring fecundity was significantly increased when the F₀ generation experienced MT treatments. The intrinsic rate of increase (r_m) was significantly decreased when the F₀ generation experienced an HT scenario. However, the MT scenario experienced by the F₀ generation did not significantly affect the r_m of their offspring. 4. The results will provide new insights into the effects of variable maternal temperature on the individual development and population dynamics of offspring under a global warming scenario.     

The effect of crowding on alate production and weight of apterous exules of the apple-grass aphid, Rhopalosiphum insertum

Offspring of Rhopalosiphum insertum are most sensitive to crowding immediately before and after birth. Crowding throughout the nymphal and adult life of the mothers irreversibly determines most offspring as alatae whether the offspring are crowded or not. Offspring of mothers having less exposure to crowding require additional stimuli after birth. The possible evolutionary advantages of such a system are discussed. Hormonal control of alary polymorphism is implicated. The weight of apterae declined with increased crowding. This is correlated with an increase in alate production, and may thus serve to limit the population increase to a level tolerable by the plant without killing it.     

Aphid colonization of spring cereals

In 1970-1, Metopolophium dirhodum, Rhopalosiphum padi and Sitobion avenae were the commonest alatae trapped from April/May to August, with most in July and early August. The first alatae appeared in the Rothamsted survey suction trap 0–34 days before aphids were found on the cereals, but during May and June no relationship was found between the numbers trapped and the number on the crop. Most species occurred first near the sheltered edge of the crop, but M. dirhodum was widespread over the field. Most infestations were quickly dispersed by the movements of older morphs; adults only stayed in one place for about 2 days. Alate M. dirhodum moved more often than apterae, but both morphs of S. avenae moved equally often and more frequently between larvipositions than did those of M. dirhodum. Apterae deposited more nymphs in a ‘group’ than alatae, and M. dirhodum deposited more than S. avenae. Few ‘groups’ persisted for more than a week. Although M. dirhodum occupied the crop area faster than S. avenae, all 0–3 m lengths of row sampled being infested within 2–5 wk of their first appearance, most or all of the tillers were colonized only in late July 1970.     

Shaped by the past,acting in the present: transgenerational plasticity of anti‐predatory traits

Phenotypic expression can be altered by direct perception of environmental cues (within‐generation phenotypic plasticity) and by the environmental cues experienced by previous generations (transgenerational plasticity). Few studies, however, have investigated how the characteristics of phenotypic traits affect their propensity to exhibit plasticity within and across generations. We tested whether plasticity differed within and across generations between morphological and behavioral anti‐predator traits of Physa acuta, a freshwater snail. We reared 18 maternal lineages of P. acuta snails over two generations using a full factorial design of exposure to predator or control cues and quantified adult F₂ shell size, shape, crush resistance, and anti‐predator behavior – all traits which potentially affect their ability to avoid or survive predation attempts. We found that most morphological traits exhibited transgenerational plasticity, with parental exposure to predator cues resulting in larger and more crush‐resistant offspring, but shell shape demonstrated within‐generation plasticity. In contrast, we found that anti‐predator behavior expressed only within‐generation plasticity such that offspring reared in predator cues responded less to the threat of predation than control offspring. We discuss the consequences of this variation in plasticity for trait evolution and ecological dynamics. Overall, our study suggests that further empirical and theoretical investigation is needed in what types of traits are more likely to be affected by within‐generational and transgenerational plasticity.     

Resistance and susceptibility of some wheat lines to bird cherry-oat aphid <Emphasis Type="Italic">Rhopalosiphum padi</Emphasis> (Hemiptera: Aphididae)

Bird cherry-oat aphid, Rhopalosiphum padi (L.), is one of the most serious pests of cereals, with an almost worldwide distribution. A primary screening test was carried out to gauge the resistance or susceptibility of 40 wheat lines to R. padi, and follow-up experiments were conducted to determine the development and fecundity rates of R. padi on eight lines. The forty wheat lines examined were classified into two major classes and four subclasses: A (e.g., ERWYT 87-1) and B (e.g., ERWYT 87-20, ERWYT 87-11), with the highest average number of aphids 14 days after infestation, subclass C (e.g., ERWYT 88-8, ERWYT 87-6, ERWYT 87-4), with the lowest average number of aphids, and subclass D (e.g., ERWYT 88-12, ERWYT 88-13), with intermediate status. Aphid nymph developmental time and survival both differed among the wheat lines. Differences were also observed in the fecundity of R. padi, the intrinsic rate of increase (r _m), and other parameters among the wheat lines tested. The highest and lowest values of r _m were obtained for ERWYT 87-1 and ERWYT 88-8 (0.36 and 0.26 nymphs/female/day, respectively). Three lines (ERWYT 87-4, ERWYT 87-6, ERWYT 88-8) were relatively resistant to R. padi, which could prove useful in the development of IPM programs for this aphid in wheat fields.     

Population differences in host use by a seed-beetle: local adaptation, phenotypic plasticity and maternal effects

For insects that develop inside discrete hosts, both host size and host quality constrain offspring growth, influencing the evolution of body size and life history traits. Using a two-generation common garden experiment, we quantified the contribution of maternal and rearing hosts to differences in growth and life history traits between populations of the seed-feeding beetle Stator limbatus that use a large-seeded host, Acacia greggii, and a small-seeded host, Pseudosamanea guachapele. Populations differed genetically for all traits when beetles were raised in a common garden. Contrary to expectations from the local adaptation hypothesis, beetles from all populations were larger, developed faster and had higher survivorship when reared on seeds of A. greggii (the larger host), irrespective of their native host. We observed two host plant-mediated maternal effects: offspring matured sooner, regardless of their rearing host, when their mothers were reared on P. guachapele (this was not caused by an effect of rearing host on egg size), and females laid larger eggs on P. guachapele. This is the first study to document plasticity by S. limbatus in response to P. guachapele, suggesting that plasticity is an ancestral trait in S. limbatus that likely plays an important role in diet expansion. Although differences between populations in growth and life history traits are likely adaptations to their host plants, host-associated maternal effects, partly mediated by maternal egg size plasticity, influence growth and life history traits and likely play an important role in the evolution of the breadth of S. limbatus’ diet. More generally, phenotypic plasticity mediates the fitness consequences of using novel hosts, likely facilitating colonization of new hosts, but also buffering herbivores from selection post-colonization. Plasticity in response to novel versus normal hosts varied among our study populations such that disentangling the historical role of plasticity in mediating diet evolution requires the consideration of evolutionary history.     

Levels and components of resistance to Amphorophora idaei in raspberry cultivars containing different resistance genes

Levels and components of resistance to Amphorophora idaei in raspberry cultivars containing different A. idaei resistance genes were studied under infestation tunnel, glasshouse and laboratory conditions. Each test consistently ranked raspberry cultivars, with increasing levels of resistance in the order, non-resistant (cv. Mailing Jewel), resistant cultivars containing minor genes, major gene A₁, gene A₁₀ (red raspberry) and gene A₁₀ (black raspberry) respectively. Resistance was expressed in three different ways; decreased alatae settling and feeding, decreased apterae settling and decreased aphid fecundity and rate of nymphal development. Following exposure to a large population of alatae, significantly fewer aphids settled on A₁, A₁₀ and to a lesser extent, minor gene-containing cultivars, compared to cv. Mailing Jewel. More alatae settled on the top than the bottom zone of non-resistant, minor gene resistant and two of four A₁-containing cultivars; alatae settling was low on all zones of A₁₀-containing raspberries. Aphid fecundity and nymph development patterns on different cultivars and resistance classes were similar to those found for alatae settling. After 7 days reproduction more than 30% of the nymphs developed to third or fourth instar on cv. Mailing Jewel, whilst on minor gene and major gene-containing cultivars the total number of nymphs and the proportion of later instars decreased. On resistant cultivars the nymphs were found mainly on the middle and bottom leaf zones, compared to cv. Mailing Jewel. A rapid (48 h) screening test using floating leaflets was developed and, on the basis of apterae settling, ranked cultivars reliably; it was particularly effective in distinguishing moderate and strong resistance to strain 1 A. idaei. Gene A₁-containing cultivars bred in England were much more resistant to strain 1 A. idaei than those bred in Scotland. Possible reasons for this difference are discussed, in relation to modified screening procedures and to the control of the viruses transmitted by this aphid vector.     

Predator-induced transgenerational phenotypic plasticity in the cotton aphid

Transgenerational phenotypic changes, whereby offspring have an altered trait or a distinct alternate phenotype, frequently occur in response to increased maternal predation risk. The cotton aphid, Aphis gossypii (Glover), is unique, however, as offspring consist of four distinct phenotypes (normal light green apterae, normal dark green apterae, dwarf yellow apterae, and alatae), all with divergent life history patterns and resulting population dynamics. Here, we show that increased predation risk induces transgenerational phenotypic changes in cotton aphids. When exposed to search tracks from larval or adult convergent ladybird beetles, Hippodamia convergens Guérin-Méneville, cotton aphids produced greater numbers of winged offspring. In a subsequent experiment, apterous and alate individuals on clean plants were found to have primarily normal and dwarf offspring, respectively. We suggest that elevated predation risk may cause phenotypic changes in aphids over multiple generations, resulting in a more precipitous decline in herbivore populations than could be explained solely by increased predation rates.     

Paternal epigenetic effects of population density on locust phase‐related characteristics associated with heat‐shock protein expression

Many species exhibit transgenerational plasticity by which environmental cues experienced by either parent can be transmitted to their offspring, resulting in phenotypic variants in offspring to match ancestral environments. However, the manner by which paternal experiences affect offspring plasticity through epigenetic inheritance in animals generally remains unclear. In this study, we examined the transgenerational effects of population density on phase‐related traits in the migratory locust Locusta migratoria. Using an experimental design that explicitly controls genetic background, we found that the effects of crowd or isolation rearing on phase plasticity could be inherited to the offspring. The isolation of gregarious locusts resulted in reduced weight in offspring eggs and altered morphometric traits in hatchlings, whereas crowding of solitarious locusts exhibited opposite effects. The consequences of density changes were transmitted by both maternal and paternal inheritance, although the expression of paternal effects was not as pronounced as that of maternal effects. Prominent expression of heat‐shock proteins (Hsps), such as Hsp90, Hsp70 and Hsp20.6, could be triggered by density changes. Hsps were significantly upregulated upon crowding but downregulated upon isolation. The variation in parental Hsp expression was also transmitted to the offspring, in which the pattern of inheritance was consistent with that of phase characteristics. These results revealed a paternal effect on phase polyphenism and Hsp expression induced by population density, and defined a model system that could be used to study the paternal epigenetic inheritance of environmental changes.     

Characterisation of bird cherry-oat aphid (Rhopalosiphum padi L.) behaviour and aphid host preference in relation to partially resistant and susceptible wheat landraces

The bird cherry-oat aphid (Rhopalosiphum padi L.) is a major pest of wheat (Triticum aestivum L.) and can cause up to 30% yield losses. Heritable plant resistance to aphids is both an economically and ecologically sound method for managing aphids. Here we report how the behaviour and performance of R. padi differs on two resistant, one susceptible wheat landrace and a susceptible elite wheat variety. Feeding behaviour differed among the genotypes, with aphids on resistant lines spending longer in the pathway phase and less time phloem feeding. These behaviours suggest that both inter- and intracellular factors encountered during pathway and phloem feeding phases could be linked to the observed aphid resistance. Locomotion and antennal positioning choice tests also revealed a clear preference for susceptible lines. Although feeding studies revealed differences in the first probe indicating that the resistance factors might also be located in the peripheral layers of the plant tissue, scanning electron microscopy revealed no difference in trichrome length and density on the surface of leaves. Aphids are phloem feeders and limiting the nutrient uptake by the aphids may negatively affect their growth and development as shown here in lower weight and survival of nymphs on resistant genotypes and decreased reproductive potential, with lowest mean numbers of nymphs produced by aphids on W064 (54.8) compared to Solstice (71.9). The results indicate that resistant lines markedly alter the behaviour, reproduction and development potential of R. padi and possess both antixenosis and antibiosis type of resistance.     

Drought stress and cereal aphid performance

The performance of clones of Rhopalosiphum padi and Sitobion avenae from England and Spain was examined on drought-stressed tillering winter wheat in an environment chamber at 14 ± 1°C. Two different levels of drought stress and an unstressed control were established by different watering regimes which resulted in drought-stressed plants being smaller at the end of the experiment. The effect of drought stress to plants on aphid performance was not significantly different between the clones tested. Drought stress had no effect on aphid development time, nymphal mortality, the weight of teneral adults and the number of embryos in teneral adults up to the onset of reproduction in the first F₁ generation. The subsequent reproductive capacity, as measured by the effective and potential fecundity, and the reproductive rate, were much reduced on drought-stressed plants. However, there was only a small decrease in the intrinsic rate of increase (r_m). Overall the clone of R. padi from Spain performed better than that from England, the development and prereproductive times being shorter and the fecundity higher in the Spanish clone, giving a higher r_m. There were no differences in the fecundity and the r_m between the Spanish and the English clones of S. avenae. The proportion of the F₂ generation that was alate differed greatly between clones, and only the English S. avenae produced significantly more alatae on drought-stressed than on unstressed plants.     

Differential colonization of wheat cultivars by two biotypes of Russian wheat aphid （Homoptera： Aphididae）

Susceptible and resistance wheat cultivars, Triticum aestivum L, were presented to two biotypes of Russian wheat aphid, Diuraphis noxia （Mordvilko）, in multiple choice tests to assay their relative acceptability as host plants. Both apterae （third and fourth instars） and alate adults were offered plants at the two-leaf stage in different cultivar combinations at 22±1℃ and 16：8 （L： D） hour photoperiod. Apterae were released from Petri dishes in the center of a circle of test plants, whereas alatae dispersed from a mature aphid colony to settle on plants arranged in rows. Both alatae and apterous nymphs of both biotypes readily colonized all cultivars tested：‘2137＇, ‘Akron＇,‘Ankor’,‘ Halt’ ,‘ Jagger’ ,‘ Prairie Red’ , ‘Stanton＇,‘TAM 107＇,‘TAM 110＇,‘Trego＇, ‘ Yuma＇, and ‘Yumar＇. Fewer biotype I apterae responded （settled and fed） in the combination containing more resistant （Dn4- and Dny-expressing） cultivars, compared to the combinations that had fewer. The reverse was true for biotype 2 apterae; more aphids responded in the combination containing the largest number of Dn4 expressing cultivars. Differential colonization of cultivars was observed in only one combination, in which biotype 2 apterae colonized Akron and Yumar in larger numbers than they did Stanton and Yuma. A separate experiment confirmed that, 48 hours after infestation, more biotype 2 apterae abandoned plants of Yuma than plants of Yumar. This differential response was likely due to genetic differences between the two ＇ near isogenic＇ lines that include the lack of Dn4 expression in Yuma. Choice tests with alatae did not result in differential rates of cultivar colonization by either biotype in any combination tested. These results suggest that young wheat plants appear to lack any meaningful antixenosis toward D. noxia, even though the aphids appear to perceive, and sometimes respond to, certain differences in cultivar suitability.     

Electronic recording of plant penetration by the cereal aphids Rhopalosiphum padi and Metopolophium dirhodum on resistant and susceptible wheat seedlings

Plant penetration by the aphids Rhopalosiphum padi and Metopolophium dirhodum was monitored using an electronic recording method. Recordings over 3 h showed that phloem ingestion was reduced on resistant wheat, when compared with a more susceptible species, but this trend was reversed when results from longer (12 h) experiments were used. Plant penetration by M. dirhodum was reduced by attachment to the electronic system, but the behaviour of R. padi was relatively unaffected. The results are discussed in relation to prospects for using electronic methods as a screening procedure.     

Honeydew production and sap ingestion by the cereal aphids Rhopalosiphum padi and Metopolophium dirhodum on seedlings of resistant and susceptible wheat species

Measurements of the frequency and rate of honeydew excretion by Rhopalosiphum padi and Metopolophium dirhodum, feeding on seedlings of resistant and susceptible wheat (Triticum monococcum and T. aestivum) were used to estimate the quantity of sap ingested over 24 h. Ingestion was significantly reduced on the more resistant T. monococcum and for M. dirhodum this may be due in part to a reduction in the success of phloem location.     

Oviposition vs. offspring fitness in Aphidius colemani parasitizing different aphid species

We measured the acceptance and suitability of four aphid species [Aphis gossypii Glover, Myzus persicae (Sulzer), Rhopalosiphum padi (L.), and Schizaphis graminum (Rondani)] (Homoptera: Aphididae) for the parasitoid Aphidius colemani Viereck (Hymenoptera: Braconidae). Female parasitoids parasitized fewer R. padi than the other three aphid species, and fewer offspring successfully completed development in R. padi than in the other three host species. Sex ratios of emerging adults were more male‐biased from R. padi than from the other three aphid species, suggesting that R. padi is a poor quality host for this population of A. colemani. Ovipositing A. colemani encountered R. padi at a slower rate, spent more time handling R. padi, and parasitoid offspring died at a higher rate in R. padi compared to A. gossypii. Our results show that oviposition behavior and offspring performance are correlated. In each experiment, we tested the effect of the host species in which the parasitoids developed (parental host) on the number of hosts attacked, the proportion of each host species accepted for oviposition and the survival of progeny. Parental host affected maternal body size and, through its effect on body size, the rate of encounter with hosts. Other than this, parental host species did not affect parasitism.     

Mapping resistance to the bird cherry-oat aphid and the greenbug in wheat using sequence-based genotyping

Key message

Identification of novel resistance QTL against wheat aphids. First QTL-resistance report for R. padi in wheat and chromosome 2DL for S. graminum . These sources have potential use in wheat breeding.

Abstract

The aphids Rhopalosiphum padi and Schizaphis graminum are important pests of common wheat (Triticum aestivum L.). Characterization of the genetic bases of resistance sources is crucial to facilitate the development of resistant wheat cultivars to these insects. We examined 140 recombinant inbred lines (RILs) from the cross of Seri M82 wheat (susceptible) with the synthetic hexaploid wheat CWI76364 (resistant). RILs were phenotyped for R. padi antibiosis and tolerance traits. Phenotyping of S. graminum resistance was based on leaf chlorosis in a greenhouse screening and the number of S. graminum/tiller in the field. RILs were also scored for pubescence. Using a sequence-based genotyping method, we located genomic regions associated with these resistance traits. A quantitative trait locus (QTL) for R. padi antibiosis (QRp.slu.4BL) that explained 10.2 % of phenotypic variation was found in chromosome 4BL and located 14.6 cM apart from the pubescence locus. We found no association between plant pubescence and the resistance traits. We found two QTLs for R. padi tolerance (QRp.slu.5AL and QRp.slu.5BL) in chromosomes 5AL and 5BL, with an epistatic interaction between a locus in chromosome 3AL (EnQRp.slu.5AL) and QRp.slu.5AL. These genomic regions explained about 35 % of the phenotypic variation. We re-mapped a previously reported gene for S. graminum resistance (putatively Gba) in 7DL and found a novel QTL associated with the number of aphids/tiller (QGb.slu-2DL) in chromosome 2DL. This is the first report on the genetic mapping of R. padi resistance in wheat and the first report where chromosome 2DL is shown to be associated with S. graminum resistance.     

Mixing barley cultivars affects aphid host plant acceptance in field experiments

Four barley varieties with no significant difference in aphid acceptance were sown in pure stands and in pairwise combinations with varieties side by side in separate rows. Settling tests were done in situ in the field plots with apterae of Rhopalosiphum padi (L.) (Homoptera:Aphididae) and showed that aphid acceptance was changed in some combinations of cultivars. In a laboratory test, in which plants of one cultivar were exposed to air from the other cultivars, aphid acceptance was significantly reduced in three of the four cultivars when treated with air from certain other cultivars. Two of these three cultivars showed the same reduction under field conditions. This supports the hypothesis that plant/plant communication may release responses in neighbouring plants that change aphid host plant acceptance. The results also show that this mechanism is not restricted to optimal growing conditions in the laboratory, although it may be modified under field conditions depending on plant genotype.     

Effects of elevated atmospheric CO2 concentrations on phloem sap composition of spring crops and aphid performance

Abstract

The concentration and composition of free amino acids and carbohydrates in the phloem sap of wheat and oilseed rape (OSR) and the effects on the performance of aphids (Rhopalosiphum padi and Myzus persicae) were determined under atmospheric carbon dioxide (CO₂) enrichment. The analysis of phloem sap showed that carbohydrates and amino acid levels of the host plants were significantly affected by elevated CO₂ level. Among carbohydrate concentrations in the phloem sap, significant increases were observed in fructose and glucose in spring wheat under CO₂ enrichment, whereas no changes were observed in OSR. These changes in plant chemistry affected the performance of herbivorous insects (i.e. aphids) in varying ways, positively affecting the relative growth rate (RGR) of R. padi in spring wheat and negatively affecting the RGR of M. persicae on OSR.     

Searching for wheat resistance to aphids and wheat bulb fly in the historical Watkins and Gediflux wheat collections

Insect pests can reduce wheat yield by direct feeding and transmission of plant viruses. Here we report results from laboratory and field phenotyping studies on a wide range of wheat, including landraces from the Watkins collection deriving from before the green revolution, more modern cultivars from the Gediflux collection (north‐western Europe) and modern UK Elite varieties, for resistance to the bird cherry‐oat aphid, Rhopalosiphum padi (Homoptera: Aphididae) and the English grain aphid, Sitobion avenae (Homoptera: Aphididae). A total of 338 lines were screened for R. padi and 340 lines for S. avenae. Field trials were also conducted on 122 Watkins lines to identify wheat bulb fly, Delia coarctata, preference on these landraces. Considerable variation was shown in insect performance among and within different wheat collections, with reduced susceptibility in a number of varieties, but phenotyping did not identify strong resistance to aphids or wheat bulb fly. Field trials showed within collection differences in aphid performance, with fewer aphids populating lines from the Watkins collection. This differs from development data in laboratory bioassays and suggests that there is a pre‐alighting cue deterring aphid settlement and demonstrates differences in aphid preference and performance on older plants in the field compared with seedlings in the laboratory, highlighting the need for phenotyping for aphid resistance at different plant growth stages. No association was identified between performance of the different insect species on individual varieties, potentially suggesting different nutritional requirements or resistance mechanisms.