The effect of cereal growth stages on the reproductive activity of Sitobion avenue and Metopolophium dirhodum

Apterous Sitobion avenae on oats were found to have a higher reproductive rate on the ears (5–95 nymphs per day) than on young leaves (3–78 nymphs per day), mature leaves (2–17 nymphs per day) or senescent leaves (2–08 nymphs per day). At each growth stage of the the host plant large aphids gave birth to more young than small aphids, but aphids of the same weight on different growth stages of the host plant had different reproductive rates. On wheat in the field S. avenae gave birth to more young on the ears but less on the leaves than Metopolophium dirhodum on the leaves.
The suitability of ears to S. avenae changed through the flowering and ripening stages. During the latter part of the milky-ripe stage the reproductive rate of S. avenae declined and its mortality increased. The consequences of the variations in reproduction and mortality are discussed in relation to the life history and population dynamics of S. avenae and the assessment of cereal varieties for aphid resistance.     

蕾铃期棉蚜种群动态及年龄结构变化和受寄生的关系

本文描述了1983年棉花蕾铃期棉蚜发生情况。从7月上旬到9月上旬发生量超过往年同期的10倍。绘出了蚜虫数量的对数值和生理时间关系图,描述了棉蚜种群下降过程中年龄结构变化及其与寄生率变化的关系。在棉蚜种群下降过程中Ⅰ龄若蚜占百分比下降表明成蚜生殖力下降。高密度组Ⅱ龄若蚜百分比下降,表明Ⅰ龄若蚜死亡率增大。高密度组成虫比例下降是因为有翅若蚜比例高。上述年龄结构变化和寄生率关系密切,表明蚜群下降到低水平时寄生率上升不完全由于寄生蜂增殖,而Ⅰ、Ⅱ龄若蚜比例下降、有翅若蚜增加和高龄若蚜比例上升中是重要原因。棉株不同位置棉蚜寄生率没有差异。寄生率和单位面积内的蚜虫密度有关。     

Natural enemy ravine revisited: the importance of sample size for determining population growth

Abstract. 1. The population growth of three aphid species, Metopolophium dirhodum (Walker), Rhopalosiphum padi (L.), and Sitobion avenae (F.), on winter wheat, was analysed by regression. The calculations were based on censuses of aphids made in 268 plots at 3- or 7-day intervals for 10 years on leaves and 6 years on ears. The calculations were made separately for each plot each year, then repeated on the pooled data from all plots monitored in a year.
2. At the level of individual plots, no population growth was detected at very low densities. At high densities, the populations grew exponentially and the growth rates did not decrease with increasing aphid density.
3. Significant growth was always detected in the pooled data. These growth rates decreased significantly at the highest densities. Field estimates of the intrinsic rate of increase derived from these data ranged from 0.010 to 0.026 for M. dirhodum , 0.0071–0.011 for R. padi , and 0.00078–0.0061 and 0.0015–0.13 for S. avenae , on leaves and ears respectively .
4. The apparent lack of growth in the individual plots at low densities is attributable to small sample size. It is concluded that the natural enemy ravine in the population dynamics of cereal aphids, identified by Southwood and Comins (1976), is a consequence of low population densities at which population increase is undetectable unless very large samples are taken.     

Comparing growth patterns among field populations of cereal aphids reveals factors limiting their maximum abundance

Cereal stands in central Europe are commonly infested with three species of aphids that may become serious pests. With increasing abundance, the proportion of a particular species in the total aphid population may remain constant, suggesting a density-independent exponential growth, or the proportion can change, suggesting density-dependent constraints on growth. The constraints that affect particular species, and thus their relative abundance, were studied. The proportionality between maximum abundances of the cereal aphids was studied using a 10-year census of the numbers of aphids infesting 268 winter wheat plots. For two species their abundance on leaves and ears was compared. With increasing aphid density the maximum abundance of Rhopalosiphum padi (Linnaeus) remained proportional, but not that of Sitobion avenae (Fabricius), which was constrained by the smaller surface area of ears compared to leaves. There was no evidence of inter-specific competition. Maximum abundance of R. padi and Metopolophium dirhodum (Walker) on leaves did not change proportionally as the proportion of M. dirhodum decreased with increasing overall aphid density. This decrease was probably caused by the restricted distribution of M. dirhodum, which is confined to leaves, where space is limiting. No change in proportion between populations was detected when the average densities were below 0.54 aphids per leaf or ear. Non-proportional relationships between aphid populations appeared to be due to spatial constraints, acting upon the more abundant population. Detecting the limitation of population growth can help with the assessment of when density-independent exponential growth is limited by density-dependent factors. This information may help in the development of models of cereal aphid population dynamics.     

Assessment of aphid lethal paralysis virus as an apparent population growth‐limiting factor in grain aphids in the presence of other natural enemies

The influence of aphid lethal paralysis virus (ALPV), Rhopalosiphum padi virus (RhPV), natural enemies and fungal infection on the population growth of Rhopalosiphum padi and Sitobion avenae in the wheat fields of the Western Cape Province of South Africa was investigated at two sites. Time‐specific life tables were compiled for R. padi at one site. During the logarithmic phase of the development of R. padi aphids, natural enemies were not present in high numbers and the apparent large‐scale mortality observed appeared to be due to other causes. During the decline phase of this aphid population, the population size was reduced by 49%. This reduction coincided with a calculated high mortality of 70 aphids per plant. A dramatic decline in R. padi numbers and a high incidence of ALPV present in the aphid population was experienced during this period. Virus assays were carried out by double‐antibody sandwich enzyme‐linked immunosorbent assay (DAS‐ELISA) and an indirect immunofluorescent technique. Entomophthorales‐type fungal infection of aphids also reached its highest level during the decline phase, but at a later stage than ALPV infection, with a calculated level of 21 aphids per plant. This suggested that the presence of ALPV limited population development in R. padi. Similar results were obtained with S. avenae.     

Seasonal dynamics of cereal aphids on Russian wheat aphid (Homoptera: Aphididae) susceptible and resistant wheats

Field experiments were conducted in 1997 and 1998 to evaluate the impact of resistance to Russian wheat aphid, Diuraphis noxia (Mordvilko), on the cereal aphid complex in wheat. Two spring wheats were planted: the variety "Centennial" (Russian wheat aphid susceptible) and the advanced line IDO488 (Russian wheat aphid resistant). IDO488 incorporates the resistance found in PI 294994 into a Centennial background. Field plots were artificially infested with adult D. noxia and sampled weekly. The most abundant aphid species in 1997 were Metopolophium dirhodum (Walker), Sitobion avenae (F.), D. noxia, and Rhopalosiphum padi (L.). In 1998, the order of abundance was M. dirhodum, R. padi, S. avenae, and D. noxia. The resistant genotype had significantly fewer D. noxia than the susceptible one during both years. However, plant genotype had no significant effect on the other aphid species in either year. Both the initial density of D. noxia and plant growth stage, had a significant effect on D. noxia population development, but had no effect on the other aphid species. There was no interaction between D. noxia resistance and the population density of the other aphid species observed.     

小麦间作大蒜或油菜对麦长管蚜及其主要天敌种群动态的影响

麦长管蚜是我国冬小麦产区的重要害虫.试验探索了不同间作方式对麦长管蚜及其主要天敌的影响.结果表明,在调查期内,麦-油(油菜)间作田和麦-蒜(大蒜)间作田中麦长管蚜无翅蚜的种群密度多显著低于单作田;麦-油间作田中有较高的瓢虫种群密度和瓢蚜比;5月2日前,麦-油间作田中蚜茧蜂的种群密度高于单作田和麦-蒜间作田;5月5日后,麦-油间作田的僵蚜率和蜂/蚜比也显著高于单作田和麦-蒜间作田;麦-蒜间作田有翅蚜的种群数量较高,但瓢虫和蚜茧蜂的种群数量无明显变化.麦-油间作和麦-蒜间作均能对麦田中麦长管蚜起到较好的控制作用.     

Testing the resource concentration hypothesis with tarnished plant bug on strawberry: density of hosts and patch size influence the interaction between abundance of nymphs and incidence of damage

Abstract. 1. Attributes of patches with strawberry plants were manipulated experimentally to assess the impact of patch size and host density on the abundance of tarnished plant bug (Hemiptera: Miridae) and incidence of fruit damage.
2. The density of nymphs per inflorescence increased with patch size and host density for some but not all generations of plant bug, providing partial support for the resource concentration hypothesis, while emphasising the importance of replicating experiments across several generations of herbivore.
3. Increasing density of nymphs with increasing patch size and host density did not translate into a high incidence of damage, which may be due to the relatively low feeding impact of nymphs in patches with numerous fruits and to the relatively high perimeter-to-area ratio in small patches resulting in a relatively high proportion of damaged fruits.
4. The decreasing proportion of damaged fruits with increasing density of fruits per patch suggests that nymphs exhibit a saturating functional response to density of strawberry fruits. Functional responses of herbivores may be a critical yet overlooked component that influences interactions between insects and their host plant. In particular, concentrated plant resources may result in increased numbers of herbivores yet sill have a negligible influence on plant damage and/or fitness.     

Cursorial spiders retard initial aphid population growth at low densities in winter wheat

Generalist predators contribute to pest suppression in agroecosystems. Spider communities, which form a substantial fraction of the generalist predator fauna in arable land, are characterized by two functional groups: web-building and cursorial (non-web-building) species. We investigated the relative impact of these two functional groups on a common pest (Sitobion avenae, Aphididae) in wheat by combining a molecular technique that revealed species-specific aphid consumption rates with a factorial field experiment that analyzed the impact, separately and together, of equal densities of these two spider functional groups on aphid population growth. Only cursorial spiders retarded aphid population growth in our cage experiment, but this effect was limited to the initial aphid-population growth period and low-to-intermediate aphid densities. The molecular analysis, which used aphid-specific primers to detect aphid DNA in predator species, detected the highest proportion of aphid-consuming individuals in two cursorial spiders: the foliage-dwelling Xysticus cristatus (Thomisidae) and the ground-active Pardosa palustris (Lycosidae). The results suggest that manipulating the community composition in favour of pest-consuming functional groups may be more important for improving biological control than fostering predator biodiversity per se. Agricultural management practices that specifically foster effective species or functional groups (e.g. mulching for cursorial spiders) should receive more attention in low-pesticide farming systems.     

Effects of variation among plant species on the interaction between a herbivore and its parasitoid

Abstract.  1. Previous studies have demonstrated that phenotypic traits of plants have the potential to affect interactions between herbivores and their natural enemies. Consequently, the impact of natural enemies on herbivore vital rates and population dynamics may vary among plant species. This study was designed to investigate the potential for density-dependent parasitism of an aphid herbivore feeding on six different host plant species.
2. Population densities of the aphid Aphis nerii B de F (Homoptera: Aphididae) and its parasitoid Lysiphlebus testaceipes Cresson (Hymenoptera: Braconidae) were recorded within a single growing season on six different species of milkweed in the genus Asclepias L. (Asclepiadaceae). Asclepias species are known to vary in their quality as food for herbivores. Although data on plant quality were not available in this study, population data were analysed to determine the effects of different Asclepias species on rates of parasitism and aphid population growth.
3. Parasitism rates of A. nerii varied among Asclepias species but were temporally density dependent over at least some range of aphid density on all plant species. Aphid population growth rates also varied among Asclepias species, and declined with an increase in the maximum parasitism rates among plant species; however, in no case was density-dependent parasitism sufficient to prevent exponential population growth of aphids within the growing season. The results serve to emphasise that, if natural enemies are to regulate herbivore populations, density-dependent mortality is a necessary, but not sufficient, condition for regulation.     

Population dynamics of cereal aphids: influence of a shared predator and weather

1 Aphid populations may show strong year-to-year fluctuations, but questions remain regarding the dominance of factors that cause this variation, especially the role of natural enemies. To better understand the dynamics of aphid species that occur as pests in cereals, we investigated the relative influence of top-down control by a predator and weather (temperature and precipitation) on population fluctuations of three cereal aphid species.
2 From 1987 to 2005, populations of Metopolophium dirhodum , Sitobion avenae and Rhopalosiphum padi in insecticide-free stands of winter wheat were monitored in the Praha-Ruzyné region of the Czech Republic. Densities of an aphidophagous predator, the ladybeetle Coccinella septempunctata , were recorded from an overwintering site in the landscape. Weather was quantified using historical records.
3 A significant bottom-up effect of densities of aphids on those of C.   septempunctata was found, but evidence of direct top-down regulation of aphids by C.   septempunctata was only significant in the case of R.   padi . There was no significant periodicity in the dynamics of the aphid or C.   septempunctata , suggesting that there was no clear predator-prey cycle. Combinations of C.   septempunctata and weather variables could be used to explain M.   dirhodum and R.   padi per capita rate of change. There were also indications that weather directly affected peak density of M.   dirhodum .
4 We conclude that regional estimates of C.   septempunctata densities are not sufficient to determine whether aphid population dynamics are driven by predator–prey interactions. Feasibility of time series analysis as an investigative tool in aphid population dynamics studies is discussed.     

The effect of parasitoid host-size preference on host population growth rates: an example of Aphidius colemani and Aphis glycines

Abstract.  1. Direct observations of Aphidius colemani foraging for Aphis glycines demonstrated that A. colemani attacks large aphid size classes selectively, in contrast to other Aphidiinae (Hymenoptera: Braconidae) species that generally prefer small or medium-sized nymphs of different host species.
2. To determine the effect of this size preference on the potential ability of A. colemani to control A. glycines outbreaks, the stage-dependent survival and fecundity of A. glycines were measured to parameterise a stage-structured demographic model for the density-independent population growth rate of the aphid.
3. Compared with hypothetical parasitoids that show either no size preference or preference for medium-sized host nymphs, the preference of A. colemani for large hosts caused a greater reduction in the population growth rate of A. glycines . This occurs in the model because, by attacking reproductive adults, A. colemani kills those aphids that have the greatest immediate effect on the population growth rate.
4. The strong effect of size preference by A. colemani is diminished somewhat by the continued reproduction of A. glycines adults, which can reproduce for up to 3 days following parasitism. Nonetheless, reproduction by parasitised aphids is not enough to compensate for the stronger, negative effect of the preference of A. colemani for large aphids, which removes individuals with the greatest reproductive value from the A. glycines population.     

Soil potassium deficiency affects soybean phloem nitrogen and soybean aphid populations

The soybean aphid is an invasive pest in the midwest United States, with frequent population outbreaks. Previous work has shown that aphid population densities are higher on potassium-deficient soybean than on healthy soybean. The experiments reported here test the hypotheses that the potassium nutrition of the host plant affects the forms of phloem nitrogen available to soybean aphids, and subsequently, their abundance. In field surveys and an exclusion cage study when aphid populations were high, soybean plants with potassium deficiency symptoms had a higher density of soybean aphids than plants without deficiency symptoms. In clip cage experiments, this effect was caused by earlier aphid reproduction and higher numbers of aphid nymphs per mother on plants growing in lower-potassium soil. In phloem exudation samples, the percentage of asparagine, an important amino acid for aphid nutrition, increased with decreasing soil potassium, perhaps because of potassium's role in the nitrogen use of the plant. Taken together, these results show that soybean potassium deficiency can lead to higher populations of soybean aphid through a bottom-up effect. A possible mechanism for this relationship is that soybean potassium deficiency improves the nitrogen nutrition of these N-limited insects. By releasing these herbivores from N limitation, host plant potassium deficiency may allow soybean aphid populations to reach higher levels more rapidly in the field.     

Costs and benefits of gregarious feeding in the meadow spittlebug, Philaenus spumarius

Abstract.  1. To study the function of group living in a non-social animal, costs and benefits of feeding in different group sizes (one to eight) were measured for nymphs of the meadow spittlebug ( Philaenus spumarius ) on tall goldenrod ( Solidago altissima ).
2. In a controlled greenhouse experiment, the number of nymphs per plant had a significant influence on their mass as adults. Adult mass increased as group size went from one to five nymphs per plant, but fell as the group size increased beyond five. In contrast, the density of nymphs on a plant had no effect on the duration of nymphal development.
3. Nymphal mortality increased linearly as the number of nymphs increased from one to eight per plant.
4. These performance results suggest that the optimal group size of spittlebug nymphs reflects a compromise between bottom-up influences (e.g. overcoming physical barriers to feeding on xylem) and within-trophic-level influences (e.g. avoiding competition for limited plant resources).
5. In a survey of local populations of goldenrod, spittlebug nymphs were found to be slightly but significantly aggregated among host-plant individuals. In addition, when more than one nymph occurred on a plant, the nymphs tended to feed in shared spittle masses.
6. It is concluded that the aggregated dispersion pattern of spittlebug nymphs reflects an adaptive balance of coping with potentially conflicting demands from different trophic levels.     

麦长管蚜的低温适应性及陕西杨凌小麦田春季虫源分析

【目的】本研究旨在探明低温条件下麦长管蚜Sitobion avenae的存活率和快速冷驯化反应,以期为该虫耐寒性的研究和准确预测预报提供依据。【方法】测定麦长管蚜实验室种群各发育阶段的过冷却点和结冰点;测定1龄若蚜和未产仔成蚜分别在-7.0～-11.0℃极端低温下暴露3 h和在0℃冷驯化1～5 h后暴露于致死温度3 h,再转移至15℃72 h后的存活率;调查自然条件下陕西杨凌小麦整个生育期麦长管蚜的种群动态。【结果】麦长管蚜1龄和2龄若蚜的过冷却点波动范围较小,分别为-27.4～-19.2℃和-27.3～-18.3℃;3龄若蚜、4龄若蚜和成蚜的过冷却点波动范围较大,分别为-27.4～-10.7℃,-26.7～-12.5℃和-26.7～-11.2℃。麦长管蚜的过冷却点和结冰点随龄期增加均显著升高,其中成蚜的过冷却点显著高于1龄和2龄若蚜。3龄若蚜、4龄若蚜和成蚜的过冷却点在不同翅型之间不存在显著性差异。低温存活率分析表明,麦长管蚜1龄若蚜和无翅成蚜的致死温度(80%死亡率)分别在-10.5℃和-8.1℃左右。0℃快速冷驯化显著提高了麦长管蚜1龄若蚜和无翅成蚜在极端低温下的存活率,其中冷驯化2 h时的存活率最高。2018-2019年小麦生育期田间调查结果表明,麦长管蚜能以孤雌生殖若蚜和成蚜在陕西杨凌越冬。【结论】麦长管蚜具有较强的低温适应能力,在陕西杨凌能以孤雌生殖蚜成功越冬。因此,其本地越冬存活个体是陕西杨凌小麦田的早春虫源之一。     

Prevalence of Pandora neoaphidis (Zygomycetes: Entomophthorales) infecting Nasonovia ribisnigri (Hemiptera: Aphididae) on lettuce crops in Argentina

Lettuce crops, Lactuca sativa, organically produced in La Plata, Argentina, were sampled in order to determine the prevalence of fungal diseased aphids. Nasonovia ribisnigri was the only aphid detected and its occurrence was highly variable. The fungal pathogen Pandora neoaphidis (Entomophthoromycotina: Entomophthorales) was the only pathogen detected. We recorded a maximum of 34.2 aphids per plant and the highest rate of fungal prevalence was 56.6% (n = 30) (aphids infected/total aphids). Infected aphids were observed in all sampling sites. No differences of infection rates were detected between the center and the edge of crops. Host density was an important factor determining infection. The majority of host population was comprised of nymphs which were the most infected in terms of individuals per habitat unit (lettuce plant), but considering the proportion of infected aphids per stage of development, the prevalence of infection in nymphs and adults was similar.     

Temperature-dependent development and population growth of Tetraneura nigriabdominalis (Homoptera: Pemphigidae) on three host plants

The root aphid Tetraneura nigriabdominalis (Sasaki) (Homoptera: Pemphigidae) is a pest of many Gramineae species; however, little is known about its biology and relationships with host plants. The objectives of this study were to quantify the effects of temperature on development, longevity, fecundity, and population growth of T. nigriabdominalis and to assess the effects of host plant on development of T. nigriabdominalis. The effects of temperature on performance of this root aphid were determined at 10, 15, 20, 25, 30, and 35 +/- 1 degrees C on rice roots, Oryza sativa L. Nymphal stages from birth to adult decreased from 46.3 d at 10 degrees C to 8.5 d at 30 degrees C. Aphid survival and development were lowest at 35 degrees C, and no aphid produced progeny at this temperature. Average adult longevity decreased from 23.3 d at 15 degrees C to 8.2 d at temperatures up to 35 degrees C. Average number of nymphs produced per female was highest at 25 degrees C; averaging near 30 nymphs per female, but it dropped to near zero at both 10 and 35 degrees C. The highest intrinsic rate of increase (r(m)) was 0.241 at 30 degrees C. Net reproductive rate (R(0)) ranged from 29.8 at 25 degrees C to 0.2 at 10 degrees C. The generation time (GT) decreased with increasing temperatures from 60.3 d at 10 degrees C to 13.8 d at 30 degrees C. In addition, root aphids reared at 30 degrees C on three host plants [O. sativa, Zea mays L. and Sorghum bicolor (L.) Moench] revealed that the developmental time of the nymphal stages averaged 6.9 d when reared on O. sativa and 10.7 d when reared on Z. mays. Comparison of the nitrogen content of the three host plants indicated that the root nitrogen content was highest in O. sativa. The effect of nitrogen content on aphid performance, however, is still not clear. Other factors, such as plant secondary chemistry, may play a role in affecting aphid performance.     

Impact of density of Aphis craccivora (Aphididae) on growth and yield of susceptible and resistant cowpea cultivars

Impact of initial density of cowpea aphid Aphis craccivora Koch (Aphididae) at infestation on the growth and yield of aphid-susceptible cowpea cultivar ICV-1 and aphid-resistant cultivar ICV-12, was investigated. Plants at the seedling, flowering and podding stages of development were infested with five aphid densities consisting of 0, 2, 5, 10 and. 20 aphids per plant and maintained for 22 days. Extended leaf heights of plants and aphid counts were recorded at 7, 12, 17 and 22 days after infestation. Two crop growth parameters (biomass duration and leaf area duration), and two plant yield parameters (number of pods per plant and number of seeds per pod) were recorded. Due to the occurrence of parthenogenesis and changes in population dynamics during infestations, aphid densities were converted into cumulative cowpea aphid-days, to facilitate data analyses and interpretation. ANOVA indicated that there was significant (P=s 0.05) difference in aphid-day accumulations between the two cultivars when infested at the seedling stage. Accumulations on cv. ICV-1 were greater than on cv. ICV-12. However, no such differences between the cultivars were detected when plants were infested at flowering and podding stages. Therefore, the seedling stage was used for comparisons of the impact of cowpea aphid-days on the growth and yield parameters of the two cultivars. At the 95% confidence intervals, ICV-12 plants were consistently taller than ICV-1 plants. Infested ICV-1 seedlings showed stunting and other growth deformities which were not observed on ICV-12 plants. Regression analyses revealed substantial reductions in the growth and yield parameters of ICV-1 relative to ICV-12. Overall, cowpea aphid-days provided a convenient and reliable method for studying the aphid population dynamics and the subsequent impact on plant growth and yield performance.     

Plant structure and the foraging success of Aphidius rhopalosiphi (Hymenoptera: Aphidiidae)

Abstract. 1. A study of searching behaviour on wheat and host preference of Aphidius rhopalosiphi (DeStefani-Perez) is described.
2. Parasitoids divided their time equally between the leaves but spent very little time on the ear.
3. After contact with honeydew or an aphid host, parasitoids were arrested in a particular area and increased the time spent searching.
4. Aphids feeding on the ear were parasitized less successfully, since their position between the grains protected them from parasitoid attacks.
5. A.rhopalosiphi exhibited no preference for Metopolophium dirhodum (Walker) or Sitobion avenae (Fabricius), although the handling time for the latter was significantly longer; this resulted in fewer S.avenae being parasitized when it was abundant. Parasitoids did not switch between hosts in these experiments.
6. As a result of its searching behaviour, A.rhopalosiphi will encounter and parasitize M.dirhodum feeding on the leaves more frequently than S.avenae, which feeds on the ear. This will limit the parasitoid's ability to regulate populations of the cereal aphid S. avenae.     

Resistance to the aphids Sitobion avenae and Rhopalosiphum padi in Gramineae in relation to hydroxamic acid levels

Aribiotic resistance to the aphid Sitobion avenae was assessed in relation to levels of hydroxamic acids (Hx) in a wide genetic range of cultivars and species of Triticum. Within hexaploid and tetraploid Triticum material, total plant concentrations of Hx explained a significant proportion of the variation in intrinsic rate of increase (r_m) of S. avenae. Significant correlations were also found between resistance to Rhopalosiphum padi and Hx levels. Although the concentrations of Hx in whole plants declined during seedling growth, concentrations of Hx in newly-emerging leaves remained high in plants of all ages, including in the emerging flag leaves of mature plants. When the mean relative growth rate of S. avenae over three days was used instead of r_m and the control of environmental conditions was improved, a higher proportion of the variation in aphid performance was explained by Hx concentrations in six cultivars.