Reproductive stragegies of the alate and apterous morphs of the grain aphid, Sitobion avenae

Differences were observed in the total fecundity and patterns of reproduction of four morphs of Sitobion avenae (F.). The reproductive rates of apterous and alate exules declined gradually with time but decreased rapidly in gynoparae and oviparae. Under short daylengths apterous exules gave birth in three phases; the first consisting mainly of gynoparae, the second of males and the third of non-gynoparous viviparae. The strategic importance of the sequence of morphs and the reproductive behaviour of each is discussed in relation to the maximization of overwintering population size.

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Stratégies reproductives des types ailés et aptères de Sitobion avenae F.

Résumé Des différences ont été observées entre les fécondités totales et les modes de reproduction de quatre types de Sitobion avenae. Les taux de reproduction diminuent graduellement dans le temps chez les aptères et ailés virginopares et rapidement chez les gynopares et ovipares. En photophases courtes, les virginopares aptères donnent naissance à trois types: le premier composé essentiellement de gynopares, le second de mâles et le troisième de vivipares. L'importance stratégique de la succession des types et du comportement reproducteur de chacun est discutée en relation avec l'accroissement maximum de la taille de la population hivernante.

     

Mobility of apterous grain aphids Sitobion avenae within wheat fields

In a field study of the grain aphid Sitobion avenae (F.) almost all apterous aphids were found to move from the natal ear before reproducing, and alatiform nymphs usually moved between ears during their development. Apterae were very mobile, with over 50% staying on one ear for less than 24 h. Movements did not appear to be a response to climatic conditions, predation or colony density.

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Résumé Lors de l'étude de Sitobion avenae dans la nature, on a constaté que presque tous les individus aptères avaient quitté leur épi d'origine avant la reproduction, et que les larves de type ailé s'étaient déplacées généralement entre les épis pendant leur développement. Les aptères étaient très mobiles, la majorité restant moins de 24 h sur le même épi. Ces déplacements n'ont pas semblé dépendre des conditions climatiques, de la prédation ou de la densité des colonies.

     

Pattern of growth and fat content of somatic and gonadal tissues of virginoparae of the vetch aphid, Megoura viciae

Throughout the development of virginoparous apterae of the vetch aphid, Megoura viciae, the percentage water content remained constant. Over the period from birth to the onset of reproduction the somatic tissues grew logistically and the gonads exponentially. During development the percentage of the weight of an aphid that consisted of gonads increased. The sigmoidal pattern of growth in weight of an aphid reflects the combined growth patterns of the somatic and gonadal components.Large adults had proportionally bigger gonads than small adults. The fat (lipid) content of both the soma and gonads increased during development; in the soma up to the adult moult, and the gonads up to the onset of reproduction.

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Résumé Le pourcentage en eau reste constant pendant tout le développement des virginipares de Megoura viciae. De la naissance au début de la reproduction, les tissus somatiques croissent logistiquement et les gonades exponentiellement. Le poids relatif des gonades augmente pendant le développement. L'allure sigmoïde de la courbe de croissance reflète la cominaison des types de croissance des tissue somatiques et des gonades.Les gros adultes ont proportionnellement de plus grosses gonades que les petits. La teneur en corps gras augmente dans le soma et les gonades pendant le développement: dans le soma jusqu'à la mue imaginale, dans les gonades jusqu'au début de la reproduction.

     

The major carotenoid pigments of the grain aphid, Sitobion avenae (F.) (Hemiptera: Aphididae)

The economically important grain aphid, Sitobion avenae (F.) shows colour polymorphism, with brown and green forms predominating. Colour is determined both genetically and in response to environmental factors, including nutrition. The biological significance of the colour polymorphism is unknown, although seasonal changes occur in the frequency of colour morphs in the field, whilst the brown morph may have adaptive significance in terms of hymenopterous endoparasitism. The ground colour of aphids is produced by haemolymph pigments, aphins (glucosides) and carotenoids. The latter may be under the synthetic control of intracellular endosymbiotic bacteria. In this study, the major carotenoid pigments of a brown and a green clone of S. avenae were examined using thin layer chromatography (TLC) and high-performance liquid chromatography (HPLC), and their absorbance spectra recorded. Using TLC, the brown clone produced five bands of different R_f, ranging from yellow, to orange-pink to pink in colour. In contrast, the green clone gave only a single yellow band of higher R_f than any of the bands of brown aphids. Following separation of carotenoids by HPLC, brown aphids gave seven peaks and green aphids five. Comparison of absorbance maxima with known published values for carotenoids provides strong evidence for the identification of four of the carotenoid pigments from brown aphids (RB-4, 3,4-didehydrolycopene; RB-5, torulene; RB-6; lycopene; RB-7, γ-carotene) and one from green aphids (RG-2, α-carotene). The other carotenoids remain unidentified. The biosynthesis and possible biological relevance of the various pigments of S. avenae are briefly discussed.     

Life-cycle variation in the aphid Sitobion avenae: costs and benefits of male production

Abstract 1. Patterns of male production and life-cycle variation were investigated in the aphid Sitobion avenae . Seventy-seven field-collected clones were subjected to a 14 °C short day-length regime to assess their reproductive modes, and three clones were used to study life-cycle inheritance. A further analysis assessed the cost of male production.
2. In comparison with those from East Anglia, there were more holocyclic and intermediate clones found in Scottish populations, and they produced significantly more mating females. In total, 44% of clones were androcyclic.
3. The inheritance of life cycle showed a greater level of complexity than could be achieved by previously suggested monohybrid inheritance mechanisms.
4. Holocyclic and intermediate clones produced a higher proportion of males than did androcyclic clones. Inter-clonal differences were reflected in the pattern of male production in the reproductive sequence.
5. A significant cost was associated with male production in terms of a reduction in both fecundity and total offspring biomass.
6. Winged females rarely gave birth to males but the pattern of mating female production differed between holocyclic and intermediate clones.
7. Patterns of male and mating female production by the different types of clone may be related to different advantages and disadvantages of dispersal and inbreeding.
8. Recent models of aphid overwintering could be enhanced by consideration of issues raised in this study, such as the cost of male production, the inheritance of life cycle, and the patterns of sexual morph production.     

Cold hardiness and overwintering of the grain aphid Sitobion avenae

ABSTRACT.

• 1 Cold hardiness as measured by supercooling ability in the active stages of the grain aphid Sitobion avenae (F.) decreased progressively with maturation.
• 2 Aphids showed no acclimation response when maintained at low temperatures.
• 3 Starvation did not improve supercooling ability.
• 4 In a single exposure, surface moisture caused inoculation above the inherent supercooling point in a small proportion of a population.
• 5 Field populations show a seasonal change in supercooling ability, which is at a maximum in summer and a minimum in late winter.
• 6 It is concluded that the act of feeding on healthy plant tissue may confer extensive supercooling ability.

     

Methods of hatching the eggs and rearing the fundatrices of the English grain aphid, Sitobion avenae

Several methods for hatching the eggs and rearing individuals of the first generation (fundatrices) of Sitobion avenae were investigated. The most successful methods were incubation of the eggs on grass seedlings at 2°C and rearing the fundatrices on grass seedlings (overall survival 66%) and incubation of the eggs in plastic boxes at 2°C and rearing the fundatrices on wheat seedlings (overall survival 62%).

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Résumé L'éclosion des oeufs de S. avenae peut être induite par le transfert à 10°C ou 12°C, après une incubation de 75–120 jours à 2°C. Le pourcentage le plus élevé d'éclosions a été obtenu quand les oeufs avaient incubé pendant 100 à 110 jours à 2°C (67% at 71.5% respectivement) dans des petites boîtes de plastique, ou pendant 100 jours à 2°C sur des pousses de graminées (73.5%). Si les oeufs sont pondus sur blé, la plante ne peut pas tolérer la période d'incubation, mais cet obstacle peut être surmonté en obligeant les ovipares à pondre leurs oeufs sur de pousses de graminées, comme Poa annua, hôte convenable pour les fondatrices. Les ovipares peuvent aussi pondre sans difficultés sur autre chose que des végétaux, et des récipients peuvent ètre mis à incuber sans contenir du matériel végétal.

     

Functional divergence of three glutathione transferases in two biotypes of the English grain aphid,Sitobion avenae

The English grain aphid, Sitobion avenae (Fabricius) (Hemiptera: Aphididae), is a significant pest of cereal crops, but molecular factors and mechanisms that underpin its ability to develop differential biotypes on variable host plants are still not well understood. In this study, we investigated the interactions between two plant secondary metabolites (i.e., gramine and gallic acid) and three glutathione S-transferases (GSTs) of S. avenae. Using artificial diets complemented or not with one of these two plant compounds, we found that gramine had relatively stronger negative effects on fitness of S. avenae biotype 3 (adapted to barley), and gallic acid on that of biotype 4 (adapted to wheat). Gramine significantly induced overexpression of SaveGST1 and SaveGST2 in biotype 4, but not in biotype 3. Gramine also reduced SaveGST3 expression in biotype 4, but not in biotype 3, suggesting biotype-specific effect of GSTs’ regulation. In the treatments with gallic acid, the overexpression of SaveGST1, but not SaveGST2 or SaveGST3, was significantly induced in both biotypes, suggesting a critical role of SaveGST1 in detoxification of phenolic compounds such as gallic acid. The total constitutive GST activity was much higher in biotype 4 than in biotype 3. Significant increase in GST activity was obtained by the addition of both secondary metabolites in biotype 4, but not in biotype 3, showing significantly higher expression plasticity of GSTs in biotype 4. Thus, both constitutive and induced expression of GSTs could affect the adaptability of S. avenae on plants with variable secondary compounds, and thus contribute to the divergence of biotypes in this aphid species.     

Insight into watery saliva proteomes of the grain aphid,Sitobion avenae

The grain aphid, Sitobion avenae, is an economically important cereal pest worldwide. Aphid saliva plays an essential role in the interaction between aphids and their host plants. However, limited information is available regarding the proteins found in the saliva of S. avenae. Here, the watery saliva proteins from S. avenae were collected in an artificial diet and identified using a liquid chromatography–mass spectrometry/mass spectrometry analysis. A total of 114 proteins were identified in S. avenae saliva, including several enzymes, binding proteins, and putative effectors, as well as other proteins with unknown functions. In comparison with salivary proteins from nine other aphid species, the most striking feature of the salivary protein from S. avenae was the different patterns of protein functions. Several orthologous proteins secreted by other aphid species such as glucose dehydrogenase, elongation factors, and effector C002 were also detected in S. avenae saliva and speculated to play a significant role in aphid–plant interactions. These results provide further insight into the molecular basis between aphids and cereal plant interactions.     

四种类型杀虫剂对麦长管蚜的温度效应及其与主要解毒酶的关系

为了明确杀虫剂毒力受温度的影响及其程度,本文测定了4大类8种药剂在10～25℃下对麦长管蚜的毒力;并测定了麦长管蚜Sitobion avenae(Fabricius)在不同温度下2个解毒酶和1个靶标酶的活性差异。结果表明,高效氯氰菊酯对麦长管蚜表现不规则负温度系数,啶虫脒表现不规则正温度系数,高效氟氯氰菊酯对麦长管蚜的毒力不受温度影响,其他药剂(辛硫磷、毒死蜱、灭多威、丁硫克百威、吡虫啉)均表现为明显的正温度系数效应,以有机磷类杀虫剂表现最为明显,毒死蜱温度系数高达57.70。酶活性实验表明:麦长管蚜在高温下GST活性增强,羧酸酯酶和乙酰胆碱酯酶活性降低。它们的变化规律表明:GST活性与负温度系数密切相关,正温度系数与羧酸酯酶活性和靶标酶乙酰胆碱酯酶活性有关。     

Antimetabolic effect of phytohemagglutinin to the grain aphid Sitobion avenae fabricius

The insecticidal activity of plant lectins against a wide range of insect species have been intensively studied. Understanding the mechanism of the toxicity of lectins is one of the studied aspects. In the present research, the first step was determine the effect of phytohemagglutinin (PHA) on the development, fecundity and mortality of grain aphid. Next, the effect of PHA lectin on the activity of such enzymes as: α- and β-glucosidases, alkaline (AkP) and acid (AcP) phosphatases, aminopeptidase N and cathepsin L involved in the metabolism of sugar, phosphorus and proteins of an adult apterae aphids was investigated. The PHA lectin added into the liquid diet increased the pre-reproductive period, mortality of Sitobion avenae, the time of generation development and decreased its fecundity and the intrinsic rate of natural increase. In addition, activity of α-glucosidase, alkaline phosphatase and aminopeptidase N of adult apterae exposed to PHA were reduced. The results indicate that the insecticidal activity of PHA on S. avenae may involve changes in activity of the enzymes in the midgut and it may be part of its toxicity.     

Biochemical basis of winter wheat resistance to the grain aphid, Sitobion avenae

Six winter wheat cultivars with differing degrees of resistance to the grain aphid, Sitobion avenae (F.), were studied under field conditions. Resistance was measured in terms of non-preference and antibiosis on plants at seven growth stages. The varieties Saga and Grana were most resistant in terms of both non-preference and antibiosis to S. avenae at all growth stages examined. The varieties Liwilla and Dana were relatively susceptible to aphid attack. The number of aphids was directly proportional to the total content of free and essential amino acids. The level of resistance showed a similar, but not identical, relationship with the observed concentration of soluble proteins. A higher degree of cultivar resistance was associated with a higher value on a toxicity index, which is the ratio between the free-phenols and free-amino acids content. Obtained results suggest that the resistance of winter wheat cultivars to the grain aphid was based mainly on the mechanism of antibiosis.

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Résumé Six cultivars de blé d'hiver, présentant différents degrés de résistance au puceron des grains, S. avenae, ont été étudiés dans la nature. La résistance a été évaluée en fonction de l'antibiose et de l'absence de préférence de plantes à 7 stades de développement. Les variétés Saga et Grana ont été les plus résistantes, suivant ces deux critères, pour tous les stades de puceron étudiés. Les variétés Liwilla et Dana ont été relativement sensibles. Les effectifs de pucerons étaient proportionnels à la teneur totale en acides aminés libres essentiels. Les concentrations en protéines solubles présentent une relation de même type, mais pas identique, avec le niveau de résistance. Un haut degré de résistance a été associé à une valeur élevée de l'indice de toxicité, rapport entre les phénols libres et la teneur en acides aminés libres. Ces résultats suggèrent une résistance des cultivars de blé d'hiver à S. avenae liée principalement aux mécanismes d'antibiose.

     

The effect of reducing growth in winter wheat on the population dynamics of the grain aphid Sitobion avenae (F.)

1 The effect of reducing the growth of winter wheat on population size and development of the grain aphid Sitobion avenae was studied. 2 Automatic, mobile, crop shading devices, which reduced radiation comparable to levels found on a cloudy day, whilst minimizing other climatic changes, were used to reduce crop growth. Shading was applied between either GS 31 and 39 (1st node detectable → flag leaf ligule just visible) or GS 39 and 55 (flag leaf just visible → 50% of inflorescence emerged). 3 Sitobion avenae populations were initiated at GS 55 and highest populations were subsequently observed on plants shaded between GS 31 and 39. 4 Individual aphids confined within clip cages were observed to measure development from birth to adult moult, adult lifespan and fecundity. Results indicated that adult lifespan was shortest on plants shaded between GS31 and 39, whilst there were no differences in development time or fecundity. 5 Possible explanations for these observations are discussed.     

Toxic and deterrent effects of phytohemagglutinin on the grain aphid Sitobion avenae

The toxicity and deterrent effects of phytoheamagglutinin (Phaseolus vulgaris agglutinin, PHA) to the grain aphid (Sitobion avenae F.) was investigated in the laboratory by use of liquid diets or gels containing various PHA concentrations. The addition of PHA to the liquid diet increased the aphid pre-reproductive period, mortality, and generation time and decreased aphid fecundity and intrinsic rate of natural increase. Aphids fed on sucrose-agarose gels with higher concentrations of PHA exhibited no phloem-feeding behavior. The number of gel penetrations was reduced and duration of the pathway phase was increased as PHA concentration increased. These results indicate that PHA is detrimental to S. avenae and that the effect of PHA is at least partially explained by suppression of feeding behaviour. PHA could be useful for managing S. avenae.     

Bio-fertilizers and micronutrients affect the digestibility,detoxification, and intermediary metabolisms of English grain aphid,Sitobion avenae,in greenhouse

The English grain aphid, Sitobion avenae (Fabricius) (Hemiptera: Aphididae), is one of the most serious pests of cereals as the transmitter of viral diseases, mainly in wheat, Triticum aestivum. This agricultural pest is controlled with chemical insecticides, which is environmental pollutants and toxic for human. This study evaluated the potential of alternative environmentally friendly compounds to control this insect pest. The activities of digestive, antioxidants, and intermediary metabolism enzymes of S. avenae reared on wheat plants treated with bio-fertilizers (Biofarm and Probio96) and micronutrients (Librel Zinc and α-Iron) in the greenhouse were evaluated. The results reveal that α-amylase, total protease, trypsin, chymotrypsin, amino- and carboxypeptidases, alanine- and aspartate- aminotransferases, and acid phosphatase enzymes had lower activities in S. avenae reared on Biofarm treated plants compared to control. On the other hand, aphids reared on treated plants with Biofarm had the highest activities of antioxidant enzymes superoxide dismutase, catalase, peroxidase, and ascorbate oxidase, γ- glutamyl transferase, as well as lactate dehydrogenase. Moreover, the oxidized/reduced thiols ratio and malondialdehyde indicated higher activity in the aphids reared on Biofarm-treated plants than control. These results suggest that the bio-fertilizer Biofarm may compromise the physiology of S. avenae as a potential alternative toll in the integrated pest management of this aphid.     

The spatial and temporal distribution of the grain aphid Sitobion avenae in winter wheat

The spatial and temporal distribution of the grain aphid Sitobion avenae F. (Homoptera: Aphididae) was studied within a field of winter wheat during the summer of 1996. Sampling was done using four nested grids comprising 133 locations. Analysis by Taylor's power law gave results typical for insect populations. Analysis by SADIE (Spatial Analysis by Distance Indices) showed spatial pattern due to edge effects and sampling scale, and positive but mild spatial association, although spatial patterns were ephemeral. Reasons for these findings and the implications for integrated crop management are discussed.     

Effects of extreme,fluctuating temperature events on life history traits of the grain aphid,Sitobion avenae

Altered temperatures affect insects’ life history traits, such as development period and fecundity, which ultimately determine population growth rates. Understanding insects’ thermal biology is therefore integral to population forecasting and pest management decision‐making such as when to utilise crop spraying or biological control. Aphids are important crop pests in temperate regions, causing considerable yield losses. The aphid thermal‐biology literature is, however, heavily biased towards the effects of rising mean temperatures, whereas the effects of fluctuating, extreme climatic events (e.g., heat waves and sub‐zero cold periods) are largely overlooked. This study assessed the effects of laboratory‐simulated heat waves and sub‐zero cold periods on the survival, development period, and fecundity of the grain aphid, Sitobion avenae (Fabricius) (Hemiptera: Aphididae: Microsiphini), in addition to assessing maternal effects on the birth weight and development period of the offspring of exposed individuals. Exposure to heat stress periods (total of 16 h at 30 °C) significantly reduced aphid fecundity and increased physiological development period (in day‐degrees) resulting in a reduced population growth rate. Cold exposure (total of 1.33 h at ?15 °C) reduced population growth rate due to an elongated development period (in days), but did not affect fecundity or physiological development period (in day‐degrees). Both cold and heat stress significantly reduced aphid survival. Maternal experience of heat stress reduced nymphal birth weight although nymphal development period was not affected by either cold or heat stress. The results suggest that including the effects of fluctuating, extreme temperature events on aphid life history in population forecast models is likely to be of great importance to pest management decision‐making. The demonstration of maternal effects on birth weight also suggests that cross‐generational effects of heat waves on population growth rates could occur.     

The role of awns in the resistance of cereals to the grain aphid, Sitobion avenae

Six genotypes of awned spring wheat were more resistant to Sitobion avenae than de-awned wheat. Aphids that fed on awns were up to 22% less fecund and were more likely to be dislodged than aphids that fed elsewhere on an ear. These two factors reduced aphid population growth on awned earing plants in the field to a third of that observed on awnless plants. As awns may also benefit yield and are easy to select for it is recommended that awned winter wheats could be bred and used to reduce the incidence of outbreaks of S. avenae.