Effects of maternal diet on offspring fitness in the bird cherry‐oat aphid

1. Maternal and offspring diet effects on life‐history traits of the bird cherry‐oat aphid Rhopalosiphum padi were tested on three wheat varieties. Using nine reciprocal combinations of wheat varieties, the effects of previous experience (maternal diet effect) on the aphid's response to resistant and susceptible varieties (offspring diet effect) were tested. Batis was susceptible, and Xiaoyan22 and Ww2730 were both resistant, but with different mechanisms. 2. Aphids produced the most alatae in the treatments with the most resistant maternal diet variety Xiaoyan22. The fecundity (F) and intrinsic rate of increase (r_m) of these alatae were at their greatest in the most resistant offspring diet variety, but these traits were not influenced in the apterae. 3. There were significant interactions in the alatae production and apterae life‐history traits, such as r_m, development time, weight gain, and mean relative growth rate, between the maternal and offspring diet varieties. The interactions in apterae responses between varieties, some of which were reciprocal, indicated phenotypic plasticity in these parthenogenetic aphids. 4. Rhopalosiphum padi produced more alatae on the most resistant variety; the alatae would disperse and were more fecund. The growth responses of the apterae showed phenotypic plasticity to the different combinations of maternal and offspring diet varieties. The phenotypic plasticity would allow R. padi to better utilise the variable environments represented by the small wheat plots of different varieties in China.     

Intra-specific mechanisms in relation to the natural regulation of numbers of Aphis fabae Scop

Parthenogenetic virginoparous apterae of Aphis fabae Scop. on field beans (Vicia faba) reproduced faster initially in populations of eight colonizing apterae than in those with 2–4 or 16–32 per plant. The aphids were at first mutually benefited but were quickly affected by competition as numbers rose above the critical density represented by about eight apterae and their first progeny. This is because the aphids remained densely aggregated and seemingly created a local shortage despite abundant food and space elsewhere on the plant. Such self-induced competition provides the basis for self-regulation of numbers of A. fabae in relation to (1) food and space provided by the growing plant and (2) mortality from natural enemies and from other causes including insecticides. As competition increased, the multiplication of A. fabae populations slowed, newly formed adult apterae emigrated and increasing numbers of alatae were formed. The mean weights of apterae decreased from about 1·8 mg. to 0·3 mg. and of alatae from 0·9 to 0·2 mg. Such decrease probably favours production of many adults that might otherwise fail to mature. Experiments in a glasshouse and in field cages indicated the success with which an A. fabae population adapts to and exploits a growing plant. Field bean plants sown in mid-March and infested as in the field produced an average of 15,000–17,000 A. fabae emigrants per plant of which 78–84% were adults (mostly alatae). This is equivalent to about 1600 million alate emigrants from 1 acre (0·4 hectare) of an infested field bean crop.     

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.     

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.     

Effect of mixed viral infections (potato virus Y-potato leafroll virus) on biology and preference of vectors Myzus persicae and Macrosiphum euphorbiae (Hemiptera: Aphididae)

Mixed viral infections of heterologous viruses such as Potato virus Y (family Potyviridae, genus Potyvirus, PVY) and Potato leafroll virus (family Luteoviridae, genus Polerovirus, PLRV) are a regular occurrence in Idaho's potato, Solanum tuberosum (L.), cropping systems. An increased number of plant samples from Idaho's potato fields over the past 2 yr has serologically tested positive for both PVY and PLRV via double antibody sandwich enzyme-linked immunosorbent assay (DAS-ELISA) and exhibited more severe symptoms than singly-infected plants (PVY or PLRV). Several studies have extensively examined the mixed infection phenomenon but to the best of our knowledge, none have examined the effect of such infections on vector biology and preference. Laboratory studies were conducted to examine the effect of mixed viral (PVY-PLRV) infection on the fecundity and preference of two of the most efficient PVY and PLRV vectors, the green peach aphid, Myzus persicae (Sulzer), and the potato aphid, Macrosiphum euphorbiae (Thomas) (Hemiptera: Aphididae). M. persicae and M. euphorbiae adults were clip-caged (one adult per cage) to leaflets of PVY, PLRV, PVY-PLRV-infected, and noninfected potato plants. The number of nymphs produced in all four treatments was recorded after 96 h. M. persicae and M. euphorbiae fecundity was significantly higher on mixed infected plants than on singly infected plants or noninfected plants. Preference of alatae and apterae of M. persicae and M. euphorbiae was determined with the use of settling bioassays. Both alatae and apterae of M. persicae and M. euphorbiae preferentially settled on PVY-PLRV-infected plants than on singly infected plants (PVY or PLRV) or noninfected plants.     

The combined use of mineral oils and pyrethroids to control plant viruses transmitted non- and semi-persistently by Myzus persicae

A mixture of the pyrethroid WL85871 (an enriched form of cypermethrin) and the mineral oil SC811 intoxicated adult apterae of an insecticide-susceptible clone of the peach-potato aphid Myzus persicae at a similar rate to a treatment containing only WL85871, but the mixture killed more. Mixtures of WL85871 andSC811 also gave better control of both acquisition and inoculation by M. persicae of the non-persistent potato virus Y (PVY) than either component alone. A mixture of the pyrethroids deltamethrin, cypermethrin and PP321 with SC811, or a mixture of WL85871 with the mineral oil Bayol 52 also decreased acquisition of PVY, and a mixture of WL85871 with SC811 decreased acquisition of another non-persistently transmitted virus, beet mosaic virus. Control with mixtures was generally better than that provided by each component applied separately. When testing acquisition or inoculation of the semi-persistent beet yellows virus, fewest plants were infected in treatments incorporating both WL85871 and SC811.     

Potato virus Y reduction by straw mulch in organic potatoes

Potato virus Y (PVY) is transmitted non-persistently by winged morphs of many aphid species and is a major problem in seed potato production. In order to evaluate the potential of straw mulch applications (4–5 t ha⁻¹) and presprouting on PVY reduction, small scale organically managed field experiments were carried out in Northern Hessen, Germany, over 3 yr. In all years mulching significantly reduced aphid infestation on leaves as well as PVY incidence in tubers. For the effect of presprouting the temporal coincidence of two factors was crucial – crop emergence and aphid flight activity. Presprouting decreased PVY incidence when in the phase of early crop emergence aphid spring flight activity was low, but increased it, although not significantly, when prominent aphid flight peaks occurred in this critical period. Straw mulch was most effective when vector pressure was concentrated early in the year acting as a PVY protectant for young plants. In later growth stages its effect declined gradually with increasing ground coverage of the crop. Combined mulching and presprouting had a synergistic, complementary effect on reduction of PVY incidence. In an on-farm experiment in 2001 scaling up the area mulched stepwise from 100 m² to 900 m² consistently kept aphid infestation at reduced levels.     

Precocene II-induced alate production in isolated and crowded alate and apterous virginoparae of the aphid, Macrosiphum euphorbiae

Precocene II was applied at doses ranging from 0.05 to 0.70 μg per individual to newly moulted adult alate and apterous virginoparae of Macrosiphum euphorbiae kept isolated or in groups of 10 per plant, under an 18L:6D photoperiodic regime. While isolated controls of both morphs produced exclusively apterous progeny, alatae virginoparae were produced in generally dose-dependent proportions by precocene-treated individuals. Grouped controls of both morphs produced alatiform progeny as expected, but in precocene—treated groups, the proportions of alate progeny generally increased as a function of dose. The overall proportions of alate offspring produced, and numbers of days after treatment when morph production was affected, were generally greater for alatae than for apterae, indicating a greater sensitivity to precocene in alatae. However during the first few days after treatment, the alatizing effect of precocene was stronger for apterae, suggesting that the first embryos produced by alatae were irreversibly determined as apterae.In an experiment where isolated alatae and apterae received 0.5 μg of precocene II at different ages ranging from 1 to 13 days after the adult moult, the alatizing effect of the compound, measured by the persistence of alate production, varied with age and morph. While in alatae, the persistence decreased more or less regularly with age, in apterae it initially increased to a maximum in the middle of reproductive life, and subsequently decreased. The results provide support for the hypothesis that juvenile hormone is involved in regulating alary dimorphism in M. euphorbiae.     

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.     

豆蚜有翅蚜产生的原因

本文研究拥挤、寄土质量、温度和蚜型等因子对大豆蚜(Ahis glyeines)有翅蚜产生的影响。结果表明：1.大豆蚜无翅胎生成好个体间的拥挤是有翅蚜产生的主要原因。在低密度下拥挤反应随密度增大而增强,但过度拥挤会导致反应的降低。无翅若好间的拥挤不能导致其本身发育为有翅胎生蚜。 2.寄主质量能改变无翅胎生成蚜对拥挤的反应。每笼2头经成熟叶片处理的无翅胎生成蚜后代中有姻蚜的比例高于幼嫩叶片和对照(无叶片)处理,且饥饿不能促进有翅蚜的产生。3.温度能影响有翅蚜的产生。较高的温度(30℃和25℃)较21℃对有翅胎生蚌的产生有较强的抑制作用。4.不同母蚜型产生有翊蚜的能力不同。有翅胎生蚜间的拥挤也能使其在后代中产生少量的有翊蚜,但对拥挤的敏感程度低于无翅胎生蚜。     

The cowpea aphid, shape Aphis craccivora, host plant odours and pheromones

Olfactometer experiments were conducted with apterae and alatae of the cowpea aphid, Aphis craccivora, Koch (Hom.:Aphidoidea). The occurrence of density dependent related pheromones and odour response to their host plant Vigna unguiculata was studied. Apterae responded with positive anemotaxis to air passed over both small groups of apterae and alatae (10 individuals) but negatively to air that passed over a bigger group (20 individuals). Alatae responded in the same way to groups of apterae but were repelled by alatae independent of the size of the group applied as odour source in the olfactometer, except when exposed to air passing ten alatae feeding on cowpea leaves. This may be due to interaction between aphid pheromones and host plant odour. Both apterae and alatae showed a general attraction to green plants and were also able to distinguish between other host plants and cowpea, which was the original host of the aphid clone used in the experiments. When attacked by aphids, the cowpea plant responded with a temporal increase in attractivity that reached a maximum after 48 h and had disappeared after one week.     

Dark respiration of leaves and traps of terrestrial carnivorous plants: are there greater energetic costs in traps?

In this study, O₂-based dark respiration rate (R_D) in leaf and trap cuttings was compared in 9 terrestrial carnivorous plant species of 5 genera to decide whether traps represent a greater energetic (maintanence) cost than leaves or photosynthetic parts of traps. R_D values of cut strips of traps or leaves of terrestrial carnivorous plants submerged in water ranged between 2.2 and 8.4 nmol g⁻¹ s⁻¹ (per unit dry weight) in pitcher traps of the genera Sarracenia, Nepenthes, and Cephalotus, while between 7.2 and 25 nmol g⁻¹ _DW s⁻¹ in fly-paper or snapping traps or leaves of Dionaea and Drosera. No clear relationship between R_D values of traps (or pitcher walls) and leaves (or pitcher wings or petioles) was found. However, RD values of separated Drosera prolifera tentacles exceeded those of leaf lamina 7.3 times.     

Resistance to the aphid Metopolophium festucae cerealium: effects of the host plant on flight and reproduction

The levels of resistance present in two cereal and four grass hosts of M. f. cerealium were assessed in terms of the percentage of alatae flying from the host and reproductive rates of alatae. Festuca rubra and F. arundinacea exhibited highest levels of resistance, inducing a greater proportion of the alatae to fly (94% and 84% respectively) after producing only a few nymphs. Although between 38% and 64% of alatae flew from L. multiflorum, oats, L. perenne and wheat these alatae deposited more nymphs on the hosts before flight than occurred on Festuca. The weight of alatae did not influence the flight response. The differences in the flight/reproductive response of M. festucae to various host plants are discussed in relation to possible host selection strategies of this species and to antixenotic/antibiotic host resistance experiments in general.     

Alata-production by an aphid: The “interval timer” concept and maternal age effects

Throughout their lives alate Acyrthosiphon pisum aphids remained unable to produce more alatae in response to crowding. Similarly, the first-born progeny of most alatae were likewise unable to respond to crowding. In contrast, the first-born progeny of apterae responded exceptionally strongly to the crowding stimulus. Later-born progeny of both morphs responded similarly. These data are inconsistent with Lees' theory of “interval timers” since ability to produce alatae did not return simultaneously in all lines founded by alatae. The theory of interval timers must be either replaced or modified to take into account differences related to maternal age.     

Embryonic growth rate and birth weight of the offspring of apterous and alate aphids: a cost of dispersal

Apterous and alate Sitobion avenae (Fab.) were dissected from the third instar onwards and the size and degree of development of their largest embryos recorded. From at least the third instar apterae had larger and more well developed embryos than alatae. Embryos in both morphs showed an exponential increase in volume with time during their mothers' nymphal development, but those in apterous mothers grew faster. Although alatae had a longer pre-reproductive development than apterae, it was not long enough to compensate for the lower embryo growth rates in alatae and as a consequence they initially produced smaller and fewer offspring than apterae. After 4 days of reproduction apterous and alate mothers produced similar sized offspring and at the same rate. The total fecundity of apterae was greater than that of alatae.

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Résumé La taille et l'état de développement des plus gros embryons de Sitobion avenae Fab ont été examinés par dissection d'ailés et aptères à partir du troisième stade. La taille des embryons des 2 morphes augmente pendant le développement larvaire des mères, mais ceux des mères aptères grossissent plus vite. Bien que la période précédant la reproduction des alates soit plus longue chez les ailés que chez les aptères, les embryons des ailés sont néanmoins incapables de l'emporter en taille sur les aptères, et les ailés produisent initialement moins de pucerons et plus petits que les aptères. Après 4 jours de reproduction, les mères ailées et aptères produisent des pucerons de même taille au même rythme. La fécondité totale des aptères est supérieure à celle des ailés; la différence est liée à la biologie des 2 morphes.

     

Alatae production and population increase of aphid vectors on virus-infected host plants

Summary Zucchini yellow mosaic virus (ZYMV) and Aphis gossypii Glover are two components of a recently identified plant-parasite system that provides an excellent opportunity to study interrelations between a virus and a vector that share the same host, but have no direct physiological interaction. In a field experiment we documented numbers of alate and apterous A. gossypii on healthy Cucurbita pepo and on plants inoculated with virus 0, 7, 14, and 21 days before aphid infestation. When plants were inoculated and infested simultaneously, more than twice as many alatae were produced after 20 days of colony growth than on any other treatment. This indicates that properties unique to the early stages of viral infection somehow stimulated wing formation. Because it is spread by the activities of alatae, virus dispersal would be greater as a result of these properties. Developmental rate, total numbers of aphids, and numbers of alatae and apterae decreased as the time between virus inoculation and aphid colonization increased.     

Observations on the morphs of Macrosiphum avenae and Metopolophium dirhodum on cereals during the summer and autumn

From June to early August 1970, populations of Macrosiphum avenae and Metopolophium dirhodum on marked tillers of field barley were compared with the numbers of alatae trapped at crop height and at 12.2 m. There were always more M. dirhodum than M. avenae on the tillers. Only apterae were produced until mid-June when both alatae and apterae occurred; after mid-July only alate M. avenae were found. Until mid-June most of the flying alatae were caught at 12.2 m as they migrated from spring hosts to cereals. Thereafter, more alate M. avenae were trapped at 12.2 m than at crop level, whereas numbers of alate M. dirhodum were usually comparable at both heights. Although crop and flying populations occasionally showed temporal similarities, insufficient is known about their field distribution and the factors affecting their alate production and flight activity to interpret this relationship. In the autumn, two consecutive reproductive phases of M. dirhodum occurred on winter wheat grown in pots outdoors. Initially, apterous virginoparae and alatae, probably sexuparae, were produced, whereas only alate males appeared during the second phase. In contrast, M. avenae deposited mainly apterous virginoparae although some oviparae developed in October to lay scattered, probably infertile eggs on the tillers.     

Effects of infection with the cabbage black ringspot strain of turnip mosaic virus on turnip as a host to Myzus persicae and Brevicoryne brassicae

When turnip plants with 3–7 leaves were inoculated with cabbage black ringspot virus (CBRSV) on the 3rd rough-leaf, symptoms only appeared on leaves that had been less than 15 mm long at the time of inoculation, although infection decreased the area and both fresh and dry weight of all leaves. Leaves were ‘aged’ by their appearance and placed in Leaf Age Categories (LACs). Leaves with symptoms senesced (‘aged’) prematurely. CBRSV-infection of cv. Green Top White did not change the distribution of populations of Myzus persicae between LACs, but increased the proportion of the plant suitable for colonisation. All suitable LACs were quickly colonised by adult apterae and nymphs. On CBRSV-infected plants the nymphal period was shorter, F₁ adults deposited larvae more frequently and the live body weight and tibial length of the F₂ generation was greater, than on healthy plants. The distribution of Brevicoryne brassicae populations on cv. Green Top White differed from that of M. persicae but was also unchanged by CBRSV-infection. On healthy plants the largest colonies were on mature leaves, so that on virus-infected plants premature senescence shortened the life of the colony. On CBRSV-infected plants the nymphal period was prolonged and the live weight of F₁ and F₂ adult apterae was less than on healthy plants. The differences between the biology of M. persicae and B. brassicae on CBRSV-infected cv. Green Top White were associated with the accelerated senescence of CBRSV-infected leaves. The possibility that CBRSV-infection might reduce the resistance of turnips to aphid infestation was tested. M. persicae and B. brassicae were cultured on two favourable and two less favourable cultivars. No improvement in population growth rate was found when the less favourable host cultivars were infected with CBRSV, but both aphid species weighed less and/or had smaller nymphal populations on cultivars showing the severest symptoms. These results are discussed in relation to the evolution of non-persistent virus transmission by aphids.     

Photosynthesis and dark respiration of leaves of terrestrial carnivorous plants

Using CO₂ gasometry, net photosynthetic (P _N) and dark respiration rates (R _D) were measured in leaves or traps of 12 terrestrial carnivorous plant species usually grown in the shade. Generally, mean maximum P _N (60 nmol CO₂ g⁻¹(DM) s⁻¹ or 2.7 μmol m⁻² s⁻¹) was low in comparison with that of vascular non-carnivorous plants but was slightly higher than that reported elsewhere for carnivorous plants. After light saturation, the facultatively heliophytic plants behaved as shade-adapted plants. Mean R _D in leaves and traps of all species reached about 50% of maximum P _N and represents the high photosynthetic (metabolic) cost of carnivory.     

Chlorophyll fluorescence imaging of photosynthetic activity in sun and shade leaves of trees

The differences in pigment levels, photosynthetic activity and the chlorophyll fluorescence decrease ratio R _Fd (as indicator of photosynthetic rates) of green sun and shade leaves of three broadleaf trees (Platanus acerifolia Willd., Populus alba L., Tilia cordata Mill.) were compared. Sun leaves were characterized by higher levels of total chlorophylls a + b and total carotenoids x + c as well as higher values for the weight ratio chlorophyll (Chl) a/b (sun leaves 3.23–3.45; shade leaves: 2.74–2.81), and lower values for the ratio chlorophylls to carotenoids (a + b)/(x + c) (with 4.44–4.70 in sun leaves and 5.04–5.72 in shade leaves). Sun leaves exhibited higher photosynthetic rates P _N on a leaf area basis (mean of 9.1–10.1 μmol CO₂ m⁻² s⁻¹) and Chl basis, which correlated well with the higher values of stomatal conductance G _s (range 105–180 mmol m⁻² s⁻¹), as compared to shade leaves (G _s range 25–77 mmol m⁻² s⁻¹; P _N: 3.2–3.7 μmol CO₂ m⁻² s⁻¹). The higher photosynthetic rates could also be detected via imaging the Chl fluorescence decrease ratio R _Fd, which possessed higher values in sun leaves (2.8–3.0) as compared to shade leaves (1.4–1.8). In addition, via R _Fd images it was shown that the photosynthetic activity of the leaves of all trees exhibits a large heterogeneity across the leaf area, and in general to a higher extent in sun leaves than in shade leaves.