Transgenic Bt maize and Rhopalosiphum padi (Hom., Aphididae) performance

Abstract.  1. The population abundance and age structure of Rhopalosiphum padi , one of the most common maize aphid species, on transgenic Bt (expressing the Cry1Ab protein) and non-Bt isogenic maize was studied in commercial plots during three crop seasons.
2. A higher density of aphids, particularly alates and young nymphs, occurred in Bt plots at very young maize development stages, corresponding to the settlement period, in the 3 years studied. Possible causes are discussed. After this period, there were no differences between Bt and non-Bt maize.
3. Mortality, development, and reproduction of the offspring of alate forms of R. padi and the offspring of different generations of apterous forms fed with Bt maize were evaluated in the laboratory under controlled conditions.
4. The developmental and pre-reproductive times of the offspring of the first generation of alatae were shorter and the intrinsic rate of natural increase ( r _m ) higher when aphids fed on Bt maize. The opposite occurs with the offspring of the first generation of apterous mothers, which have lower nymphal and adult mortality, shorter developmental and pre-reproductive times, a higher effective fecundity rate, and greater r _m , when fed on non-Bt maize. The differences in aphid development on the two cultivars may be linked to changes in host-plant quality due to pleiotropic effects of the genetic modification.
5. No differences on aphid mortality, developmental and pre-reproductive times, fecundity, and r _m were found between the offspring of apterous aphids maintained on Bt or non-Bt maize for several generations.     

Limited scope for maternal effects in aphid defence against parasitoids

Abstract.  1. A mother's environment frequently affects her offspring's phenotype. Such maternal effects may be adaptive, in particular with respect to pathogens or parasites, for example if maternal exposure increases offspring resistance.
2. In aphids, maternal effects are likely to occur as a result of their telescoping generations. This study investigated whether maternal effects influence the susceptibility of the peach-potato aphid, Myzus persicae (Sulzer) (Hemiptera: Aphididae), to its parasitoid Diaeretiella rapae (M'Intosh) (Hymenoptera: Braconidae: Aphidiinae).
3. In a first experiment, susceptibility was compared among offspring of aphid mothers that had either no contact to parasitoids, had contact but were not attacked, or were attacked but not mummified. Mothers from the last group had successfully resisted the parasitoid.
4. In a second experiment using two different clones, maternal and progeny environment were manipulated by rearing each generation either on a benign (radish) or a more stressful host plant (silver beet) before progeny exposure to parasitoids.
5. The first experiment revealed no significant effect of the maternal treatment on offspring susceptibility to parasitoids and thus no evidence for trans-generational defence. In the second experiment, maternal environment effects were also weak, yet with a trend towards less susceptible offspring of aphid mothers reared on the more stressful plant. However, there was a significant difference among clones and a strong clone × progeny host plant interaction, illustrating that the outcome of a parasitoid attack may be determined by a complex interplay of genetic and environmental factors.
6. Overall, the results suggest that there is limited scope for maternal effects in aphid defence against parasitoids.     

Impact of transgenic cotton plants on a non-target pest, Aphis gossypii Glover

Abstract.  1. Transgenic crops have shown great promise for the control of target pest insects, but in some cases they can also influence non-target species. This study investigated the impact of Bt and Bt+CpTI transgenic cottons on the non-target cotton aphid, Aphis gossypii Glover, by comparing life-table parameters, feeding behaviour, and the fluctuating asymmetry of morphological traits of aphids reared on transgenic cotton and those on untransformed control plants.
2. Aphids on the Bt+CpTI cotton showed a shorter reproductive duration and maximum lifespan, lower survival rates and potential maximum fecundity, and an earlier occurrence of peak daily mortality in the first or second generation. However, the aphid population soon developed fitness and overcame the negative effect in the second or third generation. The aphids on the Bt cotton had significantly longer reproductive durations in the first generation, higher survival rates in the third generation, and apparently larger potential maximum fecundity in all three generations.
3. The percentages of accumulated duration of feeding waveforms E1 and E2 were significantly lower in aphids on the Bt+CpTI cotton than in those on the Bt or control cotton, whereas the frequencies of moving and finding feeding sites, and probe behaviour were significantly higher.
4. Fluctuating asymmetry in three morphological characters of aphids reared on transgenic and control cotton was detected. The fluctuating asymmetry value of the third segment of antenna in aphids on Bt+CpTI cotton was significantly higher than that of aphids on Bt or control cotton. Based on the fluctuating asymmetry value, the stress of cotton on the aphids could be ranked as Bt+CpTI cotton > Bt cotton > control cotton.     

Population responses of a conifer-dwelling aphid to seasonal changes in its host

Abstract.  1. Current evidence suggests that seasonal changes in spruce needle sap nutrients have a decisive influence on green spruce aphid ( Elatobium abietinum ) population density, but the mechanisms of population change, the roles of development rate, fertility and mortality, and the existence of density-dependent processes, are not clearly understood.
2. Experimental studies of aphid populations were conducted in controlled environments to estimate seasonal patterns in aphid mean relative growth rate, prenatal development, fertility, and mortality. Studies were also made of the effect of aphid crowding on vital rates.
3. Independent of the degree of aphid crowding, seasonal changes in the amino acid concentration of needle sap were tracked by aphid growth rate, fertility (and adult size), but not by rates of aphid mortality. The most pronounced change in vital rates, and the one most likely to drive seasonal population change, was in fertility. Prenatal development time actually became shorter in periods when nutrients were scarce, but the resulting adult aphids were smaller and less fertile than during periods of improved nutrition.
4. Density dependence of vital rates was only observed during mid-summer when nutrients were least available. Mortality, growth rate, and prenatal development were the most strongly density-dependent processes. In contrast, there was no evidence that fertility rates were likely to respond to crowding.
5. There were no important differences between populations reared on small, potted spruce trees and those on plantation trees aged 25 years. This gives confidence that demographic data from a variety of field and laboratory sources could be used to compile data appropriate for population models.     

Preference and performance of the hyperparasitoid Syrphophagus aphidivorus (Hymenoptera: Encyrtidae): fitness consequences of selecting hosts in live aphids or aphid mummies

Abstract.  1. Theoretical models predict that ovipositional decisions of parasitoid females should lead to the selection of the most profitable host for parasitoid development. Most parasitoid species have evolved specific adaptations to exploit a single host stage. However, females of the aphid hyperparasitoid Syrphophagous aphidivorus (Mayr) (Hymenoptera: Encyrtidae) display a unique and atypical oviposition behaviour by attacking either primary parasitoid larvae in live aphids, or parasitoid pupae in dead, mummified aphids.
2. In the laboratory, the correlation between host suitability and host preference of S. aphidivorus on the host Aphidius nigripes Ashmead parasitising the aphid Macrosiphum euphorbiae (Thomas) was investigated.
3. The relative suitability of the two host stages was determined by measuring hyperparasitoid fitness parameters (survival, development time, fecundity, sex ratio, and adult size of progeny), and calculating the intrinsic rate of population increase ( r _m). Host preference by S. aphidivorus females and the influence of aphid defence behaviour on host selection was also examined.
4. Hyperparasitoid offspring performance was highest when developing from hosts in aphid mummies and females consistently preferred this host to hosts in parasitised aphids. Although aphid defensive behaviour may influence host selection, it was not a determining factor. Ecological and evolutionary processes that might have led to dual oviposition behaviour in S. aphidivorus are discussed.     

Effects of maternal age and environment on offspring vital rates in the oleander aphid (Hemiptera: Aphididae)

Maternal effects have the potential to affect population dynamics and evolution. To affect population dynamics, maternal effects must influence offspring vital rates (birth, death, or movement). Here, we explore the magnitude of nongenetic maternal influence on the vital rates of an insect herbivore and explore predictability of maternal effects with reference to published studies. We experimentally studied the effects of maternal age, host plant species (two Asclepias spp.), and density on offspring vital rates in Aphis nerii, the oleander aphid. Older mothers produced offspring that lived shorter lives, consistent with the "Lansing Effect." Older mothers also produced offspring that matured at a younger age. As maternal age increased, offspring mass at maturity decreased when mothers were on Asclepias syriaca. However, offspring mass was highest from intermediate aged mothers on A. viridis. The absence of maternal density effects seems to exclude maternal density as a potential source of delayed density dependence in A. nerii. Our results indicate that maternal effects have some influence on A. nerii vital rates. However, references to published studies suggest that only the Lansing Effect is a predictable response to maternal age in insects. Moreover, the magnitude of observed effects was generally low.     

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.     

The influences of host plant and genetic resistance to Bacillus thuringiensis on trade-offs between offspring number and growth rate in cabbage loopers, Trichoplusia ni

Abstract.  1. Variation in progeny size and quality is common among insects and this variation can strongly influence individual fitness. Larger progeny typically survive better and develop faster under adverse conditions and may have higher fecundity. Due to resource limitations, however, trade-offs may arise between having fewer large offspring or more smaller ones.
2. For cabbage loopers, Trichoplusia ni , pepper leaves are a poorer larval host than cucumber or tomato leaves as indicated by survival, development rate, and body size. Moths reared on cucumber produced more slower growing offspring than those that had been reared on pepper, which produced fewer, faster growing progeny.
3. Traits conferring resistance to Bacillus thuringiensis ( Bt ) generally are associated with strong deleterious effects that may influence resource allocation and reproductive trade-offs between progeny size and number.
4. Unlike the host-plant related trade-off between progeny size and fecundity observed among susceptible control moths, Bt -resistant parents had both the lowest fecundity and smallest progeny size on all host plants. This finding suggests that the progeny size–number relationship is constrained in resistant individuals.     

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.     

Migratory tendency in aging populations of the pea aphid,Acyrthosiphon pisum

Summary Sweep samples of the aphid, Acyrthosiphon pisum, were collected from six natural populations ranging in age from one to five years. Clones were established in the laboratory from the field-collected adults and tested for their migratory tendency in two subsequent generations by measuring the percentage of winged offspring produced in response to a standard stimulus. The number of aphids in sweep samples and the percentage of winged and wingless aphids were also determined. Tests on the first laboratory generation revealed a decline in migratory tendency with the age of the source population, but no such relationship was detected in tests on the second generation. These results are consistent with an explanation based on maternal age effects and differences in adult age structure among populations of different age. They are not consistent with one based on genetic differences among populations. Older populations also had higher densities and a lower percentage of alate adults. The percentage of larvae with wing buds was positively correlated with population density, but not population age.     

The role of cereal growth stages and crowding in the induction of alatae in Sitobion avenae and its consequences for population growth

Abstract. 1. Alatiform nymphs of Sitobion avenae were first recorded on wheat in 1977 and 1978 when there were 0.6 aphids per tiller and reached a maximum proportion of 0.9 of the population despite a seventeen-fold difference in aphid density between years.
2. At the same aphid density per tiller there was a higher proportion of alatiform nymphs on the tillers at an advanced than at an earlier stage of plant growth.
3. Both pre- and post-natal crowding, and late stages of plant growth, were important in inducing the development of alatae in the laboratory.
4. Changes in the quality of the host plant through its previously reported effect on reproductive and survival rates, and its effect on alate induction and emigration was important in shaping the population dynamics of S.avenae on cereals.     

Sublethal effects of selected insecticides on fecundity and wing dimorphism of green peach aphid (Hom., Aphididae)

Effects of sublethal concentrations (LC₂₅) of six insecticides (imidacloprid, rotenone, fenvalerate, abamectin, pirimicarb and azadirachtin) on fecundity and wing dimorphism of the green peach aphid, Myzus persicae (Sulzer), were studied both under laboratory and greenhouse conditions. In the laboratory, aphid reproduction reduced by 44.29% and 54.01% when rotenone and abamectin treatments were applied at sublethal dose, respectively, and sublethal fenvalerate application resulted in markedly lower average reproduction per female per day compared with control. Reproductive duration of aphid treated with abamectin significantly decreased by 44.19%. But in the greenhouse, there were no evident differences in the aphid fecundity and reproductive duration between treatments and control. Life‐table parameters also demonstrated that the six insecticides at sublethal doses did not stimulate the aphid reproductive potential. In the laboratory, after being exposed to sublethal doses of imidacloprid and fenvalerate, the proportions of alate progeny in aphid progeny were significantly higher than that of the control. In the greenhouse, percentages of alate offspring from the mother aphids treated with imidacloprid, fenvalerate and abamectin increased pronouncedly compared with control. Mortality rates of offspring in the nymphal stages from adults treated with insecticides revealed no significant changes between laboratory and greenhouse. The developmental time in days of the offspring varied in all treatments. Mechanisms of insecticide‐induced resurgence are discussed.     

Competition and dispersal in the pea aphid: clonal variation and correlations across traits

Abstract.  1. The presence of an across-species trade-off between dispersal ability and competitive ability has been proposed as a mechanism that facilitates coexistence. It is not clear if a similar trade-off exists within species. Such a trade-off would constrain the evolution of either trait and, given appropriate selection pressures, promote local adaptation in these traits.
2. This study found substantial levels of heritable variation in competitive ability of the pea aphid, Acyrthosiphon pisum Harris (Homoptera: Aphididae), measured in terms of relative survival when reared with a single clone of the vetch aphid, Megoura viciae Buckton (Homoptera: Aphididae).
3. Pea aphids can move to new patches by either flying (longer distance dispersal) or walking (local dispersal) from plant to plant. There was considerable clonal variation in dispersal ability, measured in terms of the proportion of winged offspring produced, and ability to survive away from their host plant.
4. Winged individuals showed longer off-plant survival times than wingless forms of the same pea aphid clone.
5. There was no evidence of a relationship between clonal competitive ability and either measure of dispersal ability, although the power of the test is limited by the number of pea aphid clones used in the trial.
6. However, there was a positive correlation between clonal fecundity and the proportion of winged offspring produced. Although speculative, it is suggested that clones that are more likely to either overwhelm their host plant or attract higher numbers of natural enemies as a result of having higher fecundity are more likely to produce winged morphs.     

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.     

Facultative symbionts are associated with host plant specialization in pea aphid populations

The pea aphid, Acyrthosiphon pisum, shows significant reproductive isolation and host plant specialization between populations on alfalfa and clover in New York. We examine whether specialization is seen in pea aphids in California, and whether fitness on alternative host plants is associated with the presence of bacterial symbionts. We measured the fitness of alfalfa- and clover-derived aphids on both types of plants and found no evidence for specialization when all aphid lineages were considered simultaneously. We then screened all aphids for the presence of four facultative bacterial symbionts: PAR, PASS, PABS and PAUS. Aphids with PAUS were host-plant specialized, having twice as many offspring as other aphids on clover, and dying on alfalfa. Other aphids showed no evidence of specialization. Additionally, aphids with PABS had 50% more offspring than aphids with PASS when on alfalfa. Thus, specialist and generalist aphid lineages coexist, and specialization is symbiont associated. Further work will resolve whether PAUS is directly responsible for this variation in fitness or whether PAUS is incidentally associated with host-plant specialized aphid lineages.     

The effect of past and current hosts on reproductive investment by the adult cowpea aphid Aphis craccivora

Abstract 1. A general index of accumulated investment that alate aphids make into reproduction with respect to environmental changes (specifically changes in host at adult moult) was investigated to determine whether and how reproduction in Aphis craccivora Koch changes once aphids migrate to a different host from the one on which they were born and reared. The aphids were monitored for 4 days to determine the immediate effects of a change in host.
2. A general index of accumulated investment defined as reproductive index was used; reproductive index is equal to the reproductive output (weight of nymphs born; actual fecundity) plus potential reproduction (weight of embryos within the ovarioles with red eye spots; future fecundity).
3. The results revealed that reproductive index is affected by adult weight. The host on which the aphids have been born and reared (source host) has no effect on reproduction other than its influence on adult weight.     

The significance of a facultative bacterium to natural populations of the pea aphid Acyrthosiphon pisum

Abstract. 1.  Laboratory studies have implicated various accessory bacteria of aphids as important determinants of aphid performance, especially on certain plant species and under certain thermal regimes. One of these accessory bacteria is PABS (also known as T-type), which is distributed widely but is not universal in natural populations of the pea aphid Acyrthosiphon pisum in the U.K.
2.  To explore the impact of PABS on the performance of A. pisum , the nymphal development time and fecundity of aphids collected directly from natural populations and caged on the host plant Vicia faba in the field were quantified. Over 4 consecutive months June–September 1999, the performance of PABS-positive and PABS-negative aphids did not differ significantly.
3.  Deterministic modelling of the performance data showed that the variation in simulated population increase of PABS-positive and PABS-negative aphids would overlap substantially.
4.  Analysis of aphids colonising five host plants ( Lathyrus odoratus , Medicago sativa , Pisum sativum , Trifolium pratense , Vicia faba ) between April and September 2000 and 2001, identified no robust differences between the distribution of PABS-positive and PABS-negative aphids on different plants and with season or temperature.
5.  It is concluded that PABS is not an important factor shaping the performance or plant range of A. pisum under the field conditions tested. Reasons for the discrepancies between this study and laboratory-based studies are considered.     

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.     

Effects of maternal diet quality on offspring performance in the rove beetle Tachyporus hypnorum

Abstract.  1. The reproductive output of an individual is known to be influenced by diet quality, but the quality of the parent's diet can also influence the performance of the offspring. Dietary maternal effects may interact with the effects of the offspring's diet to produce a variety of response patterns.
2. Maternal effects were investigated in a polyphagous predator, the rove beetle Tachyporus hypnorum , using three single-species diets: two low-quality diets consisting of the aphids Sitobion avenae and Rhopalosiphum padi , and the high-quality control Drosophila melanogaster diet. Offspring of females fed these diets were raised on the same monotypic diets and allowed to reproduce. Several fitness parameters were measured to indicate possible maternal effects in both F1 and F2 generations.
3. Maternal diet effects in F1 were found in egg size, hatching success, time to hatching, larval development time, larval survival, and sex ratio. Both aphid diets resulted in smaller eggs. A diet of R. padi resulted in reduced hatching success, longer time spent in the egg stage, and a female-biased sex ratio. A maternal diet of R. padi also prolonged larval development on S. avenae diet, while a maternal diet of S. avenae decreased survival on the R. padi diet. These effects were independent of egg size.
4. A maternal diet of R. padi enhanced the survival of F1 larvae raised on the same diet. Developmental selection operating through a high egg mortality may be the explanation for this seemingly positive effect.
5.  Sitobion avenae alone caused a significant reduction in the hatching success of F2 eggs, thus revealing grandmaternal effects.
6. The prediction that polyphagous predators are less likely to evolve adaptive maternal effects is supported by the fact that none of the documented maternal effects could be interpreted as adaptive.     

Juvenile hormone titres and winged offspring production do not correlate in the pea aphid, Acyrthosiphon pisum

Pea aphids, Acyrthosiphon pisum, reproduce parthenogenetically and are wing-dimorphic such that offspring can develop into winged (alate) or unwinged (apterous) adults. Alate induction is maternal and offspring phenotype is entirely determined by changes in the physiology and environment of the mother. Juvenile hormones (JHs) have been implicated in playing a role in wing differentiation in aphids, however until recently, methods were not available to accurately quantify these insect hormones in small insects such as aphids. Using a novel LC-MS approach we were able to quantify JH III in pea aphids that were either producing a high proportion of winged morphs among their offspring or mainly unwinged offspring. We measured JH III titres by pooling the hemolymph of 12 or fewer individuals (1 μL hemolymph) treated identically. Levels of JH ranged from 30 to 163 pg/μL. While aphids in the two treatments strongly differed in the proportion of winged morphs among their offspring, their JH III titres did not differ significantly. There was also no correlation between JH III titre and the proportion of winged offspring in induced aphids. This supports earlier findings that wing dimorphism in aphids may be regulated by other physiological mechanisms.