Indirect multi-trophic interactions mediated by induced plant resistance: impact of caterpillar feeding on aphid parasitoids

Cotton produces insecticidal terpenoids that are induced by tissue-feeding herbivores. Damage by Heliothis virescens caterpillars increases the terpenoid content, which reduces the abundance of aphids. This effect is not evident in Bt-transgenic cotton, which is resistant to H. virescens. We determined whether induction of terpenoids by caterpillars influences the host quality of Aphis gossypii for the parasitoid Lysiphlebus testaceipes and whether this interaction is influenced by Bt cotton. The exposure of parasitoids to terpenoids was determined by quantifying terpenoids in the aphids. We detected several terpenoids in aphids and found a positive relationship between their concentrations in plants and aphids. When L. testaceipes was allowed to parasitize aphids on Bt and non-Bt cotton that was infested or uninfested with H. virescens, fewer parasitoid mummies were found on infested non-Bt than on Bt cotton. Important parasitoid life-table parameters, however, were not influenced by induced resistance following H. virescens infestation, or the Bt trait. Our study provides an example of a tritrophic indirect interaction web, where organisms are indirectly linked through changes in plant metabolites.     

The influence of aphid-produced honeydew on parasitoid fitness and nutritional state: A comparative study

Honeydew is a sugar-rich resource excreted by many hemipteran species and is a key food source for other insect species such as ants and parasitoid wasps. Here, we evaluated the nutritional value of 14 honeydews excreted by 13 aphid species for the generalist aphid parasitoid Lysiphlebus testaceipes to test a series of hypotheses concerning variation in the nutritional value of honeydew. There was a positive correlation between the body sugar content of honeydew-fed parasitoids and their longevity. This information is valuable for biological control researchers because it demonstrates that the nutritional state of honeydew-fed parasitoids in the wild can indicate their fitness, independently of the honeydew source they have fed on.Although the carbohydrate content and longevity of L. testaceipes differed greatly among the different honeydews, we did not find a significant effect of aphid or host plant phylogeny on these traits. This result suggests that honeydew is evolutionarily labile and may be particularly subject to ecological selection pressures. This becomes apparent when considering host aphid suitability: Schizaphis graminum, one of the most suitable and commonly used hosts of L. testaceipes, produced honeydew of the poorest quality for the parasitoid whereas Uroleucon sonchi, one of the few aphids tested that cannot be parasitized by L. testaceipes, excreted the honeydew with the highest nutritional value. These data are consistent with the hypothesis that hemipterans are subject to selection pressure to minimize honeydew quality for the parasitoids that attack them.     

Influences of elevated CO2 and pest damage on the allocation of plant defense compounds in Bt‐transgenic cotton and enzymatic activity of cotton aphid

Abstract Plant allocation to defensive compounds by elevated CO₂‐grown non‐transgenic and transgenic Bt cotton in response to infestation by cotton aphid, Aphis gossypii (Glover) in open‐top chambers under elevated CO₂ were studied. The results showed that significantly lower foliar nitrogen concentration and Bt toxin protein occurred in transgenic Bt cotton with and without cotton aphid infestation under elevated CO_2. However, significantly higher carbon/nitrogen ratio, condensed tannin and gossypol were observed in transgenic Bt cotton “GK‐12” and non‐transgenic Bt cotton ‘Simian‐3’ under elevated CO_2. The CO₂ level and cotton variety significantly influenced the foliar nitrogen, condensed tannin and gossypol concentrations in the plant leaves after feeding by A. gossypii. The interaction between CO₂ level × infestation time (24 h, 48 h and 72 h) showed a significant increase in cotton condensed tannin concentrations, while the interaction between CO₂ level × cotton variety significantly decreased the true choline esterase (TChE) concentration in the body of A. gossypi. This study exemplified the complexities of predicting how transgenic and non‐transgenic plants will allocate defensive compounds in response to herbivorous insects under differing climatic conditions. Plant defensive compound allocation patterns and aphid enzyme changes observed in this study appear to be broadly applicable across a range of plant and herbivorous insect interactions as CO₂ atmosphere rises.     

Effects of elevated CO2 and plant genotype on interactions among cotton,aphids and parasitoids

Abstract Effects of CO₂ level (ambient vs. elevated) on the interactions among three cotton (Gossypium hirsutum) genotypes, the cotton aphid (Aphis gossypii Glover), and its hymenoptera parasitoid (Lysiphlebia japonica Ashrnead) were quantified. It was hypothesized that aphid‐parasitoid interactions in crop systems may be altered by elevated CO₂, and that the degree of change is influenced by plant genotype. The cotton genotypes had high (M9101), medium (HZ401) and low (ZMS13) gossypol contents, and the response to elevated CO₂ was genotype‐specific. Elevated CO₂ increased the ratio of total non‐structural carbohydrates to nitrogen (TNC: N) in the high‐gossypol genotype and the medium‐gossypol genotype. For all three genotypes, elevated CO₂ had no effect on concentrations of gossypol and condensed tannins. A. gossypii fitness declined when aphids were reared on the high‐gossypol genotype versus the low‐gossypol genotype under elevated CO₂. Furthermore, elevated CO₂ decreased the developmental time of L. japonica associated with the high‐gossypol genotype and the low‐gossypol genotype, but did not affect parasitism or emergence rates. Our study suggests that the abundance of A. gossypii on cotton will not be directly affected by increases in atmospheric CO₂. We speculate that A. gossypii may diminish in pest status in elevated CO₂ and high‐gossypol genotype environments because of reduced fitness to the high‐gossypol genotype and shorter developmental time of L. japonica.     

Effects of aphid honeydew sugars on the longevity and fecundity of the aphidophagous gall midge Aphidoletes aphidimyza

The predatory gall midge Aphidoletes aphidimyza, which is used for augmentative biological control of aphids in greenhouses, uses aphid honeydew as an energy source. Currently, Rhopalosiphum padi on barley plants and Melanaphis sacchari on sorghum plants are used as the banker plant systems for A. aphidimyza to control Aphis gossypii on eggplant in Japan. We analyzed the sugar components of the honeydew of these three species by HPLC. The major components of honeydew were sucrose, fructose, and melezitose for A. gossypii; glucose and fructose for R. padi; and glucose, fructose, and melezitose for M. sacchari. Maltose and trehalose were minor components of the honeydew of these species. We investigated the influence of sugars, including three artificial aphid honeydews, six sugar components, and distilled water, on the longevity of unmated A. aphidimyza. Both females and males attained the greatest longevity on sucrose and artificial honeydew of A. gossypii. Mean longevities of both females and males were shortest when they were provided only with water. We conducted another experiment in which a mated female was released in a cage with an eggplant seedling infested with A. gossypii and fed with sucrose or only water. Females fed with sucrose lived significantly longer and had higher lifetime fecundity than those fed only water.     

Response of amino acid changes in Aphis gossypii (Glover) to elevated CO2 levels

Effects of elevated CO₂ levels on the amino acid constituents of cotton aphid, Aphis gossypii (Glover), fed on transgenic Bacillus thuringiensis (Berliner) (Bt) cotton [Cryl A(c)], grown in ambient and double‐ambient CO₂ levels in closed‐dynamics CO₂ chambers, were investigated. Lower amounts of amino acids were found in cotton phloem under elevated CO₂ than under ambient CO₂ levels. However, higher amounts of free amino acids were found in A. gossypii fed on elevated CO₂‐grown cotton than those fed ambient CO₂‐grown cotton, and the contents of amino acids in honeydew were not significantly affected by elevated CO₂ levels. A larger amount of honeydew was produced by cotton aphids feeding on leaves under elevated CO₂ treatment than those feeding on leaves under ambient CO₂ treatment, which indicates that A. gossypii ingests more cotton phloem because of the higher C:N ratio of cotton phloem under elevated CO₂ levels. Moreover, the amino acid composition was similar in bodies of aphids ingesting leaves under both CO₂ treatments, except for two alkaline amino acids, lysine and arginine. This suggests that the nutritional constitution of the phloem sap was important for A. gossypii. Our data suggest that more phloem sap will be ingested by A. gossypii to satisfy its nutritional requirement and balance the break‐even point of amino acid in elevated CO₂. Larger amounts of honeydew produced by A. gossypii under elevated CO₂ will reduce the photosynthesis and result in the occurrence of some Entomophthora spp.     

Diurnal pattern of aphid feeding and its effect on cotton leaf physiology

Cotton aphid (Aphis gossypii G.) populations seemed to fluctuate over the past years in cotton (Gossypium hirsutum L.) perhaps as a result of excessive use of insecticides for controlling more problematic pests. Contradictory plant responses have been observed depending upon the aphid/plant system, and it is unclear if cotton aphids, abiotic stress or both are responsible for cotton yield reduction in aphid-infested fields. Our objectives were to investigate the diurnal changes in the physiology of cotton leaves following aphid herbivory, and the diurnal pattern of aphid feeding. The experiment was conducted in a growth chamber using the cotton cultivar ‘Stoneville 474’. Leaves of the same age and size were infested with wingless adults plus nymphs. Cotton aphids were allowed to increase in numbers without restriction for 9 days, after which the amounts of carbohydrates in aphid-honeydew, and the number of honeydew droplets excreted per aphid were measured. Photosynthetic rates, dark respiration rates and foliar non-structural carbohydrates were measured. The amount of individual carbohydrates found in the honeydew was significantly different with time. The total amount of carbohydrates excreted per aphid within a 24-h period averaged 2.5 μg. The number of honeydew droplets excreted per aphid varied significantly from time to time period. Cotton aphids did not significantly alter photosynthesis or respiration rates or non-structural carbohydrates on leaves. Aphid populations of approximately 300 per leaf on the 9th day of infestation did not appear to significantly alter the physiology of cotton leaves.     

High susceptibility of Bt maize to aphids enhances the performance of parasitoids of lepidopteran pests

Concerns about possible undesired environmental effects of transgenic crops have prompted numerous evaluations of such crops. So-called Bt crops receive particular attention because they carry bacteria-derived genes coding for insecticidal proteins that might negatively affect non-target arthropods. Here we show a remarkable positive effect of Bt maize on the performance of the corn leaf aphid Rhopalosiphum maidis, which in turn enhanced the performance of parasitic wasps that feed on aphid honeydew. Within five out of six pairs that were evaluated, transgenic maize lines were significantly more susceptible to aphids than their near-isogenic equivalents, with the remaining pair being equally susceptible. The aphids feed from the phloem sieve element content and analyses of this sap in selected maize lines revealed marginally, but significantly higher amino acid levels in Bt maize, which might partially explain the observed increased aphid performance. Larger colony densities of aphids on Bt plants resulted in an increased production of honeydew that can be used as food by beneficial insects. Indeed, Cotesia marginiventris, a parasitoid of lepidopteran pests, lived longer and parasitized more pest caterpillars in the presence of aphid-infested Bt maize than in the presence of aphid-infested isogenic maize. Hence, depending on aphid pest thresholds, the observed increased susceptibility of Bt maize to aphids may be either a welcome or an undesirable side effect.     

Host selection and parasitism behavior of Lysiphlebus testaceipes: Role of plant,aphid species and instar

The aphid parasitoid Lysiphlebus testaceipes is a potentially valuable biological control agent of Aphis gossypii a major worldwide pest of cotton. One means of increasing the abundance of a biological control agent is to provide an alternative host habitat adjacent to cropping, from which they can provide pest control services in the crop. Host selection and parasitism rate of an alternative host aphid, Aphis craccivora by L. testaceipes were studied in a series of experiments that tested its host suitability relative to A. gossypii on cotton, hibiscus and mungbean. Host acceptance, as measured by number of ovipositions was much greater in A. craccivora compared to A. gossypii, while more host aphids were accepted on mungbean than cotton. When given a choice L. testaceipes attacks more 4th instar and adult stages (63% and 70%, respectively) of both hosts than 2nd instar nymphs (47%). In a switching (host choice) experiment, L. testaceipes preferentially attacked A. craccivora on mungbean over A. gossypii on cotton. Observations of parasitoid contact with A. gossypii cornicle secretion suggest it provides a useful deterrent against parasitoid attack. From these experiments it appears L. testaceipes has a preference for A. craccivora and mungbean compared to A. gossypii and cotton, in this respect using A. craccivora and mungbean as alternative habitat may not work as the parasitoid is unlikely to switch away from its preferred host.     

Honeydew feeding increased the longevity of two egg parasitoids of the pine processionary moth

The longevity of a generalist (Ooencyrtus pityocampae) and a specialist (Baryscapus servadeii) egg parasitoid of the pine processionary moth (Thau‐metopoea pityocampa) was compared under laboratory feeding conditions including water and honeydew from aphid species growing on maritime pine (Pinus pinaster) or pedunculate oak (Quercus robur). The longevity of both parasitoid species increased when specimens were fed with honeydew. This increase was larger for the generalist (3.7, 32.0 and 38.0 days) than for the specialist (3.0, 23.3 and 21.5 days) parasitoid species when fed with water, oak and pine aphid honeydew respectively. The phenology of the specialist species B. servadeii is well‐adapted with its host availability with or without food supply. The generalist species O. pityocampae could overlap its host emergence curve during 14.0–20.0 days when fed with oak and pine aphid honeydew respectively, vs. no overlap when no supplementary food was provided. Analysis of honeydew composition indicated that sugars and amino acids may have distinct effects on parasitoid longevity.     

Effects of honeydew sugar composition on the longevity of Aphidius ervi

Feeding on sugar‐rich foods such as nectar and honeydew is important for survival of many adult parasitoids. Especially in agricultural systems, honeydew is often the most prevalent carbohydrate source. However, relative to plant nectar, honeydew may be relatively unsuitable, as a result of an unfavourable sugar composition or the presence of secondary plant compounds. We studied survival of the aphid parasitoid Aphidius ervi Haliday (Hymenoptera: Braconidae) on honeydew collected from various aphid species feeding on potato (Solanum tuberosum L., cv. Desiree) (Solanaceae), wheat (Triticum aestivum L., cv. Bobwhite) (Poaceae), or artificial diet, as well as the sugar composition of the different honeydews. Honeydews from the tested aphid species on potato, wheat, or artificial diet were found to be relatively suitable food sources for adult A. ervi, although not always as suitable as a 2 M sucrose solution. There were differences in honeydew sugar composition among the different aphid species on the various host plants. Multivariate statistics showed that the factor ‘aphid species’ had a significant influence on the sugar composition of the honeydew, explaining 27% of the variation in the potato system and 89% in the wheat system. When exploring the relationship between carbohydrate composition of the honeydews from aphids on potato and wheat plants, and their nutritional value for A. ervi, data revealed that differences in parasitoid longevity can to some extent be explained by carbohydrate composition. Furthermore, our results confirm that sucrose and its hexose components glucose and fructose are very suitable carbohydrate sources for hymenopteran parasitoids and show that parasitoid survival on an equimolar solution of the two monosaccharides glucose and fructose does not exceed performance on the disaccharide sucrose.     

A facultative mutualism between aphids and an invasive ant increases plant reproduction

1. The consequences to plants of ant–aphid mutualisms, particularly those involving invasive ants, are poorly studied. Ant–aphid mutualisms may increase or decrease plant fitness depending on the relative cost of herbivory by ant‐tended aphids versus the relative benefit of increased ant suppression of other (non‐aphid) herbivores. 2. We conducted field and greenhouse experiments in which we manipulated the presence and absence of cotton aphids (Aphis gossypii) on cotton plants to test the hypothesis that a mutualism between cotton aphids and an invasive ant, the red imported fire ant (Solenopsis invicta), benefits cotton plants by increasing fire ant suppression of caterpillars. We also manipulated caterpillar abundance to test whether the benefit of the mutualism varied with caterpillar density. 3. We found that more fire ants foraged on plants with cotton aphids than on plants without cotton aphids, which resulted in a significant reduction in caterpillar survival and caterpillar herbivory of leaves, flower buds, and bolls on plants with aphids. Consequently, cotton aphids indirectly increased cotton reproduction: plants with cotton aphids produced 16% more bolls, 25% more seeds, and 10% greater seedcotton mass than plants without aphids. The indirect benefit of cotton aphids, however, varied with caterpillar density: the number of bolls per plant at harvest was 32% greater on plants with aphids than on plants without aphids at high caterpillar density, versus just 3% greater at low caterpillar density. 4. Our results highlight the potential benefit to plants that host ant–hemipteran mutualisms and provide the first experimental evidence that the consequences to plants of an ant–aphid mutualism vary at different densities of non‐aphid herbivores.     

The nutritional value of aphid honeydew for non-aphid parasitoids

Intake of sugar-rich foods by adult parasitoids is crucial for their reproductive success. Hence, the availability of suitable foods should enhance the efficacy of parasitoids as biological control agents. In situations where nectar is not readily available, homopteran honeydew can be a key alternative food source. We studied the impact of honeydew feeding on the longevity of the larval endoparasitoids Cotesia marginiventris, Campoletis sonorensis and Microplitis rufiventris, all natural enemies of important lepidopteran pests. Females of these wasps lived longer when feeding on honeydew produced by the aphid Rhopalosiphum maidis on barley compared to control females provided with water only. However, they lived shorter than females fed with a sucrose solution. Further investigations with C. marginiventris showed that access to honeydew also increases the number of offspring produced, but less so than access to a sucrose solution. Moreover, it was found that females of this species need to feed several times throughout their life in order to reach optimal longevity and reproductive output. Analyses of the sugars in the honeydew produced by R. maidis on barley revealed that it contains mainly plant-derived sugars, but also several aphid-synthesized sugars. The sugar composition of the honeydew changed as a function of aphid colony size and time a colony had been feeding on a plant. In general, the higher the aphid infestation, the smaller the percentage of aphid-synthesized sugars in the honeydew. Experiments with honeydew sugar mimics allowed us to reject the hypothesis that the relatively poor performance of the parasitoid on a honeydew diet was due to the sugar composition. Instead, the results from additional feeding experiments with diluted honeydew showed that the nutritional value of pure honeydew is primarily restricted by its high viscosity. The possible consequences of these findings for biological pest control are discussed.     

Identification of Top-Down Forces Regulating Cotton Aphid Population Growth in Transgenic Bt Cotton in Central China

The cotton aphid Aphis gossypii Glover is the main aphid pest in cotton fields in the Yangtze River Valley Cotton-planting Zone (YRZ) in central China. Various natural enemies may attack the cotton aphid in Bt cotton fields but no studies have identified potential specific top-down forces that could help manage this pest in the YRZ in China. In order to identify possibilities for managing the cotton aphid, we monitored cotton aphid population dynamics and identified the effect of natural enemies on cotton aphid population growth using various exclusion cages in transgenic Cry1Ac (Bt)+CpTI (Cowpea trypsin inhibitor) cotton field in 2011. The aphid population growth in the open field (control) was significantly lower than those protected or restricted from exposure to natural enemies in the various exclusion cage types tested. The ladybird predator Propylaea japonica Thunberg represented 65% of Coccinellidae predators, and other predators consisted mainly of syrphids (2.1%) and spiders (1.5%). The aphid parasitoids Aphidiines represented 76.7% of the total count of the natural enemy guild (mainly Lysiphlebia japonica Ashmead and Binodoxys indicus Subba Rao & Sharma). Our results showed that P. japonica can effectively delay the establishment and subsequent population growth of aphids during the cotton growing season. Aphidiines could also reduce aphid density although their impact may be shadowed by the presence of coccinellids in the open field (likely both owing to resource competition and intraguild predation). The implications of these results are discussed in a framework of the compatibility of transgenic crops and top-down forces exerted by natural enemy guild.     

Effects of elevated CO2 on the interspecific competition between two sympatric species of Aphis gossypii and Bemisia tabaci fed on transgenic Bt cotton

Abstract Effects of elevated CO₂ (twice ambient vs. ambient) and Bt Cry1Ac transgene (Bt cotton cv. 33^B vs. its nontransgenic parental line cv. DP5415) on the interspecific competition between two ecologically similar species of cotton aphid Aphis gossypii and whitefly biotype‐Q Bemisia tabaci were studied in open‐top chambers. The results indicated that elevated CO₂ and Bt cotton both affected the population abundances of A. gossypii and biotype‐Q B. tabaci when introduced solely (i.e., without interspecific competition) or two species coexisted (i.e., with interspecific competition). Compared with ambient CO₂, elevated CO₂ increased the population abundances of A. gossypii and biotype‐Q B. tabaci as fed on Bt and nontransgenic cotton on 45 (i.e., seedling stage) and 60 (i.e., flowering stage) days after planting (DAP), but only significantly enhanced aphid abundance without interspecific competition on the 45‐DAP nontransgenic cotton and 60‐DAP Bt cotton, and significantly increased whitefly abundance with interspecific competition on the 45‐DAP Bt cotton and 60‐DAP nontransgenic cotton. In addition, compared with nontransgenic cotton at elevated CO₂, Bt cotton significantly reduced biotype‐Q B. tabaci abundances without and with interspecific competition during seedling and flowering stage, while only significantly decreasing A. gossypii abundances without interspecific competition during the seedling stage. When the two insect species coexisted, the proportions of biotype‐Q B. tabaci were significantly higher than those of A. gossypii on Bt and nontransgenic cotton at the same CO₂ levels, and elevated CO₂ only significantly increased the percentages of biotype‐Q B. tabaci and significantly reduced the proportions of A. gossypii on seedling and flowering nontransgenic cotton. Therefore, the effects of elevated CO₂ were favorable for biotype‐Q B. tabaci to out‐compete A. gossypii under the predicted global climate change.     

Comparing floral nectar and aphid honeydew diets on the longevity and nutrient levels of a parasitoid wasp

We compared the effects of floral nectar from buckwheat, Fagopyrum esculentum Moench, and honeydew produced by the soybean aphid, Aphis glycines Matsumura (Homoptera: Aphididae), on longevity, nutrient levels, and egg loads of the parasitoid Diadegma insulare Cresson (Hymenoptera: Ichneumonidae). Diadegma insulare lived for 2 days in control treatments of water or clean soybean leaves, for 6–7 days with honeydew, and in excess of 2 weeks with buckwheat nectar. Potential reasons for the superiority of buckwheat nectar over soybean aphid honeydew for extending the longevity of parasitoids include: (i) parasitoids ingest more sugars from floral sources, (ii) oligosaccharides in honeydew have a lower nutritional value than nectar sugars, and (iii) honeydew has antagonistic compounds. Overall sugar levels were lower in honeydew‐ vs. nectar‐fed female wasps, suggesting a lower feeding rate, but other explanations cannot be excluded. Diadegma insulare eclosed with high levels of lipids and glycogen, and low levels of gut and storage sugars. All carbohydrates increased over the life of both nectar‐ and honeydew‐fed wasps, but remained low or decreased in starved wasps. Lipid levels declined over the lifespan of female wasps, but females fed floral nectar showed the slowest rate of lipid decline. Diet did not affect egg load, probably because the females were not given hosts in the experiment.     

Nutritional ecology of a parasitic wasp: food source affects gustatory response, metabolic utilization, and survivorship

The success of biological control is partly mediated by the longevity and reproductive success of beneficial insects. Availability of nectar and honeydew can improve the nutrition of parasitic insects, and thereby increase their longevity and realized fecundity. The egg parasitoid, Anaphes iole, showed strong gustatory perception of trehalulose, a carbohydrate found in homopteran honeydew. Chromatographic analysis demonstrated that enzymatic hydrolysis of sucrose, a common nectar sugar, proceeded at a faster rate than that of melezitose, a sugar common in aphid honeydew. A long-term bioassay showed that longevity was greater at 20 °C than at 27 °C, and at both temperatures survival was generally greatest for wasps provisioned with the three major nectar sugars, sucrose, glucose, and fructose. Patterns of food acceptance and utilization showed that A. iole accepted and utilized a broad range of sugars found in nature, including those found in nectar as well as honeydew. Glucose, fructose, and several oligosaccharides composed of these monosaccharide units appear to be more suitable for A. iole than other sugars tested. Evidence suggests that individual fitness benefits afforded by food sources are important for a time-limited parasitoid, and that continued investigations on the interface between nutrition and biological control are warranted for A. iole.     

Assessment of cotton aphids, Aphis gossypii, and their natural enemies on aphid-resistant and aphid-susceptible wheat varieties in a wheat–cotton relay intercropping system

The cotton aphid, Aphis gossypii Glover (Homoptera: Aphididae), is an important cotton pest in northern China, especially in the seedling stage of cotton. After large scale commercial use of transgenic Bt cotton, cotton aphids became one of the most important cotton pests. A 2‐year study was conducted to evaluate the role of four winter wheat varieties that were resistant or susceptible to wheat aphid, Sitobion avenae Fabricius (Homoptera: Aphididae), in conserving arthropod natural enemies and suppressing cotton aphids in a wheat–cotton relay intercropping system in northern China. The results indicated that wheat–cotton intercropping preserved and augmented natural enemies more than a monoculture of cotton. The density of natural enemies in cotton was significantly different among relay‐intercropping fields with different wheat varieties. The highest density of natural enemies and low cotton aphid populations were found in the treatment of cotton in relay intercropped with the wheat variety Lovrin10, which is susceptible to wheat aphid. The lowest density of predators and parasitoids associated with high cotton aphid populations were found with the wheat variety KOK1679, which is resistant to wheat aphid. The results showed that wheat varieties that are susceptible or moderately resistant to wheat aphid might reduce cotton aphids more effectively than an aphid‐resistant variety in the intercropping system by enhancing predators to suppress cotton aphids during the cotton seedling stage.     

The parasitoid of a fruit moth caterpillar utilizes fruit components as nutrient source to increase its longevity and fertility

Longevity and fecundity of female wasps are two decisive factors for the effectiveness of parasitoid species as biological control agents. Accessibility and suitability of nutrient sources determine parasitoid survival and reproduction. Host, nectar and honeydew feeding are frequent adult parasitoid behaviors to cover nutritional needs. Here we postulate that especially parasitoid species of endophytic herbivores might use plant tissue as a nutrient source that becomes accessible upon herbivory. We investigated the influence of plant consumption and host feeding on longevity and fecundity of Hyssopus pallidus, a gregarious ecto-parasitoid of caterpillars of the codling moth that feed inside apple fruits. Longevity of unmated and mated ovipositing female parasitoids was highest in treatments with fruit pulp. While longevity in this treatment was not significantly different from that with honey, it was significantly higher than in treatments without food, with water or with a host alone.Reproduction was significantly increased by these sugar-rich nutrient sources compared to the control with a host alone. In contrast, host feeding did not yield any significant contribution to longevity and fecundity in a series of bioassays with different host–parasitoid ratios and with differently aged and sized hosts, compared to controls without food.We conclude that in this synovigenic species host feeding does not contribute to longevity and fecundity, but females can increase survival and reproduction in the field relying solely on the plant tissue damaged by their host caterpillar.     

Effects of cotton cultivars differing in gossypol content on the quality of Aphis gossypii as prey for two species of Coccinellidae

Plants can have detrimental effects on biological control agents by affecting their prey or host quality. Thus, it is important to understand the tri-trophic interactions between plants, herbivores and natural enemies when implementing biological control programmes. Studies have shown that both morphological and chemical traits of host plants can affect the third trophic level. Cotton plants are known to produce alkaloids such as gossypol, a sesquiterpene aldehyde that can confer resistance against herbivorous arthropods. Nevertheless, little is known about the effect of gossypol on biological control agents. In this study, we investigated how three cotton cultivars (BRS Rubi, BRS Safira and BRS Verde) differing in gossypol content affect development and growth of predatory coccinellids, Eriopis connexa and Harmonia axyridis, feeding on the cotton aphid Aphis gossypii reared on those cultivars. The results show that the cultivar BRS Rubi (highest gossypol content) had a sub-lethal effect on the development and growth of both Coccinellidae species compared with the other cultivars. Overall, the cultivar BRS Rubi reduced slightly fecundity, net reproductive rate and intrinsic rate of natural increase for both Coccinellidae species. However, because aphid populations stay short periods of time in the field, and adult coccinellids may supplement its diet with alternative prey and plant material this sub-lethal indirect effect of gossypol may not have a detrimental effect on field biological control of cotton aphid by either E. connexa or H. axyridis, thus suggesting a compatibility between plant resistance and biological control agents.