Aphids are frequently engaged in mutualistic associations with endosymbionts.Symbionts are bacterial or fungal microorganisms that can be obligate or facultative to aphids. Research showed facultative (or secondary) symbionts have numerous effects on their host aphids such as resistance to heat shock, parasitoids and fungus etc., which may consequently promote a co-evolution between symbionts and hosts. However, this symbiotic relation may be affected by several factors, such as the ability of symbionts to spread from aphids to others within and across populations, and the cost of infections for hosts. Moreover, aphid-symbionts interactions may be affected by aphid living environment such as its host plant, the presence of natural enemies or the temperature. Here we firstly described the functions of nine facultative symbionts (Serratia symbiotica, Hamiltonella defensa, Regiella insecticola, Rickettsia, Rickettsiella, PAXS (pea aphid X-type symbiont), Spiroplasma, Wolbachia and Arsenophonus) studied in aphids one by one, and discussed the associations between these symbionts and aphids, plants and environment. We aim to have a better knowledge of the roles the facultative symbionts play in aphid biology, ecology and evolution, which we believe can provide new inspirations for aphid control.

Odorant-binding proteins (OBPs) and chemosensory proteins (CSPs) of aphids are thought to be responsible for the initial molecular interactions during olfaction that mediate detection of chemical signals. Analysis of the diversity of proteins involved comprises critical basic research work that will facilitate the development of sustainable pest control strategies. To help us better understand differences in the olfactory system between winged and wingless grain aphids, we constructed an antennal transcriptome from winged and wingless Sitobion avenae (Fabricius), one of the most serious pests of cereal fields worldwide. Among the 133,331 unigenes in the antennal assembly, 13 OBP and 5 CSP putative transcripts were identified with 6 OBP and 3 CSP sequences representing new S. avenae annotations. We used qPCR to examine the expression profile of these genes sets across S. avenae development and in various tissues. We found 7 SaveOBPs and 1 SaveCSP were specifically or significantly elevated in antennae compared with other tissues, and that some transcripts (SaveOBP8, SaveCSP2 and SaveCSP5) were abundantly expressed in the legs of winged or wingless aphids. The expression levels of the SaveOBPs and SaveCSPs varied depending on the developmental stage. Possible physiological functions of these genes are discussed. Further molecular and functional studies of these olfactory related genes will explore their potential as novel targets for controlling S. avenae.

Wheat (Triticum aestivum L.) is one of the most cultivated crops in temperate climates. As its pests are mainly controlled with insecticides that are harmful to the environment and human health, alternative practices such as intercropping have been studied for their potential to promote biological control. Based on the published literature, this study aimed to review the effect of wheat-based intercropping systems on insect pests and their natural enemies. Fifty original research papers were obtained from a systematic search of the peer-reviewed literature. Results from a vote-counting analysis indicated that, in the majority of studies, pest abundance was significantly reduced in intercropping systems compared with pure stands. However, the occurrence of their natural enemies as well as predation and parasitism rates were not significantly increased. The country where the studies took place, the type of intercropping and the crop that was studied in the association had significant effects on these results. These findings show that intercropping is a viable practice to reduce insecticide use in wheat production systems. Nevertheless, other practices could be combined with intercropping to favour natural enemies and enhance pest control. © 2016 Society of Chemical Industry.

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Aphid saliva plays important roles in aphid-host interactions, such as assisting aphid digestion, detoxification, activating or suppressing plant defenses. The grain aphid, Sitobion avenae, is one of the most devastating pests of cereals worldwide. In this study, we performed the transcriptome analysis of salivary glands of S. avenae. A total of 33,079 assembled unigenes were identified in the salivary glands of aphids. Of the all obtained unigenes, 15,833(47.86%) and 10,829(32.73%) unigenes showed high similarity to known proteins in Nr and Swiss-Prot databases respectively. 526 unigenes were predicted to encode secretory proteins, including some digestive and detoxifying enzymes and potential effectors. The RT-PCR and RT-qPCR results showed that all of the 15 most highly expressed putative secretory proteins specifically expressed in salivary glands. Interestingly, 11 of the 15 most highly expressed putative secretory proteins were still not matched to function-known proteins. We also detected the expression of 9 interested putative secretory proteins in aphid different tissues, including some digestive and detoxifying enzymes, effectors and Ca2+ binding proteins. The results showed that only glutathione-S-transferase 1 was specifically expressed in salivary glands. These findings provide a further insight into the identification of potential effectors involving in aphid-cereals interactions.

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Infestation with Sitobion avenae induces localized defense responses in wheat; in this study, the role of S. avenae watery saliva in resistance induction was examined by infiltrating aphid saliva into wheat leaves. After feeding S. avenae on an artificial diet for 48 h, we first collected watery saliva from them and then separated the salivary proteins using one-dimensional gel electrophoresis. Gene expression studies showed that infiltration of S. avenae watery saliva in wheat leaves induced a strong salicylic acid-responsive defense but moderate jasmonic acid-dependent defense. Feeding on wheat leaves infiltrated with aphid saliva, compared with untreated leaves, significantly decreased the number of nymphs produced per day and the intrinsic rate of increase of the population of S. avenae. In a choice test against untreated wheat, saliva-infiltrated wheat had repellent effects on aphids. Additionally, electrical penetration graph results showed that the feeding behavior of S. avenae on saliva-treated wheat was negatively affected compared with that on untreated wheat. These findings provided direct evidence that salivary components of S. avenae are involved in the induction of wheat resistance against aphids and further demonstrated the important roles of watery saliva in aphid-plant interactions.

Olfaction is crucial for short distance host location and pheromone detection by insects. Complexes of olfactory receptors (ORs) are composed of odor-specific ORs and OR co-receptors (Orco). Orcos are widely co-expressed with odor-specific ORs and are conserved across insect taxa. A number of Orco orthologs have been studied to date, although none has been identified in cereal aphids. In this study, an Orco gene ortholog was cloned from the grain aphid, Sitobion avenae, and named "SaveOrco"; RNA interference (RNAi) reduced the expression of SaveOrco to 34.11% in aphids, resulting in weaker EAG (electroantennogram) responses to plant volatiles (Z-3-hexene-1-ol; methyl salicylate, MeSA) and aphid alarm pheromone (E-β-farnesene, EBF). Aphid wing differentiation induced by EBF was investigated in both RNAi treated and untreated aphids. EBF induced production of winged aphids in both pre-natal and post-natal periods in untreated aphids, but no such induction was observed in the RNAi-treated aphids. We conclude that SaveOrco is crucial for the aphid's response to pheromones and other volatiles, and is involved in wing differentiation triggered by EBF.