Diversity,frequency, and geographic distribution of facultative bacterial endosymbionts in introduced aphid pests

Facultative bacterial endosymbionts in insects have been under intense study during the last years. Endosymbionts can modify the insect's phenotype, conferring adaptive advantages under environmental stress. This seems particularly relevant for a group of worldwide agricultural aphid pests, because endosymbionts modify key fitness‐related traits, including host plant use, protection against natural enemies and heat tolerance. Aimed to understand the role of facultative endosymbionts on the success of introduced aphid pests, the distribution and abundance of 5 facultative endosymbionts (Hamiltonella defensa, Regiella insecticola, Serratia symbiotica, Rickettsia and Spiroplasma) were studied and compared in 4 cereal aphids (Sitobion avenae, Diuraphis noxia, Metopolophium dirhodum and Schizaphis graminium) and in the pea aphid Acyrthosiphon pisum complex from 2 agroclimatic zones in Chile. Overall, infections with facultative endosymbionts exhibited a highly variable and characteristic pattern depending on the aphid species/host race and geographic zone, which could explain the success of aphid pest populations after their introduction. While S. symbiotica and H. defensa were the most frequent endosymbionts carried by the A. pisum pea‐race and A. pisum alfalfa‐race aphids, respectively, the most frequent facultative endosymbiont carried by all cereal aphids was R. insecticola. Interestingly, a highly variable composition of endosymbionts carried by S. avenae was also observed between agroclimatic zones, suggesting that endosymbionts are responding differentially to abiotic variables (temperature and precipitations). In addition, our findings constitute the first report of bacterial endosymbionts in cereal aphid species not screened before, and also the first report of aphid endosymbionts in Chile.     

The price of protection:a defensive endosymbiont impairs nymph growth in the bird cherry-oat aphid,Rhopalosiphum padi

Bacterial endosymbionts have enabled aphids to adapt to a range of stressors,but their effects in many aphid species remain to be established.The bird cherry-oat aphid,Rhopalosiphum padi(Linnaeus),is an important pest of cereals worldwide and has been reported to form symbiotic associations with Serratia symbiotica and Sitobion miscanthi L-type symbiont endobacteria,although the resulting aphid phenotype has not been described.This study presents the first report of R.padi infection with the facultative bacterial endosymbiont Hamiltonella defensa.Individuals of R.padi were sampled from populations in Eastern Scotland,UK,and shown to represent seven R.padi genotypes based on the size of polymorphic microsatellite markers;two of these genotypes harbored H.defensa.In parasitism assays,survival of H.defensa-infected nymphs following attack by the parasitoid wasp Aphidius colemani(Viereck)was 5 fold higher than for uninfected nymphs.Aphid genotype was a major determinant of aphid performance on two Hordeum species,a modern cultivar of barley H.vulgare and a wild relative H.spontaneum,although aphids infected with H.defensa showed 16%lower nymph mass gain on the partially resistant wild relative compared with uninfected individuals.These findings suggest that deploying resistance traits in barley will favor the fittest R.padi genotypes,but symbiontinfected individuals will be favored when parasitoids are abundant,although these aphids will not achieve optimal performance on a poor quality host plant.     

Host genotype–endosymbiont associations and their relationship with aphid parasitism at the field level

1. The relationship between endosymbionts and insects represent complex eco‐evolutionary interactions. Vertically transmitted endosymbionts can be a source of evolutionary novelty by conferring ecologically important traits to their insect hosts, such as protection against natural enemies. Host–endosymbiont associations could constitute an adaptive complex (holobiont) on which selective pressures present in the environment can act, being transferred to the next generation. 2. Although several laboratory‐based studies have confirmed host genotype × symbiont interactions, few studies have been directed at those associations in the natural populations and their ability to protect themselves from parasitism pressure at the field level. 3. A field‐based approach to study the aphid genotype–endosymbiont associations and its relationship with the total parasitism in the grain aphid Sitobion avenae was conducted. From the field study, experiments were carried out to study the defensive effect of the two most common facultative endosymbionts (Regiella insecticola and Hamiltonella defensa) present in S. avenae against one of the most important parasitoid species, Aphidius ervi. 4. Evidence is presented here of a high specificity of the aphid clone–endosymbiont associations in the field; however, the field and experimental results here do not support a relationship between the aphid clone–endosymbiont associations and a proxy of total parasitism in S. avenae. These findings highlight the importance of particular host clone–endosymbiont couplings as a key factor in gaining an understanding of the coevolutionary dynamics of endosymbionts in nature and their effect on the invasive potential of pest insects.     

A facultative endosymbiont in aphids can provide diverse ecological benefits

Ecologically important traits of insects are often affected by facultative bacterial endosymbionts. This is best studied in the pea aphid Acyrthosiphon pisum, which is frequently infected by one or more of eight facultative symbiont species. Many of these symbiont species have been shown to provide one ecological benefit, but we have little understanding of the range of effects that a single strain can have. Here, we describe the phenotypes conferred by three strains of the recently discovered bacterium known as X‐type (Enterobacteriaceae), each in their original aphid genotype which also carries a Spiroplasma symbiont. All comparisons are made between aphids that are coinfected with Spiroplasma and X‐type and aphids of the same genotype that harbour only Spiroplasma. We show that in all cases, infection with X‐type protects aphids from the lethal fungal pathogen Pandora neoaphidis, and in two cases, resistance to the parasitoid Aphidius ervi also increases. X‐type can additionally affect aphid stress responses – the presence of X‐type increased reproduction after the aphids were heat‐stressed. Two of the three strains of X‐type are able to provide all of these benefits. Under benign conditions, the aphids tended to suffer from reduced fecundity when harbouring X‐type, a mechanism that might maintain intermediate frequencies in field populations. These findings highlight that a single strain of a facultative endosymbiont has the potential to provide diverse benefits to its aphid host.     

Aphid populations are frequently infected with facultative endosymbionts

The occurrence of facultative endosymbionts has been studied in many commercially important crop pest aphids, but their occurrence and effects in non-commercial aphid species in natural populations have received less attention. We screened 437 aphid samples belonging to 106 aphid species for the eight most common facultative aphid endosymbionts. We found one or more facultative endosymbionts in 53% (56 of 106) of the species investigated. This likely underestimates the situation in the field because facultative endosymbionts are often present in only some colonies of an aphid species. Oligophagous aphid species carried facultative endosymbionts significantly more often than monophagous species. We did not find a significant correlation between ant tending and facultative endosymbiont presence. In conclusion, we found that facultative endosymbionts are common among aphid populations. This study is, to our knowledge, the first of its kind in the Netherlands and provides a basis for future research in this field. For instance, it is still unknown in what way many of these endosymbionts affect their hosts, which is important for determining the importance of facultative endosymbionts to community dynamics.     

Co‐occurrence of defensive traits in the potato aphid Macrosiphum euphorbiae

1. Insecticide usage selects strongly for resistance in aphid populations, but this could entail fitness costs in other resistance traits. The potato aphid Macrosiphum euphorbiae Thomas exhibits intraspecific variation in susceptibility to parasitism by braconid wasps and provides a suitable species to study the relation between the defensive traits of parasitism and insecticide resistance. 2. Clonal lines (23 in total) of M. euphorbiae were established from aphids collected in 2013 from geographically separate populations in the U.K. Clonal lines belonged to five aphid genotypes, but one genotype predominated (78% of samples), and the facultative endosymbiont Hamiltonella defensa was detected in c. 40% of lines. 3. Total esterase activity in aphid tissues varied significantly between aphid genotypes and collection areas, but there was no clear pattern in relation to H. defensa infection or between collection sites likely to differ in insecticide pressures. 4. Five clonal lines representing low to moderate levels of enzyme activity, which included different aphid genotypes and presence/absence of H. defensa infection, were assayed for their susceptibility to the parasitoid wasp Aphidius ervi Haliday. Aphid mummification varied significantly between aphid genotypes, with low values in one genotype of aphids irrespective of H. defensa presence. 5. The results revealed that aphid lines belonging to the parasitism‐resistant genotype exhibited moderate levels of total esterase activity, indicating a competitve advantage for this genotype of M. euphorbiae when exposed to chemical and biological control factors in agroecosystems.     

Grain aphid clones vary in frost resistance,but this trait is not influenced by facultative endosymbionts

1. Facultative endosymbiotic bacteria of insects are known to affect life‐history traits of their hosts, and can provide important fitness benefits under certain environmental conditions. While several distinct endosymbiont‐induced effects have been reported, there is no data on whether heritable facultative endosymbionts in any species affect their hosts' performance at low temperatures, something that could have a major effect on insect physiology and survival, and thus population structure and distribution. 2. The original facultative endosymbionts were experimentally removed from five clonal genotypes of the grain aphid, Sitobion avenae Fab., which were then exposed to frost. 3. Aphid genotypes differed considerably in survival following the exposure and in fecundity of the survivors. However, the presence of the facultative symbionts had no overall effect on the studied traits. 4. The results suggest that the facultative symbionts have limited effects on the cold hardiness of their grain aphid hosts.     

Assessment of Sublethal and Transgenerational Effects of Pirimicarb on the Wheat Aphids Rhopalosiphum padi and Sitobion avenae

The wheat aphids, Rhopalosiphum padi (Linnaeus) and Sitobion avenae (Fabricius), are key pests on wheat crops worldwide. Management practices rely primarily on insecticides. The pirimicarb (carbamate) is used extensively as an effective insecticide to control these two aphids. In addition to the mortality caused by pirimicarb, various sublethal effects may occur in aphids when exposed to low lethal or sublethal doses. Understanding the general effect of pirimicarb on aphids could help increasing rational use of this insecticide. Under laboratory conditions, we assessed the sublethal effects of a low lethal concentration of pirimicarb (LC₂₅) on biological traits and acetylcholinesterase (AChE) activity of R. padi and S. avenae. Both direct and transgenerational effects, i.e. on parent and the F1 generations were assessed, respectively. We found that R. padi and S. avenae responded differentially to the LC₂₅ of pirimicarb. The parent generation of R. padi showed a 39% decrease in fecundity and multiple transgenerational effects were observed in the F1 generation; overall juvenile development, reproductive period, adult longevity and lifespan were longer than those of the control group. By contrast, LC₂₅ of pirimicarb showed almost no effects on S. avenae biological traits in both the parent and F1 generations; only the pre-reproductive duration was reduced in F1 generations. Demographic parameter estimates (e.g. r_m) showed similar trend, i.e. significant negative effect on R. padi population growth and no effect on S. avenae. However, AChE activity decreased in both R. padi and S. avenae treated by the LC₂₅ of pirimicarb. We demonstrated sublethal and transgenerational effects of pirimicarb in the two wheat aphid species; it hinted at the importance of considering sublethal effects (including hormesis) of pirimicarb for optimizing Integrated Pest Management (IPM) of wheat aphids.     

Biology and trophic interactions of lucerne aphids

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The value of three cereal aphid species as food for a generalist predator

The value of the cereal aphid species Metopolophium dirhodum (Wlk.), Sitobion avenae (F.) and Rhopalosiphum padi (L.) as prey for the linyphiid spider Erigone atra (Bl.) was assessed. Fecundity of females was determined for spiders fed on eight experimental diets: three single‐species aphid diets, a mixed diet of all three aphid species, three mixed diets with each aphid species in combination with fruit flies Drosophila melanogaster (Meig.), and pure D. melanogaster as a high quality comparison diet. The development and survival of first‐instar juveniles fed on three diets of single aphid species, and on a diet of Collembola were compared with those subjected to starvation. Prey value for adult females was assessed by egg production, hatching success and offspring size. In pure diets all three aphid species were of low value to the spiders, causing a rapid decline in egg production and supporting no growth of significance of first‐instar juveniles. No difference in value of aphid species of single‐species aphid diets was found in the fecundity experiment, while a ranking of aphid species of M. dirhodum > R. padi > S. avenae was revealed in the survivorship experiment. A mixed‐aphid diet was not found to be advantageous compared with single‐species aphid diets, and no advantage of including aphids in mixed diets with fruit flies was found. Metopolophium dirhodum and R. padi were neutral in mixed diets, while a diet of S. avenae and fruit flies caused reduced egg production compared with the pure diet of fruit flies, revealing a toxic effect of S. avenae on the spider. The value‐ranking of aphid species in mixed diets was similar to that of single‐species diets. A similar ranking of aphid species was found for different fitness parameters (fecundity of adult females and development of juveniles). A ranking of aphids by offspring size of mothers on aphid‐only diets was S. avenae > M. dirhodum > R. padi. All aphid‐fruit fly diets resulted in larger offspring than a diet of only D. melanogaster, with the overall largest offspring being produced on the diet of M. dirhodum and fruit flies.     

The impact of insecticide resistance in the currant-lettuce aphid, Nasonovia ribisnigri, on pest management in lettuce

Abstract 1 This paper reports on experiments to determine how two different insecticide resistance phenotypes in the aphid Nasonovia ribisnigri (Mosley), which is a major pest of lettuce, change its susceptibility to pyrethroid insecticides and the carbamate pirimicarb. 2 A novel statistical approach determined how the effectiveness of different insecticides was changed by the two resistance phenotypes. This compared the between‐plant distribution of aphid numbers, as opposed to the mean number of aphids per plant. 3 Results from field cage experiments showed that the effect of the resistances differed. Pyrethroid resistance resulted in lower mortality immediately after application of pyrethroids, whereas resistance to pirimicarb shortened the time over which the chemical was effective. 4 The results of laboratory bioassays suggested that these two resistances were not found together in N. ribisnigri. However, the results reported here contradict this assertion. 5 Experiments with insecticide residues showed that reproduction of resistant N. ribisnigri was greater than that of susceptible N. ribisnigri on plants with ageing insecticide residues, even in circumstances where mortality of resistant and susceptible clones of N. ribisnigri were similar. 6 If more than a few aphids are found on a plant then a whole consignment can be rejected for processing. The results reported here suggest that the effect of both insecticide resistances in N. ribsinigri will be to increase the proportion of lettuce heads with an unacceptable number of aphids on them, leading to increased rejection of plants for processing.     

A diversity of endosymbionts across Australian aphids and their persistence in aphid cultures

There is increasing interest in the use of endosymbionts in pest control, which will benefit from the identification of endosymbionts from potential donor species for transfer to pest species. Here, we screened for endosymbionts in 123 Australian aphid samples across 32 species using 16S DNA metabarcoding. We then developed a qPCR method to validate the metabarcoding data set and to monitor endosymbiont persistence in aphid cultures. Pea aphids (Acyrthosiphon pisum) were frequently coinfected with Rickettsiella and Serratia, and glasshouse potato aphids (Aulacorthum solani) were coinfected with Regiella and Spiroplasma; other secondary endosymbionts detected in samples occurred by themselves. Hamiltonella, Rickettsia and Wolbachia were restricted to a single aphid species, whereas Regiella was found in multiple species. Rickettsiella, Hamiltonella and Serratia were stably maintained in laboratory cultures, although others were lost rapidly. The overall incidence of secondary endosymbionts in Australian samples tended to be lower than recorded from aphids overseas. These results indicate that aphid endosymbionts probably exhibit different levels of infectivity and vertical transmission efficiency across hosts, which may contribute to natural infection patterns. The rapid loss of some endosymbionts in cultures raises questions about factors that maintain them under field conditions, while endosymbionts that persisted in laboratory culture provide candidates for interspecific transfers.     

Searching for wheat resistance to aphids and wheat bulb fly in the historical Watkins and Gediflux wheat collections

Insect pests can reduce wheat yield by direct feeding and transmission of plant viruses. Here we report results from laboratory and field phenotyping studies on a wide range of wheat, including landraces from the Watkins collection deriving from before the green revolution, more modern cultivars from the Gediflux collection (north‐western Europe) and modern UK Elite varieties, for resistance to the bird cherry‐oat aphid, Rhopalosiphum padi (Homoptera: Aphididae) and the English grain aphid, Sitobion avenae (Homoptera: Aphididae). A total of 338 lines were screened for R. padi and 340 lines for S. avenae. Field trials were also conducted on 122 Watkins lines to identify wheat bulb fly, Delia coarctata, preference on these landraces. Considerable variation was shown in insect performance among and within different wheat collections, with reduced susceptibility in a number of varieties, but phenotyping did not identify strong resistance to aphids or wheat bulb fly. Field trials showed within collection differences in aphid performance, with fewer aphids populating lines from the Watkins collection. This differs from development data in laboratory bioassays and suggests that there is a pre‐alighting cue deterring aphid settlement and demonstrates differences in aphid preference and performance on older plants in the field compared with seedlings in the laboratory, highlighting the need for phenotyping for aphid resistance at different plant growth stages. No association was identified between performance of the different insect species on individual varieties, potentially suggesting different nutritional requirements or resistance mechanisms.     

The influence of facultative endosymbionts on honeydew carbohydrate and amino acid composition of the black bean aphid Aphis fabae

The facultative endosymbionts Hamiltonella defensa and Regiella insecticola are commonly found in aphids. They are linked with various ecological benefits but generally occur at low prevalence, which indicates a possible harbouring cost. Little is known about how the presence of facultative endosymbionts is reflected in honeydew composition. Honeydew is the key mediator of the mutualism between aphids and their tending ants. The present study examines whether endosymbionts have an influence on aphid honeydew quality by comparing the amino acid and carbohydrate concentrations between infected and uninfected aphids. To this end, two genetic lines of the aphid Aphis fabae Scopoli are experimentally infected with different strains of Hamiltonella and Regiella. Infected aphids are shown to have reduced concentrations of amino acids in the honeydew compared with uninfected aphids. However, the presence of endosymbionts has no effect on the absolute amount of carbohydrates produced. Nevertheless, interclonal variation in honeydew composition between aphid genotypes is observed for both carbohydrate and amino acid production. These results imply that the nutritional value of honeydew depends on aphid genotype, as well as on the presence of secondary bacterial endosymbionts, which suggests that there is a physiological cost of harbouring endosymbionts and which could also impact aphid attractiveness to tending ants.     

Unrelated facultative endosymbionts protect aphids against a fungal pathogen

The importance of microbial facultative endosymbionts to insects is increasingly being recognized, but our understanding of how the fitness effects of infection are distributed across symbiont taxa is limited. In the pea aphid, some of the seven known species of facultative symbionts influence their host's resistance to natural enemies, including parasitoid wasps and a pathogenic fungus. Here we show that protection against this entomopathogen, Pandora neoaphidis, can be conferred by strains of four distantly related symbionts (in the genera Regiella, Rickettsia, Rickettsiella and Spiroplasma). They reduce mortality and also decrease fungal sporulation on dead aphids which may help protect nearby genetically identical insects. Pea aphids thus obtain protection from natural enemies through association with a wider range of microbial associates than has previously been thought. Providing resistance against natural enemies appears to be a particularly common way for facultative endosymbionts to increase in frequency within host populations.     

Prevalence,damage, management and insecticide resistance of stink bug populations (Hemiptera: Pentatomidae) in commodity crops

1. Pest management of stink bugs (Hemiptera: Pentatomidae) in soybean [Glycine max (L.) Merr.], corn (Zea mays L.) and cotton (Gossypium spp.) agroecosystems has become a major concern in several countries of the Americas.
2. In this review, we report an overview on geographical distribution, injury, damage and methods used to control (plant resistance mechanisms, biological control) the most important stink bugs in the Americas, with an emphasis on Brazil, the implications of the trend towards decreased susceptibility of stink bug populations to insecticides and the current difficulties of the management of these insect pests.
3. Currently, the Neotropical brown stink bug Euschistus heros (Fabricius) is less susceptible to organophosphate insecticides than in the past. A slight reduction in E. heros susceptibility to pyrethroids and, to a lesser extent, to neonicotinoids has also been observed. In addition, the green‐belly stink bug [Dichelops melacanthus (Dallas)] is more tolerant to the three classes of insecticides (neonicotinoids, organophosphates and pyrethroids) than E. heros.
4. Metabolic detoxification is involved in organophosphate, neonicotinoid and pyrethroid differences in susceptibility. Restricted availability of insecticides with different modes of action could favour the selection of resistant phenotypes in stink bug populations.

     

Hidden in plain sight: phylogeography of an overlooked parasitoid species Trioxys sunnysidensis Fulbright & Pike (Hymenoptera: Braconidae: Aphidiinae)

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Resistance to multiple cereal aphids in wheat–alien substitution and translocation lines

Rhopalosiphum padi, Schizaphis graminum, and Sitobion avenae are three of the most destructive aphid species of wheat (Triticum aestivum L.). They can significantly reduce wheat yields directly by feeding and indirectly by transmitting viruses. This study aimed to search for resistance to these aphid species among lines derived from different rye (Secale cereale) origins and from Aegilops speltoides, all in the genetic background of the wheat cultivar Pavon F76. Resistance was quantified as aphid weight (R. padi, S. avenae, and S. graminum) and the number of aphids and percentage of infested leaf area exhibiting chlorosis (S. graminum). The most resistant genotypes reduced R. padi and S. avenae weight by 24.2 and 34.3 %, respectively, at the seedling stage, compared with Pavon F76 control plants. Strong S. graminum resistance was found only in A. speltoides-derived lines, the most resistant of which (7A.7S-L5) sustained just 3 % chlorosis and reduced S. graminum colony weight by 67.7 %. One line carrying the 1AL.1RS_am wheat–rye translocation from Amigo wheat (originally from Insave rye) reduced S. avenae weight by 23.2 and 21.8 % in seedling and adult plants, respectively. Single genotypes carrying the complete 1R chromosome or the 1RS chromosome arm derived from E12165 wheat and Presto triticale proved to be resistant to both R. padi and S. avenae at the seedling stage. Further research should be conducted to unravel the genetic basis of resistance to these aphids in 1RS genotypes. The sources of resistance identified here may be useful for incorporating multiple aphid species resistance in wheat breeding programs, particularly for R. padi and S. avenae, to which no resistant wheats have been bred.     

Population growth of Rhopalosiphum padi under different thermal regimes: an agent-based model approach

1. Rhopalosiphum padi (Linnaeus, 1758) is abundant and has a broad geographic distribution. It is one of the most important cereal pests. In Brazil, the economic losses associated with this aphid result mainly from the transmission of the barley yellow dwarf viruses.
2. Decision-making for the adoption of management measures must consider the initial population size, the potential for population increase, and the time when this population will reach levels at which the resulting damage is equal to the costs of control measures. Consequently, the establishment of management programmes and decision support systems should be based on models that estimate the potential population growth of this pest species.
3. Temperature is one of the main factors that determine the growth rate of insect populations. Generally, controlled experiments are designed to examine the relationship of temperature at fixed intervals in relation to the development phases of insects. In nature, thermal regimes are not constant, and population growth is the result of a series of combined events.
4. In this work, the effects of different thermal regimes on the population growth of R. padi were compared.
5. An agent-based model was used to estimate population growth, and the parameters defined in controlled regimes were compared with fluctuating temperatures under natural conditions.
6. The temperature-driven model presented here can serve as a tool to predict population growth and decision-making for aphid management.
7. The model structure and the proposed experimental design allow the addition of modules and layers of factors that can progressively affect the populations of aphids to gradually improve the model.

     

Spread of barley yellow dwarf virus and relative importance of local aphid vectors in central Washington

Data from bioassays of field collected aphids, barley indicator plants exposed to natural conditions, and various types of aphid traps were used to describe the spread of barley yellow dwarf virus (BYDV) in wheat and barley near Prosser, Washington. Bioassays were also used to assess the relative importance of local vector species. Of alate aphids collected from grain in the 1982 and 1983 fall migration seasons, 3.4–14–5% transmitted BYDV. Data from concurrent and post-migration assays of resident aphids (apterae and nymphs) reflected an increase in the proportion of infected plants in the field. Maximum increase in the percentage of viruliferous aphids occurred in late November and December of 1982 and November of 1983. The 1982 increase occurred after aphid flights had ceased for the year, suggesting active secondary spread. Collections in pitfall traps and infected trap plants from November to February confirmed aphid activity and virus spread. Rhopalosiphum padi was the most important vector in central Washington in 1982 and 1983 because of its abundance and relative BYDV transmission efficiency. Metopolophium dirhodum was more winter-hardy than R. padi and equal to R. padi in its efficiency as a vector; however, it was not as abundant as R. padi except during the mild winter of 1982–83, when it was a major contributor to secondary spread. Sitobion avenae may be important in years when it is abundant, but it was only a quarter as efficient as R. padi. Rhopalosiphum maidis was a much less efficient vector than R. padi and it only reached high populations in late autumn barley.