Host range expansion of an introduced insect pest through multiple colonizations of specialized clones

Asexuality confers demographic advantages to invasive taxa, but generally limits adaptive potential for colonizing of new habitats. Therefore, pre-existing adaptations and habitat tolerance are essential in the success of asexual invaders. We investigated these key factors of invasiveness by assessing reproductive modes and host-plant adaptations in the pea aphid, Acyrthosiphon pisum, a pest recently introduced into Chile. The pea aphid encompasses lineages differing in their reproductive mode, ranging from obligatory cyclical parthenogenesis to fully asexual reproduction. This species also shows variation in host use, with distinct biotypes specialized on different species of legumes as well as more polyphagous populations. In central Chile, microsatellite genotyping of pea aphids sampled on five crops and wild legumes revealed three main clonal genotypes, which showed striking associations with particular host plants rather than sampling locations. Phenotypic analyses confirmed their strong host specialization and demonstrated parthenogenesis as their sole reproductive mode. The genetic relatedness of these clonal genotypes with corresponding host-specialized populations from the Old World indicated that each clone descended from a particular Eurasian biotype, which involved at least three successful introduction events followed by spread on different crops. This study illustrates that multiple introductions of highly specialized clones, rather than local evolution in resource use and/or selection of generalist genotypes, can explain the demographic success of a strictly asexual invader.     

Genotypic diversity of the cotton-melon aphid Aphis gossypii (Glover) in Tunisia is structured by host plants

The study of intraspecific variation with respect to host plant utilization in polyphagous insects is crucial for understanding evolutionary patterns of insect-plant interactions. Aphis gossypii (Glover) is a cosmopolitan and extremely polyphagous aphid species. If host plant species or families constitute selective regimes to these aphids, genetic differentiation and host associated adaptation may occur. In this study, we describe the genetic structure of A. gossypii collected in six localities in Tunisia on different vegetable crops, on citrus trees and on Hibiscus. The aim was to determine if the aphid populations are structured in relation to the host plants and if such differentiation is consistent among localities. The genetic variability of A. gossypii samples was examined at eight microsatellite loci. We identified only 11 multilocus genotypes among 559 individuals. Significant deviations from Hardy-Weinberg equilibrium, linkage disequilibria and absence of recombinant genotypes, confirmed that A. gossypii reproduces by continuous apomictic parthenogenesis. Genetic differentiation between localities was not significant, whereas a strong differentiation was observed between host plant families (0.175相似文献   

Population differentiation and genetic variation in performance on eight hosts in the pea aphid complex

Phytophagous insects frequently use multiple host-plant species leading to the evolution of specialized host-adapted populations and sometimes eventually to speciation. Some insects are confronted with a large number of host-plant species, which may provide complex routes of gene flow between host-adapted populations. The pea aphid (Acyrthosiphon pisum) attacks a broad range of plants in the Fabaceae and it is known that populations on Trifolium pratense and Medicago sativa can be highly specialized at exploiting these species. To find out whether adaptation to a broad range of co-occurring hosts has occurred, we tested the performance of pea aphid clones collected from eight host-plant genera on all of these plants in a reciprocal transfer experiment. We provide evidence for pervasive host-plant specialization. The high performance of all aphid clones on Vicia faba suggests that this host plant could be a site of gene flow between different populations that could limit further host-associated divergence. The genetic variance in host-plant usage was partitioned into within- and among-population components, which represent different levels of host adaptation. Little evidence of within-population trade-offs in performance on different plant species was found.     

Genetic structure and clonal diversity of an introduced pest in Chile, the cereal aphid Sitobion avenae

In Chile, the aphid Sitobion avenae is of recent introduction, lives on cultivated and wild Poaceae, and is thought to reproduce by permanent parthenogenesis. In order to study the genetic variability and population structure of this species, five microsatellite loci were typed from individual aphids collected from different cultivated and wild host plants, from different geographical zones, and years. Chilean populations showed a high degree of heterozygosity and a low genetic variability across regions and years, with four predominant genotypes representing nearly 90% of the sample. This pattern of low clonal diversity and high heterozygosity was interpreted as the result of recent founder events from a few asexually reproducing genotypes. Most geographical and temporal variation observed in the genetic composition resulted from fluctuations of a few predominant clones. In addition, comparisons of the genotypes found in Chile with those described in earlier surveys of S. avenae populations in Western Europe led us to identify 'superclones' with large geographical distribution and high ecological success, and to make a preliminary exploration of the putative origin(s) of S. avenae individuals introduced to Chile.     

Host Plant Specialization in the Sugarcane Aphid Melanaphis sacchari

Most aphids are highly specialized on one or two related plant species and generalist species often include sympatric populations adapted to different host plants. Our aim was to test the hypothesis of the existence of host specialized lineages of the aphid Melanaphis sacchari in Reunion Island. To this end, we investigated the genetic diversity of the aphid and its association with host plants by analyzing the effect of wild sorghum Sorghum bicolor subsp. verticilliflorum or sugarcane as host plants on the genetic structuring of populations and by performing laboratory host transfer experiments to detect trade-offs in host use. Genotyping of 31 samples with 10 microsatellite loci enabled identification of 13 multilocus genotypes (MLG). Three of these, Ms11, Ms16 and Ms15, were the most frequent ones. The genetic structure of the populations was linked to the host plants. Ms11 and Ms16 were significantly more frequently observed on sugarcane, while Ms15 was almost exclusively collected in colonies on wild sorghum. Laboratory transfer experiments demonstrated the existence of fitness trade-offs. An Ms11 isofemale lineage performed better on sugarcane than on sorghum, whereas an Ms15 lineage developed very poorly on sugarcane, and two Ms16 lineages showed no significant difference in performances between both hosts. Both field and laboratory results support the existence of host plant specialization in M. sacchari in Reunion Island, despite low genetic differentiation. This study illustrates the ability of asexual aphid lineages to rapidly undergo adaptive changes including shifting from one host plant to another.     

基于微卫星标记的桃蚜种群寄主遗传分化

桃蚜Myzus persicae(Sulzer)是寄主范围最广、危害最大的蚜虫种类之一。为了探明桃蚜在不同寄主上的遗传分化特点,采用微卫星分子标记技术,对西兰花、桃树、辣椒上的桃蚜种群进行遗传多样性和遗传结构研究。结果表明,在所选用的5个微卫星位点上共检测到38个等位基因,平均每个位点的等位基因数达到7.6个,桃树种群遗传多样性最高,这可能是因为各种夏寄主上的桃蚜迁回桃树上越冬,从而使多种等位基因和基因型得以聚集的原因。等位基因频率差异分析显示西兰花种群、桃树种群和辣椒种群两两之间(除了桃树06种群和辣椒06种群之间没有遗传分化外)都出现了明显遗传分化,相比之下桃树种群和辣椒种群的分化程度要比桃树种群和西兰花种群的分化程度低,这可能预示着西兰花寄主上的桃蚜正在向远离桃树和辣椒种群的方向进化。     

The genetic structure of Aphis gossypii populations in Japanese fruit orchards

The genetic structure of Aphis gossypii Glover (Hemiptera: Aphididae) populations was characterized using microsatellite analysis of insects sampled from pear and citrus orchards at three locations in the summer seasons of 2001 and 2003. These data will enable more precise forecasting of infestation and will be of value to the development of better control methods. In total, 89 genotypes were found in 500 aphid samples. The diversity indices in aphid populations on pear trees fluctuated but showed a decrease with time in citrus orchards. Japanese aphid populations have greater variation in their microsatellite loci than African, European, or Southeast Asian populations. Approximately half of the genotypes were found in only one colony, whereas the remainder were identified in two or more colonies. Fifteen genotypes appeared in different years, in samples from different areas, or on different host plants. Our data suggest the existence of aphid populations with pear‐adapted genotypes and with a permanent parthenogenetic life cycle. Some of the genotypes included genes conferring resistance to pirimicarb and pyrethroid insecticides. Japanese A. gossypii populations may maintain their large genetic variation through differences in their life cycles.     

Ecological specialization of the aphid Aphis gossypii Glover on cultivated host plants

Many plant-feeding insect species considered to be polyphagous are in fact composed of genetically differentiated sympatric populations that use different hosts and between which gene flow still exists. We studied the population genetic structure of the cotton-melon aphid Aphis gossypii that is considered as one of the most polyphagous aphid species. We used eight microsatellites to analyse the genetic diversity of numerous samples of A. gossypii collected over several years at a large geographical scale on annual crops from different plant families. The number of multilocus genotypes detected was extremely low and the genotypes were found to be associated with host plants. Five host races were unambiguously identified (Cucurbitaceae, cotton, eggplant, potato and chili- or sweet pepper). These host races were dominated by asexual clones. Plant transfer experiments using several specialized clones further confirmed the existence of host-associated trade-offs. Finally, both genetic and experimental data suggested that plants of the genus Hibiscus may be used as refuge for the specialized clones. Resource abundance is discussed as a key factor involved in the process of ecological specialization in A. gossypii.     

Does insecticide resistance alone account for the low genetic variability of asexually reproducing populations of the peach-potato aphid Myzus persicae?

The typical life cycle of aphids includes several parthenogenetic generations and a single sexual generation (cyclical parthenogenesis), but some species or populations are totally asexual (obligate parthenogenesis). Genetic variability is generally low in these asexually reproducing populations, that is, few genotypes are spread over large geographic areas. Both genetic drift and natural selection are often invoked to account for this low genetic variability. The peach-potato aphid, Myzus persicae, which encompasses both cyclical and obligate parthenogens, has developed several insecticide resistance mechanisms as a consequence of intense insecticide use since the 1950s. We collected asexually reproducing M. persicae from oilseed rape and examined genetic variability at eight microsatellite loci and three insecticide resistance genes to determine whether their genetic structure was driven by drift and/or selection. We identified only 16 multilocus microsatellite genotypes among 255 individuals. One clone, which combined two insecticide resistance mechanisms, was frequently detected in all populations whatever their location over a large geographical area (the northern half of France). These unexpected findings suggest that drift is not the unique cause of this low variability. Instead, the intensification of both insecticide treatments and oilseed rape cultivation may have favored a few genotypes. Thus, we propose that selective pressures resulting from human activities have considerably modified the genetic structure of M. persicae populations in northern France in a relatively short period of time.     

Kin structure provides no explanation for intruders in social aphids

Nontraditional social organisms have received increasing attention in recent years, because they present opportunities to study the convergent properties of social evolution. Some aphid species are social, occurring in dense clones with specialized morphs that attack predators and parasites. Little is known about how social aphid colonies resolve conflicts of interest when clonal barriers break down. Pemphigus obesinymphae is a North American gall-forming social aphid that produces both nymphal defenders that protect natal clones, and specialized intruders that invade other nearby clones on their host plants. We tested the hypothesis that clones are arranged on their host plants in spatial clusters of related family groups, such that intruders would be biased towards movement within kin groups. Movement within and not between kin groups would then provide insight into the nature of conflict in this social aphid. We sampled eight sites in the eastern United States and in Arizona, and used eight microsatellite markers to estimate pairwise relatedness between spatial groups. We found little evidence of deviation from random distributions of genotypes on their host plants. Evidently, Pem. obesinymphae intruders typically exploit unrelated clones, and spatial orientation provides no solution to the problem of 'polyclonality' in this species. We discuss implications of this result for our understanding of cooperation and conflict in social aphids.     

Elimination of a specialised facultative symbiont does not affect the reproductive mode of its aphid host

Abstract.  1. Microorganisms that manipulate the reproduction of their hosts through diverse mechanisms including the induction of parthenogenesis are widespread among arthropods.
2. The pea aphid, Acyrthosiphon pisum , shows a variation in its reproductive mode, with lineages reproducing by cyclical parthenogenesis (obligate alternation of parthenogenetic and sexual generations each year) and others by obligate parthenogenesis (continuous asexual reproduction all year round). In addition, the pea aphid harbours, along with Buchnera the primary aphid endosymbiont, several facultative symbionts whose prevalence differs among host populations.
3. The possible influence of a Rickettsia facultative symbiont on the reproductive mode of its host was tested on two pea aphid clones by comparing the response of infected and uninfected individuals with the same genetic background to conditions that typically induce the production of sexual morphs.
4. No significant effect of the Rickettsia infection was found on the type of reproductive morphs produced (sexual vs. asexual) or on their quantities for the two clones.
5. However, the Rickettsia had a detrimental effect on the fitness of its aphid host, in apparent contradiction to the high prevalence of this symbiont in some host populations. It is suggested that this negative impact may disappear under specific environmental conditions, transforming a parasitic association into a mutualistic one.     

Genetic interactions influence host preference and performance in a plant-insect system

Phytophagous insects generally feed on a restricted range of host plants, using a number of different sensory and behavioural mechanisms to locate and recognize their host plants. Phloem-feeding aphids have been shown to exhibit genetic variation for host preference of different plant species and genetic variation within a plant species can also have an effect on aphid preference and acceptance. It is known that genotypic interactions between barley genotypes and Sitobion avenae aphid genotypes influence aphid fitness, but it is unknown if these different aphid genotypes exhibit active host choice (preference) for the different barley genotypes. Active host choice by aphid genotypes for particular plant genotypes would lead to assortative association (non-random association) between the different aphid and plant genotypes. The performance of each aphid genotype on the plant genotypes also has the ability to enhance these interactions, especially if the aphid genotypes choose the plant genotype that also infers the greatest fitness. In this study, we demonstrate that different aphid genotypes exhibit differential preference and performance for different barley genotypes. Three out of four aphid genotypes exhibited preference for (or against) particular barley genotypes that were not concordant with differences in their reproductive rate on the specific barley genotype. This suggests active host choice of aphids is the primary mechanism for the observed pattern of non-random associations between aphid and barley genotypes. In a community context, such genetic associations between the aphids and barley can lead to population-level changes within the aphid species. These interactions may also have evolutionary effects on the surrounding interacting community, especially in ecosystems of limited species and genetic diversity.     

Weak spatial and temporal population genetic structure in the rosy apple aphid, Dysaphis plantaginea, in French apple orchards

We used eight microsatellite loci and a set of 20 aphid samples to investigate the spatial and temporal genetic structure of rosy apple aphid populations from 13 apple orchards situated in four different regions in France. Genetic variability was very similar between orchard populations and between winged populations collected before sexual reproduction in the fall and populations collected from colonies in the spring. A very small proportion of individuals (～2%) had identical multilocus genotypes. Genetic differentiation between orchards was low (F(ST)<0.026), with significant differentiation observed only between orchards from different regions, but no isolation by distance was detected. These results are consistent with high levels of genetic mixing in holocyclic Dysaphis plantaginae populations (host alternation through migration and sexual reproduction). These findings concerning the adaptation of the rosy apple aphid have potential consequences for pest management.     

Insecticide use and competition shape the genetic diversity of the aphid Aphis gossypii in a cotton-growing landscape

Field populations of the cotton aphid, Aphis gossypii Glover, are structured into geographically widespread host races. In the cotton-producing regions of West and Central Africa (WCA), two genotypes have been repeatedly detected within the cotton host race, one of which (Burk1) is prevalent (>90%) and resistant to several insecticides, as opposed to the second one (Ivo). Here, we conducted whole plant and field cage experiments to test hypotheses for such low genetic diversity, including selection from insecticide treatments, interclonal competition and adaptation to host plant, or climatic conditions. To assess the genetic diversity of immigrant aphids, alatae were trapped and collected on cotton and relay host plants (okra and roselle) in the early cropping season. Individuals were genotyped at eight specific microsatellite loci and characterized by a multilocus genotype (MLG). When independently transferred from cotton (Gossypium hirustum L.) leaf discs to whole plants (G. hirsutum and G. arboreum, roselle and okra), Ivo and Burk1 performed equally well. When concurrently transferred from cotton leaf discs to the same plant species, Ivo performed better than Burk1, indicating that competition favoured Ivo. This was also the case on G. hirsutum growing outdoors. Conversely, Burk1 prevailed when cotton plants were sprayed with insecticides. In experiments where aphids were allowed to move to neighbouring plants, Burk1 was better represented than Ivo on low-populated plants, suggesting that dispersal may be a way to avoid competition on crowded plants. Most cotton aphids collected on cotton or relay host plants in the early cropping season were Burk1 (>90%), indicating high dispersal ability and, probably reflecting high frequency on host plants from which they dispersed. In the agricultural landscape of WCA, the use of broad-range insecticides on both cotton and relay host plants has led to the prevalence of one genotype of A. gossypii resistant to different classes of insecticides. Deployment of widespread and integrated pest management strategies are needed to restore cotton aphid control.     

Population genetic structure and secondary symbionts in host-associated populations of the pea aphid complex

Polyphagous insect herbivores experience different selection pressures on their various host plant species. How this affects population divergence and speciation may be influenced by the bacterial endosymbionts that many harbor. Here, we study the population structure and symbiont community of the pea aphid (Acyrthosiphon pisum), which feeds on a range of legume species and is known to form genetically differentiated host-adapted populations. Aphids were collected from eight legume genera in England and Germany. Extensive host plant associated differentiation was observed with this collection of pea aphids comprising nine genetic clusters, each of which could be associated with a specific food plant. Compared to host plant, geography contributed little to genetic differentiation. The genetic clusters were differentiated to varying degrees, but this did not correlate with their degree of divergence in host use. We surveyed the pea aphid clones for the presence of six facultative (secondary) bacterial endosymbionts and found they were nonrandomly distributed across the aphid genetic clusters and this distribution was similar in the two countries. Aphid clones on average carried 1.4 species of secondary symbiont with those associated with Lathyrus having significantly fewer. The results are interpreted in the light of the evolution of specialization and ecological speciation.     

Microevolution, low clonal diversity and genetic affinities of parthenogenetic sitobion aphids in new zealand

In sharp contrast to their southeast Asian and European counterparts, Sitobion miscanthi and S. near fragariae aphids in Australia exhibit a complete absence of sexual reproduction. This demands an explanation within the context of the evolution and maintenance of sex and parthenogenesis. Accordingly, we executed a genetic analysis of the two species in neighbouring New Zealand. Microsatellites and single-stranded conformation polymorphism/sequence analysis of the nuclear gene elongation factor 1alpha were used to identify aphid clones and confirm species identification, respectively. Karyotypic variation was also investigated. The New Zealand fauna showed few (nonrecombining) genotypes and appears to have received migrants from both Australia and Asia. Other genotypes have apparently arisen in situ in New Zealand, exhibiting stepwise mutation of microsatellite alleles and also karyotypic change. Thus, these data represent rare evidence of evolution within wild-living parthenogenetic lineages. Karyotypic changes appear to occur at a rate even greater than that of microsatellite evolution. Strong geographical partitioning of genotypes/karyotypes was found, with certain ones predominating over large areas. These data suggest that clonal selection could be important in the distribution and patterning of genetic variation. We present a model to explain the genetic patterns, with particular reference to the absence of sexual reproduction in Sitobion aphids in New Zealand and Australia.     

Variation in clonal diversity in glasshouse infestations of the aphid, Aphis gossypii Glover in southern France

Aphis gossypii is an aphid species that is found throughout the world and is extremely polyphagous. It is considered a major pest of cotton and cucurbit species. In Europe, A. gossypii is assumed to reproduce exclusively by apomictic parthenogenesis. The present study investigates the genetic diversity of A. gossypii in a microgeographic, fragmented habitat consisting of eight glasshouses of cucurbit crops. This analysis, which was based on the results from seven microsatellite loci, has confirmed that A. gossypii populations in southern France are primarily asexual, as only 12 nonrecombinant genotypic classes (clones) were identified from 694 aphids. Moreover, a high proportion of the aphids (87%) had one of three common genotypes. No significant correlation was found between genotypic class and host plant species. Within a glasshouse population of A. gossypii, a significant reduction in clonal diversity was observed as the spring/summer season progressed. The final predominance of a clone could result from interclonal competition. At the microgeographic level (i.e. glasshouses within a 500-m radius), significant genetic subdivision was detected and could be attributed to founder effects and the limitation of gene flow imposed by the enclosed nature of the glasshouse structure. Finally, the three common clones of A. gossypii detected in 1996 reappeared in spring 1997 following the winter extinction, together with rare clones that had not previously been seen. The probability that A. gossypii overwinters within refuges at a microgeographic scale from which populations are renewed each spring is discussed.     

Reproductive mode and population genetic structure of the cereal aphid Sitobion avenae studied using phenotypic and microsatellite markers

As French populations of the aphid Sitobion avenae exhibit a range of reproductive modes, this species provides a good opportunity for studying the evolution of breeding system variation. The present analysis combined ecological and genetic investigations into the spatial distribution of variation in reproductive mode. Reproductive mode was characterized in 277 lineages of S. avenae from France, and these aphids were scored for five microsatellite loci. The analyses revealed strong geographical partitioning of breeding systems, with obligate asexuals mostly restricted to the south of France, while lineages producing sexual forms were more common in the north. Contrary to what might be anticipated for organisms with frequent parthenogenesis, there was substantial genic and genotypic diversity, even in the obligately asexual lineages. More than 120 different genotypes were detected among the 277 aphid lineages, with an average of 5.9 alleles per locus (range four to 16) and heterozygosity of 56.7%. As with previous studies of allozyme variation in aphids, most loci showed heterozygote deficits, and disequilibrium was common among allelic variants at different loci, even after removal of replicate copies of genotypes that might have been derived through clonal reproduction. Our results suggest that selection is important in structuring reproductive systems and genetic variation in French S. avenae. Canonical correspondence analysis was employed to examine the associations between genotypic and phenotypic variables, enabling the identification of alleles correlated with life-history traits.     

Lack of detectable genetic recombination on the X chromosome during the parthenogenetic production of female and male aphids

We used polymorphic microsatellite markers to look for recombination during parthenogenetic oogenesis between the X chromosomes of aphids of the tribe Macrosiphini. We examined the X chromosome because it comprises approximately 25 % of the genome and previous cytological observations of chromosome pairing and nucleolar organizer (NOR) heteromorphism suggest recombination, although the same is not true for autosomes. A total of 564 parthenogenetic females of Myzus clones with three distinct reproductive modes (cyclical parthenogenesis, obligate parthenogenesis and obligate parthenogenesis with male production) were genotyped at three informative X-linked loci. Also, parthenogenetically produced males from clones encompassing the full range of male-producing reproductive strategies were genotyped. These included 391 Myzus persicae males that were genotyped at three X-linked loci and 538 males from Sitobion clones that were genotyped at five informative X-linked loci. Our results show no departure from clonality in parthenogenetic generations of aphids of the tribe Macrosiphini: no recombinant genotypes were observed in parthenogenetically produced males or females.     

Association between Aphis gossypii genotype and phenotype on melon accessions

Development of molecular markers has allowed the characterization of several host–aphid interactions. We investigated the usefulness of microsatellite markers to characterize the plant resistance interaction in the model Aphis gossypii/Cucumis melo. Six aphid clones, collected in different localities and years and belonging to two multilocus genotypes (MLGs) based on eight microsatellite markers, were phenotyped on a set of 33 melon accessions, some of them known to carry the Vat gene. Three parameters were used: acceptance of plant, ability to colonize the plant and resistance to virus when inoculated by aphids. Concordance and correlation analyses showed that aphid clones sharing a same MLG exhibited a very agreeable phenotype on the set of accessions for acceptance of plant and resistance to virus when inoculated by aphids. From host point of view, melon accessions were grouped into four clear categories, resistant to aphids of both MLGs, only resistant to the NM1 MLG, only resistant to the C9 MLG, susceptible to both MLGs and another group of unclear characteristics. The four categories revealed different patterns of virulence for NM1 and C9 MLGs that are likely controlled by a single avirulence gene in accordance with a gene for gene interaction. In contrast, the ability to colonize the plant appeared slightly variable among clones sharing a same MLG. We hypothesize it is due to the putative polygenic control of this aphid trait. Because the phenotypic variability of A. gossypii matched the genetic variability revealed by eight microsatellite markers, these markers could be used to infer the frequency of biotypes in field experiments and help to elucidate the allele diversity of melon resistance genes.