Laurent Lapchin

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.

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.

Energy dynamics in a parasitoid foraging in the wild

Although parasitoids are used widely as a biological models for understanding the evolution of animal behaviour, most studies have been constrained to the laboratory. The dearth of field studies has been compounded by the almost complete ignorance of the physiological parameters involved in foraging and dispersal, in particular of the energetic constraints imposed by resource limitation. We estimated the dynamics of carbohydrates and lipids reserves of Venturia canescens (Gravenhorst) females by releasing individuals of known nutritional status in a natural environment and recapturing them using host-containing traps. The recapture rate was around 30%. These results were compared with the reserves of caged animals kept under different experimental conditions (freshly emerged, starved to death, fed ad libitum and partially starved). Wild animals were also sampled in order to estimate the resource levels of the local population. The results show that: (i) wasps are able to maintain a nearly constant level of energy over an extended foraging period; (ii) V. canescens takes sugars in the field; and (iii) the lipid reserves accumulated during the larval life may be limiting as lipogenesis does not take place in adults even under conditions of high sugar availability. These results demonstrate that wasps can forage for hosts and food and disperse in this habitat for hours and days without running into a severe risk of energy limitation.

Geographical distribution and genetic relatedness of sympatrical thelytokous and arrhenotokous populations of the parasitoid Venturia canescens (Hymenoptera)

Theory predicts that asexual reproduction has a competitive advantage over sexual reproduction because of the twofold cost of producing males. Few systems are suitable for directly testing this prediction. In the solitary parasitoid wasp Venturia canescens both arrhenotokously (sexual) and thelytokously (asexual) reproducing individuals occur sympatrically. We sampled 922 wasps from 22 localities along the coast of south‐eastern France. Thelytokous wasps were less abundant (23%) than arrhenotokous wasps and were almost always found in sympatry with arrhenotokous ones. An analysis of genetic relatedness using amplified fragment length polymorphism (AFLP) markers showed the existence of a widespread thelytokous clone. In addition, a few thelytokous individuals were found to be closely related to arrhenotokous ones and vice versa. These data suggest the occurrence of occasional gene flow between both reproductive modes and/or recurrent origin of thelytokous clones from coexisting arrhenotokous populations in the area. The results are discussed in the context of the paradox of sex.

Incidence of insecticide resistance alleles in sexually-reproducing populations of the peach-potato aphid Myzus persicae (Hemiptera: Aphididae) from southern France

Intensive chemical treatments have led to the development of a number of insecticide resistance mechanisms in the peach-potato aphid Myzus persicae (Sulzer). Some of these mechanisms are known to be associated with negative pleiotropic effects (resistance costs). Molecular and biochemical methods were used to determine the genotypes or phenotypes associated with four insecticide resistance mechanisms in single aphids from sexually-reproducing populations in southern France. The mechanisms considered were E4 and FE4 carboxylesterase overproduction, modified acetycholinesterase, and kdr and rdl resistance-associated mutations. A new method for determining individual kdr genotypes is presented. Almost all resistant individuals overproduced FE4 carboxylesterase, whereas modified acetylcholinesterase was rare. Both the kdr and rdl resistance mutations were present at high frequencies in French sexually-reproducing populations. The frequencies of insecticide resistance genes were compared before and after sexual reproduction in one peach orchard at Avignon to evaluate the potential impact of selection on the persistence of resistance alleles in the over-wintering phase. The frequencies of the kdr and rdl mutations varied significantly between autumn and spring sampling periods. The frequency of the kdr mutation increased, probably due to pyrethroid treatments at the end of the winter. Conversely, the frequency of the rdl mutation decreased significantly during winter, probably because of a fitness cost associated with this mutation.

Dispersal between host populations in field conditions: navigation rules in the parasitoid Venturia canescens.

Abstract. 1. Dispersal is a life‐history trait that can have great ecological and evolutionary consequences, however understanding of how insects disperse is limited.

Response of the melon aphid, Aphis gossypii, to host-plant resistance: evidence for high adaptive potential despite low genetic variability

In agrosystems, pests are submitted to strong human‐imposed selective pressures to which they sometimes adapt rapidly, either through selection of genotypes resulting from mutation and/or recombination events, or through phenotypic plasticity. Understanding how insects respond to such selective pressures is of great importance for sustainable pest management strategies, such as the use of resistant plants. In this study, we investigated the genetic and phenotypic variability of anholocyclic Aphis gossypii Glover (Hemiptera: Aphididae) strains, in response to the resistance gene Vat that is present in melon crops. Forty‐nine aphid colonies were sampled on several melon crops in southern France, genotyped using 15 microsatellite loci, and tested in phenotypic experiments using Vat or non‐Vat melons. The level of genetic polymorphism between these colonies was low, as only seven multilocus genotypes were detected. In contrast, the phenotypic variability for life‐history and behavioral traits between colonies, including those sharing the same genotype, was unexpectedly high, with a continuum of response to the Vat gene from complete susceptibility to strong virulence. The low genetic polymorphism associated with a strong phenotypic variability highlights the high adaptive potential of A. gossypii and the major role of environmental cues in shaping phenotypic responses of this aphid to pest management strategies.

Gene flow between arrhenotokous and thelytokous populations of Venturia canescens (Hymenoptera).

In the solitary parasitoid wasp Venturia canescens both arrhenotokously (sexual) and thelytokously (parthenogenetical) reproducing individuals occur sympatrically. We found in the laboratory that thelytokous wasps are able to mate, receive and use sperm of arrhenotokous males. Using nuclear (amplified fragment length polymorphism, virus-like protein) and mitochondrial (restriction fragment length polymorphism) markers, we show the occurrence of gene flow from the arrhenotokous to the thelytokous mode in the field. Our results reinforce the paradox of sex in this species.

Host‐Parasitoid Association and Diffuse Coevolution: When to Be a Generalist?

In host‐parasitoid communities, hosts are subjected to selective pressures from numerous parasitoid species, and parasitoids may attack several host species. The specificity of host resistance and parasitoid virulence is thus a key factor in host‐parasitoid coevolution. A continuum of strategies exists, from strict specificity to a generalist strategy. The optimal level of specificity may differ in host and parasitoid. I investigated the optimal level of resistance specificity using a model in which the host could be attacked by two parasitoid species, with variable levels of defense specificity. The fitness of a parasitoid attacking two host species with different levels of virulence specificity was also modeled. Finally, a fluctuating environment was simulated by introducing variable probabilities of encounters between antagonistic species over several generations. If the frequency of encounters with the antagonistic species is fixed, then both host and parasitoid gain from a strategy of exclusive specialization toward the most frequent antagonist. If the frequency of encounters fluctuates between generations, generalist host resistance and partially specialist parasitoid virulence are favored. Generalist host resistance may be considered to be a bet‐hedging response to an unpredictable environment. This asymmetry in host‐parasitoid coevolution may account for some of the genetic structures observed in the field for host‐parasitoid associations.

Dispersal strategies of phytophagous insects at a local scale: adaptive potential of aphids in an agricultural environment

BackgroundThe spread of agriculture greatly modified the selective pressures exerted by plants on phytophagous insects, by providing these insects with a high-level resource, structured in time and space. The life history, behavioural and physiological traits of some insect species may have evolved in response to these changes, allowing them to crowd on crops and to become agricultural pests. Dispersal, which is one of these traits, is a key concept in evolutionary biology but has been over-simplified in most theoretical studies. We evaluated the impact of the local-scale dispersal strategy of phytophagous insects on their fitness, using an individual-based model to simulate population dynamics and dispersal between leaves and plants, by walking and flying, of the aphid Aphis gossypii, a major agricultural pest, in a melon field. We compared the optimal values for dispersal parameters in the model with the corresponding observed values in experimental trials.ResultsWe show that the rates of walking and flying disperser production on leaves were the most important traits determining the fitness criteria, whereas dispersal distance and the clustering of flying dispersers on the target plant had no effect. We further show that the effect of dispersal parameters on aphid fitness depended strongly on plant characteristics.ConclusionParameters defining the dispersal strategies of aphids at a local scale are key components of the fitness of these insects and may thus be essential in the adaptation to agricultural environments that are structured in space and time. Moreover, the fact that the effect of dispersal parameters on aphid fitness depends strongly on plant characteristics suggests that traits defining aphid dispersal strategies may be a cornerstone of host-plant specialization.