Jean-Charles Bouvier

Biodiversity and pest management in orchard systems. A review

Conventional agriculture is based on a high level of chemical inputs such as pesticides and fertilisers, leading to serious environmental impacts, health risks and loss of biodiversity in agrosystems. The reduction of pesticide use is a priority for intensively sprayed agricultural systems such as orchards. The preservation and promotion of biodiversity within orchards and their boundaries is therefore an issue to explore. Indeed, orchard systems contain high plant diversity and perennial multi-strata designs that provide wealthy resources and habitats to living communities such as beneficial organisms. Orchards thus offer favourable areas to maintain food-webs within the agrosystem, provided that favourable situations are not altered by cultural practices such as applying an excess of pesticides. Here, we analysed literature on the effects of the manipulation of plant diversity and habitats on the control of pests by arthropod and bird communities in apple, pear and peach orchards. Many investigations focus on the role of plant management to enhance biodiversity in orchards but only 22 research reports presenting 30 case studies were dedicated to the study of the ecosystem service provided by plant diversity for orchard pest control. The underlying mechanisms were seldom demonstrated, and the tested grass covers and tree assemblages aimed at favouring either the beneficial complex or only some beneficial species to control one or a few pests. The effect of plant management on pest control was mostly positive (16 cases) or null (9), but also negative in some cases (5). This finding reveals the difficulties of identifying selected plants or plant assemblages for the control of key pests. We conclude that further research is needed to identify the processes involved on different scales for biological control. Orchard systems should be re-designed to optimise ecosystem services provided by biodiversity.

Deltamethrin resistance in the codling moth (Lepidoptera: Tortricidae): inheritance and number of genes involved.

The inheritance of deltamethrin resistance in Cydia pomonella (L.) has been investigated by crossing a resistant (Rv) and a susceptible (Sv) strain, derived from a population collected in south-eastern France in 1995. Deltamethrin resistance was suspected to be under the control of a kdr-type allele and an enhanced mixed-function oxidase (mfo). F1 and F2 progenies were therefore tested through dose–response and enzyme assays. Dose–response relationships indicated that resistance was inherited as an autosomal incompletely recessive (D=−0.199) character, involving at least two genes. Enzyme measures suggested the contribution of 1.2 genes to the expression of mfo, with incomplete dominance (D=0.460). Our results support the hypothesis of a polygenic response to deltamethrin selection in the Rv strain, including a major kdr-type allele with a minor effect of mfos. In the light of these findings, we consider the resistance in codling moth populations in south-eastern France as a product of an adaptive sequential selection process, occurring through the sequential addition of resistance genes.

Constraints on adaptive mutations in the codling moth Cydia pomonella (L.): measuring fitness trade-offs and natural selection

Adaptive changes in populations encountering a new environment are often constrained by deleterious pleiotropic interactions with ancestral physiological functions. Evolutionary responses of populations can thus be limited by natural selection under fluctuating environmental conditions, if the adaptive mutations are associated with pleiotropic fitness costs. In this context, we have followed the evolution of the frequencies of insecticide-resistant mutants of Cydia pomonella when reintroduced into an untreated environment. The novel set of selective forces after removal of insecticide pressure led to the decline of the frequencies of resistant phenotypes over time, suggesting that the insecticide-adapted genetic variants were selected against the absence of insecticide (with a selective coefficient estimated at 0.11). The selective coefficients were also estimated for both the major cytochrome P450-dependent monooxygenase (MFO) and the minor glutathione S-transferase (GST) systems (0.17 and negligible, respectively), which have been previously shown to be involved in resistance. The involvement of metabolic systems acting both through xenobiotic detoxification and biosynthetic pathways of endogenous compounds may be central to explaining the deleterious physiological consequences resulting from pleiotropy of adaptive changes. The estimation of the magnitude of the fitness cost associated with insecticide resistance in C. pomonella suggests that resistance management strategies exclusively based on insecticide alternations would be unlikely to delay such a selection process.

Biodiversity and Pest Management in Orchard Systems

Conventional agriculture is based on a high level of chemical inputs such as pesticides and fertilisers, leading to serious environmental impacts, health risks and loss of biodiversity in agrosystems. The reduction of pesticide use is a priority for intensively sprayed agricultural systems such as orchards. The preservation and promotion of biodiversity within orchards and their boundaries is therefore an issue to explore. Indeed, orchard systems contain high plant diversity and perennial multi-strata designs that provide wealthy resources and habitats to living communities such as beneficial organisms. Orchards thus offer favourable areas to maintain food-webs within the agrosystem, provided that favourable situations are not altered by cultural practices such as applying an excess of pesticides. Here, we analysed literature on the effects of the manipulation of plant diversity and habitats on the control of pests by arthropod and bird communities in apple, pear and peach orchards. Many investigations focus on the role of plant management to enhance biodiversity in orchards but only 22 research reports presenting 30 case studies were dedicated to the study of the ecosystem service provided by plant diversity for orchard pest control. The underlying mechanisms were seldom demonstrated, and the tested grass covers and tree assemblages aimed at favouring either the beneficial complex or only some beneficial species to control one or a few pests. The effect of plant management on pest control was mostly positive (16 cases) or null (9), but also negative in some cases (5). This finding reveals the difficulties of identifying selected plants or plant assemblages for the control of key pests. We conclude that further research is needed to identify the processes involved on different scales for biological control. Orchard systems should be re-designed to optimise ecosystem services provided by biodiversity.

Phenological segregation of insecticide resistance alleles in the codling moth Cydia pomonella (Lepidoptera : Tortricidae) : a case study of ecological divergences associated with adaptive changes in populations

Intrapopulation variability in the seasonal regulation of insect lifecycles has been shown to be due partly to genetic changes. Selection for insecticide resistance in the codling moth Cydia pomonella results from allelic substitution at two to three loci in south-eastern French populations of this species. However, such an adaptive process has been associated with an increased heterogeneity in the developmental responses to climatic factors such as temperature. In this paper, we investigate whether such pleiotropic effects of resistance on development induce a significant discrepancy in seasonal regulation in this species. The seasonal changes in a susceptible and two insecticide-resistant homozygous genotypes of C. pomonella, as well as their reciprocal F1 progeny, were followed under natural conditions during the reproductive season through the emergence events of adults, within-generation developmental rates and the number of generations. A significant delay in the occurrences of homozygous resistant genotypes resulted from significantly lower pre-imaginal developmental times relative to homozygous susceptible ones. Subsequent assessment of the number of generations indicated significantly higher diapause propensities in carriers of the resistance alleles (37.0-76.2%) than in susceptible homozygotes (6-7%), which mostly pupated towards a third generation of adults. In the light of these findings, pleiotropic effects of adaptive changes might be a crucial source of divergence in seasonal regulation at the population level, involving significant life-history trade-offs. In addition to man-made selective factors during the reproductive season, such an effect on the lifecycle could be a key component in the process of selection for resistance genes in south-eastern France C. pomonella populations.

Enzymatic diagnosis of resistance to deltamethrin in diapausing larvae of the codling moth, Cydia pomonella (L.)

The resistance to deltamethrin was evaluated in diapausing larvae of 14 field populations of the codling moth (Cydia pomonella L.) From the main French orchard areas using biological assays. Resistance to deltamethrin was compared to mixed-function-oxidase (mfo) activity measured at the individual level through ethoxycoumarin-O-deethylase (ECOD) activity using a fluorescence microplate reader. The larvae were collected in corrugated paper band traps in the autumn of 1995. Analysis was previously performed on two laboratory strains, one susceptible and one resistant to deltamethrin, in order to characterize the changes in resistance during diapause development. Resistance to deltamethrin as well as the ECOD activity were stable during the diapause, and ECOD activity was always significantly lower in the susceptible strain than in the resistant one. The ECOD activity was significantly correlated to the frequency of resistant moths in the field populations. This strongly suggested the involvement of the mfo system in the resistance to deltamethrin in these populations. The intra-strain variabilities in ECOD activity of both laboratory and field resistant insects indicated that other resistance mechanisms might also be involved. Further investigations on these mechanisms are needed in order to develop complete diagnostic methods and to define suitable control strategies against each resistant population. Arch. Insect Biochem. Physiol. 39:55–64, 1998.

Effects of apple orchard management strategies on the great tit (Parus major) in southeastern france

The impact of conventional, organic, and integrated pest management (IPM) strategies of apple orchards on the reproduction of the great tit Parus major was investigated during a three-year period in southeastern France. The colonization process, egg-laying dates, clutch sizes, and fledging success were similar among pairs of P. major nesting in orchards conducted under the three studied management strategies. However, the mean number of young produced per ha (orchard productivity) was significantly higher in organic orchards than in both conventional and IPM orchards. Such divergences between both fledging success and orchard productivity primarily resulted from higher densities of P. major nesting pairs, but also from lower rates of nest abandonment during incubation in organic orchards. We suggest that intensive pesticide use under both IPM and conventional managements may have resulted in a substantial reduction in insect prey availability that enhanced intraspecific competition, which then led to failure in reproduction in pairs with low competitive ability. Our results highlight the relevance of P. major in assessing the environmental impact of apple orchard management strategies.

Predation by generalist predators on the codling moth versus a closely-related emerging pest the oriental fruit moth: a molecular analysis

1 Biological control by conservation of native natural enemies can, at its best, reduce the need for pesticides and prevent detrimental effects upon the environment. The present study investigated the role of ground‐active generalist predators as natural enemies of two tortricid pests in apple orchards. 2 Predation rates were compared on the well established codling moth Cydia pomonella and the emerging oriental fruit moth Grapholita molesta, which has recently switched hosts to apples. 3 The present study hypothesized that the ground‐active predators consumed the two tortricid pests in significant numbers without preference, and attacked the pests at different developmental stages. 4 Using diagnostic polymerase chain reaction on the gut contents of field‐caught ground‐active predators, no difference in predation rates was found on these two pests. Spiders were the most efficient predators of emergent adult moths in spring, whereas the carabid beetles, feeding on diapausing larvae, were important in the autumn. 5 The temporal complementarity between spiders and carabid beetles, attacking different stages of the pests at different times of year, highlights the need for diverse predator assemblages to optimize biological control.

Interspecific variation in the relationship between clutch size, laying date and intensity of urbanization in four species of hole-nesting birds

Abstract The increase in size of human populations in urban and agricultural areas has resulted in considerable habitat conversion globally. Such anthropogenic areas have specific environmental characteristics, which influence the physiology, life history, and population dynamics of plants and animals. For example, the date of bud burst is advanced in urban compared to nearby natural areas. In some birds, breeding success is determined by synchrony between timing of breeding and peak food abundance. Pertinently, caterpillars are an important food source for the nestlings of many bird species, and their abundance is influenced by environmental factors such as temperature and date of bud burst. Higher temperatures and advanced date of bud burst in urban areas could advance peak caterpillar abundance and thus affect breeding phenology of birds. In order to test whether laying date advance and clutch sizes decrease with the intensity of urbanization, we analyzed the timing of breeding and clutch size in relation to intensity of urbanization as a measure of human impact in 199 nest box plots across Europe, North Africa, and the Middle East (i.e., the Western Palearctic) for four species of hole‐nesters: blue tits (Cyanistes caeruleus), great tits (Parus major), collared flycatchers (Ficedula albicollis), and pied flycatchers (Ficedula hypoleuca). Meanwhile, we estimated the intensity of urbanization as the density of buildings surrounding study plots measured on orthophotographs. For the four study species, the intensity of urbanization was not correlated with laying date. Clutch size in blue and great tits does not seem affected by the intensity of urbanization, while in collared and pied flycatchers it decreased with increasing intensity of urbanization. This is the first large‐scale study showing a species‐specific major correlation between intensity of urbanization and the ecology of breeding. The underlying mechanisms for the relationships between life history and urbanization remain to be determined. We propose that effects of food abundance or quality, temperature, noise, pollution, or disturbance by humans may on their own or in combination affect laying date and/or clutch size.

Insecticide resistance may enhance the response to a host-plant volatile kairomone for the codling moth, Cydia pomonella (L.)

The behavioral and electroantennographic responses of Cydia pomonella (L.) to the ripe pear volatile ethyl (2E,4Z)-2,4-decadienoate (Et-E,Z-DD), were compared in insecticide-susceptible and -resistant populations originating from southern France. A dose–response relationship to this kairomonal attractant was established for antennal activity and did not reveal differences between susceptible and resistant strains. Conversely, males of the laboratory strains expressing metabolic [cytochrome P450-dependent mixed-function oxidases (mfo)] or physiological (kdr-type mutation of the sodium-channel gene) resistance mechanisms exhibited a significantly higher response to Et-E,Z-DD than those of the susceptible strain in a wind tunnel experiment. No response of the females to this kairomone could be obtained in our wind-tunnel conditions. In apple orchards, mfo-resistant male moths were captured at significantly higher rates in kairomone-baited traps than in traps baited with the sex pheromone of C. pomonella. Such a differential phenomenon was not verified for the kdr-resistant insects, which exhibited a similar response to both the sex pheromone and the kairomonal attractant in apple orchards. Considering the widespread distribution of metabolic resistance in European populations of C. pomonella and the enhanced behavioral response to Et-E,Z-DD in resistant moths, the development of control measures based on this kairomonal compound would be of great interest for the management of insecticide resistance in this species.