Emergence rhythms and protandry in relation to daily patterns of locomotor activity inTrichogramma species

Summary In insect parasitoids, the mating structure of the population depends on the emergence pattern of adults and on their behaviour after emergence. In order to study some aspects of these characteristics in four species of egg parasitoid of the genusTrichogramma, laboratory experiments using automatic video analysis are performed. Adult emergence is concentrated in the morning, with an earlier time for males. The variability of the time of emergence is under genetic control. The relationships between the emergence pattern and the locomotor activity are considered by measuring individually: first, the delay between the emergence and the first locomotion which is significantly greater in females than in males and, second, the activity of newly emerged males, which is maximum at the time of the emergence of females. The locomotor activity is influenced by mating; mated females are much more active than virgin ones, mated males are less active than virgin ones. The adaptive values of the rhythmicity of emergence and of protandry are discussed together with their relationship with the patterns of locomotor activity. The results are also considered with respect to mating strategies in insect parasitoids; they actually support the hypothesis of mating occurring principally among individuals native to the same host patch inTrichogramma species.     

The joint effect of temperature and host species induce complex genotype-by-environment interactions in the larval parasitoid of Drosophila, Leptopilina heterotoma (Hymenoptera: Figitidae)

Phenotypic plasticity and genotype-by-environment interactions are usually studied by testing environmental factors separately. However, several environmental factors may vary simultaneously across the geographic range of a species, and thus interact to drive phenotypic and genotypic evolution. Here, we address the question of the interaction between two environmental factors in the case of a koinobiont parasitoid of Drosophila , Leptopilina heterotoma (Hymenoptera: Figitidae). Using two allopatric parasitoid lines, we analysed the joint effect of developmental temperature and host species on two major fitness components (pre-imaginal survival and fecundity) of emerged parasitoids. Our results indicate that both environmental factors strongly interact to shape the parasitoid phenotype. Moreover, complex genotype-by-environment interactions appeared that can even invert the relative fitness of the two parasitoid lines.     

Within-species diversity of Wolbachia-induced cytoplasmic incompatibility in haplodiploid insects

Wolbachia-induced cytoplasmic incompatibility (CI) can have two consequences in haplodiploid insects: fertilized eggs either die (female mortality, FM) or they develop into haploid males (male development, MD). Origin of this diversity remains poorly understood, but current hypotheses invoke variation in damage suffered by paternal chromosomes in incompatible eggs, thus intermediate CI types should be expected. Here, we show the existence of such a particular CI type. In the parasitoid wasp Leptopilina heterotoma, we compared CI effects in crosses involving lines derived from a single inbred line with various Wolbachia infection statuses (natural tri-infection, mono-infection, or no infection). Tri-infected males induce a FM CI type when crossed with either uninfected or mono-infected females. Crossing mono-infected males with uninfected females results in almost complete CI with both reduced offspring production, indicating partial mortality of fertilized eggs, and increased number of sons, showing haploid male development of others. Mono-infected males thus induce an intermediate Cl type when mated with uninfected females. The first evidence of this expected particular CI type demonstrates that no discontinuity separates MD and FM CI types, which appear to be end points of a phenotypic continuum. Second, different CI types can occur within a given species and even within offspring of a single pair. Third, phenotypic expression of the particular CI type induced by a given Wolbachia variant depends on other bacterial variants that co-infect the same tissues. These results support the idea that haplodiploids should be helpful in clarifying evolutionary pathways of insect-Wolbachia associations.     

Phylogenetic evidence for horizontal transmission of Wolbachia in host-parasitoid associations

Endosymbiotic Wolbachia infect a number of arthropod species in which they can affect the reproductive system. While maternally transmitted, unlike mitochondria their molecular phylogeny does not parallel that of their hosts. This strongly suggests horizontal transmission among species, the mechanisms of which remain unknown. Such transfers require intimate between-species relationships, and thus host-parasite associations are outstandingly appropriate for study. Here, we demonstrate that hymenopteran parasitoids of frugivorous Drosophila species are especially susceptible to Wolbachia infection. Of the five common European species, four proved to be infected; furthermore, multiple infections are common, with one species being doubly infected and two triply infected (first report). Phylogenetic statuses of the Wolbachia infecting the different species of the community have been studied using the gene wsp, a highly variable gene recently described. This study reveals exciting similarities between the Wolbachia variants found in parasitoids and their hosts. These arguments strongly support the hypothesis of frequent natural Wolbachia transfers into other species and open a new field for genetic exchanges among species, especially in host-parasitoid associations.     

ER stress protects from retinal degeneration

The unfolded protein response (UPR) is a specific cellular process that allows the cell to cope with the overload of unfolded/misfolded proteins in the endoplasmic reticulum (ER). ER stress is commonly associated with degenerative pathologies, but its role in disease progression is still a matter for debate. Here, we found that mutations in the ER‐resident chaperone, neither inactivation nor afterpotential A (NinaA), lead to mild ER stress, protecting photoreceptor neurons from various death stimuli in adult Drosophila. In addition, Drosophila S2 cultured cells, when pre‐exposed to mild ER stress, are protected from H₂O₂, cycloheximide‐ or ultraviolet‐induced cell death. We show that a specific ER‐mediated signal promotes antioxidant defences and inhibits caspase‐dependent cell death. We propose that an immediate consequence of the UPR not only limits the accumulation of misfolded proteins but also protects tissues from harmful exogenous stresses.     

Adaptive significance of a circadian clock: temporal segregation of activities reduces intrinsic competitive inferiority in Drosophila parasitoids

Most organisms show self-sustained circadian oscillations or biological clocks which control their daily fluctuations in behavioural and physiological activities. While extensive progress has been made in understanding the molecular mechanisms of biological clocks, there have been few clear demonstrations of the fitness value of endogenous rhythms. This study investigated the adaptive significance of circadian rhythms in a Drosophila parasitoid community. The activity rhythms of three sympatric Drosophila parasitoids are out of phase, the competitively inferior parasitoid species being active earlier than the superior competitor. This temporal segregation appears at least partially determined by endogenous periods of the clock which also vary between species and which correlate the time of activity. This earlier activity of the inferior competitor significantly reduces its intrinsic competitive disadvantage when multiparasitism occurs, thus suggesting that natural selection acting on the phase of the rhythm could substantially deviate the endogenous period from the optimal ca. 24 h period. This study demonstrates that temporal segregation of competing species could be endogenously controlled, which undoubtedly favours their coexistence in nature and also shows how natural selection can act on biological clocks to shape daily activity patterns.     

Host exploitation by parasitoids: local variations in foraging behaviour of females among populations of Trichogramma species

Using artificial patches of Ephestia kuehniella (Lepidoptera: Pyralidae) eggs, we compared two strains of Trichogramma cacoeciae Marchal and three strains of T. brassicae Bezdenko (Hymenoptera: Trichogrammatidae) for their strategies in colonizing hosts. The method by which females distribute their eggs among hosts was assessed either in terms of the average distance between parasitized hosts within a given patch, or of the average distance of attacks in discontinuous patches arranged in a circle.Within each strain, the traits increased as females aged; they vary among strains, their range of variations within T. cacoeciae falling within that of T. brassicae. Variations of both traits are correlated and strains can be easily classified according to their overall tendency to concentrate their attacks.The adaptive importance of the foraging behaviour of parasitoid insects suggests that differences reported here could reveal adaptive behavioural differentiations of natural populations in response to local variations in selective constraints, mainly those emanating from host diversity, abundance and distribution.

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Résumé Le mode de répartition des attaques parmi les hôtes est estimé par la distance moyenne entre les hôtes parasités dans une grande ooplaque homogène (500 hôtes) ou dans un système discontinu formé de six ooplaques de taille plus réduite (50 hôtes), régulièrement réparties sur un cercle.Pour chaque souche les deux caractères augmentent avec l'âge des femelles. Ils varient entre les souches, l'amplitude des variations au sein de l'espèce brassicae englobant celles observées chez cacoeciae.Les variations des deux caractères sont corrélées et les souches peuvent être classées suivant leur tendance plus ou moins marquée à concentrer leurs attaques.L'importance adaptative du comportement de colonisation des hôtes chez les insectes parasitoïdes laisse supposer que les différences constatées ici traduisent des différenciations locales des populations naturelles en réponse aux variations des contraintes sélectives, plus particulièrement celles qui sont liées à la diversité des hôtes, à leur abondance et à leur répartition.

     

Box-modeling of 15N/14N in mammals

The ¹⁵N/¹⁴N signature of animal proteins is now commonly used to understand their physiology and quantify the flows of nutrient in trophic webs. These studies assume that animals are predictably ¹⁵N-enriched relative to their food, but the isotopic mechanism which accounts for this enrichment remains unknown. We developed a box model of the nitrogen isotope cycle in mammals in order to predict the ¹⁵N/¹⁴N ratios of body reservoirs as a function of time, N intake and body mass. Results of modeling show that a combination of kinetic isotope fractionation during the N transfer between amines and equilibrium fractionation related to the reversible conversion of N-amine into ammonia is required to account for the well-established ≈4‰ ¹⁵N-enrichment of body proteins relative to the diet. This isotopic enrichment observed in proteins is due to the partial recycling of ¹⁵N-enriched urea and the urinary excretion of a fraction of the strongly ¹⁵N-depleted ammonia reservoir. For a given body mass and diet δ¹⁵N, the isotopic compositions are mainly controlled by the N intake. Increase of the urea turnover combined with a decrease of the N intake lead to calculate a δ¹⁵N increase of the proteins, in agreement with the observed increase of collagen δ¹⁵N of herbivorous animals with aridity. We further show that the low δ¹⁵N collagen values of cave bears cannot be attributed to the dormancy periods as it is commonly thought, but inversely to the hyperphagia behavior. This model highlights the need for experimental investigations performed with large mammals in order to improve our understanding of natural variations of δ¹⁵N collagen.     

Conceptual, methodological and computational issues concerning the compartmental modeling of a complex biological system: Postprandial inter-organ metabolism of dietary nitrogen in humans

A multi-compartmental model has been developed to describe dietary nitrogen (N) postprandial distribution and metabolism in humans. This paper details the entire process of model development, including the successive steps of its construction, parameter estimation and validation. The model was built using experimental data on dietary N kinetics in certain accessible pools of the intestine, blood and urine in healthy adults fed a [15N]-labeled protein meal. A 13-compartment, 21-parameter model was selected from candidate models of increasing order as being the minimum structure able to properly fit experimental data for all sampled compartments. Problems of theoretical identifiability and numerical identification of the model both constituted mathematical challenges that were difficult to solve because of the large number of unknown parameters and the few experimental data available. For this reason, new robust and reliable methods were applied, which enabled (i) a check that all model parameters could theoretically uniquely be determined and (ii) an estimation of their numerical values with satisfactory precision from the experimental data. Finally, model validation was completed by first verifying its a posteriori identifiability and then carrying out external validation.     

The nature of the dietary protein impacts the tissue-to-diet 15N discrimination factors in laboratory rats

Due to the existence of isotope effects on some metabolic pathways of amino acid and protein metabolism, animal tissues are (15)N-enriched relative to their dietary nitrogen sources and this (15)N enrichment varies among different tissues and metabolic pools. The magnitude of the tissue-to-diet discrimination (Δ(15)N) has also been shown to depend on dietary factors. Since dietary protein sources affect amino acid and protein metabolism, we hypothesized that they would impact this discrimination factor, with selective effects at the tissue level. To test this hypothesis, we investigated in rats the influence of a milk or soy protein-based diet on Δ(15)N in various nitrogen fractions (urea, protein and non-protein fractions) of blood and tissues, focusing on visceral tissues. Regardless of the diet, the different protein fractions of blood and tissues were generally (15)N-enriched relative to their non-protein fraction and to the diet (Δ(15)N>0), with large variations in the Δ(15)N between tissue proteins. Δ(15)N values were markedly lower in tissue proteins of rats fed milk proteins compared to those fed soy proteins, in all sampled tissues except in the intestine, and the amplitude of Δ(15)N differences between diets differed between tissues. Both between-tissue and between-diet Δ(15)N differences are probably related to modulations of the relative orientation of dietary and endogenous amino acids in the different metabolic pathways. More specifically, the smaller Δ(15)N values observed in tissue proteins with milk than soy dietary protein may be due to a slightly more direct channeling of dietary amino acids for tissue protein renewal and to a lower recycling of amino acids through fractionating pathways. In conclusion, the present data indicate that natural Δ(15)N of tissue are sensitive markers of the specific subtle regional modifications of the protein and amino acid metabolism induced by the protein dietary source.