Similarities and differences in altitudinal versus latitudinal variation for morphological traits in Drosophila melanogaster.

Understanding how natural environments shape phenotypic variation is a major aim in evolutionary biology. Here, we have examined clinal, likely genetically based variation in morphology among 19 populations of the fruit fly (Drosophila melanogaster) from Africa and Europe, spanning a range from sea level to 3000 m altitude and including locations approximating the southern and northern range limit. We were interested in testing whether latitude and altitude have similar phenotypic effects, as has often been postulated. Both latitude and altitude were positively correlated with wing area, ovariole number, and cell number. In contrast, latitude and altitude had opposite effects on the ratio between ovariole number and body size, which was negatively correlated with egg production rate per ovariole. We also used transgenic manipulation to examine how increased cell number affects morphology and found that larger transgenic flies, due to a higher number of cells, had more ovarioles, larger wings, and, unlike flies from natural populations, increased wing loading. Clinal patterns in morphology are thus not a simple function of changes in body size; instead, each trait might be subject to different selection pressures. Together, our results provide compelling evidence for profound similarities as well as differences between phenotypic effects of latitude and altitude.

LKB1 regulates polarity remodeling and adherens junction formation in the Drosophila eye.

The serine-threonine kinase LKB1 regulates cell polarity from Caenorhabditis elegans to man. Loss of lkb1 leads to a cancer predisposition, known as Peutz-Jeghers Syndrome. Biochemical analysis indicates that LKB1 can phosphorylate and activate a family of AMPK- like kinases, however, the precise contribution of these kinases to the establishment and maintenance of cell polarity is still unclear. Recent studies propose that LKB1 acts primarily through the AMP kinase to establish and/or maintain cell polarity. To determine whether this simple model of how LKB1 regulates cell polarity has relevance to complex tissues, we examined lkb1 mutants in the Drosophila eye. We show that adherens junctions expand and apical, junctional, and basolateral domains mix in lkb1 mutants. Surprisingly, we find LKB1 does not act primarily through AMPK to regulate cell polarity in the retina. Unlike lkb1 mutants, ampk retinas do not show elongated rhabdomeres or expansion of apical and junctional markers into the basolateral domain. In addition, nutrient deprivation does not reveal a more dramatic polarity phenotype in lkb1 photoreceptors. These data suggest that AMPK is not the primary target of LKB1 during eye development. Instead, we find that a number of other AMPK-like kinase, such as SIK, NUAK, Par-1, KP78a, and KP78b show phenotypes similar to weak lkb1 loss of function in the eye. These data suggest that in complex tissues, LKB1 acts on an array of targets to regulate cell polarity.

Evolution under monogamy feminizes gene expression in Drosophila melanogaster.

Many genes have evolved sexually dimorphic expression as a consequence of divergent selection on males and females. However, because the sexes share a genome, the extent to which evolution can shape gene expression independently in each sex is controversial. Here, we use experimental evolution to reveal suboptimal sex-specific expression for much of the genome. By enforcing a monogamous mating system in populations of Drosophila melanogaster for over 100 generations, we eliminated major components of selection on males: female choice and male-male competition. If gene expression is subject to sexually antagonistic selection, relaxed selection on males should cause evolution towards female optima. Monogamous males and females show this pattern of feminization in both the whole-body and head transcriptomes. Genes with male-biased expression patterns evolved decreased expression under monogamy, while genes with female-biased expression evolved increased expression, relative to polygamous populations. Our results demonstrate persistent and widespread evolutionary tension between male and female adaptation.

Evolutionary dynamics of molecular markers during local adaptation: a case study in Drosophila subobscura

Background: Natural selection and genetic drift are major forces responsible for temporal genetic changes in populations. Furthermore, these evolutionary forces may interact with each other. Here we study the impact of an ongoing adaptive process at the molecular genetic level by analyzing the temporal genetic changes throughout 40 generations of adaptation to a common laboratory environment. Specifically, genetic variability, population differentiation and demographic structure were compared in two replicated groups of Drosophila subobscura populations recently sampled from different wild sources. Results: We found evidence for a decline in genetic variability through time, along with an increase in genetic differentiation between all populations studied. The observed decline in genetic variability was higher during the first 14 generations of laboratory adaptation. The two groups of replicated populations showed overall similarity in variability patterns. Our results also revealed changing demographic structure of the populations during laboratory evolution, with lower effective population sizes in the early phase of the adaptive process. One of the ten microsatellites analyzed showed a clearly distinct temporal pattern of allele frequency change, suggesting the occurrence of positive selection affecting the region around that particular locus. Conclusion: Genetic drift was responsible for most of the divergence and loss of variability between and within replicates, with most changes occurring during the first generations of laboratory adaptation. We also found evidence suggesting a selective sweep, despite the low number of molecular markers analyzed. Overall, there was a similarity of evolutionary dynamics at the molecular level in our laboratory populations, despite distinct genetic backgrounds and some differences in phenotypic evolution.

Conserved chromosomal clustering of genes governed by chromatin regulators in Drosophila

Background: The trithorax group (trxG) and Polycomb group (PcG) proteins are responsible for the maintenance of stable transcriptional patterns of many developmental regulators. They bind to specific regions of DNA and direct the post-translational modifications of histones, playing a role in the dynamics of chromatin structure.Results: We have performed genome-wide expression studies of trx and ash2 mutants in Drosophila melanogaster. Using computational analysis of our microarray data, we have identified 25 clusters of genes potentially regulated by TRX. Most of these clusters consist of genes that encode structural proteins involved in cuticle formation. This organization appears to be a distinctive feature of the regulatory networks of TRX and other chromatin regulators, since we have observed the same arrangement in clusters after experiments performed with ASH2, as well as in experiments performed by others with NURF, dMyc, and ASH1. We have also found many of these clusters to be significantly conserved in D. simulans, D. yakuba, D. pseudoobscura and partially in Anopheles gambiae.Conclusion: The analysis of genes governed by chromatin regulators has led to the identification of clusters of functionally related genes conserved in other insect species, suggesting this chromosomal organization is biologically important. Moreover, our results indicate that TRX and other chromatin regulators may act globally on chromatin domains that contain transcriptionally co-regulated genes.

Distribution of the transposable elements bilbo and gypsy in original and colonizing populations of Drosophila subobscura

Background: Transposable elements (TEs) constitute a substantial amount of all eukaryotic genomes. They induce an important proportion of deleterious mutations by insertion into genes or gene regulatory regions. However, their mutational capabilities are not always adverse but can contribute to the genetic diversity and evolution of organisms. Knowledge of their distribution and activity in the genomes of populations under different environmental and demographic regimes, is important to understand their role in species evolution. In this work we study the chromosomaldistribution of two TEs, gypsy and bilbo, in original and colonizing populations of Drosophilasubobscura to reveal the putative effect of colonization on their insertion profile.Results: Chromosomal frequency distribution of two TEs in one original and three colonizingpopulations of D. subobscura, is different. Whereas the original population shows a low insertionfrequency in most TE sites, colonizing populations have a mixture of high (frequency ¿ 10%) andlow insertion sites for both TEs. Most highly occupied sites are coincident among colonizingpopulations and some of them are correlated to chromosomal arrangements. Comparisons of TEcopy number between the X chromosome and autosomes show that gypsy occupancy seems to becontrolled by negative selection, but bilbo one does not. Conclusion: These results are in accordance that TEs in Drosophila subobscura colonizing populations are submitted to a founder effect followed by genetic drift as a consequence of colonization. This would explain the high insertion frequencies of bilbo and gypsy in coincident sites of colonizing populations. High occupancy sites would represent insertion events prior to colonization. Sites of low frequency would be insertions that occurred after colonization and/orcopies from the original population whose frequency is decreasing in colonizing populations. Thiswork is a pioneer attempt to explain the chromosomal distribution of TEs in a colonizing specieswith high inversion polymorphism to reveal the putative effect of arrangements in TE insertionprofiles. In general no associations between arrangements and TE have been found, except in a fewcases where the association is very strong. Alternatively, founder drift effects, seem to play aleading role in TE genome distribution in colonizing populations.

Changes in chromosomal polymorphism and global warming: The case of Drosophila subobscura from Apatin (Serbia)

Abstract In this study, chromosomal inversion polymorphism data for a natural population of Drosophila subobscura from a swampy region near the town of Apatin (Serbia) were compared with data for the same population collected approximately 15 years earlier. The pattern of chromosomal inversion polymorphism changed over time. There were significant increases in the frequency of characteristic southern latitude ("warm" adapted) chromosomal arrangements and significant decreases in the frequency of characteristic northern latitude ("cold" adapted) chromosomal arrangements in the O and U chromosomes. The chromosomal arrangements O3+4 and O3+4+22 (derived from the O3+4 arrangement)showed significant increases in 2008 and 2009 with regard to the 1994 sample. There was also a significant increase (~50%) in the U1+2 arrangement, while U1+8+2 (a typical southern arrangement) was detected for the first time. Since the Apatin swampy population ofD. subobscura has existed for a long time in a stable habitat with high humidity that has not been changed by man our results indicate that natural selection has produced chromosomal changes in response to the increase in temperature that has occurred in the Balkan Peninsula of central southeastern European. Key words: chromosomal inversions, Drosophila subobscura, global warming, karyotypes.

Molecular evidence to suggest the origin of a colonization: Drosophila subobscura in America

Abstract The recent colonization of America by Drosophila subobscura represents a great opportunity for evolutionary biology studies. Knowledge of the populations from which the colonization started would provide an understanding of how genetic composition changed during adaptation to the new environment. Thus, a 793 nucleotide fragment of the Odh (Octanol dehydrogenase) gene was sequenced in 66 chromosomal lines from Barcelona (western Mediterranean) and in 66 from Mt. Parnes (Greece, eastern Mediterranean). No sequence of Odh fragment in Barcelona or Mt. Parnes was identical to any of those previously detected in America. However, an Odh sequence from Barcelona differed in only one nucleotide from another found in American populations. In both cases, the chromosomal lines presented the same inversion: O7, and the Odh gene was located within this inversion. This evidence suggests a possible western Mediterranean origin for the colonization. Finally, the molecular and inversion data indicate that the colonization was not characterized by multiple reintroductions.

O-chromosome lethal frequencies in Serbian and Montenegrin Drosophila subobscura populations

Lethal chromosomal frequencies were obtained from three Drosophila subobscura samples from the Mt. Avala (Serbia) population in September 2003 (0.218), June 2004 (0.204) and September 2004 (0.250). These values and those from other Balkan populations studied previously (Petnica, Kamariste, Zanjic and Djerdap) were used to analyze the possible effect of population, year, month and altitude above sea level on lethal chromosomal frequencies. According to ANOVAS no effect were observed. Furthermore, the lethal frequencies of the Balkan populations did not vary according to latitude. This is probably due to the relative proximity and high gene flow between these populations. From a joint study of all the Palearctic D. subobscura populations so far analyzed, it can be deduced that the Balkan populations are located in the central area of the species distribution. Finally, it seems that lethal chromosomal frequencies are a consequence of the genetic structure of the populations.

Thermal evolution of gene expression profiles in Drosophila subobscura

Background: Despite its pervasiveness, the genetic basis of adaptation resulting in variation directly or indirectly related to temperature (climatic) gradients is poorly understood. By using 3-fold replicated laboratory thermal stocks covering much of the physiologically tolerable temperature range for the temperate (i.e., cold tolerant) species Drosophila subobscura we have assessed whole-genome transcriptional responses after three years of thermal adaptation, when the populations had already diverged for inversion frequencies, pre-adult life history components, and morphological traits. Total mRNA from each population was compared to a reference pool mRNA in a standard, highly replicated two-colour competitive hybridization experiment using cDNA microarrays.Results: A total of 306 (6.6%) cDNA clones were identified as 'differentially expressed' (following a false discovery rate correction) after contrasting the two furthest apart thermal selection regimes (i.e., 13°C vs . 22°C), also including four previously reported candidate genes for thermotolerance in Drosophila (Hsp26, Hsp68, Fst, and Treh). On the other hand, correlated patterns of gene expression were similar in cold- and warm-adapted populations. Analysis of functional categories defined by the Gene Ontology project point to an overrepresentation of genes involved in carbohydrate metabolism, nucleic acids metabolism and regulation of transcription among other categories. Although the location of differently expressed genes was approximately at random with respect to chromosomes, a physical mapping of 88 probes to the polytene chromosomes of D. subobscura has shown that a larger than expected number mapped inside inverted chromosomal segments.Conclusion: Our data suggest that a sizeable number of genes appear to be involved in thermal adaptation in Drosophila, with a substantial fraction implicated in metabolism. This apparently illustrates the formidable challenge to understanding the adaptive evolution of complex trait variation. Furthermore, some clustering of genes within inverted chromosomal sections was detected. Disentangling the effects of inversions will be obviously required in any future approach if we want to identify the relevant candidate genes.

Comparative genomic analysis of the odorant-binding protein family in 12 Drosophila genomes: purifying selection and birth-and-death evolution

Background: Chemoreception is a widespread mechanism that is involved in critical biologic processes, including individual and social behavior. The insect peripheral olfactory system comprises three major multigene families: the olfactory receptor (Or), the gustatory receptor (Gr), and the odorant-binding protein (OBP) families. Members of the latter family establish the first contact with the odorants, and thus constitute the first step in the chemosensory transduction pathway.Results: Comparative analysis of the OBP family in 12 Drosophila genomes allowed the identification of 595 genes that encode putative functional and nonfunctional members in extant species, with 43 gene gains and 28 gene losses (15 deletions and 13 pseudogenization events). The evolution of this family shows tandem gene duplication events, progressive divergence in DNA and amino acid sequence, and prevalence of pseudogenization events in external branches of the phylogenetic tree. We observed that the OBP arrangement in clusters is maintained across the Drosophila species and that purifying selection governs the evolution of the family; nevertheless, OBP genes differ in their functional constraints levels. Finally, we detect that the OBP repertoire evolves more rapidly in the specialist lineages of the Drosophila melanogaster group (D. sechellia and D. erecta) than in their closest generalists.Conclusion: Overall, the evolution of the OBP multigene family is consistent with the birth-and-death model. We also found that members of this family exhibit different functional constraints, which is indicative of some functional divergence, and that they might be involved in some of the specialization processes that occurred through the diversification of the Drosophila genus.

Inference of chromosomal inversion dynamics from Pool-Seq data in natural and laboratory populations of Drosophila melanogaster.

Sequencing of pools of individuals (Pool-Seq) represents a reliable and cost-effective approach for estimating genome-wide SNP and transposable element insertion frequencies. However, Pool-Seq does not provide direct information on haplotypes so that, for example, obtaining inversion frequencies has not been possible until now. Here, we have developed a new set of diagnostic marker SNPs for seven cosmopolitan inversions in Drosophila melanogaster that can be used to infer inversion frequencies from Pool-Seq data. We applied our novel marker set to Pool-Seq data from an experimental evolution study and from North American and Australian latitudinal clines. In the experimental evolution data, we find evidence that positive selection has driven the frequencies of In(3R)C and In(3R)Mo to increase over time. In the clinal data, we confirm the existence of frequency clines for In(2L)t, In(3L)P and In(3R)Payne in both North America and Australia and detect a previously unknown latitudinal cline for In(3R)Mo in North America. The inversion markers developed here provide a versatile and robust tool for characterizing inversion frequencies and their dynamics in Pool-Seq data from diverse D. melanogaster populations.

Mechanosensory interactions drive collective behaviour in Drosophila.

Collective behaviour enhances environmental sensing and decision-making in groups of animals. Experimental and theoretical investigations of schooling fish, flocking birds and human crowds have demonstrated that simple interactions between individuals can explain emergent group dynamics. These findings indicate the existence of neural circuits that support distributed behaviours, but the molecular and cellular identities of relevant sensory pathways are unknown. Here we show that Drosophila melanogaster exhibits collective responses to an aversive odour: individual flies weakly avoid the stimulus, but groups show enhanced escape reactions. Using high-resolution behavioural tracking, computational simulations, genetic perturbations, neural silencing and optogenetic activation we demonstrate that this collective odour avoidance arises from cascades of appendage touch interactions between pairs of flies. Inter-fly touch sensing and collective behaviour require the activity of distal leg mechanosensory sensilla neurons and the mechanosensory channel NOMPC. Remarkably, through these inter-fly encounters, wild-type flies can elicit avoidance behaviour in mutant animals that cannot sense the odour--a basic form of communication. Our data highlight the unexpected importance of social context in the sensory responses of a solitary species and open the door to a neural-circuit-level understanding of collective behaviour in animal groups.

Natural variation in learning ability in "Drosophila melanogaster"

Learning is the ability of an organism to adapt to the changes of its environment in response to its past experience. It is a widespread ability in the animal kingdom, but its evolutionary aspects are poorly known. Learning ability is supposedly advantageous under some conditions, when environmental conditions are not too stable - because in this case there is no need to learn to predict any event in the environment - and not changing too fast - otherwise environmental cues cannot be used because they are not reliable. Nevertheless, learning ability is also known to be costly in terms of energy needed for neuronal synthesis, memory formation, initial mistakes. During my PhD, I focused on the study of genetic variability of learning ability in natural populations. Genetic variability is the basis on which natural selection and genetic drift can act. How does learning ability vary in nature? What are the roles of additive genetic variation or maternal effects in this variation? Is it involved in evolutionary trade-offs with other fitness-related traits?¦I investigated a natural population of fruit fly, Drosophila melanogaster, as a model organism. Its learning ability is easy to measure with associative memory tests. I used two research tools: multiple inbred and isofemale lines derived from a natural population as a representative sample. My work was divided into three parts.¦First, I investigated the effects of inbreeding on aversive learning (avoidance of an odor previously associated with mechanical shock). While the inbred lines consistently showed reduced egg-to-adult viability by 28 %, the effects of inbreeding on learning performance was 18 % and varied among assays, with a trend to be most pronounced for intermediate conditioning intensity. Variation among inbred lines indicates that ample genetic variance for learning was segregating in the base population, and suggests that the inbreeding depression observed in learning performance was mostly due to dominance rather than overdominance. Across the inbred lines, learning performance was positively correlated with the egg-to-adult viability. This positive genetic correlation contradicts previous studies which observed a trade-off between learning ability and lifespan or larval competitive ability. It suggests that much of the genetic variation for learning is due to pleiotropic effects of genes affecting other functions related to survival. Together with the overall mild effects of inbreeding on learning performance, this suggests that genetic variation specifically affecting learning is either very low, or is due to alleles with mostly additive (semi-dominant) effects. It also suggests that alleles reducing learning performance are on average partially recessive, because their effect does not appear in the outbred base population. Moreover, overdominance seems unlikely as major cause of the inbreeding depression, because even if the overall mean of the inbred line is smaller than the outbred base population, some of the inbred lines show the same learning score as the outbred base population. If overdominance played an important part in inbreeding depression, then all the homozygous lines should show lower learning ability than¦outbred base population.¦In the second part of my project, I sampled the same natural population again and derived isofemale lines (F=0.25) which are less adapted to laboratory conditions and therefore are more representative of the variance of the natural population. They also showed some genetic variability for learning, and for three other fitness-related traits possibly related with learning: resistance to bacterial infection, egg-to-adult viability and developmental time. Nevertheless, the genetic variance of learning ability did not appear to be smaller than the variance of the other traits. The positive correlation previously observed between learning ability and egg- to-adult viability did not appear in isofemale lines (nor a negative correlation). It suggests that there was still genetic variability within isofemale lines and that they did not fix the highly deleterious pleiotropic alleles possibly responsible for the previous correlation.¦In order to investigate the relative amount of nuclear (additive and non-additive effects) and extra-nuclear (maternal and paternal effect) components of variance in learning ability and other fitness-related traits among the inbred lines tested in part one, I performed a diallel cross between them. The nuclear additive genetic variance was higher than other components for learning ability and survival to learning ability, but in contrast, maternal effects were more variable than other effects for developmental traits. This suggests that maternal effects, which reflects effects from mitochondrial DNA, epigenetic effects, or the amount of nutrients that are invested by the mother in the egg, are more important in the early stage of life, and less at the adult stage. There was no additive genetic correlation between learning ability and other traits, indicating that the correlation between learning ability and egg-to-adult viability observed in the first pat of my project was mostly due to recessive genes.¦Finally, my results showed that learning ability is genetically variable. The diallel experiment showed additive genetic variance was the most important component of the total variance. Moreover, every inbred or isofemale line showed some learning ability. This suggested that alleles impairing learning ability are eliminated by selection, and therefore that learning ability is under strong selection in natural populations of Drosophila. My results cannot alone explain the maintenance of the observed genetic variation. Even if I cannot eliminate the hypothesis of pleiotropy between learning ability and the other fitness-related traits I measured, there is no evidence for any trade-off between these traits and learning ability. This contradicts what has been observed between learning ability and other traits like lifespan and larval competitivity.¦L'apprentissage représente la capacité d'un organisme à s'adapter aux changement de son environnement au cours de sa vie, en réponse à son expérience passée. C'est une capacité très répandue dans le règne animal, y compris pour les animaux les plus petits et les plus simples, mais les aspects évolutifs de l'apprentissage sont encore mal connus. L'apprentissage est supposé avantageux dans certaines conditions, quand l'environnement n'est ni trop stable - dans ce cas, il n'y a rien à apprendre - ni trop variable - dans ce cas, les indices sur lesquels se reposer changent trop vite pour apprendre. D'un autre côté, l'apprentissage a aussi des coûts, en terme de synthèse neuronale, pour la formation de la mémoire, ou de coûts d'erreur initiale d'apprentissage. Pendant ma thèse, j'ai étudié la variabilité génétique naturelle des capacités d'apprentissage. Comment varient les capacités d'apprentissage dans la nature ? Quelle est la part de variation additive, l'impact des effets maternel ? Est-ce que l'apprentissage est impliqué dans des interactions, de type compromis évolutifs, avec d'autres traits liés à la fitness ?¦Afin de répondre à ces questions, je me suis intéressée à la mouche du vinaigre, ou drosophile, un organisme modèle. Ses capacités d'apprentissage sont facile à étudier avec un test de mémoire reposant sur l'association entre un choc mécanique et une odeur. Pour étudier ses capacités naturelles, j'ai dérivé de types de lignées d'une population naturelle: des lignées consanguines et des lignées isofemelles.¦Dans une première partie, je me suis intéressée aux effets de la consanguinité sur les capacités d'apprentissage, qui sont peu connues. Alors que les lignées consanguines ont montré une réduction de 28% de leur viabilité (proportion d'adultes émergeants d'un nombre d'oeufs donnés), leurs capacités d'apprentissage n'ont été réduites que de 18%, la plus forte diminution étant obtenue pour un conditionnement modéré. En outre, j'ai également observé que les capacités d'apprentissage était positivement corrélée à la viabilité entre les lignées. Cette corrélation est surprenante car elle est en contradiction avec les résultats obtenus par d'autres études, qui montrent l'existence de compromis évolutifs entre les capacités d'apprentissage et d'autres traits comme le vieillissement ou la compétitivité larvaire. Elle suggère que la variation génétique des capacités d'apprentissage est due aux effets pleiotropes de gènes récessifs affectant d'autres fonctions liées à la survie. Ces résultats indiquent que la variation pour les capacités d'apprentissage est réduite comparée à celle d'autres traits ou est due à des allèles principalement récessifs. L'hypothèse de superdominance semble peu vraisemblable, car certaines des lignées consanguines ont obtenu des scores d'apprentissage égaux à ceux de la population non consanguine, alors qu'en cas de superdominance, elles auraient toutes dû obtenir des scores inférieurs.¦Dans la deuxième partie de mon projet, j'ai mesuré les capacités d'apprentissage de lignées isofemelles issues de la même population initiale que les lignées consanguines. Ces lignées sont issues chacune d'un seul couple, ce qui leur donne un taux d'hétérozygosité supérieur et évite l'élimination de lignées par fixation d'allèles délétères rares. Elles sont ainsi plus représentatives de la variabilité naturelle. Leur variabilité génétique est significative pour les capacités d'apprentissage, et trois traits liés à la fois à la fitness et à l'apprentissage: la viabilité, la résistance à l'infection bactérienne et la vitesse de développement. Cependant, la variabilité des capacités d'apprentissage n'apparaît cette fois pas inférieure à celle des autres traits et aucune corrélation n'est constatée entre les capacité d'apprentissage et les autres traits. Ceci suggère que la corrélation observée auparavant était surtout due à la fixation d'allèles récessifs délétères également responsables de la dépression de consanguinité.¦Durant la troisième partie de mon projet, je me suis penchée sur la décomposition de la variance observée entre les lignées consanguines observée en partie 1. Quatre composants ont été examinés: la variance due à des effets nucléaires (additifs et non additifs), et due à des effets parentaux (maternels et paternels). J'ai réalisé un croisement diallèle de toutes les lignées. La variance additive nucléaire s'est révélée supérieure aux autres composants pour les capacités d'apprentissage et la résistance à l'infection bactérienne. Par contre, les effets maternels étaient plus importants que les autres composants pour les traits développementaux (viabilité et vitesse de développement). Ceci suggère que les effets maternels, dus à G ADN mitochondrial, à l'épistasie ou à la quantité de nutriments investis dans l'oeuf par la mère, sont plus importants dans les premiers stades de développement et que leur effet s'estompe à l'âge adulte. Il n'y a en revanche pas de corrélation statistiquement significative entre les effets additifs des capacités d'apprentissage et des autres traits, ce qui indique encore une fois que la corrélation observée entre les capacités d'apprentissage et la viabilité dans la première partie du projet était due à des effets d'allèles partiellement récessifs.¦Au, final, mes résultats montrent bien l'existence d'une variabilité génétique pour les capacités d'apprentissage, et l'expérience du diallèle montre que la variance additive de cette capacité est importante, ce qui permet une réponse à la sélection naturelle. Toutes les lignées, consanguines ou isofemelles, ont obtenu des scores d'apprentissage supérieurs à zéro. Ceci suggère que les allèles supprimant les capacités d'apprentissage sont fortement contre-sélectionnés dans la nature Néanmoins, mes résultats ne peuvent pas expliquer le maintien de cette variabilité génétique par eux-même. Même si l'hypothèse de pléiotropie entre les capacités d'apprentissage et l'un des traits liés à la fitness que j'ai mesuré ne peut être éliminée, il n'y a aucune preuve d'un compromis évolutif pouvant contribuer au maintien de la variabilité.