The evolution of unicoloniality in ant societies : recognition, dispersal and population genetic structure

Introduction Societies of ants, bees, wasps and termites dominate many terrestrial ecosystems (Wilson 1971). Their evolutionary and ecological success is based upon the regulation of internal conflicts (e.g. Ratnieks et al. 2006), control of diseases (e.g. Schmid-Hempel 1998) and individual skills and collective intelligence in resource acquisition, nest building and defence (e.g. Camazine 2001). Individuals in social species can pass on their genes not only directly trough their own offspring, but also indirectly by favouring the reproduction of relatives. The inclusive fitness theory of Hamilton (1963; 1964) provides a powerful explanation for the evolution of reproductive altruism and cooperation in groups with related individuals. The same theory also led to the realization that insect societies are subject to internal conflicts over reproduction. Relatedness of less-than-one is not sufficient to eliminate all incentive for individual selfishness. This would indeed require a relatedness of one, as found among cells of an organism (Hardin 1968; Keller 1999). The challenge for evolutionary biology is to understand how groups can prevent or reduce the selfish exploitation of resources by group members, and how societies with low relatedness are maintained. In social insects the evolutionary shift from single- to multiple queens colonies modified the relatedness structure, the dispersal, and the mode of colony founding (e.g. (Crozier & Pamilo 1996). In ants, the most common, and presumably ancestral mode of reproduction is the emission of winged males and females, which found a new colony independently after mating and dispersal flights (Hölldobler & Wilson 1990). The alternative reproductive tactic for ant queens in multiple-queen colonies (polygyne) is to seek to be re-accepted in their natal colonies, where they may remain as additional reproductives or subsequently disperse on foot with part of the colony (budding) (Bourke & Franks 1995; Crozier & Pamilo 1996; Hölldobler & Wilson 1990). Such ant colonies can contain up to several hundred reproductive queens with an even more numerous workforce (Cherix 1980; Cherix 1983). As a consequence in polygynous ants the relatedness among nestmates is very low, and workers raise brood of queens to which they are only distantly related (Crozier & Pamilo 1996; Queller & Strassmann 1998). Therefore workers could increase their inclusive fitness by preferentially caring for their closest relatives and discriminate against less related or foreign individuals (Keller 1997; Queller & Strassmann 2002; Tarpy et al. 2004). However, the bulk of the evidence suggests that social insects do not behave nepotistically, probably because of the costs entailed by decreased colony efficiency or discrimination errors (Keller 1997). Recently, the consensus that nepotistic behaviour does not occur in insect colonies was challenged by a study in the ant Formica fusca (Hannonen & Sundström 2003b) showing that the reproductive share of queens more closely related to workers increases during brood development. However, this pattern can be explained either by nepotism with workers preferentially rearing the brood of more closely related queens or intrinsic differences in the viability of eggs laid by queens. In the first chapter, we designed an experiment to disentangle nepotism and differences in brood viability. We tested if workers prefer to rear their kin when given the choice between highly related and unrelated brood in the ant F. exsecta. We also looked for differences in egg viability among queens and simulated if such differences in egg viability may mistakenly lead to the conclusion that workers behave nepotistically. The acceptance of queens in polygnous ants raises the question whether the varying degree of relatedness affects their share in reproduction. In such colonies workers should favour nestmate queens over foreign queens. Numerous studies have investigated reproductive skew and partitioning of reproduction among queens (Bourke et al. 1997; Fournier et al. 2004; Fournier & Keller 2001; Hammond et al. 2006; Hannonen & Sundström 2003a; Heinze et al. 2001; Kümmerli & Keller 2007; Langer et al. 2004; Pamilo & Seppä 1994; Ross 1988; Ross 1993; Rüppell et al. 2002), yet almost no information is available on whether differences among queens in their relatedness to other colony members affects their share in reproduction. Such data are necessary to compare the relative reproductive success of dispersing and non-dispersing individuals. Moreover, information on whether there is a difference in reproductive success between resident and dispersing queens is also important for our understanding of the genetic structure of ant colonies and the dynamics of within group conflicts. In chapter two, we created single-queen colonies and then introduced a foreign queens originating from another colony kept under similar conditions in order to estimate the rate of queen acceptance into foreign established colonies, and to quantify the reproductive share of resident and introduced queens. An increasing number of studies have investigated the discrimination ability between ant workers (e.g. Holzer et al. 2006; Pedersen et al. 2006), but few have addressed the recognition and discrimination behaviour of workers towards reproductive individuals entering colonies (Bennett 1988; Brown et al. 2003; Evans 1996; Fortelius et al. 1993; Kikuchi et al. 2007; Rosengren & Pamilo 1986; Stuart et al. 1993; Sundström 1997; Vásquez & Silverman in press). These studies are important, because accepting new queens will generally have a large impact on colony kin structure and inclusive fitness of workers (Heinze & Keller 2000). In chapter three, we examined whether resident workers reject young foreign queens that enter into their nest. We introduced mated queens into their natal nest, a foreign-female producing nest, or a foreign male-producing nest and measured their survival. In addition, we also introduced young virgin and mated queens into their natal nest to examine whether the mating status of the queens influences their survival and acceptance by workers. On top of polgyny, some ant species have evolved an extraordinary social organization called 'unicoloniality' (Hölldobler & Wilson 1977; Pedersen et al. 2006). In unicolonial ants, intercolony borders are absent and workers and queens mix among the physically separated nests, such that nests form one large supercolony. Super-colonies can become very large, so that direct cooperative interactions are impossible between individuals of distant nests. Unicoloniality is an evolutionary paradox and a potential problem for kin selection theory because the mixing of queens and workers between nests leads to extremely low relatedness among nestmates (Bourke & Franks 1995; Crozier & Pamilo 1996; Keller 1995). A better understanding of the evolution and maintenance of unicoloniality requests detailed information on the discrimination behavior, dispersal, population structure, and the scale of competition. Cryptic genetic population structure may provide important information on the relevant scale to be considered when measuring relatedness and the role of kin selection. Theoretical studies have shown that relatedness should be measured at the level of the `economic neighborhood', which is the scale at which intraspecific competition generally takes place (Griffin & West 2002; Kelly 1994; Queller 1994; Taylor 1992). In chapter four, we conducted alarge-scale study to determine whether the unicolonial ant Formica paralugubris forms populations that are organised in discrete supercolonies or whether there is a continuous gradation in the level of aggression that may correlate with genetic isolation by distance and/or spatial distance between nests. In chapter five, we investigated the fine-scale population structure in three populations of F. paralugubris. We have developed mitochondria) markers, which together with the nuclear markers allowed us to detect cryptic genetic clusters of nests, to obtain more precise information on the genetic differentiation within populations, and to separate male and female gene flow. These new data provide important information on the scale to be considered when measuring relatedness in native unicolonial populations.

Pathway databases and tools for their exploitation: benefits, current limitations and challenges

In past years, comprehensive representations of cell signalling pathways have been developed by manual curation from literature, which requires huge effort and would benefit from information stored in databases and from automatic retrieval and integration methods. Once a reconstruction of the network of interactions is achieved, analysis of its structural features and its dynamic behaviour can take place. Mathematical modelling techniques are used to simulate the complex behaviour of cell signalling networks, which ultimately sheds light on the mechanisms leading to complex diseases or helps in the identification of drug targets. A variety of databases containing information on cell signalling pathways have been developed in conjunction with methodologies to access and analyse the data. In principle, the scenario is prepared to make the most of this information for the analysis of the dynamics of signalling pathways. However, are the knowledge repositories of signalling pathways ready to realize the systems biology promise? In this article we aim to initiate this discussion and to provide some insights on this issue.

Community structure in mediterranean shallow lentic ecosystems: size-based vs. taxon-based approaches

The main environmental variables determining the community structure and the functioning of Mediterranean shallow lentic ecosystems are described. These ecosystems are characterized by the unpredictability of their water inputs and the high variability in their water level and physical and chemical composition. Variations in flooding, salinity, and water turnover are determinant in species composition and nutrient dynamics. Taxon-based and size-based approaches to the study of the community structure of aquatic organisms that colonise these ecosystems are also compared. The conventional taxonomic approach, based on the determination of species composition, has been used for the identification of patterns in species richness, distribution and temporal dynamics, and for ecological requirements of species and their potential use as ecological indicators. This taxonbased approach has been compared with a size-based approach, where individuals are classified by their size. Size-based approach gives complementary information about community structure and dynamics, especially when communities are dominated by a single species. The use of size diversity combined with species diversity is suggested for a more complete understanding of community structuring in this type of ecosystem. Detailed examples of two Mediterranean shallow lentic ecosystems, the salt marshes of the Empordà wetlands and the Espolla temporary karstic pond, which differ in hydrology and water origin, are used to discuss the suitability of these different approaches

The fate and persistence of Leishmania major in mice of different genetic backgrounds: an example of exploitation of the immune system by intracellular parasites.

Leishmania spp. are intracellular protozoan parasites that are delivered within the dermis of their vertebrate hosts. Within this peripheral tissue and the draining lymph node, they find and/or rapidly create dynamic microenvironments that determine their ultimate fate, namely their more or less successful expansion, and favour their transmission to another vertebrate host though a blood-feeding vector. Depending on their genetic characteristics as well as the genetic make-up of their hosts, once within the dermis Leishmania spp. very rapidly drive and maintain sustained T cell-dependent immune responses that arbitrate their ultimate fate within their hosts. The analysis of the parasitism exerted by Leishmania major in mice of different genetic backgrounds has allowed us to recognize some of the early and late mechanisms driven by this parasite that lead to either uncontrolled or restricted parasitism. Uncontrolled parasitism by Leishmania major characterizing mice from a few inbred strains (e.g. BALB/c) is associated with the expansion of parasite reactive Th2 CD4 lymphocytes and results from their rapid and sustained activity. In contrast, restricted parasitism characteristic of mice from the majority of inbred strains results from the development of a polarized parasite-specific Th1 CD4 response. This murine model of infection has already been and will continue to be particularly instrumental in dissecting the rules controlling the pathway of differentiation of T cells in vivo. In the long run, the understanding of these rules should contribute to the rational development of novel immunotherapeutic interventions against severe infectious diseases.

Tropical terrestrial model ecosystems for evaluation of soil fauna and leaf litter quality effects on litter consumption, soil microbial biomass and plant growth

The aim of this work was to evaluate whether terrestrial model ecosystems (TMEs) are a useful tool for the study of the effects of litter quality, soil invertebrates and mineral fertilizer on litter decomposition and plant growth under controlled conditions in the tropics. Forty-eight intact soil cores (17.5-cm diameter, 30-cm length) were taken out from an abandoned rubber plantation on Ferralsol soil (Latossolo Amarelo) in Central Amazonia, Brazil, and kept at 28ºC in the laboratory during four months. Leaf litter of either Hevea pauciflora (rubber tree), Flemingia macrophylla (a shrubby legume) or Brachiaria decumbens (a pasture grass) was put on top of each TME. Five specimens of either Pontoscolex corethrurus or Eisenia fetida (earthworms), Porcellionides pruinosus or Circoniscus ornatus (woodlice), and Trigoniulus corallinus (millipedes) were then added to the TMEs. Leaf litter type significantly affected litter consumption, soil microbial biomass and nitrate concentration in the leachate of all TMEs, but had no measurable effect on the shoot biomass of rice seedlings planted in top soil taken from the TMEs. Feeding rates measured with bait lamina were significantly higher in TMEs with the earthworm P. corethrurus and the woodlouse C. ornatus. TMEs are an appropriate tool to assess trophic interactions in tropical soil ecossistems under controlled laboratory conditions.

Applying geostatistics for investigations of forest ecosystems using remote sensing imagery

Abstract

L'exploitation des ressources végétales dans les Pyrénées orientales durant l'holocène: analyse anthracologique des structures d'élevage, de four et de charbonnières dans l'Alt Urgell (Chaîne du Cadí) et la Vallée du Madriu (Andorre)

Dans ce travail, nous présentons le résultat des recherches anthracologiques réalisées sur des sites archéologiques de haute montagne dans les Pyrénées orientales. Cette recherche s’insère dans un projet d’archéologie du paysage. Les zones d’étude se situent dans les Pré-Pyrénées sur le versant méridional de la chaîne du Cadí (vallée de la Vansa, Alt Urgell), et dans la vallée du Madriu, située dans les Pyrénées axiales, en Andorre. Le but principal de l’analyse anthracologique a été de connaître et comprendre la dynamique et la relation qui unissent les populations qui ont occupé ces vallées avec le milieu forestier de haute montagne. Les échantillons ont été récupérés sur des structures archéologiques associés à l’exploitation forestière (charbonnières, fours pour la fabrication de résine), au pastoralisme (cabanes et enclos) et à l’exploitation minière-métallurgique (four de grillage), avec une chronologie qui débute au Néolithique Ancien et finit à l’époque moderne et contemporaine (XVIII-XIXe siècles). Les résultats indiquent une variabilité taxonomique pauvre avec la présence majoritaire du Pinus et la présence ponctuelle d’autres espèces arbustives (Ericaceae et Juniperus) et arborées, comme Betula alba ou Abies alba. Différentes formations végétales de haute montagne ont été identifiées, selon leur localisation en versant ensoleillé ou ombragé, ainsi que leur altitude. Les résultats montrent des traces d’exploitation forestière dès le Néolithique Ancien, une intensification des activités à l’Antiquité, même si l’impact anthropique majeur correspond à l’activité de charbonnage d’époque moderne.

Financial Exploitation

Financial exploitation is the unauthorized and illegal use of an individual’s funds, property or resources and includes identity theft. Financial exploitation can be committed by a family member, friend, neighbor or a complete stranger.

Détection et exploitation des traces de produits stupéfiants. Partie I : étude de leur processus d'échange et de leur utilisation dans un cadre policier

Cette publication présente une approche originale et novatrice de la recherche et de l'exploitation des traces de stupéfiants. Elle propose une description détaillée des phénomènes de transfert et de persistances de ces particules ainsi que l'interprétation des résultats obtenus par l'analyse de spectrométrie à mobilité ionique (IMS). Des simulations de manipulation et de conditionnement de produits stupéfiants ont été réalisées en distinguant plusieurs groupes de personnes en fonction de leur proximité de contact avec des stupéfiants. Ces simulations ont montré que des personnes n'étant pas en contact avec des produits stupéfiants ne présentaient pas de traces de ces derniers. Au contraire, une manipulation laisse des traces qu'il est possible de détecter et d'interpréter. Parmi les stupéfiants étudiés, le plus "contaminogène" s'est avéré être la cocaïne. Les résultats obtenus permettent enfin de proposer une démarche opérationnelle d'intégration de cette technologie dans le cadre des interventions effectuées par les brigades canines.

Les Morceaux de réception des graveurs à l'Académie royale des beaux-arts : 1655-1789 : fonds de la Bibliothèque nationale et de la chalcographie du musée du Louvre : [exposition]... 12 mai au 12 juin 1982, Musée galerie de la SEITA [Société nationale d'exploitation industrielle des tabacs et des allumettes]... Paris / catalogue [par Maxime Préaud]

[Exposition. Paris, Musée-galerie de la SEITA. 1982]