Population genetic structure in the native and introduced range of the Argentine ant/

SUMMARY : The evolution of animal societies, where some individuals forego their own reproductive opportunities to help others to reproduce, poses an evolutionary paradox that can be traced back to Darwin. Altruism may evolve through kin selection when the donor and recipient of altruistic acts are related to each other. In social insects, workers are generally highly related to the brood they rear when colonies are headed by a single queen. Yet some ants have an extraordinary social organization, called unicoloniality, whereby individuals from separate nests mix freely to form large supercolonies, which in some cases extend over hundreds of km. These supercolonies are characterised by a high number of queens (polygyny) and an absence of clear colony boundaries. This type of social organization represents an evolutionary paradox because relatedness between nestmates is effectively zero. In such conditions, kin selection cannot account for the evolution of reproductive altruism. Moreover, unicoloniality is thought to be unstable over time, because workers that can no longer aid close relatives may evolve more selfish strategies. The Argentine ant (Linepithema humile) is a highly invasive species listed among the hundred world's worst invaders by the UICN. Native from South America, L. humile has been accidentally introduced throughout the world. Native populations have been described as noninvasive with a family-based organization. In contrast, within its introduction range, they form unicolonial supercolonies that contain numerous nests without intraspecific aggression. The development of such unicolonial populations has been explained as a direct consequence of the ant's introduction into a new habitat, favouring a transition from family-based to open colonies. To determine if the social structure of the Argentine ant is fundamentally different between the native and the introduced range, we studied genetically and behaviourally native and introduced populations of L. humile over different geographic scales. Our results clearly indicated that there are no fundamental differences in the social organisation of the Argentine ant between the two ranges. Our investigations revealed that, contrary to previous claims, native populations have a unicolonial social organisation very similar to that observed in the introduced range. Consequently, the unicolonial social structure of the Argentine ant does not stem from a shift in social organization associated with introduction into new habitats but evolved in the native range and is likely a stable, evolutionarily ancient adaptation to the local environment. Our study on native populations of L. humile also gave important insight in the comprehension of the evolution of unicoloniality in the Argentine ant. Native supercolonies are relatively small compared to introduced ones and may co-habit in a same population. These supercolonies are genetically highly differentiated leading to a significant relatedness among nestmate workers when the different supercolonies of a population are taken as a reference population. This provides the necessary conditions for loin selection to operate. Furthermore, we examined a native population over time, which revealed a high supercolony extinction rate. If more competitive supercolonies are more likely to survive or replace other supercolonies, a subtle dynamical process between the spread of selfish traits within supercolony and the selective elimination of supercolonies with such traits may allow a stable equilibrium and the persistence of unicoloniality over time. Finally, a worldwide study of the Argentine ant showed that the introduced supercolonies originate from numerous independent introduction events. In conclusion, the success of the Argentine ant does not stem from a shift in social organization associated with its introduction into new habitats, but is most probably explained by the intrinsic characteristics developed in its native range. RESUME : L'altruisme de reproduction où certains individus renoncent à leur propre reproduction pour aider d'autres individus à se reproduire constitue l'un des plus grand paradoxe de l'évolution. En effet, comment expliquer l'évolution de comportements qui tendent à augmenter les chances de survie et le succès reproductif d'autres individus, alors que ces actes diminuent l'aptitude de leurs auteurs ? La théorie de la sélection de parentèle permet de résoudre ce problème. Cette théorie stipule qu'en aidant de proches parents à se reproduire, les individus peuvent promouvoir indirectement la transmission de copies de leurs propres gènes à la génération suivante. Chez les insectes sociaux, l'altruisme des ouvrières s'explique par la théorie de sélection de parentèle lorsque les colonies sont monogynes (constituées d'une seule reine) puisque les ouvrières sont fortement apparentées aux couvains dont elles s'occupent. Par contre, les espèces dites unicoloniales, dont les colonies forment des réseaux de nids appelés supercolonies, représentent toujours un paradoxe pour les théories de l'évolution puisque l'apparentement entre les différents individus d'un nid est nulle. De plus, l'unicolonialité ne devrait pas être stable sur le long terme parce que les ouvrières qui ne s'occupent plus de leur apparentés devraient développer des stratégies plus égoïstes au cours du temps. La fourmi d'Argentine (Linepithema humile) est une espèce invasive ayant un impact considérable sur son environnement. Originaire d'Amérique du Sud, elle a été introduite dans pratiquement toutes les régions du monde dont le climat est de type méditerranéen. Son incroyable succès invasif s'explique par sa structure sociale unicoloniale observée dans chacun des pays où elle a été introduite. Par contre, les rares études effectuées en Argentine ont suggéré que la fourmi d'Argentine n'était pas unicoloniale dans son aire native. L'unicolonialité chez la fourmi d'Argentine était donc considéré comme une conséquence de son introduction dans de nouveaux environnements. Durant cette thèse, nous avons vérifié si la structure sociale de cette espèce différait fondamentalement entre l'aire native et introduite. Pour cela, nous avons étudié, à différentes échelles géographiques, des populations introduites et argentines avec une approche génétique et comportementale. L'ensemble de nos résultats montrent que les différences entre les deux structure sociales ne sont pas aussi importantes que ce que l'on imaginait. Les populations natives sont aussi constituées de réseaux de nids coopérants. La taille de ses supercolonies est toutefois bien moins importante en Argentine et il n'est pas rare de trouver plusieurs supercolonies cohabitantes dans une même population. Nous avons démontré que ces réseaux de nids étaient constitués d'individus qui sont plus apparentés entre eux qu'ils ne le sont avec les individus d'autres supercolonies, ainsi l'unicolonialité dans son aire d'origine ne représente pas un réel paradoxe pour les théories de l'évolution. Finalement nous avons étudié la même population en Argentine à six ans d'intervalle et avons constaté que les supercolonies avaient un taux de survie très faible ce qui pourrait expliquer la stabilité de l'unicolonialité au cours du temps. Si les supercolonies les plus compétitives survivent mieux que les supercolonies dans lesquelles apparaissent des traits égoïstes, on devrait alors observer une dynamique entre l'apparition de traits égoïstes et l'élimination des supercolonies dans lesquelles ces traits égoïstes évolueraient. Finalement, une étude mondiale nous a montré que les supercolonies étaient originaires de nombreux événements d'introductions indépendants. En conclusion, le succès invasif de la fourmi d'Argentine n'est donc pas dû à un changement de comportement associé à son introduction mais est lié aux caractéristiques qu'elle a développées en Argentine.

Cooperació i teoria de jocs, el programari lliure o el #15m, dos exemples per l'universitat

Xerrada de cloenda de la Setmana internacional d'accés obert 2011 a la UOC, a càrrec de l'advocat Josep Jover. Per què les estratègies altruistes guanyen les egoistes en el programari lliure i en el #15m? El moviment #15m, igual que el programari, a diferència dels béns materials, no es pot posseir, ja que en pot gaudir (formant-ne part) un nombre indeterminat de persones sense que per això hagi de privar ningú de tenir-lo al seu torn. I això porta a girar com un mitjó la manera com manegen la informació les universitats, i quina és la missió de la universitat en la nova societat. En el futur immediat, valorarem les universitats no per la informació que guarden, que fora sempre serà millor i més extensa, sinó per la capacitat de crear masses crítiques, sia de recerca de coneixement, de capacitació humana, d'enllaç entre iguals... Les universitats hauran d'implantar el model o quedaran relegades.

Cooperación y teoría de juegos, el software libre o el #15m, dos ejemplos para la universidad

Xerrada de cloenda de la Setmana internacional d'accés obert 2011 a la UOC, a càrrec de l'advocat Josep Jover. Per què les estratègies altruistes guanyen les egoistes en el programari lliure i en el #15m? El moviment #15m, igual que el programari, a diferència dels béns materials, no es pot posseir, ja que en pot gaudir (formant-ne part) un nombre indeterminat de persones sense que per això hagi de privar ningú de tenir-lo al seu torn. I això porta a girar com un mitjó la manera com manegen la informació les universitats, i quina és la missió de la universitat en la nova societat. En el futur immediat, valorarem les universitats no per la informació que guarden, que fora sempre serà millor i més extensa, sinó per la capacitat de crear masses crítiques, sia de recerca de coneixement, de capacitació humana, d'enllaç entre iguals... Les universitats hauran d'implantar el model o quedaran relegades.

Cooperation and game theory, free software or the #15m movement: Two examples for universities

Closing talk of the Open Access Week 2011 at the UOC, by Josep Jover. Why do altruistic strategies beat selfish ones in the spheres of both free software and the #15m movement? The #15m movement, like software but unlike tangible goods, cannot be owned. It can be used (by joining it) by an indeterminate number of people without depriving anyone else of the chance to do the same. And that turns everything on its head: how universities manage information and what their mission is in this new society. In the immediate future, universities will be valued not for the information they harbour, which will always be richer and more extensive beyond their walls, but rather for their capacity to create critical masses, whether of knowledge research, skill-building, or networks of peers... universities must implement the new model or risk becoming obsolete.

Rousseau's Child: Preschoolers Expect Strangers to Favor Prosocial Actions

Modern thinking about human nature is notoriously divided between two contradictory notions: The Hobbesian tradition portrays men as driven by selfish desires, while the Rousseauian tradition recognizes altruistic proclivities as truemotivations to cooperate. We tested preschoolers' predictions about the prosocial or antisocial manner in which people would behave toward each other. Four stories were presented to 3- and 4-year-old children. In each story, the protagonists could either cooperate, act in terms of their own interests, or adopt a behavior unrelated to the ongoing scenario. Children as young as 3 years of age expected the protagonists to behave prosocially - and even more so if the protagonists were female. The results suggest that, even at an early age, children are inclined to adopt a "Rousseau-like" stance rather than a "suspicious" or "pessimistic" Hobbesian stance.

Altruism, spite and choosy females

Natural selection favours the genes which are able to introduce replicates of themselves in the next generation with higher certainty than do rival genes (Hamilton 1963). The fitness of an individual, it?s ability to produce future parents, depends on it?s own behaviour as well as on the behaviour of other individuals in the population. For instance, the intensity of competition an individual experience depends on the exploitation of resources by neighbours. The fitness is thus frequency dependent on what neighbours do. Behaviours can be classified according to the costs and benefits they have on the fitness of the behaver and it?s neighbours (Hamilton 1964, Hamilton 1975). According to this classification there exist four distinct social behaviours. (1) A gene confering the ability to use a new ressource is called selfish because it has a positive e_ect on the bearer of the gene but a negative e_ect on neighbours by the concomitant increase in competition. (2) An altruistic behaviour is defined as an action where an individual increases the fitness of a neighbour at the expense of it?s own. The e_ect is deleterious for the actor but positive for the receptor. (3) More surprinsingly, an individual might sacrifice a fraction of it?s ressources to harm another at no direct benefits. This spitefull behaviour incurs a cost for the actor but is also deleterious for the receptor. (4) Finally a cooperative behaviour breeds benefits for both actors and neighbours. In this thesis I will continue on the path traced by numerous evolutionnary biologist which attempt to fine tune our understanding of the evolution of social behaviours since Hamilton?s foundation (1963, 1964). A critical development over the last 40 years has been the realisation that competition between kin can partly or completely cancel out the role of relatedness as an agent favouring altruism (Wilson et al., 1992; Taylor, 1992a,b). Of importance is thus to determine the scale at which competition and altruism occur. One mechanism avoiding the complete dilution of relatedness by competition is the conditionnal expression of the social behaviors. Focus will be given in this thesis at the role played by di_erent recognition mechanism in paving the way to altruism (Komdeur and Hatchwell, 1999) when the population has a spatial structure. Further, the evolution of spite will also be considered in these settings. The thesis is fractionated into two parts. First, di_erent models promoting altruism cooperation and spite will be compared under the same theoretical umbrella. This is a rather informal and more personnal part of my thesis. It also serve as a justification and basis to "Altruism among kin and non-kin individuals" which is an article attempting to clas- sify the mechanisms leading to altruism and cooperation. Second, in the annexe, there are three research papers about kin selection, altruism and dispersal: "Is sociality driven by the costs of dispersal or the benefits of philopatry?: A role for kin-discrimination mechanism", "Altruism, dispersal and phenotype kin recognition" and "Inbreeding avoidance through kin recognition: choosy female boost male dispersal" this last paper incorporates kin recognition as an agent favoring sex-biased dispersal.

Evolutionary Games in Networked Populations: Models and Experiments

Cooperation and coordination are desirable behaviors that are fundamental for the harmonious development of society. People need to rely on cooperation with other individuals in many aspects of everyday life, such as teamwork and economic exchange in anonymous markets. However, cooperation may easily fall prey to exploitation by selfish individuals who only care about short- term gain. For cooperation to evolve, specific conditions and mechanisms are required, such as kinship, direct and indirect reciprocity through repeated interactions, or external interventions such as punishment. In this dissertation we investigate the effect of the network structure of the population on the evolution of cooperation and coordination. We consider several kinds of static and dynamical network topologies, such as Baraba´si-Albert, social network models and spatial networks. We perform numerical simulations and laboratory experiments using the Prisoner's Dilemma and co- ordination games in order to contrast human behavior with theoretical results. We show by numerical simulations that even a moderate amount of random noise on the Baraba´si-Albert scale-free network links causes a significant loss of cooperation, to the point that cooperation almost vanishes altogether in the Prisoner's Dilemma when the noise rate is high enough. Moreover, when we consider fixed social-like networks we find that current models of social networks may allow cooperation to emerge and to be robust at least as much as in scale-free networks. In the framework of spatial networks, we investigate whether cooperation can evolve and be stable when agents move randomly or performing Le´vy flights in a continuous space. We also consider discrete space adopting purposeful mobility and binary birth-death process to dis- cover emergent cooperative patterns. The fundamental result is that cooperation may be enhanced when this migration is opportunistic or even when agents follow very simple heuristics. In the experimental laboratory, we investigate the issue of social coordination between indi- viduals located on networks of contacts. In contrast to simulations, we find that human players dynamics do not converge to the efficient outcome more often in a social-like network than in a random network. In another experiment, we study the behavior of people who play a pure co- ordination game in a spatial environment in which they can move around and when changing convention is costly. We find that each convention forms homogeneous clusters and is adopted by approximately half of the individuals. When we provide them with global information, i.e., the number of subjects currently adopting one of the conventions, global consensus is reached in most, but not all, cases. Our results allow us to extract the heuristics used by the participants and to build a numerical simulation model that agrees very well with the experiments. Our findings have important implications for policymakers intending to promote specific, desired behaviors in a mobile population. Furthermore, we carry out an experiment with human subjects playing the Prisoner's Dilemma game in a diluted grid where people are able to move around. In contrast to previous results on purposeful rewiring in relational networks, we find no noticeable effect of mobility in space on the level of cooperation. Clusters of cooperators form momentarily but in a few rounds they dissolve as cooperators at the boundaries stop tolerating being cheated upon. Our results highlight the difficulties that mobile agents have to establish a cooperative environment in a spatial setting without a device such as reputation or the possibility of retaliation. i.e. punishment. Finally, we test experimentally the evolution of cooperation in social networks taking into ac- count a setting where we allow people to make or break links at their will. In this work we give particular attention to whether information on an individual's actions is freely available to poten- tial partners or not. Studying the role of information is relevant as information on other people's actions is often not available for free: a recruiting firm may need to call a job candidate's refer- ences, a bank may need to find out about the credit history of a new client, etc. We find that people cooperate almost fully when information on their actions is freely available to their potential part- ners. Cooperation is less likely, however, if people have to pay about half of what they gain from cooperating with a cooperator. Cooperation declines even further if people have to pay a cost that is almost equivalent to the gain from cooperating with a cooperator. Thus, costly information on potential neighbors' actions can undermine the incentive to cooperate in dynamical networks.

Plant Mating Systems: Female Sterility in the Driver's Seat.

Violation of Mendel's Law of Segregation by selfish X chromosomes that favour their own transmission is known for a number of organisms. Now, a new study reveals sex-ratio distortion favouring males and explains previously puzzling sex ratios in a Mediterranean shrub.

The hpx genetic system for hypoxanthine assimilation as a nitrogen source in Klebsiella pneumoniae: gene organization and transcriptional regulation.

Growth experiments showed that adenine and hypoxanthine can be used as nitrogen sources by several strains of K. pneumoniae under aerobic conditions. The assimilation of all nitrogens from these purines indicates that the catabolic pathway is complete and proceeds past allantoin. Here we identify the genetic system responsible for the oxidation of hypoxanthine to allantoin in K. pneumoniae. The hpx cluster consists of seven genes, for which an organization in four transcriptional units, hpxDE, hpxR, hpxO and hpxPQT, is proposed. The proteins involved in the oxidation of hypoxanthine (HpxDE) or uric acid (HpxO) did not display any similarity to other reported enzymes known to catalyze these reactions, but instead are similar to oxygenases acting on aromatic compounds. Expression of the hpx system is activated by nitrogen limitation and by the presence of specific substrates, with hpxDE and hpxPQT controlled by both signals. Nitrogen control of hpxPQT transcription, which depends on 54, is mediated by the Ntr system. In contrast, neither NtrC nor NAC is involved in the nitrogen control of hpxDE, which is dependent on 70 for transcription. Activation of these operons by the specific substrates is also mediated by different effectors and regulatory proteins. Induction of hpxPQT requires uric acid formation, whereas expression of hpxDE is induced by the presence of hypoxanthine through the regulatory protein HpxR. This LysR-type regulator binds to a TCTGC-N4-GCAAA site in the intergenic hpxD-hpxR region. When bound to this site for hpxDE activation, HpxR negatively controls its own transcription.

Stochasticity in economic losses increases the value of reputation in indirect reciprocity.

Recent theory predicts harsh and stochastic conditions to generally promote the evolution of cooperation. Here, we test experimentally whether stochasticity in economic losses also affects the value of reputation in indirect reciprocity, a type of cooperation that is very typical for humans. We used a repeated helping game with observers. One subject (the "Unlucky") lost some money, another one (the "Passer-by") could reduce this loss by accepting a cost to herself, thereby building up a reputation that could be used by others in later interactions. The losses were either stable or stochastic, but the average loss over time and the average efficiency gains of helping were kept constant in both treatments. We found that players with a reputation of being generous were generally more likely to receive help by others, such that investing into a good reputation generated long-term benefits that compensated for the immediate costs of helping. Helping frequencies were similar in both treatments, but players with a reputation to be selfish lost more resources under stochastic conditions. Hence, returns on investment were steeper when losses varied than when they did not. We conclude that this type of stochasticity increases the value of reputation in indirect reciprocity.

Linking brain structure and activation in the temporoparietal junction to explain the neurobiology of human altruism

Human altruism shaped our evolutionary history and pervades social and political life. There are, however, enormous individual differences in altruism. Some people are almost completely selfish, while others display strong altruism, and the factors behind this heterogeneity are only poorly understood. We examine the neuroanatomical basis of these differences with voxel-based morphometry and show that gray matter (GM) volume in the right temporoparietal junction (TPJ) is strongly associated with both individuals' altruism and the individual-specific conditions under which this brain region is recruited during altruistic decision making. Thus, individual differences in GM volume in TPJ not only translate into individual differences in the general propensity to behave altruistically, but they also create a link between brain structure and brain function by indicating the conditions under which individuals are likely to recruit this region when they face a conflict between altruistic and selfish acts.

A Forwarding Cooperation Protocol for Plain and Cluster-based Ad Hoc Networks

Peer-reviewed

A Fair and Secure Cluster Formation Process for Ad Hoc Networks

An efficient approach for organizing large ad hoc networks is to divide the nodesinto multiple clusters and designate, for each cluster, a clusterhead which is responsible forholding intercluster control information. The role of a clusterhead entails rights and duties.On the one hand, it has a dominant position in front of the others because it manages theconnectivity and has access to other node¿s sensitive information. But on the other hand, theclusterhead role also has some associated costs. Hence, in order to prevent malicious nodesfrom taking control of the group in a fraudulent way and avoid selfish attacks from suitablenodes, the clusterhead needs to be elected in a secure way. In this paper we present a novelsolution that guarantees the clusterhead is elected in a cheat-proof manner.

Mathematics inspired by Darwin. Adaptive dynamics of dispersal and cooperation

In 1859, Charles Darwin published his theory of evolution by natural selection, the process occurring based on fitness benefits and fitness costs at the individual level. Traditionally, evolution has been investigated by biologists, but it has induced mathematical approaches, too. For example, adaptive dynamics has proven to be a very applicable framework to the purpose. Its core concept is the invasion fitness, the sign of which tells whether a mutant phenotype can invade the prevalent phenotype. In this thesis, four real-world applications on evolutionary questions are provided. Inspiration for the first two studies arose from a cold-adapted species, American pika. First, it is studied how the global climate change may affect the evolution of dispersal and viability of pika metapopulations. Based on the results gained here, it is shown that the evolution of dispersal can result in extinction and indeed, evolution of dispersalshould be incorporated into the viability analysis of species living in fragmented habitats. The second study is focused on the evolution of densitydependent dispersal in metapopulations with small habitat patches. It resulted a very surprising unintuitive evolutionary phenomenon, how a non-monotone density-dependent dispersal may evolve. Cooperation is surprisingly common in many levels of life, despite of its obvious vulnerability to selfish cheating. This motivated two applications. First, it is shown that density-dependent cooperative investment can evolve to have a qualitatively different, monotone or non-monotone, form depending on modelling details. The last study investigates the evolution of investing into two public-goods resources. The results suggest one general path by which labour division can arise via evolutionary branching. In addition to applications, two novel methodological derivations of fitness measures in structured metapopulations are given.

Conflicting genomes, the demic theory & biodiversity

Organismic-centered Darwinism, in order to use direct phenotypes to measure natural selection's effect, necessitates genome's harmony and uniform coherence plus large population sizes. However, modern gene-centered Darwinism has found new interpretations to data that speak of genomic incoherence and disharmony. As a result of these two conflicting positions a conceptual crisis in Biology has arisen. My position is that the presence of small, even pocket-size, demes is instrumental in generating divergence and phenotypic crisis. Moreover, the presence of parasitic genomes as in acanthocephalan worms, which even manipulate suicidal behavior in their hosts; segregation distorters that change meiosis and Mendelian ratios; selfish genes and selfish whole chromosomes, such as the case of B-chromosomes in grasshoppers; P-elements in Drosophila; driving Y-chromosomes that manipulate sex ratios making males more frequent, as in Hamilton's X-linked drive; male strategists and outlaw genes, are eloquent examples of the presence of real conflicting genomes and of a non-uniform phenotypic coherence and genome harmony. Thus, we are proposing that overall incoherence and disharmony generate disorder but also more biodiversity and creativeness. Finally, if genes can manipulate natural selection, they can multiply mutations or undesirable characteristics and even lethal or detrimental ones, hence the accumulation of genetic loads. Outlaw genes can change what is adaptively convenient even in the direction of the trait that is away from the optimum. The optimum can be "negotiated" among the variants, not only because pleiotropic effects demand it, but also, in some cases, because selfish, outlaw, P-elements or extended phenotypic manipulation require it. With organismic Darwinism the genome in the population and in the individual was thought to act harmoniously without conflicts, and genotypes were thought to march towards greater adaptability. Modern Darwinism has a gene-centered vision in which genes, as natural selection's objects can move in dissonance in the direction which benefits their multiplication. Thus, we have greater opportunities for genomes in permanent conflict.