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Bshary, Redouan
Nom
Bshary, Redouan
Affiliation principale
Fonction
Professeur ordinaire
Email
redouan.bshary@unine.ch
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Résultat de la recherche
Voici les éléments 1 - 10 sur 11
- PublicationAccès libreWhat makes a cleaner a cleaner ?(2016)
; In his last presidential address to the Royal Society in 2005, Robert M. May stated that “The most important unanswered question in evolutionary biology, and more generally in the social sciences, is how cooperative behaviour evolved and can be maintained”. My thesis provides a contribution to answering this big question by investigating how one particular species evolved for an ecology that rests heavily upon cooperative interactions: the bluestreak cleaner wrasse Labroides dimidiatus. This species engages in up to 2000 cooperative interactions per day with dozens of other ‘client’ coral reef fishes, thus making it a prime system for studying cooperation between unrelated individuals. Conflicts between cleaners and clients arise because the former prefer to bite clients to eat their protective mucus rather than focusing on ectoparasites. In response to such exploitation, clients use various forms of partner control mechanisms that promote cooperative behaviour in cleaners. As a result, cleaners are known to use a diversity of strategic behaviours to determine when to cheat and when to cooperate. Cleaners’ behaviour thus appears very well adapted to the demands of their peculiar ecology.
In my thesis, I used a comparative approach to investigate which traits appear to be associated with the highly social ecology of cleaners, or in other words: What makes a cleaner a cleaner? Through a series of experiments, I compared the cognitive skills, escape performance, foraging ecology and vision of L. dimidiatus with other species of labrids that do not engage in cleaning (or only occasionally). These comparisons allowed me to identify some of the characteristics that set cleaners apart, and thus further our understanding of how social ecology can affect the evolution of a species.
In the first chapter, I showed that L. dimidiatus was able to fine-tune its level of cooperation to the specifics of different cleaning-related situations, an ability that was absent in the closely related Halichoeres melanurus. In the second chapter, I investigated whether the demands of a highly social lifestyle led to an overall increase of cognitive performance in L. dimidiatus, or whether performance was tightly linked to ecological demands. L. dimidiatus outperformed five other species of labrids in two ecologically relevant tasks. However, all species performed similarly in a task with little ecological relevance, suggesting that cognition in cleaners is tightly linked to the challenges faced in nature. In the third chapter, results from foraging experiments suggest that L. dimidiatus evolved a foraging position that allows for increased efficiency in cleaning interactions. In comparison with four other species, L. dimidiatus adopted a significantly lower body angle with regards to the substrate when foraging. Furthermore, this species experienced almost no reduction in efficiency when searching for cryptic food items in comparison with conspicuous ones, while the other species all performed worse in the cryptic condition. It thus appears that the peculiar foraging posture of L. dimidiatus is well suited for cleaning interactions. Unfortunately, my current data did not allow assessing whether the visual system also adapted for a low body angle while foraging. Finally, in the last chapter, I asked whether because of the service they provide to predators, a reduction in predation pressures led to the decay of escape performance in L. dimidiatus. Interestingly, measures of escape performance in a controlled laboratory setup showed that L. dimidiatus was among the top performers in comparison with 5 other labrids. These results suggest that the risks associated with cleaning interactions are sufficient to maintain a high escape performance in cleaners, despite their privileged relationship with predatory clients.
In summary, a wide range of characteristics appear to be important for cleaning interactions, and species that specialized in this activity seem to have undergone very different selective pressures than fishes with more standard ecologies. Identifying some of the key aspects related to the ecology of cleaners provides a good example of how the evolution of a species can be affected by the demands of a highly social life. In this system, I argue that competition among service providers and conflicts of interests between cleaners and clients appear to be the major drivers of adaptation. - PublicationAccès libreAdjustments of levels of cooperation in cleaner wrasse "Labroides dimidiatus": the effects of an audience and satiation(2015)
Mon système modèle a consisté d’un mutualisme de nettoyage marin impliquant des labres nettoyeurs (Labroides dimidiatus) et ses poissons de récif dit «clients» qui leur rendent visite pour se faire enlever les ectoparasites. Cependant, des conflits surgissent car les nettoyeurs préfèrent la couche de mucus protectrice des clients aux ectoparasites, où se nourrir de mucus est préjudiciable au client et est donc fonctionnellement considéré comme de la tricherie. Par conséquent, les clients doivent faire en sorte que les nettoyeurs mangent contre leur préférence afin de pouvoir bénéficier d'un bon service de nettoyage. Des observations sur le terrain et une expérience de laboratoire utilisant des plaques en plexiglas en remplacement de clients avaient déjà suggéré que les clients spectateurs étaient attentifs à la manière dont un nettoyeur traite son client actuel et que les nettoyeurs sont plus coopératifs s’ils sont observés. J'ai pu démontrer ce concept de prestige social pour la première fois dans le cadre d'une expérience de laboratoire contrôlée utilisant de véritables interactions client-nettoyeur. Dans deux autres expériences utilisant soit des plaques en plexiglas soit de vrais clients, j'ai montré que les nettoyeurs peuvent ajuster la qualité de leurs services à l'importance relative du client actuel par rapport au client spectateur: plus le spectateur a de la valeur, plus le service envers le client actuel est peaufiné. Dans une troisième expérience, j’ai manipulé le niveau de satiété des nettoyeurs afin de tester la prédiction de la théorie du marché biologique selon laquelle un besoin d’interactions temporairement faible entraîne une baisse de la qualité du service. De manière quelque peu surprenante, cette prédiction n’a pas été confirmée, les nettoyeurs rassassiés ayant augmenté leur niveau de coopération envers leurs clients, c’est-à-dire qu’ils ont moins triché lors des interactions avec leurs clients et se sont nourris davantage contre leur préférence sur les plaques en plexiglas. Ainsi, les nettoyeurs rassassiés investissent fonctionnellement dans leur relation avec les clients pour des avantages futurs.
En conclusion, mon travail de recherche démontre que le labre nettoyeur L. dimidiatus, est capable de prendre des décisions sophistiquées, adaptées aux spécificités de la situation. Les résultats soulèvent des questions concernant les processus cognitifs sous-jacents, car ils remettent en cause la notion selon laquelle des cerveaux plus larges sont nécessaires pour une coopération sophistiquée. Au lieu de cela, il semble qu'une approche écologique par rapport à la cognition soit plus appropriée pour expliquer mes résultats. Mon étude s'inscrit dans une longue tradition selon laquelle les poissons constituent des systèmes modèles idéaux pour tester la théorie de la coopération, dont les résultats prometteurs devraient inspirer de nouvelles analyses théoriques., Cooperation, the mutual helping between unrelated individuals, is an evolutionary puzzle. This is because helping is often an investment that must yield future benefits in order to be under positive selection. Generally, natural selection favours individuals that perform self-serving behaviour, and hence cheating is a major conceptual issue. However, examples of cooperation are abundant in nature. As such, finding explanations that reconcile cooperation with evolutionary theory has long been a major focus in biology but also in the social sciences with their aim to explain the social complexity in humans. A variety of so-called partner control mechanisms – behavioural responses that cause a reduction in the payoffs of a cheating partner such that a cooperating partner gains more – have been shown to stabilise cooperation. My research has focused in particular on the role of social prestige in a communication network. Many animal interactions are observed by third parties (“bystanders”), who may gain valuable information about the interactants. In the context of cooperation, bystanders should try to identify particularly cooperative individuals as future partners, which selects for individuals being particularly cooperative if they are observed.
My model system has been marine cleaning mutualism involving bluestreak cleaner wrasses (Labroides dimidiatus) and their so-called “client” reef fishes that visit to have ectoparasites removed. However, conflict arises as cleaners prefer the protective mucus layer of clients over ectoparasites, where mucus feeding is detrimental to the client and hence functionally constitutes cheating. Thus, clients have to make cleaners feed against their preference in order to receive a good cleaning service. Field observations and a laboratory experiment using Plexiglas plates as client surrogates had already suggested that bystander clients pay attention to how a cleaner treats its current client, and that cleaners are more cooperative if observed. I could demonstrate this social prestige concept for the first time in a controlled laboratory experiment in real cleaner-client interactions. In two further experiments using either plates or real clients I showed that cleaners can fine-tune their service quality to the relative importance of current client versus bystander: the more valuable the bystander the better the current service. Finally, I manipulated the cleaners’ level of satiation in order to test the prediction from biological market theory that a temporarily low need for interactions causes a decrease in service quality. Somewhat surprisingly, this prediction was not met as satiated cleaners increased their cooperation levels towards their clients, i.e. they caused less jolts during interactions with their clients and fed more against their preference on plates. Thus, satiated cleaners functionally invest into their relationship with clients for future benefits.
In conclusion, my work shows that cleaner wrasse L. dimidiatus show sophisticated decision rules that are fine-tuned to the specifics of the situation. The results raise questions concerning the underlying cognitive processes as they challenge the notion that large brains are necessary for sophisticated cooperation. Instead, it appears that an ecological approach to cognition is better suited to explain my results. My study fits into a long tradition that fish yield ideal model systems to test cooperation theory, where results hopefully inspire further theoretical analyses. - PublicationAccès libreSignalling by the cleaner shrimp Periclimenes longicarpus(2010)
- PublicationAccès libreThe effect of nectar reduction in Petunia Axillaris on foraging behavior of noctural hawkmoths, observed in laboratory and field behavioral assays(2009)
; Kuhlemeier, CrisA key component shaping plant-pollinator interactions is nectar. Its volume can regulate the length and frequency of pollination events. Nectar provisioning can be costly for the plant. Once secreted by the nectaries, the sugar-rich solution is usually consumed by a floral visitor and lost for “recycling” within the plant. Nectar reduction should thus be advantageous for the plant: non-secreted carbohydrates can be reallocated within the plant to other structures promoting growth, reproduction or attraction. However, most angiosperms provide nectar. It is assumed that certain pollinator behaviors, so called partner control mechanisms, favor nectariferous over deceptive plants and ultimately prevent the spread of “cheaters”. The partner control mechanisms identified in the context of plant-pollinator mutualisms are avoidance of nectarless species, reduction of drinking time and number of flowers visited on nectarless plants. Not all behaviors are performed simultaneously, and external conditions such as plant density as well as intrinsic factors of the foraging insect can determine to which extent certain behaviors are exerted. In the present study, we analyze foraging behavior of nocturnal hawkmoths on cheating Petunia axillaris axillaris plants under several conditions. The aim of this thesis was to assess which partner control mechanisms are executed by pollinators facing nectarless/low nectar plants. We observed hawkmoth behavior in two field sites and conducted experiments with naïve and experienced hawkmoths Manduca sexta under controlled conditions. We investigated which of the foraging decision rules might potentially reduce the fitness of cheaters and thus limit their spread in a population. In field assays, we observed that the density of naturally occurring P. axillaris plants and the presence of alternative food sources can influence hawkmoth behavior on nectarless Petunias: only when food plants were abundant and dense, pollinators would reduce the number of flowers on nectarless Petunias, whereas in the lower density there seemed to be no selection against cheaters. In learning experiments under controlled conditions, we observed that none of the tested behaviors (reduction of drinking duration on nectarless plants, avoidance of nectarless plants, reduction of number of flowers visited on nectarless plants) were improved over the course of the experiment. However, in all learning trials there was a significant reduction of drinking duration on nectarless plants, indicating that this control mechanism of hawkmoths is always exerted innately. Learning might therefore not be of major importance in discrimination against cheaters in our system. We constructed a plant with extremely high phenotypic similarity to P. axillaris yet only a third of the regular nectar volume (F25). Genotyping of F25 revealed a high genotypic imilarity to its parental plant but failed to answer questions about the genetic background of low nectar volume. The low nectar line was used in behavioral experiments with Manduca sexta. A major goal was to find out how pollination behavior affects female reproductive success of F25. Analogous to previous experiments, we found that the drinking duration was significantly reduced on cheaters. In hand pollination assays, F25 produced significantly more seeds than P. axillaris, however this effect was neutralized when pollinated by Manduca sexta. The benefits of nectar reduction are thus counterbalanced by a change in pollinator foraging behavior. In the future, we would like to assess which other fitness parameters are concerned when a plant ceases its nectar production. Altogether, we were able to show which foraging rules are exerted by hawkmoths on cheating P. axillaris and how one partner control mechanism, namely drinking duration, affects seed set of a plant with reduced nectar offerings. We hope that this work has contributed to answering questions about the costs and benefits of cheating. - PublicationAccès libreStrategic adjustment of service quality to client identity in the cleaner shrimp, Periclimenes longicarpus(2009)
- PublicationAccès libreMutualism, market effects and partner control(2008)
- PublicationAccès librePunishment and partner switching cause cooperative behaviour in a cleaning mutualism(2005)
; Grutter, Alexandra S.What are the mechanisms that prevent partners from cheating in potentially cooperative interactions between unrelated individuals? The cleaner fish Labroides dimidiatus and client reef fish both benefit from an interaction as long as the cleaner eats ectoparasites. However, the cleaner fish prefers some client mucus, which constitutes cheating. Field observations suggested that clients control such cheating by using punishment (chasing the cleaner) or by switching partners (fleeing from the cleaner). Here, we tested experimentally whether such client behaviours result in cooperative cleaner fish. Cleaners were allowed to feed from Plexiglas plates containing prawn items and fish flake items. A lever attached to the plates allowed us to mimic the behaviours of clients. As cleaners showed a strong preference for prawn over flakes, we taught them that eating their preferred food would cause the plate to either chase them or to flee, while feeding on flakes had no negative consequences. We found a significant shift in cleaner fish foraging behaviour towards flake feeding after six learning trials. As punishment and terminating an interaction resulted in the cleaners feeding against their preferences in our experiment, we propose that the same behaviours in clients improve the service quality of cleaners under natural conditions. - PublicationMétadonnées seulementFrom parasitism to mutualism: partner control in asymmetric interactions(2002)
- PublicationMétadonnées seulementCleaner fish Labroides dimidiatus manipulate client reef fish by providing tactile stimulation(2001)
; Wurth, ManuelaThe cleaner wrasse Labroides dimidiatus often touches 'client' reef fish dorsal fin areas with its pelvic and pectoral fins. The relative spatial positions of cleaner and client remain constant and the cleaner's head points away from the client's body. Therefore, this behaviour is not compatible with foraging and the removal of client ectoparasites. As clients seek such 'tactile stimulation', it can be classified as an interspecific socio-positive behaviour. Our field observations on 12 cleaners (observation time of 112 h) suggest that cleaners use tactile stimulation in order to successfully (i) alter client decisions over how long to stay for an inspection, and (ii) stop clients from fleeing or aggressive chasing of the cleaner in response to a cleaner fish bite that made them jolt. Finally predatory clients receive tactile stimulation more often than non-predatory clients, which might be interpreted as an extra service that cleaners give to specific partners as pre-conflict management, as these partners would be particularly dangerous if they started a conflict. We therefore propose that cleaner fish use interspecific social strategies, which have so far been reported only from mammals, particularly primates. - PublicationAccès libreBacterial farming by the fungus Morchella crassipes(The Royal Society of London, )
; ; ; ; The interactions between bacteria and fungi, the main actors of the soil microbiome, remain poorly studied. Here, we show that the saprotrophic and ectomycorrhizal soil fungus Morchella crassipes acts as a bacterial farmer of Pseudomonas putida, which serves as a model soil bacterium. Farming by M. crassipes consists of bacterial dispersal, bacterial rearing with fungal exudates, as well as harvesting and translocation of bacterial carbon. The different phases were confirmed experimentally using cell counting and 13C probing. Common criteria met by other non-human farming systems are also valid for M. crassipes farming, including habitual planting, cultivation and harvesting. Specific traits include delocalization of food production and consumption and separation of roles in the colony (source versus sink areas), which are also found in human agriculture. Our study evidences a hitherto unknown mutualistic association in which bacteria gain through dispersal and rearing, while the fungus gains through the harvesting of an additional carbon source and increased stress resistance of the mycelium. This type of interaction between fungi and bacteria may play a key role in soils.