Colette Rivault

Social Integration of Robots into Groups of Cockroaches to Control Self-Organized Choices

Collective behavior based on self-organization has been shown in group-living animals from insects to vertebrates. These findings have stimulated engineers to investigate approaches for the coordination of autonomous multirobot systems based on self-organization. In this experimental study, we show collective decision-making by mixed groups of cockroaches and socially integrated autonomous robots, leading to shared shelter selection. Individuals, natural or artificial, are perceived as equivalent, and the collective decision emerges from nonlinear feedbacks based on local interactions. Even when in the minority, robots can modulate the collective decision-making process and produce a global pattern not observed in their absence. These results demonstrate the possibility of using intelligent autonomous devices to study and control self-organized behavioral patterns in group-living animals.

Self-organized aggregation in cockroaches

Aggregation is widespread in invertebrate societies and can appear in response to environmental heterogeneities or by attraction between individuals. We performed experiments with cockroach, Blattella germanica, larvae in a homogeneous environment to investigate the influence of interactions between individuals on aggregations. Different densities were tested. A first phase led to radial dispersion of larvae in relation to wall-following behaviours; the consequence of this process was a homogeneous distribution of larvae around the periphery of the arena. A second phase corresponded to angular reorganization of larvae leading to the formation of aggregates. The phenomenon was analysed both at the individual and collective levels. Individual cockroaches modulated their behaviour depending on the presence of other larvae in their vicinity: probabilities of stopping and resting times were both higher when the numbers of larvae were greater. We then developed an agent-based model implementing individual behavioural rules, all derived from experiments, to explain the aggregation dynamics at the collective level. This study supports evidence that aggregation relies on mechanisms of amplification, supported by interactions between individuals that follow simple rules based on local information and without knowledge of the global structure.

Cockroach aggregation based on strain odour recognition

Blattella germanica (L.) cockroaches are gregarious during their resting period. Binary choice tests with groups of larvae indicated a strong tendency to aggregate on a single resting site. This was observed even when all sites were identical or when larvae came from either one or two strains. Previous results showed that gregarious behaviour is mainly based on recognition of cuticular hydrocarbons characterizing strain odour and that larvae prefer their own strain odour to that of another strain. Nevertheless, when groups in tests came from two different strains and had the choice between their two strain odours, they aggregated on only one of the sites. A mathematical model relying on a minimum number of functioning rules was devised to reproduce these experimental collective responses. The model is based on a fundamental parameter representing individual variation in the resting period on a given site in relation to the number of individuals on that site. Taking only this parameter into account, the model predicted that different strains are able to aggregate on the same site despite a weak interstrain interattraction parameter. Blattella germanica is thus an interesting biological model to investigate different aspects of aggregation and interindividual recognition.

Cockroaches as carriers of bacteria in multi-family dwellings

The potential risk of bacterial dissemination due to the presence of cockroaches (Blattella germanica, Blattellidae) in low-income flats was investigated. Cockroaches can carry a great variety of bacterial species; we identified 30 different species from 52 different flats. Klebsiella oxycytoca, K. pneumoniae and Enterobacter cloacae were the most frequently found. Pathogenic and potentially pathogenic bacteria represented 54% of all the bacterial identifications. Bacteria were carried either on the cuticle or in the gut. Contamination through external contact is sufficient to insure bacterial diffusion. There was a very low level of overlap estimated by Piankas index (a) between the bacterial flora of neighbouring blocks of flats, and (b) between bacterial flora of different flats in the same block.

Cuticular extracts inducing aggregation in the German cockroach, Blattella germanica (L.).

German cockroaches Blattella germanica (L.) are gregarious insects. An aggregation pheromone contributes to the maintenance of aggregates. Choice experiments checked the efficiency of different solvents, i.e. dichloromethane, methanol and pentane, in extracting attractive substances and compared the attractiveness of extracts of different parts of the body. Dichloromethane and pentane were the most efficient solvents tested for extracting the attractive substances. Methanol whole body extracts appeared inefficient to induce aggregation. The proportion of larvae attracted to conditioned papers decreased in relation to the size of cuticular surface washed, from whole body to half-body and again to a section of the body cut in three. Attractive substances appear to be present on all parts of the body. Chemical analysis by gas chromatography (GC) and gas chromatography-mass spectrometry (GC-MS) showed that the active extracts contained only cuticular hydrocarbons. In addition to behavioural tests, differences between the composition of methanol extracts and that of the extracts for the other two solvents were revealed by GC. These results indicated that the cuticular hydrocarbons operate as an aggregation pheromone in Blattella germanica.

Mutual Mate Choice: When it Pays Both Sexes to Avoid Inbreeding

Theoretical models of sexual selection predict that both males and females of many species should benefit by selecting their mating partners. However, empirical evidence testing and validating this prediction is scarce. In particular, whereas inbreeding avoidance is expected to induce sexual conflicts, in some cases both partners could benefit by acting in concert and exerting mutual mate choice for non-assortative pairings. We tested this prediction with the gregarious cockroach Blattella germanica (L.). We demonstrated that males and females base their mate choice on different criteria and that choice occurs at different steps during the mating sequence. Males assess their relatedness to females through antennal contacts before deciding to court preferentially non-siblings. Conversely, females biased their choice towards the most vigorously courting males that happened to be non-siblings. This study is the first to demonstrate mutual mate choice leading to close inbreeding avoidance. The fact that outbred pairs were more fertile than inbred pairs strongly supports the adaptive value of this mating system, which includes no “best phenotype” as the quality of two mating partners is primarily linked to their relatedness. We discuss the implications of our results in the light of inbreeding conflict models.

Spatial distribution of the cockroach, Blattella germanica, in a swimming-bath facility

A population of Blattella germanica L. was studied in a swimming‐bath facility using a sampling method. A set of traps deposited weekly, over a period of 50 weeks, allowed us to follow the variations of the temporal and spatial distributions of the population inside the building. A Factorial Correspondance Analysis, performed on the data, led us to the conclusion that the population was stable in time. The spatial distribution of the insects was contiguous. Inside an aggregate, we were able to define a circular movement from the center towards the periphery and vice versa; this movement can be superimposed up on the biological development on the individuals. After birth in the middle of the aggregate, during development, larvae explore further a field as far as the border of the aggregate; this exploratory phase, related to foraging behavior or search for new shelters when these become limiting factors, could be considered as the spreading phase of the population. Then, with the imminence of the imaginai moult, old larvae tend to gather in high density areas in the middle of the aggregate. Newly molted adults look for a sexual partner. And finally females remain in the shelters during oothecae maturation.

Bacterial Load of Cockroaches in Relation to Urban-Environment

Sanitation is an important problem in relation to the control of pests in urban environments. This investigation analysed the potential risk related to the presence of cockroaches and their capacity for disseminating bacteria in six different types of buildings: hospital nursing area and out-patient area, swimming-pool pool-side and toilet area, low-income flats and food-handling places. Fifty-six species of bacteria were identified from 157 samples, 14 of these have previously been reported as potentially pathogenic for man and vertebrates. Similarities were found between samples collected in (a) the hospital out-patient area and food-handling establishments and (b) the hospital nursing area and flats. Pool-sides possessed a poorer bacterial flora. There was a greater bacterial specific diversity in food-handling establishments, flats and swimming-bath toilet area. Enterobacter cloacae. Klebsiella pneumoniae and Klebsiella oxytoca were dominant species in flats and the hospital nursing area. Therefore, cockroaches can play a role in disseminating bacteria, which they can carry passively on their cuticle.

The weight of the clan: even in insects, social isolation can induce a behavioural syndrome.

Social isolation has dramatic consequences on the development of individuals of many vertebrate species, and it induces a set of behavioural disturbances rending them unable to process environmental as well as social stimuli appropriately. We hypothesized that isolation syndrome is a ubiquitous trait of social life that can be observed in a wide array of species, including invertebrates. Here we report that gregarious cockroaches (Blattella germanica) reared in isolation showed (i) stronger exploration-avoidance, (ii) reduced foraging activity, (iii) reduced willingness to interact socially, and (iv) reduced ability to assess mating partner quality than conspecifics reared in groups. We demonstrate the occurrence of a behavioural syndrome induced by social isolation, similar to syndromes described in vertebrates, revealing the importance of social interactions and group-living in this non-eusocial insect species. We suggest that investigating social isolation effects on individual development should provide interesting results to assess social cohesion of species and thus constitute an additional tool for comparative studies focusing on the evolution of social life.

Collective foraging decision in a gregarious insect

Group foraging by eusocial insects implies sophisticated recruitment processes that often result in collective decisions to exploit the most profitable sources. These advanced levels of cooperation, however, remain limited to a small range of species, and we still know little about the mechanisms underlying group foraging behaviours in the great mass of animals exhibiting lower levels of social complexity. In this paper, we report, for the first time in a gregarious insect, the cockroach Blattella germanica (L.), a collective foraging decision whereby the selection of food sources is reached without requiring active recruitment. Groups of cockroaches given a binary choice between identical food sources exhibited exploitation asymmetries whose amplitude increases with group size. By coupling behavioural observations to computer simulations, we demonstrate that selection of food sources relies uniquely on a retention effect of feeding individuals on newcomers without comparison between available opportunities. This self-organised pattern presents similarities with the foraging dynamics of eusocial species, thus stressing the generic dimension of collective decision-making mechanisms based on social amplification rules despite fundamental differences in recruitment processes. We hypothesise that such parsimony could apply to a wide range of species and help understand the emergence of collective behaviours in simple social systems.