3D cell segregation geometry and dynamics are governed by tissue surface tension regulation.

Tissue morphogenesis and patterning during development involve the segregation of cell types. Segregation is driven by differential tissue surface tensions generated by cell types through controlling cell-cell contact formation by regulating adhesion and actomyosin contractility-based cellular cortical tensions. We use vertebrate tissue cell types and zebrafish germ layer progenitors as in vitro models of 3-dimensional heterotypic segregation and developed a quantitative analysis of their dynamics based on 3D time-lapse microscopy. We show that general inhibition of actomyosin contractility by the Rho kinase inhibitor Y27632 delays segregation. Cell type-specific inhibition of non-muscle myosin2 activity by overexpression of myosin assembly inhibitor S100A4 reduces tissue surface tension, manifested in decreased compaction during aggregation and inverted geometry observed during segregation. The same is observed when we express a constitutively active Rho kinase isoform to ubiquitously keep actomyosin contractility high at cell-cell and cell-medium interfaces and thus overriding the interface-specific regulation of cortical tensions. Tissue surface tension regulation can become an effective tool in tissue engineering.

Synergistic Benefits of Group Search in Rats.

Locating unpredictable but essential resources is a task that all mobile animals have to perform in order to survive and reproduce. Research on search strategies has focused largely on independent individuals [1-3], but many organisms display collective behaviors, including during group search and foraging [4-6]. One classical experimental search task, informing studies of navigation, memory, and learning, is the location of a reward in a confined, complex maze setting [7, 8]. Rats (Rattus norvegicus) have been paradigmatic in psychological and biological studies [9, 10], but despite rats being highly social [11, 12], their group search behavior has not been investigated. Here, we explore the decision making of rats searching individually, or in groups, for a reward in a complex maze environment. Using automated video tracking, we find that rats exhibit-even when alone-a partially systematic search, leading to a continuous increase in their chance of finding the reward because of increased attraction to unexplored regions. When searching together, however, synergistic group advantages arise through integration of individual exploratory and social behavior. The superior search performances result from a strategy that represents a hierarchy of influential preferences in response to social and asocial cues. Furthermore, we present a computational model to compare the essential factors that influence how collective search operates and to validate that the collective search strategy increases the search efficiency of individuals in groups. This strategy can serve as direct inspiration for designing computational search algorithms and systems, such as autonomous robot groups, to explore areas inaccessible to humans. VIDEO ABSTRACT.

Adaptive leadership overcomes persistence-responsivity trade-off in flocking.

The living world is full of cohesive collectives that have evolved to move together with high efficiency. Schools of fish or flocks of birds maintain their global direction despite significant noise perturbing the individuals, yet they are capable of performing abrupt collective turns when relevant agitation alters the state of a few members. Ruling local fluctuations out of global movement leads to persistence and requires overdamped interaction dynamics, while propagating swift turns throughout the group leads to responsivity and requires underdamped interaction dynamics. In this paper we show a way to avoid this conflict by introducing a time-dependent leadership hierarchy that adapts locally to will: agents' intention of changing direction. Integrating our new concept of will-based inter-agent behaviour highly enhances the responsivity of standard collective motion models, thus enables breaking out of their former limit, the persistence-responsivity trade-off. We also show that the increased responsivity to environmental cues scales well with growing flock size. Our solution relies on active communication or advanced cognition for the perception of will. The incorporation of these into collective motion is a plausible hypothesis in higher order species, while it is a realizable feature for artificial robots, as demonstrated by our swarm of 52 drones.

Optimized flocking of autonomous drones in confined environments.

We address a fundamental issue of collective motion of aerial robots: how to ensure that large flocks of autonomous drones seamlessly navigate in confined spaces. The numerous existing flocking models are rarely tested on actual hardware because they typically neglect some crucial aspects of multirobot systems. Constrained motion and communication capabilities, delays, perturbations, or the presence of barriers should be modeled and treated explicitly because they have large effects on collective behavior during the cooperation of real agents. Handling these issues properly results in additional model complexity and a natural increase in the number of tunable parameters, which calls for appropriate optimization methods to be coupled tightly to model development. In this paper, we propose such a flocking model for real drones incorporating an evolutionary optimization framework with carefully chosen order parameters and fitness functions. We numerically demonstrated that the induced swarm behavior remained stable under realistic conditions for large flock sizes and notably for large velocities. We showed that coherent and realistic collective motion patterns persisted even around perturbing obstacles. Furthermore, we validated our model on real hardware, carrying out field experiments with a self-organized swarm of 30 drones. This is the largest of such aerial outdoor systems without central control reported to date exhibiting flocking with collective collision and object avoidance. The results confirmed the adequacy of our approach. Successfully controlling dozens of quadcopters will enable substantially more efficient task management in various contexts involving drones.

Glassy nature of hierarchical organizations.

The question of why and how animal and human groups form temporarily stable hierarchical organizations has long been a great challenge from the point of quantitative interpretations. The prevailing observation/consensus is that a hierarchical social or technological structure is optimal considering a variety of aspects. Here we introduce a simple quantitative interpretation of this situation using a statistical mechanics-type approach. We look for the optimum of the efficiency function [Formula: see text] with J ij denoting the nature of the interaction between the units i and j and a i standing for the ability of member i to contribute to the efficiency of the system. Notably, this expression for E eff has a similar structure to that of the energy as defined for spin-glasses. Unconventionally, we assume that J ij -s can have the values 0 (no interaction), +1 and -1; furthermore, a direction is associated with each edge. The essential and novel feature of our approach is that instead of optimizing the state of the nodes of a pre-defined network, we search for extrema for given a i -s in the complex efficiency landscape by finding locally optimal network topologies for a given number of edges of the subgraphs considered.

Friction forces position the neural anlage.

During embryonic development, mechanical forces are essential for cellular rearrangements driving tissue morphogenesis. Here, we show that in the early zebrafish embryo, friction forces are generated at the interface between anterior axial mesoderm (prechordal plate, ppl) progenitors migrating towards the animal pole and neurectoderm progenitors moving in the opposite direction towards the vegetal pole of the embryo. These friction forces lead to global rearrangement of cells within the neurectoderm and determine the position of the neural anlage. Using a combination of experiments and simulations, we show that this process depends on hydrodynamic coupling between neurectoderm and ppl as a result of E-cadherin-mediated adhesion between those tissues. Our data thus establish the emergence of friction forces at the interface between moving tissues as a critical force-generating process shaping the embryo.

A nationwide study of the epidemiology of relapsing polychondritis.

OBJECTIVE: Relapsing polychondritis (RP) is a rare autoimmune inflammatory disease that attacks mainly cartilaginous structures or causes serious damage in proteoglycan-rich structures (the eyes, heart, blood vessels, inner ear). This study shows results regarding the epidemiology, progression, and associations of this highly variable disease by collecting all cases from a 124-million-person-year Central European nationwide cohort. METHODS: We used the Hungarian Health Care Database to identify all persons with possible RP infection. We followed patients who had International Classification of Diseases 10th edition code M94.1 at least once in their inpatient or outpatient records between January 1, 2002 and December 31, 2013 in Hungary. We classified these patients into disease severity groups by their drug consumption patterns between January 1, 2010 and December 31, 2013. We analyzed the regional distribution of RP incidences as well. Overall maps of comorbidity are presented with network layouts. RESULTS: We identified 256 patients with RP among cumulatively 11.5 million registered inhabitants. We classified these patients into four severity classes as "extremely mild" (n=144), "mild" (n=22), "moderate" (n=41), and "severe" (n=4). Two additional groups were defined for patients without available drug data as "suspected only" (n=23) and "confirmed but unknown treatment" (n=22). The age and sex distributions of patients were similar to worldwide statistics. Indeed, the overall survival was good (95% confidence interval for 5 years was 83.6%-92.9% and for 10 years was 75.0%-88.3% which corresponds to the overall survival of the general population in Hungary), and the associations with other autoimmune disorders were high (56%) in Hungary. Almost any disease can occur with RP; however, the symptoms of chromosomal abnormalities are only incidental. Spondylosis can be a sign of the activation of RP, while Sjögren syndrome is the most frequent autoimmune association. Regional distribution of incidences suggests arsenic drinking water and sunlight exposure as possible triggering factors. CONCLUSION: The good survival rate of RP in Hungary is probably associated with the early diagnosis of the disease.

Speed Determines Leadership and Leadership Determines Learning during Pigeon Flocking.

A key question in collective behavior is how individual differences structure animal groups, affect the flow of information, and give some group members greater weight in decisions. Depending on what factors contribute to leadership, despotic decisions could either improve decision accuracy or interfere with swarm intelligence. The mechanisms behind leadership are therefore important for understanding its functional significance. In this study, we compared pigeons' relative influence over flock direction to their solo flight characteristics. A pigeon's degree of leadership was predicted by its ground speeds from earlier solo flights, but not by the straightness of its previous solo route. By testing the birds individually after a series of flock flights, we found that leaders had learned straighter homing routes than followers, as we would expect if followers attended less to the landscape and more to conspecifics. We repeated the experiment from three homing sites using multiple independent flocks and found individual consistency in leadership and speed. Our results suggest that the leadership hierarchies observed in previous studies could arise from differences in the birds' typical speeds. Rather than reflecting social preferences that optimize group decisions, leadership may be an inevitable consequence of heterogeneous flight characteristics within self-organized flocks. We also found that leaders learn faster and become better navigators, even if leadership is not initially due to navigational ability. The roles that individuals fall into during collective motion might therefore have far-reaching effects on how they learn about the environment and use social information.

Anomalous segregation dynamics of self-propelled particles.

A number of novel experimental and theoretical results have recently been obtained on active soft matter, demonstrating the various interesting universal and anomalous features of this kind of driven systems. Here we consider the adhesion difference-driven segregation of actively moving units, a fundamental but still poorly explored aspect of collective motility. In particular, we propose a model in which particles have a tendency to adhere through a mechanism which makes them both stay in touch and synchronize their direction of motion - but the interaction is limited to particles of the same kind. The calculations corresponding to the related differential equations can be made in parallel, thus a powerful GPU card allows large scale simulations. We find that in a very large system of particles, interacting without explicit alignment rule, three basic segregation regimes seem to exist as a function of time: i) at the beginning the time dependence of the correlation length is analogous to that predicted by the Cahn-Hillard theory, ii) next rapid segregation occurs characterized with a separation of the different kinds of units being faster than any previously suggested speed, finally, iii) the growth of the characteristic sizes in the system slows down due to a new regime in which self-confined, rotating, splitting and re-joining clusters appear. Our results can explain recent observations of segregating tissue cells in vitro.

HIV competition dynamics over sexual networks: first comer advantage conserves founder effects.

Outside Africa, the global phylogeography of HIV is characterized by compartmentalized local epidemics that are typically dominated by a single subtype, which indicates strong founder effects. We hypothesized that the competition of viral strains at the epidemic level may involve an advantage of the resident strain that was the first to colonize a population. Such an effect would slow down the invasion of new strains, and thus also the diversification of the epidemic. We developed a stochastic modelling framework to simulate HIV epidemics over dynamic contact networks. We simulated epidemics in which the second strain was introduced into a population where the first strain had established a steady-state epidemic, and assessed whether, and on what time scale, the second strain was able to spread in the population. Simulations were parameterized based on empirical data; we tested scenarios with varying levels of overall prevalence. The spread of the second strain occurred on a much slower time scale compared with the initial expansion of the first strain. With strains of equal transmission efficiency, the second strain was unable to invade on a time scale relevant for the history of the HIV pandemic. To become dominant over a time scale of decades, the second strain needed considerable (>25%) advantage in transmission efficiency over the resident strain. The inhibition effect was weaker if the second strain was introduced while the first strain was still in its growth phase. We also tested how possible mechanisms of interference (inhibition of superinfection, depletion of highly connected hubs in the network, one-time acute peak of infectiousness) contribute to the inhibition effect. Our simulations confirmed a strong first comer advantage in the competition dynamics of HIV at the population level, which may explain the global phylogeography of the virus and may influence the future evolution of the pandemic.

Collective foraging in heterogeneous landscapes.

Animals foraging alone are hypothesized to optimize the encounter rates with resources through Lévy walks. However, the issue of how the interactions between multiple foragers influence their search efficiency is still not completely understood. To address this, we consider a model to study the optimal strategy for a group of foragers searching for targets distributed heterogeneously. In our model, foragers move on a square lattice containing immobile but regenerative targets. At any instant, a forager is able to detect only those targets that happen to be in the same site. However, we allow the foragers to have information about the state of other foragers. A forager who has not detected any target walks towards the nearest location, where another forager has detected a target, with a probability exp(-αd), where d is the distance between the foragers and α is a parameter characterizing the propensity of the foragers to aggregate. The model reveals that neither overcrowding (α → 0) nor independent searching (α → ∞) is beneficial for the foragers. For a patchy distribution of targets, the efficiency is maximum for intermediate values of α. In addition, in the limit α → 0, the length of the walks can become scale-free.

Route-dependent switch between hierarchical and egalitarian strategies in pigeon flocks.

The mechanisms that underlie fascinating inter-individual interactions among animal groups have attracted increasing attention from biologists, physicists, and system scientists. There are two well-known types of interaction patterns: hierarchical and egalitarian. In the former type, individuals follow their leaders, whereas they follow their neighbors in the latter. Using high-resolution spatiotemporal data derived from the free flights of a flock of pigeons, we show that pigeon flocks actually adopt a mode that switches between the two aforementioned strategies. To determine its flight direction, each pigeon tends to follow the average of its neighbors while moving along a smooth trajectory, whereas it switches to follow its leaders when sudden turns or zigzags occur. By contrast, when deciding how fast to fly, each pigeon synthesizes the average velocity of its neighbors. This switching mechanism is promising for possible industrial applications in multi-robot system coordination, unmanned vehicle formation control, and other areas.

Collective motion of cells: from experiments to models.

Swarming or collective motion of living entities is one of the most common and spectacular manifestations of living systems that have been extensively studied in recent years. A number of general principles have been established. The interactions at the level of cells are quite different from those among individual animals, therefore the study of collective motion of cells is likely to reveal some specific important features which we plan to overview in this paper. In addition to presenting the most appealing results from the quickly growing related literature we also deliver a critical discussion of the emerging picture and summarize our present understanding of collective motion at the cellular level. Collective motion of cells plays an essential role in a number of experimental and real-life situations. In most cases the coordinated motion is a helpful aspect of the given phenomenon and results in making a related process more efficient (e.g., embryogenesis or wound healing), while in the case of tumor cell invasion it appears to speed up the progression of the disease. In these mechanisms cells both have to be motile and adhere to one another, the adherence feature being the most specific to this sort of collective behavior. One of the central aims of this review is to present the related experimental observations and treat them in light of a few basic computational models so as to make an interpretation of the phenomena at a quantitative level as well.

Flocking algorithm for autonomous flying robots.

Animal swarms displaying a variety of typical flocking patterns would not exist without the underlying safe, optimal and stable dynamics of the individuals. The emergence of these universal patterns can be efficiently reconstructed with agent-based models. If we want to reproduce these patterns with artificial systems, such as autonomous aerial robots, agent-based models can also be used in their control algorithms. However, finding the proper algorithms and thus understanding the essential characteristics of the emergent collective behaviour requires thorough and realistic modeling of the robot and also the environment. In this paper, we first present an abstract mathematical model of an autonomous flying robot. The model takes into account several realistic features, such as time delay and locality of communication, inaccuracy of the on-board sensors and inertial effects. We present two decentralized control algorithms. One is based on a simple self-propelled flocking model of animal collective motion, the other is a collective target tracking algorithm. Both algorithms contain a viscous friction-like term, which aligns the velocities of neighbouring agents parallel to each other. We show that this term can be essential for reducing the inherent instabilities of such a noisy and delayed realistic system. We discuss simulation results on the stability of the control algorithms, and perform real experiments to show the applicability of the algorithms on a group of autonomous quadcopters. In our case, bio-inspiration works in two ways. On the one hand, the whole idea of trying to build and control a swarm of robots comes from the observation that birds tend to flock to optimize their behaviour as a group. On the other hand, by using a realistic simulation framework and studying the group behaviour of autonomous robots we can learn about the major factors influencing the flight of bird flocks.

Shock waves on complex networks.

Power grids, road maps, and river streams are examples of infrastructural networks which are highly vulnerable to external perturbations. An abrupt local change of load (voltage, traffic density, or water level) might propagate in a cascading way and affect a significant fraction of the network. Almost discontinuous perturbations can be modeled by shock waves which can eventually interfere constructively and endanger the normal functionality of the infrastructure. We study their dynamics by solving the Burgers equation under random perturbations on several real and artificial directed graphs. Even for graphs with a narrow distribution of node properties (e.g., degree or betweenness), a steady state is reached exhibiting a heterogeneous load distribution, having a difference of one order of magnitude between the highest and average loads. Unexpectedly we find for the European power grid and for finite Watts-Strogatz networks a broad pronounced bimodal distribution for the loads. To identify the most vulnerable nodes, we introduce the concept of node-basin size, a purely topological property which we show to be strongly correlated to the average load of a node.

Leadership and path characteristics during walks are linked to dominance order and individual traits in dogs.

Movement interactions and the underlying social structure in groups have relevance across many social-living species. Collective motion of groups could be based on an "egalitarian" decision system, but in practice it is often influenced by underlying social network structures and by individual characteristics. We investigated whether dominance rank and personality traits are linked to leader and follower roles during joint motion of family dogs. We obtained high-resolution spatio-temporal GPS trajectory data (823,148 data points) from six dogs belonging to the same household and their owner during 14 30-40 min unleashed walks. We identified several features of the dogs' paths (e.g., running speed or distance from the owner) which are characteristic of a given dog. A directional correlation analysis quantifies interactions between pairs of dogs that run loops jointly. We found that dogs play the role of the leader about 50-85% of the time, i.e. the leader and follower roles in a given pair are dynamically interchangable. However, on a longer timescale tendencies to lead differ consistently. The network constructed from these loose leader-follower relations is hierarchical, and the dogs' positions in the network correlates with the age, dominance rank, trainability, controllability, and aggression measures derived from personality questionnaires. We demonstrated the possibility of determining dominance rank and personality traits of an individual based only on its logged movement data. The collective motion of dogs is influenced by underlying social network structures and by characteristics such as personality differences. Our findings could pave the way for automated animal personality and human social interaction measurements.

Hierarchical self-organization of non-cooperating individuals.

Hierarchy is one of the most conspicuous features of numerous natural, technological and social systems. The underlying structures are typically complex and their most relevant organizational principle is the ordering of the ties among the units they are made of according to a network displaying hierarchical features. In spite of the abundant presence of hierarchy no quantitative theoretical interpretation of the origins of a multi-level, knowledge-based social network exists. Here we introduce an approach which is capable of reproducing the emergence of a multi-levelled network structure based on the plausible assumption that the individuals (representing the nodes of the network) can make the right estimate about the state of their changing environment to a varying degree. Our model accounts for a fundamental feature of knowledge-based organizations: the less capable individuals tend to follow those who are better at solving the problems they all face. We find that relatively simple rules lead to hierarchical self-organization and the specific structures we obtain possess the two, perhaps most important features of complex systems: a simultaneous presence of adaptability and stability. In addition, the performance (success score) of the emerging networks is significantly higher than the average expected score of the individuals without letting them copy the decisions of the others. The results of our calculations are in agreement with a related experiment and can be useful from the point of designing the optimal conditions for constructing a given complex social structure as well as understanding the hierarchical organization of such biological structures of major importance as the regulatory pathways or the dynamics of neural networks.

Identification of behaviour in freely moving dogs (Canis familiaris) using inertial sensors.

Monitoring and describing the physical movements and body postures of animals is one of the most fundamental tasks of ethology. The more precise the observations are the more sophisticated the interpretations can be about the biology of a certain individual or species. Animal-borne data loggers have recently contributed much to the collection of motion-data from individuals, however, the problem of translating these measurements to distinct behavioural categories to create an ethogram is not overcome yet. The objective of the present study was to develop a "behaviour tracker": a system composed of a multiple sensor data-logger device (with a tri-axial accelerometer and a tri-axial gyroscope) and a supervised learning algorithm as means of automated identification of the behaviour of freely moving dogs. We collected parallel sensor measurements and video recordings of each of our subjects (Belgian Malinois, N=12; Labrador Retrievers, N=12) that were guided through a predetermined series of standard activities. Seven behavioural categories (lay, sit, stand, walk, trot, gallop, canter) were pre-defined and each video recording was tagged accordingly. Evaluation of the measurements was performed by support vector machine (SVM) classification. During the analysis we used different combinations of independent measurements for training and validation (belonging to the same or different individuals or using different training data size) to determine the robustness of the application. We reached an overall accuracy of above 90% perfect identification of all the defined seven categories of behaviour when both training and validation data belonged to the same individual, and over 80% perfect recognition rate using a generalized training data set of multiple subjects. Our results indicate that the present method provides a good model for an easily applicable, fast, automatic behaviour classification system that can be trained with arbitrary motion patterns and potentially be applied to a wide range of species and situations.

Group performance is maximized by hierarchical competence distribution.

Groups of people or even robots often face problems they need to solve together. Examples include collectively searching for resources, choosing when and where to invest time and effort, and many more. Although a hierarchical ordering of the relevance of the group members' inputs during collective decision making is abundant, a quantitative demonstration of its origin and advantages using a generic approach has not been described yet. Here we introduce a family of models based on the most general features of group decision making, and show that the optimal distribution of competences is a highly skewed function with a structured fat tail. Our results are obtained by optimizing the groups' compositions through identifying the best-performing distributions for both the competences and for the members' flexibilities/pliancies. Potential applications include choosing the best composition for a group intended to solve a given task.