Raphaël Candelier
Current Organizations / Institutes
The organizations/institutes from the scholar's latest article
Previous Organizations / Institutes
The organizations / institutes from the scholar's other articles in the past ten years
Research Subjects
Research Interests
Raphaël Candelier has a total of 102 co-authors. Here are the top co-authors by number of shared publications:
Georges Debrégeas
Laboratoire Jean Perrin (LJP), Institut de Biologie Paris-Seine (IBPS), Sorbonne Université, CNRS, Paris, France
6 shared publications- Olivier Dauchot
Gulliver UMR CNRS 7083, ESPCI Paris, Université PSL, Paris, France
6 shared publications Benjamin Gallois
Laboratoire Jean Perrin (LJP), Institut de Biologie Paris-Seine (IBPS), Sorbonne Université, CNRS, Paris, France
4 shared publicationsG. Biroli
4 shared publications- Georges Debrégeas
Institut de Biologie Paris-Seine, Laboratoire Jean Perrin, Sorbonne Université, CNRS
4 shared publications - Germán Sumbre
Institut de Biologie de l’ENS (IBENS), Département de biologie, École normale supérieure, CNRS, INSERM, Université PSL, Paris, France
3 shared publications
Latest Publications
- Open Access
Maze-solving with density-driven swarms
We propose a new kind of collective motion where swarms of simple agents are able to find and fix the solution of two-dimensional mazes. The model consists of active memoryless particles interacting exclusively via short-ranged perception of local density and orientations. This system generates traveling density waves when constrained in one dimension, and self-organized swarms exploring local branches in two-dimensional mazes. Depending on a single kinetic parameter, the swarms can develop large tails and further gain long-term persistence, which ultimately allows them to robustly solve mazes of virtually any kind and size. By systematic exploration of the parameter space, we show that there exists a fast solving regime where the resolution time is linear in number of squares, hence making it an efficient maze-solving algorithm. Our model represents a new class of active systems with unprecedented contrast between the minimality of the processed information and the complexity of the resolved task, which is of prime importance for the interpretation and modeling of collective intelligence in living systems as well as for the design of future swarms of active particles and robots.
Open AccessMaze-solving with density-driven swarms
We propose a new kind of collective motion where swarms of simple agents are able to find and fix the solution of two-dimensional mazes. The model consists of active memoryless particles interacting exclusively via short-ranged perception of local density and orientations. This system generates traveling density waves when constrained in one dimension, and self-organized swarms exploring local branches in two-dimensional mazes. Depending on a single kinetic parameter, the swarms can develop large tails and further gain long-term persistence, which ultimately allows them to robustly solve mazes of virtually any kind and size. By systematic exploration of the parameter space, we show that there exists a fast solving regime where the resolution time is linear in number of squares, hence making it an efficient maze-solving algorithm. Our model represents a new class of active systems with unprecedented contrast between the minimality of the processed information and the complexity of the resolved task, which is of prime importance for the interpretation and modeling of collective intelligence in living systems as well as for the design of future swarms of active particles and robots.
- Open Access
- 11 March 2025
- Published by Journal of Graph Algorithms and Applications in Journal of Graph Algorithms and Applications
- Vol. 29 (1) , 289-320
Graph Matching Based on Similarities in Structure and Attributes
Finding vertex-to-vertex correspondences in real-world graphs is a challenging task with applications in a wide variety of domains. Structural matching based on graphs connectivities has attracted considerable attention, while the integration of all the other information stemming from vertices and edges attributes has been mostly left aside. Here we present the Graph Attributes and Structure Matching (GASM) algorithm, which provides high-quality solutions by integrating all the available information in a unified framework. Parameters quantifying the reliability of the attributes can tune how much the solutions should rely on the structure or on the attributes. We further show that even without attributes GASM consistently finds as-good-as or better solutions than state-of-the-art algorithms, with similar processing times.
- 11 March 2025
- Published by Journal of Graph Algorithms and Applications in Journal of Graph Algorithms and Applications
- Vol. 29 (1) , 289-320
Open AccessGraph Matching Based on Similarities in Structure and Attributes
Finding vertex-to-vertex correspondences in real-world graphs is a challenging task with applications in a wide variety of domains. Structural matching based on graphs connectivities has attracted considerable attention, while the integration of all the other information stemming from vertices and edges attributes has been mostly left aside. Here we present the Graph Attributes and Structure Matching (GASM) algorithm, which provides high-quality solutions by integrating all the available information in a unified framework. Parameters quantifying the reliability of the attributes can tune how much the solutions should rely on the structure or on the attributes. We further show that even without attributes GASM consistently finds as-good-as or better solutions than state-of-the-art algorithms, with similar processing times.
- Open Access
- 23 September 2022
- Published by Frontiers Media SA in Frontiers in Behavioral Neuroscience
- Vol. 16, 990792
A scalable assay for chemical preference of small freshwater fish
Sensing the chemical world is of primary importance for aquatic organisms, and small freshwater fish are increasingly used in toxicology, ethology, and neuroscience by virtue of their ease of manipulation, tissue imaging amenability, and genetic tractability. However, precise behavioral analyses are generally challenging to perform due to the lack of knowledge of what chemical the fish are exposed to at any given moment. Here we developed a behavioral assay and a specific infrared dye to probe the preference of young zebrafish for virtually any compound. We found that the innate aversion of zebrafish to citric acid is not mediated by modulation of the swim but rather by immediate avoidance reactions when the product is sensed and that the preference of juvenile zebrafish for ATP changes from repulsion to attraction during successive exposures. We propose an information-based behavioral model for which an exploration index emerges as a relevant behavioral descriptor, complementary to the standard preference index. Our setup features a high versatility in protocols and is automatic and scalable, which paves the way for high-throughput preference compound screening at different ages.
- 23 September 2022
- Published by Frontiers Media SA in Frontiers in Behavioral Neuroscience
- Vol. 16, 990792
Open AccessA scalable assay for chemical preference of small freshwater fish
Sensing the chemical world is of primary importance for aquatic organisms, and small freshwater fish are increasingly used in toxicology, ethology, and neuroscience by virtue of their ease of manipulation, tissue imaging amenability, and genetic tractability. However, precise behavioral analyses are generally challenging to perform due to the lack of knowledge of what chemical the fish are exposed to at any given moment. Here we developed a behavioral assay and a specific infrared dye to probe the preference of young zebrafish for virtually any compound. We found that the innate aversion of zebrafish to citric acid is not mediated by modulation of the swim but rather by immediate avoidance reactions when the product is sensed and that the preference of juvenile zebrafish for ATP changes from repulsion to attraction during successive exposures. We propose an information-based behavioral model for which an exploration index emerges as a relevant behavioral descriptor, complementary to the standard preference index. Our setup features a high versatility in protocols and is automatic and scalable, which paves the way for high-throughput preference compound screening at different ages.
- Open Access
A scalable assay for chemical preference of small freshwater fish
Sensing the chemical world is of primary importance for aquatic organisms, and small freshwater fish are increasingly used in toxicology, ethology, and neuroscience by virtue of their ease of manipulation, tissue imaging amenability, and genetic tractability. However, precise behavioral analyses are generally challenging to perform due to the lack of knowledge of what chemical the fish are exposed to at any given moment. Here we developed a behavioral assay and a specific infrared dye to probe the preference of young zebrafish for virtually any compound. We found that the innate aversion of zebrafish to citric acid is not mediated by modulation of the swim but rather by immediate avoidance reactions when the product is sensed and that the preference of juvenile zebrafish for ATP changes from repulsion to attraction during successive exposures. We propose an information-based behavioral model for which an exploration index emerges as a relevant behavioral descriptor, complementary to the standard preference index. Our setup features a high versatility in protocols and is automatic and scalable, which paves the way for high-throughput preference compound screening at different ages.
Open AccessA scalable assay for chemical preference of small freshwater fish
Sensing the chemical world is of primary importance for aquatic organisms, and small freshwater fish are increasingly used in toxicology, ethology, and neuroscience by virtue of their ease of manipulation, tissue imaging amenability, and genetic tractability. However, precise behavioral analyses are generally challenging to perform due to the lack of knowledge of what chemical the fish are exposed to at any given moment. Here we developed a behavioral assay and a specific infrared dye to probe the preference of young zebrafish for virtually any compound. We found that the innate aversion of zebrafish to citric acid is not mediated by modulation of the swim but rather by immediate avoidance reactions when the product is sensed and that the preference of juvenile zebrafish for ATP changes from repulsion to attraction during successive exposures. We propose an information-based behavioral model for which an exploration index emerges as a relevant behavioral descriptor, complementary to the standard preference index. Our setup features a high versatility in protocols and is automatic and scalable, which paves the way for high-throughput preference compound screening at different ages.
- 15 September 2021
- Published by Elsevier in Current Biology
- Vol. 31 (21) , 4762-4772.e5
Trans-inhibition of axon terminals underlies competition in the habenulo-interpeduncular pathway
- 15 September 2021
- Published by Elsevier in Current Biology
- Vol. 31 (21) , 4762-4772.e5
Trans-inhibition of axon terminals underlies competition in the habenulo-interpeduncular pathway
- Open Access
Thermal modulation of Zebrafish exploratory statistics reveals constraints on individual behavioral variability
Background: Variability is a hallmark of animal behavior. It contributes to survival by endowing individuals and populations with the capacity to adapt to ever-changing environmental conditions. Intra-individual variability is thought to reflect both endogenous and exogenous modulations of the neural dynamics of the central nervous system. However, how variability is internally regulated and modulated by external cues remains elusive. Here we address this question by analyzing the statistics of spontaneous exploration of freely swimming zebrafish larvae, and by probing how these locomotor patterns are impacted when changing the water temperatures within an ethologically relevant range. Results: We show that, for this simple animal model, five short-term kinematic parameters - interbout interval, turn amplitude, travelled distance, turn probability and orientational flipping rate - together control the long-term exploratory dynamics. We establish that the bath temperature consistently impacts the means of these parameters, but leave their pairwise covariance unchanged. These results indicate that the temperature merely controls the sampling statistics within a well-defined kinematic space delineated by this robust statistical structure. At a given temperature, individual animals explore the behavioral space over a timescale of tens of minutes, suggestive of a slow internal state modulation that could be externally biased through the bath temperature. By combining these various observations into a minimal stochastic model of navigation, we show that this thermal modulation of locomotor kinematics results in a thermophobic behavior, complementing direct gradient-sensing mechanisms. Conclusions: This study establishes the existence of a well-defined locomotor space accessible to zebrafish larvae during spontaneous exploration, and quantifies self-generated modulation of locomotor patterns. Intra-individual variability reflects a slow diffusive-like probing of this space by the animal. The bath temperature in turn restricts the sampling statistics to sub-regions, endowing the animal with basic thermophobicity. This study suggests that in Zebrafish, as well as in other ectothermic animals, ambient temperature could be used to efficiently manipulate internal states in a simple and ethological way.
Open AccessThermal modulation of Zebrafish exploratory statistics reveals constraints on individual behavioral variability
Background: Variability is a hallmark of animal behavior. It contributes to survival by endowing individuals and populations with the capacity to adapt to ever-changing environmental conditions. Intra-individual variability is thought to reflect both endogenous and exogenous modulations of the neural dynamics of the central nervous system. However, how variability is internally regulated and modulated by external cues remains elusive. Here we address this question by analyzing the statistics of spontaneous exploration of freely swimming zebrafish larvae, and by probing how these locomotor patterns are impacted when changing the water temperatures within an ethologically relevant range. Results: We show that, for this simple animal model, five short-term kinematic parameters - interbout interval, turn amplitude, travelled distance, turn probability and orientational flipping rate - together control the long-term exploratory dynamics. We establish that the bath temperature consistently impacts the means of these parameters, but leave their pairwise covariance unchanged. These results indicate that the temperature merely controls the sampling statistics within a well-defined kinematic space delineated by this robust statistical structure. At a given temperature, individual animals explore the behavioral space over a timescale of tens of minutes, suggestive of a slow internal state modulation that could be externally biased through the bath temperature. By combining these various observations into a minimal stochastic model of navigation, we show that this thermal modulation of locomotor kinematics results in a thermophobic behavior, complementing direct gradient-sensing mechanisms. Conclusions: This study establishes the existence of a well-defined locomotor space accessible to zebrafish larvae during spontaneous exploration, and quantifies self-generated modulation of locomotor patterns. Intra-individual variability reflects a slow diffusive-like probing of this space by the animal. The bath temperature in turn restricts the sampling statistics to sub-regions, endowing the animal with basic thermophobicity. This study suggests that in Zebrafish, as well as in other ectothermic animals, ambient temperature could be used to efficiently manipulate internal states in a simple and ethological way.
- Open Access
Evolutionary divergence of locomotion in two related vertebrate species
Locomotion exists in diverse forms in nature and is adapted to the environmental constraints of each species 1 . However, little is known about how closely related species with similar neuronal circuitry can evolve different navigational strategies to explore their environments. We established a powerful approach in comparative neuroethology to investigate evolution of neuronal circuits in vertebrates by comparing divergent swimming pattern of two closely related larval fish species, Danionella translucida (DT) and Danio rerio or zebrafish (ZF) 2,3 . During swimming, we demonstrate that DT utilizes lower half tail-beat frequency and amplitude to generate a slower and continuous swimming pattern when compared to the burst-and-glide swimming pattern in ZF. We found a high degree of conservation in the brain anatomy between the two species. However, we revealed that the activity of a higher motor region, referred here as the Mesencephalic Locomotion Maintenance Neurons (MLMN) correlates with the duration of swim events and differs strikingly between DT and ZF. Using holographic stimulation, we show that the activation of the MLMN is sufficient to increase the frequency and duration of swim events in ZF. Moreover, we propose two characteristics, availability of dissolved oxygen and timing of swim bladder inflation, which drive the observed differences in the swim pattern. Our findings uncover the neuronal circuit substrate underlying the evolutionary divergence of navigational strategies and how they are adapted to their respective environmental constraints.
Open AccessEvolutionary divergence of locomotion in two related vertebrate species
Locomotion exists in diverse forms in nature and is adapted to the environmental constraints of each species 1 . However, little is known about how closely related species with similar neuronal circuitry can evolve different navigational strategies to explore their environments. We established a powerful approach in comparative neuroethology to investigate evolution of neuronal circuits in vertebrates by comparing divergent swimming pattern of two closely related larval fish species, Danionella translucida (DT) and Danio rerio or zebrafish (ZF) 2,3 . During swimming, we demonstrate that DT utilizes lower half tail-beat frequency and amplitude to generate a slower and continuous swimming pattern when compared to the burst-and-glide swimming pattern in ZF. We found a high degree of conservation in the brain anatomy between the two species. However, we revealed that the activity of a higher motor region, referred here as the Mesencephalic Locomotion Maintenance Neurons (MLMN) correlates with the duration of swim events and differs strikingly between DT and ZF. Using holographic stimulation, we show that the activation of the MLMN is sufficient to increase the frequency and duration of swim events in ZF. Moreover, we propose two characteristics, availability of dissolved oxygen and timing of swim bladder inflation, which drive the observed differences in the swim pattern. Our findings uncover the neuronal circuit substrate underlying the evolutionary divergence of navigational strategies and how they are adapted to their respective environmental constraints.
- Open Access
- 11 February 2021
- Published by Public Library of Science (PLoS) in PLOS Computational Biology
- Vol. 17 (2) , e1008697
FastTrack: An open-source software for tracking varying numbers of deformable objects
Analyzing the dynamical properties of mobile objects requires to extract trajectories from recordings, which is often done by tracking movies. We compiled a database of two-dimensional movies for very different biological and physical systems spanning a wide range of length scales and developed a general-purpose, optimized, open-source, cross-platform, easy to install and use, self-updating software called FastTrack. It can handle a changing number of deformable objects in a region of interest, and is particularly suitable for animal and cell tracking in two-dimensions. Furthermore, we introduce the probability of incursions as a new measure of a movie’s trackability that doesn’t require the knowledge of ground truth trajectories, since it is resilient to small amounts of errors and can be computed on the basis of an ad hoc tracking. We also leveraged the versatility and speed of FastTrack to implement an iterative algorithm determining a set of nearly-optimized tracking parameters—yet further reducing the amount of human intervention—and demonstrate that FastTrack can be used to explore the space of tracking parameters to optimize the number of swaps for a batch of similar movies. A benchmark shows that FastTrack is orders of magnitude faster than state-of-the-art tracking algorithms, with a comparable tracking accuracy. The source code is available under the GNU GPLv3 at https://github.com/FastTrackOrg/FastTrack and pre-compiled binaries for Windows, Mac and Linux are available at http://www.fasttrack.sh.
- 11 February 2021
- Published by Public Library of Science (PLoS) in PLOS Computational Biology
- Vol. 17 (2) , e1008697
Open AccessFastTrack: An open-source software for tracking varying numbers of deformable objects
Analyzing the dynamical properties of mobile objects requires to extract trajectories from recordings, which is often done by tracking movies. We compiled a database of two-dimensional movies for very different biological and physical systems spanning a wide range of length scales and developed a general-purpose, optimized, open-source, cross-platform, easy to install and use, self-updating software called FastTrack. It can handle a changing number of deformable objects in a region of interest, and is particularly suitable for animal and cell tracking in two-dimensions. Furthermore, we introduce the probability of incursions as a new measure of a movie’s trackability that doesn’t require the knowledge of ground truth trajectories, since it is resilient to small amounts of errors and can be computed on the basis of an ad hoc tracking. We also leveraged the versatility and speed of FastTrack to implement an iterative algorithm determining a set of nearly-optimized tracking parameters—yet further reducing the amount of human intervention—and demonstrate that FastTrack can be used to explore the space of tracking parameters to optimize the number of swaps for a batch of similar movies. A benchmark shows that FastTrack is orders of magnitude faster than state-of-the-art tracking algorithms, with a comparable tracking accuracy. The source code is available under the GNU GPLv3 at https://github.com/FastTrackOrg/FastTrack and pre-compiled binaries for Windows, Mac and Linux are available at http://www.fasttrack.sh.
- Open Access
FastTrack: an open-source software for tracking varying numbers of deformable objects
Analyzing the dynamical properties of mobile objects requires to extract trajectories from recordings, which is often done by tracking movies. We compiled a database of two-dimensional movies for very different biological and physical systems spanning a wide range of length scales and developed a general-purpose, optimized, open-source, cross-platform, easy to install and use, self-updating software called FastTrack. It can handle a changing number of deformable objects in a region of interest, and is particularly suitable for animal and cell tracking in two-dimensions. Furthermore, we introduce the probability of incursions as a new measure of a movie's trackability that doesn't require the knowledge of ground truth trajectories, since it is resilient to small amounts of errors and can be computed on the basis of an ad hoc tracking. We also leveraged the versatility and speed of FastTrack to implement an iterative algorithm determining a set of nearly-optimized tracking parameters -- yet further reducing the amount of human intervention -- and demonstrate that FastTrack can be used to explore the space of tracking parameters to optimize the number of swaps for a batch of similar movies. A benchmark shows that FastTrack is orders of magnitude faster than state-of-the-art tracking algorithms, with a comparable tracking accuracy. The source code is available under the GNU GPLv3 at https://github.com/FastTrackOrg/FastTrack and pre-compiled binaries for Windows, Mac and Linux are available at http://www.fasttrack.sh.
Open AccessFastTrack: an open-source software for tracking varying numbers of deformable objects
Analyzing the dynamical properties of mobile objects requires to extract trajectories from recordings, which is often done by tracking movies. We compiled a database of two-dimensional movies for very different biological and physical systems spanning a wide range of length scales and developed a general-purpose, optimized, open-source, cross-platform, easy to install and use, self-updating software called FastTrack. It can handle a changing number of deformable objects in a region of interest, and is particularly suitable for animal and cell tracking in two-dimensions. Furthermore, we introduce the probability of incursions as a new measure of a movie's trackability that doesn't require the knowledge of ground truth trajectories, since it is resilient to small amounts of errors and can be computed on the basis of an ad hoc tracking. We also leveraged the versatility and speed of FastTrack to implement an iterative algorithm determining a set of nearly-optimized tracking parameters -- yet further reducing the amount of human intervention -- and demonstrate that FastTrack can be used to explore the space of tracking parameters to optimize the number of swaps for a batch of similar movies. A benchmark shows that FastTrack is orders of magnitude faster than state-of-the-art tracking algorithms, with a comparable tracking accuracy. The source code is available under the GNU GPLv3 at https://github.com/FastTrackOrg/FastTrack and pre-compiled binaries for Windows, Mac and Linux are available at http://www.fasttrack.sh.
- Open Access
- 2 January 2020
- Published by eLife Sciences Publications, Ltd in eLife
- Vol. 9
From behavior to circuit modeling of light-seeking navigation in zebrafish larvae
Bridging brain-scale circuit dynamics and organism-scale behavior is a central challenge in neuroscience. It requires the concurrent development of minimal behavioral and neural circuit models that can quantitatively capture basic sensorimotor operations. Here, we focus on light-seeking navigation in zebrafish larvae. Using a virtual reality assay, we first characterize how motor and visual stimulation sequences govern the selection of discrete swim-bout events that subserve the fish navigation in the presence of a distant light source. These mechanisms are combined into a comprehensive Markov-chain model of navigation that quantitatively predicts the stationary distribution of the fish’s body orientation under any given illumination profile. We then map this behavioral description onto a neuronal model of the ARTR, a small neural circuit involved in the orientation-selection of swim bouts. We demonstrate that this visually-biased decision-making circuit can capture the statistics of both spontaneous and contrast-driven navigation.
- 2 January 2020
- Published by eLife Sciences Publications, Ltd in eLife
- Vol. 9
Open AccessFrom behavior to circuit modeling of light-seeking navigation in zebrafish larvae
Bridging brain-scale circuit dynamics and organism-scale behavior is a central challenge in neuroscience. It requires the concurrent development of minimal behavioral and neural circuit models that can quantitatively capture basic sensorimotor operations. Here, we focus on light-seeking navigation in zebrafish larvae. Using a virtual reality assay, we first characterize how motor and visual stimulation sequences govern the selection of discrete swim-bout events that subserve the fish navigation in the presence of a distant light source. These mechanisms are combined into a comprehensive Markov-chain model of navigation that quantitatively predicts the stationary distribution of the fish’s body orientation under any given illumination profile. We then map this behavioral description onto a neuronal model of the ARTR, a small neural circuit involved in the orientation-selection of swim bouts. We demonstrate that this visually-biased decision-making circuit can capture the statistics of both spontaneous and contrast-driven navigation.