Emergence of grid-like representations by training recurrent neural networks to perform spatial localization
Not on the map. Why?
Abstract (arXiv)
Decades of research on the neural code underlying spatial navigation have revealed a diverse set of neural response properties. The Entorhinal Cortex (EC) of the mammalian brain contains a rich set of spatial correlates, including grid cells which encode space using tessellating patterns. However, the mechanisms and functional significance of these spatial representations remain largely mysterious. As a new way to understand these neural representations, we trained recurrent neural networks (RNNs) to perform navigation tasks in 2D arenas based on velocity inputs. Surprisingly, we find that grid-like spatial response patterns emerge in trained networks, along with units that exhibit other spatial correlates, including border cells and band-like cells. All these different functional types of neurons have been observed experimentally. The order of the emergence of grid-like and border cells is also consistent with observations from developmental studies. Together, our results suggest that grid cells, border cells and others as observed in EC may be a natural solution for representing space efficiently given the predominant recurrent connections in the neural circuits.
Key facts. Cited by 52 papers in the map (data through 7 Oct 2026). First posted on arXiv 21 Mar 2018.
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Citing papers per year: 2019: 1; 2020: 0; 2021: 6; 2022: 11; 2023: 12; 2024: 7; 2025: 11; 2026: 4.
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- 2021Relating transformers to models and neural representations of the hippocampal formationcited by 62
- 2023Sphere2Vec: A General-Purpose Location Representation Learning over a Spherical Surface for Large-Scale Geospatial Predictionscited by 38
- 2023Emergence of Maps in the Memories of Blind Navigation Agentscited by 31
- 2022Disentanglement with Biological Constraints: A Theory of Functional Cell Typescited by 30
- 2023Brain-inspired learning in artificial neural networks: a reviewcited by 26
- 2023Harmonics of Learning: Universal Fourier Features Emerge in Invariant Networkscited by 23
- 2022Multi-Object Navigation with dynamically learned neural implicit representationscited by 21
- 2021Reverse engineering recurrent neural networks with Jacobian switching linear dynamical systemscited by 17
- 2024Linking In-context Learning in Transformers to Human Episodic Memorycited by 15
- 2025Better artificial intelligence does not mean better models of biologycited by 10
- 2023Self-Supervised Learning of Representations for Space Generates Multi-Modular Grid Cellscited by 8
- 2022A Topological Deep Learning Framework for Neural Spike Decodingcited by 6
38 more
- 2023Learning with a Mole: Transferable latent spatial representations for navigation without reconstructioncited by 6
- 2024On Conformal Isometry of Grid Cells: Learning Distance-Preserving Position Embeddingcited by 6
- 2021Hippocampal formation-inspired probabilistic generative modelcited by 5
- 2021Graph augmented Deep Reinforcement Learning in the GameRLand3D environmentcited by 5
- 2022How to build a cognitive map: insights from models of the hippocampal formationcited by 5
- 2022Conformal Isometry of Lie Group Representation in Recurrent Network of Grid Cellscited by 5
- 2021SpikePropamine: Differentiable Plasticity in Spiking Neural Networkscited by 4
- 2022Human-AI Shared Control via Policy Dissectioncited by 4
- 2024Time Makes Space: Emergence of Place Fields in Networks Encoding Temporally Continuous Sensory Experiencescited by 4
- 2025EDEN: Entorhinal Driven Egocentric Navigation Toward Robotic Deploymentcited by 3
- 2021Emergent behavior and neural dynamics in artificial agents tracking turbulent plumescited by 2
- 2022Sphere2Vec: Multi-Scale Representation Learning over a Spherical Surface for Geospatial Predictionscited by 2
- 2022The Frost Hollow Experiments: Pavlovian Signalling as a Path to Coordination and Communication Between Agentscited by 2
- 2023Emergence of Grid-like Representations by Training Recurrent Networks with Conformal Normalizationcited by 2
- 2025Sufficient conditions for offline reactivation in recurrent neural networkscited by 2
- 2025REMI: Reconstructing Episodic Memory During Internally Driven Path Planningcited by 2
- 2022Emergence of Double-slit Interference by Representing Visual Space in Artificial Neural Networkscited by 1
- 2023From Data-Fitting to Discovery: Interpreting the Neural Dynamics of Motor Control through Reinforcement Learningcited by 1
- 2024Geometric sparsification in recurrent neural networkscited by 1
- 2025Building spatial world models from sparse transitional episodic memoriescited by 1
- 2019Imitation Learning of Factored Multi-agent Reactive Modelscited by 0
- 2022A Computational Theory of Learning Flexible Reward-Seeking Behavior with Place Cellscited by 0
- 2022Learning Parsimonious Dynamics for Generalization in Reinforcement Learningcited by 0
- 2023Determinantal Point Process Attention Over Grid Cell Code Supports Out of Distribution Generalizationcited by 0
- 2023A Population-Level Analysis of Neural Dynamics in Robust Legged Robotscited by 0
- 2023A Versatile Hub Model For Efficient Information Propagation And Feature Selectioncited by 0
- 2023Automated mapping of virtual environments with visual predictive codingcited by 0
- 2024Spectral-Stimulus Information for Self-Supervised Stimulus Encodingcited by 0
- 2024Visuospatial navigation from the bottom-up: without vestibular integration, distance prediction, or mapscited by 0
- 2024Spatial-Aware Decision-Making with Ring Attractors in Reinforcement Learning Systemscited by 0
- 2025SegDesicNet: Lightweight Semantic Segmentation in Remote Sensing with Geo-Coordinate Embeddings for Domain Adaptationcited by 0
- 2025Flexible inference for animal learning rules using neural networkscited by 0
- 2025Predictive Spike Timing Enables Distributed Shortest Path Computation in Spiking Neural Networkscited by 0
- 2025Hierarchical Learning for Maze Navigation: Emergence of Mental Representations via Second-Order Learningcited by 0
- 2025Vector Quantization in the Brain: Grid-like Codes in World Modelscited by 0
- 2025Boosting Brain-inspired Path Integration Efficiency via Learning-based Replication of Continuous Attractor Neurodynamicscited by 0
- 2026Paradoxical noise preference in RNNscited by 0
- 2026Path Integration and Object-Location Binding Emerge in an Action-Conditioned Predictive Sequence Networkcited by 0
50 most cited of 52 All 52 (JSON)
- 2026When and Where: A Model Hippocampal Network Unifies Formation of Time Cells and Place Cells22 Mar 2026
- 2026Leakage and Second-Order Dynamics Improve Hippocampal RNN Replay20 Feb 2026
- 2026Path Integration and Object-Location Binding Emerge in an Action-Conditioned Predictive Sequence Network3 Feb 2026
- 2026Paradoxical noise preference in RNNs8 Jan 2026
- 2025Boosting Brain-inspired Path Integration Efficiency via Learning-based Replication of Continuous Attractor Neurodynamics21 Nov 2025
- 2025Vector Quantization in the Brain: Grid-like Codes in World Models16 Oct 2025
- 2025Hierarchical Learning for Maze Navigation: Emergence of Mental Representations via Second-Order Learning17 Sep 2025
- 2025Predictive Spike Timing Enables Distributed Shortest Path Computation in Spiking Neural Networks12 Sep 2025
- 2025Flexible inference for animal learning rules using neural networks4 Sep 2025
- 2025REMI: Reconstructing Episodic Memory During Internally Driven Path Planning2 Jul 2025
- 2025EDEN: Entorhinal Driven Egocentric Navigation Toward Robotic Deployment3 Jun 2025
- 2025Sufficient conditions for offline reactivation in recurrent neural networks22 May 2025
38 more
- 2025Building spatial world models from sparse transitional episodic memories19 May 2025
- 2025Better artificial intelligence does not mean better models of biology8 Apr 2025
- 2025SegDesicNet: Lightweight Semantic Segmentation in Remote Sensing with Geo-Coordinate Embeddings for Domain Adaptation11 Mar 2025
- 2024Spatial-Aware Decision-Making with Ring Attractors in Reinforcement Learning Systems4 Oct 2024
- 2024Time Makes Space: Emergence of Place Fields in Networks Encoding Temporally Continuous Sensory Experiences11 Aug 2024
- 2024Visuospatial navigation from the bottom-up: without vestibular integration, distance prediction, or maps18 Jul 2024
- 2024Spectral-Stimulus Information for Self-Supervised Stimulus Encoding10 Jun 2024
- 2024Geometric sparsification in recurrent neural networks10 Jun 2024
- 2024On Conformal Isometry of Grid Cells: Learning Distance-Preserving Position Embedding27 May 2024
- 2024Linking In-context Learning in Transformers to Human Episodic Memory23 May 2024
- 2023Harmonics of Learning: Universal Fourier Features Emerge in Invariant Networks13 Dec 2023
- 2023Self-Supervised Learning of Representations for Space Generates Multi-Modular Grid Cells4 Nov 2023
- 2023Emergence of Grid-like Representations by Training Recurrent Networks with Conformal Normalization29 Oct 2023
- 2023Automated mapping of virtual environments with visual predictive coding20 Aug 2023
- 2023A Versatile Hub Model For Efficient Information Propagation And Feature Selection5 Jul 2023
- 2023Sphere2Vec: A General-Purpose Location Representation Learning over a Spherical Surface for Large-Scale Geospatial Predictions30 Jun 2023
- 2023A Population-Level Analysis of Neural Dynamics in Robust Legged Robots27 Jun 2023
- 2023Learning with a Mole: Transferable latent spatial representations for navigation without reconstruction6 Jun 2023
- 2023Determinantal Point Process Attention Over Grid Cell Code Supports Out of Distribution Generalization28 May 2023
- 2023Brain-inspired learning in artificial neural networks: a review18 May 2023
- 2023From Data-Fitting to Discovery: Interpreting the Neural Dynamics of Motor Control through Reinforcement Learning18 May 2023
- 2023Emergence of Maps in the Memories of Blind Navigation Agents30 Jan 2023
- 2022A Topological Deep Learning Framework for Neural Spike Decoding1 Dec 2022
- 2022Multi-Object Navigation with dynamically learned neural implicit representations11 Oct 2022
- 2022Conformal Isometry of Lie Group Representation in Recurrent Network of Grid Cells6 Oct 2022
- 2022Disentanglement with Biological Constraints: A Theory of Functional Cell Types30 Sep 2022
- 2022Learning Parsimonious Dynamics for Generalization in Reinforcement Learning29 Sep 2022
- 2022Human-AI Shared Control via Policy Dissection31 May 2022
- 2022Emergence of Double-slit Interference by Representing Visual Space in Artificial Neural Networks20 May 2022
- 2022A Computational Theory of Learning Flexible Reward-Seeking Behavior with Place Cells22 Apr 2022
- 2022The Frost Hollow Experiments: Pavlovian Signalling as a Path to Coordination and Communication Between Agents17 Mar 2022
- 2022How to build a cognitive map: insights from models of the hippocampal formation3 Feb 2022
- 2022Sphere2Vec: Multi-Scale Representation Learning over a Spherical Surface for Geospatial Predictions25 Jan 2022
- 2021Graph augmented Deep Reinforcement Learning in the GameRLand3D environment22 Dec 2021
- 2021Relating transformers to models and neural representations of the hippocampal formation7 Dec 2021
- 2021Reverse engineering recurrent neural networks with Jacobian switching linear dynamical systems1 Nov 2021
- 2021Emergent behavior and neural dynamics in artificial agents tracking turbulent plumes25 Sep 2021
- 2021SpikePropamine: Differentiable Plasticity in Spiking Neural Networks4 Jun 2021
50 newest of 52 All 52 (JSON)
- 2019Imitation Learning of Factored Multi-agent Reactive Models12 Mar 2019
- 2021Relating transformers to models and neural representations of the hippocampal formationcited by 62
- 2023Sphere2Vec: A General-Purpose Location Representation Learning over a Spherical Surface for Large-Scale Geospatial Predictionscited by 38
- 2023Emergence of Maps in the Memories of Blind Navigation Agentscited by 31
- 2022Disentanglement with Biological Constraints: A Theory of Functional Cell Typescited by 30
- 2023Brain-inspired learning in artificial neural networks: a reviewcited by 26
- 2023Harmonics of Learning: Universal Fourier Features Emerge in Invariant Networkscited by 23
- 2022Multi-Object Navigation with dynamically learned neural implicit representationscited by 21
- 2021Reverse engineering recurrent neural networks with Jacobian switching linear dynamical systemscited by 17
- 2024Linking In-context Learning in Transformers to Human Episodic Memorycited by 15
- 2025Better artificial intelligence does not mean better models of biologycited by 10
- 2023Self-Supervised Learning of Representations for Space Generates Multi-Modular Grid Cellscited by 8
- 2022A Topological Deep Learning Framework for Neural Spike Decodingcited by 6
12 most cited of 48 All 48 (JSON)
- 2026When and Where: A Model Hippocampal Network Unifies Formation of Time Cells and Place Cells22 Mar 2026
- 2026Leakage and Second-Order Dynamics Improve Hippocampal RNN Replay20 Feb 2026
- 2026Paradoxical noise preference in RNNs8 Jan 2026
- 2025Boosting Brain-inspired Path Integration Efficiency via Learning-based Replication of Continuous Attractor Neurodynamics21 Nov 2025
- 2025Hierarchical Learning for Maze Navigation: Emergence of Mental Representations via Second-Order Learning17 Sep 2025
- 2025Predictive Spike Timing Enables Distributed Shortest Path Computation in Spiking Neural Networks12 Sep 2025
- 2025Flexible inference for animal learning rules using neural networks4 Sep 2025
- 2025REMI: Reconstructing Episodic Memory During Internally Driven Path Planning2 Jul 2025
- 2025EDEN: Entorhinal Driven Egocentric Navigation Toward Robotic Deployment3 Jun 2025
- 2025Sufficient conditions for offline reactivation in recurrent neural networks22 May 2025
- 2025Better artificial intelligence does not mean better models of biology8 Apr 2025
- 2025SegDesicNet: Lightweight Semantic Segmentation in Remote Sensing with Geo-Coordinate Embeddings for Domain Adaptation11 Mar 2025
12 newest of 48 All 48 (JSON)