Emergence of Grid-like Representations by Training Recurrent Networks with Conformal Normalization
Abstract: Grid cells in the entorhinal cortex of mammalian brains exhibit striking hexagon grid firing patterns in their response maps as the animal (e.g., a rat) navigates in a 2D open environment. In this paper, we study the emergence of the hexagon grid patterns of grid cells based on a general recurrent neural network (RNN) model that captures the navigation process. The responses of grid cells collectively form a high dimensional vector, representing the 2D self-position of the agent. As the agent moves, the vector is transformed by an RNN that takes the velocity of the agent as input. We propose a simple yet general conformal normalization of the input velocity of the RNN, so that the local displacement of the position vector in the high-dimensional neural space is proportional to the local displacement of the agent in the 2D physical space, regardless of the direction of the input velocity. We apply this mechanism to both a linear RNN and nonlinear RNNs. Theoretically, we provide an understanding that explains the connection between conformal normalization and the emergence of hexagon grid patterns. Empirically, we conduct extensive experiments to verify that conformal normalization is crucial for the emergence of hexagon grid patterns, across various types of RNNs. The learned patterns share similar profiles to biological grid cells, and the topological properties of the patterns also align with our theoretical understanding.
- A theory of joint attractor dynamics in the hippocampus and the entorhinal cortex accounts for artificial remapping and grid cell field-to-field variability. eLife, 9:e56894, 2020.
- Amit, D. J. Modeling brain function: The world of attractor neural networks. Cambridge university press, 1992.
- Solid state physics, 1976.
- Layer normalization. arXiv preprint arXiv:1607.06450, 2016.
- Banchoff, T. http://www.math.brown.edu/tbanchof/gc/script/b3d/hypertorus.html, 1968.
- Vector-based navigation using grid-like representations in artificial agents. Nature, 557(7705):429, 2018.
- Experience-dependent rescaling of entorhinal grids. Nature neuroscience, 10(6):682–684, 2007.
- Navigating cognition: Spatial codes for human thinking. Science, 362(6415):eaat6766, 2018.
- Scale-invariant memory representations emerge from moire interference between grid fields that produce theta oscillations: a computational model. Journal of Neuroscience, 27(12):3211–3229, 2007.
- Accurate path integration in continuous attractor network models of grid cells. PLoS computational biology, 5(2):e1000291, 2009.
- Normalization as a canonical neural computation. Nature Reviews Neuroscience, 13(1):51–62, 2012.
- Organizing conceptual knowledge in humans with a gridlike code. Science, 352(6292):1464–1468, 2016.
- Recurrent inhibitory circuitry as a mechanism for grid formation. Nature neuroscience, 16(3):318–324, 2013.
- Emergence of grid-like representations by training recurrent neural networks to perform spatial localization. arXiv preprint arXiv:1803.07770, 2018.
- Emergence of functional and structural properties of the head direction system by optimization of recurrent neural networks. International Conferences on Learning Representations (ICLR), 2020.
- The input–output transformation of the hippocampal granule cells: from grid cells to place fields. Journal of Neuroscience, 29(23):7504–7512, 2009.
- Evidence for grid cells in a human memory network. Nature, 463(7281):657, 2010.
- Extracting grid cell characteristics from place cell inputs using non-negative principal component analysis. Elife, 5:e10094, 2016.
- Actionable neural representations: Grid cells from minimal constraints. arXiv preprint arXiv:2209.15563, 2022.
- The elementary geometric structure of compact lie groups. Bulletin of the London Mathematical Society, 30(4):337–364, 1998.
- What grid cells convey about rat location. Journal of Neuroscience, 28(27):6858–6871, 2008.
- A spin glass model of path integration in rat medial entorhinal cortex. Journal of Neuroscience, 26(16):4266–4276, 2006.
- Spatial representation in the entorhinal cortex. Science, 305(5688):1258–1264, 2004.
- Grid cells in mice. Hippocampus, 18(12):1230–1238, 2008.
- Learning grid cells as vector representation of self-position coupled with matrix representation of self-motion. In International Conference on Learning Representations, 2019.
- On path integration of grid cells: group representation and isotropic scaling. In Neural Information Processing Systems, 2021.
- Toroidal topology of population activity in grid cells. Nature, 602(7895):123–128, 2022.
- Cortical neurons: isolation of contrast gain control. Vision research, 32(8):1409–1410, 1992.
- Impaired path integration in mice with disrupted grid cell firing. Nature neuroscience, 21(1):81–91, 2018.
- Microstructure of a spatial map in the entorhinal cortex. Nature, 436(7052):801, 2005.
- Heeger, D. J. Normalization of cell responses in cat striate cortex. Visual neuroscience, 9(2):181–197, 1992.
- Gaussian error linear units (gelus). arXiv preprint arXiv:1606.08415, 2016.
- Grid-like processing of imagined navigation. Current Biology, 26(6):842–847, 2016.
- Ioffe, S. Batch renormalization: Towards reducing minibatch dependence in batch-normalized models. Advances in neural information processing systems, 30, 2017.
- Batch normalization: Accelerating deep network training by reducing internal covariate shift. arXiv preprint arXiv:1502.03167, 2015.
- Direct recordings of grid-like neuronal activity in human spatial navigation. Nature neuroscience, 16(9):1188, 2013.
- A map of visual space in the primate entorhinal cortex. Nature, 491(7426):761, 2012.
- Langley, P. Crafting papers on machine learning. In Langley, P. (ed.), Proceedings of the 17th International Conference on Machine Learning (ICML 2000), pp. 1207–1216, Stanford, CA, 2000. Morgan Kaufmann.
- Development of the spatial representation system in the rat. Science, 328(5985):1576–1580, 2010.
- Probable nature of higher-dimensional symmetries underlying mammalian grid-cell activity patterns. Elife, 4:e05979, 2015.
- Path integration and the neural basis of the’cognitive map’. Nature Reviews Neuroscience, 7(8):663, 2006.
- Network mechanisms of grid cells. Philosophical Transactions of the Royal Society B: Biological Sciences, 369(1635):20120511, 2014.
- Where am i? where am i going? Scientific American, 314(1):26–33, 2016.
- Explaining heterogeneity in medial entorhinal cortex with task-driven neural networks. Advances in Neural Information Processing Systems, 34:12167–12179, 2021.
- Niell, C. M. Cell types, circuits, and receptive fields in the mouse visual cortex. Annual review of neuroscience, 38:413–431, 2015.
- The hippocampus as a spatial map: preliminary evidence from unit activity in the freely-moving rat. Brain research, 1971.
- Précis of o’keefe & nadel’s the hippocampus as a cognitive map. Behavioral and Brain Sciences, 2(4):487–494, 1979.
- Feedback inhibition enables theta-nested gamma oscillations and grid firing fields. Neuron, 77(1):141–154, 2013.
- Impaired speed encoding is associated with reduced grid cell periodicity in a mouse model of tauopathy. bioRxiv, pp. 595652, 2019.
- Functional properties of stellate cells in medial entorhinal cortex layer ii. Elife, 7:e36664, 2018.
- The stabilized supralinear network: a unifying circuit motif underlying multi-input integration in sensory cortex. Neuron, 85(2):402–417, 2015.
- Conjunctive representation of position, direction, and velocity in entorhinal cortex. Science, 312(5774):758–762, 2006.
- Spectral methods for dimensionality reduction. Semi-supervised learning, 3, 2006.
- No free lunch from deep learning in neuroscience: A case study through models of the entorhinal-hippocampal circuit. Advances in Neural Information Processing Systems, 35:16052–16067, 2022a.
- No free lunch from deep learning in neuroscience: A case study through models of the entorhinal-hippocampal circuit. bioRxiv, 2022b.
- Self-supervised learning of representations for space generates multi-modular grid cells. arXiv preprint arXiv:2311.02316, 2023.
- Coherently remapping toroidal cells but not grid cells are responsible for path integration in virtual agents. bioRxiv, pp. 2022–08, 2022.
- Natural signal statistics and sensory gain control. Nature neuroscience, 4(8):819–825, 2001.
- A unified theory for the origin of grid cells through the lens of pattern formation. 2019.
- A unified theory for the computational and mechanistic origins of grid cells. Neuron, 111(1):121–137, 2023.
- Grid cells generate an analog error-correcting code for singularly precise neural computation. Nature neuroscience, 14(10):1330, 2011.
- The hippocampus as a predictive map. Nature neuroscience, 20(11):1643, 2017.
- Connecting multiple spatial scales to decode the population activity of grid cells. Science Advances, 1(11):e1500816, 2015.
- The entorhinal grid map is discretized. Nature, 492(7427):72, 2012.
- Taylor, M. Lectures on lie groups. Lecture Notes, available at http://www. unc. edu/math/Faculty/met/lieg. html, 2002.
- Tolman, E. C. Cognitive maps in rats and men. Psychological review, 55(4):189, 1948.
- Attention is all you need. arXiv preprint arXiv:1706.03762, 2017.
- A principle of economy predicts the functional architecture of grid cells. Elife, 4:e08362, 2015a.
- A principle of economy predicts the functional architecture of grid cells. Elife, 4:e08362, 2015b.
- Relating transformers to models and neural representations of the hippocampal formation. arXiv preprint arXiv:2112.04035, 2021.
- Group normalization. In Proceedings of the European conference on computer vision (ECCV), pp. 3–19, 2018.
- Conformal isometry of lie group representation in recurrent network of grid cells. arXiv preprint arXiv:2210.02684, 2022.
- Grid cells without theta oscillations in the entorhinal cortex of bats. Nature, 479(7371):103, 2011.
- Optogenetic dissection of entorhinal-hippocampal functional connectivity. Science, 340(6128), 2013.
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