Rao, R. P. N. & Ballard, D. H. Predictive coding in the visual cortex: a functional interpretation of some extra-classical receptive-field effects. Nat. Neurosci. 2, 79–87 (1999).

Article  CAS  PubMed  Google Scholar 

Keller, G. B. & Mrsic-Flogel, T. D. Predictive processing: a canonical cortical computation. Neuron 100, 424–435 (2018).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Friston, K. & Kiebel, S. Predictive coding under the free-energy principle. Philos. Trans. R. Soc. London B Biol. Sci. 364, 1211–1221 (2009).

Article  PubMed  PubMed Central  Google Scholar 

Jiang, L. P. & Rao, R. P. N. Predictive coding theories of cortical function. Oxford Research Encyclopedia of Neuroscience https://doi.org/10.1093/acrefore/9780190264086.013.328 (Oxford Univ. Press, 2022).

Halpern, B. P. Tasting and smelling as active, exploratory sensory processes. Am. J. Otolaryngol. 4, 246–249 (1983).

Article  CAS  PubMed  Google Scholar 

Lederman, S. J. & Klatzky, R. L. Hand movements: a window into haptic object recognition. Cogn. Psychol. 19, 342–368 (1987).

Article  CAS  PubMed  Google Scholar 

Ahissar, E. & Assa, E. Perception as a closed-loop convergence process. eLife 5, e12830 (2016).

Article  PubMed  PubMed Central  Google Scholar 

Friston, K. The free-energy principle: a unified brain theory? Nat. Rev. Neurosci. 11, 127–138 (2010).

Article  CAS  PubMed  Google Scholar 

Zatka-Haas, P., Steinmetz, N. A., Carandini, M. & Harris, K. D. Sensory coding and the causal impact of mouse cortex in a visual decision. eLife 10, e63163 (2021).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Steinmetz, N. A., Zatka-Haas, P., Carandini, M. & Harris, K. D. Distributed coding of choice, action and engagement across the mouse brain. Nature 576, 266–273 (2019).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Stringer, C. et al. Spontaneous behaviors drive multidimensional, brainwide activity. Science 364, eaav7893 (2019).

Article  CAS  Google Scholar 

Talluri, B. C. et al. Activity in primate visual cortex is minimally driven by spontaneous movements. Nat. Neurosci. 26, 1953–1959 (2023).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Jordan, R. & Keller, G. B. Opposing influence of top–down and bottom–up input on excitatory layer 2/3 neurons in mouse primary visual cortex. Neuron 108, 1194–1206 (2020).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Nakamura, K. & Colby, C. L. Updating of the visual representation in monkey striate and extrastriate cortex during saccades. Proc. Natl Acad. Sci. USA 99, 4026–4031 (2002).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Duhamel, J. R., Colby, C. L. & Goldberg, M. E. The updating of the representation of visual space in parietal cortex by intended eye movements. Science 255, 90–92 (1992).

Article  CAS  PubMed  Google Scholar 

Umeno, M. M. & Goldberg, M. E. Spatial processing in the monkey frontal eye field. I. Predictive visual responses. J. Neurophysiol. 78, 1373–1383 (1997).

Article  CAS  PubMed  Google Scholar 

Wurtz, R. H., McAlonan, K., Cavanaugh, J. & Berman, R. A. Thalamic pathways for active vision. Trends Cogn. Sci. 15, 177–184 (2011).

Article  PubMed  PubMed Central  Google Scholar 

Sherman, S. M. & Guillery, R. W. Functional Connections of Cortical Areas: A New View from the Thalamus (MIT, 2013).

Prasad, J., Carroll, B. & Sherman, S. Layer 5 corticofugal projections from diverse cortical areas: variations on a pattern of thalamic and extrathalamic targets. J. Neurosci. 40, 5785–5796 (2020).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Mountcastle, V. in the Mindful Brain (eds Edelman, G. & Mountcastle, V.) 7–50 (MIT, 1978).

Creutzfeldt, O. D. Generality of the functional structure of the neocortex. Naturwissenschaften 64, 507–517 (1977).

Article  CAS  PubMed  Google Scholar 

Mumford, D. On the computational architecture of the neocortex. II. The role of cortico–cortical loops. Biol. Cybern. 66, 241–251 (1992).

Article  CAS  PubMed  Google Scholar 

Hawkins, J. A Thousand Brains: A New Theory of Intelligence (Basic Books, 2021).

Douglas, R. J. & Martin, K. A. Neuronal circuits of the neocortex. Annu. Rev. Neurosci. 27, 419–451 (2004).

Article  CAS  PubMed  Google Scholar 

Harris, K. D. & Shepherd, G. M. The neocortical circuit: themes and variations. Nat. Neurosci. 18, 170–181 (2015).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Roe, A. W., Pallas, S. L., Kwon, Y. H. & Sur, M. Visual projections routed to the auditory pathway in ferrets: receptive fields of visual neurons in primary auditory cortex. J. Neurosci. 12, 3651–3664 (1992).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Felleman, D. & Essen, D. V. Distributed hierarchical processing in the primate cerebral cortex. Cereb. Cortex 1, 1–47 (1991).

Article  PubMed  Google Scholar 

Murray, J. D. et al. A hierarchy of intrinsic timescales across primate cortex. Nat. Neurosci. 17, 1661–1663 (2014).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Siegle, J. H. et al. Survey of spiking in the mouse visual system reveals functional hierarchy. Nature 592, 86–92 (2021).

Article  CAS  PubMed  PubMed Central  Google Scholar 

Grafton, S. T. & de C. Hamilton, A. F. Evidence for a distributed hierarchy of action representation in the brain. Hum. Mov. Sci. 26, 590–616 (2007).

Article  PubMed  PubMed Central  Google Scholar 

Gklezakos, D. C. & Rao, R. P. N. Active predictive coding networks: a neural solution to the problem of learning reference frames and part–whole hierarchies. Preprint at arxiv.org/abs/2201.08813 (2022).

Rao, R. P. N., Gklezakos, D. C. & Sathish, V. Active predictive coding: a unifying neural model for active perception, compositional learning, and hierarchical planning. Neural Comput. 36, 1–32 (2024).

Article  Google Scholar 

Fisher, A. & Rao, R. P. N. Recursive neural programs: a differentiable framework for learning compositional part–whole hierarchies and image grammars. PNAS Nexus 2, pgad337 (2023).

Article  PubMed  PubMed Central  Google Scholar 

Kasper, E., Larkman, A., Lübke, J. & Blakemore, C. Pyramidal neurons in layer 5 of the rat visual cortex. I. Correlation among cell morphology, intrinsic electrophysiological properties, and axon targets. J. Comp. Neurol. 339, 459–474 (1994).

Article  CAS  PubMed  Google Scholar 

Stebbings, K., Lesicko, A. & Llano, D. The auditory corticocollicular system: molecular and circuit-level considerations. Hear. Res. 314, 51–59 (2014).

Article