{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,17]],"date-time":"2026-10-17T01:42:30Z","timestamp":1792201350443,"version":"4.4.0"},"reference-count":123,"publisher":"MIT Press","issue":"11","license":[{"start":{"date-parts":[[2026,9,22]],"date-time":"2026-09-22T00:00:00Z","timestamp":1790035200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["direct.mit.edu"],"crossmark-restriction":true},"short-container-title":[],"published-print":{"date-parts":[[2026,10,16]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Vernon Mountcastle hypothesized that the basis for intelligence in mammals is the replication of a general computational unit, the cortical column. The Thousand Brains Theory proposed that each column is a sensorimotor system, capable of learning structured models of objects by integrating sensory input over multiple movements. Previous papers on the Thousand Brains Theory focused on the computations that occur within individual cortical columns and how columns can use long-range connections to rapidly reach a consensus. However, several prominent long-range connection types in the neocortex were not addressed by the theory. These include hierarchical feedforward and feedback connections, as well as those that go through the thalamus. In addition, several theoretical requirements were not addressed. These include how the cortex learns compositional objects and how information is converted from the egocentric perspective of sensors to the allocentric perspective of models in the cortex. In this letter, we extend the Thousand Brains Theory to address these issues. We begin by reviewing the anatomy of long-range neocortical connections, arguing that they form a heterarchy, rather than hierarchy, which has made their functions challenging to understand through existing theoretical models. We then propose specific roles for each of these connections. First, the thalamus converts the orientation of features and movement information from an egocentric perspective to the allocentric perspective of learned models. Second, hierarchical feedforward, feedback, and cortico-thalamo-cortical projections enable columns to learn compositional models. We discuss the relationship of our proposals to experimental findings at the levels of anatomy, neurophysiology, and behavior, along with testable predictions for future experimental work. We conclude by discussing the implications of the extended Thousand Brains Theory for the design of artificial intelligence.<\/jats:p>","DOI":"10.1162\/neco.a.1579","type":"journal-article","created":{"date-parts":[[2026,9,22]],"date-time":"2026-09-22T17:41:58Z","timestamp":1790098918000},"page":"2042-2078","update-policy":"https:\/\/doi.org\/10.1162\/mitpressjournals.corrections.policy","source":"Crossref","is-referenced-by-count":0,"title":["The Thousand Brains Theory 2.0: An Extension for the Long-Range Connections of the Neocortical Heterarchy"],"prefix":"10.1162","volume":"38","author":[{"given":"Jeff","family":"Hawkins","sequence":"first","affiliation":[{"name":"Thousand Brains Project, Santa Rosa, CA 95401, U.S.A. jhawkins@thousandbrains.org"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Niels","family":"Leadholm","sequence":"additional","affiliation":[{"name":"Thousand Brains Project, Santa Rosa, CA 95401, U.S.A. nleadholm@thousandbrains.org"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Viviane","family":"Clay","sequence":"additional","affiliation":[{"name":"Thousand Brains Project, Santa Rosa, CA 95401, U.S.A. vclay@thousandbrains.org"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"281","published-online":{"date-parts":[[2026,10,16]]},"reference":[{"issue":"19","key":"2026101621153260800_bib1","doi-asserted-by":"publisher","first-page":"8633","DOI":"10.1523\/JNEUROSCI.22-19-08633.2002","article-title":"Circuits for local and global signal integration in primary visual cortex","volume":"22","author":"Angelucci","year":"2002","journal-title":"Journal of Neuroscience"},{"issue":"9143","key":"2026101621153260800_bib2","doi-asserted-by":"publisher","first-page":"1841","DOI":"10.1016\/S0140-6736(98)05430-0","article-title":"Vertigo","volume":"352","author":"Baloh","year":"1998","journal-title":"Lancet"},{"issue":"39","key":"2026101621153260800_bib3","doi-asserted-by":"publisher","first-page":"8441","DOI":"10.1523\/JNEUROSCI.1400-04.2004","article-title":"A quantitative map of the circuit of cat primary visual cortex","volume":"24","author":"Binzegger","year":"2004","journal-title":"Journal of Neuroscience"},{"issue":"8","key":"2026101621153260800_bib4","doi-asserted-by":"publisher","DOI":"10.1038\/nn1278","article-title":"Underlying principles of visual shape selectivity in posterior inferotemporal cortex","volume":"7","author":"Brincat","year":"2004","journal-title":"Nature Neuroscience"},{"issue":"1","key":"2026101621153260800_bib5","doi-asserted-by":"publisher","DOI":"10.1007\/BF00239409","article-title":"Projection of the lateral geniculate nucleus onto cortical area V2 in the macaque monkey","volume":"53","author":"Bullier","year":"1983","journal-title":"Experimental Brain Research"},{"issue":"9","key":"2026101621153260800_bib6","doi-asserted-by":"publisher","first-page":"801","DOI":"10.1002\/hipo.20327","article-title":"An oscillatory interference model of grid cell firing","volume":"17","author":"Burgess","year":"2007","journal-title":"Hippocampus"},{"issue":"3","key":"2026101621153260800_bib7","doi-asserted-by":"publisher","first-page":"1071","DOI":"10.1073\/pnas.86.3.1071","article-title":"Organization of corticocortical connections in human visual cortex","volume":"86","author":"Burkhalter","year":"1989","journal-title":"Proceedings of the National Academy of Sciences"},{"key":"2026101621153260800_bib8","volume-title":"Comparative study of the sensory areas of the human cortex","author":"Cajal","year":"1899"},{"key":"2026101621153260800_bib9","author":"Clay","year":"2024","journal-title":"The Thousand Brains Project: A new paradigm for sensorimotor intelligence"},{"issue":"23","key":"2026101621153260800_bib10","doi-asserted-by":"publisher","first-page":"191","DOI":"10.1038\/newbio231191a0","article-title":"Simultaneous recording of input and output of lateral geniculate neurones","volume":"231","author":"Cleland","year":"1971","journal-title":"Nature New Biology"},{"issue":"6292","key":"2026101621153260800_bib11","doi-asserted-by":"publisher","first-page":"1464","DOI":"10.1126\/science.aaf0941","article-title":"Organizing conceptual knowledge in humans with a gridlike code","volume":"352","author":"Constantinescu","year":"2016","journal-title":"Science"},{"issue":"6140","key":"2026101621153260800_bib12","doi-asserted-by":"publisher","first-page":"1591","DOI":"10.1126\/science.1236425","article-title":"Deep cortical layers are activated directly by thalamus","volume":"340","author":"Constantinople","year":"2013","journal-title":"Science"},{"key":"2026101621153260800_bib13","article-title":"Whose cortical column would that be?","volume":"4","author":"Da Costa","year":"2010","journal-title":"Frontiers in Neuroanatomy"},{"issue":"9","key":"2026101621153260800_bib14","doi-asserted-by":"publisher","first-page":"1103","DOI":"10.1038\/s41592-021-01239-8","article-title":"High-speed, cortex-wide volumetric recording of neuroactivity at cellular resolution using light beads microscopy","volume":"18","author":"Demas","year":"2021","journal-title":"Nature Methods"},{"issue":"11","key":"2026101621153260800_bib15","doi-asserted-by":"publisher","first-page":"1687","DOI":"10.1038\/nn.4131","article-title":"Connectivity of mouse somatosensory and prefrontal cortex examined with trans-synaptic tracing","volume":"18","author":"DeNardo","year":"2015","journal-title":"Nature Neuroscience"},{"issue":"3","key":"2026101621153260800_bib16","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2012.01.010","article-title":"How does the brain solve visual object recognition?","volume":"73","author":"DiCarlo","year":"2012","journal-title":"Neuron"},{"key":"2026101621153260800_bib17","doi-asserted-by":"publisher","DOI":"10.1038\/nature08704","article-title":"Evidence for grid cells in a human memory network","volume":"463","author":"Doeller","year":"2010","journal-title":"Nature"},{"issue":"1","key":"2026101621153260800_bib18","doi-asserted-by":"publisher","first-page":"419","DOI":"10.1146\/annurev.neuro.27.070203.144152","article-title":"Neuronal circuits of the neocortex","volume":"27","author":"Douglas","year":"2004","journal-title":"Annual Review of Neuroscience"},{"issue":"7","key":"2026101621153260800_bib19","doi-asserted-by":"publisher","first-page":"1529","DOI":"10.1093\/cercor\/bhp213","article-title":"Projection from visual areas V2 and prostriata to caudal auditory cortex in the monkey","volume":"20","author":"Falchier","year":"2010","journal-title":"Cerebral Cortex"},{"issue":"1","key":"2026101621153260800_bib20","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1016\/j.tins.2010.10.002","article-title":"Development, specification, and diversity of callosal projection neurons","volume":"34","author":"Fame","year":"2011","journal-title":"Trends in Neurosciences"},{"issue":"1","key":"2026101621153260800_bib21","doi-asserted-by":"publisher","DOI":"10.1093\/cercor\/1.1.1","article-title":"Distributed hierarchical processing in the primate cerebral cortex","volume":"1","author":"Felleman","year":"1991","journal-title":"Cerebral Cortex"},{"key":"2026101621153260800_bib22","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.72573","article-title":"Columnar processing of border ownership in primate visual cortex","volume":"10","author":"Franken","year":"2021","journal-title":"eLife"},{"issue":"1","key":"2026101621153260800_bib23","doi-asserted-by":"publisher","first-page":"tgaa052","DOI":"10.1093\/texcom\/tgaa052","article-title":"In-depth characterization of layer 5 output neurons of the primary somatosensory cortex innervating the mouse dorsal spinal cord","volume":"1","author":"Frezel","year":"2020","journal-title":"Cerebral Cortex Communications"},{"issue":"4","key":"2026101621153260800_bib24","doi-asserted-by":"publisher","DOI":"10.1007\/BF00344251","article-title":"Neocognitron: A self-organizing neural network model for a mechanism of pattern recognition unaffected by shift in position","volume":"36","author":"Fukushima","year":"1980","journal-title":"Biological Cybernetics"},{"issue":"7","key":"2026101621153260800_bib25","doi-asserted-by":"publisher","first-page":"2432","DOI":"10.1523\/JNEUROSCI.09-07-02432.1989","article-title":"Columnar specificity of intrinsic horizontal and corticocortical connections in cat visual cortex","volume":"9","author":"Gilbert","year":"1989","journal-title":"Journal of Neuroscience"},{"issue":"2","key":"2026101621153260800_bib26","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.1977.sp011863","article-title":"Laminar differences in receptive field properties of cells in cat primary visual cortex","volume":"268","author":"Gilbert","year":"1977","journal-title":"Journal of Physiology"},{"key":"2026101621153260800_bib27","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-021-03783-x","article-title":"Locally ordered representation of 3D space in the entorhinal cortex","volume":"596","author":"Ginosar","year":"2021","journal-title":"Nature"},{"issue":"3882","key":"2026101621153260800_bib28","doi-asserted-by":"publisher","first-page":"917","DOI":"10.1126\/science.164.3882.917","article-title":"Organization of the visual pathways: Evidence is discussed concerning parallel pathways from the eye to the brain","volume":"164","author":"Glickstein","year":"1969","journal-title":"Science"},{"issue":"3","key":"2026101621153260800_bib29","doi-asserted-by":"crossref","first-page":"e30","DOI":"10.23915\/distill.00030","article-title":"Multimodal neurons in artificial neural networks","volume":"6","author":"Goh","year":"2021","journal-title":"Distill"},{"key":"2026101621153260800_bib30","doi-asserted-by":"publisher","DOI":"10.1038\/s41593-021-00907-4","article-title":"Irregular distribution of grid cell firing fields in rats exploring a 3D volumetric space","volume":"24","author":"Grieves","year":"2021","journal-title":"Nature Neuroscience"},{"key":"2026101621153260800_bib31","doi-asserted-by":"crossref","DOI":"10.3389\/fnsys.2021.650263","article-title":"A canonical laminar neocortical circuit whose bottom-up, horizontal, and top-down pathways control attention, learning, and prediction","volume":"15","author":"Grossberg","year":"2021","journal-title":"Frontiers in Systems Neuroscience"},{"key":"2026101621153260800_bib32","doi-asserted-by":"crossref","DOI":"10.1038\/nature03721","article-title":"Microstructure of a spatial map in the entorhinal cortex","author":"Hafting","year":"2005"},{"issue":"10","key":"2026101621153260800_bib33","doi-asserted-by":"publisher","first-page":"1575","DOI":"10.1016\/j.celrep.2015.08.003","article-title":"Multiple retinal axons converge onto relay cells in the adult mouse thalamus","volume":"12","author":"Hammer","year":"2015","journal-title":"Cell Reports"},{"issue":"7474","key":"2026101621153260800_bib34","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1038\/nature12654","article-title":"Cortical connectivityand sensory coding","volume":"503","author":"Harris","year":"2013","journal-title":"Nature"},{"key":"2026101621153260800_bib35","volume-title":"A thousand brains: A new theory of intelligence","author":"Hawkins","year":"2021"},{"key":"2026101621153260800_bib36","doi-asserted-by":"publisher","DOI":"10.3389\/fncir.2016.00023","article-title":"Why neurons have thousands of synapses: A theory of sequence memory in neocortex","volume":"10","author":"Hawkins","year":"2016","journal-title":"Frontiers in Neural Circuits"},{"issue":"October","key":"2026101621153260800_bib37","doi-asserted-by":"publisher","first-page":"1","DOI":"10.3389\/fncir.2017.00081","article-title":"A theory of how columns in the neocortex enable learning the structure of the world","volume":"11","author":"Hawkins","year":"2017","journal-title":"Frontiers in Neural Circuits"},{"key":"2026101621153260800_bib38","doi-asserted-by":"publisher","DOI":"10.3389\/fncir.2018.00121","article-title":"A framework for intelligence and cortical function based on grid cells in the neocortex","volume":"12","author":"Hawkins","year":"2019","journal-title":"Frontiers in Neural Circuits"},{"issue":"5","key":"2026101621153260800_bib39","doi-asserted-by":"crossref","first-page":"416","DOI":"10.1177\/1073858407305201","article-title":"Reappraising the functional implications of the primate visual anatomical hierarchy","volume":"13","author":"Hegd\u00e9","year":"2007","journal-title":"Neuroscientist"},{"key":"2026101621153260800_bib40","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-019-13996-4","article-title":"Word contexts enhance the neural representation of individual letters in early visual cortex","volume":"11","author":"Heilbron","year":"2020","journal-title":"Nature Communications"},{"issue":"2","key":"2026101621153260800_bib41","doi-asserted-by":"publisher","first-page":"955","DOI":"10.1007\/s00429-013-0694-4","article-title":"Possible anatomical pathways for short-latency multisensory integration processes in primary sensory cortices","volume":"220","author":"Henschke","year":"2015","journal-title":"Brain Structure and Function"},{"key":"2026101621153260800_bib42","doi-asserted-by":"publisher","first-page":"836498","DOI":"10.3389\/fncom.2022.836498","article-title":"Symmetry-based representations for artificial and biological general intelligence","volume":"16","author":"Higgins","year":"2022","journal-title":"Frontiers in Computational Neuroscience"},{"issue":"4","key":"2026101621153260800_bib43","doi-asserted-by":"publisher","first-page":"377","DOI":"10.1016\/j.conb.2006.06.014","article-title":"Laminar processing in the visual cortical column","volume":"16","author":"Hirsch","year":"2006","journal-title":"Current Opinion in Neurobiology"},{"issue":"5","key":"2026101621153260800_bib44","doi-asserted-by":"publisher","first-page":"1882","DOI":"10.1093\/cercor\/bhy036","article-title":"Subset of cortical layer 6b neurons selectively innervates higher order thalamic nuclei in mice","volume":"28","author":"Hoerder-Suabedissen","year":"2018","journal-title":"Cerebral Cortex"},{"issue":"3","key":"2026101621153260800_bib45","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.1959.sp006308","article-title":"Receptive fields of single neurones in the cat\u2019s striate cortex","volume":"148","author":"Hubel","year":"1959","journal-title":"Journal of Physiology"},{"issue":"1","key":"2026101621153260800_bib46","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.1962.sp006837","article-title":"Receptive fields, binocular interaction and functional architecture in the cat\u2019s visual cortex","volume":"160","author":"Hubel","year":"1962","journal-title":"Journal of Physiology"},{"issue":"31","key":"2026101621153260800_bib47","doi-asserted-by":"publisher","DOI":"10.3389\/fncom.2020.00031","article-title":"Contextual integration in cortical and convolutional neural networks","volume":"14","author":"Iyer","year":"2020","journal-title":"Frontiers in Computational Neuroscience"},{"issue":"9","key":"2026101621153260800_bib48","doi-asserted-by":"publisher","first-page":"1188","DOI":"10.1038\/nn.3466","article-title":"Direct recordings of grid-like neuronal activity in human spatial navigation","volume":"16","author":"Jacobs","year":"2013","journal-title":"Nature Neuroscience"},{"issue":"2","key":"2026101621153260800_bib49","doi-asserted-by":"publisher","DOI":"10.1038\/s41593-017-0049-1","article-title":"Human entorhinal cortex represents visual space using a boundary-anchored grid","volume":"21","author":"Julian","year":"2018","journal-title":"Nature Neuroscience"},{"issue":"29","key":"2026101621153260800_bib50","doi-asserted-by":"publisher","first-page":"9656","DOI":"10.1523\/JNEUROSCI.1325-14.2014","article-title":"Layer 6 corticothalamic neurons activate a cortical output layer, layer 5a","volume":"34","author":"Kim","year":"2014","journal-title":"Journal of Neuroscience"},{"key":"2026101621153260800_bib51","article-title":"Brain-like object recognition with high-performing shallow recurrent ANNs","volume-title":"Advances in neural information processing systems","author":"Kubilius","year":"2019"},{"issue":"6266","key":"2026101621153260800_bib52","doi-asserted-by":"publisher","DOI":"10.1126\/science.aab3050","article-title":"Human-level concept learning through probabilistic program induction","volume":"350","author":"Lake","year":"2015","journal-title":"Science"},{"issue":"8086","key":"2026101621153260800_bib53","doi-asserted-by":"publisher","first-page":"872","DOI":"10.1038\/s41586-025-09441-w","article-title":"A compressed hierarchy for visual form processing in the tree shrew","volume":"646","author":"Lanfranchi","year":"2025","journal-title":"Nature"},{"key":"2026101621153260800_bib54","author":"Leadholm","year":"2025","journal-title":"Thousand-brains systems: Sensorimotor intelligence for rapid, robust learning and inference"},{"key":"2026101621153260800_bib55","doi-asserted-by":"crossref","first-page":"436","DOI":"10.1038\/nature14539","article-title":"Deep learning","volume":"521","author":"Lecun","year":"2015","journal-title":"Nature"},{"issue":"11","key":"2026101621153260800_bib56","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1109\/5.726791","article-title":"Gradient-based learning applied to document recognition","volume":"86","author":"Lecun","year":"1998","journal-title":"Proceedings of the IEEE"},{"issue":"6","key":"2026101621153260800_bib57","doi-asserted-by":"publisher","first-page":"1420","DOI":"10.1016\/j.neuron.2017.08.036","article-title":"A sensorimotor circuit in mouse cortex for visual flow predictions","volume":"95","author":"Leinweber","year":"2017","journal-title":"Neuron"},{"key":"2026101621153260800_bib58","doi-asserted-by":"publisher","DOI":"10.3389\/fncir.2019.00022","article-title":"Locations in the neocortex: A theory of sensorimotor object recognition using cortical grid cells","volume":"13","author":"Lewis","year":"2019","journal-title":"Frontiers in Neural Circuits"},{"key":"2026101621153260800_bib59","doi-asserted-by":"publisher","first-page":"RP89025","DOI":"10.7554\/eLife.89025","article-title":"Distance and grid-like codes support the navigation of abstract social space in the human brain","volume":"12","author":"Liang","year":"2024","journal-title":"eLife"},{"key":"2026101621153260800_bib60","doi-asserted-by":"publisher","DOI":"10.1007\/s00221-013-3565-6","article-title":"Adapting to inversion of the visual field: A new twist on an old problem","volume":"228","author":"Lillicrap","year":"2013","journal-title":"Experimental Brain Research"},{"issue":"6","key":"2026101621153260800_bib61","doi-asserted-by":"publisher","first-page":"335","DOI":"10.1038\/s41583-020-0277-3","article-title":"Backpropagation and the brain","volume":"21","author":"Lillicrap","year":"2020","journal-title":"Nature Reviews Neuroscience"},{"issue":"12","key":"2026101621153260800_bib62","doi-asserted-by":"publisher","first-page":"1519","DOI":"10.1007\/s12264-022-00858-1","article-title":"The superior colliculus: Cell types, connectivity, and behavior","volume":"38","author":"Liu","year":"2022","journal-title":"Neuroscience Bulletin"},{"issue":"4853","key":"2026101621153260800_bib63","doi-asserted-by":"publisher","first-page":"740","DOI":"10.1126\/science.3283936","article-title":"Segregation of form, color, movement, and depth: Anatomy, physiology, and perception","volume":"240","author":"Livingstone","year":"1988","journal-title":"Science"},{"key":"2026101621153260800_bib64","article-title":"Object-centric learning with slot attention","author":"Locatello","year":"2020","journal-title":"Proceedings of the 34th Conference on Neural Information Processing Systems"},{"issue":"6","key":"2026101621153260800_bib65","doi-asserted-by":"publisher","first-page":"649","DOI":"10.1038\/s41422-020-00448-8","article-title":"A novel somatosensory spatial navigation system outside the hippocampal formation","volume":"31","author":"Long","year":"2021","journal-title":"Cell Research"},{"issue":"1\u20132","key":"2026101621153260800_bib66","doi-asserted-by":"publisher","first-page":"119","DOI":"10.1016\/j.brainresrev.2010.12.002","article-title":"The thalamocortical vestibular system in animals and humans","volume":"67","author":"Lopez","year":"2011","journal-title":"Brain Research Reviews"},{"issue":"1","key":"2026101621153260800_bib67","doi-asserted-by":"publisher","first-page":"225","DOI":"10.1002\/cne.23458","article-title":"Anatomy of hierarchy: Feedforward and feedback pathways in macaque visual cortex","volume":"522","author":"Markov","year":"2014","journal-title":"Journal of Comparative Neurology"},{"key":"2026101621153260800_bib68","doi-asserted-by":"crossref","DOI":"10.1038\/nrn1519","article-title":"Interneurons of the neocortical inhibitory system","author":"Markram","year":"2004","journal-title":"Nature Reviews Neuroscience"},{"issue":"C","key":"2026101621153260800_bib69","doi-asserted-by":"crossref","DOI":"10.1016\/S0079-7421(08)60536-8","article-title":"Catastrophic interference in connectionist networks: The sequential learning problem","volume":"24","author":"McCloskey","year":"1989","journal-title":"Psychology of Learning and Motivation-Advances in Research and Theory"},{"issue":"4","key":"2026101621153260800_bib70","doi-asserted-by":"publisher","first-page":"408","DOI":"10.1152\/jn.1957.20.4.408","article-title":"Modality and topographic properties of single neurons of cat\u2019s somatic sensory cortex","volume":"20","author":"Mountcastle","year":"1957","journal-title":"Journal of Neurophysiology"},{"issue":"4","key":"2026101621153260800_bib71","doi-asserted-by":"publisher","DOI":"10.1093\/brain\/120.4.701","article-title":"The columnar organization of the neocortex","volume":"120","author":"Mountcastle","year":"1997","journal-title":"Brain"},{"key":"2026101621153260800_bib72","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1016\/S0079-6123(01)34005-0","article-title":"Thalamic relay functions","volume-title":"Vision: From neurons to cognition","author":"Murray Sherman","year":"2001"},{"issue":"36","key":"2026101621153260800_bib73","doi-asserted-by":"publisher","first-page":"14342","DOI":"10.1523\/JNEUROSCI.2275-13.2013","article-title":"A circuit for motor cortical modulation of auditory cortical activity","volume":"33","author":"Nelson","year":"2013","journal-title":"Journal of Neuroscience"},{"issue":"10","key":"2026101621153260800_bib74","doi-asserted-by":"publisher","first-page":"2375","DOI":"10.1093\/cercor\/bhr317","article-title":"Cell type-specificc three-dimensional structure of thalamocortical circuits in a column of rat vibrissal cortex","volume":"22","author":"Oberlaender","year":"2012","journal-title":"Cerebral Cortex"},{"issue":"11","key":"2026101621153260800_bib75","doi-asserted-by":"crossref","first-page":"e7","DOI":"10.23915\/distill.00007","article-title":"Feature visualization","volume":"2","author":"Olah","year":"2017","journal-title":"Distill"},{"key":"2026101621153260800_bib76","author":"Ollikka","year":"2024","journal-title":"A comparison between humans and AI at recognizing objects in unusual poses."},{"issue":"9","key":"2026101621153260800_bib77","doi-asserted-by":"publisher","first-page":"1292","DOI":"10.1038\/s41593-021-00916-3","article-title":"Inferences on a multidimensional social hierarchy use a grid-like code","volume":"24","author":"Park","year":"2021","journal-title":"Nature Neuroscience"},{"issue":"5","key":"2026101621153260800_bib78","doi-asserted-by":"publisher","DOI":"10.1152\/jn.1999.82.5.2490","article-title":"Responses to contour features in macaque area V4","volume":"82","author":"Pasupathy","year":"1999","journal-title":"Journal of Neurophysiology"},{"issue":"3","key":"2026101621153260800_bib79","doi-asserted-by":"publisher","first-page":"283","DOI":"10.1017\/S0952523800006283","article-title":"Complex transcallosal interactions in visual cortex","volume":"6","author":"Payne","year":"1991","journal-title":"Visual Neuroscience"},{"issue":"10","key":"2026101621153260800_bib80","doi-asserted-by":"publisher","first-page":"2092","DOI":"10.1038\/s41593-025-02054-6","article-title":"Grid cells accurately track movement during path integration-based navigation despite switching reference frames","volume":"28","author":"Peng","year":"2025","journal-title":"Nature Neuroscience"},{"key":"2026101621153260800_bib81","article-title":"Towards biologically plausible convolutional networks","author":"Pogodin","year":"2021","journal-title":"Proceedings of the 35th Conference on Neural Information Processing Systems"},{"issue":"435","key":"2026101621153260800_bib82","first-page":"7045","article-title":"Invariant visual representation by single neurons in the human brain","volume":"2005","author":"Quiroga","year":"2005","journal-title":"Nature"},{"issue":"7","key":"2026101621153260800_bib83","doi-asserted-by":"publisher","first-page":"1221","DOI":"10.1038\/s41593-024-01673-9","article-title":"A sensory\u2013motor theory of the neocortex","volume":"27","author":"Rao","year":"2024","journal-title":"Nature Neuroscience"},{"issue":"1","key":"2026101621153260800_bib84","doi-asserted-by":"publisher","first-page":"79","DOI":"10.1038\/4580","article-title":"Predictive coding in the visual cortex: A functional interpretation of some extra-classical receptive-field effects","volume":"2","author":"Rao","year":"1999","journal-title":"Nature Neuroscience"},{"issue":"11","key":"2026101621153260800_bib85","doi-asserted-by":"publisher","DOI":"10.1038\/14819","article-title":"Hierarchical models of object recognition in cortex","volume":"2","author":"Riesenhuber","year":"1999","journal-title":"Nature Neuroscience"},{"key":"2026101621153260800_bib86","doi-asserted-by":"publisher","first-page":"772","DOI":"10.1016\/j.neuroimage.2017.07.032","article-title":"What do we know about laminar connectivity?","volume":"197","author":"Rockland","year":"2019","journal-title":"NeuroImage"},{"issue":"4","key":"2026101621153260800_bib87","doi-asserted-by":"crossref","first-page":"529","DOI":"10.1002\/(SICI)1096-9861(19960930)373:4<529::AID-CNE5>3.0.CO;2-3","article-title":"Collateralized divergent feedback connections that target multiple cortical areas","volume":"373","author":"Rockland","year":"1996","journal-title":"Journal of Comparative Neurology"},{"issue":"1\u20132","key":"2026101621153260800_bib88","doi-asserted-by":"publisher","first-page":"19","DOI":"10.1016\/S0167-8760(03)00121-1","article-title":"Multisensory convergence in calcarine visual areas in macaque monkey","volume":"50","author":"Rockland","year":"2003","journal-title":"International Journal of Psychophysiology"},{"issue":"1","key":"2026101621153260800_bib89","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1016\/0006-8993(79)90485-2","article-title":"Laminar origins and terminations of cortical connections of the occipital lobe in the rhesus monkey","volume":"179","author":"Rockland","year":"1979","journal-title":"Brain Research"},{"issue":"1","key":"2026101621153260800_bib90","doi-asserted-by":"publisher","first-page":"54","DOI":"10.1002\/cne.902850106","article-title":"Terminal arbors of individual \u201cfeedback\u201d axons projecting from area V2 to V1 in the macaque monkey: A study using immunohistochemistry of anterogradely transported Phaseolus vulgaris leucoagglutinin","volume":"285","author":"Rockland","year":"1989","journal-title":"Journal of Comparative Neurology"},{"key":"2026101621153260800_bib91","doi-asserted-by":"crossref","DOI":"10.1093\/acprof:oso\/9780198784852.001.0001","volume-title":"Cerebral cortex: Principles of operation","author":"Rolls","year":"2016"},{"key":"2026101621153260800_bib92","doi-asserted-by":"publisher","DOI":"10.1007\/s00429-021-02347-z","article-title":"The connections of neocortical pyramidal cells can implement the learning of new categories, attractor memory, and top\u2013down recall and attention","volume":"226","author":"Rolls","year":"2021","journal-title":"Brain Structure and Function"},{"key":"2026101621153260800_bib93","doi-asserted-by":"publisher","DOI":"10.1080\/713756803","article-title":"Orientation constancy in neurons of monkey visual cortex","volume":"6","author":"Sauvan","year":"1999","journal-title":"Visual Cognition"},{"key":"2026101621153260800_bib94","doi-asserted-by":"publisher","first-page":"32916","DOI":"10.52202\/079017-1036","article-title":"The surprising ineffectiveness of pre-trained visual representations for model-based reinforcement learning","volume-title":"Advances in neural information processing systems","author":"Schneider","year":"2024"},{"key":"2026101621153260800_bib95","doi-asserted-by":"publisher","DOI":"10.1101\/407007","author":"Schrimpf","year":"2018","journal-title":"Brain-score: Which artificial neural network for object recognition is most brain-like?"},{"issue":"1","key":"2026101621153260800_bib96","doi-asserted-by":"publisher","first-page":"221","DOI":"10.1146\/annurev-neuro-100520-012117","article-title":"Neocortical Layer 1: An elegant solution to top-down and bottom-up integration","volume":"44","author":"Schuman","year":"2021","journal-title":"Annual Review of Neuroscience"},{"issue":"36","key":"2026101621153260800_bib97","doi-asserted-by":"publisher","first-page":"9117","DOI":"10.1523\/JNEUROSCI.1253-06.2006","article-title":"The role of feedback in shaping the extra-classical receptive field of cortical neurons: A recurrent network model","volume":"26","author":"Schwabe","year":"2006","journal-title":"Journal of Neuroscience"},{"key":"2026101621153260800_bib98","article-title":"Assessment of ocular counter roll during head tilt using binocular video oculography","volume":"4","author":"Schworm","year":"2002","journal-title":"Investigative Ophthalmology and Visual Science"},{"key":"2026101621153260800_bib99","doi-asserted-by":"crossref","DOI":"10.1109\/TPAMI.2007.56","article-title":"Robust object recognition with cortex-like mechanisms","volume-title":"IEEE Transactions on Pattern Analysis and Machine Intelligence","author":"Serre","year":"2007"},{"key":"2026101621153260800_bib100","doi-asserted-by":"publisher","first-page":"107","DOI":"10.1016\/S0079-6123(05)49009-3","article-title":"Thalamic relays and cortical functioning","volume":"149","author":"Sherman","year":"2005","journal-title":"Progress in Brain Research"},{"key":"2026101621153260800_bib101","doi-asserted-by":"crossref","DOI":"10.7551\/mitpress\/9780262019309.001.0001","author":"Sherman","year":"2013","journal-title":"Functional connections of cortical areas: A new view from the thalamus"},{"issue":"12","key":"2026101621153260800_bib102","doi-asserted-by":"publisher","first-page":"R443","DOI":"10.1016\/j.cub.2007.03.044","article-title":"Structure and function of the cerebral cortex","volume":"17","author":"Shipp","year":"2007","journal-title":"Current Biology"},{"issue":"4","key":"2026101621153260800_bib103","doi-asserted-by":"publisher","first-page":"1020","DOI":"10.1093\/cercor\/bht292","article-title":"Decoding visual object categories in early somatosensory cortex","volume":"25","author":"Smith","year":"2015","journal-title":"Cerebral Cortex"},{"issue":"18","key":"2026101621153260800_bib104","doi-asserted-by":"publisher","first-page":"3679","DOI":"10.1002\/cne.22411","article-title":"Evaluation of inputs to rat primary auditory cortex from the suprageniculate nucleus and extrastriate visual cortex","volume":"518","author":"Smith","year":"2010","journal-title":"Journal of Comparative Neurology"},{"issue":"1","key":"2026101621153260800_bib105","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1207\/s15516709cog1401_3","article-title":"Principles of object perception","volume":"14","author":"Spelke","year":"1990","journal-title":"Cognitive Science"},{"issue":"4","key":"2026101621153260800_bib106","doi-asserted-by":"publisher","first-page":"739","DOI":"10.1016\/S0896-6273(02)01029-2","article-title":"Lateral connectivity and contextual interactions in macaque primary visual cortex","volume":"36","author":"Stettler","year":"2002","journal-title":"Neuron"},{"key":"2026101621153260800_bib107","doi-asserted-by":"publisher","DOI":"10.1037\/h0075482","article-title":"Vision without inversion of the retinal image","volume":"4","author":"Stratton","year":"1897","journal-title":"Psychological Review"},{"issue":"12","key":"2026101621153260800_bib108","doi-asserted-by":"publisher","first-page":"778","DOI":"10.1038\/s41583-023-00756-z","article-title":"How deep is the brain? The shallow brain hypothesis","volume":"24","author":"Suzuki","year":"2023","journal-title":"Nature Reviews Neuroscience."},{"key":"2026101621153260800_bib109","article-title":"Inferotemporal cortex and object vision","volume":"19","author":"Tanaka","year":"1996","journal-title":"Annual Review of Neuro-science"},{"issue":"1","key":"2026101621153260800_bib110","doi-asserted-by":"publisher","DOI":"10.1152\/jn.1991.66.1.170","article-title":"Coding visual images of objects in the inferotemporal cortex of the macaque monkey","volume":"66","author":"Tanaka","year":"1991","journal-title":"Journal of Neurophysiology"},{"key":"2026101621153260800_bib111","doi-asserted-by":"publisher","DOI":"10.3389\/fnana.2010.00013","author":"Thomson","year":"2010","journal-title":"Neocortical layer 6, a review"},{"issue":"1","key":"2026101621153260800_bib112","doi-asserted-by":"publisher","first-page":"105","DOI":"10.1016\/S0896-6273(00)80826-0","article-title":"Solutions to the binding problem: Progress through controversy and convergence","volume":"24","author":"Treisman","year":"1999","journal-title":"Neuron"},{"issue":"3","key":"2026101621153260800_bib113","doi-asserted-by":"publisher","first-page":"640","DOI":"10.1002\/cne.24423","article-title":"Corticofugal circuits: Communication lines from the cortex to the rest of the brain","volume":"527","author":"Usrey","year":"2019","journal-title":"Journal of Comparative Neurology"},{"key":"2026101621153260800_bib117","doi-asserted-by":"crossref","DOI":"10.1016\/j.neuron.2018.02.023","article-title":"A circuit for integration of head visual-motion signals in layer 6 of mouse primary visual cortex","volume":"98","author":"V\u00e9lez-Fort","year":"2018","journal-title":"Neuron"},{"issue":"3","key":"2026101621153260800_bib114","doi-asserted-by":"publisher","first-page":"277","DOI":"10.1016\/j.pneurobio.2010.05.001","article-title":"A modeler\u2019s view on the spatial structure of intrinsic horizontal connectivity in the neocortex","volume":"92","author":"Voges","year":"2010","journal-title":"Progress in Neurobiology"},{"key":"2026101621153260800_bib115","doi-asserted-by":"publisher","first-page":"1136987","DOI":"10.3389\/fcomp.2023.1136987","article-title":"Visual cortical processing\u2014from image to object representation","volume":"5","author":"von der Heydt","year":"2023","journal-title":"Frontiers in Computer Science"},{"issue":"1","key":"2026101621153260800_bib116","doi-asserted-by":"publisher","first-page":"95","DOI":"10.1016\/S0896-6273(00)80825-9","article-title":"The what and why of binding: The modeler\u2019s perspective","volume":"24","author":"Von Der Malsburg","year":"1999","journal-title":"Neuron"},{"key":"2026101621153260800_bib118","article-title":"Retinal afferents synapse with relay cells targeting the middle temporal area in the pulvinar and lateral geniculate nuclei","volume":"4","author":"Warner","year":"2010","journal-title":"Frontiers in Neuroanatomy"},{"issue":"23","key":"2026101621153260800_bib119","article-title":"Performance-optimized hierarchical models predict neural responses in higher visual cortex","volume":"111","author":"Yamins","year":"2014","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"key":"2026101621153260800_bib120","author":"Yu","year":"2025","journal-title":"A parietal grid-like code rotates with cognitive maps but lags rapid behavioral transfer"},{"issue":"17","key":"2026101621153260800_bib121","doi-asserted-by":"publisher","first-page":"6594","DOI":"10.1523\/JNEUROSCI.20-17-06594.2000","article-title":"Coding of border ownership in monkey visual cortex","volume":"20","author":"Zhou","year":"2000","journal-title":"Journal of Neuroscience"},{"key":"2026101621153260800_bib122","author":"Zimmermann","year":"2023","journal-title":"Sensitivity of slot-based object-centric models to their number of slots"},{"issue":"4","key":"2026101621153260800_bib123","doi-asserted-by":"publisher","first-page":"565","DOI":"10.1016\/j.neuron.2025.11.024","article-title":"The layer 6b theory of attention","volume":"114","author":"Zolnik","year":"2026","journal-title":"Neuron"}],"container-title":["Neural Computation"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/direct.mit.edu\/neco\/article-pdf\/38\/11\/2042\/2629850\/neco.a.1579.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/direct.mit.edu\/neco\/article-pdf\/38\/11\/2042\/2629850\/neco.a.1579.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,17]],"date-time":"2026-10-17T01:15:42Z","timestamp":1792199742000},"score":1,"resource":{"primary":{"URL":"https:\/\/direct.mit.edu\/neco\/article\/38\/11\/2042\/139102\/The-Thousand-Brains-Theory-2-0-An-Extension-for"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10,16]]},"references-count":123,"journal-issue":{"issue":"11","published-online":{"date-parts":[[2026,10,16]]},"published-print":{"date-parts":[[2026,10,16]]}},"URL":"https:\/\/doi.org\/10.1162\/neco.a.1579","relation":{},"ISSN":["0899-7667","1530-888X"],"issn-type":[{"value":"0899-7667","type":"print"},{"value":"1530-888X","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2026,11]]},"published":{"date-parts":[[2026,10,16]]}}}