{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T20:49:23Z","timestamp":1791492563586,"version":"4.3.4"},"reference-count":133,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T00:00:00Z","timestamp":1791417600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T00:00:00Z","timestamp":1791417600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Cogn Neurodyn"],"published-print":{"date-parts":[[2026,12]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Neuroscience has long informed the development of artificial neural networks, but the success of modern architectures invites, in turn, the converse: can modern networks teach us lessons about brain function? Here, we examine the structure of the cortical column and propose that the transformer provides a natural computational analogy for multiple elements of cortical microcircuit organization. Rather than claiming a literal implementation of transformer equations in cortex, we develop a hypothetical mapping between transformer operations and laminar cortical features, using the analogy as an orienting framework for analysis and discussion. This mapping allows us to examine in greater depth how contextual selection, content routing, recurrent integration, and interlaminar transformations may be distributed across cortical circuitry. In doing so, we generate a broad set of predictions and experimentally testable hypotheses concerning laminar specialization, contextual modulation, dendritic integration, oscillatory coordination, and the effective connectivity of cortical columns. This proposal is intended as a structured hypothesis rather than a definitive account of cortical computation. Placing transformer operations and cortical architectonics into a common descriptive framework sharpens questions, reveals new functional correspondences, and opens a productive route for reciprocal exchange between systems neuroscience and modern AI. More broadly, this perspective suggests that comparing brains and architectures at the level of computational organization can yield genuine insight into both.<\/jats:p>","DOI":"10.1007\/s11571-026-10566-5","type":"journal-article","created":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T07:16:18Z","timestamp":1791443778000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["The neuroscience of transformers"],"prefix":"10.1007","volume":"20","author":[{"given":"Peter","family":"K\u00f6nig","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8527-4259","authenticated-orcid":false,"given":"Mario","family":"Negrello","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,10,8]]},"reference":[{"issue":"1","key":"10566_CR1","doi-asserted-by":"publisher","first-page":"39","DOI":"10.1002\/cne.903410105","volume":"341","author":"B Ahmed","year":"1994","unstructured":"Ahmed B, Anderson JC, Douglas RJ, Martin KAC, Nelson JC (1994) Polyneuronal innervation of spiny stellate neurons in cat visual cortex. J Comp Neurol 341(1):39\u201349. https:\/\/doi.org\/10.1002\/cne.903410105","journal-title":"J Comp Neurol"},{"key":"10566_CR2","doi-asserted-by":"publisher","first-page":"93","DOI":"10.1016\/S0079-6123(06)54005-1","volume":"154","author":"A Angelucci","year":"2006","unstructured":"Angelucci A, Bressloff PC (2006) Contribution of feedforward, lateral and feedback connections to the classical receptive field center and extra-classical receptive field surround of primate v1 neurons. Prog Brain Res 154:93\u2013120","journal-title":"Prog Brain Res"},{"key":"10566_CR3","unstructured":"Bahdanau D, Cho K, Bengio Y (2015) Neural machine translation by jointly learning to align and translate. ICLR"},{"key":"10566_CR4","doi-asserted-by":"publisher","first-page":"566","DOI":"10.1038\/s41583-018-0038-8","volume":"19","author":"DS Bassett","year":"2018","unstructured":"Bassett DS, Zurn P, Gold JI (2018) On the nature and use of models in network neuroscience. Nat Rev Neurosci 19:566\u2013578","journal-title":"Nat Rev Neurosci"},{"issue":"49","key":"10566_CR5","doi-asserted-by":"publisher","first-page":"31459","DOI":"10.1073\/pnas.2014868117","volume":"117","author":"AM Bastos","year":"2020","unstructured":"Bastos AM, Lundqvist M, Waite AS, Kopell N, Miller EK (2020) Layer and rhythm specificity for predictive routing. Proc Natl Acad Sci 117(49):31459\u201331469. https:\/\/doi.org\/10.1073\/pnas.2014868117","journal-title":"Proc Natl Acad Sci"},{"issue":"4","key":"10566_CR6","doi-asserted-by":"publisher","first-page":"695","DOI":"10.1016\/j.neuron.2012.10.038","volume":"76","author":"AM Bastos","year":"2012","unstructured":"Bastos AM, Usrey WM, Adams RA, Mangun GR, Fries P, Friston KJ (2012) Canonical microcircuits for predictive coding. Neuron 76(4):695\u2013711","journal-title":"Neuron"},{"issue":"2","key":"10566_CR7","doi-asserted-by":"publisher","first-page":"390","DOI":"10.1016\/j.neuron.2014.12.018","volume":"85","author":"AM Bastos","year":"2015","unstructured":"Bastos AM, Vezoli J, Fries P (2015) Visual areas exert feedforward and feedback influences through distinct frequency channels. Neuron 85(2):390\u2013401. https:\/\/doi.org\/10.1016\/j.neuron.2014.12.018","journal-title":"Neuron"},{"key":"10566_CR8","doi-asserted-by":"crossref","unstructured":"Bender EM, Gebru T, McMillan-Major A, Shmitchell S (2021) On the dangers of stochastic parrots: can language models be too big? facct. InProceedings of the 2021 ACM conference on fairness, accountability, and transparency. pp. 610-623.","DOI":"10.1145\/3442188.3445922"},{"issue":"6013","key":"10566_CR9","doi-asserted-by":"publisher","first-page":"83","DOI":"10.1126\/science.1195870","volume":"331","author":"P Berkes","year":"2011","unstructured":"Berkes P, Orb\u00e1n G, Lengyel M, Fiser J (2011) Spontaneous cortical activity reveals hallmarks of an optimal internal model of the environment. Science 331(6013):83\u201387","journal-title":"Science"},{"issue":"1","key":"10566_CR10","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1007\/s00422-003-0434-6","volume":"90","author":"BY Betsch","year":"2004","unstructured":"Betsch BY, Einh\u00e4user W, K\u00f6rding KP, K\u00f6nig P (2004) The world from a cat\u2019s perspective: Statistics of natural videos. Biol Cybern 90(1):41\u201350. https:\/\/doi.org\/10.1007\/s00422-003-0434-6","journal-title":"Biol Cybern"},{"issue":"24","key":"10566_CR11","doi-asserted-by":"publisher","first-page":"10464","DOI":"10.1523\/JNEUROSCI.18-24-10464.1998","volume":"18","author":"G-Q Bi","year":"1998","unstructured":"Bi G-Q, Poo M-M (1998) Synaptic modifications in cultured hippocampal neurons: Dependence on spike timing, synaptic strength, and postsynaptic cell type. J Neurosci 18(24):10464\u201310472","journal-title":"J Neurosci"},{"issue":"39","key":"10566_CR12","doi-asserted-by":"publisher","first-page":"8441","DOI":"10.1523\/JNEUROSCI.1400-04.2004","volume":"24","author":"T Binzegger","year":"2004","unstructured":"Binzegger T, Douglas RJ, Martin KAC (2004) A quantitative map of the circuit of cat primary visual cortex. J Neurosci 24(39):8441\u20138453","journal-title":"J Neurosci"},{"key":"10566_CR13","unstructured":"Bommasani R, Hudson DA, Adeli E, Altman R, Arora S, Von Arx S, Bernstein MS, Bohg J, Bosselut A, Brunskill E, Brynjolfsson E, Buch S, Card D, Castellon R, Chatterji N, Chen A, Creel K, Davis JQ, Demszky D, Donahue C, Doumbouya M, Durmus E, Ermon S, Etchemendy J, Ethayarajh K, Fei-Fei L, Finn C, Gale T, Gillespie LE, Goel K, Goodman N, Grossman S, Guha N, Hashimoto T, Henderson P, Hewitt J, Ho DE, Hong J, Jurafsky D, Khan A, Khattab O, Koh PW, Krass M, Krishna R, Kuditipudi R, Kumar A, Ladhak F, Lee M, Lee T, Leskovec J, Levent I, Li XL, Ma T, Malik A, Manning CD, Mirchandani S, Mitchell E, Munyikwa Z, Nair S, Narayanan D, Newman B, Nie A, Niebles JC, Nilforoshan H, Nyarko JF, Ogut G, Orr L, Papadimitriou I, Park JS, Piech C, Portelance E, Potts C, Raghunathan A, Reich R, Ren H, Rong F, Roohani Y, Ruiz C, Ryan J, Sadigh D, Sagawa S, Santhanam K, Shih A, Srinivasan K, Tamkin A, Taori R, Thomas AW, Tramer F, Wang RE, Wang W, Wu B, Wu J, Wu Y, Xie SM, Yasunaga M, You J, Zaharia M, Zhang M, Zhang T, Zhang X, Zhang Y, Zheng L, Zhou K, Liang P (2021) On the opportunities and risks of foundation models. arXiv:2108.07258"},{"key":"10566_CR14","first-page":"1877","volume":"33","author":"TB Brown","year":"2020","unstructured":"Brown TB, Mann B, Ryder N, Subbiah M, Kaplan J, Dhariwal P, Neelakantan A, Shyam P, Sastry G, Askell A, Agarwal S, Herbert-Voss A, Krueger G, Henighan T, Child R, Ramesh A, Ziegler D, Wu J, Winter C, Hesse C, Chen M, Sigler E, Litwin M, Gray S, Chess B, Clark J, Berner C, McCandlish S, Radford A, Sutskever I, Amodei D (2020) Language models are few-shot learners. In NeurIPS 33:1877\u20131901","journal-title":"In NeurIPS"},{"key":"10566_CR15","doi-asserted-by":"publisher","first-page":"25","DOI":"10.1146\/annurev.neuro.31.060407.125639","volume":"31","author":"N Caporale","year":"2008","unstructured":"Caporale N, Dan Y (2008) Spike timing\u2013dependent plasticity: a hebbian learning rule. Annu Rev Neurosci 31:25\u201346","journal-title":"Annu Rev Neurosci"},{"issue":"1","key":"10566_CR16","doi-asserted-by":"publisher","first-page":"54","DOI":"10.1016\/S0734-189X(87)80014-2","volume":"37","author":"GA Carpenter","year":"1987","unstructured":"Carpenter GA, Grossberg S (1987) A massively parallel architecture for a self-organizing neural pattern recognition machine. Comp Vision Graph Image Process 37(1):54\u2013115","journal-title":"Comp Vision Graph Image Process"},{"key":"10566_CR17","doi-asserted-by":"publisher","first-page":"134","DOI":"10.1038\/s42003-022-03036-1","volume":"5","author":"C Caucheteux","year":"2022","unstructured":"Caucheteux C, Gramfort A, King J-R (2022) Brains and algorithms partially converge in natural language processing. Comms Biol 5:134","journal-title":"Comms Biol"},{"key":"10566_CR18","unstructured":"Choromanski K, Likhosherstov V, Dohan D, Song X, Gane A, Sarlos T, Hawkins P, Davis JQ, Mohiuddin A, Kaiser L, Belanger D, Colwell L, Weller A (2021) Rethinking attention with performers. ICLR"},{"issue":"7539","key":"10566_CR19","doi-asserted-by":"publisher","first-page":"399","DOI":"10.1038\/nature14182","volume":"518","author":"L Cossell","year":"2015","unstructured":"Cossell L, Iacaruso MF, Muir DR, Houlton R, Sader EN, Ko H, Hofer SB, Mrsic-Flogel TD (2015) Functional organization of excitatory synaptic strength in primary visual cortex. Nature 518(7539):399\u2013403. https:\/\/doi.org\/10.1038\/nature14182","journal-title":"Nature"},{"issue":"6203","key":"10566_CR20","doi-asserted-by":"publisher","first-page":"129","DOI":"10.1038\/337129a0","volume":"337","author":"F Crick","year":"1989","unstructured":"Crick F (1989) The recent excitement about neural networks. Nature 337(6203):129\u2013132","journal-title":"Nature"},{"issue":"6737","key":"10566_CR21","doi-asserted-by":"publisher","first-page":"655","DOI":"10.1038\/21371","volume":"399","author":"A Das","year":"1999","unstructured":"Das A, Gilbert CD (1999) Topography of bounding contour orientation and spatial affinity in primary visual cortex. Nature 399(6737):655\u2013661","journal-title":"Nature"},{"key":"10566_CR22","volume-title":"Theoretical Neuroscience: Computational and Mathematical Modeling of Neural Systems","author":"P Dayan","year":"2001","unstructured":"Dayan P, Abbott LF (2001) Theoretical Neuroscience: Computational and Mathematical Modeling of Neural Systems. MIT Press"},{"issue":"10","key":"10566_CR23","doi-asserted-by":"publisher","first-page":"4046","DOI":"10.1523\/JNEUROSCI.19-10-04046.1999","volume":"19","author":"GC DeAngelis","year":"1999","unstructured":"DeAngelis GC, Ghose GM, Ohzawa I, Freeman RD (1999) Functional micro-organization of primary visual cortex: receptive field analysis of nearby neurons. J Neurosci 19(10):4046\u20134064","journal-title":"J Neurosci"},{"key":"10566_CR24","unstructured":"Dehghani M, Gouws S, Vinyals O, Uszkoreit J, Kaiser L (2019) Universal transformers. In International Conference on Learning Representations. URL https:\/\/arxiv.org\/abs\/1807.03819"},{"key":"10566_CR25","unstructured":"Dosovitskiy A, Beyer L, Kolesnikov A, Weissenborn D, Zhai X, Unterthiner T, Dehghani M, Minderer M, Heigold G, Gelly S, Uszkoreit J, Houlsby N (2021) An image is worth 16x16 words: Transformers for image recognition at scale. ICLR"},{"key":"10566_CR26","doi-asserted-by":"publisher","first-page":"419","DOI":"10.1146\/annurev.neuro.27.070203.144152","volume":"27","author":"RJ Douglas","year":"2004","unstructured":"Douglas RJ, Martin KAC (2004) Neuronal circuits of the neocortex. Annu Rev Neurosci 27:419\u2013451","journal-title":"Annu Rev Neurosci"},{"issue":"13","key":"10566_CR27","doi-asserted-by":"publisher","first-page":"R496","DOI":"10.1016\/j.cub.2007.04.024","volume":"17","author":"RJ Douglas","year":"2007","unstructured":"Douglas RJ, Martin KAC (2007) Recurrent neuronal circuits in the neocortex. Curr Biol 17(13):R496\u2013R500","journal-title":"Curr Biol"},{"key":"10566_CR28","doi-asserted-by":"publisher","first-page":"475","DOI":"10.1046\/j.0953-816x.2001.01885.x","volume":"15","author":"W Einh\u00e4user","year":"2002","unstructured":"Einh\u00e4user W, Kayser C, K\u00f6nig P, K\u00f6rding KP (2002) Learning the invariance properties of complex cells from their responses to natural stimuli. Eur J Neurosci 15:475\u2013486","journal-title":"Eur J Neurosci"},{"issue":"2","key":"10566_CR29","doi-asserted-by":"publisher","first-page":"179","DOI":"10.1207\/s15516709cog1402_1","volume":"14","author":"JL Elman","year":"1990","unstructured":"Elman JL (1990) Finding structure in time. Cogn Sci 14(2):179\u2013211","journal-title":"Cogn Sci"},{"issue":"1","key":"10566_CR30","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1093\/cercor\/1.1.1","volume":"1","author":"DJ Felleman","year":"1991","unstructured":"Felleman DJ, Van Essen DC (1991) Distributed hierarchical processing in the primate cerebral cortex. Cereb Cortex 1(1):1\u201347","journal-title":"Cereb Cortex"},{"issue":"4","key":"10566_CR31","doi-asserted-by":"publisher","first-page":"438","DOI":"10.1016\/S0959-4388(00)00113-6","volume":"10","author":"D Fitzpatrick","year":"2000","unstructured":"Fitzpatrick D (2000) Seeing beyond the receptive field in primary visual cortex. Curr Opin Neurobiol 10(4):438\u2013443","journal-title":"Curr Opin Neurobiol"},{"issue":"1","key":"10566_CR32","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1162\/0898929052880093","volume":"17","author":"MJ Frank","year":"2005","unstructured":"Frank MJ (2005) Dynamic dopamine modulation in the basal ganglia: a neurocomputational account of cognitive deficits in medicated and nonmedicated parkinsonism. J Cogn Neurosci 17(1):51\u201372","journal-title":"J Cogn Neurosci"},{"issue":"1","key":"10566_CR33","doi-asserted-by":"publisher","first-page":"220","DOI":"10.1016\/j.neuron.2015.09.034","volume":"88","author":"P Fries","year":"2015","unstructured":"Fries P (2015) Rhythms for cognition: Communication through coherence. Neuron 88(1):220\u2013235","journal-title":"Neuron"},{"key":"10566_CR34","doi-asserted-by":"publisher","first-page":"193","DOI":"10.1007\/BF00344251","volume":"36","author":"K Fukushima","year":"1980","unstructured":"Fukushima K (1980) Neocognitron: a self-organizing neural network model for a mechanism of pattern recognition unaffected by shift in position. Biol Cybern 36:193\u2013202","journal-title":"Biol Cybern"},{"key":"10566_CR35","doi-asserted-by":"publisher","DOI":"10.1017\/CBO9780511815706","volume-title":"Spiking Neuron Models: Single Neurons, Populations","author":"W Gerstner","year":"2002","unstructured":"Gerstner W, Kistler WM (2002) Spiking Neuron Models: Single Neurons, Populations. Cambridge University Press, Plasticity"},{"key":"10566_CR36","doi-asserted-by":"publisher","DOI":"10.1017\/CBO9781107447615","volume-title":"Neuronal Dynamics: From Single Neurons to Networks and Models of Cognition","author":"W Gerstner","year":"2014","unstructured":"Gerstner W, Kistler WM, Naud R, Paninski L (2014) Neuronal Dynamics: From Single Neurons to Networks and Models of Cognition. Cambridge University Press"},{"key":"10566_CR37","doi-asserted-by":"crossref","unstructured":"Geva M, Schuster R, Berant J, Levy O (2021) Transformer feed-forward layers are key\u2013value memories. EMNLP, pp 5484\u20135495","DOI":"10.18653\/v1\/2021.emnlp-main.446"},{"issue":"5","key":"10566_CR38","doi-asserted-by":"publisher","first-page":"1116","DOI":"10.1523\/JNEUROSCI.03-05-01116.1983","volume":"3","author":"CD Gilbert","year":"1983","unstructured":"Gilbert CD, Wiesel TN (1983) Clustered intrinsic connections in cat visual cortex. J Neurosci 3(5):1116\u20131133","journal-title":"J Neurosci"},{"key":"10566_CR39","volume-title":"Deep Learning","author":"I Goodfellow","year":"2016","unstructured":"Goodfellow I, Bengio Y, Courville A (2016) Deep Learning. MIT Press"},{"key":"10566_CR40","unstructured":"Granier A, Senn W (2025) Multihead self-attention in cortico-thalamic circuits. arXiv preprint . 10.48550\/arXiv.2504.06354"},{"issue":"4","key":"10566_CR41","doi-asserted-by":"publisher","first-page":"404","DOI":"10.1037\/h0059831","volume":"38","author":"DA Grant","year":"1948","unstructured":"Grant DA, Berg EA (1948) A behavioral analysis of degree of reinforcement and ease of shifting to new responses in a Weigl-type card-sorting problem. J Exp Psychol 38(4):404\u2013411. https:\/\/doi.org\/10.1037\/h0059831","journal-title":"J Exp Psychol"},{"issue":"6213","key":"10566_CR42","doi-asserted-by":"publisher","first-page":"334","DOI":"10.1038\/338334a0","volume":"338","author":"CM Gray","year":"1989","unstructured":"Gray CM, K\u00f6nig P, Engel AK, Singer W (1989) Oscillatory responses in cat visual cortex exhibit inter-columnar synchronization which reflects global stimulus properties. Nature 338(6213):334\u2013337","journal-title":"Nature"},{"issue":"6","key":"10566_CR43","doi-asserted-by":"publisher","first-page":"401","DOI":"10.1007\/PL00007984","volume":"84","author":"K Gurney","year":"2001","unstructured":"Gurney K, Prescott TJ, Redgrave P (2001) A computational model of action selection in the basal ganglia. Biol Cybern 84(6):401\u2013410","journal-title":"Biol Cybern"},{"key":"10566_CR44","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1038\/nature12654","volume":"503","author":"KD Harris","year":"2013","unstructured":"Harris KD, Mrsic-Flogel TD (2013) Cortical connectivity and sensory coding. Nature 503:51\u201358","journal-title":"Nature"},{"key":"10566_CR45","doi-asserted-by":"publisher","first-page":"170","DOI":"10.1038\/nn.3917","volume":"18","author":"KD Harris","year":"2015","unstructured":"Harris KD, Shepherd GMG (2015) The neocortical circuit: themes and variations. Nat Neurosci 18:170\u2013181","journal-title":"Nat Neurosci"},{"issue":"2","key":"10566_CR46","doi-asserted-by":"publisher","first-page":"245","DOI":"10.1016\/j.neuron.2017.06.011","volume":"95","author":"D Hassabis","year":"2017","unstructured":"Hassabis D, Kumaran D, Summerfield C, Botvinick M (2017) Neuroscience-inspired artificial intelligence. Neuron 95(2):245\u2013258","journal-title":"Neuron"},{"issue":"3","key":"10566_CR47","doi-asserted-by":"publisher","first-page":"416","DOI":"10.1016\/j.neuron.2019.12.002","volume":"105","author":"U Hasson","year":"2020","unstructured":"Hasson U, Nastase SA, Goldstein A (2020) Direct fit to nature: an evolutionary perspective on biological and artificial neural networks. Neuron 105(3):416\u2013434","journal-title":"Neuron"},{"key":"10566_CR48","doi-asserted-by":"crossref","unstructured":"He K, Zhang X, Ren S, Sun J (2016) Deep residual learning for image recognition. IEEE Conference on Computer Vision and Pattern Recognition (CVPR). pp 770\u2013778","DOI":"10.1109\/CVPR.2016.90"},{"key":"10566_CR49","doi-asserted-by":"crossref","unstructured":"Hilgetag CC, Goulas A (1796) hierarchy in the organization of brain networks. Philos Trans R Soc B 375:2020","DOI":"10.1098\/rstb.2019.0319"},{"issue":"8","key":"10566_CR50","doi-asserted-by":"publisher","first-page":"1735","DOI":"10.1162\/neco.1997.9.8.1735","volume":"9","author":"S Hochreiter","year":"1997","unstructured":"Hochreiter S, Schmidhuber J (1997) Long short-term memory. Neural Comput 9(8):1735\u20131780","journal-title":"Neural Comput"},{"key":"10566_CR51","doi-asserted-by":"crossref","unstructured":"Huang G, Liu Z, Van Der Maaten L, Weinberger KQ (2017) Densely connected convolutional networks. IEEE Conference on Computer Vision and Pattern Recognition (CVPR). pp 4700\u20134708","DOI":"10.1109\/CVPR.2017.243"},{"key":"10566_CR52","doi-asserted-by":"publisher","first-page":"106","DOI":"10.1113\/jphysiol.1962.sp006837","volume":"160","author":"DH Hubel","year":"1962","unstructured":"Hubel DH, Wiesel TN (1962) Receptive fields, binocular interaction and functional architecture in the cat\u2019s visual cortex. J Physiol 160:106\u2013154","journal-title":"J Physiol"},{"key":"10566_CR53","doi-asserted-by":"publisher","first-page":"215","DOI":"10.1113\/jphysiol.1968.sp008455","volume":"195","author":"DH Hubel","year":"1968","unstructured":"Hubel DH, Wiesel TN (1968) Receptive fields and functional architecture of monkey striate cortex. J Physiol 195:215\u2013243","journal-title":"J Physiol"},{"key":"10566_CR54","doi-asserted-by":"crossref","unstructured":"J Devlin, Chang M-W, Lee K, Toutanova K (2019) Bert: Pre-training of deep bidirectional transformers for language understanding. Proceedings of the 2019 Conference of the North American Chapter of the Association for Computational Linguistics: Human Language Technologies (NAACL-HLT), pp. 4171\u20134186","DOI":"10.18653\/v1\/N19-1423"},{"issue":"6","key":"10566_CR55","doi-asserted-by":"publisher","first-page":"743","DOI":"10.1038\/nn1909","volume":"10","author":"C Kapfer","year":"2007","unstructured":"Kapfer C, Glickfeld LL, Atallah BV, Scanziani M (2007) Supralinear increase of recurrent inhibition during sparse activity in the somatosensory cortex. Nat Neurosci 10(6):743\u2013753","journal-title":"Nat Neurosci"},{"issue":"11","key":"10566_CR56","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pcbi.1003915","volume":"10","author":"S-M Khaligh-Razavi","year":"2014","unstructured":"Khaligh-Razavi S-M, Kriegeskorte N (2014) Deep supervised, but not unsupervised, models may explain it cortical representation. PLoS Comput Biol 10(11):e1003915","journal-title":"PLoS Comput Biol"},{"issue":"10s","key":"10566_CR57","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1145\/3505244","volume":"54","author":"S Khan","year":"2022","unstructured":"Khan S, Naseer M, Hayat M, Zamir SW, Khan FS, Shah M (2022) Transformers in vision: a survey. ACM Comput Surv 54(10s):1\u201341","journal-title":"ACM Comput Surv"},{"issue":"43","key":"10566_CR58","doi-asserted-by":"publisher","first-page":"21854","DOI":"10.1073\/pnas.1905544116","volume":"116","author":"TC Kietzmann","year":"2019","unstructured":"Kietzmann TC, Spoerer CJ, S\u00f6rensen LKA, Cichy RM, Hauk O, Kriegeskorte N (2019) Recurrence is required to capture the representational dynamics of the human visual system. Proc Natl Acad Sci U S A 116(43):21854\u201321863","journal-title":"Proc Natl Acad Sci U S A"},{"key":"10566_CR59","doi-asserted-by":"publisher","first-page":"14637","DOI":"10.52202\/075280-0644","volume":"36","author":"DK Kim","year":"2023","unstructured":"Kim DK, Kwon J, Cha M, Lee C (2023) Transformer as a hippocampal memory consolidation model based on NMDAR-inspired nonlinearity. Adv Neural Inf Process Syst 36:14637\u201314664","journal-title":"Adv Neural Inf Process Syst"},{"issue":"7345","key":"10566_CR60","doi-asserted-by":"publisher","first-page":"87","DOI":"10.1038\/nature09880","volume":"473","author":"H Ko","year":"2011","unstructured":"Ko H, Hofer SB, Pichler B, Buchanan KA, Sj\u00f6str\u00f6m PJ, Mrsic-Flogel TD (2011) Functional specificity of local synaptic connections in neocortical networks. Nature 473(7345):87\u201391. https:\/\/doi.org\/10.1038\/nature09880","journal-title":"Nature"},{"key":"10566_CR61","doi-asserted-by":"publisher","first-page":"206","DOI":"10.1152\/jn.00149.2003","volume":"91","author":"KP K\u00f6rding","year":"2004","unstructured":"K\u00f6rding KP, Kayser C, Einh\u00e4user W, K\u00f6nig P (2004) How are complex cell properties adapted to the statistics of natural stimuli? J Neurophysiol 91:206\u2013212","journal-title":"J Neurophysiol"},{"key":"10566_CR62","doi-asserted-by":"crossref","unstructured":"K\u00f6rding KP, K\u00f6nig P (2000) Learning with two sites of synaptic integration. Netw Comput Neural Syst 11:1\u201315","DOI":"10.1088\/0954-898X_11_1_302"},{"key":"10566_CR63","doi-asserted-by":"publisher","first-page":"2823","DOI":"10.1162\/089976601317098547","volume":"13","author":"KP K\u00f6rding","year":"2001","unstructured":"K\u00f6rding KP, K\u00f6nig P (2001) Generation of invariances with a neuron model with two sites of synaptic integration. Neural Comput 13:2823\u20132849","journal-title":"Neural Comput"},{"key":"10566_CR64","doi-asserted-by":"publisher","unstructured":"Kozachkov L, Kastanenka KV, Krotov D (2023) Building transformers from neurons and astrocytes. Proceedings of the National Academy of Sciences of the USA 120:e2219150120. https:\/\/doi.org\/10.1073\/pnas.2219150120","DOI":"10.1073\/pnas.2219150120"},{"key":"10566_CR65","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2417788122","volume":"122","author":"L Kozachkov","year":"2025","unstructured":"Kozachkov L, Slotine J-J, Krotov D (2025) Neuron\u2013astrocyte associative memory. Proc Natl Acad Sci U S A 122:e2417788122","journal-title":"Proc Natl Acad Sci U S A"},{"key":"10566_CR66","doi-asserted-by":"publisher","first-page":"417","DOI":"10.1146\/annurev-vision-082114-035447","volume":"1","author":"N Kriegeskorte","year":"2015","unstructured":"Kriegeskorte N (2015) Deep neural networks: a new framework for modeling biological vision and brain information processing. Annual Rev Vision Sci 1:417\u2013446","journal-title":"Annual Rev Vision Sci"},{"key":"10566_CR67","first-page":"1097","volume":"25","author":"A Krizhevsky","year":"2012","unstructured":"Krizhevsky A, Sutskever I, Hinton GE (2012) Imagenet classification with deep convolutional neural networks. In Adv Neural Inform Process Syst 25:1097\u20131105","journal-title":"In Adv Neural Inform Process Syst"},{"key":"10566_CR68","unstructured":"L.\u00a0Kozachkov, J.-J. Slotine, and D.\u00a0Krotov. Neuron\u2013astrocyte associative memory, 2023b. arXiv preprint, arXiv:2311.08135"},{"issue":"11","key":"10566_CR69","doi-asserted-by":"publisher","first-page":"571","DOI":"10.1016\/S0166-2236(00)01657-X","volume":"23","author":"VAF Lamme","year":"2000","unstructured":"Lamme VAF, Roelfsema PR (2000) The distinct modes of vision offered by feedforward and recurrent processing. Trends Neurosci 23(11):571\u2013579","journal-title":"Trends Neurosci"},{"issue":"12","key":"10566_CR70","doi-asserted-by":"publisher","DOI":"10.1101\/cshperspect.a009621","volume":"2","author":"JL Lanciego","year":"2012","unstructured":"Lanciego JL, Luquin N, Obeso JA (2012) Functional neuroanatomy of the basal ganglia. Cold Spring Harb Perspect Med 2(12):a009621","journal-title":"Cold Spring Harb Perspect Med"},{"issue":"3","key":"10566_CR71","doi-asserted-by":"publisher","first-page":"141","DOI":"10.1016\/j.tins.2012.11.006","volume":"36","author":"ME Larkum","year":"2013","unstructured":"Larkum ME (2013) A cellular mechanism for cortical associations: an organizing principle for the cerebral cortex. Trends Neurosci 36(3):141\u2013151","journal-title":"Trends Neurosci"},{"issue":"10","key":"10566_CR72","doi-asserted-by":"publisher","first-page":"1059","DOI":"10.1093\/cercor\/bhh065","volume":"14","author":"ME Larkum","year":"2004","unstructured":"Larkum ME, Senn W, L\u00fcscher H-R (2004) Top-down dendritic input increases the gain of layer 5 pyramidal neurons. Cereb Cortex 14(10):1059\u20131070","journal-title":"Cereb Cortex"},{"issue":"6725","key":"10566_CR73","doi-asserted-by":"publisher","first-page":"338","DOI":"10.1038\/18686","volume":"398","author":"ME Larkum","year":"1999","unstructured":"Larkum ME, Zhu JJ, Sakmann B (1999) A new cellular mechanism for coupling inputs arriving at different cortical layers. Nature 398(6725):338\u2013341","journal-title":"Nature"},{"key":"10566_CR74","doi-asserted-by":"publisher","first-page":"436","DOI":"10.1038\/nature14539","volume":"521","author":"Y LeCun","year":"2015","unstructured":"LeCun Y, Bengio Y, Hinton G (2015) Deep learning. Nature 521:436\u2013444","journal-title":"Nature"},{"key":"10566_CR75","doi-asserted-by":"crossref","unstructured":"LeCun Y, Bottou L, Bengio Y, Haffner P (1998) Gradient-based learning applied to document recognition. In Proceedings of the IEEE 86(11):2278\u20132324","DOI":"10.1109\/5.726791"},{"issue":"4","key":"10566_CR76","doi-asserted-by":"publisher","first-page":"235","DOI":"10.1038\/s41583-020-0277-3","volume":"21","author":"TP Lillicrap","year":"2020","unstructured":"Lillicrap TP, Santoro A, Marris L, Akerman CJ, Hinton G (2020) Backpropagation and the brain. Nat Rev Neurosci 21(4):235\u2013245","journal-title":"Nat Rev Neurosci"},{"issue":"6","key":"10566_CR77","doi-asserted-by":"publisher","first-page":"1002","DOI":"10.1016\/j.neuron.2013.03.007","volume":"77","author":"JE Lisman","year":"2013","unstructured":"Lisman JE, Jensen O (2013) The theta-gamma neural code. Neuron 77(6):1002\u20131016. https:\/\/doi.org\/10.1016\/j.neuron.2013.03.007","journal-title":"Neuron"},{"issue":"9","key":"10566_CR78","doi-asserted-by":"publisher","first-page":"1659","DOI":"10.1016\/S0893-6080(97)00011-7","volume":"10","author":"W Maass","year":"1997","unstructured":"Maass W (1997) Networks of spiking neurons: The third generation of neural network models. Neural Netw 10(9):1659\u20131671","journal-title":"Neural Netw"},{"key":"10566_CR79","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1146\/annurev-neuro-062111-150343","volume":"36","author":"G Major","year":"2013","unstructured":"Major G, Larkum ME, Schiller J (2013) Active properties of neocortical pyramidal neuron dendrites. Annu Rev Neurosci 36:1\u201324","journal-title":"Annu Rev Neurosci"},{"key":"10566_CR80","doi-asserted-by":"publisher","first-page":"94","DOI":"10.3389\/fncom.2016.00094","volume":"10","author":"AH Marblestone","year":"2016","unstructured":"Marblestone AH, Wayne G, Kording KP (2016) Toward an integration of deep learning and neuroscience. Front Comput Neurosci 10:94","journal-title":"Front Comput Neurosci"},{"key":"10566_CR81","unstructured":"Marcus G (2018) Deep learning: A critical appraisal.arXiv:1801.00631"},{"issue":"1","key":"10566_CR82","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.neuron.2012.09.010","volume":"76","author":"E Marder","year":"2012","unstructured":"Marder E (2012) Neuromodulation of neuronal circuits: Back to the future. Neuron 76(1):1\u201311","journal-title":"Neuron"},{"issue":"1","key":"10566_CR83","doi-asserted-by":"publisher","first-page":"225","DOI":"10.1002\/cne.23458","volume":"522","author":"NT Markov","year":"2014","unstructured":"Markov NT, Ercsey-Ravasz M, Van Essen DC, Knoblauch K, Toroczkai Z, Kennedy H (2014) Anatomy of hierarchy: feedforward and feedback pathways in macaque visual cortex. J Comp Neurol 522(1):225\u2013259. https:\/\/doi.org\/10.1002\/cne.23458","journal-title":"J Comp Neurol"},{"key":"10566_CR84","doi-asserted-by":"publisher","first-page":"153","DOI":"10.1038\/nrn1848","volume":"7","author":"H Markram","year":"2006","unstructured":"Markram H (2006) The blue brain project. Nat Rev Neurosci 7:153\u2013160","journal-title":"Nat Rev Neurosci"},{"issue":"5297","key":"10566_CR85","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1126\/science.275.5297.213","volume":"275","author":"H Markram","year":"1997","unstructured":"Markram H, L\u00fcbke J, Frotscher M, Sakmann B (1997) Regulation of synaptic efficacy by coincidence of postsynaptic aps and epsps. Science 275(5297):213\u2013215","journal-title":"Science"},{"key":"10566_CR86","first-page":"479","volume-title":"The Cognitive Neurosciences V","author":"KAC Martin","year":"2014","unstructured":"Martin KAC (2014) Microcircuits in visual cortex. In: Gazzaniga MS (ed) The Cognitive Neurosciences V. MIT Press, pp 479\u2013492"},{"key":"10566_CR87","doi-asserted-by":"publisher","first-page":"115","DOI":"10.1007\/BF02478259","volume":"5","author":"WS McCulloch","year":"1943","unstructured":"McCulloch WS, Pitts W (1943) A logical calculus of the ideas immanent in nervous activity. Bulletin Math Biophys 5:115\u2013133","journal-title":"Bulletin Math Biophys"},{"key":"10566_CR88","doi-asserted-by":"crossref","unstructured":"Mondal K, Kumar A (2025) Attention via synaptic plasticity is all you need: a biologically inspired spiking neuromorphic transformer.arXiv:2511.14691 arXiv preprint","DOI":"10.2139\/ssrn.6096874"},{"issue":"4","key":"10566_CR89","doi-asserted-by":"publisher","first-page":"701","DOI":"10.1093\/brain\/120.4.701","volume":"120","author":"VB Mountcastle","year":"1997","unstructured":"Mountcastle VB (1997) The columnar organization of the neocortex. Brain 120(4):701\u2013722","journal-title":"Brain"},{"issue":"2","key":"10566_CR90","doi-asserted-by":"publisher","first-page":"283","DOI":"10.1162\/089976606775093909","volume":"18","author":"RC O\u2019Reilly","year":"2006","unstructured":"O\u2019Reilly RC, Frank MJ (2006) Making working memory work: a computational model of learning in the prefrontal cortex and basal ganglia. Neural Comput 18(2):283\u2013328","journal-title":"Neural Comput"},{"issue":"1","key":"10566_CR91","doi-asserted-by":"publisher","first-page":"69","DOI":"10.1093\/cercor\/3.1.69","volume":"3","author":"A Peters","year":"1993","unstructured":"Peters A, Payne BR (1993) Numerical relationships between geniculocortical afferents and pyramidal cell modules in cat primary visual cortex. Cereb Cortex 3(1):69\u201378","journal-title":"Cereb Cortex"},{"key":"10566_CR92","doi-asserted-by":"publisher","DOI":"10.3389\/fnins.2018.00774","volume":"12","author":"M Pfeiffer","year":"2018","unstructured":"Pfeiffer M, Pfeil T (2018) Deep learning with spiking neurons: opportunities and challenges. Front Neurosci 12:774","journal-title":"Front Neurosci"},{"key":"10566_CR93","doi-asserted-by":"publisher","first-page":"50","DOI":"10.3389\/fncir.2018.00050","volume":"12","author":"WA Phillips","year":"2018","unstructured":"Phillips WA, Bachmann T, Storm JF (2018) Apical function in neocortical pyramidal cells: A common pathway by which general anesthetics can affect consciousness and other cognitive processes. Front Neural Circuits 12:50","journal-title":"Front Neural Circuits"},{"key":"10566_CR94","volume":"38","author":"WA Phillips","year":"2015","unstructured":"Phillips WA, Clark A, Silverstein SM (2015) On the functions, mechanisms, and malfunctions of buffering in the brain. Behav Brain Sci 38:e3","journal-title":"Behav Brain Sci"},{"issue":"3","key":"10566_CR95","doi-asserted-by":"crossref","first-page":"211","DOI":"10.1017\/S0140525X12002257","volume":"36","author":"WA Phillips","year":"2013","unstructured":"Phillips WA, Silverstein SM (2013) Convergence of biological and psychological perspectives on cognitive coordination in schizophrenia. Behav Brain Sci 36(3):211\u2013212","journal-title":"Behav Brain Sci"},{"issue":"2","key":"10566_CR96","doi-asserted-by":"publisher","first-page":"222","DOI":"10.1162\/jocn.1997.9.2.222","volume":"9","author":"A Pouget","year":"1997","unstructured":"Pouget A, Sejnowski TJ (1997) Spatial transformations in the parietal cortex using basis functions. J Cogn Neurosci 9(2):222\u2013237","journal-title":"J Cogn Neurosci"},{"issue":"5885","key":"10566_CR97","doi-asserted-by":"publisher","first-page":"48","DOI":"10.1126\/science.1155564","volume":"321","author":"MI Rabinovich","year":"2008","unstructured":"Rabinovich MI, Huerta R, Laurent G (2008) Transient dynamics for neural processing. Science 321(5885):48\u201350","journal-title":"Science"},{"issue":"1","key":"10566_CR98","doi-asserted-by":"publisher","first-page":"79","DOI":"10.1038\/4580","volume":"2","author":"RPN Rao","year":"1999","unstructured":"Rao RPN, Ballard DH (1999) Predictive coding in the visual cortex: A functional interpretation of some extra-classical receptive-field effects. Nat Neurosci 2(1):79\u201387","journal-title":"Nat Neurosci"},{"issue":"2","key":"10566_CR99","doi-asserted-by":"publisher","first-page":"168","DOI":"10.1016\/j.neuron.2009.01.002","volume":"61","author":"JH Reynolds","year":"2009","unstructured":"Reynolds JH, Heeger DJ (2009) The normalization model of attention. Neuron 61(2):168\u2013185","journal-title":"Neuron"},{"key":"10566_CR100","doi-asserted-by":"publisher","first-page":"1761","DOI":"10.1038\/s41593-019-0520-2","volume":"22","author":"BA Richards","year":"2019","unstructured":"Richards BA, Lillicrap TP, Beaudoin P, Bengio Y, Bogacz R, Christensen A, Clopath C, Costa RP, de Berker A, Ganguli S, Gillon CJ, Hafner D, Kepecs A, Kriegeskorte N, Latham P, Lindsay GW, Miller KD, Naud R, Pack CC, Poirazi P, Roelfsema P, Sacramento J, Saxe A, Scellier B, Schapiro AC, Senn W, Wayne G, Yamins D, Zenke F, Zylberberg J, Therien D (2019) A deep learning framework for neuroscience. Nat Neurosci 22:1761\u20131770","journal-title":"Nat Neurosci"},{"issue":"7451","key":"10566_CR101","doi-asserted-by":"publisher","first-page":"585","DOI":"10.1038\/nature12160","volume":"497","author":"M Rigotti","year":"2013","unstructured":"Rigotti M, Barak O, Warden MR, Wang X-J, Daw ND, Miller EK, Fusi S (2013) The importance of mixed selectivity in complex cognitive tasks. Nature 497(7451):585\u2013590","journal-title":"Nature"},{"issue":"13","key":"10566_CR102","doi-asserted-by":"publisher","first-page":"5639","DOI":"10.1523\/JNEUROSCI.22-13-05639.2002","volume":"22","author":"DL Ringach","year":"2002","unstructured":"Ringach DL, Shapley RM, Hawken MJ (2002) Orientation selectivity in macaque v1: Diversity and laminar dependence. J Neurosci 22(13):5639\u20135651. https:\/\/doi.org\/10.1523\/JNEUROSCI.22-13-05639.2002","journal-title":"J Neurosci"},{"issue":"3","key":"10566_CR103","doi-asserted-by":"publisher","first-page":"303","DOI":"10.1002\/cne.902160307","volume":"216","author":"KS Rockland","year":"1983","unstructured":"Rockland KS, Lund JS (1983) Intrinsic laminar lattice connections in primate visual cortex. J Comp Neurol 216(3):303\u2013318","journal-title":"J Comp Neurol"},{"key":"10566_CR104","doi-asserted-by":"publisher","first-page":"131","DOI":"10.1146\/annurev-vision-111815-114443","volume":"2","author":"PR Roelfsema","year":"2016","unstructured":"Roelfsema PR, de Lange FP (2016) Early visual cortex as a multiscale cognitive blackboard. Annual Rev Vision Sci 2:131\u2013151","journal-title":"Annual Rev Vision Sci"},{"issue":"11","key":"10566_CR105","doi-asserted-by":"publisher","DOI":"10.1016\/j.heliyon.2024.e31965","volume":"10","author":"ET Rolls","year":"2024","unstructured":"Rolls ET (2024) The memory systems of the human brain and generative artificial intelligence. Heliyon 10(11):e31965. https:\/\/doi.org\/10.1016\/j.heliyon.2024.e31965","journal-title":"Heliyon"},{"issue":"6","key":"10566_CR106","doi-asserted-by":"publisher","first-page":"386","DOI":"10.1037\/h0042519","volume":"65","author":"F Rosenblatt","year":"1958","unstructured":"Rosenblatt F (1958) The perceptron: A probabilistic model for information storage and organization in the brain. Psychol Rev 65(6):386\u2013408","journal-title":"Psychol Rev"},{"key":"10566_CR107","doi-asserted-by":"publisher","first-page":"533","DOI":"10.1038\/323533a0","volume":"323","author":"DE Rumelhart","year":"1986","unstructured":"Rumelhart DE, Hinton GE, Williams RJ (1986) Learning representations by back-propagating errors. Nature 323:533\u2013536","journal-title":"Nature"},{"issue":"3","key":"10566_CR108","doi-asserted-by":"publisher","first-page":"211","DOI":"10.1007\/s11263-015-0816-y","volume":"115","author":"O Russakovsky","year":"2015","unstructured":"Russakovsky O, Deng J, Su H, Krause J, Satheesh S, Ma S, Huang Z, Karpathy A, Khosla A, Bernstein M, Berg AC, Fei-Fei L (2015) Imagenet large scale visual recognition challenge. Int J Comput Vision 115(3):211\u2013252","journal-title":"Int J Comput Vision"},{"issue":"1","key":"10566_CR109","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1016\/S0896-6273(00)00004-0","volume":"27","author":"E Salinas","year":"2000","unstructured":"Salinas E, Thier P (2000) Gain modulation: a major computational principle of the central nervous system. Neuron 27(1):15\u201321","journal-title":"Neuron"},{"key":"10566_CR110","doi-asserted-by":"publisher","first-page":"55","DOI":"10.1038\/s41583-020-00395-8","volume":"22","author":"A Saxe","year":"2021","unstructured":"Saxe A, Nelli S, Summerfield C (2021) If deep learning is the answer, what is the question? Nat Rev Neurosci 22:55\u201367","journal-title":"Nat Rev Neurosci"},{"key":"10566_CR111","unstructured":"Schrimpf M, Blank IA, Tuckute G, Kauf C, Hosseini EA, Kanwisher N, Tenenbaum JB, Fedorenko E (2020) Artificial neural networks accurately predict language processing in the brain. bioRxiv, page 2020.06.26.174482"},{"issue":"5306","key":"10566_CR112","doi-asserted-by":"publisher","first-page":"1593","DOI":"10.1126\/science.275.5306.1593","volume":"275","author":"W Schultz","year":"1997","unstructured":"Schultz W, Dayan P, Montague PR (1997) A neural substrate of prediction and reward. Science 275(5306):1593\u20131599","journal-title":"Science"},{"issue":"21","key":"10566_CR113","doi-asserted-by":"publisher","first-page":"2121","DOI":"10.1016\/j.cub.2013.09.013","volume":"23","author":"MW Self","year":"2013","unstructured":"Self MW, van Kerkoerle T, Sup\u00e8r H, Roelfsema PR (2013) Distinct roles of the cortical layers of area v1 in figure\u2013ground segregation. Curr Biol 23(21):2121\u20132129","journal-title":"Curr Biol"},{"issue":"5","key":"10566_CR114","doi-asserted-by":"publisher","first-page":"735","DOI":"10.1016\/j.neuron.2007.02.012","volume":"53","author":"G Silberberg","year":"2007","unstructured":"Silberberg G, Markram H (2007) Disynaptic inhibition between neocortical pyramidal cells mediated by martinotti cells. Neuron 53(5):735\u2013746","journal-title":"Neuron"},{"key":"10566_CR115","unstructured":"Simonyan K, Zisserman A (2015) Very deep convolutional networks for large-scale image recognition. International Conference on Learning Representations (ICLR)"},{"issue":"1","key":"10566_CR116","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1016\/S0896-6273(00)80821-1","volume":"24","author":"W Singer","year":"1999","unstructured":"Singer W (1999) Neuronal synchrony: a versatile code for the definition of relations? Neuron 24(1):49\u201365","journal-title":"Neuron"},{"key":"10566_CR117","doi-asserted-by":"publisher","first-page":"1551","DOI":"10.3389\/fpsyg.2017.01551","volume":"8","author":"CJ Spoerer","year":"2017","unstructured":"Spoerer CJ, McClure P, Kriegeskorte N (2017) Recurrent convolutional neural networks: a better model of biological object recognition. Front Psychol 8:1551","journal-title":"Front Psychol"},{"issue":"4","key":"10566_CR118","doi-asserted-by":"publisher","first-page":"739","DOI":"10.1016\/S0896-6273(02)01029-2","volume":"36","author":"DD Stettler","year":"2002","unstructured":"Stettler DD, Das A, Bennett J, Gilbert CD (2002) Lateral connectivity and contextual interactions in macaque primary visual cortex. Neuron 36(4):739\u2013750","journal-title":"Neuron"},{"issue":"6","key":"10566_CR119","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pcbi.1014378","volume":"22","author":"F Tahvili","year":"2026","unstructured":"Tahvili F, Vinck M, Di Volo M (2026) A mean-field model of neural networks with PV and SOM interneurons reveals connectivity-based mechanisms of gamma oscillations. PLoS Comput Biol 22(6):e1014378. https:\/\/doi.org\/10.1371\/journal.pcbi.1014378","journal-title":"PLoS Comput Biol"},{"issue":"1","key":"10566_CR120","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1093\/cercor\/13.1.5","volume":"13","author":"AM Thomson","year":"2003","unstructured":"Thomson AM, Bannister AP (2003) Interlaminar connections in the neocortex. Cereb Cortex 13(1):5\u201314. https:\/\/doi.org\/10.1093\/cercor\/13.1.5","journal-title":"Cereb Cortex"},{"issue":"1","key":"10566_CR121","doi-asserted-by":"publisher","first-page":"42","DOI":"10.1186\/1471-2202-5-42","volume":"5","author":"G Tononi","year":"2004","unstructured":"Tononi G (2004) An information integration theory of consciousness. BMC Neurosci 5(1):42","journal-title":"BMC Neurosci"},{"key":"10566_CR122","doi-asserted-by":"publisher","first-page":"575","DOI":"10.1038\/21176","volume":"399","author":"S Treue","year":"1999","unstructured":"Treue S, Mart\u00ednez Trujillo JC (1999) Feature-based attention influences motion processing gain in macaque visual cortex. Nature 399:575\u2013579","journal-title":"Nature"},{"issue":"3","key":"10566_CR123","doi-asserted-by":"publisher","first-page":"521","DOI":"10.1016\/j.neuron.2013.11.030","volume":"81","author":"R Urbanczik","year":"2014","unstructured":"Urbanczik R, Senn W (2014) Learning by the dendritic prediction of somatic spiking. Neuron 81(3):521\u2013528","journal-title":"Neuron"},{"key":"10566_CR124","unstructured":"Vaswani A, Shazeer N, Parmar N, Uszkoreit J, Jones L, Gomez AN, Kaiser L, Polosukhin I (2017) Attention is all you need. NeurIPS. pp 5998\u20136008"},{"issue":"1","key":"10566_CR125","first-page":"1","volume":"56","author":"R Wang","year":"2023","unstructured":"Wang R, Zhang Z (2023) Neural energy theory. Artif Intell Rev 56(1):1\u201345","journal-title":"Artif Intell Rev"},{"issue":"5","key":"10566_CR126","doi-asserted-by":"publisher","first-page":"955","DOI":"10.1016\/S0896-6273(02)01092-9","volume":"36","author":"X-J Wang","year":"2002","unstructured":"Wang X-J (2002) Probabilistic decision making by slow reverberation in cortical circuits. Neuron 36(5):955\u2013968","journal-title":"Neuron"},{"key":"10566_CR127","unstructured":"Whittington JCR, Warren J, Behrens TEJ (2022) Relating transformers to models and neural representations of the hippocampal formation. International Conference on Learning Representations"},{"issue":"5561","key":"10566_CR128","doi-asserted-by":"publisher","first-page":"1907","DOI":"10.1126\/science.1067903","volume":"295","author":"SR Williams","year":"2002","unstructured":"Williams SR, Stuart GJ (2002) Dependence of epsp efficacy on synapse location in neocortical pyramidal neurons. Science 295(5561):1907\u20131910","journal-title":"Science"},{"issue":"4","key":"10566_CR129","doi-asserted-by":"publisher","first-page":"1314","DOI":"10.1523\/JNEUROSCI.3733-05.2006","volume":"26","author":"K-F Wong","year":"2006","unstructured":"Wong K-F, Wang X-J (2006) A recurrent network mechanism of time integration in perceptual decisions. J Neurosci 26(4):1314\u20131328","journal-title":"J Neurosci"},{"issue":"3","key":"10566_CR130","doi-asserted-by":"publisher","first-page":"356","DOI":"10.1038\/nn.4244","volume":"19","author":"DLK Yamins","year":"2016","unstructured":"Yamins DLK, DiCarlo JJ (2016) Using goal-driven deep learning models to understand sensory cortex. Nat Neurosci 19(3):356\u2013365","journal-title":"Nat Neurosci"},{"key":"10566_CR131","doi-asserted-by":"crossref","unstructured":"Yamins DLK, Hong H, Cadieu CF, Solomon EA, Seibert D, DiCarlo JJ (2014) Performance-optimized hierarchical models predict neural responses in higher visual cortex. In Proceedings of the National Academy of Sciences 111(23):8619\u20138624","DOI":"10.1073\/pnas.1403112111"},{"key":"10566_CR132","unstructured":"Zhou Z, Zhu Y, He C, Wang Y, Su F, Tian Y (2023) Spikformer: When spiking neural network meets transformer. International Conference on Learning Representations (ICLR"},{"issue":"4","key":"10566_CR133","doi-asserted-by":"publisher","first-page":"565","DOI":"10.1016\/j.neuron.2025.11.024","volume":"114","author":"TA Zolnik","year":"2026","unstructured":"Zolnik TA, Eickholt BJ, Moln\u00e1r Z, Larkum ME (2026) The layer 6b theory of attention. Neuron 114(4):565\u2013582. https:\/\/doi.org\/10.1016\/j.neuron.2025.11.024","journal-title":"Neuron"}],"container-title":["Cognitive Neurodynamics"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s11571-026-10566-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s11571-026-10566-5","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s11571-026-10566-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T20:02:29Z","timestamp":1791489749000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s11571-026-10566-5"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10,8]]},"references-count":133,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2026,12]]}},"alternative-id":["10566"],"URL":"https:\/\/doi.org\/10.1007\/s11571-026-10566-5","relation":{},"ISSN":["1871-4080","1871-4099"],"issn-type":[{"value":"1871-4080","type":"print"},{"value":"1871-4099","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,10,8]]},"assertion":[{"value":"13 May 2026","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"8 September 2026","order":2,"name":"revised","label":"Revised","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"21 September 2026","order":3,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"8 October 2026","order":4,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Declarations"}}],"article-number":"197"}}