The DOI matches its Crossref record; 205 of the 205 references it cites were found in a registry or an index.
Neocortical Layer 1: An Elegant Solution to Top-Down and Bottom-Up Integration
Open accessAbstract
Many of our daily activities, such as riding a bike to work or reading a book in a noisy cafe, and highly skilled activities, such as a professional playing a tennis match or a violin concerto, depend upon the ability of the brain to quickly make moment-to-moment adjustments to our behavior in response to the results of our actions. Particularly, they depend upon the ability of the neocortex to integrate the information provided by the sensory organs (bottom-up information) with internally generated signals such as expectations or attentional signals (top-down information). This integration occurs in pyramidal cells (PCs) and their long apical dendrite, which branches extensively into a dendritic tuft in layer 1 (L1). The outermost layer of the neocortex, L1 is highly conserved across cortical areas and species. Importantly, L1 is the predominant input layer for top-down information, relayed by a rich, dense mesh of long-range projections that provide signals to the tuft branches of the PCs. Here, we discuss recent progress in our understanding of the composition of L1 and review evidence that L1 processing contributes to functions such as sensory perception, cross-modal integration, controlling states of consciousness, attention, and learning.
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“the distal compartment receives input from neuromodulatory centers [36], other cortical areas [37–39], and the non-lemniscal thalamus [40,41]”
From Testing circuit-level theories of consciousness in humans (2026) · cites this paper as [36]
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Cites
Neural Mechanisms of Selective Visual Attention
Desimone, R., & Duncan, J. (1995) · Annual Review of Neuroscience · doi:10.1146/annurev.ne.18.030195.001205 · Cited by 8,388
Distributed hierarchical processing in the primate cerebral cortex
Felleman, D. J., & Van Essen, D. C. (1991) · Cerebral Cortex · doi:10.1093/cercor/1.1.1 · Cited by 7,783
Rao, R. P. N., & Ballard, D. H. (1999) · Nature Neuroscience · doi:10.1038/4580 · Cited by 5,483
Preface: Cerebral Cortex Has Come of Age
Goldman, P. S., & RAKIC, P. T. (1991) · Cerebral Cortex · doi:10.1093/cercor/1.1.1-a · Cited by 3,537
Canonical Microcircuits for Predictive Coding
Bastos, A. M., Usrey, W. M., Adams, R. A., Mangun, G. R., Fries, P., & Friston, K. J. (2012) · Neuron · doi:10.1016/j.neuron.2012.10.038 · Cited by 2,421
Mechanisms of visual attention in the human cortex
Ungerleider, S. K. A. L. G. (2000) · Annual Review of Neuroscience · doi:10.1146/annurev.neuro.23.1.315 · Cited by 2,159
GABAergic Interneurons in the Neocortex: From Cellular Properties to Circuits
Tremblay, R., Lee, S., & Rudy, B. (2016) · Neuron · doi:10.1016/j.neuron.2016.06.033 · Cited by 2,033
Self-citationNEURONAL CIRCUITS OF THE NEOCORTEX
Douglas, R. J., & Martin, K. A. C. (2004) · Annual Review of Neuroscience · doi:10.1146/annurev.neuro.27.070203.144152 · Cited by 1,886
Maunsell, J., & van Essen, D. (1983) · The Journal of Neuroscience · doi:10.1523/jneurosci.03-12-02563.1983 · Cited by 1,602
Pfeffer, C. K., Xue, M., He, M., Huang, Z. J., & Scanziani, M. (2013) · Nature Neuroscience · doi:10.1038/nn.3446 · Cited by 1,353
Research works printed but not linked: 7
- Chen Z, 2015, Front. Neural. Circuits, 9, 83
- Chou XL, 2020, eLife, 9:e54157
- Guillery RW., 1995, J. Anat., 187, 583
- Jones EG., 2007, The Thalamus
- Kerlin A, 2019, eLife, 8:e46966
- Krnjevic K, 1965, J. Anat., 99, 711
- Zhang W, 2019, eLife, 8:e44158
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Moreno, C., Riquelme, D., Cornejo, C., Leyton, P., & Leiva‐Salcedo, E. (2026) · Biological Research · doi:10.1186/s40659-026-00673-2
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Martinez, M. (2025) · theses.fr (ABES) · doi:10.70675/d05e33fcz2cd4z45d4zbe74z5e57fadd4292
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Zhao, H., Anderson, T. R., & Schmidt, E. R. E. (2026) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.64898/2026.06.24.734394
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Liu, N., Mais, L.-T., Mustin, M., Yun, S. D., Shah, N. J., Fink, G. R., Grefkes, C., & Tscherpel, C. (2026) · medRxiv · doi:10.64898/2026.07.30.26359200
Shin, H., Kwon, S. Y., Hwang, H., Ko, T., Choi, E. B., Kim, Y., Han, Y. M., Kim, J. G., Zhou, Q., Yang, S., et al. (2026) · International Journal of Molecular Sciences · doi:10.3390/ijms27177564 · Cited by 1
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Metadata
- Authors
- Schuman, B., Dellal, S., Prönneke, A., Machold, R., & Rudy, B.
- Affiliations
- New York University (US)
- Corresponding author
- Rudy, Bernardo
- Venue
- Annual Review of Neuroscience
- Publisher
- Annual Reviews
- Published
- 2021-07-08
- Volume
- 44
- Issue
- 1
- Pages
- 221-252
- ISSN
- 0147-006X (ISSN-L)
- 1545-4126
- PMC
- PMC9012327
- OpenAlex
- W3138514668
- Cited by
- 148
- References
- 204
- Citations by year
- 2026: 22 · 2025: 28 · 2024: 31 · 2023: 47 · 2022: 12 · 2021: 8
- FWCI
- 3.62
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Locations
- Annual Review of Neuroscience (publishedVersion)
- PubMed (publishedVersion)
- PubMed Central (acceptedVersion)
- Funders
- National Institutes of Health
- National Institute of Mental Health (T32MH096331)
- National Institute of Neurological Disorders and Stroke (R01NS107257, P01NS074972, R01NS110079, F31NS106793)
- Topics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Neural dynamics and brain function
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neuroscience and Neuropharmacology Research
- Social Sciences › Psychology › Experimental and Cognitive Psychology › Multisensory perception and integration
- Keywords
- top-down processing, bottom-up processing, pyramidal neurons, apical dendrites, sensory perception, cross-modal integration, consciousness, attention, learning, cortical integration
- MeSH terms
- Dendrites, Learning, Pyramidal Cells, Neocortex
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