A genuine layer 4 in motor cortex with prototypical synaptic circuit connectivity
Open accessAbstract
The motor cortex (M1) is classically considered an agranular area, lacking a distinct layer 4 (L4). Here, we tested the idea that M1, despite lacking a cytoarchitecturally visible L4, nevertheless possesses its equivalent in the form of excitatory neurons with input–output circuits like those of the L4 neurons in sensory areas. Consistent with this idea, we found that neurons located in a thin laminar zone at the L3/5A border in the forelimb area of mouse M1 have multiple L4-like synaptic connections: excitatory input from thalamus, largely unidirectional excitatory outputs to L2/3 pyramidal neurons, and relatively weak long-range corticocortical inputs and outputs. M1-L4 neurons were electrophysiologically diverse but morphologically uniform, with pyramidal-type dendritic arbors and locally ramifying axons, including branches extending into L2/3. Our findings therefore identify pyramidal neurons in M1 with the expected prototypical circuit properties of excitatory L4 neurons, and question the traditional assumption that motor cortex lacks this layer.
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A t-type, L4/5 IT_1, was located on the boundary between L2/3 and L5 and probably corresponds to the quasi-L4 neurons described previously in motor cortex.
“We were also able to identify a t-type, L4/5 IT_1, that was located on the boundary between L2/3 and L5 and probably corresponds to the quasi-L4 neurons described previously in motor cortex31.”
From Phenotypic variation of transcriptomic cell types in mouse motor cortex (2021) · cites this paper as [31]
Previous studies suggested that L4 neurons similar to those typically found in sensory cortical areas are present in mouse MOp and macaque M1.
“However, previous studies have suggested that L4 neurons similar to those typically found in sensory cortical areas are also present in mouse MOp and macaque M165,66.”
From A multimodal cell census and atlas of the mammalian primary motor cortex (2021) · cites this paper as [65]
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- Authors
- Yamawaki, N., Borges, K., Suter, B. A., Harris, K. D., & Shepherd, G. M. G.
- Affiliations
- Northwestern University (US)
- UCL Queen Square Institute of Neurology (GB)
- Corresponding author
- Shepherd, Gordon M G
- Venue
- eLife
- Publisher
- eLife Sciences Publications, Ltd
- Published
- 2014-12-19
- Volume
- 3
- Pages
- e05422-e05422
- ISSN
- 2050-084X (ISSN-L)
- PMC
- PMC4290446
- OpenAlex
- W2171385325
- Cited by
- 141
- References
- 77
- Citations by year
- 2026: 8 · 2025: 11 · 2024: 8 · 2023: 16 · 2022: 17 · 2021: 19 · 2020: 11 · 2019: 14 · 2018: 8 · 2017: 14 · 2016: 7 · 2015: 8
- FWCI
- 4.42
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Licence
- Locations
- eLife (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
- UCL Discovery (University College London) (submittedVersion)
- Funders
- National Institutes of Health (NS061963)
- Whitehall Foundation
- Wellcome Trust
- Topics
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neuroscience and Neuropharmacology Research
- Life Sciences › Neuroscience › Cognitive Neuroscience › Motor Control and Adaptation
- Life Sciences › Neuroscience › Cognitive Neuroscience › Neural dynamics and brain function
- Keywords
- motor cortex, layer 4, pyramidal neurons, synaptic connectivity, thalamocortical input, corticocortical connections, neuronal morphology, laminar organization
- MeSH terms
- Action Potentials, Adenoviridae, Animals, Axons, Dendrites, Fluorescent Dyes, Genetic Vectors, Microspheres, Microtomy, Motor Cortex, Synaptic Transmission, Stereotaxic Techniques, Synapses, Thalamus, Pyramidal Cells, Tissue Culture Techniques, Mice, Synaptic Potentials
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
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