cyto7: an open, surface-native cytoarchitectural type atlas of the human cortex with multimodal validation and a per-vertex confidence map
Ricardo Salvador, Borja Mercadal, Francesca Castaldo, ★ Giulio Ruffini
★ guarantor: Giulio Ruffini · vouches for the paper per WP0084 §6
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The cerebral cortex is organized along an axis of laminar differentiation, the cortical-type axis of the Structural Model, that predicts laminar connectivity, plasticity, and disease vulnerability. Existing digital cortical-type maps are area-level: they assign one type per classical area and cannot represent a type boundary that runs through an area. We release , an open, vertex-level, seven-type cytoarchitectural atlas painted on the fsaverage surface following the García-Cabezas protocol and revised by the protocol's authors.
We distribute the atlas with a reusable quality-control and support framework. A topological consistency audit reports the map against the four spatial rules the Structural Model predicts, which were imposed as constraints during construction rather than tested, and quantifies what imposing them cost: 2,368 vertices (0.78% of labelled cortex) were relabelled to satisfy the sequential-gradient rule, of which only 42 were strictly required. A vertex-wise benchmark against the existing area-level map, one type per von Economo--Koskinas area, shows exact agreement at 52.5% of vertices (Cohen's (\kappa = 0.36), quadratic-weighted (\kappa_w = 0.70)), with disagreement almost always a single level and concentrated inside the large, heterogeneous areas that a single areal label cannot resolve, the within-area refinement a vertex map is meant to provide. Where the two maps disagree, 's label better matches the independent multimodal profile in 60% of vertices, against a chance level of 46% under a rotation null (spin (p = 0.001)) and of 54% under 200 granularity-matched alternative partitions ((p = 0.020)), the harder of the two comparisons. A per-vertex anatomical support score, computed from anatomy alone, states how strongly each hand-drawn label is backed by the anatomical evidence, and is deliberately not a calibrated error probability.
We then validate against independent brain measures not used to define it, under a spatial-autocorrelation (spin) null with FDR control. Intracortical myelin (T1w/T2w) rises, cortical thickness falls, and the principal functional-connectivity gradient reverses along the type axis (all (q \lesssim 0.006)). Source-reconstructed resting-state MEG dynamics also track the axis: the intrinsic timescale shortens toward koniocortex ((\rho \approx -0.46)) and survives the null, as do the spectral centroid and the slow/fast band-power ratio, whereas the aperiodic 1/f exponent does not. The laminar marker genes are what separate from a quantisation of any single continuous map: it tracks the layer-IV marker RORB (ρ = +0.78) and the layer-V marker FEZF2 (−0.59) more closely than seven-level septiles of myelin, of the functional gradient or of a histological gradient, while tying or losing to those septiles on the macroscale proxies. Defined from anatomy alone, recovers not only the sensorimotor-to-association hierarchy but the laminar basis of it, and is released with its support score and full pipeline as a reusable, uncertainty-quantified substrate for structure--function and modelling studies.
{introduction}{
A vertex-resolution map of human cortical cell-layer types, validated against imaging, genetics, and brain dynamics — and shipped with per-vertex uncertainty estimates.
The cerebral cortex isn't uniform. It varies systematically in how many distinct layers it has, and especially in how developed its fourth layer (the "granular" layer) is. This gradient — from the nearly layerless allocortex near the hippocampus, through transitional limbic cortex, up to the hyper-granular primary sensory areas — is called the cortical-type axis. It predicts how areas connect, how plastic they are, and which diseases hit them hardest. The problem is that existing digital maps of this axis assign one type per classical brain area, like painting a country a single color. A type boundary that runs through an area is invisible. cyto7 fixes this by painting types vertex-by-vertex on the standard fsaverage surface — roughly 164,000 points — following an established expert protocol, then having the protocol's own authors revise the result.
The map enforces four spatial rules the theory predicts: types must change gradually between neighbors (no skipping a level), the least-differentiated limbic cortex must form a continuous belt around the medial wall, and the most differentiated types must appear as isolated islands. Satisfying these rules required relabeling only 0.78% of vertices, and removing those relabeled vertices barely moves any correlation. The map is then benchmarked against the older area-level map: they agree at 52.5% of vertices, with disagreement almost always just one ordinal step and concentrated inside the large heterogeneous areas that a single areal label can't resolve. Where they disagree, cyto7's label better matches independent MRI features (myelin, thickness, functional connectivity gradient) at 60% of disagreement vertices versus a chance rate of ~46–54% under conservative null tests.
The multimodal validation is the external, independent leg. Intracortical myelin rises along the type axis, cortical thickness falls, and the principal functional-connectivity gradient reverses — all surviving a spatial-autocorrelation null with FDR control. Resting-state MEG dynamics also track the axis: neural timescales shorten toward more granular cortex (ρ ≈ −0.46), and the spectral centroid rises, consistent with faster dynamics in more differentiated areas. Notably, the aperiodic 1/f exponent — often interpreted as an excitation-inhibition proxy — does not track type, a reliable null given good spectral fits and high reproducibility. The clearest separation from simply quantizing a continuous MRI map comes from laminar marker genes: cyto7 tracks the layer-IV marker RORB (ρ = +0.78) and the layer-V marker FEZF2 (ρ = −0.59) more closely than seven-level bins of myelin or the functional gradient, because those continuous maps have no way to represent a marker that runs against myelin.
Each vertex also gets an anatomical support score — a number between 0 and 1 built purely from anatomy (atlas concordance, topological consistency, geometry, and a per-type provenance prior), deliberately excluding the imaging data used for validation. Eulaminate and koniocortex vertices score highest; the developmentally-defined allocortex scores lowest, flagging exactly the tissue where the typing rests on embryological reasoning rather than surface atlases. The honest limitations are stated plainly: single rater, no independent second painting, and the one direct histological check (BigBrain) is inconclusive at single-specimen scale. The map is released open, with the full change log, per-vertex support components, and a crossed parcellation for region-based models.
- WP ID
- WP0226
- Lifecycle
- ongoing
- Visibility
- internal
- Access level
- open
- Embargo until
- —
- Priority
- —
- Collab
- open
- Venue
- —
- DOI
- —
- Deadline
- —
- Owner
- giulio.ruffini@bcom.one
- Source
- drive_legacy
- Repo path
- WP0226
- 0.1.0 (draft) · auto-run-placeholder
