The Control of Cortical Folding: Multiple Mechanisms, Multiple Models.

The cerebral cortex develops through a carefully conscripted series of cellular and molecular events that culminate in the production of highly specialized neuronal and glial cells. During development, cortical neurons and glia acquire a precise cellular arrangement and architecture to support higher-order cognitive functioning. Decades of study using rodent models, naturally gyrencephalic animal models, human pathology specimens, and, recently, human cerebral organoids, reveal that rodents recapitulate some but not all the cellular and molecular features of human cortices. Whereas rodent cortices are smooth-surfaced or lissencephalic, larger mammals, including humans and nonhuman primates, have highly folded/gyrencephalic cortices that accommodate an expansion in neuronal mass and increase in surface area. Several genes have evolved to drive cortical gyrification, arising from gene duplications or de novo origins, or by alterations to the structure/function of ancestral genes or their gene regulatory regions. Primary cortical folds arise in stereotypical locations, prefigured by a molecular "blueprint" that is set up by several signaling pathways (e.g., Notch, Fgf, Wnt, PI3K, Shh) and influenced by the extracellular matrix. Mutations that affect neural progenitor cell proliferation and/or neurogenesis, predominantly of upper-layer neurons, perturb cortical gyrification. Below we review the molecular drivers of cortical folding and their roles in disease.

Shifting attention does not influence numerical processing.

Many theories of numerical cognition assume that numbers and space share a common representation at the response level. For example, observers are faster to respond to small numbers with their left hand and large numbers with their right hand (the SNARC effect). There is also evidence that viewing numbers can produce spatial shifts of attention, suggesting that attention may play a role in the spatial representation of numbers. In the present study, we assessed whether shifts of attention can influence numerical processing. Participants viewed a leftward or rightward peripheral cue followed by a centrally presented number, then judged whether the number was odd or even. Participants responded faster and made fewer errors when the number magnitude and response side were compatible, revealing a response-based SNARC effect. Participants also responded faster when the cue direction and response side were compatible, revealing a Simon effect. However, participants did not respond faster when the cue direction and number magnitude were compatible. Similar findings were observed when the association between numbers and space was relatively explicit. Moreover, although we failed to observe a response-based SNARC effect when number magnitude was directly relevant to observers' task, we observed a large Simon effect. Together, these findings suggest that although numbers and space share a common representation at the response level, attention does not play a substantial role in the spatial representation of numbers.