title: Cellular scaling rules for the brain of Artiodactyla include a highly folded cortex with few neurons.
abstract: Quantitative analysis of the cellular composition of rodent, primate, insectivore, and afrotherian brains has shown that non-neuronal scaling rules are similar across these mammalian orders that diverged about 95 million years ago, and therefore appear to be conserved in evolution, while neuronal scaling rules appear to be free to vary in a clade-specific manner. Here we analyze the cellular scaling rules that apply to the brain of artiodactyls, a group within the order Cetartiodactyla, believed to be a relatively recent radiation from the common Eutherian ancestor. We find that artiodactyls share non-neuronal scaling rules with all groups analyzed previously. Artiodactyls share with afrotherians and rodents, but not with primates, the neuronal scaling rules that apply to the cerebral cortex and cerebellum. The neuronal scaling rules that apply to the remaining brain areas are, however, distinct in artiodactyls. Importantly, we show that the folding index of the cerebral cortex scales with the number of neurons in the cerebral cortex in distinct fashions across artiodactyls, afrotherians, rodents, and primates, such that the artiodactyl cerebral cortex is more convoluted than primate cortices of similar numbers of neurons. Our findings suggest that the scaling rules found to be shared across modern afrotherians, glires, and artiodactyls applied to the common Eutherian ancestor, such as the relationship between the mass of the cerebral cortex as a whole and its number of neurons. In turn, the distribution of neurons along the surface of the cerebral cortex, which is related to its degree of gyrification, appears to be a clade-specific characteristic. If the neuronal scaling rules for artiodactyls extend to all cetartiodactyls, we predict that the large cerebral cortex of cetaceans will still have fewer neurons than the human cerebral cortex.
authors.affiliation: Instituto de Ciências Biomédicas, Universidade Federal do Rio de Janeiro Rio de Janeiro, Brazil ; Instituto Nacional de Neurociência Translacional, CNPq/MCT, São Paulo, Brazil.
authors.forename: Rodrigo S
authors.initials: RS
authors.lastname: Kazu
authors.affiliation: Microbrightfield Biosciences, Burlington VT, USA.
authors.forename: José
authors.initials: J
authors.lastname: Maldonado
authors.affiliation: Instituto de Física, Universidade Federal do Rio de Janeiro Rio de Janeiro, Brazil.
authors.forename: Bruno
authors.initials: B
authors.lastname: Mota
authors.affiliation: School of Anatomical Sciences, University of the Witwatersrand Johannesburg, South Africa.
authors.forename: Paul R
authors.initials: PR
authors.lastname: Manger
authors.affiliation: Instituto de Ciências Biomédicas, Universidade Federal do Rio de Janeiro Rio de Janeiro, Brazil ; Instituto Nacional de Neurociência Translacional, CNPq/MCT, São Paulo, Brazil.
authors.forename: Suzana
authors.initials: S
authors.lastname: Herculano-Houzel
keywords: brain size; cortical expansion; evolution; gyrification; numbers of neurons