What nervous systems do: Early evolution, input-output, and the skin brain thesis

Nervous systems are standardly interpreted as information processing input-output devices. They receive environmental information from their sensors as input, subsequently process or adjust this information, and use the result to control effectors, providing output. Through-conducting activity is here the key organizational feature of nervous systems. In this paper, we argue that this input-output interpretation is not the most fundamental feature of nervous system organization. Building on biological work on the early evolution of nervous systems, we provide an alternative proposal: the skin brain thesis (SBT). The SBT postulates that early nervous systems evolved to organize a new multicellular effector: muscle tissue, the primary source of animal motility. Early nervous systems provided a new way of inducing and coordinating self-organized contractile activity across an extensive muscle surface underneath the skin. The main connectivity in such nervous systems runs across a spread out effector and is transverse to sensor-effector signaling. The SBT therefore constitutes a fundamental conceptual shift in understanding both nervous system operation and what nervous systems are. Nervous systems are foremost spatial organizers that turn large multi-cellular animal bodies into dynamic self-moving units. At the end, we briefly discuss some theoretical connections to central issues within the behavioral, cognitive and neurosciences.