Reconciling the functions of even-skipped interneurons during crawling, swimming, and walking

In all bilaterally symmetric animals, movements across the body are coordinated by interneurons that traverse the midline. Recent work is beginning to tease apart the functional complexity of interneurons labeled by the homeodomain transcription factor even-skipped, which provide a phylogenetically-conserved source of commissural excitation during locomotion in both vertebrates and invertebrates. Here we review recent studies of the roles of even-skipped neurons during locomotion in flies (EL neurons), fishes, frogs, and mice (V0v neurons). Comparisons across species reveal commonalities, which include the functional organization of even-skipped circuits based on birth order, the link between increased muscular complexity and even-skipped neuron diversity, and the hierarchical organization of even-skipped circuits based on their control of escape versus exploratory movements. We discuss how stronger links between different species enable testable predictions to further the discovery of principles of locomotor network organization.