Walking in circles: Linking high- and low-level parameter scaling of visually guided and spontaneous turning behaviour

The ability of animals to adjust their heading, i.e. to turn, is essential for all walking animals. While several studies have addressed how leg movement or inter-leg coordination may change during turning, relatively little is known about how turning-related changes scale with turn magnitude. Here, we used spontaneous and visually induced turns of unrestrained walking stick insects to test (i) how high-level parameters of unrestrained turning scale with low-level parameters of leg movement, and (ii) the effect of visual guidance on turning parameters. To this end, we used a step change in stationary landmark position in an open-field arena to constrain timing and magnitude of target-directed turns. These visually guided turns were compared with spontaneous turns in an all-white condition. We show that visually induced turns were walked at a larger forward velocity and had fewer short steps than spontaneous turns. The scaling of turning responses was dominated by an increase in turning duration (factor 1.87) rather than turning speed (factor 1.32). Increased rotational velocity correlated with reduced forward velocity, though with flexible timing of both effects. These changes were accompanied by larger shifts in step direction, as well as an increased asymmetry of step types between inner and outer legs, suggesting a mix of distinct turning strategies, that depend on overall turn angle. Future models on six-legged locomotion should thus consider the incorporation of more than one mechanism to govern turning.