The femur-tibia control system in a proscopiid (Caelifera, Orthoptera): a test for assumptions on the functional basis and evolution of twig mimesis in stick insects.

The extremely slow return movements observed in stick insects (phasmids) after imposed changes in posture are termed catalepsy. In the literature, catalepsy is treated as a behavioural component of the twig mimesis observed in walking stick insects. It is produced by the high gain of the velocity-sensitive component of the relevant joint control systems and by the non-linear dependency of its time constant on movement velocity. The high gain, in turn, causes the system to work close to instability, and this may have driven the evolution of gain control mechanisms. Although these statements represent plausible assumptions, based on correlated occurrence, they remain largely hypothetical like many ideas concerning evolutionary tendencies. To test these hypotheses, we studied catalepsy and the relevant properties of the femur-tibia control system in the middle and hind legs of Prosarthria teretrirostris.cf. Prosarthria teretrirostris is a proscopiid closely related to grasshoppers and locusts. With its slender, green-to-brown body and legs, it shows clear morphological twig mimesis, which has evolved independently of the well-known twig mimesis in stick insects. The animals show clear catalepsy. The main properties of femur-tibia joint control are remarkably similar between proscopiids and stick insects (e.g. the marked sensitivity to movement velocity rather than to joint position and the non-linear dependency of the time constants of response decay on movement velocity), but there are also important differences (habituation and activity-related mechanisms of gain control are absent). Together, these results validate the main concepts that have been developed concerning the neural basis and evolution of catalepsy in stick insects and its relationship to twig mimesis, while demonstrating that ideas on the role of habituation and gain control should be refined.