The dynamic components of the fixation behaviour of walking Colorado beetles were examined. The beetles walked on top of a locomotion compensator under closed-loop conditions. The fixation target was a yellow disk with small black stripes (‘Colorado dummy’). The paths of beetles walking towards the stationary dummy were characterized by oscillations, and were investigated by applying a fast Fourier transform to the turning-velocity time course. The power density peaks near 0.3 Hz with varying amplitude. Films of the fixation reaction revealed that this frequency was much slower than the stepping-pattern frequency of the beetles. During orientation to other stimuli (wind or gravity), the oscillations were not present, showing that they were not an artefact of the motor output. However, they were present in other forms of visual orientation, such as the optomotor response and during positive phototaxis.
A frequency-response curve (Bode diagram) under closed-loop conditions was obtained for the fixation reaction by changing the dummy position sinusoidally and measuring the phase and amplitude of the tracking response. A simple model of this behaviour, including a low-pass filter and an integrator, describes the reaction. Hence, the reaction can be understood as the simple resonance behaviour of an underdamped closed-position loop. The question of whether this can explain zigzagging paths in other species is discussed.
Present address: Universität Kaiserslautern, FB Biologie, Abteilung Allgemeine Zoologie, D-6750 Kaiserslautern, Germany.