Active touch sensing in the rat: anticipatory and regulatory control of whisker movements during surface exploration.

Animals actively regulate the position and movement of their sensory systems to boost the quality and quantity of the sensory information they obtain. The rat vibrissal system is recognized to be an important model system in which to investigate such "active sensing" capabilities. The current study used high-speed video analysis to investigate whisker movements in untrained, freely moving rats encountering unexpected, vertical surfaces. A prominent feature of rat vibrissal movement is the repeated posterior-anterior sweep of the whiskers in which the macrovibrissae are seen to move largely in synchrony. Here we show that a second significant component of whisking behavior is the size of the arc, or "spread," between the whiskers. Observed spread is shown to vary over the whisk cycle and to substantially decrease during exploration of an unexpected surface. We further show that the timing of whisker movements is affected by surface contact such that 1) the whiskers rapidly cease forward protraction following an initial, unexpected contact, and may do so even more rapidly following contact with the same surface in the subsequent whisk cycle, and 2) retraction velocity is reduced following this latter contact, leading to longer second-contact durations. This evidence is taken to support two hypotheses: 1) that the relative velocities of different whiskers may be actively controlled by the rat and 2) that control of whisker velocity and timing may serve to increase the number and duration of whisker-surface contacts while ensuring that such contacts are made with a light touch.