Animal navigation: general properties of directed walks.

The ability to locomote is a defining characteristic of all animals. Yet, all but the most trivial forms of navigation are poorly understood. Here we report and discuss the analytical results of an in-depth study of a simple navigation problem. In principle, there are two strategies for navigating a straight course. One is to use an external directional reference and to continually reorient with reference to it. The other is to monitor body rotations from internal sensory information only. We showed previously that, at least for simple representations of locomotion, the first strategy will enable an animal or mobile agent to move arbitrarily far away from its starting point, but the second strategy will not do so, even after an infinite number of steps. This paper extends and generalizes the earlier results by demonstrating that these findings are true even when a very general model of locomotion is used. In this general model, error components within individual steps are not independent, and directional errors may be biased. In the absence of a compass, the expected path of a directed walk in general approximates a logarithmic spiral. Some examples are given to illustrate potential applications of the quantitative results derived here. Motivated by the analytical results developed in this work, a nomenclature for directed walks is proposed and discussed. Issues related to path integration in mammals and robots, and measuring the curvature of a noisy path are also addressed using directed walk theory.