We investigated the transduction operation and function of the mystacial vibrissae, using a comparative morphological analysis and behavioral experiments in rats. Vibrissal architecture was documented in a series of mammals to identify evolutionary conserved features of vibrissal organization. As a result of this analysis, we distinguish between a frontal microvibrissal system and macrovibrissal system of the mystacial pad. The latter was invariably comprised of whiskers aligned in regular rows. In each row, whiskers were oriented perpendicular to the animal's rostrocaudal axis; all shared a specific dorsoventral orientation. In all species, progressing from rostral to caudal in any vibrissal row, there was a precisely exponential increase in whisker length. Each whisker appeared to act as a lever-like transducer, providing information as to whether or not--but not where--an individual vibrissa had been deflected. The rat's frontal microvibrissae system was found to have a vibrissa tip density that was about 40 times higher than that of the mystacial macrovibrissae. In behavioral studies spatial tasks and object recognition tasks were used to investigate (a) search behaviors; (b) single whisker movements; (c) object recognition ability; and (d) effects of selective macro- or microvibrissae removal on task performances. A clear distinction between the functional roles of macro- and microvibrissae was demonstrated in these studies. Mystacial macrovibrissae were critically involved in spatial tasks, but were not essential for object recognition. Microvibrissae were critically involved in object recognition tasks, but were not essential for spatial tasks. A synthesis of these morphological and behavioral data led to the following functional concept: The mystacial macrovibrissae row is a distance decoder. Its function is to derive head centered obstacle/opening contours at the various dorsoventral angles represented by vibrissal rows. This distance detector model is functionally very different from traditional concepts of whisker function, in which the mystacial whiskers were hypothesized to form a fine grain skin-like object-recognizing tactile surface.