“The Proprioceptive Ability of the White Rat”, 1931 (; backlinks):
With no exteroceptive differential stimuli beyond the starting alley, the rat cannot except infrequently run to a spot a fixed distance ahead as is required by any maze alley. It is certain that the rat cannot run a maze consistently when limited to proprioceptive differential cues. Proprioception may be an important factor in the sensory control of the maze habit, but in any case other cues as well as proprioceptive cues must be present.
The problem was to determine whether a simple maze could be accurately run upon the basis of proprioceptive cues alone. In the first experiment the rats were required to locate a wire ladder in the center of an open space, the only orientation being that given them by a short alley through which they entered the space. At the top of the ladder they found an elevated pathway leading to food. The apparatus was constantly rotated to eliminate fixed extra-apparatus stimuli.
Rats trained in the light could not directly locate the ladder in darkness. Blind rats were unable to exceed an accuracy of 7% in locating the ladder. Similar results were obtained when the animals were placed in the center of a circular apparatus and required to locate food at the edge of the circle straight ahead from the small entrance alley. The small central alley was oriented so that the only constant feature of the situation was that of going straight ahead after leaving the alley.
Normal animals learned the problem, but blinded animals had an average error of 15.2°. The author claims that proprioception is not accurate enough to explain maze orientation. Exteroceptive stimuli are essential for accurate performance.