III. ROTATION EXPERIMENTS
It has been found that the higher vertebrates, on being rotated on a turn-table, exhibit all the symptoms which accompany the sensation of dizziness in man. The question arises, to what degree and in what manner do invertebrates respond to rotation? Schaefer,[243] the first to take up this question, denied on rather meagre observations that Crustacea respond in any way to rotation on the turn-table. Kreidl[244] showed that this statement was altogether too sweeping, that Palæmon reacts very definitely to rotation by running in the opposite direction. Bunting[245] tried the crayfish, but all the rotation experiments resulted negatively, so she was led to confirm Schaefer's statement so far as the crayfish is concerned. Bethe[246] found that Carcinus behaved in a very definite manner on being rotated, that during the rotation the animals ran in the opposite direction to that in which they were turned, and as soon as the motion ceased they began running in the other direction. Finally Lyon,[247] while agreeing with Bunting that adult crayfish do not react to rotation, discovered that young animals two or three centimetres long react very prettily to the movement, going in a direction opposite to the turn. To confirm and if possible extend these observations on the crayfish was the purpose of the following experiments.
It was soon found that a great deal depended on the method of experimentation. None of the experimenters mentioned above gives any detailed description of the manner in which the experiments were carried out. One is left uncertain whether the animals were placed on the periphery of the turn-table or over the centre, whether in the former case they were set with their heads toward the centre or away from it, or placed at right angles to a radius, or whether they were merely set down in any chance fashion and whirled about. The same indefiniteness exists in most of the accounts as to how fast they were turned, and whether the experiments were performed in the air or in the water. Finally it is not stated whether the rotation was always in the same direction, or whether its direction was alternated.
The turn-table used in the following experiments was one that had to be turned by hand, so that it was impossible to regulate the speed accurately. The crank, however, was not attached directly to the rotating board, but was connected with it by means of a gearing so that one turn of the crank produced about ten turns of the table. This gearing gave a steadying effect to the motion so that the speed could be kept tolerably constant. A circular pan, about 15 cm. in diameter at the bottom with the sides slightly sloping outward, was set so that its centre coincided with the axis of the rotating table. It was in this pan that all the experiments were tried. Through various preliminary experiments to determine the most favorable speed, it was found that a rotation rate of over one turn of the table per second produced such a strong centrifugal force that unless the animals were set exactly over the centre they were swept off against the side of the pan in such a manner that it was difficult to decide whether the rotation as such had any effect upon their movements. It was finally decided that the best results were obtained from a rate of approximately one rotation in two seconds.
It soon became evident that when the larger and more sluggish crayfish were merely dropped in the pan and rotated there was no particular reaction. This was true whether the animals rotated were in the air or in the water. The smaller and more active crayfish, however, showed a decided tendency to run either with or against the direction of the rotation, especially when the experiments were carried on in the water. In no case was there any tendency to go in the opposite direction when the rotation ceased, except in so far as the animals were carried along by the water. To get a quantitative expression for these tendencies a more delicate method of experimentation was resorted to. If there was a tendency on the part of the active animal to move either with or against the rotation, such a tendency might also be supposed to exist in the sluggish animal, only in the latter the inertia was sufficiently strong to prevent its appearance. If, however, the animals should be set radially to the periphery of the pan, the tendency to go with or against the rotation would be exhibited in the direction in which they turned out of the radial position. For it was found that no animal would remain in that position for any great length of time. Two groups of animals were used for these experiments, five animals in each group, and the first group was selected from the smallest and most active animals, the second from the largest and most sluggish. Each animal was set in two positions, position I, with the head toward the centre, position II, with the head away from the centre. Each animal was given ten trials in each position, and the number of times it turned in a direction with the rotation is set down in the + column, the number of times it turned against the rotation is indicated in the - column. In general from 5 to 15 turns were necessary for the orientation of the animal, though sometimes the number ran up to 30 or 40. Each trial was made in the opposite direction to the preceding one, in order to avoid the formation of any habit in turning. All these experiments were carried out in water, the depth of which in the pan was about 4 cm. In order that there should be no difference between the velocity of the water and that of the pan, the table was rotated a few times before the animal was put in. As a check a series of experiments of 5 in each position was performed in the air on the more active group.
The following table shows the results of these experiments in rotation: