TABLE XII. THIGMOTAXIS REACTIONS

Animal45913212729313335363738394142434445464748
I49210417166 6103 21193219115
II 5213189 5245221141136111947
III 2125193231122471 414 915
IV111 2 1 24412
Animal495152545658606264 Sum
Totals
I 611312471012 195
II 837581321 190
III 1 169510322 154
IV 11 21 33
572

In order to appreciate the significance of the figures in this table it is necessary to consider the amount of lateral surface with which it was possible to come in contact in each case. In IV of course it was zero, in III it was 76 cm. with four corners in close proximity, in II it was only 38 cm. and one corner, but the space was so narrow that there was practically a contact-surface on both sides, and in I there was 212 cm. of lateral surface with three corners. I mention corners in this connection because they were almost invariably occupied. If we examine the table with these facts in mind, we find, (1) that the number of animals resting freely without contact with any lateral surface is very small, only about 6% of the whole; (2) that the number of animals found in the narrow space between the box and the walls of the aquarium is very large in proportion to the length of the space: indeed the animals were frequently found wedged into this space three or four deep; (3) that the number of animals found in the box was probably due largely to the fact that they found in it a greater lateral contact-surface, particularly in the corners, than was possible outside.

Two or three minor considerations are of interest. The animals were frequently observed "on edge" about half out of the water, that is, with the ventral surface of the body pressed against the vertical surface against which they were resting. This was also observed where the water was so deep that none of the members could touch the bottom. It was perhaps on account of the quality of the surface affording a rougher contact that so large a number of the animals were found in contact with the wooden box rather than the smooth, slippery surface of the glass. In the centre of the aquarium a wooden stopper 2 cm. in diameter projected about 15 cm. above the surface of the water. Very often a crayfish would be found almost at the top of this stopper, completely out of the water. This tendency to climb was frequently observed in the light-reaction experiments, where the animals would climb up on any piece of wood that chanced to be left in the box. It reminds one of the tree-climbing crabs of the West and East Indies. Along the creeks of Ohio I have frequently seen crayfish that had climbed up on logs or sticks that projected some feet out of the water.

In the table we see decided evidences of "habit" in the sense of an animal returning to the same place which it had occupied. No. 5 has almost half the observations in the open, nos. 21, 37, and 39 showed a decided preference for the space between the box and the aquarium wall, while nos. 38, 44, and 52 were more frequently found on the inside of the box. This recurrence to a particular position also came out in the light-reaction work, where an individual would return to the same spot in the box for days at a time as soon as released.

From the above considerations we conclude that the crayfish is strongly positively thigmotactic and that this thigmotaxis probably plays a most important part in the life of the animal.

(2) Touch Reactions. Lemoine[250] investigated the reactions of crayfish to touch-stimuli and found that the plates of the telson, the sternal portions of the thorax, the abdominal pleopods, the chelæ, and particularly the antennæ toward their points are especially sensitive, but that nowhere, even on the back of the carapace, is a touch-stimulus altogether devoid of reaction. Gulland[251] found that a needle could be inserted between the tufts of setæ on the chelæ without causing any reaction, but as soon as one of the hairs was touched, the chelæ closed with a snap. Considering the setæ as the organs of touch, he claimed to have found that the eyes, eye-stalks, and carapace (which he says have no setæ) are impervious to tactile impressions. This claim of Gulland's is strangely at variance with the facts. In no case have I been able to bring about retraction of the eye-stalk by visual stimulation, but a very light touch-stimulus on the eye itself or on the eye-stalk or a stronger stimulus on some portion of the head will cause the eye to be drawn in. It is true that after repeated stimulation the eye is retracted no longer, and with a heavy bristle one can make a perceptible indentation in the corneal surface without the eye being withdrawn.

The antennæ, from their anatomical structure, their position, and the manner in which they are carried, are generally considered the special organs of touch. Nevertheless, as far as the reactions of the animal are concerned, a stimulation of the antennæ by touch produces a less decided response than almost any other portion of the body. If the stimulus is very light no reaction at all is observed in most cases, and if stronger the antennæ are moved away, but that is all. A stimulation of the edge of the telson produces a more decided reaction. Either the animal folds it under the abdomen at once or faces about like a flash in an attitude of defence; frequently both reactions occur. While the response to stimulation of the chelæ was decided, that to touch on the first chelipedes was quicker and more accurate. The mouth-parts are also very sensitive to touch. I cannot agree with Gulland's assertions as to the insensitiveness of the carapace, for I have been able to find no place upon it where a light-stimulation would not produce a reaction. In this connection a curious phenomenon is characteristic of the animal. If the carapace or the front portion of the abdomen be lightly stroked with a solid object such as a pencil, the animal will slowly turn toward the stimulus on its antero-posterior axis. If, now, a like stimulus be applied on the other side, the animal will roll back through the normal position to a like inclination toward the stimulus on the other side. If the alternation be kept up and the change made quickly, a continuous and curious rolling movement is maintained, the animal growing more and more excited until it scampers off with a kind of cramp-like motion. With some animals this rolling reflex is more marked than with others, but in no case is it altogether lacking. Some animals have been known to roll so far over that they topple over on their backs. Dr. Yerkes informs me that he has observed the same phenomenon in a less degree with turtles when the edge of their shell is stimulated by scratching. The movement seems to be caused by the reflex stimulation of the extensor muscles on the opposite side of the body from the part stimulated. The thrust of the legs thereby brought about raises that side of the body and thus causes a rotation to some extent about the antero-posterior axis. But how was this connection between the stimulation of one side of the body and the contraction of the extensor muscles of the other side established? I have no doubt that it is intimately connected with the positive thigmotaxis described above. These animals live under loose stones for the most part, and thus the carapace gets a great deal of stimulation. If the animal is stimulated on one side, a contraction of the extensor muscles of the opposite side tends to roll the animal toward the source of the stimulus, and hence to increase the contact. In the race-history of the animal this has doubtless been advantageous in enabling it to escape the dangers of its habitat.