The Dancing Mouse A Study in Animal Behavior - Chapter III

BEHAVIOR: DANCE MOVEMENTS

The peculiarities of behavior of the dancing mouse are responsible alike for the widespread interest which it has aroused, and for its name. In a little book on fancy varieties of mice, in which there is much valuable information concerning the care of the animals, one who styles himself "An old fancier" writes thus of the behavior of the dancer: "I believe most people have an idea that the waltzing is a stately dance executed on the hind feet; this is not so. The performer simply goes round and round on all fours, as fast as possible, the head pointing inwards. The giddy whirl, after continuing for about a dozen turns, is then reversed in direction, and each performance usually occupies from one to two minutes. Whether it is voluntary or not, is difficult to determine, but I am inclined to think the mouse can refrain if it wishes to do so, because I never see them drop any food they may be eating, and begin to waltz in the midst of their meal. The dance, if such it can be called, generally seizes the mouse when it first emerges from its darkened sleeping place, and this would lead one to suppose that the light conveys an impression of shock to the brain, through the eyes, which disturbs the diseased centers and starts the giddy gyrations. The mice can walk or run in a fairly straight line when they wish to do so." Some of the old fancier's statements are true, others are mere guesses. Those who have studied the mice carefully will doubtless agree that he has not adequately described the various forms of behavior of which they are capable. I have quoted his description as an illustration of the weakness which is characteristic of most popular accounts of animal behavior. It proves that it is not sufficient to watch and then describe. The fact is that he who adequately describes the behavior of any animal watches again and again under natural and experimental conditions, and by prolonged and patient observation makes himself so familiar with his subject that it comes to possess an individuality as distinctive as that of his human companions. To the casual observer the individuals of a strange race are almost indistinguishable. Similarly, the behavior of all the animals of a particular species seems the same to all except the observer who has devoted himself whole-heartedly to the study of the subject and who has thus become as familiar with their life of action as most of us are with that of our fellow-men; for him each individual has its own unmistakable characteristics.

I shall now describe the behavior of the dancing mouse in the light of the results of the observation of scores of individuals for months at a time, and of a large number of experiments. From time to time I shall refer to points in the accounts of the subject previously given by Rawitz (25 p. 236), Cyon (9 p. 214), Alexander and Kreidl (1 p. 542), Zoth (31 p. 147), and Kishi (21 p. 479).

The most striking features of the ordinary behavior of the dancer are restlessness and movements in circles. The true dancer seldom runs in a straight line for more than a few centimeters, although, contrary to the statements of Rawitz and Cyon, it is able to do so on occasion for longer distances. Even before it is old enough to escape from the nest it begins to move in circles and to exhibit the quick, jerky head movements which are characteristic of the race. At the age of three weeks it is able to dance vigorously, and is incessantly active when not washing itself, eating, or sleeping. According to Zoth (31 p. 149) the sense of sight and especially the sense of smell of the dancer "seem to be keenly developed; one can seldom remain for some time near the cage without one or another of the animals growing lively, looking out of the nest, and beginning to sniff around in the air (windet). They also seem to have strongly developed cutaneous sensitiveness, and a considerable amount of curiosity, if one may call it such, in common with their cousin, the white mouse." I shall reserve what I have to say concerning the sense of sight for later chapters. As for the sense of smell and the cutaneous sensitiveness, Zoth is undoubtedly right in inferring from the behavior of the animal that it is sensitive to certain odors and to changes in temperature. One of the most noticeable and characteristic activities of the dancer is its sniffing. Frequently in the midst of its dancing it stops suddenly, raises its head so that the nose is pointed upward, as in the case of one of the mice of the frontispiece, and remains in that position for a second or two, as if sniffing the air.

The restlessness, the varied and almost incessant movements, and the peculiar excitability of the dancer have repeatedly suggested to casual observers the question, why does it move about in that aimless, useless fashion? To this query Rawitz has replied that the lack of certain senses compels the animal to strive through varied movements to use to the greatest advantage those senses which it does possess. In Rawitz's opinion the lack of hearing and orientation is compensated for by the continuous use of sight and smell. The mouse runs about rapidly, moves its head from side to side, and sniffs the air, in order that it may see and smell as much as possible. In support of this interpretation of the restlessness of the dancer, Rawitz states that he once observed similar behavior in an albino dog which was deaf. This suggestion is not absurd, for it seems quite probable that the dancer has to depend for the guidance of its movements upon sense data which are relatively unimportant in the common mouse, and that by its varied and restless movements it does in part make up for its deficiency in sense equipment.

The dancing, waltzing, or circus course movement, as it is variously known, varies in form from moment to moment. Now an individual moves its head rapidly from side to side, perhaps backing a little at the same time, now it spins around like a top with such speed that head and tail are almost indistinguishable, now it runs in circles of from 5 cm. to 30 cm. in diameter. If there are any objects in the cage about or through which it may run, they are sure to direct the expression of activity. A tunnel or a hole in a box calls forth endless repetitions of the act of passing through. When two individuals are in the same cage, they frequently dance together, sometimes moving in the same direction, sometimes in opposite directions. Often, as one spins rapidly about a vertical axis, the other runs around the first in small circles; or again, both may run in a small circle in the same direction, so that their bodies form a living ring, which, because of the rapidity of their movements, appears perfectly continuous. The three most clearly distinguishable forms of dance are (1) movement in circles with all the feet close together under the body, (2) movement in circles, which vary in diameter from 5 cm. to 30 cm., with the feet spread widely, and (3) movement now to the right, now to the left, in figure eights ([Symbol: figure eight]). For convenience of reference these types of dance may be called whirling, circling, and the figure eight dance. Zoth, in an excellent account of the behavior of the dancer (31 p. 156), describes "manège movements," "solo dances," and "centre dances." Of these the first is whirling, the second one form of circling, and the third the dancing of two individuals together in the manner described above.

Both the whirling and the circling occur to the right (clockwise) and to the left (anticlockwise). As certain observers have stated that it is chiefly to the left and others that it is as frequently to the right, I have attempted to get definite information concerning the matter by observing a number of individuals systematically and at stated intervals. My study of this subject soon convinced me that a true conception of the facts cannot be got simply by noting the direction of turning from time to time. I therefore planned and carried out a series of experimental observations with twenty dancers, ten of each sex. One at a time these individuals were placed in a glass jar, 26 cm. in diameter, and the number of circle movements executed to the right and to the left during a period of five minutes was determined as accurately as possible. This was repeated at six hours of the day: 9 and 11 o'clock A.M., and 2, 4, 6, and 8 o'clock P.M. In order that habituation to the conditions under which the counts of turning were made might hot influence the results for the group, with ten individuals the morning counts were made first, and with the others the afternoon counts. No attempt was made in the counting to keep a separate record of the whirling and circling, although had it been practicable this would have been desirable, for, as soon became evident to the observer, some individuals which whirl in only one direction, circle in both.

In Table 2 the results of the counts for the males are recorded; in Table 3 those for the females. Each number in the column headed "right" and "left" indicates the total number of circles executed by a certain dancer in a period of five minutes at the hour of the day named at the head of the column. I may point out briefly the curiously interesting and entirely unexpected new facts which this method of observation revealed to me.

First, there are three kinds of dancers: those which whirl almost uniformly toward the right, those which whirl just as uniformly toward the left, and those which whirl about as frequently in one direction as in the other. To illustrate, No. 2 of Table 2 may be characterized as a "right whirler," for he turned to the right almost uniformly. In the case of the 6 P.M. count, for example, he turned 285 times to the right, not once to the left. No. 152, on the contrary, should be characterized as a "left whirler," since he almost always turned to the left. From both of these individuals No. 210 is distinguished by the fact that he turned now to the left, now to the right. For him the name "mixed whirler" seems appropriate.

Second, the amount of activity, as indicated by the number of times an individual turns in a circle within five minutes, increases regularly and rapidly from 9 A.M. to 8 P.M. According to the general averages which appear at the bottom of Table 2, the average number of circles executed by the males at 9 A.M. was 89.8 as compared with 207.1 at 8 P.M. In other words, the mice dance more in the evening than during the day.

Third, as it appears in a comparison of the general averages of Tables 2 and 3, the females dance more than the males, under the conditions of observation. At 9 A.M. the males circled 89.8 times, the females 151.0 times; at 8 P.M. the males circled 207.1 times, the females, 279.0 times.

Fourth, according to the averages for the six counts made with each individual, as they appear in Table 4, the males turn somewhat more frequently to the left than to the right (the difference, however, is not sufficient to be considered significant); whereas, the females turn much more frequently to the right than to the left. I do not wish to emphasize the importance of this difference, for it is not improbable that counts made with a larger number of animals, or even with another group of twenty, would yield different results.

TABLE 2

NUMBER or WHIRLS TO THE RIGHT AND TO THE LEFT DURING
FIVE-MINUTE INTERVALS AS DETERMINED BY COUNTS MADE AT
SIX DIFFERENT HOURS, FOR EACH OF TEN MALE DANCERS

NUMBER 9 A.M 11 A.M. 2 P.M. OF ANIMAL RIGHT LEFT RIGHT LEFT RIGHT LEFT

2 11 2 23 4 194 1 30 20 1 134 1 109 2 34 2 16 2 48 4 92 36 194 21 180 11 143 65 152 7 48 3 171 6 79 156 63 8 53 9 27 6 210 3 9 7 41 225 21 220 168 105 39 43 47 5 410 2 67 10 27 8 103 420 15 142 5 214 16 238

Averages 48.5 41.3 45.6 56.9 77.9 61.2

Gen. Av. 89.8 102.5 139.1

NUMBER 4 P.M 6 P.M. 8 P.M. OF ANIMAL RIGHT LEFT RIGHT LEFT RIGHT LEFT

2 70 3 285 0 237 10 30 154 0 107 6 134 5 34 7 158 5 118 6 147 36 173 14 170 11 325 19 152 0 91 16 210 9 223 156 85 2 72 26 139 26 210 159 18 31 82 47 201 220 45 38 78 17 69 33 410 9 155 9 394 24 94 420 18 243 16 291 3 320

Averages 72.0 72.2 78.9 115.5 99.3 107.8

Gen. Av. 144.2 194.4 207.1

TABLE 3

NUMBER OF WHIRLS TO THE RIGHT AND TO THE LEFT DURING FIVE-MINUTE INTERVALS AS DETERMINED BY COUNTS MADE AT SIX DIFFERENT HOURS, FOR EACH OF TEN FEMALE DANCERS

NUMBER 9 A.M. 11 A.M. 2 P.M. OF ANIMAL RIGHT LEFT RIGHT LEFT RIGHT LEFT

29 9 18 17 30 7 22 33 287 0 329 1 352 3 35 48 15 198 46 208 14 151 13 88 7 75 3 167 157 57 6 50 45 53 12 211 218 21 31 55 66 5 215 67 216 33 105 37 226 225 46 39 72 49 143 44 415 23 0 156 0 34 3 425 43 296 12 201 12 210

Averages 81.1 69.9 90.5 60.7 91.5 70.6

Gen. Av. 151.0 151.2 162.1

NUMBER 4 P.M. 6 P.M. 8 P.M. OF ANIMAL RIGHT LEFT RIGHT LEFT RIGHT LEFT

29 33 114 31 36 45 99 33 436 7 408 3 364 2 35 279 6 165 24 353 10 151 3 8 2 285 2 217 157 52 15 19 125 51 104 211 190 7 86 31 67 250 215 15 292 45 336 150 232 225 133 86 48 39 177 81 415 268 3 437 7 382 8 425 12 242 19 210 4 192

Averages 142.1 78.0 126.0 109.6 159.5 119.5

Gen. Av. 220.1 235.6 279.0

The most important results of this statistical study of turning are the demonstration of the existence of individual tendencies to turn in a particular direction, and of the fact that the whirling increases in amount from morning to evening.

In order to discover whether the distribution of the dancers among the three groups which have been designated as right, left, and mixed whirlers agrees in general with that indicated by Table 4 (approximately the same number in each group) I have observed the direction of turning in the case of one hundred dancers, including those of the foregoing tables, and have classified them in accordance with their behavior as is indicated below.

RIGHT LEFT MIXED WHIRLERS WHIRLERS WHIRLERS

Males 19 19 12
Females 12 23 15

Totals 31 42 27

The left whirlers occur in excess of both the right and the mixed whirlers. This fact, together with the results which have already been considered in connection with the counts of turning, suggests that a tendency to whirl in a certain way may be inherited. I have examined my data and conducted breeding experiments for the purpose of ascertaining whether this is true. But as the results of this part of the investigation more properly belong in a special chapter on the inheritance of behavior (XVIII), the discussion of the subject may be closed for the present with the statement that the preponderance of left whirlers indicated above is due to a strong tendency to turn to the left which was exhibited by the individuals of one line of descent.

TABLE 4

AVERAGE NUMBER OF WHIRLS TO THE RIGHT AND TO THE LEFT FOR THE SIX INTERVALS OF TABLES 2 AND 3, WITH A CHARACTERIZATION OF THE ANIMALS AS RIGHT WHIRLERS, LEFT WHIRLERS, OR MIXED WHIRLERS.

AVERAGE NO. AVERAGE NO. MALES AGE OF WHIRLS OF WHIRLS CHARACTERIZATION

    2 12 mo. 136.7 3.3 Right whirler
   30 2 mo. 109.7 2.5 Right whirler
   34 2 mo. 4.3 96.5 Left whirler
   36 2 mo. 197.5 23.5 Right whirler
  152 6 mo. 6.8 137.0 Left whirler
  156 1 mo. 73.2 12.8 Right whirler
  210 3 mo. 78.7 62.0 Mixed whirler
  220 4 mo. 74.3 40.2 Mixed whirler
  410 3 mo. 10.3 139.0 Left whirler
  420 3 mo. 12.2 241.3 Left whirler

Average 70.4 75.8 4 Right whirlers 4 Left whirlers 2 Mixed whirlers

FEMALES

29 2 mo. 23.7 53.2 Left whirler 33 2 mo. 362.7 2.7 Right whirler 35 2 mo. 208.5 19.2 Left whirler 151 6 mo. 5.0 140.0 Right whirler 157 1 mo. 47.0 51.2 Left whirler 211 3 mo. 109.7 61.5 Right whirler 215 3 mo. 57.8 234.5 Mixed whirler 225 4 mo. 103.2 56.3 Mixed whirler 415 3 mo. 216.7 3.5 Left whirler 425 3 mo. 17.0 225.2 Left whirler

Average 115.1 84.7 3 Right whirlers 4 Left whirlers 3 Mixed whirlers

The tendency of the dancer's activity to increase in amount toward evening, which the results of Tables 2, 3, and 4 exhibit, demands further consideration. Haacke (7 p. 337) and Kishi (21 p. 458) agree that the dancing is most vigorous in the evening; but Alexander and Kreidl (i p. 544) assert, on the contrary, that the whirling of the individuals which they observed bore no definite relation to the time of day and apparently was not influenced in intensity thereby. Since the results of my own observations contradict many of the statements made by the latter authors, I suspect that they may not have watched their animals long enough to discover the truth. The systematic records which I have kept indicate that the mice remain quietly in their nests during the greater part of the day, unless they are disturbed or come out to obtain food. Toward dusk they emerge and dance with varying intensity for several hours. I have seldom discovered one of them outside the nest between midnight and daylight. The period of greatest activity is between 5 and 10 o'clock P.M.

Zoth states that he has observed the adult dancer whirl 79 times without an instant's interruption, and I have counted as many as 110 whirls. It seems rather absurd to say that an animal which can do this is weak. Evidently the dancer is exceptionally strong in certain respects, although it may be weak in others. Such general statements as are usually made fail to do justice to the facts.

The supposition that light determines the periodicity of dancing is not borne out by my observations, for I have found that the animals continue to dance most vigorously toward evening, even when they are kept in a room which is constantly illuminated. In all probability the periodicity of activity is an expression of the habits of the mouse race rather than of the immediate influence of any environmental condition. At some time in the history of the dancer light probably did have an influence upon the period of activity; but at present, as a result of the persistence of a well-established racial tendency, the periodicity of dancing depends to a greater extent upon internal than upon external conditions. During its hours of quiescence it is possible to arouse the dancer and cause it to whirl more or less vigorously by stimulating it strongly with intense light, a weak electric current, or by placing two individuals which are strangers to one another in the same cage; but the dancing thus induced is seldom as rapid, varied, or as long-continued as that which is characteristic of the evening hours.

One of the most interesting results of this study of the direction of turning, from the observer's point of view, is the demonstration of the fact that the truth concerning even so simple a matter as this can be discovered only by long and careful observation. The casual observer of the dancer gets an impression that it turns to the left more often than to the right; he verifies his observation a few times and then asserts with confidence that such is the truth about turning. That such a method of getting knowledge of the behavior of the animal is worse than valueless is clear in the light of the results of the systematic observations which have just been reported. But, however important the progress which we may have made by means of systematic observation of the phenomenon of turning, it must not for one moment be supposed that the whole truth has been discovered. Continued observation will undoubtedly reveal other important facts concerning circling, whirling, and the periodicity of dancing, not to mention the inheritance of peculiarities of dancing and the significance of the various forms of activity.

CHAPTER IV

BEHAVIOR: EQUILIBRATION AND DIZZINESS

Quite as interesting and important as the general facts of behavior which we have been considering are the results of experimental tests of the dancer's ability to maintain its position under unusual spatial conditions—to climb, cross narrow bridges, balance itself on high places. Because of its tendency to circle and whirl, to dart hither and thither rapidly and apparently without control of its movements, the study of the mouse's ability to perform movements which demand accurate and delicate muscular coördination, and to control its expressions of activity, are of peculiar scientific interest.

That observers do not entirely agree as to the facts in this field is apparent from the following comparison of the statements made by Cyon and Zoth (31 p. 174).

Cyon states that the dancer

Cannot run in a straight line,
Cannot turn in a narrow space,
Cannot run backward,
Cannot run up an incline,
Cannot move about safely when above the ground, because of
  fear and visual dizziness,
Can hear certain tones.

Zoth, on the contrary, maintains that the animal

Can run in a straight line for at least 20 cm.,
Can and repeatedly does turn in a narrow space,
Can run backward, for he has observed it do so,
Can run up an incline unless the surface is too smooth for it to
  gain a foothold,
Can move about safely when above the ground, and gives no
  signs of fear or dizziness,
Cannot hear, or at least gives no signs of sensitiveness to sounds.

Such contradictory statements (and unfortunately they are exceedingly common) stimulated me to the repetition of many of the experiments which have been made by other investigators to test the dancer's behavior in unusual spatial relations. I shall state very briefly the general conclusions to which these experiments have led me, with only sufficient reference to methods and details of results to enable any one who wishes to repeat the tests for himself to do so. For the sake of convenience of presentation and clearness, the facts have been arranged under three rubrics: equilibrational ability, dizziness, and behavior when blinded. To our knowledge of each of these three groups of facts important contributions have come from the experiments of Cyon (9 p. 220), Alexander and Kreidl (1 p. 545), Zoth (31 p. 157), and Kishi (21 p. 482), although, as has been stated, in many instances their results are so contradictory as to demand reexamination. All in all, Zoth has given the most satisfactory account of the behavior and motor capacity of the dancer.

If the surface upon which it is moving be sufficiently soft or rough to furnish it a foothold, the dancer is able to run up or down inclines, even though they be very steep, to cross narrow bridges, to balance itself at heights of at least 30 cm. above the ground, and even to climb up and down on rods, as is shown by certain of Zoth's photographs which are reproduced in Figure 4. Zoth himself says, and in this I am able fully to agree with him on the basis of my own observations, "that the power of equilibration in the dancing mouse, is, in general, very complete. The seeming reduction which appears under certain conditions should be attributed, not to visual dizziness, but in part to excitability and restlessness, and in part to a reduced muscular power" (31 p. 161). The dancer certainly has far less grasping power than the common mouse, and is therefore at a disadvantage in moving about on sloping surfaces. One evidence of this fact is the character of the tracks made by the animal. Instead of raising its feet from the substratum and placing them neatly, as does the common mouse (Figure 5), it tends to shuffle along, dragging its toes and thus producing on smoked paper such tracks as are seen in Figure 6. From my own observations I am confident that these figures exaggerate the differences. My dancers, unless they were greatly excited or moving under conditions of stress, never dragged their toes as much as is indicated in Figure 6. However, there can be no doubt that they possess less power of grasping with their toes than do common mice. The animal is still further incapacitated for movement on inclined surfaces or narrow places by its tendency to move in circles and zigzags. The results of my own experiments indicate that the timidity of the adult is greater than that of the immature animal when it is placed on a bridge 1 or 2 cm. wide at a distance of 20 cm. from the ground. Individuals three weeks old showed less hesitation about trying to creep along such a narrow pathway than did full-grown dancers three or four months old; and these, in turn, were not so timid apparently as an individual one year old. But the younger animals fell off more frequently than did the older ones.

[Illustration: FIGURE 4.—Zoth's photographs of dancers crossing bridges and climbing rods. Reproduced from Pfluger's Archiv, Bd. 86.]

[Illustration: FIGURE 5—Tracks of common mouse Reproduced from Alexander and Kreidl's figure in Pfluger's Archiv, Bd 82]

[Illustration: FIGURE 6—Tracks of dancing mouse Reproduced from Alexander and Kreidl's figure in Pfluger's Archiv Bd 82]

Additional support for these statements concerning equilibrational ability is furnished by the observations of Kishi (21 p. 482). He built a wooden bridge 60 cm. long, 1 cm. wide at one end, and 1/2 cm. at the other, and supported it at a height of 30 cm. above the ground by posts at the ends. On this bridge ten dancers were tested. Some attempted to move sidewise, others began to whirl and fell to the ground; only one of the ten succeeded in getting all the way across the bridge on the first trial. The second time he was tested this individual crossed the bridge and found the post; and the third time he crossed the bridge and climbed down the post directly. The others did not succeed in descending the post even after having crossed the bridge safely, but, instead, finally fell to the floor from awkwardness or exhaustion. On the basis of these and other similar observations, Kishi says that the dancer possesses a fair degree of ability to orient and balance itself.

Inasmuch as equilibration occurs similarly in darkness and in daylight, Zoth thinks that there is neither visual dizziness nor fear of heights. But it is doubtful whether he is right concerning fear. There is no doubt in my mind, in view of the way the mice behave when placed on an elevated surface, that they are timid; but this is due probably to the uncomfortable and unusual position rather than to perception of their distance from the ground. That they lack visual dizziness seems fairly well established.

When rotated in a cyclostat[1] the dancer, unlike the common mouse, does not exhibit symptoms of dizziness. The following vivid description of the behavior of both kinds of mice when rotated is given by Alexander and Kreidl (1 p. 548). I have not verified their observations.

[Footnote 1: An apparatus consisting of a glass cylinder with a mechanism for turning it steadily and at different speeds about its vertical axis.]

The common mouse at first runs with increasing rapidity, as the speed of rotation of the cyclostat cylinder is increased, in the direction opposite to that of the cylinder itself. This continues until the speed of rotation has increased to about 60 revolutions per minute. As the rotation becomes still more rapid the mouse begins to crawl along the floor, its body stretched out and clinging to the floor. At a speed of 250 revolutions per minute it lies flat on the floor with its limbs extended obliquely to the movement of rotation, and at times with its back bent against the axis of the cylinder; in this position it makes but few and feeble efforts to crawl forward. When the rotation is suddenly stopped, the animal pulls itself together, remains for some seconds with extended limbs lying on the floor, and then suddenly falls into convulsions and trembles violently. After several attacks of this kind, cramps appear and, despite its resistance, the animal is thrown about, even into the air at times, as if by an external force. This picture of the position assumed during rapid rotation, and of cramps after the cessation of rotation (the typical picture of rotation dizziness), is repeated with great uniformity in the case of the common mouse. Within fifteen minutes after being returned to its cage the animal recovers from the effects of its experience. This description of the symptoms of rotation dizziness in the common mouse applies equally well to the blinded and the seeing animal.

In sharp contrast with the behavior of the common mouse in the cyclostat is that of the dancer. As the cylinder begins to rotate the dancer runs about as usual in circles, zigzags, and figure-eights. As the speed becomes greater it naturally becomes increasingly difficult for the mouse to do this, but it shows neither discomfort nor fear, as does the common mouse. Finally the centrifugal force becomes so great that the animal is thrown against the wall of the cylinder, where it remains quietly without taking the oblique position. When the cyclostat is stopped suddenly, it resumes its dance movements as if nothing unusual had occurred. It exhibits no signs of dizziness, and apparently lacks the exhaustion which is manifest in the case of other kinds of mice after several repetitions of the experiment. The behavior of the blinded dancer is very similar.

If these statements are true, there is no reason to believe that the dancer is capable of turning or rotation dizziness. If it were, its daily life would be rendered very uncomfortable thereby, for its whirling would constantly bring about the condition of dizziness. Apparently, then, the dancer differs radically from most mammals in that it lacks visual and rotational dizziness. In the next chapter we shall have to seek for the structural causes for these facts.

The behavior of the blinded animal is so important in its bearings upon the facts of orientation and equilibration that it must be considered in connection with them. Cyon insists that the sense of vision is of great importance to the dancer in orienting and equilibrating itself. When the eyes are covered with cotton wads fastened by collodion, this writer states (9 p. 223) that the mice behave as do pigeons and frogs whose semicircular canals have been destroyed. They perform violent forced movements, turn somersaults forward and backward, run up inclines and fall over the edges, and roll over and over. In a word, they show precisely the kind of disturbances of behavior which are characteristic of animals whose semicircular canals are not functioning normally. Cyon, however, observed that in certain dancers these peculiarities of behavior did not appear when they were blinded, but that, instead, the animals gave no other indication of being inconvenienced by the lack of sight than do common white mice. This matter of individual differences we shall have to consider more fully later.

No other observer agrees with Cyon in his conclusions concerning vision, or, for that matter, in his statements concerning the behavior of the blind dancer. Alexander and Kreidl (1 p. 550) contrast in the following respects the behavior of the white mouse and that of the dancer when they are blinded. The white mouse runs less securely and avoids obstacles less certainly when deprived of vision. The dancer is much disturbed at first by the shock caused by the removal of its eyes, or in case they are covered, by the presence of the unusual obstruction. It soon recovers sufficiently to become active, but it staggers, swerves often from side to side, and frequently falls over. It moves clumsily and more slowly than usual. Later these early indications of blindness may wholly disappear, and only a slightly impaired ability to avoid obstacles remains.

It was noted by Kishi (21 p. 484), that the dancer when first blinded trembles violently, jumps about wildly, and rolls over repeatedly, as Cyon has stated; but Kishi believes that these disturbances of behavior are temporary effects of the strong stimulation of certain reflex centers in the nervous system. After having been blinded for only a few minutes the dancers observed by him became fairly normal in their behavior. They moved about somewhat more slowly than usually, especially when in a position which required accurately coordinated movements. He therefore fully agrees with Alexander and Kreidl in their conclusion that vision is not so important for the guidance of the movements of the dancer as Cyon believes.

In summing up the results of his investigation of this subject Zoth well says (31 p. 168), "the orientation of the positions of the body with respect to the horizontal and vertical planes seems to take place without the assistance of the sense of sight." And, as I have already stated, this excellent observer insists that the ability of the dancer to place its body in a particular position (orientation), and its ability to maintain its normal relations to its surroundings (equilibration) are excellent in darkness and in daylight, provided only the substratum be not too smooth for it to gain a foothold.

It must be admitted that the contradictions which exist in the several accounts of the behavior of the dancer are too numerous and too serious to be explained on the basis of careless observation. Only the assumption of striking individual differences among dancers or of the existence of two or more varieties of the animal suffices to account for the discrepancies. That there are individual or variety differences is rendered practically certain by the fact that Cyon himself worked with two groups of dancers whose peculiarities he has described in detail, both as to structure and behavior.

In the case of the first group, which consisted of three individuals, the snout was more rounded than in the four individuals of the second group, and there were present on the head three large tufts of bristly black hair which gave the mice a very comical appearance. The animals of the second group resembled more closely in appearance the common albino mouse. They possessed the same pointed snout and long body, and only the presence of black spots on the head and hips rendered them visibly different from the albino mouse.

In behavior the individuals of these two groups differed strikingly. Those of the first group danced frequently, violently, and in a variety of ways; they seldom climbed on a vertical surface and when forced to move on an incline they usually descended by sliding down backwards or sidewise instead of turning around and coming down head first; they gave no signs whatever of hearing sounds. Those of the second group, on the contrary, danced very moderately and in few ways; they climbed the vertical walls of their cage readily and willingly, and when descending from a height they usually turned around and came down head first; two of the four evidently heard certain sounds very well. No wonder that Cyon suggests the possibility of a different origin! It seems not improbable that the individuals of the second group were of mixed blood, possibly the result of crosses with common mice.

As I shall hope to make clear in a subsequent discussion of the dancer's peculiarities of behavior, in a chapter on individual differences, there is no sufficient reason for doubting the general truth of Cyon's description, although there is abundant evidence of his inaccuracy in details. If, for the present, we accept without further evidence the statement that there is more than one variety of dancer, we shall be able to account for many of the apparent inaccuracies of description which are to be found in the literature on the animal.

As a result of the examination of the facts which this chapter presents we have discovered at least six important peculiarities of behavior of the dancer which demand an explanation in terms of structure. These are: (1) the dance movements—whirling, circling, figure-eights, zigzags; (2) restlessness and the quick, jerky movements of the head; (3) lack of responsiveness to sounds; (4) more or less pronounced deficiency in orientational and equilibrational power; (5) lack of visual dizziness; (6) lack of rotational dizziness.

Naturally enough, biologists from the first appearance of the dancing mouse in Europe have been deeply interested in what we usually speak of as the causes of these peculiarities of behavior. As a result, the structure of those portions of the body which are supposed to have to do with the control of movement, with the phenomena of dizziness, and with the ability to respond to sounds, have been studied thoroughly. In the next chapter we shall examine such facts of structure as have been discovered and attempt to correlate them with the facts of behavior.

The Dancing Mouse A Study in Animal Behavior - Chapter II

FEEDING, BREEDING, AND DEVELOPMENT OF THE YOUNG

In this chapter I shall report, for the benefit of those who may wish to know how to take care of dancing mice, my experience in keeping and breeding the animals, and my observations concerning the development of the young. It is commonly stated that the dancer is extremely delicate, subject to diseases to an unusual degree and difficult to breed. I have not found this to be true. At first I failed to get them to breed, but this was due, as I discovered later, to the lack of proper food. For three years my mice have bred frequently and reared almost all of their young. During one year, after I had learned how to care for the animals, when the maximum number under observation at any time was fifty and the total number for the year about one hundred, I lost two by disease and one by an accident. I very much doubt whether I could have done better with any species of mouse. There can be no doubt, however, that the dancer is delicate and demands more careful attention than do most mice. In March, 1907, I lost almost all of my dancers from what appeared to be an intestinal trouble, but with this exception I have had remarkably good luck in breeding and rearing them.

My dancers usually were kept in the type of cage of which Figure 2 is a photograph.[1] Four of these double cages, 70 cm. long, 45 cm. wide, and 10 cm. deep in front, were supported by a frame as is shown in Figure 3. The fact that the covers of these cages cannot be left open is of practical importance. A similar type of cage, which I have used to some extent, consists of a wooden box 30 by 30 cm. by 15 cm. deep, without any bottom, and with a hinged cover made in part of 1 cm. mesh wire netting. Such a cage may be placed upon a piece of tin or board, or simply on a newspaper spread out on a table. The advantage of the loose bottom is that the box may be lifted off at any time, and the bottom thoroughly cleansed. I have had this type of cage constructed in blocks of four so that a single bottom and cover sufficed for the block. If the mice are being kept for show or for the observation of their movements, at least one side of the cages should be of wire netting, and, as Kishi suggests, such objects as a wheel, a tower, a tunnel, a bridge, and a turntable, if placed in the cage, will give the animals excellent opportunity to exhibit their capacity for varied forms of activity.

[Footnote 1: This cage was devised by Professors W.E. Castle and E.L.
Mark, and has been used in the Zoological Laboratories of Harvard
University for several years.]

[Illustration: FIGURE 2.—Double cage, with nest boxes and water dishes.]

The floors of the cages were covered with a thin layer of sawdust for the sake of cleanliness, and in one corner of each cage a nest box of some sort was placed. During the warm months I found it convenient and satisfactory to use berry boxes, such as appear in Figure 2, with a small entrance hole cut in one side; and during the cold months cigar boxes, with an entrance hole not more than 5 cm. in diameter at one end. In the nest box a quantity of tissue paper, torn into fragments, furnished material for a nest in which the adults could make themselves comfortable or the female care for her young. Cotton should never be used in the nest boxes, for the mice are likely to get it wound about their legs with serious results. Apparently they are quite unable to free themselves from such an incumbrance, and their spinning motion soon winds the threads so tightly that the circulation of the blood is stopped.

[Illustration: FIGURE 3.—Double cages in frame.]

The cages and nest boxes were emptied and thoroughly cleaned once a week with an emulsion made by heating together one part of kerosene and one part of water containing a little soap. This served to destroy whatever odor the cages had acquired and to prevent vermin from infesting the nests. In hot weather far greater cleanliness is necessary for the welfare of the mice than in cold weather. The animals attend faithfully to their own toilets, and usually keep themselves scrupulously clean.

For water and food dishes I have used heavy watch glasses[1] 5 cm. in diameter and 1/2 cm. deep. They are convenient because they are durable, easily cleaned, and not large enough for the young mice to drown in when they happen to spin into one which contains water. It is said that mice do not need water, but as the dancers seem very fond of a little, I have made it a rule to wash the watch glasses thoroughly and fill them with pure fresh water daily. The food, when moist, may be placed in the cages in the same kind of watch glass.

[Footnote 1: Minot watch glasses.]

There is no need of feeding the animals oftener than once a day, and as they eat mostly in the evening and during the night, it is desirable that the food should be placed in the cage late in the afternoon. For almost a year I kept a pair of dancers on "force"[1] and water. They seemed perfectly healthy and were active during the whole time, but they produced no young. If the animals are kept as pets, and breeding is not desired, a diet of "force," "egg-o-see,"[1] and crackers, with some bird-seed every few days, is likely to prove satisfactory. As with other animals, a variety of food is beneficial, but it appears to be quite unnecessary. Too much rich food should not be given, and the mice should be permitted to dictate their own diet by revealing their preferences. They eat surprisingly little for the amount of their activity. I have had excellent success in breeding the mice by feeding them a mixture of dry bread- crumbs, "force," and sweet, clean oats slightly moistened with milk. The food should never be made soppy. A little milk added thus to the food every other day greatly increases fertility. About once a week a small quantity of some green food, lettuce for example, should be given. It is well, I have found, to vary the diet by replacing the bread and "force" at intervals with crackers and seeds. Usually I give the food dry every other day, except in the case of mice which are nursing litters. One person to whom I suggested that lettuce was good for the dancers lost four, apparently because of too much of what the mice seemed to consider a good thing. This suggests that it should be used sparingly.

[Footnote 1: A cereal food.]

Success in keeping and breeding dancing mice depends upon three things: cleanliness, warmth, and food supply. The temperature should be fairly constant, between 60° and 70° Fahr. They cannot stand exposure to cold or lack of food. If one obtains good healthy, fertile individuals, keeps them in perfectly clean cages with soft nesting materials, maintains a temperature of not far above or below 65°, and regularly supplies them with pure water and food which they like, there is not likely to be trouble either in keeping or breeding these delicate little creatures. Several persons who have reported to me difficulty in rearing the young or in keeping the adults for long periods have been unable to maintain a sufficiently high or constant temperature, or have given them food which caused intestinal trouble.

The males are likely to fight if kept together, and they may even kill one another. A male may be kept with one or more females, or several females may be kept together, for the females rarely, in my experience, fight, and the males seldom harm the females. Unless the male is removed from the cage in which the female is kept before the young are born, he is likely to kill the newborn animals. When a female is seen to be building a nest in preparation for a litter, it is best to place her in a cage by herself so that she may not be disturbed.

The sex of individuals may be determined easily in most cases, at the age of 10 to 12 days, by the appearance of teats in the case of females.

The period of gestation is from 18 to 21 days. The maximum number born by my dancers in any single litter was 9, the minimum number 3. In 25 litters of which I have accurate records, 135 individuals were born, an average of 5.4. The average number of males per litter was precisely the same, 2.7, as the number of females.

On the birth of a litter it is well to see that the female has made a nest from which the young are not likely to escape, for at times, if the nest is carelessly made, they get out of it or under some of the pieces of paper which are used in its construction, and perish. Several times I have observed nests so poorly built that almost all of the young perished because they got too far away to find their way back to the mother. It is surprising that the female should not take more pains to keep her young safe by picking them up in her mouth, as does the common mouse, and carrying them to a place where they can obtain warmth and nourishment. This I have never seen a dancing mouse do. For the first day or two after the birth of a litter the female usually remains in the nest box almost constantly and eats little. About the second day she begins to eat ravenously, and for the next three or four weeks she consumes at least twice as much food as ordinarily. Alexander and Kreidl (3 p. 567) state that the female does not dance during the first two weeks after the birth of a litter, but my experience contradicts their statement. There is a decreased amount of activity during this period, and usually the whirling movement appears but rarely; but in some cases I have seen vigorous and long-continued dancing within a few hours after the birth of a litter. There is a wide range of variability in this matter, and the only safe statement, in the light of my observations, is that the mother dances less than usual for a few days after a litter is born to her.

The development of the young, as I have observed it in the cases of twenty litters, for ten of which (Table I) systematic daily records were kept, may be sketched as follows. At birth the mice have a rosy pink skin which is devoid of hair and perfectly smooth; they are blind, deaf, and irresponsive to stimulation of the vibrissae on the nose. During the first week of post-natal life the members of a litter remain closely huddled together in the nest, and no dance movements are exhibited. The mother stays with them most of the time. On the fourth or fifth day colorless hairs are visible, and by the end of the week the body is covered with a coat which rapidly assumes the characteristic black and white markings of the race. For the first few days the hind legs are too weak to support the body weight, and whatever movements appear are the result of the use of the fore legs. As soon as the young mice are able to stand, circling movements are exhibited, and by the end of the second week they are pronounced. Somewhere about the tenth day the appearance of the teats in the case of the females serves to distinguish the sexes plainly. Between the tenth and fifteenth days excitability, as indicated by restless jerky movements in the presence of a disturbing condition, increases markedly; the auditory meatus opens, and, in the case of some individuals, there are signs of hearing. On or after the fifteenth day the eyes open and the efforts to escape from the nest box rapidly become more vigorous. About this time the mother resumes her dancing with customary vigor, and the young, when they have opportunity, begin to eat of the food which is given to her. They now dance essentially as do the adults. From the end of the third week growth continues without noteworthy external changes until sexual maturity is attained, between the fourth and the sixth week. For several weeks after they are sexually mature the mice continue to increase in size.
TABLE I
DEVELOPMENT OF THE YOUNG
NUMBER JERKY REACT IN HAIR TEATS MOVE- EARS TO EYES PARENTS LITTER VISIBLE VISIBLE MENTS OPEN SOUND OPEN M F APPEAR

152+151 5 0 4th day — 13th day 14th day 14th day 16th day
152+151 1 3 4th day 9th day 10th day 12th day 13th day 15th day
410+415 4 1 5th day 11th day 14th day 15th day 15th day 17th day
410+415 2 4 5th day 10th day 13th day 14th day 14th day 16th day
420+425 0 2 4th day 10th day 12th day 14th day 14th day 16th day
210+215 4 1 — — 17th day 13th day 17th day 15th day
210+215 3 3 5th day 11th day 11th day 14th day No 16th day
212+211 1 3 4th day 10th day 15th day 14th day No 15th day
220+225 2 4 4th day 10th day 16th day 14th day No 15th day
220+225 3 3 4th day 10th day 17th day 13th day No 15th day

A course of development very similar to that just described was observed by Alexander and Kreidl (3 p. 565) in three litters of dancing mice which contained 3, 5, and 7 individuals respectively. These authors, in comparing the development of the dancer with that of the common mouse, say that at birth the young in both cases are about 24 mm. in length. The young common mouse grows much more rapidly than the dancer, and by the ninth day its length is about 43 mm. as compared with 31 mm. in the case of the dancer. According to Zoth (31 p. 148) the adult dancer has a body length of from 7 to 7.5 cm., a length from tip of nose to tip of tail of from 12 to 13 cm., and a weight of about 18 grams. The movement of the dancer from the first tends to take the form of circles toward the middle of the nest; that of the common mouse has no definite tendency as to direction. When the common mouse does move in circles, it goes first in one direction, then in the other, and not for any considerable period in one direction as does the true dancer. Neither the young dancer nor the common mouse is able to equilibrate itself well for the first few days after birth, but the latter can follow a narrow path with far greater accuracy and steadiness than the former. The uncertain and irregular movements of the common mouse are due to muscular weakness and to blindness, but the bizarre movements of the young dancer seem to demand some additional facts as an explanation.

A brief account of the development of the dancer given by Zoth (31 p. 149) adds nothing of importance to the description given by Alexander and Kreidl. As my own observations disagree with their accounts in certain respects, I shall now give, in the form of a diary, a description of the important changes observed from day to day in a normal litter. The litter which I have selected as typical of the course of development in the dancer grew rapidly under favorable conditions. I have observed many litters which passed through the various stages of development mentioned in this description anywhere from a day to a week later. This was usually due to some such obviously unfavorable condition as too little food or slight digestive or bowel troubles. According to the nature of the conditions of growth the eyes of the dancer open anywhere from the fourteenth to the twentieth day. This statement may serve to indicate the degree of variability as to the time at which a given stage of development is reached by different litters.

On July 14, 1906, No. 151 (female) and No. 152 (male) were mated, and on August 3 a litter of six was born to them. The course of the development of this litter during the first three weeks was as follows:—

First day The skin is pink and hairless, several vibrissae are visible on the nose and lips, but there is no definite response when they are touched. The mice are both blind and deaf, but they are able to squeak vigorously. The mother was not seen to dance or eat during the day.

Second day. There is a very noticeable increase in size. The vibrissae are longer, but touching them still fails to cause a reaction. No hairs are visible on the body. The mother danced rapidly for periods of a minute several times while the record was being made. She ate very little to-day.

Third day. Scales began to appear on the skin to-day. The animals are rapidly increasing in strength; they can now crawl about the nest easily, but they are too weak to stand, and constantly roll over upon their sides or backs when they are placed on a smooth surface. Because of their inability to progress it is impossible to determine with certainty whether they have a tendency to move in circles. The mother was seen out of the nest dancing once to-day. She now eats ravenously.

Fourth day. One of the six young mice was found under a corner of the nest this morning dead, and the others were scattered about the nest box. I gathered them together into a nest which I made out of bits of tissue paper, and the mother immediately began to suckle them. They are very sensitive to currents of air, but they do not respond to light or sound and seldom to contact with the vibrissae.

Fifth day. When placed on a smooth surface, they tend to move in circles, frequently rolling over. When placed on their sides or backs, they immediately try to right themselves. They do not walk, for their legs are still too weak to support the weight of the body; instead they drag themselves about by the use of the fore legs. Fine colorless hairs are visible over the entire body surface. When the vibrissae are touched, the head is moved noticeably. The mother dances a great deal and eats about twice as much as she did before the birth of the litter.

Sixth day. Certain regions of the skin, which were slightly darker than the remainder on the fourth and fifth days, are now almost black. It is evident that they are the regions in which the black hair is to appear. The movement in circles is much more definite today, although most of the individuals are still too weak to stand on their feet steadily for more than a few seconds at a time. Most of their time, when they are first taken from the nest, is spent in trying to maintain or regain an upright position. The hair is now easily visible, and the skin begins to have a white appearance as a result.

Seventh day. Although they are strong enough to move about the nest readily, none of the young has attempted to leave the nest. They huddle together in the middle of it for warmth. The epidermal scales, which have increased in number since the third day, are dropping off rapidly. Contact with the vibrissae or with the surface of the body, frequently calls forth a motor reaction but neither light nor sound produces any visible change in behavior. The black and white regions of the skin are sufficiently definite now to enable one to distinguish the various individuals by their markings. The mother was seen to dance repeatedly today, and she ate all the food that was given to her.

Eighth day. A fold is plainly visible where later the eyelids will separate. The black pigment in the skin has increased markedly.

Ninth day. The eyelids are taking form rapidly, but they I have not separated. The body is covered with a thick coat of hair which is either pure white or black. Standing on the four legs is still a difficult task.

Tenth day. To-day teats are plainly visible in the case of four of the five individuals of the litter. Up to this time I had thought, from structural indications, that there were three males and two females; it is now evident that there are four females and one male. The external ear, the pinna, is well formed, and has begun to stand out from the head, but no opening to the inner portion of the ear is present. The eyelids appear to be almost fully formed.

Eleventh day. There are no very noticeable changes in appearance except in size, which continues to increase rapidly. They are able to regain their normal upright position almost immediately when they happen to roll over. The mother dances as usual.

Twelfth day. It appears to-day as if the eyes were about to open. The ears are still closed, and there is no evidence of a sense of hearing. They squeaked considerably when in the nest, but not at all when I took them out to note their development. The mother stays outside of the nest box much of the time now, probably to prevent the young ones from sucking continuously.

Thirteenth day. One of the little mice came out of the nest box while I was watching the litter this morning, and was able to find his way back directly despite the lack of sight. The mice are still dependent upon the mother for nourishment. I have not seen any of them attempt to eat the food which is given to the mother. They are extremely neat and clean. I watched one of them wash himself this morning. Each foot was carefully licked with the tongue. There seems to be special care taken to keep the toes perfectly clean.

Fourteenth day. An opening into the ear is visible to-day. When tested with the Galton whistle, all five responded with quick, jerky movements of the head and legs. They evidently hear certain tones. During the past two days the ears have changed rapidly. In one of the females, which seems to be a little in advance of the others in development, certain peculiarities of behavior appeared to-day. She jumped and squeaked sharply when touched and sprang out of my hand when I attempted to take her up. This is in marked contrast with her behavior previously.

Fifteenth day. The eyes are partly opened. All of the members of the litter came out of the nest box this morning and ran around the cage, dancing frequently and trying to eat with the mother. Three out of the five gave auditory reactions on first being stimulated; none of them responded to repetitions of the stimulus. All appeared to be less sensitive to sounds than yesterday. The quick, nervous, jerky movements are very noticeable.

Sixteenth day. The eyes of all five are fully opened. They dance vigorously and are outside the nest much of the time.

Seventeenth day. No reactions to sound could be detected to-day. The sense of sight gives evidence of being well developed. The nervous jumping movements persist.

Eighteenth day. The young mice continue to suck, although they eat of the food which is given to the mother. They are now able to take care of themselves.

Nineteenth day. There are no noteworthy changes except increase in size and strength.

Twentieth day. No auditory reactions were obtained today, but other forms of stimulation brought about unmistakable responses.

Twenty-first day. They are now about half grown and there is no other change of special interest to be recorded. Growth continues for several weeks. The statement made by Alexander and Kreidl to the effect that the dancer is almost full grown by the thirty-first day of life is false. At that age they may be sexually mature, but usually they are far from full grown.

The Dancing Mouse A Study in Animal Behavior - Chapter I

CHARACTERISTICS, ORIGIN, AND HISTORY

The variety of mouse which is known as the Japanese dancing or waltzing mouse has been of special interest to biologists and to lovers of pets because of its curious movements. Haacke in Brehm's "Life of Animals" (7 p. 337)[1] writes as follows concerning certain mice which were brought to Europe from China and Japan: "From time to time a Hamburg dealer in animals sends me two breeds of common mice, which he calls Chinese climbing mice (Chinesische Klettermäuse) and Japanese dancing mice (Japanische Tanzmäuse). It is true that the first are distinguished only by their different colors, for their climbing accomplishments are not greater than those of other mice. The color, however, is subject to many variations. Besides individuals of uniform gray, light yellow, and white color, I have had specimens mottled with gray and white, and blue and white. Tricolored mice seem to be very rare. It is a known fact that we also have white, black, and yellow mice and occasionally pied ones, and the Chinese have profited by these variations of the common mouse also, to satisfy their fancy in breeding animals. The Japanese, however, who are no less enthusiastic on this point, know how to transform the common mouse into a really admirable animal. The Japanese dancing mice, which perfectly justify their appellation, also occur in all the described colors. But what distinguishes them most is their innate habit of running around, describing greater or smaller circles or more frequently whirling around on the same spot with incredible rapidity. Sometimes two or, more rarely, three mice join in such a dance, which usually begins at dusk and is at intervals resumed during the night, but it is usually executed by a single individual."

[Footnote 1: The reference numbers, of which 7 is an example, refer to the numbers in the bibliographic list which precedes this chapter.]

As a rule the dancing mouse is considerably smaller than the common mouse, and observers agree that there are also certain characteristic peculiarities in the shape of the head. One of the earliest accounts of the animal which I have found, that of Landois (22 p. 62), states, however, that the peculiarities of external form are not remarkable. Landois further remarks, with reason, that the name dancing mouse is ill chosen, since the human dance movement is rather a rhythmic hopping motion than regular movement in a circle. As he suggests, they might more appropriately be called "circus course mice" (22 p. 63).

Since 1903 I have had under observation constantly from two to one hundred dancing mice. The original pair was presented to the Harvard Psychological Laboratory by Doctor A.G. Cleghorn of Cambridge. I have obtained specimens, all strikingly alike in markings, size, and general behavior, from animal dealers in Washington, Philadelphia, and Boston. Almost all of the dancers which I have had, and they now number about four hundred, were white with patches, streaks, or spots of black. The black markings occurred most frequently on the neck, ears, face, thighs, hind legs, about the root of the tail, and occasionally on the tail itself. In only one instance were the ears white, and that in the case of one of the offspring of a male which was distinguished from most of his fellows by the possession of one white ear. I have had a few individuals whose markings were white and gray instead of white and black.

The method by which I was able to keep an accurate record of each of my dancers for purposes of identification and reference is illustrated in Figure 1. As this method has proved very convenient and satisfactory, I may briefly describe it. With a rubber stamp[1] a rough outline of a mouse, like that of Figure 1 A, was made in my record book. On this outline I then indicated the black markings of the individual to be described. Beside this drawing of the animal I recorded its number, sex,[2] date of birth, parentage, and history. B, C, and D of Figure 1 represent typical color patterns. D indicates the markings of an individual whose ears were almost entirely white. The pattern varies so much from individual to individual that I have had no trouble whatever in identifying my mice by means of such records as these.

[Footnote 1: For the use of the plate from which this stamp was made, I am indebted to Professor W.E. Castle, who in turn makes acknowledgment to Doctor G.M. Allen for the original drawing.]

[Footnote 2: I have found it convenient to use the even numbers for the males and the odd numbers for the females. Throughout this book this usage is followed. Wherever the sex of an individual is not specially given, the reader therefore may infer that it is a male if the number is even; a female if the number is odd.]

All of my dancers had black eyes and were smaller as well as weaker than the albino mouse and the gray house mouse. The weakness indicated by their inability to hold up their own weight or to cling to an object curiously enough does not manifest itself in their dancing; in this they are indefatigable. Frequently they run in circles or whirl about with astonishing rapidity for several minutes at a time. Zoth (31 p. 173), who measured the strength of the dancer in comparison with that of the common mouse, found that it can hold up only about 2.8 times its own weight, whereas the common white mouse can hold up 4.4 times its weight. No other accurate measurements of the strength, endurance, or hardiness of the dancer are available. They are usually supposed to be weak and delicate, but my own observations cause me to regard them as exceptionally strong in certain respects and weak in others.

[Illustration: FIGURE I.—Typical markings of dancers. A, blank outline of mouse for record. B, markings of No. 2 [symbol for male], born September 7, 1905, of unknown parents, died March 30,1907. C, markings of No 43 [symbol for female], born November 10, 1906, of 212 and 211. D, markings of No. 151 [symbol for female], born February 28, 1906, of 1000 and 5, died February 26, 1907.]

What the Japanese have to say about the dancing mouse is of special importance because Japan is rather commonly supposed to be its home. For this reason, as well as because of the peculiar interest of the facts mentioned, I quote at length from Doctor Kishi (21 p. 457). "The dancing mouse has received in Europe this name which it does not bear in its own home, because of the fact that the circular movements which it makes are similar to the European (human) dance. Sometimes it is also called the Japanese or Chinese mouse; originally, however, China must have been its home, since in Japan it is mostly called 'Nankin nesumi,' the mouse from Nankin. When this animal came from China to Japan I shall inquire at a later opportunity. There were originally in Japan two different species of mouse, the gray and the white; therefore in order to distinguish our dancing mouse from these it was necessary to use the name of its native city.

"In Japan, as in Europe, the animal lives as a house animal in small cages, but the interest which is taken in it there is shown in quite another way than in Europe, where the whirling movements, to which the name dancing mouse is due, are of chief interest. For this reason in Europe it is given as much room as possible in its cage that it may dance conveniently. In Japan also the circular movements have been known for a long time, but this has had no influence upon our interest in the animal, for the human fashion of dancing with us is quite different from that in Europe. What has lent interest to the creature for us are its prettiness, its cleverness in tricks, and its activity. It is liked, therefore, as an amusement for children. For this purpose it is kept in a small cage, usually fifteen centimeters square, sometimes in a somewhat broader wooden box one of whose walls is of wire netting. In this box are built usually a tower, a tunnel, a bridge, and a wheel. The wheel is rather broad, being made in the form of a drum and pierced with holes on one side through which the animal can slip in and out. Running around on the inside, the mouse moves the wheel often for hours at a time, especially in the evening. Moreover, there are found in the box other arrangements of different kinds which may be set in motion by the turning of the wheel. No space remains in the box in which the animal may move about freely, and therefore one does not easily or often have an opportunity to observe that the animal makes circular movements, whether voluntarily or involuntarily. This is the reason that in its home this interesting little animal has never been studied by any one in this respect."

It is odd indeed that the remarkable capacity of the dancer for the execution of quick, graceful, dextrous, bizarre, and oft-repeated movements has not been utilized in America as it has in Japan. The mice are inexhaustible sources of amusement as well as invaluable material for studies in animal behavior and intelligence.

Concerning the origin and history of this curious variety of mouse little is definitely known. I have found no mention of the animal in scientific literature previous to 1890. The fact that it is called the Chinese dancing mouse, the Japanese dancing mouse, and the Japanese waltzing mouse is indicative of the existing uncertainty concerning the origin of the race.

Thinking that Japanese literature might furnish more information bearing on the question of racial history than was available from European sources, I wrote to Professor Mitsukuri of the University of Tokyo, asking him whether any reliable records of the dancer existed in Japan. He replied as follows: "I have tried to find what is known in Japan about the history of the Japanese waltzing mice, but I am sorry to say that the results are wholly negative. I cannot find any account of the origin of this freak, either authentic or fictitious, and, strange as it may seem to you, no study of the mice in a modern sense has been made, so you may consider the literature on the mouse in the Japanese language as absolutely nil." In explanation of this somewhat surprising ignorance of the origin of the race in what is commonly supposed to be its native land, Professor Mitsukuri adds: "The breeders of the mice have mostly been ignorant men to whom writing is anything but easy."

In response to similar inquiries, I received the following letter, confirmatory of Professor Mitsukuri's statements, from Doctor S. Hatai of Wistar Institute, Philadelphia: "If I remember rightly the so-called Japanese dancing mouse is usually called by us Nankin-nedzumi. Nankin means anything which has been imported from China, and nedzumi means rat-like animal, or in this case mouse, or Chinese mouse. I referred to one of the standard Japanese dictionaries and found the following statement: 'The Nankin-nedzumi is one of the varieties of Mus spiciosus (Hatszuka-nedzumi), and is variously colored. It was imported from China. These mice are kept in cages for the amusement of children, who watch their play.' Mus spiciosus, if I remember correctly, is very much like Mus musculus in color, size, and several other characteristics, if not the same altogether."

In Swinhoe's list of the mammals of China, which appeared in the Proceedings of the Zoological Society of London for 1870, Mus musculus L. is mentioned as occurring in houses in South China and in Formosa. It is further stated that black and white varieties which are brought from the Straits are often kept by the Chinese (p. 637).

The statements of Kishi, Mitsukuri, and Hatai which have been quoted, taken in connection with the opinions expressed by various European scientists who have studied the dancer, make it seem highly probable that the race appeared first in China, and was thence introduced into Japan, from which country it has been brought to Europe and America. Accepting for the present this conclusion with reference to the place of origin of the dancer, we may now inquire, how and when did this curious freak, as Professor Mitsukuri has called it, come into existence? Concerning these matters there is wide divergence of opinion.

Haacke (6 p. 514), as quoted in Brehm's "Tierleben," says that an animal dealer with whom he discussed the question of the possible origin of the dancer maintained that it came from Peru, where it nests in the full cotton capsules, arranging the cotton fibers in the form of a nest by running about among them in small circles. Hence the name cotton mouse is sometimes applied to it. Haacke himself believes, however, that the race originated either in China or Japan as the result of systematic selectional breeding. Of this he has no certainty, for he states that he failed to find any literature on the "beautiful mice of China and Japan." Whether Haacke's description of the dancing mouse was published elsewhere previous to its appearance in Brehm's "Tierleben" I am unable to state; I have found nothing written on the subject by him before 1890. Zoth (31 p. 176) also thinks that the race was developed by systematic breeding, or in other words, that it is a product of the skill of the Asiatic animal breeders.

Another account of the origin of the race is that accepted by Kishi (21 p. 481) and some other Japanese biologists. It is their belief that the forms of movement acquired by the individual as the result of confinement in narrow cages are inherited. Thus centuries of subjection to the conditions which Kishi has described (p. 6) finally resulted in a race of mice which breed true to the dance movement. It is only fair to add, although Kishi does not emphasize the fact, that in all probability those individuals in which the dancing tendency was most pronounced would naturally be selected by the breeders who kept these animals as pets, and thus it would come about that selectional breeding would supplement the inheritance of an acquired character. Few indeed will be willing to accept this explanation of the origin of the dancer so long as the inheritance of acquired characters remains, as at present, unproved.

Still another mode of origin of the mice is suggested by the following facts. In 1893 Saint Loup (28 p. 85) advanced the opinion that dancing individuals appear from time to time among races of common mice. The peculiarity of movement may be due, he thinks, to an accidental nervous defect which possibly might be transmissible to the offspring of the exceptional individual. Saint Loup for several months had under observation a litter of common mice whose quick, jerky, nervous movements of the head, continuous activity, and rapid whirling closely resembled the characteristic movements of the true dancers of China. He states that these mice ran around in circles of from 1 to 20 cm. in diameter. They turned in either direction, but more frequently to the left, that is, anticlockwise. At intervals they ran in figure-eights ([Symbol: figure eight]) as do the true dancers. According to Saint Loup these exceptional individuals were healthy, active, tame, and not markedly different in general intelligence from the ordinary mouse. One of these mice produced a litter of seven young, in which, however, none of the peculiarities of behavior of the parents appeared.

In view of this proof of the occurrence of dancing individuals among common mice, Saint Loup believes that the race of dancers has resulted from the inheritance and accentuation of an "accidental" deviation from the usual mode of behavior. It is scarcely necessary to say that this opinion would be of far greater weight had he observed, instead of postulating, the inheritance of the peculiarities of movement which he has described. It might be objected, to the first of his so-called facts, that the litter resulted from the mating of mice which possessed dancer blood. Until the occurrence of dancers among varieties of mice which are known to be unmixed with true dancers is established, and further, until the inheritance of this peculiar deviation from the normal is proved, Saint Loup's account of the origin of the dancing mouse race must be regarded as an hypothesis.

The occurrence of dancing individuals among common mice has been recorded by several other observers. Kammerer (20 p. 389) reports that he found a litter of young wood mice (Mus sylvaticus L.) which behaved much as do the spotted dancers of China. He also observed, among a lot of true dancers, a gray individual which, instead of spinning around after the manner of the race, turned somersaults at frequent intervals. It is Kammerer's opinion, as a result of these observations, that the black and white dancers of China and Japan have been produced by selectional breeding on the basis of this occasional tendency to move in circles. Among albino mice Rawitz (25 p. 238) has found individuals which whirled about rapidly in small circles. He states, however, that they lacked the restlessness of the Chinese dancers. Some shrews (Sorex vulgaris L.) which exhibited whirling movements and in certain other respects resembled the dancing mouse were studied for a time by Professor Häcker of Freiburg in Baden, according to a report by von Guaita (17 p. 317, footnote). Doctor G. M. Allen of Cambridge has reported to me that he noticed among a large number of mice kept by him for the investigation of problems of heredity[1] individuals which ran in circles; and Miss Abbie Lathrop of Granby, Massachusetts, who has raised thousands of mice for the market, has written me of the appearance of an individual, in a race which she feels confident possessed no dancer blood, which whirled and ran about in small circles much as do the true dancers.

[Footnote 1: Allen, G.M. "The Heredity of Coat Color in Mice." Proc. Amer.
Academy, Vol. 40, 59-163, 1904.]

Although it is possible that some of these cases of the unexpected appearance of individuals with certain of the dancer's peculiarities of behavior may have been due to the presence of dancer blood in the parents, it is not at all probable that this is true of all of them. We may, therefore, accept the statement that dancing individuals now and then appear in various races of mice. They are usually spoken of as freaks, and, because of their inability to thrive under the conditions of life of the race in which they happen to appear, they soon perish.

Another and a strikingly different notion of the origin of the race of dancers from those already mentioned is that of Cyon (11 p. 443) who argues that it is not a natural variety of mouse, as one might at first suppose it to be, but instead a pathological variation. The pathological nature of the animals is indicated, he points out, by the exceptionally high degree of variability of certain portions of the body. According to this view the dancing is due to certain pathological structural conditions which are inherited. Cyon's belief raises the interesting question, are the mice normal or abnormal, healthy or pathological? That the question cannot be answered with certainty off-hand will be apparent after we have considered the facts of structure and function which this volume presents.

Everything organic sooner or later is accounted for, in some one's mind, by the action of natural selection. The dancing mouse is no exception, for Landois (22 p. 62) thinks that it is the product of natural selection and heredity, favored, possibly, by selectional breeding in China. He further maintains that the Chinese dancer is a variety of Mus musculus L. in which certain peculiarities of behavior appear because of bilateral defects in the brain. This author is not alone in his belief that the brain of the dancer is defective, but so far as I have been able to discover he is the only scientist who has had the temerity to appeal to natural selection as an explanation of the origin of the race.

Milne-Edwards, as quoted by Schlumberger (29 p. 63), is of the opinion that the Chinese dancer is not a natural wild mouse race, but instead the product of rigid artificial selection. And in connection with this statement Schlumberger describes a discovery of his own which seems to have some bearing upon the problem of origin. In an old Japanese wood carving which came into his possession he found a group of dancing mice. The artist had represented in minute detail the characteristics of the members of the group, which consisted of the parents and eight young. The father and mother as well as four of the little mice are represented as white spotted with black. Of the four remaining young mice, two are entirely black and two entirely white. The two pure white individuals have pink eyes, as has also the mother. The eyes of all the others are black. From these facts Schlumberger infers that the dancer has resulted from the crossing of a race of black mice with a race of albinos; the two original types appear among the offspring in the carving.

Experimental studies of the inheritance of the tendency to dance are of interest in their bearing upon the question of origin. Such studies have been made by Haacke (19), von Guaita (17, 18), and Darbishire (13, 14, 15, 16), and the important results of their investigations have been well summarized by Bateson (5).

By crossing dancing mice with common white mice both Haacke and von Guaita obtained gray or black mice which are very similar to the wild house mouse in general appearance and behavior. The characteristic movements of the dancers do not appear. As the result of a long series of breeding experiments, Darbishire (16 pp. 26, 27) says: "When the race of waltzing mice is crossed with albino mice which do not waltz, the waltzing habit disappears in the resulting young, so that waltzing is completely recessive in Mendel's sense; the eye-color of the hybrids is always dark; the coat-color is variable, generally a mixture of wild-gray and white, the character of the coat being distinctly correlated with characters transmitted both by the albino and by the colored parent." When hybrids produced by the cross described by Darbishire are paired, they produce dancers in the proportion of about one to five.

Bateson (5 p. 93, footnote), in discussing the results obtained by Haacke, von Guaita, and Darbishire, writes: "As regards the waltzing character, von Guaita's experiments agree with Darbishire's in showing that it was always recessive to the normal. No individual in F1 [thus the first hybrid generation is designated] or in families produced by crossing F1 with the pure normal, waltzed. In Darbishire's experiments F1 x F1 [first hybrids mated] gave 8 waltzers in 37 offspring, indicating 1 in 4 as the probable average. From von Guaita's matings in the form DR x DR the totals of families were 117 normal and 21 waltzers…. There is therefore a large excess of normals over the expected 3 to 1. This is possibly due to the delicacy of the waltzers, which are certainly much more difficult to rear than normals are. The small number in von Guaita's litters makes it very likely that many were lost before such a character as this could be determined."

Bateson does not hazard a guess at the origin of the dancer, but merely remarks (5 p. 86) that the exact physiological basis of the dancing character is uncertain and the origin of this curious variation in behavior still more obscure. "Mouse fanciers have assured me," he continues, "that something like it may appear in strains inbred from the normal type, though I cannot find an indubitable case. Such an occurrence may be nothing but the appearance of a rare recessive form. Certainly it is not a necessary consequence of inbreeding, witness von Guaita's long series of inbred albinos." (von Guaita (17 p. 319) inbred for twenty-eight generations.)

From the foregoing survey of the available sources of information concerning the origin and history of the race of dancing mice the following important facts appear. There are four theories of the origin of the race: (1) origin by selectional breeding (Haacke, Zoth, Milne- Edwards); (2) origin through the inheritance of an acquired character (Kishi); (3) origin by mutation, inheritance, and selectional breeding (Saint Loup, Kammerer, Cyon); (4) origin by natural selection, and inheritance, favored by selectional breeding (Landois). Everything indicates that the race originated in China. It is fairly certain that individuals with a tendency to move in circles appear at rare intervals in races of common mice. It seems highly probable, in view of these facts, that the Chinese took advantage of a deviation from the usual form of behavior to develop by means of careful and patient selectional breeding a race of mice which is remarkable for its dancing. Even if it should be proved that the mutation as it appears among common mice is not inherited, the view that slight deviations were taken advantage of by the breeders would still be tenable. The dancing tendency is such in nature as to unfit an individual for the usual conditions of mouse existence, hence, in all probability human care alone could have produced and preserved the race of dancers.

In answer to the question, how and when did the race of dancers originate, it may be said that historical research indicates that a structural variation or mutation which occasionally appears in Mus musculus, and causes those peculiarities of movement which are known as dancing, has been preserved and accentuated through selectional breeding by the Chinese and Japanese, until finally a distinct race of mice which breeds true to the dance character has been established. The age of the race is not definitely known, but it is supposed to have existed for several centuries.

Biology - A lecture delivered at Columbia University in the series on Science Philosophy and Art November 20, 1907

I must at the outset remark that among the many sciences that are occupied with the study of the living world there is no one that may properly lay exclusive claim to the name of Biology. The word does not, in fact, denote any particular science but is a generic term applied to a large group of biological sciences all of which alike are concerned with the phenomena of life. To present in a single address, even in rudimentary outline, the specific results of these sciences is obviously an impossible task, and one that I have no intention of attempting. I shall offer no more than a kind of preface or introduction to those who will speak after me on the biological sciences of physiology, botany and zoology; and I shall confine it to what seem to me the most essential and characteristic of the general problems towards which all lines of biological inquiry must sooner or later converge.
It is the general aim of the biological sciences to learn something of the order of nature in the living world. Perhaps it is not amiss to remark that the biologist may not hope to solve the ultimate problems of life any more than the chemist and physicist may hope to penetrate the final mysteries of existence in the non-living world. What he can do is to observe, compare and experiment with phenomena, to resolve more complex phenomena into simpler components, and to this extent, as he says, to "explain" them; but he knows in advance that his explanations will [6]never be in the full sense of the word final or complete. Investigation can do no more than push forward the limits of knowledge.
The task of the biologist is a double one. His more immediate effort is to inquire into the nature of the existing organism, to ascertain in what measure the complex phenomena of life as they now appear are capable of resolution into simpler factors or components, and to determine as far as he can what is the relation of these factors to other natural phenomena. It is often practically convenient to consider the organism as presenting two different aspects—a structural or morphological one, and a functional or physiological—and biologists often call themselves accordingly morphologists or physiologists. Morphological investigation has in the past largely followed the method of observation and comparison, physiological investigation that of experiment; but it is one of the best signs of progress that in recent years the fact has come clearly into view that morphology and physiology are really inseparable, and in consequence the distinctions between them, in respect both to subject matter and to method, have largely disappeared in a greater community of aim. Morphology and physiology alike were profoundly transformed by the introduction into biological studies of the genetic or historical point of view by Darwin, who did more than any other to establish the fact, suspected by many earlier naturalists, that existing vital phenomena are the outcome of a definite process of evolution; and it was he who first fully brought home to us how defective and one-sided is our view of the organism so long as we do not consider it as a product of the past. It is the second and perhaps greater task of the biologist to study the organism from the historical point of view, considering it as the product of a continuous process of evolution that has been in operation since life began. [7]In its widest scope this genetic inquiry involves not only the evolution of higher forms from lower ones, but also the still larger question of the primordial relation of living things to the non-living world. Here is involved the possibility so strikingly expressed many years ago by Tyndall in that eloquent passage in the Belfast address, where he declared himself driven by an intellectual necessity to cross the boundary line of the experimental evidence and to discern in non-living matter, as he said, the promise and potency of every form and quality of terrestrial life. This intellectual necessity was created by a conviction of the continuity and consistency of natural phenomena, which is almost inseparable from the scientific attitude towards nature. But Tyndall's words stood after all for a confession of faith, not for a statement of fact; and they soared far above the terra firma of the actual evidence. At the present day we too may find ourselves logically driven to the view that living things first arose as a product of non-living matter. We must fully recognize the extraordinary progress that has been made by the chemist in the artificial synthesis of compounds formerly known only as the direct products of living protoplasm. But it must also be admitted that we are still wholly without evidence of the origin of any living thing, at any period of the earth's history, save from some other living thing; and after more than two centuries Redi's aphorism omne vivum e vivo retains to-day its full force. It is my impression therefore that the time has not yet come when hypotheses regarding a different origin of life can be considered as practically useful.
If I have the temerity to ask your attention to the fundamental problem towards which all lines of biological inquiry sooner or later lead us it is not with the delusion that I can contribute anything new to the prolonged discussions [8]and controversies to which it has given rise. I desire only to indicate in what way it affects the practical efforts of biologists to gain a better understanding of the living organism, whether regarded as a group of existing phenomena or as a product of the evolutionary process; and I shall speak of it, not in any abstract or speculative way, but from the standpoint of the working naturalist. The problem of which I speak is that of organic mechanism and its relation to that of organic adaptation. How in general are the phenomena of life related to those of the non-living world? How far can we profitably employ the hypothesis that the living body is essentially an automaton or machine, a configuration of material particles, which, like an engine or a piece of clockwork, owes its mode of operation to its physical and chemical construction? It is not open to doubt that the living body is a machine. It is a complex chemical engine that applies the energy of the food-stuffs to the performance of the work of life. But is it something more than a machine? If we may imagine the physico-chemical analysis of the body to be carried through to the very end, may we expect to find at last an unknown something that transcends such analysis and is neither a form of physical energy nor anything given in the physical or chemical configuration of the body? Shall we find anything corresponding to the usual popular conception—which was also along the view of physiologists—that the body is "animated" by a specific "vital principle," or "vital force," a dominating "archæus" that exists only in the realm of organic nature? If such a principle exists, then the mechanistic hypothesis fails and the fundamental problem of biology becomes a problem sui generis.
In its bearing on man's place in nature this question is one of the most momentous with which natural science has to deal, and it has occupied the attention of thinking men in every age. I cannot trace its history, but it will be [9]worth our while to place side by side the words of three of the great leaders of modern scientific and philosophic thought. The saying has been attributed to Descartes, "Give me matter and I will construct the world"—meaning by this the living world as well as the non-living; but Descartes specifically excepted the human mind. I do not know whether the great French philosopher actually used these particular words, but they express the essence of the mechanistic hypothesis that he adopted. Kant utterly repudiated such a conception in the following well known passage: "It is quite certain that we cannot become adequately acquainted with organized creatures and their hidden potentialities by means of the merely mechanical principles of nature, much less can we explain them; and this is so certain that we may boldly assert that it is absurd for man even to make such an attempt or to hope that a Newton may one day arise who will make the production of a blade of grass comprehensible to us according to natural laws that have not been ordered by design. Such an insight we must absolutely deny to man." Still, in another place Kant admitted that the facts of comparative anatomy give us "a ray of hope, however faint, that something may be accomplished by the aid of the principle of the mechanism of nature, without which there can be no science in general." It is interesting to turn from this to the bold and aggressive assertion of Huxley: "Living matter differs from other matter in degree and not in kind, the microcosm repeats the macrocosm; and one chain of causation connects the nebulous origin of suns and planetary systems with the protoplasmic foundations of life and organization."
Do not expect me to decide where such learned doctors disagree; but I will at this point venture on one comment which may sound the key-note of this address. Perhaps we shall find that in the long run and in the large sense [10]Kant was right; but it is certain that to-day we know very much more about the formation of the living body, whether a blade of grass or a man, than did the naturalists of Kant's time; and for better or for worse the human mind seems to be so constituted that it will continue its efforts to explain such matters, however difficult they may seem to be. But I return to our more specific inquiry with the remark that the history of physiology in the past two hundred years has been the history of a progressive restriction of the notion of a "vital force" or "vital principle" within narrower and narrower limits, until at present it may seem to many physiologists that no room for it remains within the limits of our biological philosophy. One after another the vital activities have been shown to be in greater or less degree explicable or comprehensible considered as physico-chemical operations of various degrees of complexity. Every physiologist will maintain that we cannot name one of these activities, not even thought, that is not carried on by a physical mechanism. He will maintain further that in most cases the vital actions are not merely accompanied by physico-chemical operations but actually consist of them; and he may go so far as definitely to maintain that we have no evidence that life itself can be regarded as anything more than their sum total. He is able to bring forward cogent evidence that all modes of vital activity are carried on by means of energy that is set free in protoplasm or its products by means of definite chemical processes collectively known as metabolism. When the matter is reduced to its lowest terms, life, as thus viewed, seems to have its root in chemical change; and we can understand how an eminent German physiologist offers us a definition or characterization of life that runs: "The life-process consists in the metabolism of proteids." I ask your particular attention to this definition since I now wish to contrast with it another and very different one.
[11]I shall introduce it to your attention by asking a very simple question. We may admit that digestion, for example, is a purely chemical operation, and one that may be exactly imitated outside the living body in a glass flask. My question is, how does it come to pass that an animal has a stomach?—and, pursuing the inquiry, how does it happen that the human stomach is practically incapable of digesting cellulose, while the stomachs of some lower animals, such as the goat, readily digest this substance? The earlier naturalists, such as Linnaeus, Cuvier or Agassiz, were ready with a reply which seemed so simple, adequate and final that the plodding modern naturalist cannot repress a feeling of envy. In their view plants and animals are made as they were originally created, each according to its kind. The biologist of to-day views the matter differently; and I shall give his answer in the form in which I now and then make it to a student who may chance to ask why an insect has six legs and a spider eight, or why a yellowbird is yellow and a bluebird blue. The answer is: "For the same reason that the elephant has a trunk." I trust that a certain rugged pedagogical virtue in this reply may atone for its lack of elegance. The elephant has a trunk, as the insect has six legs, for the reason that such is the specific nature of the animal; and we may assert with a degree of probability that amounts to practical certainty that this specific nature is the outcome of a definite evolutionary process, the nature and causes of which it is our tremendous task to determine to such extent as we may be able. But this does not yet touch the most essential side of the problem. What is most significant is that the clumsy, short-necked elephant has been endowed—"by nature," as we say—with precisely such an organ, the trunk, as he needs to compensate for his lack of flexibility and agility in other respects. If we are asked why the elephant has a trunk, we must answer because the animal [12]needs it. But does such a reply in itself explain the fact? Evidently not. The question which science must seek to answer, is how came the elephant to have a trunk; and we do not properly answer it by saying that it has developed in the course of evolution. It has been well said that even the most complete knowledge of the genealogy of plants and animals would give us no more than an ancestral portrait-gallery. We must determine the causes and conditions that have cooperated to produce this particular result if our answer is to constitute a true scientific explanation. And evidently he who adopts the machine-theory as a general interpretation of vital phenomena must make clear to us how the machine was built before we can admit the validity of his theory, even in a single case. Our apparently simple question as to why the animal has a stomach has thus revealed to us the full magnitude of the task with which the mechanist is confronted; and it has brought us to that part of our problem that is concerned with the nature and origin of organic adaptations. Without tarrying to attempt a definition of adaptation I will only emphasize the fact that many of the great naturalists, from Aristotle onward, have recognized the purposeful or design-like quality of vital phenomena as their most essential and fundamental characteristic. Herbert Spencer defined life as the continuous adjustment of internal relations to external relations. It is one of the best that has been given, though I am not sure that Professor Brooks has not improved upon it when he says that life is "response to the order of nature." This seems a long way from the definition of Verworn, heretofore cited, as the "metabolism of proteids." To this Brooks opposes the telling epigram: "The essence of life is not protoplasm but purpose."
Without attempting adequately to illustrate the nature of organic adaptations, I will direct your attention to what [13]seems to me one of their most striking features regarded from the mechanistic position. This is the fact that adaptations so often run counter to direct or obvious mechanical conditions. Nature is crammed with devices to protect and maintain the organism against the stress of the environment. Some of these are given in the obvious structure of the organism, such as the tendrils by means of which the climbing plant sustains itself against the action of gravity or the winds, the protective shell of the snail, the protective colors and shapes of animals, and the like. Any structural feature that is useful because of its construction is a structural adaptation; and when such adaptations are given the mechanist has for the most part a relatively easy task in his interpretation. He has a far more difficult knot to disentangle in the case of the so-called functional adaptations, where the organism modifies its activities (and often also its structure) in response to changed conditions. The nature of these phenomena may be illustrated by a few examples so chosen as to form a progressive series. If a spot on the skin be rubbed for some time the first result is a direct and obviously mechanical one; the skin is worn away. But if the rubbing be continued long enough, and is not too severe, an indirect effect is produced that is precisely the opposite of the initial direct one; the skin is replaced, becomes thicker than before, and a callus is produced that protects the spot from further injury. The healing of a wound involves a similar action. Again, remove one kidney or one lung and the remaining one will in time enlarge to assume, as far as it is able, the functions of both. If the leg of a salamander or a lobster be amputated, the wound not only heals but a new leg is regenerated in place of that which has been lost. If a flatworm be cut in two, the front piece grows out a new tail, the hind piece a new head, and two perfect worms result. Finally, it has been found in certain cases, including animals as highly organized as [14]salamanders, that if the egg be separated into two parts at an early period of development each part develops into a perfect embryo animal of half the usual size, and a pair of twins results. In each of these cases the astonishing fact is that a mechanical injury sets up in the organism a complicated adaptive response in the form of operations which in the end counteract the initial mechanical effect. It is no doubt true that somewhat similar self-adjustments or responses may be said to take place in certain non-living mechanical systems, such as the spinning top or the gyroscope; but those that occur in the living body are of such general occurrence, of such complexity and variety, and of so design-like a quality, that they may fairly be regarded as among the most characteristic of the vital activities. It is precisely this characteristic of many vital phenomena that renders their accurate analysis so difficult and complex a task; and it is largely for this reason that the biological sciences, as a whole, still stand far behind the physical sciences, both in precision and in completeness of analysis.
What is the actual working attitude of naturalists towards the general problem that I have endeavored to outline? It would be a piece of presumption for me to speak for the body of working biologists, and I will therefore speak for only one of them. It is my own conviction that whatever be the difficulties that the mechanistic hypothesis has to face, it has established itself as the most useful working hypothesis that we can at present employ. I do not mean to assert that it is adequate, or even true. I believe only that we should make use of it as a working program, because the history of biological research proves it to have been a more effective and fruitful means of advancing knowledge than the vitalistic hypothesis. We should therefore continue to employ it for this purpose until it is clearly shown to be untenable. Whether [15]we must in the end adopt it will depend on whether it proves the simplest hypothesis in the large sense, the one most in harmony with our knowledge of nature in general. If such is the outcome, we shall be bound by a deeply lying instinct that is almost a law of our intellectual being to accept it, as we have accepted the Copernican system rather than the Ptolemaic. I believe I am right in saying that the attitude I have indicated as a more or less personal one is also that of the body of working biologists, though there are some conspicuous exceptions.
In endeavoring to illustrate how this question actually affects research I will offer two illustrative cases, one of which may indicate the fruitfulness of the mechanistic conception in the analysis of complex and apparently mysterious phenomena, the other the nature of the difficulties that have in recent years led to attempts to re-establish the vitalistic view. The first example is given by the so-called law or principle of Mendel in heredity. The principle revealed by Mendel's wonderful discovery is not shown in all the phenomena of heredity and is probably of more or less limited application. It possesses however a profound significance because it gives almost a demonstration that a definite, and perhaps a relatively simple, mechanism must lie behind the phenomena of heredity in general. Hereditary characters that conform to this law undergo combinations, disassociations and recombinations which in certain way suggest those that take place in chemical reactions; and like the latter they conform to definite quantitative rules that are capable of arithmetical formulation. This analogy must not be pressed too far; for chemical reactions are individually definite and fixed, while those of the hereditary characters involve a fortuitous element of such a nature that the numerical result is not fixed or constant in the individual case but follows the law of probability in the aggregate of individuals. Nevertheless, it is possible, and [16]has already become the custom, to designate the hereditary organization by symbols or formulas that resemble those of the chemist in that they imply the quantitative results of heredity that follow the union of compounds of known composition. Quantitative prediction—not precisely accurate, but in accordance with the law of probability—has thus become possible to the biological experimenter on heredity. I will give one example of such a prediction made by Professor Cuénot in experimenting on the heredity of color in mice (see the following table). The experiment extended through three generations. Of the four grandparents three were pure white albinos, identical in outward appearance, but of different hereditary capacity, while the fourth was a pure black mouse. The first pair of grandparents consisted of an albino of gray ancestry, AG, and one of black ancestry, AB. The second pair consisted of an albino of yellow ancestry, AY, and a black mouse, CB. The result of the first union, AG x AB is to produce again pure white mice of the composition AGAB. The second union, AY x CB is to produce mice that appear pure yellow, and have the formula AYCB. What, now, [17]will be the result of uniting the two forms thus produced—i.e. AGAB × AYCB? Cuénot's prediction was that they should yield eight different kinds of mice, of which four should be white, two yellow, one black and one gray. The actual aggregate result of such unions, repeatedly performed, compared with the theoretic expectation, is shown in the foregoing table. As will be seen, the correspondence, though close, is not absolutely exact, yet is near enough to prove the validity of the principle on which the prediction was based, and we may be certain that had a much larger number of these mice been reared the correspondence would have been still closer. I have purposely selected a somewhat complicated example, and time will not admit of a full explanation of the manner in which this particular result was reached. I will however attempt to give an indication of the general Mendelian principle by means of which predictions of this kind are made. This principle appears in its simplest form in the behavior of two contrasting characters of the same general type—for instance two colors, such as gray and white in mice. If two animals, which show respectively two such characters are bred together, only one of the characters (known as the "dominant") appears in the offspring, while the other (known as the "recessive") disappears from view. In the next generation, obtained by breeding these hybrids together, both characters appear separately and in a definite ratio, there being in the long run three individuals that show the dominant character to one that shows the recessive. Thus, in the case of gray and white mice, the first cross is always gray, while the next generation includes three grays to one white. This is the fundamental Mendelian ratio for a single pair of characters; and from it may readily be deduced the more complicated combinations that appear when two or more pairs of characters are considered together. Such combinations appear in definite series, the nature of which may be worked out by [18]a simple method of binomial expansion. By the use of this principle astonishingly accurate numerical predictions may be made, even of rather complex combinations; and furthermore, new combinations may be, and have been, artificially produced, the number, character and hereditary capacity of which are known in advance. The fundamental ratio for a single pair of characters is explained by a very simple assumption. When a dominant and a recessive character are associated in a hybrid, the two must undergo in some sense a disjunction or separation in the formation of the germ-cells of the hybrid. This takes place in a quite definite way, exactly half the germ-cells in each sex receiving the potentiality of the dominant character, the other half the potentiality of the recessive. This is roughly expressed by saying that the germ-cells are no longer hybrid, like the body in which they arise, but bear one character or the other; and although in a technical sense this is probably not precisely accurate, it will sufficiently answer our purpose. If, now, it be assumed that fertilization takes place fortuitously—that is that union is equally probable between germ-cells bearing the same character and those bearing opposite characters,—the observed numerical ratio in the following generation follows according to the law of probability. Thus is explained both the fortuitous element that differentiates these cases from exact chemical combinations, and the definite numerical relations that appear in the aggregate of individuals.

Grandparents	AG (white)		AB (white)		AY (white)		CB (black)
Parents		AGAB (white)				AYCB (yellow)
							Observed	Calculated
			{	AGAY ABAY AGAB ABAB		} } (White) }	81	76
			{
			{
Offspring			{
			{	AGCY ABCY		} (Yellow)	34	38
			{
			{	ABCB		(Black)	20	19
			{	AGCB		(Gray)	16	19
							151	152

Now, the point that I desire to emphasize is that one or two very simple mechanistic assumptions give a luminously clear explanation of the behavior of the hereditary characters according to Mendel's law, and at one stroke bring order out of the chaos in which facts of this kind at first sight seem to be. Not less significant is the fact that direct microscopical investigation is actually revealing in the germ-cells a physical mechanism that seems adequate [19]to explain the disjunction of characters on which Mendel's law depends; and this mechanism probably gives us also at least a key to the long standing riddle of the determination and heredity of sex. These phenomena are therefore becoming intelligible from the mechanistic point of view. From any other they appear as an insoluble enigma. When such progress as this is being made, have we not a right to believe that we are employing a useful working hypothesis?
But let us now turn to a second example that will illustrate a class of phenomena which have thus far almost wholly eluded all attempts to explain them. The one that I select is at present one of the most enigmatical cases known, namely, the regeneration of the lens of the eye in the tadpoles of salamanders. If the lens be removed from the eye of a young tadpole, the animal proceeds to manufacture a new one to take its place, and the eye becomes as perfect as before. That such a process should take place at all is remarkable enough; but from a technical point of view this is not the extraordinary feature of the case. What fills the embryologist with astonishment is the fact that the new lens is not formed in the same way or from the same material as the old one. In the normal development of the tadpole from the egg, as in all other vertebrate animals, the lens is formed from the outer skin or ectoderm of the head. In the replacement of the lens after removal it arises from the cells of the iris, which form the edge of the optic cup, and this originates in the embryo not from the outer skin but as an outgrowth from the brain. As far as we can see, neither the animal itself nor any of its ancestors can have had experience of such a process. How, then, can such a power have been acquired, and how does it inhere in the structure of the organism? If the process of repair be due to some kind of intelligent action, as some naturalists have supposed, why should not the higher [20]animals and man possess a similar useful capacity? To these questions biology can at present give no reply. In the face of such a case the mechanist must simply confess himself for the time being brought to a standstill; and there are some able naturalists who have in recent years argued that by the very nature of the case such phenomena are incapable of a rational explanation along the lines of a physico-chemical or mechanistic analysis. These writers have urged, accordingly, that we must postulate in the living organism some form of controlling or regulating agency which does not lie in its physico-chemical configuration and is not a form of physical energy—something that may be akin to a form of intelligence (conscious or unconscious), and to which the physical energies are in some fashion subject. To this supposed factor in the vital processes have been applied such terms as the "entelechy" (from Aristotle), or the "psychoid"; and some writers have even employed the word "soul" in this sense—though this technical and limited use of the word should not be confounded with the more usual and general one with which we are familiar. Views of this kind represent a return, in some measure, to earlier vitalistic conceptions, but differ from the latter in that they are an outcome of definite and exact experimental work. They are therefore often spoken of collectively as "neo-vitalism."
It is not my purpose to enter upon a detailed critique of this doctrine. To me it seems not to be science, but either a kind of metaphysics or an act of faith. I must own to complete inability to see how our scientific understanding of the matter is in any way advanced by applying such names as "entelechy" or "psychoid" to the unknown factors of the vital activities. They are words that have been written into certain spaces that are otherwise blank in our record of knowledge, and as far as I can see no more than this. It is my impression that we shall do [21]better as investigators of natural phenomena frankly to admit that they stand for matters that we do not yet understand, and continue our efforts to make them known. And have we any other way of doing this than by observation, experiment, comparison and the resolution of more complex phenomena into simpler components? I say again, with all possible emphasis, that the mechanistic hypothesis or machine-theory of living beings is not fully established, that it may not be adequate or even true; yet I can only believe that until every other possibility has realty been exhausted scientific biologists should hold fast to the working program that has created the sciences of biology. The vitalistic hypothesis may be held, and is held, as a matter of faith; but we cannot call it science without misuse of the word.
When we turn, finally, to the genetic or historical part of our task, we find ourselves confronted with precisely the same general problem as in case of the existing organism. Biological investigators have long since ceased to regard the fact of organic evolution as open to serious discussion. The transmutation of species is not an hypothesis or assumption, it is a fact accurately observed in our laboratories; and the theory of evolution is only questioned in the same very general way in which all the great generalizations of science are held open to modification as knowledge advances. But it is a very large question what has caused and determined evolution. Here, too, the fundamental problem is, how far the process may be mechanically explicable or comprehensible, how far it is susceptible of formulation in physico-chemical or mechanistic terms. The most essential part of this problem relates to the origin of organic adaptations, the production of the fit. With Kant, Cuvier and Linnaeus believed this problem scientifically insoluble. Lamarck attempted to find a solution in his theory of the [22]inheritance of the effects of use, disuse and other "acquired characters"; but his theory was insecurely based and also begged the question, since the power of adaptation through which use, disuse and the like produce their effects is precisely that which must be explained. Darwin believed he had found a partial solution in his theory of natural selection, and he was hailed by Haeckel as the biological Newton who had set at naught the obiter dictum of Kant. But Darwin himself did not consider natural selection as an adequate explanation, since he called to its aid the subsidiary hypotheses of sexual selection and the inheritance of acquired characters. If I correctly judge, the first of these hypotheses must be considered as of limited application if it is not seriously discredited, while the second can at best receive the Scotch verdict, not proven. In any case, natural selection must fight its own battles.
Latter day biologists have come to see clearly that the inadequacy of natural selection lies in its failure to explain the origin of the fit; and Darwin himself recognized clearly enough that it is not an originative or creative principle. It is only a condition of survival, and hence a condition of progress. But whether we conceive with Darwin that selection has acted mainly upon slight individual variations, or with DeVries that it has operated with larger and more stable mutations, any adequate general theory of evolution must explain the origin of the fit. Now, under the theory of natural selection, pure and simple, adaptation or fitness has a merely casual or accidental character. In itself the fit has no more significance than the unfit. It is only one out of many possibilities of change, and evolution by natural selection resolves itself into a series of lucky accidents. For Agassiz or Cuvier the fit is that which was designed to fit. For natural selection, pure and simple, the fit is that which happens to fit. I, for one, am [23]unable to find a logical flaw in this conception of the fit; and perhaps we may be forced to accept it as sufficient. But I believe that naturalists do not yet rest content with it. Darwin himself was repeatedly brought to a standstill, not merely by specific difficulties in the application of his theory, but also by a certain instinctive or temperamental dissatisfaction with such a general conclusion as the one I have indicated; and many able naturalists feel the same difficulty to-day. Whether this be justified or not, it is undoubtedly the fact that few working naturalists feel convinced that the problem of organic evolution has been fully solved. One of the questions with which research is seriously engaged is whether variations or mutations are indeterminate, as Darwin on the whole believed, or whether they may be in greater or less degree determinate, proceeding along definite lines as if impelled by a vis a tergo. The theory of "orthogenesis," proposed by Naegeli and Eimer, makes the latter assumption; and it has found a considerable number of adherents among recent biological investigators, including some of our own colleagues, who have made important contributions to the investigation of this fundamental question. It is too soon to venture a prediction as to the ultimate result. That evolution has been orthogenetic in the case of certain groups, seems to be well established, but many difficulties stand in the way of its acceptance as a general principle of explanation. The uncertainty that still hangs over this question and that of the heredity of acquired characters bears witness to the unsettled state of opinion regarding the whole problem, and to the inadequacy of the attempts thus far made to find its consistent and adequate solution.
Here, too, accordingly, we find ourselves confronted with wide gaps in our knowledge which open the way to vitalistic or transcendental theories of development. I think we should resist the temptation to seek such refuge. It is [24]more than probable that there are factors of evolution still unknown. We can but seek for them. Nothing is more certain than that life and the evolution of life are natural phenomena. We must approach them, and as far as I can see must attempt to analyze them, by the same methods that are employed in the study of other natural phenomena. The student of nature can do no more than strive towards the truth. When he does not find the whole truth there is but one gospel for his salvation—still to strive towards the truth. He knows that each forward step on the highway of discovery will bring to view a new horizon of regions still unknown. It will be an ill day for science when it can find no more fields to conquer. And so, if you ask whether I look to a day when we shall know the whole truth in regard to organic mechanism and organic evolution, I answer: No! But let us go forward.