STUDIES IN ANIMAL PSYCHOLOGY. — часть 2

It is almost impossible to take up a discussion of the time relations of neural processes without having to read of physiological and psychological time. The time of nerve transmission, we are told, is pure physiological time and has nothing whatever to do with psychic processes; the time occupied by the changes in brain centers is, on the contrary, psychological time. At the very beginning of my discussion of this subject I wish to have it clearly understood that I make no such distinction. If one phase of the neural process be called physiological time, with as good reason may all be so named. I prefer, therefore, to speak of the time relations of the neural process.

Of the value of reaction-time studies, one may well believe that it lies chiefly in the way of approach which they open to the understanding of the biological significance of the nervous system. Certainly they are not important as giving us knowledge of the time of perception, cognition, or association, except in so far as we discover the relations of these various processes and the conditions under which they occur most satisfactorily. To determine how this or that factor in the environment influences the activities of the nervous system, and in what way system may be adjusted to system or part-process to whole, is the task of the reaction-time investigator.

The problems of reaction time naturally fall within three classes: Those which deal with (1) nerve transmission rates; (2) the time relations of the spinal center activities, and (3) brain processes. Within each of these groups there are innumerable special problems for the comparative physiologist or psychologist. Under class 1, for instance, there is the determining of the rates of impulse transmission in the sensory and the motor nerves, (a) for a variety of stimuli, (b) for different strengths of each stimulus, (c) for different conditions of temperature, moisture, nourishment, fatigue, etc., in case of each stimulus, (d) and all this for hundreds of representative animals. From this it is clear that lines of work are not lacking.

Closely related to these problems of rate of transmission are certain fundamental problems concerning the nature of the nerve impulse or wave. Whether there is a nerve wave, the reaction-time worker has as favorable an opportunity to determine as anyone, and we have a right to expect him to do something along this line. The relations of the form of the nerve impulse to the rhythm of vital action, to fatigue and to inhibition are awaiting investigation. Some of the most important unsettled points of psychology depend upon those aspects of neural activities which we ordinarily refer to as phenomena of inhibition, and which the psychologist is helpless to explain so long as the physiological basis and conditions are not known.

Then, too, in the study of animals the relation of reaction time to instincts, habits, and the surroundings of the subject are to be noted. Variability and adaptability offer chances for extended biological inquiries; and it is from just such investigations as these that biology has reason to expect much. The development of activity, the relation of reflex action to instinctive, of impulsive to volitional, and the value of all to the organism, should be made clear by reaction-time study. Such are a few of the broad lines of inquiry which are before the comparative student of animal reaction time. It is useless to dwell upon the possibilities and difficulties of the work, they will be recognized by all who are familiar with the results of human studies.

In the study of the time relations of neural processes Helmholtz was the pioneer. By him, in 1850, the rate of transmission of the nerve impulse in the sciatic nerve of the frog was found to be about 27 meters per second4. Later Exner5 studied the time occupied by various processes in the nervous system of the frog by stimulating the exposed brain in different regions and noting the time which intervened before a contraction of the gastrocnemius in each case. Further investigation of the frog's reflex reaction time has been made by Wundt6, Krawzoff and Langendorff7, Wilson8 and others, but in no case has the method of study been that of the psychologist. Most of the work has been done by physiologists who relied upon vivisectional methods. The general physiology of the nervous system of the frog has been very thoroughly worked up and the papers of Sanders-Ezn9, Goltz10, Steiner11, Schrader12 and Merzbacher13, 14 furnish an excellent basis for the interpretation of the results of the reaction-time studies.

In the present investigation it has been my purpose to study the reactions of the normal frog by the reaction-time methods of the psychologist. Hitherto the amount of work done, the extent of movements or some other change has been taken as a measure of the influence of a stimulus. My problem is, What are the time relations of all these reactions? With this problem in mind I enter upon the following program: (1) Determination of reaction time to electrical stimuli: (a) qualitative, (b) quantitative, (c) for different strengths of current; (2) Determination of reaction time to tactual stimuli (with the same variations); (3) Auditory: (a) qualitative, (b) quantitative, with studies on the sense of hearing; (4) Visual: (a) qualitative, (b) quantitative, with observations concerning the importance of this sense in the life of the frog, and (5) Olfactory: (a) qualitative, (b) quantitative.

The present paper presents in rather bare form the results thus far obtained on electrical, tactual, and auditory reaction time; discussion of them will be deferred until a comparison of the results for the five kinds of stimuli can be given.

V. METHOD OF STUDY.

The measurements of reaction time herein considered were made with the Hipp Chronoscope. Cattell's 'Falling Screen' or 'Gravity Chronoscope' was used as a control for the Hipp. The Gravity Chronoscope consists of a heavy metal plate which slides easily between two vertical posts, with electrical connections so arranged that the plate, when released from the magnet at the top of the apparatus, in its fall, at a certain point breaks an electric circuit and at another point further down makes the same circuit. The rate of fall of the plate is so nearly constant that this instrument furnishes an accurate standard time with which Hipp readings may be compared, and in accordance with which the Hipp may be regulated. For, since the rate of a chronoscope varies with the strength of the current in use, with the variations in temperature and with the positions of the springs on the magnetic bar, it is always necessary to have some standard for corrections. In these experiments the time of fall of the gravity chronoscope plate, as determined by the graphic method with a 500 S.V. electric tuning fork, was 125σ (i.e., thousandths of a second).

This period, 125σ, was taken as a standard, and each hour, before the beginning of reaction-time experiments, the time of the plate's fall was measured ten times with the Hipp, and for any variation of the average thus obtained from 125σ, the standard, the necessary corrections were made by changing the position of the chronoscope springs or the strength of the current.

The standard of comparison, 125σ, is shorter than most of the reaction times recorded, but since the time measured was always that from the breaking to the making of the circuit passing through the chronoscope it cannot be urged that there were errors resulting from the difference of magnetization which was caused by variations in the reaction time. But it is evident that the danger from differences in magnetization, if such exists, is not avoided in this way; instead, it is transferred from the reaction time proper to the period of preparation immediately preceding the reaction; for, from the moment the chronoscope is started until the stimulus is given a current is necessarily passing through the instrument. At a verbal signal from the operator the assistant started the chronoscope; the stimulus was then given by the operator, and the instrument recorded the time from the breaking of the circuit, effected by the stimulating apparatus, to the making of the circuit by the reaction of the animal. Despite precautions to prevent it, the period from the starting of the chronoscope to the giving of the stimulus was variable, and errors were anticipated, but a number of the tests proved that variations of even a second did not cause any considerable error.

A fairly constant current for the chronoscope was supplied by a six-cell 'gravity battery' in connection with two storage cells, GB (Fig. 6). This current could be used for two hours at a time without any objectionable diminution in its strength. The introduction of resistance by means of the rheostat, R, was frequently a convenient method of correcting the chronoscope.

Fig. 6. General Plan of Apparatus in Diagram. H, Hipp Chronoscope; R, rheostat; C, commutator; SC, storage cells; GB, 'Excello' gravity battery; F, Cattell's falling screen; T, reaction table; RK, reaction key; SK, Stimulating apparatus; K, key in chronoscope circuit; S, stimulus circuit.

Fig. 6 represents the general plan of the apparatus used in these experiments.

The general method of experimentation is in outline as follows:

1. At a 'ready' signal from the operator the assistant makes the chronoscope circuit by closing a key, K (Fig. 6), and then immediately starts the chronoscope.

2. Stimulus is given by the operator as soon as the chronoscope is started, and by this act the chronoscope circuit is broken and the record begun.

3. Animal reacts and by its movements turns a key, RK (Fig. 6), thus making the chronoscope circuit and stopping the record.

4. Assistant stops chronoscope and takes reading.

Fig. 7. Reaction Key. l, lever swung on pivot; p, p, posts for contacts with platinum plates on base; b, upright bar for string; s, spring for clamping string; w, wheel to carry string; c, c, chronoscope circuit; 1 and 2, points which are brought into contact by animal's reaction.

The steps of this process and the parts of the apparatus concerned in each may be clearly conceived by reference to the diagram given in Fig. 6. The various forms of stimulating apparatus used and the modification of the method will be described in the sections dealing with results. The same reaction key was used throughout (see Fig. 7). Its essential features are a lever l, pivoted in the middle and bearing a post at either end, p, p. From the middle of this lever there projected upward a small metal bar, b, through the upper part of which a string to the animal ran freely except when it was clamped by the spring, s. This string, which was attached to the subject's leg by means of a light elastic band, after passing through the bar ran over a wheel, w, and hung tense by reason of a five-gram weight attached to the end. Until everything was in readiness for an experiment the string was left free to move through the bar so that movement of the animal was not hindered, but the instant before the ready-signal was given it was clamped by pressure on s. The diagram shows the apparatus arranged for a reaction. The current is broken, since 1 and 2 are not in contact, but a slight movement of the animal turns the lever enough to bring 1 against 2, thus making the circuit and stopping the chronoscope. When the motor reaction of the subject was violent the string pulled out of the clamp so that the animal was free from resistance, except such as the string and weight offered. The five-gram weight served to give a constant tension and thus avoided the danger of error from this source. Between experiments the weight was placed on the table in order that there might be no strain upon the subject.

That the subject might be brought into a favorable position for an experiment without being touched by the operator a special reaction box was devised.

The animals used in these studies were specimens of Rana clamitans which were kept in a tank in the laboratory throughout the year.

VI. ELECTRIC REACTION TIME.

The reaction time to electrical stimuli was determined first because it seemed probable that this form of the pain reaction would be most useful for comparison with the auditory, visual, olfactory and tactual reactions. In this paper only the electrical and the tactual reaction times will be considered. The former will be divided into two groups: (1) Those resulting from a stimulus given by touching electrodes to the leg of the frog, and (2) those gotten by having the frog resting upon wires through which a current could be passed at any time.

Group 1 of the electrical reactions were taken under the following conditions. A reaction box about 40 cm. in diameter was used. The mean temperature of the experimenting room was about 20° C. In all cases the string was attached to the left hind leg of the frog, and the stimulus applied to the middle of the gastrocnemius muscle of the right hind leg. Reaction times were taken in series of ten, excluding those which were imperfect. As the moistness of the skin affects the strength of the electric stimulus received, it was necessary to moisten the animal occasionally, but as it did not seem advisable to disturb it after each experiment this was done at intervals of five minutes throughout the series. Were it not for this precaution it might be said that lengthening of the reaction times toward the end of a series simply indicated the weakening of the stimulus which resulted from the gradual drying of the skin. The stimulus in this group was applied by means of the stimulating apparatus of Fig. 6. It is merely two wire electrodes which could be placed upon the animal, with the additional device of a key for the breaking of the chronoscope circuit the instant the stimulus was given. The most serious objection to this method of stimulating is that there is a tactual as well as an electrical stimulus.

Before presenting averages, two representative series of reactions may be considered.

SERIES I. FROG B. APRIL 9, 1900. 10 A.M.

Temperature 19° C. String to left hind leg. Stimulus to right hind leg.

Strength of stimulating current 1.0 volt, .0001 ampère.

Average of 10, 147.4σ

SERIES 2. FROG F. ELECTRICAL STIMULUS.

Both are fairly representative series. They show the extremely large variations, in the case of series 1, from 102 to 179σ. In all these experiments such variation is unavoidable because it is impossible to have the conditions uniform. A very slight difference in the frog's position, which could not be detected by the operator, might cause considerable difference in the time recorded. Efforts were made to get uniform conditions, but the results seem to show that there is still much to be desired in this direction.

Tables VII. contains the results of four series of ten reactions each for frog A. It will be noticed that the time for the first five in each series is much shorter than that for the last five; this is probably indicative of fatigue.

TABLE VII.

REACTION TIME OF FROG A TO ELECTRICAL STIMULI.

TABLE VIII.

REACTION TIME OF FROG B TO ELECTRICAL STIMULI.

TABLE IX.

NORMAL AND REFLEX REACTION TIME OF SIX ANIMALS TO ELECTRICAL STIMULUS.

For D the average is for ten reactions.

B and E were males, F a female; the sex of the others was not determined by dissection and is uncertain.

Early in the experiments it became evident that there were three clearly defined types of reactions: there were a number of reactions whose time was shorter than that of the ordinary quick voluntary pain reaction, and there were also many whose time was considerably longer. The first type it was thought might represent the spinal reflex reaction time. For the purpose of determining whether the supposition was true, at the end of the series of experiments three of the frogs were killed and their reflex reaction time noted. This was done by cutting the spinal cord just back of the medulla, placing the animal on an experimenting board close to the reaction key with the thread from the key fastened to the left leg as in case of the previous work and stimulating the gastrocnemius with an induced current by the application of wire electrodes.

In Table IX. the reflex reaction times for the three animals are given.

The following results obtained with frog E show that the time of reaction increases with the increase in the time after death. The average of 20 reactions by E taken an hour after the cord had been cut was 45.5σ; the average of 20 taken twenty hours later was 55.85σ.

As a rule the reflex reactions were but slightly variable in time as is indicated by the accompanying series.

SERIES OF REFLEX REACTIONS OF FROG F.

Taken at rate of one per minute.

Throughout these experiments it was noticed that any stimulus might cause (1) a twitch in the limb stimulated, or (2) a twitch followed by a jump, or (3) a sudden jump previous to which no twitch could be detected. And it soon appeared that these types of reaction, as it seems proper to call them, would have to be considered in any determination of the mean reaction time. As proof of the type theory there is given (Fig. 8) a graphic representation of 277 reactions to the electrical stimulus.

Fig 8: Distribution of 277 reactions.

The column of figures at the left indicates the number of reactions at any point. Below the base line are the classes. For convenience of plotting the reactions have been grouped into classes which are separated by 25σ. Class 1 includes all reactions between 1σ and 25σ, class 2 all from 25σ to 50σ, and so on to 400σ, thereafter the classes are separated by 100σ. It is noticeable that there is one well-marked mode at 75σ. A second mode occurs at 175σ. This is the primary and in our present work the chiefly significant mode, since it is that of the quick instinctive reaction to a stimulus. At 500σ there is a third mode; but as such this has little meaning, since the reactions are usually pretty evenly distributed from 300σ on to 2000σ; if there is any grouping, however, it appears to be about 500σ and 800σ.

The first mode has already been called the reflex mode. The short reactions referred to usually lie between 40σ and 80σ, and since experiment has shown conclusively that the spinal reflex occupies about 50σ, there can be little doubt that the first mode is that of the reflex reaction time.

The second mode represents those reactions which are the result of central activity and control. I should be inclined to argue that they are what we usually call the instinctive and impulsive actions. And the remaining reactions represent such as are either purely voluntary, if any frog action can be so described, or, in other words, depend upon such a balancing of forces in the brain as leads to delay and gives the appearance of deliberate choice.

Everything points to some such classification of the types as follows: (1) Stimuli strong enough to be injurious cause the shortest possible reaction by calling the spinal centers into action, or if not spinal centers some other reflex centers; (2) slightly weaker stimuli are not sufficient to affect the reflex mechanism, but their impulse passes on to the brain and quickly discharges the primary center. There is no hesitation, but an immediate and only slightly variable reaction; just the kind that is described as instinctive. As would be expected, the majority of the frog's responses are either of the reflex or of this instinctive type. (3) There is that strength of stimulus which is not sufficient to discharge the primary center, but may pass to centers of higher tension and thus cause a response. This increase in the complexity of the process means a slower reaction, and it is such we call a deliberate response. Precisely this kind of change in neural action and in reaction time is at the basis of voluntary action. And (4) finally, the stimulus may be so weak that it will not induce a reaction except by repetition. Just above this point lies the threshold of sensibility, the determination of which is of considerable interest and importance.

Group 2 of the electrical reactions consists of three series taken to determine the relation of strength of stimulus to reaction time. The conditions of experimentation differed from those for group 1 in the following points: (1) The stimulus was applied directly by the making of a circuit through wires upon which the subject rested (Fig. 9); (2) the thread was attached to the right hind leg; (3) the thread, instead of being kept at the tension given by the 5-gram weight as in the former reactions, was slackened by pushing the upright lever of the reaction key one eighth of an inch toward the animal. This was done in order to avoid the records given by the slight twitches of the legs which precede the motor reaction proper. For this reason the reactions of group 2 are not directly comparable with those of group 1. Fig. 9 is the plan of the bottom of a reaction box 15 cm. at one end, 30 cm. at the other, 60 cm. long and 45 cm. deep. On the bottom of this, at one end, a series of interrupted circuits were arranged as shown in the figure. The wires were 1.2 cm. apart, and an animal sitting anywhere on the series necessarily touched two or more, so that when the stimulus key, X, was closed the circuit was completed by the animal's body; hence, a stimulus resulted. The stimulus key, X, was a simple device by which the chronoscope circuit, c, c, was broken at the instant the stimulus circuit, s, c, was made.

Cells of 'The 1900 Dry Battery' furnished the current used as a stimulus. Three different strengths of stimulus whose relative values were 1, 2 and 4, were employed in the series 1, 2 and 3. Careful measurement by means of one of Weston's direct-reading voltmeters gave the following values: 1 cell, 0.2 to 0.5 volt, 0.00001 to 0.00003 ampère. This was used as the stimulus for series 1. 2 cells, 0.5 to 1.0 volt, 0.00003 to 0.00006 ampère. This was used for series 2. 4 cells, 1.2 to 1.8 volt, 0.00007 to 0.0001 ampère. This was used for series 3.

Fig. 9. Ground Plan of Reaction Box for Electrical Stimuli (Group 2). IC, interrupted circuits; CC, chronoscope circuit; X, key for making stimulus circuit and breaking chronoscope circuit; B, stimulus battery; S, string from reaction key to animal. Scale 1/2.

The reactions now under consideration were taken in sets of 24 in order to furnish evidence on the problem of fatigue. The stimulus was given at intervals of one minute, and the subject was moistened at intervals of ten minutes. To obtain 24 satisfactory reactions it was usually necessary to give from thirty to forty stimulations. Five animals, numbers 1, 2, 4, 5, and 6, served as subjects. They were green frogs whose size and sex were as follows:

For most of these frogs a one-cell stimulus was near the threshold, and consequently the reaction time is extremely variable. In Table X. an analysis of the reactions according to the number of repetitions of the stimulus requisite for a motor reaction has been made. Numbers 1 and 5 it will be noticed reacted most frequently to the first stimulus, and for them 48 satisfactory records were obtained; but in case of the others there were fewer responses to the first stimulus, and in the tabulation of series 1 (Table XI.) averages are given for less than the regular sets of 24 reactions each.

TABLE X.

ANALYSIS OF REACTIONS TO ONE-CELL STIMULUS.

Table XI. is self-explanatory. In addition to the usual averages, there is given the average for each half of the sets, in order that the effect of fatigue may be noted. In general, for this series, the second half is in its average about one third longer than the first half. There is, therefore, marked evidence of tiring. The mean reaction time for this strength of stimulus is difficult to determine because of the extremely great variations. At one time a subject may react immediately, with a time of not over a fifth of a second, and at another it may hesitate for as much as a second or two before reacting, thus giving a time of unusual length. Just how many and which of these delayed responses should be included in a series for the obtaining of the mean reaction time to this particular stimulus is an extremely troublesome question. It is evident that the mode should be considered in this case rather than the mean, or at least that the mean should be gotten by reference to the mode. For example, although the reaction times for the one-cell stimulus vary all the way from 150σ to 1000σ or more, the great majority of them lie between 200σ and 400σ. The question is, how much deviation from the mode should be allowed? Frequently the inclusion of a single long reaction will lengthen the mean by 10σ or even 20σ. What is meant by the modal condition and the deviation therefrom is illustrated by the accompanying curve of a series of reaction times for the electric stimulus of group I.

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100  110  120  130  140  150  160  170  180  190  200  210  220  230230

The column of figures at the left indicates the number of reactions; that below the base line gives the reaction times in classes separated by 10σ. Of thirty-one reactions, seven are here in the class 170σ. This is the model class, and the mean gotten by taking the average of 31 reactions is 162σ. If the mode had been taken to represent the usual reaction time in this case, there would have been no considerable error. But suppose now that in the series there had occurred a reaction of 800σ. Should it have been used in the determination of the mean? If so, it would have made it almost 30σ greater, thus removing it considerably from the mode. If not, on what grounds should it be discarded? The fact that widely varying results are gotten in any series of reactions, points, it would seem, not so much to the normal variability as to accidental differences in conditions; and the best explanation for isolated reactions available is that they are due to such disturbing factors as would decrease the strength of the stimulus or temporarily inhibit the response. During experimentation it was possible to detect many reactions which were unsatisfactory because of some defect in the method, but occasionally when everything appeared to be all right an exceptional result was gotten. There is the possibility of any or all such results being due to internal factors whose influence it should be one of the objects of reaction-time work to determine; but in view of the fact that there were very few of these questionable cases, and that in series I, for instance, the inclusion of two or three reactions which stood isolated by several tenths of a second from the mode would have given a mean so far from the modal condition that the results would not have been in any wise comparable with those of other series, those reactions which were entirely isolated from the mode and removed therefrom by 200σ have been omitted. In series I alone was this needful, for in the other series there was comparatively little irregularity.

The results of studies of the reaction time for the one-cell electric stimulus appear in Table XI. The first column of this table contains the average reaction time or mean for each subject. Nos. 2 and 4 appeared to be much less sensitive to the current than the others, and few responses to the first stimulus could be obtained. Their time is longer than that of the others, and their variability on the whole greater. Individual differences are very prominent in the studies thus far made on the frog. The one-cell stimulus is so near the threshold that it is no easy matter to get a mean which is significant. Could the conditions be as fully controlled as in human reaction time it would not be difficult, but in animal work that is impossible. No attempt has thus far been made to get the reaction time in case of summation effects except in occasional instances, and in so far as those are available they indicate no great difference between the normal threshold reaction and the summation reaction, but on this problem more work is planned.

There are large mean variations in Table XI., as would be anticipated. Since the reactions were taken in sets of 24, the means of each set as well as that of the total are given, and also, in columns 4 and 5, the means of the first half and the last half of each set.

A comparison of Tables XI., XII. and XIII. makes clear the differences in reaction time correlated with differences in the strength of an electric stimulus. For Table XI., series I, the relative value of the stimulus was I; for Table XII., series 2, it was 2, and for Table XIII., series 3, it was 4. Throughout the series from I to 3 there is a rapid decrease in the reaction time and in the variability of the same. The reaction time for stimulus I, the so-called threshold, is given as 300.9σ; but of the three it is probably the least valuable, for reasons already mentioned. The mean of the second series, stimulus 2, is 231.5σ while that of the third, stimulus 4, is only 103.1σ. This great reduction in reaction time for the four-cell stimulus apparently shows the gradual transition from the deliberate motor reaction, which occurs only after complex and varied central neural activities, and the purely reflex reaction, which takes place as soon as the efferent impulse can cause changes in the spinal centers and be transmitted as an afferent impulse to the muscular system.

TABLE XI.

ELECTRICAL STIMULUS REACTION TIME. SERIES 1.

TABLE XII.

ELECTRICAL STIMULUS REACTION TIME. SERIES 2.

For No. 5 the averages are for two sets of 18 each; for all the others there are 24 in each set.

TABLE XIII.

ELECTRICAL STIMULUS REACTION TIME. SERIES 3.

For each animal there are two sets of 24 reactions each.

The spinal reflex for a decapitated frog, as results previously discussed show, is approximately 50σ; and every time the four-cell stimulus is given this kind of a reaction results. There is a slight twitch of the legs, immediately after which the animal jumps. Now for all these series the thread was slackened by one eighth of an inch, but the reflex time was determined without this slack. Calculation of the lengthening of the reaction time due to the slack indicated it to be between 20 and 30σ, so if allowance be made in case of the reactions to the four-cell stimulus, the mean becomes about 70σ, or, in other words, nearly the same as the spinal reflex. The conclusion seems forced, therefore, that when a stimulus reaches a certain intensity it produces the cord response, while until that particular point is reached it calls forth central activities which result in much longer and more variable reaction times. It was said above that the series under consideration gave evidence of the gradual transition from the reflex to the volitional in reaction time. Is this true, or do we find that there are well-marked types, between which reactions are comparatively rare? Examination of the tables VII., VIII., IX., XI., XII. and XIII. will show that between 70σ and 150σ there is a break. (In tables XI., XII. and XIII., allowance must always be made for the slack in the thread, by subtracting 30σ.) All the evidence furnished on this problem by the electrical reaction-time studies is in favor of the type theory, and it appears fairly clear that there is a jump in the reaction time from the reflex time of 50-80σ, to 140 or 150σ, which may perhaps be taken as the typical instinctive reaction time. From 150σ up there appears to be a gradual lengthening of the time as the strength of the stimulus is decreased, until finally the threshold is reached, and only by summation effect can a response be obtained.

The most important averages for the three series have been arranged in Table XIV. for the comparison of the different subjects. Usually the reaction time for series 3 is about one half as long as that for series 2, and its variability is also not more than half as large. In the small variability of series 3 we have additional reason for thinking that it represents reflexes, for Table IX. gives the mean variation of the reflex as not more than 8σ, and the fact that the means of this series are in certain cases much larger is fully explained by the greater opportunity for variation afforded by the slack in the thread.

TABLE XIV.

MEANS, ETC., FOR EACH SUBJECT FOR THE THREE SERIES. (TIME IN σ)

A striking fact is that the averages for the first and last half of sets of reactions differ more for the weak than for the strong stimulus. One would naturally expect, if the increase were a fatigue phenomenon purely, that it would be greatest for the strongest stimulus; but the results force us to look for some other conditions than fatigue. A stimulus that is sufficiently strong to be painful and injurious to an animal forces an immediate response so long as the muscular system is not exhausted; but where, as in series 1 and 2 of the electrical stimulus, the stimulus is not harmful, the reason for a sudden reaction is lacking unless fear enters as an additional cause. Just as long as an animal is fresh and unfamiliar with the stimulus there is a quick reaction to any stimulus above the threshold, and as soon as a few experiences have destroyed this freshness and taught the subject that there is no immediate danger the response becomes deliberate. In other words, there is a gradual transition from the flash-like instinctive reaction, which is of vast importance in the life of such an animal as the frog, to the volitional and summation responses. The threshold electrical stimulus does not force reactions; it is a request for action rather than a demand, and the subject, although startled at first, soon becomes accustomed to the experience and responds, if at all, in a very leisurely fashion. The reaction time to tactual stimuli, soon to be considered, was determined by giving a subject only three or four stimulations a day; if more were given the responses failed except on repetition or pressure; for this reason the data on fatigue, or lengthening of reaction time toward the end of a series, are wanting in touch. A few tests for the purpose of discovering whether the time would lengthen in a series were made with results very similar to those of the threshold electrical stimulus; the chief difference lies in the fact that the responses to touch fail altogether much sooner than do those to the electrical stimulus. This, however, is explicable on the ground that the latter is a stimulus to which the animal would not be likely to become accustomed so soon as to the tactual.

If pure fatigue, that is, the exhaustion of the nervous or muscular system, appears anywhere in this work, it is doubtless in series 3, for there we have a stimulus which is so strong as to force response on penalty of death; the reaction is necessarily the shortest possible, and, as a matter of fact, the motor reaction (jump forward) here occupies little more time than the leg-jerk of a decapitated frog. This probably indicates that the reaction is a reflex, and that the slight increase in its length over that of the spinal reflex is due to occasional cerebellar origin; but of this there can be no certainly from the evidence herewith presented. At any rate, there is no possibility of a voluntary reaction to the strong current, and any changes in the general character of the reaction time in a series will have to be attributed to fatigue of the nervous or muscular systems. The second halves of the sets of series 3 are 5 per cent. longer than the first, and unless this is due to the partial exhaustion of the nervous system it is hard to find an explanation of the fact. Fatigue of the muscles concerned seems out of the question because the reactions occur at the rate of only one per minute, and during the rest interval any healthy and well-nourished muscle would so far recover from the effect of contraction that it would be able to continue the rhythmic action for long periods.

To the inquiry, Does fatigue in the experiments mean tiring by the exhaustion of nerve energy, or is the lengthening in reaction time which would naturally be attributed to tiring due to the fact that experience has shown quick reaction to be unnecessary? we shall have to reply that there is evidence in favor of both as factors. There can be little doubt that in case of the strong stimuli there is genuine fatigue which makes quick reaction impossible; but at the same time it is certain that the 40 to 50 per cent. increase of the second half of sets in series 1 over the first half can not be due to fatigue, for the strain is here evidently much less than for series 3. Rather, it would seem that habituation instead of exhaustion is the all-important cause of the difference in series 1 and 2. It becomes clear from these considerations that the repetition of a stimulus can never mean the repetition of an effect.

VII. TACTUAL REACTION TIME.

In the following work on the reactions to tactual stimulation the subject was placed in a large reaction box with a thread attached to one of its legs and passing to a reaction key, as in the experiments already described. The box in which the subject was confined was surrounded by movable cloth curtains to prevent the animal's escape and at the same time permit the experimenter to work without being seen by the frog.

Tactual stimulation was given by means of a hand key15 similar to that used for electrical stimulation which is represented in Fig. 6. The touch key ended in a hard-rubber knob which could be brought in contact with the skin of the subject. This key was fixed to a handle of sufficient length to enable the operator to reach the animal wherever it chanced to be sitting in the reaction box. Stimulation was given by allowing the rubber point of the touch key to come in contact with the skin in the middle region of the subject's back. As soon as the point touched the animal the chronoscope circuit was broken by the raising of the upper arm of the key.

As a precaution against reactions to visual stimuli, which it might well be supposed would appear since the subject could not in every case be prevented from seeing the approaching apparatus, the frog was always placed with its head away from the experimenter so that the eyes could not readily be directed toward the touch apparatus. Notwithstanding care in this matter, a reaction occasionally appeared which was evidently due to some disturbance preceding the tactual stimulus which served as a warning or preparation for the latter. All such responses were at once marked as questionable visual reactions and were not included in the series of touch reactions proper.

As has been mentioned in connection with the discussion of fatigue, it was found absolutely necessary to have the subjects perfectly fresh and active, and for this purpose it was advisable to give not more than three or four stimulations at any one time. The subject was usually kept in the reaction box from 30 to 45 minutes, dependent upon the success of the experiments. As the work progressed it became evident that the responses to the stimulus were becoming less and less certain and slower, that the subjects were becoming accustomed to the novel experience and no longer suffered the surprise which had been the cause of the prompt reactions at first. It seemed best for this reason not to continue the work longer than two weeks, and as a consequence it was impossible to base the averages on more than twenty reactions for each subject.

So far as the tension of the thread is concerned, the condition for the tactual reaction time was the same as that for the first group of electrical reaction-time experiments. In comparing the tactual with the electrical of series 1, 2 and 3, allowance must be made for the slack in the latter cases.

Selection of the tactual reaction times upon which the mean is based, has been made with reference to the mode for each set of experiments. Inspection of the curves given by the reactions of each subject indicated that the great majority of the responses lay between 100 and 300σ, and that those which were beyond these limits were isolated and, in all probability, exceptional reactions due to some undetected variation in conditions which should throw them out of the regular series. On this account it was thought best to use only reactions between 100 and 300σ.

For convenience of comparison, again, the averages for the electrical reaction time of subjects A, B, C, D, E and F, and the same for the tactual reaction time of subjects 1, 2, 3, 4, 5 and 6 are herewith given together. All averages are for twenty reactions, except for D and 5, for which there are ten.

Besides the usual determination for the tactual reaction-time work on the six subjects named, there is given in Table XVI. the electrical reaction time of these animals to a two-cell current. Comparison of the electrical and tactual results are of interest in this case because the mean variation for each is about 34σ, being 34.3σ, for the electrical and 33.8σ, for the tactual.

TABLE XV.

TABLE XVI.

REACTION TIME FOR TACTUAL AND ELECTRICAL STIMULI.

¹For 5 the average of ten instead of twenty is given.

VIII. EQUAL VARIABILITY AS A CRITERION OF COMPARABILITY OF REACTION TIME FOR DIFFERENT KINDS OF STIMULI.

Since variability as indicated in the study of the influence of different strengths of electrical stimulus becomes less as the stimulus increases, parity in variability for different stimuli offers a basis for the comparison of reaction times. Certain it is that there is no use in comparing the reaction times for different senses or different qualities of stimuli unless the relative values of the stimuli are taken into consideration; but how are these values to be determined unless some such index as variability is available? If the reaction time to tactual stimuli as here presented is to be studied in its relation to the electrical reaction time, it will mean little simply to say that the former is longer than the latter, because the electrical reaction time for a one-cell stimulus happens to be somewhat less than that for the particular tactual stimulus used. For it is clear that this tactual reaction time is really shorter than the reaction time to a weak current. In making variability a basis of comparison it must be assumed that the strength of stimulus is the important factor, and that all other variable conditions are, so far as possible, excluded. If, now, on the basis of parity in variability we compare the tactual and electrical reaction times, it is apparent that the tactual is considerably longer. The tactual average of Table XV. is 205.7σ, while the electrical reaction time which has approximately the same variability is 172.1σ. It may well be objected that I have no right to make variability the basis of my comparison in these experiments, because the work for the various kinds of stimuli was done under different conditions. Admitting the force of this objection, and at the same time calling attention to the fact that I do not wish to lay any stress on the results of the comparisons here made, I take this opportunity to call attention to the possibility of this criterion.

The use of variability as a basis of comparison would involve the assumptions (1) that a certain intensity of every stimulus which is to be considered is capable of producing the shortest possible, or reflex reaction, and that this reaction is at the same time the least variable; (2) that as the strength of a stimulus decreases the variability increases until the threshold is reached.

Suppose, now, it is our desire to compare the results of reactions to different intensities of electrical and tactual stimuli; let the figures be as follows:

In the double columns the results for electrical stimuli are given first, and in the second column are the tactual. Stimulus 8 is assumed to be of sufficient strength to induce what may be designated as forced movement, and whatever the quality of the stimulus this reaction time is constant. I make this statement theoretically, although all the evidence which this work furnishes is in support of it. So, likewise, is the variability of this type of reaction time small and nearly constant. At the other extreme, stimulus 1 is so weak as to be just sufficient to call forth a response; it is the so-called threshold stimulus. Whether all qualities of stimulus will give the same result here is a question to be settled by experimentation. Wundt contends that such is the case, but the observations I have made on the electrical and tactual reactions of the frog cause me to doubt this assumption. It seems probable that the 'just perceptible stimulus reaction time' is by no means the same thing for different qualities of stimulus. Those modifications of the vital processes which alone enable organisms to survive, make their appearance even in the response to the minimal stimulus. In one case the just perceptible stimulus may cause nothing more than slight local changes in circulation, excretion, muscular action; in another it may produce, just because of the particular significance of the stimulus to the life of the organism, a violent and sudden motor reaction. But grant, if you will, that the threshold reaction time is the same for all kinds of stimuli, and suppose that the variability is fairly constant, then, between the two extremes of stimuli, there are gradations in strength which give reaction times of widely differing variabilities. If, now, at some point in the series, as, for instance, to stimulus 2, the variability for different kinds of stimuli is the same either with reference to the reaction time (ratio) or absolutely, what interpretation is to be put upon the fact? Is it to be regarded as merely a matter of chance, and unworthy of any special attention, or should it be studied with a view to finding out precisely what variability itself signifies? It is obvious that any discussion of this subject, even of the possible or probable value of variability as a criterion for the comparative study of stimuli, can be of little value so long as we do not know what are the determining factors of variations of this sort. The only suggestion as to the meaning of such a condition (i.e., equal variability at some point)—and our studies seem to show it for touch and electrical stimulation—which I feel justified in offering at present, is that parity in variability indicates equality in strength of stimuli, that is, the electrical stimulus which has a reaction time of the same variability as a tactual stimulus has the same effect upon the peripheral nervous system as the tactual, it produces the same amplitude and perhaps the same form of wave, but the reaction times for the two stimuli differ because of the biological significance of the stimuli. The chances are that this is wholly dependent upon the central nervous system.

IX. SUMMARY.

1. This paper gives the results of some experiments on the frog to determine its electrical and tactual reaction time. It is the beginning of comparative reaction-time studies by which it is hoped important information may be gained concerning the significance and modes of action of the nervous system. Comparative physiology has already made clear that the time relations of neural processes deserve careful study.

2. According to the strength of the stimulus, electric stimulation of the frog causes three types of reaction: (1) A very weak or threshold stimulus results in a deliberate or delayed reaction, the time of which may be anywhere from 300σ (thousandths of a second) to 2,000σ. (2) A very strong stimulus causes a spinal reflex, whose time is from 50 to 80σ; and (3) a stimulus of intermediate strength causes a quick instinctive reaction of from 150 to 170σ in duration.

3. The reaction time for electric stimuli whose relative values were 1, 2 and 4 were found to be 300.9σ, 231.5σ and 103.1σ.

4. The reaction time of the frog to a tactual stimulus (contact of a rubber point) is about 200σ.

5. The variability of reaction times of the frog is great, and increases as the strength of the stimulus decreases.

6. When two kinds of stimuli (e.g., electrical and tactual) give reaction times of equal variability, I consider them directly comparable.

7. According to this criterion of comparability the reaction time to electric stimulation which is comparable with that to tactual is 172.1σ; and it is to be compared with 205.7σ. Both of these have a variability of approximately 34σ. On this basis one may say that the tactual reaction time is considerably longer than the electrical.

PART III. AUDITORY REACTIONS OF FROGS.

X. HEARING IN THE FROG.

A. Influences of Sounds in the Laboratory.

After determining the simple reaction time of the green frog to tactual and electrical stimulation, I attempted to do the same in case of auditory stimuli. In this I was unsuccessful because of failure to get the animal to give a motor response which could be recorded. The animal was placed in an experimenting box with a string attached to one hind leg as in the experiments described in Part II., and after it had become accustomed to the situation a sound was made. A wide range of sounds were tried, but to none except the croak of another frog was a motor reaction frequently given. Even a loud noise, such as the explosion of a large pistol cap, caused a visible motor reaction only in rare cases. In fifty trials with this stimulus I succeeded in getting three reactions, and since all of them measured between 230 and 240σ it is perhaps worth while to record the result as indicative of the auditory reaction time. As these were the only measurements obtained, I have no satisfactory basis for the comparison of auditory with other reaction times.

The remarkable inhibition of movement shown by the frog in the presence of strong auditory stimulation, at least what is for the human being a strong stimulus, led me to inquire concerning the limits and delicacy of the sense of hearing in frogs. In the vast quantity of literature on the structure and functions of the sense organs of the animal I have been able to find only a few casual remarks concerning hearing.

In approaching the problem of frog audition we may first examine the structure of the ear for the purpose of ascertaining what sounds are likely to affect the organ. There is no outer ear, but the membrana tympani, or ear drum, covered with skin, appears as a flat disc from 5 to 10 mm. in diameter on the side of the head just back of the eye and a little below it. In the middle ear there is but one bone, the columella, forming the connecting link between the tympanum and the internal ear. The inner ear, which contains the sense organs, consists of a membranous bag, the chief parts of which are the utriculus, the sacculus, the lagena, and the three semicircular canals. The cavity of this membranous labyrinth is filled with a fluid, the endolymph; and within the utriculus, sacculus and lagena are masses of inorganic matter called the otoliths. The auditory nerve terminates in eight sense organs, which contain hair cells. There is no cochlea as in the mammalian ear. The assumption commonly made is that vibrations in the water or air by direct contact cause the tympanic membrane to vibrate; this in turn causes a movement of the columella, which is transmitted to the perilymphatic fluid of the inner ear. The sensory hair cells are disturbed by the movements of the otoliths in the endolymph, and thus an impulse is originated in the auditory nerve which results in a sensation more or less resembling our auditory sensation. It is quite probable that the frog's sense of hearing is very different from ours, and that it is affected only by gross air vibrations. This conclusion the anatomy of the ear supports.

Although there does not seem to be a structural basis for a delicate sense of hearing, one must examine the physiological facts at hand before concluding that frogs do not possess a sense of hearing similar to our own. First, the fact that frogs make vocal sounds is evidence in favor of the hearing of such sounds at least, since it is difficult to explain the origin of the ability to make a sound except through its utility to the species. Granting, however, that a frog is able to hear the croaks or pain-screams of its own species, the range of the sense still remains very small, for although the race of frogs makes a great variety of sounds, any one species croaks within a narrow range.

Having satisfied myself that motor reactions for reaction-time measurements could not be gotten to any ordinary sounds in the laboratory, I tried the effect of the reflex croaking of another frog of the same species. In attempting to get frogs to croak regularly, I tested the effect of removing the brain. The animals are said to croak reflexly after this operation whenever the back is stroked; but for some reason I have never been successful in getting the reaction uniformly. In many cases I was able to make normal animals croak by rubbing the back or flanks, and to this sound the animals under observation occasionally responded by taking what looked like an attitude of attention. They straightened up and raised the head as if listening. In no case have other motor responses been noticed; and the above response was so rare that no reaction-time measurements could be made.

Again, while working with the green frog on habit formation, I one day placed two animals in a labyrinth from which they could escape by jumping into a tank of water. Several times when one frog jumped into the water I noticed the other one straighten up and hold the 'listening' or 'attentive' attitude for some seconds. As the animals could not see one another this is good evidence of their ability to hear the splash made by a frog when it strikes the water.

B. Influence of Sounds in Nature.

In order to learn how far fear and artificial conditions were causes of the inhibition of response to sounds in the laboratory, and how far the phenomenon was indicative of the animal's inability to perceive sounds, I observed frogs in their native haunts.

By approaching a pond quietly, it is easy to get within a few yards of frogs sitting on the banks. In most cases they will not jump until they have evidence of being noticed. Repeatedly I have noted that it is never possible to get near to any frogs in the same region after one has jumped in. In this we have additional proof that they hear the splash-sound. To make sure that sight was not responsible for this on-guard condition in which one finds the frogs after one of their number has jumped into the water, I made observations on animals that were hidden from one another. The results were the same. I therefore conclude that the splash of a frog jumping into the water is not only perceived by other frogs in the vicinity, but that it is a peculiarly significant sound for them, since it is indicative of danger, and serves to put them 'on watch.'