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diff --git a/.gitattributes b/.gitattributes new file mode 100644 index 0000000..6833f05 --- /dev/null +++ b/.gitattributes @@ -0,0 +1,3 @@ +* text=auto +*.txt text +*.md text diff --git a/8729-8.txt b/8729-8.txt new file mode 100644 index 0000000..7e477a9 --- /dev/null +++ b/8729-8.txt @@ -0,0 +1,10329 @@ +The Project Gutenberg EBook of The Dancing Mouse, by Robert M. Yerkes + +Copyright laws are changing all over the world. Be sure to check the +copyright laws for your country before downloading or redistributing +this or any other Project Gutenberg eBook. + +This header should be the first thing seen when viewing this Project +Gutenberg file. Please do not remove it. Do not change or edit the +header without written permission. + +Please read the "legal small print," and other information about the +eBook and Project Gutenberg at the bottom of this file. Included is +important information about your specific rights and restrictions in +how the file may be used. You can also find out about how to make a +donation to Project Gutenberg, and how to get involved. + + +**Welcome To The World of Free Plain Vanilla Electronic Texts** + +**eBooks Readable By Both Humans and By Computers, Since 1971** + +*****These eBooks Were Prepared By Thousands of Volunteers!***** + + +Title: The Dancing Mouse + A Study in Animal Behavior + +Author: Robert M. Yerkes + +Release Date: August, 2005 [EBook #8729] +[Yes, we are more than one year ahead of schedule] +[This file was first posted on August 4, 2003] + +Edition: 10 + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THE PROJECT GUTENBERG EBOOK THE DANCING MOUSE *** + + + + +Produced by Juliet Sutherland, Michael Oltz, +Charles Franks and the Online Distributed Proofreading Team. + + + + +[Illustration: DANCING MICE--SNIFFING AND EATING.] + + + +THE ANIMAL BEHAVIOR SERIES. VOLUME I + + +THE DANCING MOUSE + +A Study in Animal Behavior + +BY + +ROBERT M. YERKES, Ph.D. +INSTRUCTOR IN COMPARATIVE PSYCHOLOGY IN HARVARD +UNIVERSITY + +The Cartwright Prize of the Alumni Association of the College of +Physicians and Surgeons, Columbia University, was awarded, in 1907, for an +Essay which comprised the first twelve chapters of this volume. + + +1907 + + + +IN LOVE AND GRATITUDE +THIS BOOK IS DEDICATED TO +MY MOTHER + + + + + +PREFACE + + +This book is the direct result of what, at the time of its occurrence, +seemed to be an unimportant incident in the course of my scientific work-- +the presentation of a pair of dancing mice to the Harvard Psychological +Laboratory. My interest in the peculiarities of behavior which the +creatures exhibited, as I watched them casually from day to day, soon +became experiment-impelling, and almost before I realized it, I was in the +midst of an investigation of their senses and intelligence. + +The longer I observed and experimented with them, the more numerous became +the problems which the dancers presented to me for solution. From a study +of the senses of hearing and sight I was led to investigate, in turn, the +various forms of activity of which the mice are capable; the ways in which +they learn to react adaptively to new or novel situations; the facility +with which they acquire habits; the duration of habits; the roles of the +various senses in the acquisition and performance of certain habitual +acts; the efficiency of different methods of training; and the inheritance +of racial and individually acquired forms of behavior. + +In the course of my experimental work I discovered, much to my surprise, +that no accurate and detailed account of this curiously interesting animal +existed in the English language, and that in no other language were all +the facts concerning it available in a single book. This fact, in +connection with my appreciation of the exceptional value of the dancer as +a pet and as material for the scientific study of animal behavior, has led +me to supplement the results of my own observation by presenting in this +little book a brief and not too highly technical description of the +general characteristics and history of the dancer. + +The purposes which I have had in mind as I planned and wrote the book are +three: first, to present directly, clearly, and briefly the results of my +investigation; second, to give as complete an account of the dancing mouse +as a thorough study of the literature on the animal and long-continued +observation on my own part should make possible; third, to provide a +supplementary text-book on mammalian behavior and on methods of studying +animal behavior for use in connection with courses in Comparative +Psychology, Comparative Physiology, and Animal Behavior. + +It is my conviction that the scientific study of animal behavior and of +animal mind can be furthered more just at present by intensive special +investigations than by extensive general books. Methods of research in +this field are few and surprisingly crude, for the majority of +investigators have been more deeply interested in getting results than in +perfecting methods. In writing this account of the dancing mouse I have +attempted to lay as much stress upon the development of my methods of work +as upon the results which the methods yielded. In fact, I have used the +dancer as a means of exhibiting a variety of methods by which the behavior +and intelligence of animals may be studied. As it happens the dancer is an +ideal subject for the experimental study of many of the problems of animal +behavior. It is small, easily cared for, readily tamed, harmless, +incessantly active, and it lends itself satisfactorily to a large number +of experimental situations. For laboratory courses in Comparative +Psychology or Comparative Physiology it well might hold the place which +the frog now holds in courses in Comparative Anatomy. + +Gratefully, and with this expression of my thanks, I acknowledge my +indebtedness to Professor Hugo Münsterberg for placing at my command the +resources of the Harvard Psychological Laboratory and for advice and +encouragement throughout my investigation; to Professor Edwin B. Holt for +valuable assistance in more ways than I can mention; to Professor Wallace +C. Sabine for generous aid in connection with the experiments on hearing; +to Professor Theobald Smith for the examination of pathological dancers; +to Miss Mary C. Dickerson for the photographs of dancing mice which are +reproduced in the frontispiece; to Mr. Frank Ashmore for additional +photographs which I have been unable to use in this volume; to Mr. C. H. +Toll for the drawings for Figures 14 and 20; to Doctors H. W. Rand and C. +S. Berry for valuable suggestions on the basis of a critical reading of +the proof sheets; and to my wife, Ada Watterson Yerkes, for constant aid +throughout the experimental work and in the preparation of this volume. + +R. M. Y. + +CAMBRIDGE, MASSACHUSETTS, + +August, 1907. + + + + + +CONTENTS + + +LIST OF ILLUSTRATIONS + +LITERATURE ON THE DANCING MOUSE + + +CHAPTER I + +CHARACTERISTICS, ORIGIN, AND HISTORY + +Peculiarities of the dancing mouse--Markings and method of keeping record +of individuals--The dancer in China and Japan (Kishi, Mitsukuri, Hatai)-- +Theories concerning the origin of the race: selectional breeding; the +inheritance of an acquired character; mutation, inheritance, and +selectional breeding; pathological changes; natural selection--Instances +of the occurrence of dancers among other kinds of mice--Results of +crossing dancer with other kinds of mice. + + +CHAPTER II + +FEEDING, BREEDING, AND DEVELOPMENT OF THE YOUNG + +Methods of keeping and caring for dancers--Cages, nest-boxes, and +materials for nest--Cleansing cages--Food supply and feeding--Importance +of cleanliness, warmth, and pure food--Relations of males and females, +fighting--The young, number in a litter--Care of young--Course of +development--Comparison of young of dancer with young of common mouse-- +Diary account of the course of development of a typical litter of dancers. + + +CHAPTER III + +BEHAVIOR: DANCE MOVEMENTS + +Dancing--Restlessness and excitability--Significance of restlessness-- +Forms of dance: whirling, circling, and figure-eights--Direction of +whirling and circling: right whirlers, left whirlers, and mixed whirlers-- +Sex differences in dancing--Time and periodicity of dancing--Influence of +light on activity--Necessity for prolonged observation of behavior. + + +CHAPTER IV + +BEHAVIOR: EQUILIBRATION AND DIZZINESS + +Muscular coordination--Statements of Cyon and Zoth concerning behavior-- +Control of movements, orientation, equilibration, movement on inclined +surfaces, climbing--The tracks of the dancer--Absence of visual +dizziness--Comparison of the behavior of the dancer with that of the +common mouse when they are rotated in a cyclostat--Behavior of blinded +dancers (Cyon, Alexander and Kreidl, Kishi)--Cyon's two types of dancer-- +Phenomena of behavior for which structural bases are sought: dance +movements; lack of response to sounds; deficiency in equilibrational +ability; lack of visual and rotational dizziness. + + +CHAPTER V + +STRUCTURAL PECULIARITIES AND BEHAVIOR + +The functions of the ear--Structure of the ear of the dancer as described +by Rawitz, by Panse, by Baginsky, by Alexander and Kreidl, and by Kishi-- +Cyon's theory of the relation of the semicircular canals to space +perception--Condition of the auditory organs--Condition of the +equilibrational organs--Condition of the sound-transmitting organs--The +bearing of the results of anatomical investigations upon the facts of +behavior. + + +CHAPTER VI + +THE SENSE OF HEARING + +Experiments on hearing in the dancer made by Rawitz, by Panse, by Cyon, by +Alexander and Kreidl, by Zoth, and by Kishi--Hearing and the voice-- +Methods of testing sensitiveness to sounds--Results of tests with adults-- +Importance of indirect method of experimentation--Results of tests with +young--The period of auditory sensitiveness--Individual differences. + + +CHAPTER VII + +THE SENSE OF SIGHT: BRIGHTNESS VISION + +What is known concerning sight in the dancer--Brightness vision and color +vision--Methods of testing brightness vision, the visual discrimination +apparatus--Motives for discrimination and choice--Punishment versus reward +as an incentive in animal experiments--Hunger as an incentive--An electric +stimulus as an incentive--Conditions for brightness vision tests-- +White-black vision--Evidence of preference--Check experiments--Conclusion. + + +CHAPTER VIII + + +THE SENSE OF SIGHT: BRIGHTNESS VISION (_Continued_) + +The delicacy of brightness discrimination--Methods of testing the dancer's +ability to detect slight differences in brightness--Results of tests with +gray papers--Relation of intensity of visual stimuli to the threshold of +discrimination--Weber's law apparatus and method of experimentation-- +Results of Weber's law tests--Practice effects, the training of vision-- +Description of the behavior of the dancer in the discrimination box +experiments--Modes of choice: by affirmation; by negation; by comparison-- +Evidence of indiscriminable visual conditions. + + +CHAPTER IX + +THE SENSE OF SIGHT: COLOR VISION + +Does the dancer see colors?--The food-box method of testing color vision-- +Waugh's food-box method--Results of tests--Tests by the use of colored +papers in the visual discrimination box--Yellow-red vision--Blue-orange +vision--Brightness vision _versus_ color vision--Brightness check +tests--Green-blue vision--Violet-red vision--Conclusions. + + +CHAPTER X + +THE SENSE OF SIGHT: COLOR VISION (_Continued_) + +The use of color filters--Testing color vision by the use of transmitted +light--Green-blue vision--Green-red vision--Blue-red vision--Stimulating +value of different portions of the spectrum--Does red appear darker to the +dancer than to us?--Conclusions concerning color vision--Structure of the +retina of the dancer and its significance. + + +CHAPTER XI + +THE ROLE OF SIGHT IN THE DAILY LIFE OF THE DANCER. + +Sight and general behavior--Behavior of blinded dancers--Experimental +tests of ability to perceive form--Visual guidance in mazes--Following +labyrinth paths in the dark--The relative importance of visual, olfactory, +and kinaesthetic stimuli--Conditions for the acquisition of a motor +habit--Conditions for the execution of an habitual act. + + +CHAPTER XII + +EDUCABILITY: METHODS OF LEARNING + +The modifiability of behavior--Educational value of experimental studies +of modifiability--Methods: the problem method; the labyrinth method; the +discrimination method--Relation of method to characteristics of animal-- +Simple test of the docility of the dancer--Lack of imitative tendency-- +Persistence of useless acts--Manner of profiting by experience--Individual +differences in initiative. + + +CHAPTER XIII + +HABIT FORMATION: THE LABYRINTH HABIT + +The labyrinth method--Problems--Preliminary tests--Comparison of the +behavior of the dancer in a maze with that of the common mouse--Evolution +of a labyrinth method--Records of time and records of errors--Simple and +effective method of recording the path--Curves of habit formation--Regular +and irregular labyrinths--Points for a standard labyrinth--Values and +defects of the labyrinth method. + + +CHAPTER XIV + +HABIT FORMATION: THE DISCRIMINATION METHOD + +Quantitative _versus_ qualitative results--Motives--Precautions-- +Preference--Results of systematic habit-forming experiments--Curves of +habit formation--Meaning of irregularity in curve--Individual +differences--Comparison of curves for discrimination habits with those for +labyrinth habits--Averages--The index of modifiability as a measure of +docility--Reliability of the index. + + +CHAPTER XV + +THE EFFICIENCY OF TRAINING METHODS + +Importance of measuring the efficiency of educational methods--Rapidity of +learning and permanency of modifications wrought by training--Results of a +study of the efficiency of discrimination methods--Comparison by means of +indices of modifiability--Number of tests per series versus number of +series--Efficiency as measured by memory tests. + + +CHAPTER XVI + +THE DURATION OF HABITS: MEMORY AND RE-LEARNING + +Measures of the permanency of modifications in behavior--The duration of +brightness and color discrimination habits--The relation of learning to +re-learning--Can a habit which has been lost completely be re-acquired +with greater facility than it was originally acquired?--Relation of +special training to general efficiency--Does the training in one form of +labyrinth aid the dancer in acquiring other labyrinth habits? + + +CHAPTER XVII + +INDIVIDUAL, AGE, AND SEX DIFFERENCES IN BEHAVIOR + +Individual peculiarities in sensitiveness, docility, and initiative--The +relation of docility to age--The individual result and the average--How +averages conceal facts--Sex differences in docility and initiative-- +Individual differences of motor capacity which seem to indicate +varieties--Is the dancer pathological? + + +CHAPTER XVIII + +THE INHERITANCE OF FORMS OF BEHAVIOR + +Characteristics of the race--Inheritance of the tendency to whirl in a +particular way--Tests of the inheritance of individually acquired forms of +behavior. + + +INDEX + + + + + + +ILLUSTRATIONS + +Dancing Mice--sniffing and eating _Frontispiece_ + +FIGURE + +1. Color patterns of dancers. Record blanks + +2. Double cage, with nest-boxes and water dishes + +3. Double cages in frame + +4. Photographs of dancers climbing (After Zoth) + +5. Tracks of common mouse (After Alexander and Kreidl) + +6. Tracks of dancer (After Alexander and Kreidl) + +7. The inner ear of the rabbit (Retzius) + +8. The membranous labyrinth of the ear of the dancer (After Rawitz) + +9. Same + +10. Same + +it. Model of the ear of the dancer (After Baginsky) + +12. Ear of the dancer (After Kishi) + +13. Ear of the dancer (After Kishi) + +14. Discrimination box + +15. Ground plan of discrimination box + +16. Nendel's gray papers + +17. Weber's law apparatus + +18. Food-box apparatus + +19. Waugh's food-box apparatus + +20. Color discrimination apparatus + +21. Ground plan of color discrimination apparatus + +22. Cards for form discrimination + +23. Labyrinth B + +24. Labyrinth B on electric wires + +25. Labyrinth A + +26. Curves of habit formation for labyrinth B + +27. Plan of labyrinth C, and path records + +28. Labyrinth D + +29. Curve of learning for white-black discrimination, twenty individuals + +30. Curve of learning for white-black discrimination, thirty individuals + +31. Curve of habit formation for labyrinth D + +32. Curves of learning and re-learning + +33. Plasticity curves + + + + + +LITERATURE ON THE DANCING MOUSE + +1. ALEXANDER, G. UND KREIDL, A. "Zur Physiologie des Labyrinths der +Tanzmaus." _Archiv für die gesammte Physiologie_, Bd. 82: 541-552. +1900. + +2. ALEXANDER, G. UND KREIDL, A. "Anatomisch-physiologische Studien über +das Ohrlabyrinth der Tanzmaus." II Mittheilung. _Archiv für die gesammte +Physiologie_. Bd. 88: 509-563. 1902. + +3. ALEXANDER, G. UND KREIDL, A. "Anatomisch-physiologische Studien über +das Ohrlabyrinth der Tanzmaus." III Mittheilung. _Archiv für die +gesammte Physiologie_, Bd. 88: 564-574. 1902. + +4. BAGINSKY, B. "Zur Frage über die Zahl der Bogengänge bei japanischen +Tanzmäusen." _Centralblatt für Physiologie_, Bd. 16: 2-4. 1902. + +5. BATESON, W. "The present state of knowledge of colour-heredity in mice +and rats." _Proceedings of the Zoölogical Society of London_, Vol. 2: +71-99. 1903. + +6. BREHM, A. E. "Tierleben." Dritte Auflage. Saugetiere, Bd. 2: 513-514. +1890. + +7. BREHM, A. E. "Life of Animals." Translated from the third German +edition of the "Tierleben" by G. R. Schmidtlein. Mammalia, p. 338. +Marquis, Chicago. 1895. + +8. CYON, E. DE. "Le sens de l'espace chez les souris dansantes +japonaises." _Cinquantenaire de la Société de Biologie_ (Volume +jubilaire). p. 544-546. Paris. 1899. + +9. CYON, E. VON. "Ohrlabyrinth, Raumsinn und Orientirung." _Archiv für +die gesammte Physiologie_, Bd. 79: 211-302. 1900. + +10. CYON, E. DE. "Presentation de souris dansantes japonaises." _Comptes +rendus du XIII Congrès International de Paris, Section de +physiologie_, p. 160-161. 1900. + +11. CYON, E. VON. "Beiträge zur Physiologie des Raumsinns." I Theil. "Neue +Beobachtungen an den japanischen Tanzmäusen." _Archiv für die gesammte +Physiologie_, Bd. 89: 427-453. 1902. + +12. CYON, E. DE. "Le sens de l'espace." Richet's "Dictionnaire de +physiologie," T. 5: 570-571. 1901. + +13. DARBISHIRE, A. D. Note on the results of crossing Japanese waltzing +mice with European albino races. _Biometrica_, Vol. 2: 101-104. 1902. + +14. DARBISHIRE, A. D. Second report on the result of crossing Japanese +waltzing mice with European albino races. _Biometrica_, Vol.2: +165-173. 1903. + +15. DARBISHIRE, A. D. Third report on hybrids between waltzing mice and +albino races. _Biometrica_, Vol. 2: 282-285. 1903. + +16. DARBISHIRE, A. D. On the result of crossing Japanese waltzing with +albino mice. _Biometrica_, Vol 3: 1-51. 1904. + +17. GUAITA, G. v. "Versuche mit Kreuzungen von verschiedenen Rassen der +Hausmaus." _Berichte der naturforschenden Gesellschaft zu Freiburg i. +B_., Bd. 10: 317-332. 1898. + +18. GUAITA, G. v. "Zweite Mitteilung uber Versuche mit Kreuzungen von +verschiedenen Hausmausrassen." _Berichte der naturforschenden +Gesellschaft zu Freiburg i. B_., Bd. 11: 131-138. 1900. + +19. HAACKE, W. "Ueber Wesen, Ursachen und Vererbung von Albinismus und +Scheckung und über deren Bedeutung für vererbungstheoretische und +entwicklungsmechanische Fragen." _Biologisches Centralblatt_, Bd. +15: 44-78. 1895. + +19a. HUNTER, M. S. "A Pair of Waltzing Mice." _The Century Magazine_, +Vol. 73: 889-893. April, 1907. + +20. KAMMERER, P. "Tanzende Waldmaus und radschlagende Hausmaus." +_Zoölogische Garten_, Bd. 41: 389-390. 1900. + +21. KISHI, K. "Das Gehörorgan der sogenannten Tanzmaus." _Zeitschrift +für wissenschaftliche Zoölogie_, Bd. 71: 457-485. 1902. + +22. LANDOIS, H. "Chinesische Tanzmäuse." _Jahresbericht des Westfähschen +Provinzial-Vereins_, Munster, 1893-1894: 62-64. + +22a. LOSE, J. "Waltzing Mice." _Country Life in America_, September, +1904. p. 447. + +23. PANSE, R. Zu Herrn Bernhard Rawitz' Arbeit: "Das Gehörorgan der +japanischen Tanzmäuse." _Archiv für Anatomie und Physiologie_, +Physiologische Abtheilung, 1901: 139-140. + +24. PANSE, R. "Das Gleichgewichts- und Gehörorgan der japanischen +Tanzmäuse." _Münchener medicinische Wochenschrift_, Jahrgang 48, Bd. +I: 498-499. 1901. + +25. RAWITZ, B. "Das Gehörorgan der japanischen Tanzmäuse." _Archiv für +Anatomie und Physiologie_, Physiologische Abtheilung, 1899: 236-243. + +26. RAWITZ, B. "Neue Beobachtungen über das Gehörorgan japanischer +Tanzmäuse." _Archiv für Anatomie und Physiologie_, Physiologische +Abtheilung, 1901, Supplement: 171-176. + +27. RAWITZ, B. "Zur Frage über die Zahl der Bogengänge bei japanischen +Tanzmäusen." _Centralblatt für Physiologie_, Bd. 15: 649-651. 1902. + +28. SAINT-LOUP, R. "Sur le mouvement de manège chez les souris." +_Bulletin de la Société Zoölogique de France_, T. 18: 85-88. 1893. + +29. SCHLUMBERGER, C. "A propos d'un netzukè japonais." _Memoires de la +Société Zoölogique de France_, T. 7: 63-64. 1894. + +30. WELDON, W. F. R. Mr. Bateson's revisions of Mendel's theory of +heredity. _Biometrica_, Vol. 2: 286-298. 1903. + +31. ZOTH, O. "Ein Beitrag zu den Beobachtungen und Versuchen an +japanischen Tanzmäusen." _Archiv für die gesammte Physiologie_, Bd. +86: 147-176. 1901. + +32. ANONYMOUS. "Fancy Mice: Their Varieties, Management, and Breeding." +Fourth edition. London: L. Upcott Gill. No date. + + + + + + + + +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. + + + + +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. + + + + +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. + + + + +CHAPTER V + + +STRUCTURAL PECULIARITIES AND BEHAVIOR + +The activities of an animal are expressions of changes which occur in its +structure, and they can be explained satisfactorily only when the facts of +structure are known. Such peculiarities of activity as are exhibited by +the dancing mouse, as contrasted with the common mouse, suggest at once +that this creature has a body which differs in important respects from +that of the ordinary mouse. In this chapter I shall present what is known +concerning the structural bases for the whirling, the lack of +equilibrational ability and of dizziness, the quick jerky head movements, +the restlessness, and the partial or total deafness of the dancing mouse. + +Comparative physiologists have discovered that the ability of animals to +regulate the position of the body with respect to external objects and to +respond to sounds is dependent in large measure upon the groups of sense +organs which collectively are called the ear. Hence, with reason, +investigators who sought structural facts with which to explain the forms +of behavior characteristic of the dancer turned their attention first of +all to the study of the ear. But the ear of the animal is not, as might be +supposed on superficial examination, a perfectly satisfactory natural +experiment on the functions of this group of sensory structures, for it is +extremely uncertain whether any one of the usual functions of the organ is +totally lacking. Dizziness may be lacking, and in the adult hearing also, +but in general the functional facts lead the investigator to expect +modifications of the sense organs rather than their absence. + +I shall now give an account of the results of studies concerning the +structure of the ear and brain of the dancer. Since the descriptions given +by different anatomists contradict one another in many important points, +the several investigations which have been made may best be considered +chronologically. + +Bernhard Rawitz (25 p. 239) was the first investigator to describe the +structure of the ear of the Japanese or Chinese dancers, as he calls them. +The definite problem which he proposed to himself at the beginning of his +study was, what is the structural basis of the whirling movement and of +the deafness of the mice? + +In his first paper Rawitz described the form of the ears of five dancers. +His method of work was to make microscopic preparations of the ears, and +from the sections, by the use of the Born method, to reconstruct the ear +in wax. These wax models were then drawn for the illustration of the +author's papers (Figures 8, 9, 10). + +The principal results of the early work of Rawitz are summed up in the +following quotation from his paper: "The Japanese dancing mice have only +one normal canal and that is the anterior vertical. The horizontal and +posterior vertical canals are crippled, and frequently they are grown +together. The utriculus is a warped, irregular bag, whose sections have +become unrecognizable. The utriculus and sacculus are in wide-open +communication with each other and have almost become one. The utriculus +opens broadly into the scala tympani, and the nervous elements of the +cochlea are degenerate. + +"The last-mentioned degeneration explains the deafness of the dancing +mice; but in my opinion it is a change of secondary nature. The primary +change is the broad opening between the utriculus and the scala tympani +from which results the streaming of the endolymph from the semicircular +canals into the cochlea. When, as a consequence of the rapid whirling +movements, a great part of the endolymph is hurled into the scala tympani, +the organ of Corti in the scala vestibuli is fixed and its parts are +rendered incapable of vibration. The condition of atrophy which is +observable in the sense cells and in the nerve elements is probably due to +the impossibility of functional activity; it is an atrophy caused by +disuse "(25 p. 242). + + +Ampulla externa +Ampulla anterior +Ramus utriculi + +Membrana basilaris + +Lagena + +Canalis utriculo-saccularis + +Membrana basilaris +Ampulla posterior +Macula acustica sacculi + +[Illustration: FIGURE 7.--The inner ear of the rabbit. Reproduced from +Selenka after Retzius.] + + +To render the terms which occur in this and subsequent descriptions of the +ear of the dancer somewhat more intelligible to those who are not familiar +with the general anatomy of the vertebrate ear, a side view of the inner +ear of the rabbit is reproduced from a drawing by Retzius (Figure 7). I +have chosen the ear of the rabbit for this purpose, not in preference to +that of the common mouse, but simply because I failed to find any reliable +description of the latter with drawings which could be reproduced. The +rabbit's ear, however, is sufficiently like that of the mouse to make it +perfectly satisfactory for our present purpose. + +This drawing of the rabbit's ear represents the three semicircular canals, +which occur in the ear of all mammals, and which are called, by reason of +their positions, the anterior vertical, the posterior vertical, and the +horizontal. Each of these membranous canals possesses at one end, in an +enlargement called the ampulla, a group of sense cells. In Figure 7 the +ampullae of the three canals are marked respectively, ampulla anterior, +ampulla posterior, and ampulla externa. This figure shows also the +cochlea, marked lagena, in which the organ of hearing of mammals (the +organ of Corti) is located. The ear sac, of which the chief divisions are +the utriculus and the sacculus, with which the canals communicate, is not +shown well in this drawing. + +Within a few months after the publication of Rawitz's first paper on the +structure of the dancer's ear, another European investigator, Panse (23 +and 24) published a short paper in which he claimed that previous to the +appearance of Rawitz's paper he had sectioned and mounted ears of the +common white mouse and the dancing mouse side by side, and, as the result +of careful comparison, found such slight differences in structure that he +considered them unworthy of mention. Panse, therefore, directly +contradicts the statements made by Rawitz. In fact, he goes so far as to +say that he found even greater differences between the ears of different +white mice than between them and the ears of the dancer (23 p. 140). + +In a somewhat later paper Panse (24 p. 498) expresses his belief that, +since there are no peculiarities in the general form, sensory structures, +or nerve supply of the ear of the dancer, which serve to explain the +behavior of the animal, it is probable that there are unusual structural +conditions in the brain, perhaps in the cerebellum, to which are due the +dance movements and the deafness. The work of Panse is not very +convincing, however, for his figures are poor and his descriptions meager; +nevertheless, it casts a certain amount of doubt upon the reliability of +the descriptions given by Rawitz. + +[Illustration: FIGURE 8.--The membranous labyrinth of the dancer's ear. +Type I. This figure, as well as 9 and 10, are reproduced from Rawitz's +figures in the _Archiv für Anatomie und Physiologie, Physiologische +Abtheilung_, 1899. _C.s._, anterior vertical canal; _C.p._, posterior +vertical canal; _C.e._, horizontal canal; _U._, utriculus.] + +The unfavorable light in which his report was placed by Panse's statements +led Rawitz to examine additional preparations of the ear of the dancer. +Again he used the reconstruction method. The mice whose ears he studied +were sent to him by the physiologist Cyon. + +As has been noted in Chapter IV, Cyon discovered certain differences in +the structure and in the behavior of these dancers (11 p. 431), which led +him to classify them in two groups. The individuals of one group climbed +readily on the vertical walls of their cages and responded vigorously to +sounds; those of the other group could not climb at all and gave no +evidences of hearing. After he had completed his study of their behavior, +Cyon killed the mice and sent their heads to Rawitz; but unfortunately +those of the two groups became mixed, and Rawitz was unable to distinguish +them. When he examined the structure of the ears of these mice, Rawitz did +find, according to his accounts, two structural types between which very +marked differences existed. Were it not for the carelessness which is +indicated by the confusion of the materials, and the influence of Cyon's +suggestion that there should be different structures to account for the +differences in behavior, Rawitz's statements might be accepted. As matters +stand there can be no doubt of individual differences in behavior, +external appearance, and the structure of the ear; but until these have +been correlated on the basis of thoroughgoing, careful observation, it is +scarcely worth while to discuss their relations. + +[Illustration: FIGURE 9.--The membranous labyrinth of the dancer's ear. +Type II.] + +[Illustration: FIGURE 10.--The membranous labyrinth of the dancer's ear. +Type III.] + +To his previous description of the conditions of the ear sacs, sense +organs, and nerve elements of the dancer's ear, Rawitz adds nothing of +importance in his second paper (26 p. 171). He merely reiterates his +previous statements concerning the form of the canals, on the basis of his +findings in the case of six additional dancers. Figures 8, 9, and 10 are +reproduced from Rawitz to show the anatomical conditions which he claims +that he found. As these figures indicate, the canals were found to be +extremely variable, as well as unusual in form, and the sacs distorted. In +the ears of some specimens there were only two canals, and in all cases +they were more or less reduced in size, distorted, or grown together. + +[Illustration: FIGURE 11.--Photograph of a wax model of the membranous +labyrinth of the ear of the dancer. Reproduced from Baginsky's figure in +the _Centralblatt für Physiologie_, Bd. 16.] + +The work of Rawitz was unfavorably criticised by Alexander and Kreidl (2), +Kishi (21), and Baginsky (4), as well as by Panse (23 and 24). To their +criticisms Rawitz replied by insisting that the other investigators could +not with right attack his statements because they had not used the +reconstruction method. In order to test the value of this contention, and +if possible settle the question of fact, Baginsky had a model of the ear +of the dancer constructed by a skilled preparator (Herr Spitz) from +sections which had been prepared by the best neurological methods. This +model was made eighty times the size of the ear. It was then reduced in +the process of photographic reproduction to sixteen times the natural size +of the ear in the mouse. Figure 11 is a photograph of Baginsky's model. It +shows beyond question the presence of three canals of the same general +form and relations as those of the common mouse and of other mammals. +Baginsky's paper is brief and to the point. His criticisms of the work of +both Cyon and Rawitz are severe, but they are justified in all probability +by the carelessness of these investigators in the fixation of their +materials. Of the five skilled histologists who have examined the ear of +the dancer, Rawitz alone found markedly abnormal canals. It is highly +probable, therefore, that the canals in his preparations in some way +became distorted before the ears were sectioned. He doubtless described +accurately the conditions which he found, but the chances are that those +conditions never existed in the living animals. + +The conflicting statements of Rawitz and Panse stimulated interest, and as +a result two other investigators, without knowledge of one another's work, +began careful researches on the dancer's ear. One, Alexander (2 and 3), +worked in coöperation with the physiologist Kreidl; the other, Kishi (21), +worked independently. The anatomical papers of Alexander and Kishi +appeared at about the same time, and since neither contains a reference to +the other, it is evident that the investigations were carried on almost +simultaneously. Alexander's descriptions are more detailed than those of +Rawitz and Panse, and in certain respects Kishi's are even more +thoroughgoing. The first paper published by Alexander and Kreidl (1) +contains the results of observations on the habits and behavior of the +dancers. Having examined the chief facts of function, these investigators +attempted to discover the structural conditions for the peculiarities of +behavior which they had observed. + +As material for their anatomical work they made use of four dancers, one +albino mouse, and four common gray mice. The ears of these individuals +were fixed, sectioned, and examined microscopically in connection with +parts of the brain. In all, eight dancer ears and six common mouse ears +were studied. + +Very extensive descriptions of these preparations, together with +measurements of many important portions of the ear, are presented in their +paper, the chief conclusions of which are the following:-- + +1. The semicircular canals, the ampullae, the utriculus, and the cristae +acusticae of the canals are normal in their general form and relations to +one another as well as in their histological conditions (2 p. 529). This +is contradictory of the statements made by Rawitz. + +2. There is destruction of the macula sacculi (2 p. 534). + +3. There is destruction also of the papilla basilaris cochleae, with +encroachment of the surrounding tissues in varying degrees. + +4. There is diminution in the number of fibers of the branches and roots +of the ramus superior and ramus medius of the eighth nerve, and the fiber +bundles are very loosely bound together. + +5. Similarly the number of fibers in the inferior branch (the cochlear +nerve) of the eighth nerve is very much reduced. + +6. There is moderate reduction in the size of the two vestibular ganglia +as a result of the unusually small number of nerve cells. + +7. The ganglion spirale is extremely degenerate. + +There is therefore atrophy of the branches, ganglia, and roots of the +entire eighth nerve, together with atrophy and degeneration of the pars +inferior labyrinthii. The nerve endings are especially degenerate (2 p. +534). + +The above structural deviations of the ear of the dancer from that of the +common mouse may be considered as primary or secondary according as they +are inherited or acquired. Since, according to Alexander and Kreidl, the +dancers' peculiarities of behavior and deafness are directly and uniformly +inherited, it is obvious that certain primary structural deviations must +serve as a basis for these functional facts. But it is equally clear, in +the opinion of Alexander and Kreidl (2 p. 536), that other structural +peculiarities of the dancer are the result of the primary changes, and in +no way the conditions for either the dancing or the deafness. These +authors feel confident that the facts of behavior which are to be +accounted for are almost certainly due to the pathological changes which +they have discovered in the nerves, ganglia, and especially in the +peripheral nerve endings of the ear of the mouse (2 p. 537). + +It is further claimed by Alexander and Kreidl that there are very marked +individual differences among the dancers in the structure of the ear. In +some cases the otoliths and the sensory hairs are lacking; in others, they +are present in the state of development in which they are found in other +varieties of mouse. Sometimes the cochlea is much reduced in size; at +other times it is found to be of normal size (2 p. 538). These variations +in structure, if they really exist, go far toward justifying the tendency +of Cyon and Alexander and Kreidl, as well as many other investigators, to +regard the dancer as abnormal or even pathological. + +The functions of the ear as at present known to the comparative +physiologist are grouped as the acoustic and the non-acoustic. The cochlea +is supposed on very good grounds to have to do with the acoustic +functions, and the organs of the semicircular canals on equally good +evidence are thought to have to do with such of the non-acoustic functions +as equilibration and orientation. Just what the functions of the organs of +the ear sacs are is not certainly known. These facts are of importance +when we consider the attempts made by Alexander and Kreidl to correlate +the various peculiarities of behavior shown by the dancer with the +structural facts which their work has revealed. This correlation is +indicated schematically below. The physiological facts to be accounted for +in terms of structure are presented in the first column, and the +anatomical facts which are thought to be explanatory, in the second (2 p. +539). + + +FUNCTION + +1 Lack of sensitiveness to auditory stimuli. {Structure 1,2,3 below} + +2 Defective equilibrational ability. {Structure 4,5,6 below} + +3 Lack of turning dizziness. {Structure 4,5,6 below} + +4 Normal reactions to galvanic stimulation. (not related in table to any +Structure) + + +STRUCTURE + +1 Destruction of the papilla basilaris cochleae, etc. + +2 Diminution of the inferior branch of the eighth nerve. + +3 Marked degeneration of the ganglion spirale. + +4 Destruction of the macula sacculi. + +5 Diminution of the branches and roots of the superior and middle branches +of the eighth nerve. + +6 Diminution of both ganglia vestibulii and of the nerve cells. + + +Alexander and Kreidl themselves believe that the partial deafness of the +dancers (for they admit that the total lack of hearing has not been +satisfactorily proved) is due to the defective condition of the cochlea. +They account for the imperfect equilibrational ability of the animals by +pointing out the structural peculiarities of the sacculus, the vestibular +ganglia, and the peripheral nerves. Similarly, the lack of dizziness they +suppose to be due to the diminution of the fibers of the nerves which +supply the canal organs, the atrophied condition of the vestibular +ganglia, and a disturbance of the peripheral sense organs. Furthermore, +there are no anatomical facts which would indicate a lack of galvanic +dizziness (2 p. 552). + +Despite the fact that they seem to explain all the functional +peculiarities of the dancer, the statements made by Alexander and Kreidl +are neither satisfying nor convincing. Their statements concerning the +structure of the ear have not been verified by other investigators, and +their correlation of structural with functional facts lacks an +experimental basis. + +In this connection it may be worth while to mention that a beautiful +theory of space perception which Cyon (9) had constructed, largely on the +basis of the demonstration by Rawitz that the dancers have only one normal +canal, is totally destroyed by Panse, Baginsky, Alexander and Kreidl, and +Kishi, for all of these observers found in the dancer three canals of +normal shape. Cyon had noted that the most abnormal of the voluntary as +well as of the forced movements of the dancer occur in the plane of the +canal which Rawitz found to be most strikingly defective. This fact he +connected with his observation that the fish Petromyzon, which possesses +only two canals, moves in only two spatial dimensions. The dancer with +only one functional canal in each ear moves in only one plane, and neither +it nor Petromyzon is able to move far in a straight line (11 p. 444). From +these and similar surmises, which his eagerness to construct an ingenious +theory led him to accept as facts quite uncritically, Cyon concluded that +the perception of space depends upon the number and arrangement of the +semicircular canals, and that the dancer behaves as it does because it +possesses canals of unusual shape and relations to one another. The +absurdity of Cyon's position becomes obvious when it is shown that the +structural conditions of which he was making use do not exist in the +dancer. + +The results obtained by Kishi in his study of the ear of the dancer differ +in many important respects from those of all other investigators, but +especially from those of Rawitz and Alexander and Kreidl. + +Kishi's work was evidently done with admirable carefulness. His methods in +the preparation of his materials, so far as can be judged from his report, +were safe and satisfactory, and his descriptions of results are minute and +give evidence of accuracy and conscientious thoughtfulness. The material +for his histological work he obtained from three different animal dealers. +It consisted of fifteen adult and nineteen young dancers, and, as material +for comparison, ten common gray mice. The animals were studied first +biologically, that their habits and behavior might be described accurately +and so far as possible accounted for in the light of whatever histological +results might be obtained subsequently; then they were studied +physiologically, that the functional importance of various organs which +would naturally be supposed to have to do with the peculiarities of the +mouse might be understood; and, finally, they were killed and their ears +and portions of their brains were studied microscopically, that structural +conditions for the biological and physiological facts might be discovered. + +The ear, which was studied by the use of several series of sections, as +well as in gross dissections, is described by Kishi under three +headings:-- + +(1) The sound-receiving apparatus (auditory organs). + +(2) The static apparatus (equilibrational organs). + +(3) The sound-transmitting apparatus (ear drum, ear bones, etc.). + +The chief results of his structural investigation may be stated briefly +under these three headings. In the sound-receiving or auditory apparatus, +Kishi failed to find the important deviations from the usual structure of +the mammalian ear which had been described by Rawitz. The latter +distinctly says that although the organ of Corti is present in all of the +whirls of the cochlea, the auditory cells in it are noticeably degenerate. +Kishi does not agree with Panse's statement (21 p. 476) that the auditory +organ of the dancer differs in no important respects from that of the +common mouse, for he found that in certain regions the hair cells of the +organ of Corti were fewer and smaller in the dancer. He therefore +concludes that the auditory organ is not entirely normal, but at the same +time he emphasizes the serious discrepancy between his results and those +of Rawitz. In not one of the ears of the twelve dancers which he studied +did Kishi find the direct communication between the utriculus and the +scala tympani which Rawitz described, and such differences as appeared in +the organ of Corti were in the nature of slight deviations rather than +marked degenerations. + +In the outer wall of the ductus cochlearis of the dancer the stria +vasculosa is almost or totally lacking, while in the gray mouse it is +prominent. This condition of the stria vasculosa Kishi was the first to +notice in the dancer; Alexander and Kreidl had previously described a +similar condition in an albino cat. If, as has been supposed by some +physiologists, the stria vasculosa is really the source of the endolymph, +this state of affairs must have a marked influence on the functions of the +auditory apparatus and the static apparatus, for pressure differences +between the endolymph and the perilymph spaces must be present. And, as +Kishi points out, should such pressure differences be proved to exist, the +functional disturbance in the organ of hearing which the lack of responses +to sounds seems to indicate might better be ascribed to them than to the +streaming of the endolymph from the canals into the cochlea as assumed by +Rawitz (21 p. 477). Kishi merely suggests that the condition of the stria +may account for the deafness of the mouse; he does not feel at all +confident of the truth of his explanation, and he therefore promises in +his first paper a continuation of his work in an investigation of the +functions of the stria. This, however, he seems not to have accomplished +thus far. + +[Illustration: FIGURE 12.--The inner ear of the dancer. Reproduced from +Kishi's figure in the _Zeitschrift für wissenschaftliche Zoölogie_, Bd. +71. _c.c._ crus simplex; o.b. anterior vertical canal; _h.b._ posterior +vertical canal; _a.b._ horizontal canal.] + +The static apparatus Kishi describes as closely similar in form to that of +the gray mouse. In none of his twelve preparations of the ear of the +dancer did he find such abnormalities of form and connections in the +semicircular canals as Rawitz's figures and descriptions represent. Rawitz +states that the anterior canal is normal except in its lack of connection +with the posterior and that the posterior and horizontal are much reduced +in size. Kishi, on the contrary, insists that all of the three canals are +normal in shape and that the usual connection between the anterior and the +posterior canals, the crus simplex, exists. He justifies these statements +by presenting photographs of two dancer ears which he carefully removed +from the head. Comparison of these photographs (Figures 12 and 13) with +Rawitz's drawings of the conditions of the canals and sacs as he found +them (Figures 8, 9, and 10), and of both with the condition in the typical +mammalian ear as shown by Figure 7, will at once make clear the meaning of +Kishi's statements. That Rawitz's descriptions of the canals are not +correct is rendered almost certain by the fact that Panse, Baginsky, +Alexander and Kreidl, and Kishi all agree in describing them as normal in +form. + +The only important respects in which Kishi found the membranous labyrinth, +that is, the canals and the ear sacs, of the dancer to differ from that of +the gray mouse are the following. In the dancer the cupola of the crista +acustica is not so plainly marked and not so highly developed, and the +raphae of the ampullae and canals, which frequently are clearly visible in +the gray mouse, are lacking (21 p. 478). + +[Illustration: FIGURE 13.--The inner ear of the dancer, showing the spiral +form of the cochlea. After Kishi.] + +The sound-transmitting apparatus of the dancer, according to Kishi, +differs only very slightly from that of the gray mouse, and there is no +reason to consider the differences which appear as important (21 p. 478). + +Almost as amusing as the way in which Cyon's theory of space perception +disappears in the light of critical research is Panse's explanation of the +deafness of the dancer. Failing to find any defects in the auditory +apparatus of the inner ear which seemed adequate to account for the +obvious lack of responsiveness to sounds, this investigator concluded that +plugs of wax which he had noticed in the auditory meatus of the dancer +excluded sounds or in some way interfered with the functioning of the +tympanic membrane. Kishi reports that he found such plugs of wax in the +ears of one gray mouse, but in none of the dancers which he examined did +he discover them (21 p. 479). Panse's explanation of the defective hearing +of the dancer neither needs nor deserves further comment. + +As one result of his investigation, Kishi is convinced that the dance +movements are not due to peculiarities in the semicircular canals and +their sense organs, as Rawitz claimed, for the general form and finer +structure of these organs in the dancer is practically the same as in the +common mouse. Kishi is just as certain that the whirling is not due to +defects in the canal organs, as Rawitz is that it is due to such +structural conditions! It is rather surprising that any one should feel +confident of the power of the microscope to reveal all those structural +conditions which are important as conditions of function. Probably there +are histological differences between the ear of the dancer and that of the +gray mouse, which, although undetectable by scientific means at present, +furnish the structural basis for the marked differences in behavior. As +has been set forth already (p. 9), Kishi accounts for the dance movements +by assuming the inheritance of an acquired character of behavior. This +inherited tendency to dance, he thinks, has been accentuated by the +confinement of the mice in narrow cages and their long-continued movement +in the wheels which are placed in the cages (21 p. 481). + +Rawitz, Cyon, and Alexander and Kreidl felt themselves under the necessity +of finding peculiarities of behavior in the dancer which could be referred +to the various abnormalities of structure which they had either seen or +accepted on faith; Kishi found himself in a very different predicament, +for he had on his hands the commonly accepted statement that the animals +are deaf, without being able to find any structural basis for this defect. +To avoid the difficulty he questions the existence of deafness! If +perchance they are deaf, he thinks that it is possibly because of the +defect in the stria vasculosa. This suggestion Kishi makes despite the +fact that our ignorance of the function of the stria renders it impossible +for us to do otherwise than guess at its relation to hearing. + +We have now briefly reviewed the results of the various important +investigations of the behavior and structure of the dancer. + +The observations of Cyon, Zoth, and the writer establish beyond doubt the +existence of important individual differences in behavior if not of +distinct divisions within the species of mouse, and the general results of +the several anatomical investigations make it seem highly probable that +the structure of the ear, as well as the externally visible structural +features of the animals, vary widely. Unfortunately, the lack of agreement +in the descriptions of the ear given by the different students of the +subject renders impossible any certain correlation of structural and +functional facts. That the whirling and the lack of dizziness and of +hearing have their structural bases no one doubts, but whether it is in +the brain itself, in the sense organs, or in the labyrinth, our knowledge +does not permit us to say. With this statement Rawitz, Cyon, and Alexander +and Kreidl would not agree, for they believe that they have discovered +structural peculiarities which fully explain the behavior of the dancer. +Panse and Kishi, on the other hand, contend that the ear gives no +structural signs of such peculiarities as the dancing and deafness +suggest; they therefore look to the cerebellum for the seat of the +disturbance. With the same possibility in mind the author of "Fancy +Varieties of Mice" writes: "These quaint little creatures make amusing +pets for any one who is not scientific, or very fond of knowing 'the +reason why.' In their case, the reason of the peculiarity which gives them +their name is rather a sad one. It is now pretty conclusively established +that they are no more Japanese than they are of any other country in +particular, but that the originators of the breed were common fancy mice +which were suffering from a disease of the brain analogous to the 'gid' in +sheep. In the latter, the complaint is caused by a parasite in the brain; +in the case of the Waltzing Mouse, it is probably due to an hereditary +malformation therein. Be this as it may, the breed is now a firmly +established one, and the children of waltzing mice waltz like their +parents" (32 p. 45). Although it is quite possible that peculiarities in +the central nervous system, rather than in the peripheral nervous system, +may be responsible for the forms of behavior exhibited by the dancer, it +must be remembered that no such peculiarities have been revealed by the +examination of the central nervous system. The old fancier has neither +better nor worse grounds for his belief than have Panse and Kishi. + +So far as the reliability of the anatomical work which has been discussed +is in question, it would seem that Rawitz's results are rendered somewhat +unsatisfactory by the carelessness of Cyon in fixing the materials; that +Panse's descriptions and comparisons are neither careful nor detailed +enough to be convincing; that the work of Alexander and Kreidl, as well as +that of Kishi, gives evidence of accuracy and trustworthiness. The fact +that the statements of Alexander and Kreidl frequently do not agree with +those of Kishi proves that there are serious errors in the work of one or +another of these investigators. Cyon's discussion of the anatomy of the +dancer is not to be taken too seriously, for by his theory of space +perception and of a sixth sense he was unduly biased in favor of the +structural peculiarities described by Rawitz. Nevertheless, his discussion +is not without interest, for the way in which he succeeded in making every +structural fact which Rawitz suggested fit into his theories and help to +account for the functional peculiarities which he had himself observed, is +extremely clever and indicates a splendid scientific imagination. + +To sum up: All the facts of behavior and physiology which have been +established lead us to expect certain marked structural differences +between the dancer and the common mouse. The bizarre movements, lack of +equilibrational ability, and the nervous shaking of the head suggest the +presence of peculiar conditions in the semicircular canals or their sense +organs; and the lack of sensitiveness to sounds indicates defects in the +cochlea. Yet, strange as it may seem to those who are not familiar with +the difficulties of the study of the minute structure of these organs, no +structural conditions have been discovered which account satisfactorily +for the dancer's peculiarities of behavior. That the ear is unusual in +form is highly probable, since three of the four investigators who have +studied it carefully agree that it differs more or less markedly from that +of the common mouse. But, on the other hand, the serious lack of agreement +in their several descriptions of the conditions which they observed +renders their results utterly inconclusive and extremely unsatisfactory. +The status of our knowledge of the structure of the central nervous system +is even less satisfactory, if possible, than that of our knowledge of +those portions of the peripheral nervous system which would naturally be +supposed to have to do with such functional peculiarities as the dancer +exhibits. So far as I have been able to learn, no investigator has +carefully examined the brain and spinal cord in comparison with those of +the common mouse, and only those who have failed to find any structural +basis for the facts of behavior in the organs of the ear have attempted to +account for the dancer's whirling and deafness by assuming that the +cerebellum is unusual in structure. We are, therefore, forced to conclude +that our knowledge of the nervous system of the dancing mouse does not at +present enable us to explain the behavior of the animal. + +It seems highly probable to me, in the light of my observation of the +dancer and my study of the entire literature concerning the animal, that +no adequate explanation of its activities can be given in terms of the +structure of the peripheral or the central nervous system, or of both, but +that the structure of the entire organism will have to be taken into +account. The dancer's physiological characteristics, in fact, suggest +multitudinous structural peculiarities. I have confined my study to its +behavior, not because the problems of structure seemed less interesting or +less important, but simply because I found it necessary thus to limit the +field of research in order to accomplish what I wished within a limited +period. + +That there are structural bases for the forms of behavior which this book +describes is as certain as it could be were they definitely known; that +they, or at least some of them, are discoverable by means of our present- +day histological methods is almost as certain. It is, therefore, obvious +that this is an excellent field for further research. It is not an +agreeable task to report inconclusive and contradictory results, and I +have devoted this chapter to a brief account of the work that has been +done by others on the structure of the ear of the dancer rather for the +sake of presenting a complete account of the animal as it is known to-day +than because of the value of the facts which could be stated. + + + + +CHAPTER VI + +THE SENSE OF HEARING + +Repeatedly in the foregoing chapters mention has been made of the dancer's +irresponsiveness to sounds, but it has not been definitely stated whether +this peculiarity of behavior is due to deafness or to the inhibition of +reaction. This chapter is concerned with the evidence which bears upon the +problem of the existence of a sense of hearing. Again I may be permitted +to call attention to the observations of other investigators before +presenting the results of my own experiments and stating the conclusions +which I have reached through the consideration of all available facts. + +By the results of various simple tests which he made, Rawitz (25 p. 238) +was convinced that the adult dancer is totally deaf. He did not experiment +with the young, but he says he thinks they may be able to hear, since the +necessary structural conditions are present. This guess which Rawitz made +on the basis of very indefinite and uncertain knowledge of the histology +of the ear of the young dancer is of special interest in the light of +facts revealed by my own experiments. Unfortunately the study of hearing +made by Rawitz is casual rather than thorough, and although it may turn +out that all of his statements are justified by his observations, the +reader is not likely to get much satisfaction from his discussion of the +subject. + +Inasmuch as he could discover no structural basis for deafness, Panse (23 +p. 140) expressed himself as unwilling to believe that the mice are deaf, +and this despite the fact that he observed no responses to the sounds made +by a series of tuning forks ranging from C5 to C8. He believes rather that +they are strangely irresponsive to sounds and that their sensitiveness is +dulled, possibly, by the presence of plugs of wax in the ears. Since +another investigator, Kishi, has observed the presence of similar plugs of +wax in the ears of common mice which could hear, there is but slight +probability that Panse is right in considering the plugs of wax as the +cause of the dancer's irresponsiveness to sounds. + +Far more thoroughgoing tests than those of Rawitz or Panse were made by +Cyon (9 p. 218), who holds the unique position of being the only person on +record who has observed the adult dancer give definite reactions to +sounds. To a König Galton whistle so adjusted that it gave a tone of about +7000 complete vibrations per second, which is said to be about the pitch +of the voice of the dancer, some of the animals tested by Cyon responded +unmistakably, others not at all. In one group of four mice, two not only +reacted markedly to the sound of the whistle but apparently listened +intently, for as soon as the whistle was blown they ran to the side of the +cage and pressed their noses against the walls as if attempting to +approach the source of the stimulus. The remaining two mice gave not the +slightest indication that the sound acted as a stimulus. By the repetition +of this sound from eight to twelve times Cyon states that he was able to +arouse the mice from sleep. When thus disturbed, the female came out of +the nest box before the male. Similarly when the mice were disturbed by +the whistle in the midst of their dancing, the female was first to retreat +into the nest box. There is thus, according to Cyon, some indication of +sex, as well as individual, differences in sensitiveness to the sound of +the whistle. Cyon's statement that in order to evoke a response the +whistle must be held above the head of the dancer suggests at once the +possibility that currents of air or odors instead of sounds may have been +responsible for the reactions which he observed. The work of this +investigator justifies caution in the acceptance of his statements. +Neither the conditions under which the auditory tests were made nor the +condition of the animals is described with sufficient accuracy to make +possible the comparison of Cyon's work with that of other investigators. +As will appear later, it is of the utmost importance that the influence of +other stimuli than sound be avoided during the tests and that the age of +the mouse be known. The conclusion reached by Cyon is that some dancers +are able to hear sounds of about the pitch of their own cries. + +The fact, emphasized by Cyon, that the mice respond to tones of about the +pitch of their own voice is of peculiar interest in its relation to the +additional statements made by the same author to the effect that the +female dancer is more sensitive to sounds than the male, and that the +males either do not possess a voice or are much less sensitive to +disagreeable stimuli than the females. In the case of the dancers which he +first studied (9 p. 218), Cyon observed that certain strong stimuli evoked +pain cries; but later in his investigation he noticed that four +individuals, all of which were males, never responded thus to disagreeable +stimulation (11 p.431). He asks, therefore, does this mean that the males +lack a voice or that they are less sensitive than the females? The fact +that he did not succeed in getting a definite answer to this simple +question is indicative of the character of Cyon's work. My dancers have +provided me with ample evidence concerning the matter. Both the males and +the females, among the dancers which I have studied, possess a voice. + +The females, especially during periods of sexual excitement, are much more +likely to squeak than the males. At such times they give their shrill cry +whenever they are touched by another mouse or by the human hand. A slight +pinching of the tail will frequently cause the female to squeak, but the +male seldom responds to the same stimulus by crying out. The most +satisfactory way to demonstrate the existence of a voice in the male is to +subject him to the stimulating effect of an induced current, so weak that +it is barely appreciable to the human hand. To this unexpected stimulus +even the male usually responds by a sudden squeak. There can be no doubt, +then, of the possession of a voice by both males and females. The males +may be either less sensitive or less given to vocal expression, but they +are quite able to squeak when favorable conditions are presented. The +possession of a voice by an animal is presumptive evidence in favor of a +sense of hearing, but it would scarcely be safe to say that the mice must +be able to hear their own voices. Cyon, however, thinks that some dancers +can. What further evidence is to be had? + +Although they obtained no visible motor reactions to such noises as the +clapping of the hands, the snapping of the fingers, or to the tones of +tuning forks of different pitches and the shrill tones of the Galton +whistle, Alexander and Kreidl (1 p. 547) are not convinced of the total +deafness of the dancer, for, as they remark, common mice which undoubtedly +hear do not invariably respond visibly to sounds. Furthermore, the +anatomical conditions revealed by their investigation of the ear of the +dancer are not such as to render sensitiveness to sounds impossible. They +recognize also that the existence of the ability to produce sounds is an +indication of hearing. They have no confidence in the results reported by +Cyon, for they feel that he did not take adequate precautions to guard +against the action of other than auditory stimuli. + +Zoth (31 p. 170) has pointed out with reason and force that testing the +sensitiveness of the mice is especially difficult because of their +restlessness. They are almost constantly executing quick, jerky movements, +starting, stopping, or changing the direction of movement, and it is +therefore extremely difficult to tell with even a fair degree of certainty +whether a given movement which occurs simultaneously with a sound is a +response to the sound or merely coincident with it. With great care in the +exclusion of the influence of extraneous stimuli, Zoth tried a large +number of experiments to test the hearing of both young and adult dancers. +Not once did he observe an indubitable auditory reaction. As he says, "I +have performed numerous experiments with the Galton whistle, with a +squeaking glass stopper, with caps and cartridges, without being able to +come to any certain conclusion. With reference to the Galton whistle and +particularly to the tone which was said to have been heard extremely well +by Cyon's mice, I believe I am rather safe in asserting that my mice, +young (12-13 days) as well as old, do not react to the König Galton +whistle (7210 Vs.). They could not be awakened out of sleep by repetitions +of the sound, nor enticed out of their nests, and their dancing could not +be interrupted" (31 p. 170). Zoth's experiments appear to be the most +careful and critical of those thus far considered. + +Last to be mentioned, but in many respects of greatest interest and value, +is the work of Kishi (21 p. 482) on the problem of hearing. To this acute +observer belongs the credit of calling attention emphatically to the ear +movements which are exhibited by the dancer. Frequently, as he remarks, +the ears move as if the animal were listening or trying to determine the +direction whence comes a sound, yet usually the mouse gives no other sign +of hearing. That the absence of ordinary reactions to sounds is due to +deafness, Kishi, like Panse, is led to doubt because his anatomical +studies have not revealed any defects in the organs of hearing which would +seem to indicate the lack of this sense. + +This historical survey of the problem of hearing has brought out a few +important facts. No one of the several investigators of the subject, with +the exception of Cyon, is certain that the dancer can hear, and no one of +them, with the exception of Rawitz, is certain that it cannot hear! Cyon +almost certainly observed two kinds of dancing mice. Those of his dancers +which exhibited exceptional ability to climb in the vertical direction and +which also gave good evidence of hearing certain sounds may have been +hybrids resulting from the crossing of the dancer with a common mouse, or +they may have been exceptional specimens of the true dancer variety. A +third possibility is suggested by Rawitz's belief in the ability of the +young dancer to hear. Cyon's positive results may have been obtained with +immature individuals. I am strongly inclined to believe that Cyon did +observe two types of dancer, and to accept his statement that some of the +mice could hear, whereas others could not. It is evident, in the light of +our examination of the experimental results thus far obtained by other +investigators, that neither the total lack of sensitiveness to sounds in +the adult nor the presence of such sensitiveness in the young dancer has +been satisfactorily proved. + +I shall now report in detail the results of my own study of the sense of +hearing in the dancer. As the behavior of the young differs greatly from +that of the adult, by which is meant the sexually mature animal, I shall +present first the results of my experiments with adults and later, in +contrast, the results obtained with mice from one to twenty-eight days +old. + +My preliminary tests were made with noises. While carefully guarding +against the interference of visual, tactual, temperature, and olfactory +stimuli, I produced noises of varying degrees of loudness by clapping the +hands together suddenly, by shouting, whistling, exploding pistol caps, +striking steel bars, ringing an electric bell, and causing another mouse +to squeak. To these sounds a common mouse usually responds either by +starting violently, or by trembling and remaining perfectly quiet for a +few seconds, as if frightened. The adult dancers which I have tested, and +I have repeated the experiment scores of times during the last three years +with more than a hundred different individuals, have never given +unmistakable evidence of hearing. Either they are totally deaf or there is +a most surprising lack of motor reactions. + +Precisely the same results were obtained in tests made with the Galton +whistle throughout its range of pitches, and with Appuun whistles which, +according to their markings, ranged from 2000 Vs. (C_4) to 48,000 (G_9), +but which undoubtedly did not correspond at all exactly to this range, and +with a series of König tuning forks which gave tones varying in pitch from +1024 to 16,382 complete vibrations. + +I am willing to trust these experimental results the more fully because +during all the time I have had adult dancers under observation I have +never once seen a reaction which could with any fair degree of certainty +be referred to an auditory stimulus. Never once, although I have tried +repeatedly, have I succeeded in arousing a dancer from sleep by producing +noises or tones, nor have I ever been able to observe any influence of +sounds on the dance movements. All of Cyon's signs have failed with my +mice. Occasionally what looked like a response to some sound appeared, but +critical observation invariably proved it to be due to some other cause +than the auditory stimulus. A sound produced above the animal is very +likely to bring about a motor reaction, as Cyon claims; but I have always +found it to be the result of the currents of air or odors, which usually +influence the animal when the experimenter is holding any object above it. +I do not wish to maintain that Cyon's conclusions are false; I merely +emphasize the necessity for care in the exclusion of other stimuli. The +mice are extremely sensitive to changes in temperature, such, for example, +as are produced by the breath of the experimenter, and one must constantly +guard against the misinterpretation of behavior. + +In a single experiment with mice over a month old, I observed what might +possibly indicate sensitiveness to sound. While holding a mouse, thirty- +five days old, in my hand I pursed my lips and made a very shrill sound by +drawing in air; the mouse seemed to start perceptibly according to the +indications given by my sense of touch. I repeated the stimulus several +times and each time I could see and feel the animal start slightly. With +two other individuals which I tested the reaction was less certain, and +with several others I failed to get any indication of response. This would +seem to prove that the three individuals which responded happened to be +sensitive to that particular tone at the age of five weeks. The test is +unsatisfactory because the vibrations from my own body may have brought +about the reaction instead of the air vibrations produced by my lips, and +I therefore merely mention it in the enumeration of the various +experimental tests which I have made. + +If we should conclude from all the negative evidence that is available, or +that could be obtained, that the dancer is totally deaf, it might fairly +be objected that the conclusion is unsafe, since an animal does not +necessarily respond to stimuli by a visible change in the position or +relations of its body. Death feigning may fairly be considered a response +to a stimulus or stimulus complex, yet there may be no sign of movement. +The green frog when observed in the laboratory usually gives no indication +whatever, by movements that are readily observable, that it hears sounds +which occur about it, but I have been able to show by means of indirect +methods of study that it is stimulated by these same sounds.[1] Its rate +of respiration is changed by the sounds, and although a sound does not +bring about a bodily movement, it does very noticeably influence movements +in response to other stimuli which occur simultaneously with the sound. I +discovered that under certain rather simple experimental conditions the +green frog would regularly respond to a touch on the back by drawing its +hind leg up toward the body. Under the same conditions the sound of an +electric bell caused no visible movement of the leg, but if at the instant +the back was touched the bell was rung, the leg movement was much greater +than that brought about by the touch alone. This suggests at once the +desirability of studying the sense of hearing in the dancer by some +indirect method. The animal may be stimulated, and yet it may not give any +visible sign of the influence of the auditory stimulus. + +[Footnote 1: "The Sense of Hearing in Frogs." _Journal of Comparative +Neurology and Psychology_, Vol. XV, p. 288, 1905.] + +Were not the dancing so extremely variable in rapidity and duration, it +might be used as an index of the influence of auditory stimuli. Cyon's +statements would indicate that sounds interfere with the dancing, but as I +obtained no evidence of this, I worked instead with the following indirect +method, which may be called the method of auditory choice. + +The apparatus which was used is described in detail in Chapter VII. +Figures 14 and 15 will greatly aid the reader in understanding its +essential features. Two small wooden boxes, identical in form and as +closely similar as possible in general appearance, were placed in a larger +box in such positions that a mouse was forced to enter and pass through +one of them in order to get to the nest-box. On the bottom of each of +these small boxes was a series of wires through which an electric current +could be made to pass at the will of the experimenter. The boxes could +readily be interchanged in position. At one side of the large wooden box +and beyond the range of vision of the mouse was an electric bell which +could be caused to ring whenever the mouse approached the entrance to one +of the small boxes. The point of the experiment was to determine whether +the dancer could learn to avoid the box-which-rang when it was approached. +The method of conducting the tests was as follows. Each day at a certain +hour the mouse was placed in that part of the large box whence it could +escape to the nest-box only by passing through one of the small boxes. If +it approached the wrong box (whether it happened to be the one on the +right or the one on the left depended upon the experimenter's decision), +the bell began to ring as a warning against entering; if it approached the +other box, all was silent. As motives for the choice of the box-which-did- +not-ring both reward and punishment were employed. The reward consisted of +freedom to return to the nest-box _via_ the passage which led from the +box-which-did-not-ring; the punishment, which consisted of a disagreeable +electric shock, was given whenever the mouse entered the wrong box, that +is, the one which had the sound as a warning. Entering the wrong box +resulted in a disagreeable stimulus and in the necessity of returning to +the large box, for the exit to the nest-box by way of the passage from +this box was closed. My assumption, on the basis of extended study of the +ability of the dancer to profit by experience, was that if it could hear +the sound of the bell it would soon learn to avoid the box-that-rang and +enter instead the one which had no sound associated with it. + +Systematic tests were made with No. 4 from the 3d to the 12th of February, +inclusive, 1906. Each day the mouse was permitted to find his way to the +nest-box through one of the small boxes ten times in succession. Usually +the experimenter rang the bell alternately for the box on the left and the +box on the right. The time required for such a series of experiments +varied, according to the rapidity with which the mouse made his choice, +from ten to thirty minutes. If in these experiments the animal approached +and entered the right, or soundless box, directly, the choice indicated +nothing so far as ability to hear is concerned; if it entered the wrong, +or sounding box, despite the ringing of the bell, it indicated either the +lack of the influence of experience or inability to hear the sound; but if +it regularly avoided the box-which-sounded it thus gave evidence of +ability to hear the sound of the bell. The purpose of the test was to +determine, not whether the mouse could learn, but whether it could hear. + +For ten successive days this experiment was carried on with No. 4 without +the least indication of increasing ability to avoid the wrong box by the +association of the sound of the bell with the disagreeable electric shock +and failure to escape to the nest-box. In fact, the experiment was +discontinued because it became evident that an impossible task had been +set for the mouse. Day by day as the tests were in progress I noticed that +the animal became increasingly afraid of the entrances to the small boxes; +it seemed absolutely helpless in the face of the situation. Partly because +of the definiteness of the negative results obtained with No. 4 and partly +because of the cruelty of subjecting an animal to disagreeable conditions +which it is unable to avoid, the experiment was not repeated with other +individuals. I have never conducted an experiment which gave me as much +discomfort as this; it was like being set to whip a deaf child because it +did not learn to respond to stimuli which it could not feel. + +By a very similar method No. 18 was tested for his sensitiveness to the +noise and jar from the induction apparatus which was used in connection +with many of my experiments on vision and the modifiability of behavior. +In this experiment the wrong box was indicated by the buzzing sound of the +apparatus and the slight vibrations which resulted from it. Although No. +18 was tested, as was No. 4, for ten successive days, ten trials each day, +it gave no evidence of ability to avoid the box-which-buzzed. + +Since both direct and indirect methods of testing the hearing of the +dancer have uniformly given negative results, in the case of mice more +than five weeks old, I feel justified in concluding that they are totally +deaf and not merely irresponsive to sounds. + +Rawitz's statements, and the fact that what may have been auditory +reactions were obtained with a few individuals of five weeks of age, +suggest that the mice may be able to hear at certain periods of life. To +discover whether this is true I have tested the young of twenty different +litters from the first day to the twenty-eighth, either daily or at +intervals of two or three days. In these tests König forks, steel bars, +and a Galton whistle were used. The results obtained are curiously +interesting. + +During the first two weeks of life none of the mice which I tested gave +any visible motor response to the various sounds used. During the third +week certain of the individuals responded vigorously to sudden high tones +and loud noises. After the third week I have seen only doubtful signs of +hearing. I shall now describe in detail the method of experimentation, the +condition of the animals, and the nature of the auditory reactions. + +Between the twelfth and the eighteenth day the auditory canal becomes open +to the exterior. The time is very variable in different litters, for their +rate of growth depends upon the amount of nourishment which the mother is +able to supply. Without exception, in my experience, the opening to the +ear appears before the eyes are open. Consequently visual stimuli usually +are not disturbing factors in the auditory tests with mice less than +sixteen days old. There is also a sudden and marked change in the behavior +of the mice during the third week. Whereas, for the first fourteen or +eighteen days they are rather quiet and deliberate in their movements when +removed from the nest, some time in the third week their behavior suddenly +changes and they act as if frightened when taken up by the experimenter. +They jump out of his hand, squeak, and sometimes show fight. This is so +pronounced that it has attracted my attention many times and I have +studied it carefully to determine, if possible, whether it is due to some +profound change in the nervous system which thus suddenly increases the +sensitiveness of the animal or to the development of the sexual organs. I +am inclined to think that it is a nervous phenomenon which is intimately +connected with the sexual condition. Within a day or two after it appears +the mice usually begin to show auditory reactions and continue to do so +for three to five days. + +I shall now describe the results obtained with a few typical litters. A +litter born of Nos. 151 and 152 gave uniformly negative results in all +auditory tests up to the fourteenth day. On that day the ears were open, +and the following observations were recorded. The five individuals of the +litter, four females and one male, were taken from the nest one at a time +at 7 A.M. and placed on a piece of paper in the bright sunlight. The +warmth of the sun soon quieted them so that auditory tests could be made +to advantage. As soon as an individual had become perfectly still, the +Galton whistle was held at a distance of about four inches from its head +in such a position that it could not be seen nor the currents of air +caused by it felt, and suddenly blown. Each of the five mice responded to +the first few repetitions of this stimulus by movements of the ears, +twitchings of the body, and jerky movements of the legs. The most violent +reactions resulted when the individual was lying on its back with its legs +extended free in the air. Under such circumstances the four legs were +often drawn together suddenly when the whistle was sounded. Similar +responses were obtained with the lip sound already mentioned. Two other +observers saw these experiments, and they agreed that there can be no +doubt that the mice responded to the sound. The sounds which were +effective lay between 5000 and 10,000 complete vibrations. + +On the fifteenth day the eyes were just beginning to open. Three of the +mice responded definitely to the sounds, but the other two slightly, if at +all. On the sixteenth day they were all too persistently active for +satisfactory auditory tests, and on the seventeenth, although they were +tested repeatedly under what appeared to be favorable conditions, no signs +of sensitiveness were noted. Although I continued to test this litter, at +intervals of three or four days, for two weeks longer, I did not once +observe a response to sound. + +This was the first litter with which I obtained perfectly definite, clear- +cut responses to sounds. That the reactive ability had not been present +earlier than the fourteenth day I am confident, for I had conducted the +tests in precisely the same manner daily up to the time of the appearance +of the reactions. To argue that the mice heard before the fourteenth day, +but were unable to react because the proper motor mechanism had not +developed sufficiently would be short-sighted, for if the response +depended upon the development of such a mechanism, it is not likely that +it would disappear so quickly. I am therefore satisfied that these +reactions indicate hearing. + +With another litter the following results were obtained. On the thirteenth +day each of the eight members of the litter responded definitely and +uniformly to the Galton whistle, set at 5 (probably about 8000 complete +vibrations), and to a König steel bar of a vibration rate of 4096 Vs. The +largest individuals, for almost always there are noticeable differences in +size among the members of a litter, appeared to be most sensitive to +sounds. + +On the fifteenth day and again on the seventeenth unmistakable responses +to sound were observed; on the eighteenth the responses were indefinite, +and on the nineteenth none were obtained. I continued the tests up to the +twenty-eighth day without further indications of hearing. + +Certain individuals in this litter reacted so vigorously to the loud sound +produced by striking the steel bar a sharp blow and also to the Galton +whistle, during a period of five days, that I have no hesitation in saying +that they evidently heard during that period of their lives. Other members +of the litter seemed to be less sensitive; their reactions were sometimes +so indefinite as to leave the experimenter in doubt about the presence of +hearing. + +A third litter, which developed very slowly because of lack of sufficient +food, first showed unmistakable reactions to sound on the twenty-first +day. On this day only two of the five individuals reacted. The reactions +were much more obvious on the twenty-second day, but thereafter they +became indefinite. + +Still another litter, which consisted of one female and four males, began +to exhibit the quick, jerky movements, already mentioned, on the +fourteenth day. On the morning of the fifteenth day three members of the +litter definitely reacted to the tone of the steel bar, and also to the +hammer blow when the bar was held tightly in the hand of the experimenter. +My observations were verified by another experimenter. Two individuals +which appeared to be very sensitive were selected for special tests. Their +reactions were obvious on the sixteenth, seventeenth, and eighteenth days; +on the nineteenth day they were indefinite, and on the twentieth none +could be detected. Some individuals of this litter certainly had the +ability to hear for at least five days. + +A sixth litter of four females and two males first gave indications of the +change in behavior which by this time I had come to interpret as a sign of +the approach of the period of auditory sensitiveness, on the seventeenth +day. I had tested them almost every day previous to this time without +obtaining evidence of hearing. The tests with the steel bar and the Galton +whistle were continued each day until the end of the fourth week without +positive results. To all appearances the individuals of this litter were +unable to hear at any time during the first month of life. + +Practically the same results were obtained with another litter of four +females. The change in their behavior was obvious on the eighteenth day, +but at no time during the first month did they give any satisfactory +indications of hearing. + +In the accompanying table, I have presented in condensed form the results +of my auditory tests in the case of twelve litters of young dancers. + + + + +TABLE 5 + +PERIOD OF AUDITORY REACTION IN YOUNG DANCERS + +PARENTS No. in Change in Ears Auditory Reactions + Litter Behavior Open Appear Disappear + +152+151 5 13th day 14th day 14th day 16th day +152+15l 8 (?) 13th day 13th day 17th day +152+151 5 13th day 13th day 13th day 17th day +152+151 4 10th day 12th day 13th day 15th day +410+415 5 14th day 15th day 15th day 19th day +410+415 6 13th day 14th day 14th day 18th day +420+425 2 12th day 14th day 14th day 17th day +210+215 5 17th day 13th day 17th day 19th day +210+215 6 11th day 14th day No reactions +220+225 6 16th day 14th day No reactions +220+225 6 17th day 13th day No reactions +212+211 4 15th day 14th day No reactions + + + + +Certain of the litters tested responded definitely to sounds, others gave +no sign of hearing at any time during the first four weeks of life. Of the +twelve litters for which the results of auditory tests are presented in +Table 5, eight evidently passed through an auditory period. It is +important to note that all except one of these were the offspring of Nos. +151 and 152, or of their descendants Nos. 410 and 415 and Nos. 420 and +425. In fact every one of the litters in this line of descent which I have +tested, and they now number fifteen, has given indications of auditory +sensitiveness. And, on the other hand, only in a single instance have the +litters born of Nos. 210 and 215, or of their descendants, given evidence +of ability to hear. + +These two distinct lines of descent may be referred to hereafter as the +400 and the 200 lines. I have observed several important differences +between the individuals of these groups in addition to the one already +mentioned. The 200 mice were sometimes gray and white instead of black and +white; they climbed much more readily and danced less vigorously than +those of the 400 group. These facts are particularly interesting in +connection with Cyon's descriptions of the two types of dancer which he +observed. + +In criticism of my conclusion that the young dancers are able to hear +certain sounds for a few days early in life, and then become deaf, it has +been suggested that they cease to react because they rapidly become +accustomed to the sounds. That this is not the case, is evident from the +fact that the reactions often increase in definiteness during the first +two or three days and then suddenly disappear entirely. But even if this +were not true, it would seem extremely improbable that the mouse should +become accustomed to a sudden and startlingly loud sound with so few +repetitions as occurred in these tests. On any one day the sounds were not +made more than five to ten times. Moreover, under the same external +condition, the common mouse reacts unmistakably to these sounds day after +day when they are first produced, although with repetition of the stimulus +at short intervals, the reactions soon become indefinite or disappear. + +The chief results of my study of hearing in the dancer may be summed up in +a very few words. The young dancer, in some instances, hears sounds for a +few days during the third week of life. The adult is totally deaf. Shortly +before the period of auditory sensitiveness, the young dancer becomes +extremely excitable and pugnacious. + + + + +CHAPTER VII + +THE SENSE OF SIGHT: BRIGHTNESS VISION + +The sense of sight in the dancer has received little attention hitherto. +In the literature there are a few casual statements to the effect that it +is of importance. Zoth, for example (31 p. 149), remarks that it seems to +be keenly developed; and other writers, on the basis of their observation +of the animal's behavior, hazard similar statements. The descriptions of +the behavior of blinded mice, as given by Cyon, Alexander and Kreidl, and +Kishi (p.47), apparently indicate that the sense is of some value; they do +not, however, furnish definite information concerning its nature and its +role in the daily life of the animal. + +The experimental study of this subject which is now to be described was +undertaken, after careful and long-continued observation of the general +behavior of the dancer, in order that our knowledge of the nature and +value of the sense of sight in this representative of the Mammalia might +be increased in scope and definiteness. The results of this study +naturally fall into three groups: (1) those which concern brightness +vision, (2) those which concern color vision, and (3) those which indicate +the role of sight in the life of the dancer. + +Too frequently investigators, in their work on vision in animals, have +assumed that brightness vision and color vision are inseparable; or, if +not making this assumption, they have failed to realize that the same +wave-length probably has markedly different effects upon the retinal +elements of the eyes of unlike organisms. In a study of the sense of sight +it is extremely important to discover whether difference in the quality, +as well as in the intensity, of a visual stimulus influences the organism; +in other words, whether color sensitiveness, as well as brightness +sensitiveness, is present. If the dancer perceives only brightness or +luminosity, and not color, it is evident that its visual world is +strikingly different from that of the normal human being. The experiments +now to be described were planned to show what the facts really are. + +[Illustration: Figure 14.--Discrimination box. _W_, electric-box with +white cardboards; B, electric-box with black cardboards. Drawn by Mr. C.H. +Toll.] + +As a means of testing the ability of the dancer to distinguish differences +in brightness, the experiment box represented by Figures 14 and 15 was +devised. Figure 14 is the box as seen from the position of the +experimenter during the tests. Figure 15 is its ground plan. This box, +which was made of wood, was 98 cm. long, 38 cm. wide, and 17 cm. deep, as +measured on the outside. The plan of construction and its significance in +connection with these experiments on vision will be clear from the +following account of the experimental procedure. A mouse whose brightness +vision was to be tested was placed in the nest-box, A (Figure 15). Thence +by pushing open the swinging door at _I_, it could pass into the entrance +chamber, _B_. Having entered _B_ it could return to _A_ only by passing +through one of the electric-boxes, marked _W_, and following the alley to +_O_, where by pushing open the swing door it could enter the nest-box. The +door at _I_ swung inward, toward _B_, only; those at _O_, right and left, +swung outward, toward _A_, only. It was therefore impossible for the mouse +to follow any other course than _A-I-B-L-W-E-O_ or _A-I-B-R-W-E-O_. The +doors at _I_ and _O_ were pieces of wire netting of 1/2 cm. mesh, hinged +at the top so that a mouse could readily open them, in one direction, by +pushing with its nose at any point along the bottom. On the floor of each +of the electric-boxes, _W_, was an oak board 1 cm. in thickness, which +carried electric wires by means of which the mouse could be shocked in _W_ +when the tests demanded it. The interrupted circuit constituted by the +wires in the two electric-boxes, in connection with the induction +apparatus, _IC_, the dry battery, _C_, and the hand key, _K_, was made by +taking two pieces of No. 20 American standard gauge copper wire and +winding them around the oak board which was to be placed on the floor of +each electric-box. The wires, which ran parallel with one another, 1/2 cm. +apart, fitted into shallow grooves in the edges of the board, and thus, as +well as by being drawn taut, they were held firmly in position. The coils +of the two pieces of wire alternated, forming an interrupted circuit +which, when the key _K_ was closed, was completed if the feet of a mouse +rested on points of both pieces of wire. Since copper wire stretches +easily and becomes loose on the wooden base, it is better to use phosphor +bronze wire of about the same size, if the surface covered by the +interrupted circuit is more than three or four inches in width. The +phosphor bronze wire is more difficult to wind satisfactorily, for it is +harder to bend than the copper wire, and it has the further disadvantage +of being more brittle. But when once placed properly, it forms a far more +lasting and satisfactory interrupted circuit for such experiments as those +to be described than does copper wire. In the case of the electric-boxes +under consideration, the oak boards which carried the interrupted circuits +were separate, and the two circuits were joined by the union of the wires +between the boxes. The free ends of the two pieces of wire which +constituted the interrupted circuit were connected with the secondary coil +of a Porter inductorium whose primary coil was in circuit with a No. 6 +Columbia dry battery. In the light of preliminary experiments, made in +preparation for the tests of vision, the strength of the induced current +received by the mouse was so regulated, by changing the position of the +secondary coil with reference to the primary, that it was disagreeable but +not injurious to the animal. What part the disagreeable shock played in +the test of brightness vision will now be explained. + +[Illustration: FIGURE 15.--Ground plan of discrimination box. _A_, nest- +box; _B_, entrance chamber; _W,W_, electric-boxes; _L_, doorway of left +electric-box; _R_, doorway of right electric-box; _E_, exit from electric- +box to alley; _I_, swinging door between _A_ and _B_; _O_, swinging door +between alley and _A_; _IC_, induction apparatus; _C_, electric cell; _K_, +key in circuit.] + +An opportunity for visual discrimination by brightness difference was +provided by placing dead black cardboard at the entrance and on the inside +of one of the electric-boxes, as shown in Figure 14, _B_, and white +cardboard similarly in the other box. These cardboards were movable and +could be changed from one box to the other at the will of the +experimenter. The test consisted in requiring the mouse to choose a +certain brightness, for example, the white cardboard side, in order to +return to the nest-box without receiving an electric shock. The question +which the experimenter asked in connection with this test really is, Can a +dancer learn to go to the white box and thus avoid discomfort? If we +assume its ability to profit by experience within the limits of the number +of experiences which it was given, such a modification of behavior would +indicate discrimination of brightness. Can the dancer distinguish white +from black; light gray from dark gray; two grays which are almost of the +same brightness? The results which make up the remainder of this and the +following chapter furnish a definite answer to these questions. + +To return to the experimental procedure, the mouse which is being tested +is placed by the experimenter in the nest-box, where frequently in the +early tests food and a comfortable nest were attractions. If it does not +of its own accord, as a result of its abundant random activity, pass +through _I_ into _B_ within a few seconds, it is directed to the doorway +and urged through. A choice is now demanded of the animal; to return to +the nest-box it must enter either the white electric-box or the black one. +Should it choose the white box, it is permitted to return directly to _A_ +by way of the doorway _E_, the alley, and the swinging door at _O_, and it +thus gets the satisfaction of unobstructed activity, freedom to whirl, to +feed, and to retreat for a time to the nest. Should it choose to attempt +to enter the black box, as it touches the wires of the interrupted circuit +it receives a shock as a result of the closing of the key in the circuit +by the experimenter, and further, if it continues its forward course +instead of retreating from the "stinging" black box, its passage through +_E_ is blocked by a barrier of glass temporarily placed there by the +experimenter, and the only way of escape to the nest-box is an indirect +route by way of _B_ and the white box. Ordinarily the shock was given only +when the mouse entered the wrong box, not when it retreated from it; it +was never given when the right box was chosen. The box to be chosen, +whether it was white, gray, or black, will be called the right box. The +electric shock served as a means of forcing the animal to use its +discriminating ability. But the question of motives in the tests is not so +simple as might appear from this statement. + +The reader will wonder why the mouse should have any tendency to enter +_B_, and why after so doing, it should trouble to go further, knowing, as +it does from previous experiences, that entering one of the electric-boxes +may result in discomfort. The fact is, a dancer has no very constant +tendency to go from _A_ to _B_ at the beginning of the tests, but after it +has become accustomed to the box and has learned what the situation +demands, it shows eagerness to make the trip from _A_ to _B_, and thence +by way of either the right or the left route to _A_. That the mouse should +be willing to enter either of the electric-boxes, after it has experienced +the shock, is even more surprising than its eagerness to run from _A_ to +_B_. When first tested for brightness discrimination in this apparatus, a +dancer usually hesitated at the entrance to the electric-boxes, and this +hesitation increased rapidly unless it were able to discriminate the boxes +by their difference in brightness and thus to choose the right one. During +the period of increasing hesitancy in making the choice, the experimenter, +by carefully moving from _I_ toward the entrances to the electric-boxes a +piece of cardboard which extended all the way across _B_, greatly +increased the mouse's desire to enter one of the boxes by depriving it of +dancing space in _B_. If an individual which did not know which entrance +to choose were permitted to run about in _B_, it would often do so for +minutes at a time without approaching the entrance to the boxes; but the +same individual, when confined to a dancing space 4 or 5 cm. wide in front +of the entrances, would enter one of the electric-boxes almost +immediately. This facilitation of choice by decrease in the amount of +space for whirling was not to any considerable extent the result of fear, +for all the dancers experimented with were tame, and instead of forcing +them to rush into one of the boxes blindly and without attempt at +discrimination, the narrowing of the space simply increased their efforts +to discriminate. The common mouse when subjected to similar experimental +conditions is likely to be frightened by being forced to approach the +entrances to the boxes, and fails to choose; it rushes into one box +directly, and in consequence it is as often wrong as right. The dancer +always chooses, but its eagerness to choose is markedly increased by the +restriction of its movements to a narrow space in front of the entrances +between which it is required to discriminate. It is evident that the +animal is uncomfortable in a space which is too narrow for it to whirl in +freely. It must have room to dance. This furnished a sufficiently strong +motive for the entering of the electric-boxes. It must avoid disagreeable +and unfavorable stimuli. This is a basis for attempts to choose, by visual +discrimination, the electric-box in which the shock is not given. It may +safely be said that the success of the majority of the experiments of this +book depended upon three facts: (1) the dancer's tendency to avoid +disagreeable external conditions, (2) its escape-from-confinement- +impelling need of space in which to dance freely, and (3) its abundant and +incessant activity. + +Of these three conditions of success in the experiments, the second and +third made possible the advantageous use of the first. For the avoidance +of a disagreeable stimulus could be made use of effectively in the tests +just because the mice are so restless and so active. In fact their +eagerness to do things is so great that the experimenter, instead of +having to wait for them to perform the desired act, often is forced to +make them wait while he completes his observation and record. In this +respect they are unlike most other animals. + +My experiments with the dancer differ from those which have been made by +most students of mammalian behavior in one important respect. I have used +punishment instead of reward as the chief motive for the proper +performance of the required act. Usually in experiments with mammals +hunger has been the motive depended upon. The animals have been required +to follow a certain devious path, to escape from a box by working a +button, a bolt, a lever, or to gain entrance to a box by the use of teeth, +claws, hands, or body weight and thus obtain food as a reward. There are +two very serious objections to the use of the desire for food as a motive +in animal behavior experiments--objections which in my opinion render it +almost worthless in the case of many mammals. These are the discomfort of +the animal and the impossibility of keeping the motive even fairly +constant. However prevalent the experience of starvation may be in the +life of an animal, it is not pleasant to think of subjecting it to extreme +hunger in the laboratory for the sake of finding out what it can do to +obtain food. Satisfactory results can be obtained in an experiment whose +success depends chiefly upon hunger only when the animal is so hungry that +it constantly does its best to obtain food, and when the desire for food +is equally strong and equally effective as a spur to action in the +repetitions of the experiment day after day. It is easy enough to get +almost any mammal into a condition of utter hunger, but it is practically +impossible to have the desire for food of the same strength day after day. +In short, the desire for food is unsatisfactory as a motive in animal +behavior work, first, because a condition of utter hunger, as has been +demonstrated with certain mammals, is unfavorable for the performance of +complex acts, second, because it is impossible to control the strength of +the motive, and finally, because it is an inhumane method of +experimentation. + +In general, the method of punishment is more satisfactory than the method +of reward, because it can be controlled to a greater extent. The +experimenter cannot force his subject to desire food; he can, however, +force it to discriminate between conditions to the best of its knowledge +and ability by giving it a disagreeable stimulus every time it makes a +mistake. In other words, the conditions upon which the avoidance of a +disagreeable factor in the environment depends are far simpler and much +more constant than those upon which the seeking of an agreeable factor +depends. Situations which are potentially beneficial to the animal attract +it in varying degrees according to its internal condition; situations +which are potentially disagreeable or injurious repel it with a constancy +which is remarkable. The favorable stimulus solicits a positive response; +the unfavorable stimulus demands a negative response. + +Finally, in connection with the discussion of motives, it is an important +fact that forms of reward are far harder to find than forms of punishment. +Many animals feed only at long intervals, are inactive, do not try to +escape from confinement, cannot be induced to seek a particular spot, in a +word, do not react positively to any of the situations or conditions which +are employed usually in behavior experiments. It is, however, almost +always possible to find some disagreeable stimulus which such an animal +will attempt to avoid. + +As it happens, the dancer is an animal which does not stand the lack of +food well enough to make hunger a possible motive. I was driven to make +use of the avoiding reaction, and it has proved so satisfactory that I am +now using it widely in connection with experiments on other animals. The +use of the induction shock, upon which I depended almost wholly in the +discrimination experiments with the dancer, requires care; but I am +confident that no reasonable objection to the conduct of the experiments +could be made on the ground of cruelty, for the strength of the current +was carefully regulated and the shocks were given only for an instant at +intervals. The best proof of the humaneness of the method is the fact that +the animals continued in perfect health during months of experimentation. + +The brightness discrimination tests demanded, in addition to motives for +choice, adequate precautions against discrimination by other than visual +factors, and, for that matter, by other visual factors than that of +brightness. The mouse might choose, for example, not the white or the +black box, but the box which was to the right or to the left, in +accordance with its experience in the previous test. This would be +discrimination by position. As a matter of fact, the animals have a strong +tendency at first to go uniformly either to the right or to the left +entrance. This tendency will be exhibited in the results of the tests. +Again, discrimination might depend upon the odors of the cardboards or +upon slight differences in their shape, texture, or position. Before +conclusive evidence of brightness discrimination could be obtained, all of +these and other possibilities of discrimination had to be eliminated by +check tests. I shall describe the various precautions taken in the +experiments to guard against errors in interpretation, in order to show +the lengths to which an experimenter may be driven in his search for +safely interpretable results. + +To exclude choice by position, the cardboards were moved from one +electric-box to the other. When the change was made regularly, so that +white was alternately on the right and the left, the mouse soon learned to +go alternately to the right box and the left without stopping to notice +the visual factor. This was prevented by changing the position of the +cardboards irregularly. + +Discrimination by the odor, texture, shape, and position of the cardboards +was excluded by the use of different kinds of cardboards, by changing the +form and position of them in check tests, and by coating them with +shellac. + +The brightness vision tests described in this chapter were made in a room +which is lighted from the south only, with the experiment box directed +away from the windows. The light from the windows shone upon the +cardboards at the entrances to the electric-boxes, not into the eyes of +the mouse as it approached them. Each mouse used in the experiments was +given a series of ten tests in succession daily. The experiment was +conducted as follows. A dancer was placed in _A_, where it usually ran +about restlessly until it happened to find its way into _B_. Having +discovered that the swing door at _I_ could be pushed open, the animal +seemed to take satisfaction in passing through into _B_ as soon as it had +been placed in or had returned to _A_. In _B_, choice of two entrances, +one of which was brighter than the other, was forced by the animal's need +of space for free movement. If the right box happened to be chosen, the +mouse returned to _A_ and was ready for another test; if it entered the +wrong box, the electric shock was given, and it was compelled to retreat +from the box and enter the other one instead. In the early tests with an +individual, a series sometimes covered from twenty to thirty minutes; in +later tests, provided the condition of discrimination was favorable, it +did not occupy more than ten minutes. + +To exhibit the methods of keeping the records of these experiments and +certain features of the results, two sample record sheets are reproduced +below. The first of these sheets, Table 6, represents the results given by +No. 5, a female,[1] in her first series of white-black tests. Table 7 +presents the results of the eleventh series of tests given to the same +individual. + +[Footnote 1: It is to be remembered that the even numbers always designate +males; the odd numbers, females.] + +In the descriptions of the various visual experiments of this and the +following chapters, the first word of the couplet which describes the +condition of the experiment, for example, white-black, always designates +the visual condition which the animal was to choose, the second that which +it was to avoid on penalty of an electric shock. In the case of Tables 6 +and 7, for example, white cardboard was placed in one box, black in the +other, and the animal was required to enter the white box. In the tables +the first column at the left gives the number of the test, the second the +positions of the cardboards, and the third and fourth the result of the +choice. The first test of Table 6 was made with the white cardboard on the +box which stood at the left of the mouse as it approached from _A_, and, +consequently, with the black cardboard on the right. As is indicated by +the record in the "wrong" column, the mouse chose the black instead of the +white. The result of this first series was choice of the white box four +times as compared with choice of the black box six times. On the eleventh +day, that is, after No. 5 had been given 100 tests in this brightness +vision experiment, she made no mistakes in a series of ten trials (Table +7). + + +TABLE 6 + +BRIGHTNESS DISCRIMINATION + +White-Black, Series 1 + +Experimented on No. 5 January 15, 1906 + POSITION OF +TEST CARDBOARDS RIGHT WRONG + +1 White left -- Wrong +2 White right -- Wrong +3 White left -- Wrong +4 White right -- Wrong +5 White left Right -- +6 White right Right -- +7 White left -- Wrong +8 White right Right -- +9 White left -- Wrong +10 White right Right -- + +Totals 4 6 + + +Before tests, such as have been described, can be presented as conclusive +proof of discrimination, it must be shown that the mouse has no preference +for the particular brightness which the arrangement of the test requires +it to select. That any preference which the mouse to be tested might have +for white, rather than black, or for a light gray rather than a dark gray, +might be discovered, what may be called preference test series were given +before the discrimination tests were begun. These series, two of which +were given usually, consisted of ten tests each, with the white +alternately on the left and on the right. The mouse was permitted to enter +either the white or the black box, as it chose, and to pass through to the +nest-box without receiving a shock and without having its way blocked by +the glass plate. The conditions of these preference tests may be referred +to hereafter briefly as "No shock, open passages." The preference tests, +which of course would be valueless as such unless they preceded the +training tests, were given as preliminary experiments, in order that the +experimenter might know how to plan his discrimination tests, and how to +interpret his results. + + + +TABLE 7 + +BRIGHTNESS DISCRIMINATION + +White-Black, Series II + +Experimented on No. 5 February 2, 1906 + + POSITION +TEST OF CARDBOARDS RIGHT WRONG + + 1 White left Right -- + 2 White left Right -- + 3 White right Right -- + 4 White right Right -- + 5 White right Right -- + 6 White left Right -- + 7 White left Right -- + 8 White left Right -- + 9 White right Right -- +10 White right Right -- + +Totals 10 0 + + + +The results given in the white-black preference tests by ten males and ten +females are presented in Table 8. Three facts which bear upon the +brightness discrimination tests appear from this table: (1) black is +preferred by both males and females, (2) this preference is more marked in +the first series of tests than in the second, and (3) it is slightly +stronger for the first series in the case of females than in the case of +males. + + +TABLE 8 + +WHITE-BLACK PREFERENCE TESTS + +MALES FIRST SERIES SECOND SERIES + WHITE BLACK WHITE BLACK + +No. 10 3 7 3 7 + 18 5 5 5 5 + 20 2 8 4 6 + 152 4 6 6 4 + 210 4 6 4 6 + 214 6 4 3 7 + 220 5 5 3 7 + 230 4 6 2 8 + 410 4 6 5 5 + 420 4 6 9 1 + +Averages 4.1 5.9 4.4 5.6 + + +FEMALES FIRST SERIES SECOND SERIES + WHITE BLACK WHITE BLACK + +No. 11 5 5 4 6 + 151 6 4 5 5 + 215 2 8 2 8 + 213 2 8 5 5 + 225 4 6 2 8 + 227 4 6 6 4 + 235 6 4 4 6 + 415 2 8 4 6 + 425 5 5 8 2 + 229 2 8 5 5 + +Averages 3.8 6.2 4.5 5.5 + + +That the dancers should prefer to enter the dark rather than the light box +is not surprising in view of the fact that the nests in which they were +kept were ordinarily rather dark. But whatever the basis of the +preference, it is clear that it must be taken account of in the visual +discrimination experiments, for an individual which strongly preferred +black might choose correctly, to all appearances, in its first black-white +series. Such a result would demonstrate preference, and therefore one kind +of discrimination, but not the formation of a habit of choice by +discrimination. The preference for black is less marked in the second +series of tests because the mouse as it becomes more accustomed to the +experiment box tends more and more to be influenced by other conditions +than those of brightness. The record sheets for both series almost +invariably indicate a strong tendency to continue to go to the left or the +right entrance according to the way by which the animal escaped the first +time. This cannot properly be described as visual choice, for the mouse +apparently followed the previous course without regard to the conditions +of illumination. We have here an expression of the tendency to the +repetition of an act. It is only after an animal acquires considerable +familiarity with a situation that it begins to vary its behavior in +accordance with relatively unimportant factors in the situation. It is +this fact, illustrations of which may be seen in human life, as well as +throughout the realm of animal behavior, that renders it imperative that +an animal be thoroughly acquainted with the apparatus for experimentation +and with the experimenter before regular experiments are begun. Any animal +will do things under most experimental conditions, but to discover the +nature and scope of its ability it is necessary to make it thoroughly at +home in the experimental situation. As the dancer began to feel at home in +the visual discrimination apparatus it began to exercise its +discriminating ability, the first form of which was choice according to +position. + +Since there appears to be a slight preference on the part of most dancers' +for the black box in comparison with the white box, white-black training +tests were given to fifty mice, and black-white to only four. The tests +with each individual were continued until it had chosen correctly in all +of the tests of three successive series (thirty tests). As the +reproduction of all the record sheets of these experiments would fill +hundreds of pages and would provide most readers with little more +information than is obtainable from a simple statement of the number of +right and wrong choices, only the brightness discrimination records of +Tables 6 and 7 are given in full. + +As a basis for the comparison of the results of the white-black tests with +those of the black-white tests, two representative sets of data for each +of these conditions of brightness discrimination are presented (Tables 9 +and 10). In these tables only the number of right and wrong choices for +each series of ten tests appears. + +Tables 9 and 10 indicate--if we grant that the precautionary tests to be +described later exclude the possibility of other forms of discrimination-- +that the dancer is able to tell white from black; that it is somewhat +easier, as the preference tests might lead us to expect, for it to learn +to go to the black than to the white, and that the male forms the habit of +choosing on the basis of brightness discrimination more quickly than the +female. + + + +TABLE 9 +WHITE-BLACK TESTS + + + No. 210 No. 215 + AGE, 28 DAYS AGE, 28 DAYS +SERIES DATE RIGHT WRONG RIGHT WRONG + (WHITE) (BLACK) (WHITE) (BLACK) + A June 22 4 6 2 8 + B 23 4 6 2 8 + + 1 24 4 6 3 7 + 2 25 6 4 5 5 + 3 26 7 3 7 3 + 4 27 5 5 8 2 + 5 28 7 3 9 1 + 6 29 8 2 8 2 + 7 30 9 1 9 1 + 8 July 1 10 0 10 0 + 9 2 10 0 9 1 + 10 3 10 0 10 0 + 11 4 -- -- 10 0 + 12 5 -- -- 10 0 + + + + +TABLE 10 +WHITE-BLACK TESTS + + No. 14 No. 13 + AGE, 32 DAYS AGE, 32 DAYS +SERIES DATE RIGHT WRONG RIGHT WRONG + (WHITE) (BLACK) (WHITE) (BLACK) + 1 May 13[1] 5 5 7 3 + 2 14 8 2 6 4 + 3 15 7 3 9 1 + 4 16 9 1 9 1 + 5 17 10 0 10 0 + 6 18 10 0 9 1 + 7 19 10 0 10 0 + 8 20 -- -- 10 0 + 9 21 -- -- 10 0 + +[Footnote 1: No preference tests were given.] + + +It is now necessary to justify the interpretation of these results as +evidence of brightness discrimination by proving that all other conditions +for choice except brightness difference may be excluded without +interfering with the animal's ability to select the right box. We shall +consider in order the possibility of discrimination by position, by odor, +and by texture and form of the cardboards. + +The tendency which the dancer has in common with many, if not all, animals +to perform the same movement or follow the same path under uniform +conditions is an important source of error in many habit-formation +experiments. This tendency is evident even from casual observation of the +behavior of the dancer. The ease with which the habit of choosing the box +on the left or the box on the right is formed in comparison with that of +choosing the white box or the black box is strikingly shown by the +following experiment. Five mice were given one series of ten trials each +in the discrimination box of Figure 14 without the presence of cardboards +or of other means of visual discrimination. The electric shock was given +whenever the box on the left was entered. Thus without other guidance than +that of direction, for the boxes themselves were interchanged in position, +and, as was proved by additional tests, the animals were utterly unable to +tell one from the other, the mouse was required to choose the box on its +right. Only one of the five animals went to the box on the left after once +experiencing the electric shock. The results of the series are given in +Table 11. + + + +TABLE 11 + +CHOICE BY POSITION + Choices of Choices of + Box on Right Box on Left +First mouse 9 1 +Second mouse 8 2 +Third mouse 9 1 +Fourth mouse 9 1 +Fifth mouse 9 1 + + + +This conclusively proves that the habit of turning in a certain direction +or of choosing by position can be formed more readily than a habit which +depends upon visual discrimination. A rough comparison justifies the +statement that it takes from six to ten times as long for the dancer to +learn to choose the white box as it does to learn to choose the box on the +right. Since this is true, it is exceedingly important that the +possibility of choice by position or direction of movement be excluded in +the case of tests of brightness discrimination. To indicate how this was +effectively accomplished in the experiments, the changes in the position +of the cardboards made in the case of a standard set of white-black series +are shown in Table 12. The number of the series, beginning at the top of +the table with the two lettered preference series, is given in the first +column at the left, the number of the tests at the top of the table, and +the position of the white cardboard, left or right, is indicated below by +the letters l (left) and r (right). + + +TABLE 12 + +POSITION OF WHITE CARDBOARDS FOR A SET OF 150 TESTS + + +SERIES 1 2 3 4 5 6 7 8 9 10 + +Preference + A l r l r l r l r l r + B r l r l r l r l r l + + 1 r l r l r l r l r l + 2 l l r r l r l l r r + 3 r r l r l l r l r l + 4 l l l r r r l r r l + 5 r l r l r l r l r l + 6 l l r l r r l r l r + 7 r l l l r r r l r l + 8 r r l l r l r l r l + 9 r r r l l l r l r l + 10 l l l l r r r r l r + 11 r l r r r l l l r l + 12 r l r l r r l l r l + 13 r l r l l l r r r l + 14 l l l l r r r r l r + 15 r l r r r l l l r l + + +It is to be noted that in the case of each series of ten tests the white +cardboard was on the left five times and on the right five times. Thus the +establishment of a tendency in favor of one side was avoided. The +irregularity of the changes in position rendered it impossible for the +mouse to depend upon position in its choice. It is an interesting fact +that the dancer quickly learns to choose correctly by position if the +cardboards are alternately on the left box and on the right. + +The prevalent, although ill-founded, impression that mice have an +exceedingly keen sense of smell might lead a critic of these experiments +to claim that discrimination in all probability was olfactory rather than +visual. As precautions against this possibility the cardboards were +renewed frequently, so that no odor from the body of the mouse itself +should serve as a guiding condition, different kinds of cardboard were +used from time to time, and, as a final test, the cardboards were coated +with shellac so that whatever characteristic odor they may have had for +the dancer was disguised if not totally destroyed. Despite all these +precautions the discrimination of the boxes continued. A still more +conclusive proof that we have to do with brightness discrimination, and +that alone, in these experiments is furnished by the results of white- +black tests made with an apparatus which was so arranged that light was +transmitted into the two electric-boxes through a ground glass plate in +the end of each box. No cardboards were used. The illumination of each box +was controlled by changes in the position of the sources of light. Under +these conditions, so far as could be ascertained by critical examination +of the results, in addition to careful observation of the behavior of the +animals as they made their choices, there was no other guiding factor than +brightness difference. Nevertheless the mice discriminated the white from +the black perfectly. This renders unnecessary any discussion of the +possibility of discrimination by the texture or form of the cardboards. + +Since a variety of precautionary tests failed to reveal the presence, in +these experiments, of any condition other than brightness difference by +which the mice were enabled to choose correctly, and since evidence of +ability to discriminate brightness differences was obtained by the use of +both reflected light (cardboards) and transmitted light (lamps behind +ground glass), it is necessary to conclude that the dancer possesses +brightness vision. + + + +CHAPTER VIII + +THE SENSE OF SIGHT: BRIGHTNESS VISION (Continued) + +Since the ability of the dancer to perceive brightness has been +demonstrated by the experiments of the previous chapter, the next step in +this investigation of the nature of vision is a study of the delicacy of +brightness discrimination, and of the relation of the just perceivable +difference to brightness value. Expressed in another way, the problems of +this portion of the investigation are to determine how slight a difference +in brightness enables the dancer to discriminate one light from another, +and what is the relation between the absolute brightnesses of two lights +and that amount of difference which is just sufficient to render the +lights distinguishable. It has been discovered in the case of the human +being that a stimulus must be increased by a certain definite fraction of +its own value if it is to seem different. For brightness, within certain +intensity limits, this increase must be about one one-hundredth; a +brightness of 100 units, for example, is just perceivably different from +one of 101 units. The formulation of this relation between the amount of a +stimulus and the amount of change which is necessary that a difference be +noted is known as Weber's law. Does this law, in any form, hold for the +brightness vision of the dancing mouse? + +Two methods were used in the study of these problems. For the first +problem, that of the delicacy of brightness discrimination, I first used +light which was reflected from gray papers, according to the method of +Chapter VII. For the second, the Weber's law test, transmitted light was +used, in an apparatus which will be described later. Either of these +methods might have been used for the solution of both problems. Which of +them is the more satisfactory is definitely decided by the results which +make up the material of this chapter, Under natural conditions the dancer +probably sees objects which reflect light more frequently than it does +those which transmit it; it would seem fairer, therefore, to require it to +discriminate surfaces which differ in brightness. This the use of gray +papers does. But, on the other hand, gray papers are open to the +objections that they may not be entirely colorless (neutral), and that +their brightness values cannot be changed readily by the experimenter. As +will be made clear in the subsequent description of the experiments with +transmitted light, neither of these objections can be raised in connection +with the second method of experimentation. + +To determine the delicacy of discrimination with reflected light it is +necessary to have a series of neutral grays (colorless) whose adjacent +members differ from one another in brightness by less than the threshold +of discrimination of the animal to be tested. A series which promised to +fulfill these conditions was that of Richard Nendel of Berlin. It consists +of fifty papers, beginning with pure white, numbered 1, and passing by +almost imperceptible steps of decrease in brightness through the grays to +black, which is numbered 50. For the present we may assume that these +papers are so nearly neutral that whatever discrimination occurs is due to +brightness. The differences between successive papers of the series are +perceptible to man. The question is, can they, under favorable conditions +of illumination, be perceived by the dancer? + +On the basis of the fact that the dancer can discriminate between white +and black, two grays which differed from one another in brightness by a +considerable amount were chosen from the Nendel series; these were numbers +10 and 20. It seemed certain, from the results of previous experiments, +that the discrimination of these papers by brightness difference would be +possible, and that therefore the use of papers between these two extremes +would suffice to demonstrate the delicacy of discrimination. In Figure 16 +we have a fairly accurate representation of the relative brightness of the +Nendel papers Nos. 10, 15, and 20. + +[Illustration: FIGURE 16. Three of Nendel's gray papers: Nos. 10, 15, and +20. To exhibit differences in brightness.] + +Pieces of the gray papers were pasted upon cardboard carriers so that they +might be placed in the discrimination box as were the white and black +cardboards in the tests of brightness vision previously described. Mice +which had been trained to choose the white box by series of white-black +tests were now tested with light gray (No. 10) and dark gray (No. 20), my +assumption being that they would immediately choose the brighter of the +two if they were able to detect the difference. As a matter of fact this +did not always occur; some individuals had to be trained to discriminate +gray No. 10 from gray No. 20. As soon as an individual had been so trained +that the ability to choose the lighter of these grays was perfect, it was +tested with No. 10 in combination with No. 15. If these in turn proved to +be discriminable, No. 10 could be used with No. 14, with No. 13, and so on +until either the limit of discrimination or that of the series had been +reached. + +That it was not necessary to use other combinations than 10 with 20, and +10 with 15 is demonstrated by the results of Table 13. Mouse No. 420, +whose behavior was not essentially different from that of three other +individuals which were tested for gray discrimination, learned with +difficulty to choose gray No. 10 even when it was used with No. 20. Two +series of ten tests each were given to this mouse daily, and not until the +twentieth of these series (200 tests) did he succeed in making ten correct +choices in succession. Immediately after this series of correct choices, +tests with grays No. 10 and No. 15 were begun. In the case of this amount +of brightness difference twenty series failed to reveal discrimination. +The average number of right choices in the series is slightly in excess of +the mistakes, 5.8 as compared with 4.2. + +From the experiments with gray papers we may conclude that under the +conditions of the tests the amount by which Nendel's gray No. 10 differs +in brightness from No. 20 is near the threshold of discrimination, or, in +other words, that the difference in the brightness of the adjacent grays +of Figure 16 is scarcely sufficient to enable the dancer to distinguish +them. + + +TABLE 13 + +GRAY DISCRIMINATION + +The Delicacy of Brightness Discrimination + +No. 420 + + + + GRAYS NOS. 10 GRAYS NOS. 20 + AND 20 AND 15 +SERIES DATE DATE + NO. 10 NO. 2 NO. 10 NO. 15 + (RIGHT) (WRONG) (RIGHT) (WRONG) + + 1 Jan. 26 5 5 Feb. 6 8 2 + 2 27 8 2 6 5 5 + 3 28 6 4 7 9 1 + 4 28 2 8 7 7 3 + 5 29 1 9 8 5 5 + 6 29 6 4 8 6 4 + 7 30 9 1 9 5 5 + 8 30 7 3 9 6 4 + 9 31 6 4 10 8 2 + 10 31 4 6 10 3 7 + 11 Feb. 1 7 3 11 4 6 + 12 1 8 2 11 4 6 + 13 2 7 3 12 7 3 + 14 2 8 2 12 7 3 + 15 3 9 1 13 6 4 + 16 3 9 1 13 4 6 + 17 4 6 4 14 4 6 + 18 4 9 1 14 5 5 + 19 5 6 4 15 5 5 + 20 5 10 0 15 8 2 + + Averages 6.6 3.4 5.8 4.2 + + + + +This result of the tests with gray papers surprised me very much at the +time of the experiments, for all my previous observation of the dancer had +led me to believe that it is very sensitive to light. It was only after a +long series of tests with transmitted light, in what is now to be +described as the Weber's law apparatus, that I was able to account for the +meager power of discrimination which the mice exhibited in the gray tests. +As it happened, the Weber's law experiment contributed quite as +importantly to the solution of our first problem as to that of the second, +for which it was especially planned. + +For the Weber's law experiment a box similar to that used in the previous +brightness discrimination experiments (Figure 14) was so arranged that its +two electric-boxes could be illuminated independently by the light from +incandescent lamps directly above them. The arrangements of the light-box +and the lamps, as well as their relations to the other important parts of +the apparatus, are shown in Figure 17. The light-box consisted of two +compartments, of which one may be considered as the upward extension of +the left electric-box and the other of the right electric-box. The light- +box was pivoted at A and could be turned through an angle of 180° by the +experimenter. Thus, by the turning of the light-box, the lamp which in the +case of one test illuminated the left electric-box could be brought into +such a position that in the case of the next test it illuminated the right +electric-box. The practical convenience of this will be appreciated when +the number of times that the brightnesses of the two boxes had to be +reversed is considered. The light-box was left open at the top for +ventilation and the prevention of any considerable increase in the +temperature of the experiment box. In one side of each of the compartments +of the light-box a slit (B, B of the figure) was cut out for an +incandescent lamp holder. A strip of leatherette, fitted closely into inch +grooves at the edges of the slit, prevented light from escaping through +these openings in the sides of the light-box. By moving the strips of +leatherette, one of which appears in the figure, C, the lamps could be +changed in position with reference to the bottom of the electric-box. A +scale, S, at the edge of each slit enabled the experimenter to determine +the distance of the lamp from the floor of the electric-box. The front of +the light-box was closed, instead of being open as it appears in the +figure. + +[Illustration: Figure l7.--Weber's law apparatus for testing brightness +discrimination. Lower part, discrimination box similar to that of Figure +14. Upper part, rotatory light-box, pivoted at A, and divided into two +compartments by a partition P in the middle. L, L, incandescent lamps +movable in slits, B, B, in which a narrow strip of leatherette, C, serves +to prevent the escape of light. S, scale.] + +This apparatus has the following advantages. First, the electric-boxes, +between which the mouse is expected to discriminate by means of their +difference in brightness, are illuminated from above and the light +therefore does not shine directly from the lamps into the eyes of the +animal, as it approaches the entrances to the boxes. Choice is required, +therefore, between illuminated spaces instead of between two directly +illuminated surfaces. Second, the amount of illumination of each electric- +box can be accurately measured by the use of a photometer. Third, since +the same kind of lamp is used in each box, and further, since the lamps +may be interchanged at any time, discrimination by qualitative instead of +quantitative difference in illumination is excluded. And finally, fourth, +the tests can be made expeditiously, conveniently, and under such simple +conditions that there should be no considerable error of measurement or of +observation within the range of brightness which must be used. + +It was my purpose in the experiment with this apparatus to ascertain how +great the difference in the illumination of the two electric-boxes must be +in order that the mouse should be able to choose the brighter of them. +This I attempted to do by fixing an incandescent lamp of a certain known +illuminating power at such a position in one compartment of the light-box +that the electric-box below it was illuminated by what I call a standard +value, and by moving the incandescent lamp in the other compartment of the +light-box until the illumination of the electric-box below it was just +sufficiently less than that of the standard to enable the dancer to +distinguish them, and thereby to choose the brighter one. The light which +was changed from series to series I shall call the _variable_, in contrast +with the _standard_, which was unchanged. + +The tests, which were made in a dark-room under uniform conditions, were +given in series of fifty each; usually only one such series was given per +day, but sometimes one was given in the morning and another in the +afternoon of the same day. To prevent choice by position the lights were +reversed in position irregularly, first one, then the other, illuminating +the right electric-box. For the fifty tests of each initial series the +order of the changes in position was as follows: standard (brighter light) +on the _l_ (left), _l, r_ (right), _r, l, l, r, r, l, r, l, r, l, l, r, r, +l, l, r, r, l, l, l, r, r, r, l, r, l, r, r, r, l, l, l, r, r, r, l, l, r, +l, r, l, r, l, r, l, r, l_. Twenty-five times in fifty the standard light +illuminated the right electric-box, and the same number of times it +illuminated the left electric-box. When a second series was given under +the same conditions of illumination, a different order of change was +followed. + +In order to discover whether Weber's law holds in the case of the +brightness vision of the dancer it was necessary for me to determine the +just perceivable difference between the standard and the variable lights +for two or more standard values. I chose to work with three values, 5, 20, +and 80 hefners, and I was able to discover with a fair degree of accuracy +how much less than 5, 20, or 80 hefners, as the case might be, the +variable light had to be in order that it should be discriminable from the +other. For the work with the 5 hefner standard I used 2-candle-power +lamps,[1] for the 20, 4-candle-power, and for the 80, 16-candle-power. + +[Footnote 1: I give merely the commercial markings of the lamps. They had +been photometered carefully by two observers by means of a Lummer-Brodhun +photometer and a Hefner amyl acetate lamp previous to their use in the +experiment. For the photometric measurements in connection with the +Weber's law tests I made use of the Hefner lamp with the hope of attaining +greater accuracy than had been possible with a standard paraffine candle, +in the case of measurements which I had made in connection with the +experiments on color vision that are reported in Chapters IX and X. The +Hefner unit is the amount of light produced by an amyl acetate lamp at a +flame height of 40 mm. (See Stine's "Photometrical Measurements.") A +paraffine candle at a flame height of 50 mm. is equal to 1.2 Hefner +units.] + +For reasons which will soon appear, Weber's law tests were made with only +one dancer. This individual, No. 51, had been thoroughly trained in white- +black discrimination previous to the experiments in the apparatus which is +represented in Figure 17. Having given No. 51 more than two hundred +preliminary tests in the Weber's law apparatus with the electric-boxes +sufficiently different in brightness to enable her to discriminate +readily, I began my experiments by trying to ascertain how much less the +value of the illumination of one electric-box must be in order that it +should be discriminable from a value of 20 hefners in the other electric- +box. In recording the several series of tests and their results hereafter, +I shall state in Hefner units the value of the fixed or standard light and +the value of the variable light, the difference between the two in terms +of the former, and the average number of wrong choices in per cent. + +With the lamps so placed that the difference in the illumination of the +two electric-boxes was .53 of the value of the standard, that is about one +half, No. 51 made twenty wrong choices in one hundred, or 20 per cent. +When the difference was reduced to .36 (one third) the number of errors +increased to 36 per cent, and with an intermediate difference of .48 there +were 26 per cent of errors (see Table 14). + +Are these results indicative of discrimination, or are the errors in +choice too numerous to justify the statement that the dancer was able to +distinguish the boxes by their difference in brightness? Evidently this +question cannot be answered satisfactorily until we have decided what the +percentage of correct choices should be in order that it be accepted as +evidence of ability to discriminate, or, to put it in terms of errors, +what percentage of wrong choices is indicative of the point of just +perceivable difference in brightness. Theoretically, there should be as +many mistakes as right choices, 50 per cent of each, when the two +electric-boxes are equally illuminated (indiscriminable), but in practice +this does not prove to be the case because the dancer tends to return to +that electric-box through which in the previous test it passed safely, +whereas it does not tend in similar fashion to reënter the box in which it +has just received an electric shock. The result is that the percentage of +right choices, especially in the case of series which have the right box +in the same position two, three, or four times in succession, rises as +high as 60 or 70, even when the visual conditions are indiscriminable. +Abundant evidence in support of this statement is presented in Chapters +VII and IX, but at this point I may further call attention to the results +of an experiment in the Weber's law apparatus which was made especially to +test the matter. The results appear under the date of May 27 in Table 14. +In this experiment, despite the fact that both boxes were illuminated by +80 hefners, the mouse chose the standard (the illumination in which it was +not shocked) 59 times in 100. In other words the percentage of error was +41 instead of 50. It is evident, therefore, that as low a percentage of +errors as 40 is not necessarily indicative of discrimination. Anything +below 40 per cent is likely, however, to be the result of ability to +distinguish the brighter from the darker box. To be on the safe side we +may agree to consider 25 wrong choices per 100 as indicative of a just +perceivable difference in illumination. Fewer mistakes we shall consider +indicative of a difference in illumination which is readily perceivable, +and more as indicative of a difference which the mouse cannot detect. The +reader will bear in mind as he examines Table 14 that 25 per cent of wrong +choices indicates the point of just perceivable difference in brightness. + + + +TABLE 14 + +RESULTS OF WEBER'S LAW EXPERIMENTS +Brightness vision + +DATE NUMBER STANDARD VARIABLE DIFFERENCE % OF ERRORS + OF TESTS LIGHT LIGHT + +May 13 100 20 9.4 .53 20 + 15 100 20 12.8 .36 36 + 16 100 20 10.8 .46 26 + 20 50 80 37.6 .53 6 + 21 50 80 51.3 .36 10 + 22 100 80 71.1 .11 35 + 24 100 80 60.0 .25 21 + 25 100 80 65.0 .19 25 + 27 100 80 80 0 41 + 28 50 5 2.5 .50 18 + 29 50 5 4.0 .20 14 + 29 100 5 4.5 .10 25 + 31 50 5 4.25 .15 20 +June 1 50 5 4.85 .03 48 + 2 50 20 15.0 .25 16 + 3 50 20 17.4 .13 22 + 3 100 20 18.0 .10 22 + 4 100 80 72.0 .10 18 + 5 100 5 4.5 .10 12 + 7 100 5 4.67 .067 46 + 8 50 80 74.67 .067 56 + 9 50 20 18.67 .067 44 + + + +If we apply this rule to the results of the first tests, reported above, +it appears that a standard of 20 hefners was distinguished from a variable +of 9.4 hefners (.53 difference), for the percentage of errors was only 20. +But in the case of a difference of .36 in the illuminations lack of +discrimination is indicated by 36 per cent of errors. A difference of .46 +gave a frequency of error so close to the required 25 (26 per cent) that I +accepted the result as a satisfactory determination of the just +perceivable difference for the 20 hefner standard and proceeded to +experiment with another standard value. + +The results which were obtained in the case of this second standard, the +value of which was 80 hefners, are strikingly different from those for the +20 hefner standard. Naturally I began the tests with this new standard by +making the differences the same as those for which determinations had been +made in the case of the 20 standard. Much to my surprise only 6 per cent +of errors resulted when the difference in illumination was .53. I finally +discovered that about .19 difference (about one fifth) could be +discriminated with that degree of accuracy which is indicated by 25 per +cent of mistakes. + +So far as I could judge from the results of determinations for the 20 and +the 80 hefner standards, Weber's law does not hold for the dancer. With +the former a difference of almost one half was necessary for +discrimination; with the latter a difference of about one fifth could be +perceived. But before presenting additional results I should explain the +construction of Table 14, and comment upon the number of experiments which +constitutes a set. + +The table contains the condensed results of several weeks of difficult +experimentation. From left to right the columns give the date of the +initial series of a given set of experiments, the number of experiments in +the set, the value of the standard light in hefners, the value of the +variable light, the difference between the lights in terms of the standard +(the variable was always less than the standard), and the percentage of +errors or wrong choices. Very early in the investigation I discovered that +one hundred tests with any given values of the lights sufficed to reveal +whatever discriminating ability the mouse possessed at the time. In some +instances either the presence or the lack of discrimination was so clear, +as the result of 50 tests (first series), that the second series of 50 was +not given. Consequently in the table the number of tests for the various +values of the lights is sometimes 100, sometimes 50. + +After finishing the experiments with the 80 standard on May 27 (see table) +I decided, in spite of the evidence against Weber's law, to make tests +with 5 as the standard, for it seemed not impossible that the lights were +too bright for the dancer to discriminate readily. I even suspected that I +might have been working outside of the brightness limits in which Weber's +law holds, if it holds at all. The tests soon showed that a difference of +one tenth made discrimination possible in the case of this standard. If +the reader will examine the data of the table, he will note that a +difference of .20 gave 14 per cent of mistakes; a difference of .03, 48 +per cent. Evidently the former difference is above the threshold, the +latter below it. But what of the interpretation of the results in terms of +Weber's law? The difference instead of being one half or one fifth, as it +was in the cases of the 20 and 80 standards respectively, has now become +one tenth. Another surprise and another contradiction! + +Had these three differences either increased or decreased regularly with +the value of the standard I should have suspected that they indicated a +principle or relationship which is different from but no less interesting +than that which Weber's law expresses. But instead of reading 5 standard, +difference one tenth; 20 standard, difference one fifth; 80 standard, +difference one half: or 5 standard, difference one half; 20 standard, +difference one fifth; 80 standard, difference one tenth: they read 5 +standard, difference one tenth; 20 standard, difference one half; 80 +standard, difference one fifth. What does this mean? I could think of no +other explanation than that of the influence of training. It seemed not +impossible, although not probable, that the mouse had been improving in +ability to discriminate day by day. It is true that this in itself would +be quite as interesting a fact as any which the experiment might reveal. + +To test the value of my supposition, I made additional experiments with +the 20 standard, the results of which appear under the dates June 2 and 3 +of the table. These results indicate quite definitely that the animal had +been, and still was, improving in her ability to discriminate. For instead +of requiring a difference of about one half in order that she might +distinguish the 20 standard from the variable light she was now able to +discriminate with only 22 per cent of errors when the difference was one +tenth. + +As it seemed most improbable that improvement by training should continue +much longer, I next gave additional tests with the 80 standard. Again a +difference of one tenth was sufficient for accurate discrimination (18 per +cent of errors). These series were followed immediately by further tests +with the 5 standard. As the results indicated greater ease of +discrimination with a difference of one tenth in the case of this standard +than in the case of either of the others I was at first uncertain whether +the results which I have tabulated under the dates June 3, 4, and 5 of the +table should be interpreted in terms of Weber's law. + +Up to this point the experiments had definitely established two facts: +that the dancer's ability to discriminate by means of brightness +differences improves with training for a much longer period and to a far +greater extent than I had supposed it would; and that a difference of one +tenth is sufficient to enable the animal to distinguish two lights in the +case of the three standard values, 5, 20, and 80 hefners. The question +remains, is this satisfactory evidence that Weber's law holds with respect +to the brightness vision of the dancer, or do the results indicate rather, +that this difference is more readily detected in the case of 5 as a +standard (12 per cent error) than in the case of 20 as a standard (22 per +cent error)? + +For the purpose of settling this point I made tests for each of the three +standards with a difference of only one fifteenth. In no instance did I +obtain the least evidence of ability to discriminate. These final tests, +in addition to establishing the fact that the limit of discrimination for +No. 51, after she had been subjected to about two thousand tests, lay +between one tenth and one fifteenth, proved to my satisfaction, when taken +in connection with the results already discussed, that Weber's law does +hold for the brightness vision of the dancer. + +In concluding this discussion of the Weber's law experiment I wish to call +attention to the chief facts which have been revealed, and to make a +critical comment. In my opinion it is extremely important that the student +of animal behavior should note the fact that the dancer with which I +worked week after week in the Weber's law investigation gradually improved +in her ability to discriminate on the basis of brightness differences +until she was able to distinguish from one another two boxes whose +difference in illumination was less than one tenth[1] that of the brighter +box. At the beginning of the experiments a difference of one half did not +enable her to choose as certainly as did a difference of one tenth after +she had chosen several hundred times. Evidently we are prone to +underestimate the educability of our animal subjects. + +[Footnote 1: Under the conditions of the experiment I was unable to +distinguish the electric-boxes when they differed by less than one +twentieth.] + + The reason that the experiments were carried out with only one mouse must +now be apparent. It was a matter of time. The reader must not suppose that +my study of this subject is completed. It is merely well begun, and I +report it here in its unfinished state for the sake of the value of the +method which I have worked out, rather than for the purpose of presenting +the definite results which I obtained with No. 51. + +The critical comment which I wish to make for the benefit of those who are +working on similar problems is this. The phosphor bronze wires, on the +bottom of the electric-boxes, by means of which an electric shock could be +given to the mouse when it chose the wrong box, are needless sources of +variability in the illumination of the boxes. They reflect the light into +the eyes of the mouse too strongly, and unless they are kept perfectly +clean and bright, serious inequalities of illumination appear. To avoid +these undesirable conditions I propose hereafter to use a box within a +box, so that the wires shall be hidden from the view of the animal as it +attempts to discriminate. + +A brief description of the behavior of the dancer in the brightness +discrimination experiments which have been described may very +appropriately form the closing section of this chapter. For the +experimenter, the incessant activity and inexhaustible energy of the +animal are a never-failing source of interest and surprise. When a dancer +is inactive in the experiment box, it is a good indication either of +indisposition or of too low a temperature in the room. In no animal with +which I am familiar is activity so much an end in itself as in this odd +species of mouse. With striking facility most of the mice learn to open +the wire swing doors from either side. They push them open with their +noses in the direction in which they were intended by the experimenter to +work, and with almost equal ease they pull them open with their teeth in +the direction in which they were not intended to work. In the rapidity +with which this trick is learned, there are very noticeable individual +differences. The pulling of these doors furnished an excellent opportunity +for the study of the imitative tendency. + +When confronted with the two entrances of the electric-boxes, the dancer +manifested at first only the hesitation caused by being in a strange +place. It did not seem much afraid, and usually did not hesitate long +before entering one of the boxes. The first choice often determined the +majority of the choices of the preference series. If the mouse happened to +enter the left box, it kept on doing so until, having become so accustomed +to its surroundings that it could take time from its strenuous running +from _A_ by way of the left box to the alley and thence to _A_, to examine +things in _B_ a little, it observed the other entrance and in a seemingly +half-curious, half-venturesome way entered it. In the case of other +individuals, the cardboards themselves seemed to determine the choices +from the first. + +The electric shock, as punishment for entering the wrong box, came as a +surprise. At times an individual would persistently attempt to enter, or +even enter and retreat from the wrong box repeatedly, in spite of the +shock. This may have been due in some instances to the effects of fright, +but in others it certainly was due to the strength of the tendency to +follow the course which had been taken most often previously. The next +effect of the shock was to cause the animal to hesitate before the +entrances to the boxes, to run from one to the other, poking its head into +each and peering about cautiously, touching the cardboards at the +entrances, apparently smelling of them, and in every way attempting to +determine which box could be entered safely. I have at times seen a mouse +run from one entrance to the other twenty times before making its choice; +now and then it would start to enter one and, when halfway in, draw back +as if it had been shocked. Possibly merely touching the wires with its +fore paws was responsible for this simulation of a reaction to the shock. +The gradual waning of this inhibition of the forward movement was one of +the most interesting features of the experiment. Could we but discover +what the psychical states and the physiological conditions of the animal +were during this period of choosing, comparative psychology and physiology +would advance by a bound. + +If the conditions at the entrances of the two boxes were discriminable, +the mouse usually learned within one hundred experiences to choose the +right box without much hesitation. Three distinct methods of choice were +exhibited by different individuals, and to a certain extent by the same +individual from time to time. These methods, which I have designated +_choice by affirmation_, _choice by negation_, and _choice by comparison_, +are of peculiar interest to the psychologist and logician. + +When an individual runs directly to the entrance of the right box, and, +after stopping for an instant to examine it, enters, the choice may be +described as recognition of the right box. I call it choice by affirmation +because the act of the animal is equivalent to the judgment--"this is it." +If instead it runs directly to the wrong box, and, after examining it, +turns to the other box and enters without pause for examination, its +behavior may be described as recognition of the wrong box. This I call +choice by negation because the act seems equivalent to the judgment--"this +is not it." Further, it seems to imply the judgment--"therefore the other +is it." In the light of this fact, this type of choice might appropriately +be called choice by exclusion. Finally, when the mouse runs first to one +box and then to the other, and repeats this anywhere from one to fifty +times, the choice may be described as comparison of the boxes; therefore, +I call it choice by comparison. Certain individuals choose first by +comparison, and later almost uniformly by affirmation and negation. +Whenever the conditions are difficult to discriminate, choice by +comparison occurs most frequently and persistently. If, however, the +conditions happen to be absolutely indiscriminable, as was true, for +example, in the case of the sound tests, in certain of the Weber's law +tests, and in the plain electric-box tests, the period of hesitation +rapidly increases during the first three or four series of tests, then the +mouse seems to lessen its efforts to discriminate and more and more tends +to rush into one of the boxes without hesitation or examination, and +apparently with the expectation of a shock, but with the intention of +getting it over as soon as possible. Now and then under such conditions +there is a marked tendency to enter the same box each time. +Indiscriminable conditions are likely to render the animals fearful of the +experiment; instead of going from _A_ to _A_ willingly, they fight against +making the trip. They refuse to pass from _A_ to _B_; and when in _B_, +they fight against being driven toward the entrances to the electric- +boxes. + +In marked contrast with this behavior on the part of the mouse under +conditions which do not permit it to choose correctly is that of the +animal which has learned what is expected of it. The latter, far from +holding back or fighting against the conditions which urge it forward, is +so eager to make the trip that it sometimes has to be forced to wait while +the experimenter records the results of the tests. There is evidence of +delight in the freedom of movement and in the variety of activity which +the experiment furnishes. The thoroughly trained dancer runs into _B_ +almost as soon as it has been placed in _A_ by the experimenter; it +chooses the right entrance by one of the three methods described above, +immediately, or after whirling about a few times in _B_; it runs through +_E_ and back to _A_ as quickly as it can, and almost before the +experimenter has had time to record the result of the choice it is again +in _B_ ready for another choice. + + + + +CHAPTER IX + +THE SENSE OF SIGHT: COLOR VISION + +Is the dancing mouse able to discriminate colors as we do? Does it possess +anything which may properly be called color vision? If so, what is the +nature of its ability in this sense field? Early in my study of the mice I +attempted to answer these and similar questions, for the fact that they +are completely deaf during the whole or the greater part of their lives +suggested to me the query, are they otherwise defective in sense +equipment? In the following account of my study of color vision, I shall +describe the evolution of my methods in addition to stating the results +which were obtained and the conclusions to which they led me. For in this +case the development of a method of research is quite as interesting as +the facts which the method in its various stages of evolution revealed. + +Observation of the behavior of the dancer under natural conditions caused +me to suspect that it is either defective in color vision or possesses a +sense which is very different from human vision. I therefore devised the +following extremely simple method of testing the animal's ability to +distinguish one color from another. In opposite corners of a wooden box 26 +cm. long, 23 cm. wide, and 11 cm. deep, two tin boxes 5 cm. in diameter +and 1.5 cm. deep were placed, as is shown in part I of Figure 18. One of +these boxes was covered on the outside with blue paper (_B_ of Figure 18), +and the other with orange[1] (_O_ of Figure 18). A small quantity of +"force" was placed in the orange box. As the purpose of the test was to +discover whether the animals could learn to go directly to the box which +contained the food, the experiments were made each morning before the mice +had been fed. The experimental procedure consisted in placing the +individual to be tested in the end of the large wooden box opposite the +color boxes, and then permitting it to run about exploring the box until +it found the food in the orange box. While it was busily engaged in eating +a piece of "force" which it had taken from the box and was holding in its +fore paws, squirrel fashion, the color boxes were quickly and without +disturbance shifted in the directions indicated by the arrows of Figure +18, I. Consequently, when the animal was ready for another piece of +"force," the food-box was in the corresponding corner of the opposite end +of the experiment box (position 2, 18, II). After the mouse had again +succeeded in finding it, the orange box was shifted in position as is +indicated by the arrows in Figure 18, II. Thus the tests were continued, +the boxes being shifted after each success on the part of the animal in +such a way that for no two successive tests was the position of the food- +box the same; it occupied successively the positions 1, 2, 3, and 4 of the +figure, and then returned to 1. Each series consisted of 20 tests. + +[Footnote 1: These were the Milton Bradley blue and orange papers.] + +[Illustration: FIGURE 18.--Food-box apparatus for color discrimination +experiments. _O_, orange food-box; _B_, blue food-box; 1, 2, 3, 4, +different positions of the food-boxes, _O_ and _B_; I, II, III, IV, +figures in which the arrows indicate the direction in which the food-boxes +were moved.] + +[Illustration: FIGURE 19.--Food-box apparatus with movable partitions. +_O_, orange food-box; _B_, blue food-box; _X_, starting point for mouse; +_A_, point at which both food-boxes become visible to the mouse as it +approaches them; 1, 2, two different positions of the food-boxes; _T_, +_T_, movable partitions. (After Doctor Waugh.)] + +An improvement on this method, which was suggested by Doctor Karl Waugh, +has been used by him in a study of the sense of vision in the common +mouse. It consisted in the introduction, at the middle of the experiment +box, of two wooden partitions which were pivoted on their mid-vertical +axes so that they could be placed in either of the positions indicated in +Figure 19. Let us suppose that a mouse to be tested for color vision in +this apparatus has been placed at _X_. In order to obtain food it must +pass through _A_ and choose either the orange or the blue box. If it +chooses the former, the test is recorded as correct; if it goes to the +blue box first, and then to the orange, it is counted an error. While the +animal is eating, the experimenter shifts the boxes to position 1 of +Figure 19, and at the same time moves the partitions so that they occupy +the position indicated by the dotted lines. The chief advantage of this +improvement in method is that the animal is forced to approach the color +boxes from a point midway between them, instead of following the sides of +the experiment box, as it is inclined to do, until it happens to come to +the food-box. This renders the test fairer, for presumably the animal has +an opportunity to see both boxes from _A_ and can make its choice at that +point of vantage. + +Two males, A and B, of whose age I am ignorant, were each given seventeen +series of tests in the apparatus of Figure 18. A single series, consisting +of twenty choices, was given daily. Whether the animal chose correctly or +not, it was allowed to get food; that is, if it went first to the blue +box, thus furnishing the condition for a record of error, it was permitted +to pass on to the orange box and take a piece of "force." No attempt was +made to increase the animal's desire for food by starving it. Usually it +sought the food-box eagerly; when it would not do so, the series was +abandoned and work postponed. "Force" proved a very convenient form of +food in these tests. The mice are fond of it, and they quickly learned to +take a flake out of the box instead of trying to get into the box and sit +there eating, for when they attempted the latter they were promptly pushed +to one side by the experimenter and the box, as well as the food, was +removed to a new position. + +The results of the tests appear in Table 15. No record of the choices in +the first two of the 17 series was kept. The totals therefore include 15 +series, or 300 tests, with each individual. Neither the daily records nor +the totals of this table demonstrate choice on the basis of color +discrimination. Either the dancers were not able to tell one box from the +other, or they did not learn to go directly to the orange box. It might be +urged with reason that there is no sufficiently strong motive for the +avoidance of an incorrect choice. A mistake simply means a moment's delay +in finding food, and this is not so serious a matter as stopping to +discriminate. I am inclined, in the light of result of other experiments, +to believe that there is a great deal in this objection to the method. +Reward for a correct choice should be supplemented by some form of +punishment for a mistake. This conclusion was forced upon me by the +results of these preliminary experiments on color vision and by my +observation of the behavior of the animals in the apparatus. At the time +the above tests were made I believed that I had demonstrated the inability +of the dancer to distinguish orange from blue, but now, after two years' +additional work on the subject, I believe instead that the method was +defective. + +The next step in the evolution of a method of testing the dancer's color +vision was the construction of the apparatus (Figures 14 and 15) which was +described in Chapter VII. In connection with this experiment box the basis +for a new motive was introduced, namely, the punishment of mistakes by an +electric shock. Colored cardboards, instead of the white, black, or grays +of the brightness tests, were placed in the electric-boxes. + + + +TABLE 15 + +ORANGE-BLUE TESTS, WITH FOOD-BOX + + + + MOUSE A MOUSE B +SERIES DATE + 1904 RIGHT WRONG RIGHT WRONG + (ORANGE) (BLUE) (ORANGE) (BLUE) + + 1 Dec. 6 -- -- -- -- + 2 7 -- -- -- -- + 3 8 12 8 12 8 + 4 9 10 10 9 11 + 5 10 15 5 10 10 + 6 11 10 10 12 8 + 7 12 9 11 9 11 + 8 13 10 10 9 11 + 9 14 12 8 12 8 + 10 15 13 7 12 8 + 11 16 13 7 10 10 + 12 17 12 8 10 10 + 13 18 11 9 10 10 + 14 19 13 7 8 12 + 15 20 13 7 9 11 + 16 22 14 6 12 8 + 17 23 10 10 9 11 + + TOTALS 177 123 153 147 + + + + +In preliminary tests, at the rate of four per day, the colored cardboards +were placed only at the entrances to the boxes, not inside, and as was +true also in the case of brightness tests under like conditions, no +evidence of discrimination was obtained from ten days' training. This +seemed to indicate that a considerable area of the colored surface should +be exposed to the mouse's view, if discrimination were to be made +reasonably easy. + +This conclusion was supported by the results of other preliminary +experiments in which rectangular pieces of colored papers[1] 6 by 3 cm., +were placed on the floor at the entrances to the electric-boxes, instead +of on the walls of the boxes. Mouse No. 2 was given five series of ten +tests each with a yellow card to indicate the right box and a red card at +the entrance to the wrong box. At first he chose the red almost uniformly, +and at no time during these fifty tests did he exhibit ability to choose +the right box by color discrimination. I present the results of these +series in Table 16, because they indicate a fact to which I shall have to +refer repeatedly later, namely, that the brightness values of different +portions of the spectrum are not the same for the dancer as for us. +Previous to this yellow-red training, No. 2, as a result of ten days of +white-black training (two tests per day), had partially learned to go to +the brighter of the two electric-boxes. It is possible therefore that the +choice of the box in the case of these color experiments was in reality +the choice of what appeared to the mouse to be the brighter box. If this +were not true, how are the results of Table 16 to be accounted for? + +[Footnote 1: These were the only Hering papers used in my experiments.] + +TABLE 16 + +YELLOW-RED TESTS + +In Color Discrimination Box with 6 by 3 cm. Pieces of Hering +Papers at Entrances to Boxes + +No. 2 + + SERIES DATE RIGHT WRONG + 1906 (Yellow) (Red) + 1 Jan. 16 1 9 + 2 17 3 7 + 3 18 4 6 + 4 19 5 5 + 5 20 5 5 + + + +Without further mention of the many experiments which were necessary for +the perfecting of this method of testing color vision, I may at once +present the final results of the tests which were made with reflected +light. These tests were made with the discrimination apparatus in +essentially the same way as were the brightness discrimination tests of +Chapter VII. + +In all of the color experiments, unless otherwise stated, a series of ten +tests each day was given, until satisfactory evidence of discrimination or +proof of the lack of the ability to discriminate had been obtained. The +difficulties of getting conclusive evidence in either direction will be +considered in connection with the results themselves. For all of these +tests with reflected light the Milton Bradley colored papers were used. +These colored papers were pasted on white cardboard carriers. I shall +designate, in the Bradley nomenclature, the papers used in each +experiment. + +With colored cardboards inside the electric-boxes as well as at their +entrances (see Figure 14 for position of cardboards) blue-orange tests +were given to Nos. 2 and 3 until they discriminated perfectly. The papers +were Bradley's blue tint No. 1 and orange. Number 2 was perfect in the +twelfth series (Table 17), No 3 in the fourteenth and again in the +sixteenth. They were then tested with a special brightness check series +which was intended by the experimenter to reveal any dependence upon a +possible brightness difference rather than upon the color difference of +the boxes. + + + +TABLE 17 + +LIGHT BLUE-ORANGE TESTS IN COLOR DISCRIMINATION BOX + +SERIES DATE NO. 2 NO. 3 + 1906 RIGHT WRONG RIGHT WRONG + (LIGHT (ORANGE) (LIGHT (ORANGE) + BLUE) BLUE) + + + 1 Jan. 26 7 3 1 9 + 2 27 7 3 5 5 + 3 28 7 3 6 4 + 4 29 7 3 7 3 + 5 30 7 3 4 6 + 6 31 10 0 7 3 + 7 Feb. 1 9 1 7 3 + 8 2 8 2 6 4 + 9 3 9 1 9 1 + 10 5 7 3 5 5 + 11 6 8 2 5 5 + 12 7 10 0 5 5 +Special brightness check series (see Table 18) + 13 8 10 0 7 3 +Special light blue-dark blue series + 14 9 8 2 10 0 + 15 10 9 1 9 1 +Special light blue-dark blue series + 16 11 9 1 10 0 + Special brightness + check series + 17 12 10 0 9 1 + + + + +TABLE 18 + +LIGHT BLUE-ORANGE + +Brightness check series Mouse No. 2, Series 13 + Feb. 8, 1906 + +TEST CONDITION RIGHT WRONG + +1 Light blue on right + Orange on left Right ____ + +2 Light blue on left + Orange on right Right ____ + +3 Light blue on right + Red substituted for orange Right ____ + +4 Light blur on left + Red substituted for orange Right ____ + +5 Dark blue on right + Orange on left Right ____ + +6 Dark blue on left + Orange on right Right ____ + +7 Dark blue on left + Orange on right Right ____ + +8 Dark blue on right + Red substituted for orange Right ____ + +9 Dark blue on left + Red substituted for orange Right ____ + +10 Dark blue on left + Red substituted for orange Right ____ + + Totals 10 0 + + + +The nature of this brightness check series, as well as the results which +No. 2 gave when tested by it, may be appreciated readily by reference to +Table 18. Tint No. 1 of the blue, which is considerably brighter, in my +judgment, than the Bradley blue, was replaced at intervals in this series +by the latter. For it was thought that in case the mouse were choosing the +blue of the series because it seemed brighter than the orange, this +substitution might mislead it into choosing the orange. These blues are +referred to in the table as light blue (tint No. 1) and dark blue +(standard blue). Again a change in the opposite direction was made by +substituting Bradley red for orange. As this was for the human eye the +substitution of a color whose brightness was considerably less than that +of the orange, it seemed possible that the mouse, if it had formed the +habit of choosing the box which seemed the darker, might by this change be +misled into choosing the red instead of the light blue. In a word, changes +in the conditions of the experiments were made in such a way that now one +color, now the other, appeared to be the brighter. But I did not attempt +to exclude brightness discrimination on the part of the mouse by +dependence upon the human judgment of brightness equality, for it is +manifestly unsafe to assume that two colors which are of the same +brightness for the human eye have a like relation for the eye of the +dancer or of any other animal. My tests of color vision have been +conducted without other reference to human standards of judgment or +comparisons than was necessary for the description of the experimental +conditions. In planning the experiments I assumed neither likeness nor +difference between the human retinal processes and those of the dancer. It +was my purpose to discover the nature of the mouse's visual discriminative +ability. + +As is indicated in the tables, neither the substitution of dark blue for +light blue, nor the replacement of the orange by red or dark blue rendered +correct choice impossible, although certain of the combinations did render +choice extremely difficult. In other words, despite all of the changes +which were made in the brightness of the cardboards in connection with the +light blue-orange tests, the mice continued to make almost perfect +records. What are we to conclude from this? Either that the ability to +discriminate certain colors is possessed by the dancer, or that for some +reason the tests are unsatisfactory. If it be granted that the possibility +of brightness discrimination was excluded in the check series, the first +of these alternatives apparently is forced upon us. That such a +possibility was not excluded, later experiments make perfectly clear. The +fact was that not even in the check series was the brightness value of the +orange as great as that of the blue. Consequently the mice may have chosen +the brighter box each time while apparently choosing the blue. + +Although conclusive proof of the truth of this statement is furnished only +by later experiments, the results of the light blue-orange series, as +given in Table 17, strongly suggest such a possibility. Mouse No. 3 had +not been experimented with previous to these color discrimination tests. +Her preference for the orange, which in the case of the first series was 9 +to 1, consequently demands an explanation. If she had been trained +previously to choose the white instead of the black, as was true of No. 2, +it might be inferred that she went to the orange box because it appeared +brighter than the blue. As this explanation is not available, we are +driven back upon the results of the white-black preference tests in +Chapter VII, which proved that many dancers prefer the black to the white. +This may mean that they prefer the lower degree of brightness or +illumination, and if so it might be argued, in turn, that the orange was +chosen by No. 3 because it appeared darker than the blue. Since, as has +already been stated, the orange was far brighter for me than the blue, +this would also mean that the brightness values of different colors are +not the same for man and mouse. + +Practically the same kind of color tests as those described for Nos. 2 and +3 were given to Nos. 1000 and 5. The results appear in Table 19. These +tests followed upon the formation of a habit to choose white instead of +black (that is, the greater brightness). From the first both No. 1000 and +No. 5 chose the light blue in preference to the orange or the red. It +therefore seems probable that the former was considerably brighter than +the latter. Number 1000, to be sure, was led into three erroneous choices +by the brightness check series (series 7), but, on the other hand, No. 5 +was not at all disturbed in her choices by similar check tests. It seems +natural to conclude from these facts that both of these mice chose the +blue at first because of its relatively greater brightness, and that they +continued to do so for the same reason. In other words, their behavior +indicates that the brightness check tests were valueless because not +enough allowance had been made for the possible differences between the +vision of mouse and man. + + + TABLE 19 +LIGHT BLUE-ORANGE AND DARK BLUE-RED TESTS + No. 1000 No. 5 +SERIES DATE Condition Right Wrong Right Wrong + (Light (Orange (Light (Orange + Blue or or Blue or + Dark Red) Dark Red) + Blue) Blue) + 1 Jan. 25 Blue-red 8 2 10 0 + 2 26 Blue-red or + Light blue-orange 10 0 10 0 + 3 27 Light blue-orange 10 0 5 5 + 4 29 Light blue-orange 9 1 8 2 + 5 30 Light blue-orange 10 0 8 2 + 6 31 Light blue-orange 10 0 10 0 + 7 Feb. 1 Light blue-orange + or Dark blue-red 7 3 10 0 + + +If only the final results of my experiments with the dancer and the +conclusions to which they lead were of interest, all of this description +of experiments which served merely to clear the ground and thus make +possible crucial tests might be omitted. It has seemed to me, however, +that the history of the investigation is valuable, and I am therefore +presenting the evolution of my methods step by step. To be sure, not every +detail of this process can be mentioned, and only a few of the individual +results can be stated, but my purpose will have been fulfilled if I +succeed in showing how one method of experimentation pointed the way to +another, and how one set of results made possible the interpretation of +others. + +As the results of my color vision experiments seemed to indicate that the +red end of the spectrum appears much darker to the dancer than to us, +tests were now arranged with colors from adjacent regions of the spectrum, +green and blue. The papers used were the Bradley green and tint No. 1 of +the blue. They were not noticeably different in brightness for the human +eye. Green marked the box to be chosen. Three of the individuals which had +previously been used in the light blue-orange series, and which therefore +had perfect habits of going to the light blue, were used for the green- +light blue tests. Of these individuals, No. 1000 became inactive on the +fifth day of the experiment, and the tests with him were discontinued. +Twenty series were given to each of the other mice, with the results which +appear in Table 20. To begin with, both No. 4 and No. 5 exhibited a +preference for the light blue, as a result of the previous light blue- +orange training. As this preference was gradually destroyed by the +electric shock which was received each time the light blue box was +entered, they seemed utterly at a loss to know which box to enter. +Occasionally a record of six, seven, or even eight right choices would be +made in a series, but in no case was this unquestionably due to color +discrimination; usually it could be explained in the light of the order of +the changes in the positions of the cardboards. For example, series 9, in +which No. 5 made a record of 8 right and 2 wrong, had green on the right +for the first three tests. The animal happened to choose correctly in the +first test, and continued to do so three times in succession simply +because there was no change in the position of the cardboards. I have +occasionally observed a record of seven right choices result when it was +perfectly evident to the observer that the mouse could not discriminate +visually. It was to avoid unsafe conclusions and unfair comparisons, as +the result of such misleading series, that three perfect series in +succession were required as evidence of a perfectly formed habit of +discrimination. + + +TABLE 20 + +GREEN-LIGHT BLUE TESTS + + Date No. 1000 No. 4 No. 5 +SERIES 1906 RIGHT WRONG RIGHT WRONG RIGHT WRONG + (GREEN) (BLUE) (GREEN) (BLUE) (GREEN) (BLUE) + +1 Feb.3 2 8 3 7 3 7 + +2 5 7 3 5 5 5 5 + +3 6 5 5 6 4 5 5 + +4 7 5 5 5 5 5 5 + +5 8 2 8 5 5 4 6 + +6 9 7 3 7 3 + +7 10 4 6 3 7 + +8 10 6 4 4 6 + +9 12 6 4 8 2 + +10 13 6 4 6 4 + +11 14 5 5 3 7 + +12 15 6 4 7 3 + +13 16 5 5 7 3 + +14 17 3 7 6 4 + +15 19 6 4 6 4 + +16 20 7 3 5 5 + +17 21 4 6 8 2 + +18 22 3 7 4 6 + +19 23 6 4 4 6 + +20 24 6 4 5 5 + + + +Twenty series, 200 tests for each of the individuals in the experiment, +yielded no evidence whatever of the dancer's ability to tell green from +blue. As it has already been proved that they readily learn to choose the +right box under discriminable conditions, it seems reasonable to conclude +either that they lack green-blue vision, or that they have it in a +relatively undeveloped state. + +If it be objected that the number of training tests given was too small, +and that the dancer probably would exhibit discrimination if it were given +1000 instead of 200 tests in such an experiment, I must reply that the +behavior of the animal in the tests is even more satisfactory evidence of +its inability to choose than are the results of Table 20. Had there been +the least indication of improvement as the result of 200 tests, I should +have continued the experiment; as a matter of fact, the mice each day +hesitated more and more before choosing, and fought against being driven +toward the entrance to the experiment box. That they were helpless was so +evident that it would have been manifestly cruel to continue the +experiment. + + + TABLE 21 + VIOLET-RED TESTS + With Odor of All Cardboards the Same + +SERIES DATE NO. 7 NO. 998 + RIGHT WRONG RIGHT WRONG + (VIOLET) (RED) (VIOLET) (RED) + A MAR. 7 8 2 5 5 + B 7 3 7 2 8 + 1 14 3 7 6 4 + 2 15 4 6 4 6 + 3 16 5 5 5 5 + 4 19 4 6 4 6 + 5 20 5 5 6 4 + 6 21 4 6 8 2 + 7 22 8 2 4 6 + 8 23 4 6 6 4 + 9 24 6 4 4 6 + 10 25 4 6 6 4 + + + +Further color tests with reflected light were made with violet and red. +Two dancers, Nos. 998 and 7, neither of which had been in any experiment +previously, were subjected to the ten series of tests whose results are to +be found in Table 21. In this experiment the cardboards used had been +coated with shellac to obviate discrimination by means of odor. It is +therefore impossible to give a precise description of the color or +brightness by referring to the Bradley papers.[1] Both the violet and the +red were rendered darker, and apparently less saturated, by the coating. + +[Footnote 1: The violet was darker than Bradley's shade No. 2, and the red +was lighter than Bradley's red.] + +These violet-red tests were preceded by two series of preference tests +(_A_ and _B_), in which no shock was given and escape was possible through +either electric-box. Although the results of these preference tests as +they appear in Table 21 seem to indicate a preference for the red on the +part of No. 998, examination of the record sheets reveals the fact that +neither animal exhibited color preference, but that instead both chose by +position. Number 998 chose the box on the right 15 times in 20, and No. 7 +chose the box on the left 15 times in 20. + +Ten series of tests with the violet-red cardboards failed to furnish the +least indication of discrimination. The experiment was discontinued +because the mice had ceased to try to discriminate and dashed into one or +the other of the boxes on the chance of guessing correctly. When wrong +they whirled about, rushed out of the red box and into the violet +immediately. They had learned perfectly as much as they were able to learn +of what the experiment required of them. Although we are not justified in +concluding from this experiment that dancers cannot be taught to +distinguish violet from red, there certainly is good ground for the +statement that they do not readily discriminate between these colors. + +The experiments on color vision which have been described and the records +which have been presented will suffice to give the reader an accurate +knowledge of the nature of the results, only a few of which could be +printed, and of the methods by which they were obtained. + +In brief, these results show that the dancer, under the conditions of the +experiments, is not able to tell green from blue, or violet from red. The +evidence of discrimination furnished by the light blue-orange tests is not +satisfactory because the conditions of the experiment did not permit the +use of a sufficiently wide range of brightnesses. It is obvious, +therefore, that a method of experimentation should be devised in which the +experimenter can more fully control the brightness of the colors which he +is using. I shall now describe a method in which this was possible. + + + + +CHAPTER X + +THE SENSE OF SIGHT: COLOR VISION (_Continued_) + +There are three well-known ways in which colors may be used as stimuli in +experiments on animals: by the use of colored papers (reflected light); by +the use of a prism (the spectrum which is obtained may be used as directly +transmitted or as reflected light); and by the use of light filters +(transmitted light). In the experiments on the color vision of the dancer +which have thus far been described only the first of these three methods +has been employed. Its advantages are that it enables the experimenter to +work in a sunlit room, with relatively simple, cheap, and easily +manipulated apparatus. Its chief disadvantages are that the brightness of +the light can neither be regulated nor measured with ease and accuracy. +The use of the second method, which in many respects is the most desirable +of the three, is impracticable for experiments which require as large an +illuminated region as do those with the mouse; I was therefore limited to +the employment of light filters in my further tests of color +discrimination. + +The form of filter which is most conveniently handled is the colored +glass, but unfortunately few glasses which are monochromatic are +manufactured. Almost all of our so-called colored glasses transmit the +light of two or more regions of the spectrum. After making spectroscopic +examinations of all the colored glasses which were available, I decided +that only the ruby glass could be satisfactorily used in my experiments. +With this it was possible to get a pure red. Each of the other colors was +obtained by means of a filter, which consisted of a glass box filled with +a chemical solution which transmitted light of a certain wave length. + +For the tests with transmitted light the apparatus of Figures 20 and 21 +was constructed. It consisted of a reaction-box essentially the same as +that used in the brightness vision tests, except that holes were cut in +the ends of the electric-boxes, at the positions _G and R_ of Figure 20, +to permit the light to enter the boxes. Beyond the reaction-box was a long +light-box which was divided lengthwise into two compartments by a +partition in the middle. A slit in the cover of each of these compartments +carried an incandescent lamp _L_ (Figure 20). Between the two lamps, _L, +L_, and directly over the partition in the light-box was fastened a +millimeter scale, _S_, by means of which the experimenter could determine +the position of the lights with reference to the reaction-box. The light- +box was separated from the reaction-box by a space 6 cm. wide in which +moved a narrow wooden carrier for the filter boxes. This carrier, as shown +in Figure 20, could be moved readily from side to side through a distance +of 20 cm. The filter boxes, which are represented in place in Figures 20 +and 21, consisted of three parallel-sided glass boxes 15 cm. long, 5 cm. +wide, and 15 cm. deep. Each box contained a substance which acted as a ray +filter. Tightly fitted glass covers prevented the entrance of dust and the +evaporation of the solutions in the boxes. Figures 20 and 21 represent the +two end boxes, _R, R_, as red light filters and the middle one, _G_, as a +green light filter. Three filters were used thus side by side in order +that the position of a given color with reference to the electric-boxes +might be changed readily. As the apparatus was arranged, all the +experimenter had to do when he wished to change from green-left, red-right +to green-right, red-left was to push the carrier towards the right until +the green filter covered the hole on the right at the end of the electric- +box. When this had been accomplished the red filter at the left end of the +carrier covered the hole on the left at the end of the electric-box. Thus +quickly, noiselessly, easily, and without introducing any other change in +conditions than that of the interchange of lights, the experimenter was +able to shift the positions of his colored lights at will. + +[Illustration: FIGURE 20.--Color discrimination apparatus. _A,_ nest-box; +_B,_ entrance chamber; _R, R,_ red filters; _G,_ green filter; _L, L,_ +incandescent lamps in light-box; _S,_ millimeter scale on light-box; _I,_ +door between _A_ and _B; O, O,_ doors between alleys and _A_.] + +[Illustration: FIGURE 21--Ground plan of color discrimination apparatus. +_E, E_, exits from electric-boxes. _LB_, light-box; _R, G, R_, filter +boxes on carrier; _L_, left electric-box; _R_, right electric-box; _IC_ +induction apparatus; _C_, electric cell; _K_, key; _S_, millimeter scale.] + +In the tests which are now to be reported, three portions of the spectrum +were used: the red end, the blue-violet end, and a middle region, chiefly +green. The red light was obtained by the use of a filter which was made by +placing two plates of ruby glass in one of the glass boxes, filling the +box with filtered water and then sealing it to prevent evaporation. The +blue-violet was obtained by the use of a filter box which contained a 5 +per cent solution of copper ammonium sulphate. The green, which, however, +was not monochromatic, was obtained by the use of a filter box which +contained a saturated solution of nickel nitrate. These three sets of +filters were examined spectroscopically both before the experiments had +been made and after their completion.[1] The red filters, of which I had +two for shifting the lights, transmitted only red light. The blue-violet +filters, two also, at first appeared to transmit only portions of the blue +and violet of the spectrum, but my later examination revealed a trace of +green. It is important to note, however, that the red and the blue-violet +filters were mutually exclusive in the portions of the spectrum which they +transmitted. Of all the filters used the green finally proved the least +satisfactory. I detected some yellow and blue in addition to green in my +first examination, and later I discovered a trace of red. Apparently the +transmitting power of the solutions changed slightly during the course of +the experiments. On this account certain solutions are undesirable for +experiments on color vision, for one must be certain of the constancy of +the condition of stimulation. It is to be understood, of course, that each +of the three filters transmitted, so far as the eye is concerned, only the +color named. I consider the red filter perfectly satisfactory, the blue- +violet very good, and the green poor. Henceforth, in testing color vision +in animals, I shall make use of colored glasses as filters, if it is in +any way possible to obtain or have manufactured blue, green, and yellow +glasses which are as satisfactory as the ruby. + +[Footnote 1: A Janssen-Hoffman spectroscope was used.] + +The apparatus needs no further description, as its other important +features were identical with those of the reflected light experiment box. +The use of artificial light for the illumination of the electric-boxes +made it necessary to conduct all of the following tests in a dark-room. +The method of experimentation was practically the same as that already +described. A mouse which had been placed in _A_ by the experimenter was +permitted to enter _B_ and thence to return to _A_ by entering one of the +electric-boxes, the red or blue or green one, as the case might be. +Mistakes in choice were punished by an electric shock. One further point +in the method demands description and discussion before the results of the +tests are considered, namely, the manner of regulating and measuring the +brightness of the lights. + +Regulating brightness with this apparatus was easy enough; measuring it +accurately was extremely difficult. The experimenter was able to control +the brightness of each of the two colored lights which he was using by +changing the position or the power of the incandescent lamps in the light- +box. The position of a lamp could be changed easily between tests simply +by moving it along toward or away from the electric-box in the slit which +served as a lamp carrier. As the distance from the entrances of the +electric-boxes to the further end of the light-box was 120 cm., a +considerable range or variation in brightness was possible without change +of lamps. Ordinarily it was not necessary to change the power of the +lamps, by replacing one of a given candle power by a higher or lower, +during a series of tests. Both the candle power of the lamps and their +distance from the filters were recorded in the case of each test, but for +the convenience of the reader I have reduced these measurements to candle +meters[1] and report them thus in the descriptions of the experiments. + +[Footnote 1: The illuminating power of a standard candle at a distance of +one meter.] + +But measuring the actual brightness of the red light or the green light +which was used for a particular series of tests, and the variations in +their brightnesses, was not so simple a matter as might appear from the +statements which have just been made. The influence of the light filters +themselves upon the brightness must be taken into account. The two red +filters were alike in their influence upon the light which entered them, +for they were precisely alike in construction, and the same was true of +the two blue-violet filters. The same kind of ruby glass was placed in +each of the former, and a portion of the same solution of copper ammonium +sulphate was put into each of the filter boxes for the latter. But it is +difficult to say what relation the diminution in brightness caused by a +red filter bore to that caused by a blue-violet or a green filter. My only +means of comparison was my eye, and as subjective measurement was +unsatisfactory for the purposes of the experiment, no attempt was made to +equalize the amounts of brightness reduction caused by the several +filters. So far as the value of the tests themselves, as indications of +the condition of color vision in the dancer is concerned, I have no +apology for this lack of measurement, but I do regret my inability to give +that accurate objective statement of brightness values which would enable +another experimenter with ease and certainty to repeat my tests. The +nearest approach that it is possible for me to make to such an objective +measurement is a statement of the composition and thickness of the filters +and of the candle-meter value of the light when it entered the filter. The +distance from this point to the entrance to the electric-box was 20 cm. + +To sum up and state clearly the method of defining the brightness of the +light in the following experiments: the candle-meter value of each light +by which an electric-box was illuminated, as determined by the use of a +Lummer-Brodhun photometer and measurements of the distance of the source +of light from the filter, is given in connection with each of the +experiments. This brightness value less the diminution caused by the +passage of the light through a filter, which has been defined as to +composition and thickness of the layer of solution, gives that degree of +brightness by which the electric-box was illuminated. + +Tests of the dancer's ability to discriminate green and blue[1] in the +transmitted light apparatus were made with four animals. An incandescent +lamp marked 16-candle-power was set in each of the light-boxes. These +lamps were then so placed that the green and the blue seemed to be of +equal brightness to three persons who were asked to compare them +carefully. Their candle-meter values in the positions selected were +respectively 18 and 64, as appears from the statement of conditions at the +top of Table 22. + +[Footnote 1: Hereafter the light transmitted by the blue-violet filter +will be referred to for convenience as blue.] + + + + TABLE 22 + + GREEN-BLUE TESTS + + Brightnesses Equal for Human Eye + +Green 18 candle meters Blue 64 candle meters + +SERIES DATE NO. 10 NO. 11 + 1906 RIGHT WRONG RIGHT WRONG + (GREEN) (BLUE) (GREEN) (BLUE) +A and B[1] April 2 10 10 12 8 + 1 3 6 4 5 5 + 2 4 5 5 6 4 + 3 5 5 5 5 5 + 4 6 5 5 5 5 + 5 7 7 3 5 5 + 6 8 7 3 3 7 + 7 9 7 3 5 5 + 8 10 3 7 7 3 + 9 11 5 5 4 6 + 10 12 5 5 6 4 +[Footnote: A single preference series of twenty tests.] + + + +Numbers 10 and 11 exhibited no preference for either of these colors in +the series of 20 tests which preceded the training tests, and neither of +them gave evidence of ability to discriminate as the result of ten series +of training tests. In this case, again, the behavior of the animals was as +strongly against the inference that they can tell green from blue as are +the records of choices which appear in the table. Granted, that they are +unable to discriminate green from blue when these colors are of about the +same brightness for the human eye, what results when they differ markedly +in brightness? Table 23 furnishes a definite answer to this question. +Numbers 5 and 12 were given eight series of green-blue tests with each +light at 18 candle meters. Little, if any, evidence of discrimination +appeared. Then, on the supposition that the difference was not great +enough for easy discrimination, the blue light was reduced almost to 0, +the green being left at 18. The tests (series 9) immediately indicated +discrimination. For series 10 the green was made 64 candle meters, the +blue 18, and again there was discrimination. These results were so +conclusively indicative of the lack of color vision and the presence of +brightness vision, that there appeared to be no need of continuing the +experiment further. + +Accepting provisionally the conclusion that the dancers cannot tell green +from blue except by brightness differences, we may proceed to inquire +whether they can discriminate other colors. Are green and red +distinguishable? + +Green-red discrimination now was tested by a method which it was hoped +might from the first prevent dependence upon brightness. The light in the +light-box on the left was so placed that it had a value of 18 candle +meters, that in the light-box on the right so that it had a value of 1800 +candle meters. Neither light was moved during the first four series of the +green-red tests which were given to Nos. 151 and 152. + + + +TABLE 23 + +GREEN-BLUE TESTS + +Brightnesses Different for Human Eye + +Green 18 candle meters Blue 18 candle meters + + + No. 5 No. 12 + DATE +SERIES 1906 RIGHT WRONG RIGHT WRONG + (GREEN) (BLUE) (GREEN) (BLUE) + + 1 April 10 6 4 5 5 + 2 11 5 5 7 3 + 3 12 6 4 7 3 + 4 13 4 6 7 3 + 5 14 7 3 5 5 + 6 15 4 6 6 4 + 7 16 6 4 8 2 + 8 17 5 5 4 6 + + + +As it was now evident that the intensity difference was not sufficient to +render discrimination easy, the blue was reduced to 0 and the green left +at 18. + + + 9 17 7 3 8 2 + + +Now the brightnesses were made, green 64, blue 18, just the reverse of +those of series of Table 22. + + + 10 17 8 2 8 2 + + + +Each of these series consisted of 20 tests instead of 10. As a result of +the arrangement of the lights just mentioned, the green appeared to me +very much brighter than the red when it was on the right and very much +darker when it was on the left. If this were true for the mouse also, it +is difficult to see how it could successfully depend upon brightness for +guidance in its choices. Such dependence would cause it to choose now the +green, now the red. + +The first four series of green-red tests so clearly demonstrated +discrimination, of some sort, that it was at once necessary to alter the +conditions of the experiment. The only criticism of the above method of +excluding brightness discrimination, of which I could think, was that the +red at no time had been brighter than the green. In other words, that +despite a value of 1800 candle meters for the red and only 18 candle +meters for the green, the latter still appeared the brighter to the mouse. +To meet this objection, I made the extreme brightness values 1 and 1800 +candle meters in some of the later series, of which the results appear in +Table 24. From day to day different degrees of brightness were used, as is +indicated in the second column of the table. Instead of having first one +color and then the other the brighter, after the fourth series I changed +the position of the lights each time the position of the filters was +changed; hence, the table states a certain brightness value for each color +instead of for each electric-box. + +Series 5 to 14 so clearly indicated discrimination, that it seemed +necessary to devise some other means than that of changing the +brightnesses of the colored lights themselves to test the assumption that +the animals were choosing the brighter light. I therefore removed the +light filters so that the colors which had been present as conditions of +discrimination were lacking, and arranged the apparatus so that first one +box, then the other, was illuminated the more brightly. The purpose of +this was to discover whether as the result of their green-red training the +mice had acquired the habit of choosing uniformly either the lighter or +the darker box. One series was given under the conditions of illumination +specified in Table 24 with the result that the brighter box was chosen +eight times in ten by No. 151 and every time by No. 152. Since neither of +these individuals had previously been trained by white-black tests to go +to the white, and since, furthermore, the dancers usually manifest a +slight preference for the lower instead of the higher illumination, this +result may be interpreted as indicative of dependence upon brightness in +the previous color tests. It looks very much indeed as if the green had +been chosen, not because of its greenness, but on account of its +relatively greater brightness. + +This test of brightness preference was followed by two series, 16 and 17, +under conditions similar to those of the first four series of the table. +For series 16 the value of the light in the left box was 1 candle meter, +that of the light in the right box 1800 candle meters. Discrimination was +perfect. For series 17 the value for the left remained at 1 candle meter, +but that of the right box was decreased to 0. In this series No. 152 was +entirely at a loss to know which box to choose. Of course this was an +entirely new set of conditions for choice, namely, a colored box, the +green or the red as the case might be, beside a dark box, the one which +was not illuminated. If the mice really had been choosing correctly +because of a habit of avoiding the red or of seeking the green, this +method should bring out the fact, for the red box, since with it the +disagreeable electric shock had always been associated, should be a box to +be avoided. For No. 151 this seemed to be the case. + +Series 23 to 27 of Table 24 were given as final and crucial tests of the +relation of brightness discrimination to color discrimination. As it is +not possible to express in a simple formula the conditions of the tests, a +sample series which indicates the brightness of the colors in each of the +twenty tests of a series, and in addition the results given by No. 151 in +the first of these final series, is reproduced in Table 25. For an animal +which had presumably learned perfectly to choose green in preference to +red, the record of 8 mistakes in 20 choices as a result of changes in +relative brightness is rather bad, and it renders doubtful the existence +of color discrimination in any of these experiments. No. 152 showed no +ability whatever to choose the green in the first of the series (series 23 +of Table 24) of which that of Table 25 is a sample. His record, 10 +mistakes in 20 choices, was even poorer than that of No. 151. That both of +these mice learned to choose fairly accurately in these final tests is +shown by the results of series 24, 25, 26, and 27. I must admit, however, +that these records indicate little ability on the part of the animals to +discriminate colors. + + + +TABLE 24 + +GREEN-RED TESTS + +Brightnesses Extremely Different for Human Eye +Intensities are given in candle meters (c.m.) + + + NO. 151 NO. 152 +SERIES DATE CONDITIONS + RIGHT WRONG RIGHT WRONG + (GREEN) (RED) (GREEN) (RED) + + 1 April 26 18 c.m. on left + 1800 c.m. on right 11 9 7 13 + 2 27 Same 16 4 16 4 + 3 28 Same 20 0 17 3 + 4 29 Same 19 1 19 1 + 5 30 Green 18 c.m. + Red 18 c.m. 9 1 10 0 + 6 30 Green 64 c.m. + Red 18 c.m. 9 1 8 2 + 7 May 1 Green 6 c.m. + Red 1500 c.m. 7 3 9 1 + 8 1 Green 4 c.m. + Red 1500 c.m. 8 2 7 3 + 9 2 Both varied from + 4 to 1500 c.m. 18 2 18 2 + 10 3 Green 2 c.m. + Red 1800 c.m. 6 4 7 3 + 11 3 Same 10 0 10 0 + 12 4 Same 7 3 8 2 + 13 4 Same 8 2 6 4 + 14 5 Green 1 c.m. + Red 1800 c.m. 19 1 19 1 + + + +Filters were now removed. An illumination of 15 c.m. was established on +one side and an illumination of 0 on the other side, in order to ascertain +whether the mice would choose the brighter box. This was done to test the +assumption that the green in the previous tests had always appeared +brighter to the mice than did the red, and that in consequence they had +chosen the brighter box instead of the green box. + + +TABLE 24--CONTINUED + + + No. 151 No. 152 + +SERIES DATE CONDITIONS RIGHT WRONG RIGHT WRONG + (GREEN) (RED) (GREEN) (RED) + + 15 May 5 Brighter 15 c.m. 8[1] 2[2] 10[1] 0[2] + Darker 0 c.m. + 16 5 1 c.m. on left + 1800 c.m. on right 10 0 10 0 + 17 5 1 c.m. on left + 0 c.m. on right 9 1 4 6 + 18 5 Green 18 c.m. + Red 18 c.m. 19 1 17 3 + 19 9 Same 9 1 9 1 + 20 9 Same 10 0 10 0 + 21 10 Same 10 0 10 0 + 22 11 Same 10 0 10 0 + 23 June 1 Both varied from + 1 to 1800 c.m. 12 8 10 10 + 24 2 Same 18 2 14 6 + 25 June 3 Both varied from + 2 to 1800 c.m. 19 1 17 3 + 26 4 Same 17 3 17 3 + 27 5 Same 18 2 18 2 + +[Footnote 1: Brighter.] +[Footnote 2: Darker.] + + + +These long-continued and varied tests with Nos. 151 and 152 revealed three +facts: that the mice depend chiefly upon brightness differences in visual +discrimination; that they probably have something which corresponds to our +red-green vision, although their color experience may be totally unlike +ours; and that the red end of the spectrum seems much darker to them than +to us, or, in other words, that the least refrangible rays are of lower +stimulating value for them than for us. + + + +TABLE 25 + +GREEN-RED TESTS + +June 1, 1906 No. 151 + + BRIGHTNESS VALUE IN CANDLE RIGHT WRONG +TEST POSITION METERS (GREEN) (RED) + 1 Green on left Green 4, Red 448 Right -- + 2 Green on right Green 448, Red 4 Right -- + 3 Green on right Green 4, Red 448 Right -- + 4 Green on left Green 448, Red 4 Right -- + 5 Green on left Green 3, Red 1800 -- Wrong + 6 Green on right Green 1800, Red 3 -- Wrong + 7 Green on right Green 3, Red 1800 -- Wrong + 8 Green on left Green 1800, Red 3 Right -- + 9 Green on right Green 5, Red 34 Right -- + 10 Green on left Green 34, Red 5 Right -- + 11 Green on right Green 6, Red 74 Right -- + 12 Green on left Green 74, Red 6 Right -- + 13 Green on left Green 4, Red 448 -- Wrong + 14 Green on right Green 448, Red 4 Right -- + 15 Green on right Green 4, Red 448 -- Wrong + 16 Green on left Green 448, Red 4 Right -- + 17 Green on right Green 3, Red 1800 -- Wrong + 18 Green on left Green 1800, Red 3 -- Wrong + 19 Green on right Green 1800, Red 3 -- Wrong + 20 Green on left Green 3, Red 1800 Right -- + + Totals 12 8 + + + +So many of the results of my color experiments have indicated the all- +important role of brightness vision that I have hesitated to interpret any +of them as indicative of true color discrimination. But after I had made +all the variations in brightness by which it seemed reasonable to suppose +that the mouse would be influenced under ordinary conditions, and after I +had introduced all the check tests which seemed worth while, there still +remained so large a proportion of correct choices that I was forced to +admit the influence of the quality as well as of the intensity of the +visual stimulus. + +The first of the facts mentioned above, that brightness discrimination is +more important in the life of the mouse than color discrimination, is +attested by almost all of the experiments whose results have been +reported. The second fact, namely, that the dancer possesses something +which for the present we may call red-green vision, also has been proved +in a fairly satisfactory manner by both the reflected and the transmitted +light experiments. I wish now to present, in Table 26, results which +strikingly prove the truth of the statement that red appears darker to the +dancer than to us. + +The brightness conditions which appeared to make the discrimination +between green and red most difficult were, so far as my experiments permit +the measurement thereof, green from 1 to 4 candle meters with red from +1200 to 1600. Under these conditions the red appeared extremely bright, +the green very dark, to the human subject. + +According to the description of conditions in Table 26, Nos. 2 and 5 were +required to distinguish green from red with the former about 3 candle +meters in brightness and the latter about 1800 candle meters. In the +eighth series of 20 tests, each of these animals made a perfect record. As +it seemed possible that they had learned to go to the darker of the two +boxes instead of to the green box, I arranged the following check test. +The filters were removed, the illumination of one electric-box was made 74 +candle meters, that of the other 3, and the changes of the lighter box +from left to right were made at irregular intervals. In February, No. 2 +had been trained to go to the black in black-white tests, and at the same +time No. 5 had been trained to go to the white in white-black tests. The +results of these brightness check tests, as they appear in the table, +series 8 _a_, are indeed striking. Number 2 chose the darker box each +time; No. 5 chose it eight times out of ten. Were it not for the fact that +memory tests four weeks after his black-white training had proved that No. +2 had entirely lost the influence of his previous experience (he chose +white nine times out of ten in the memory series), it might reasonably be +urged that this individual chose the darker box because of his experience +in the black-white experiment. And what can be said in explanation of the +choices of No. 5? I can think of no more reasonable way of accounting for +this most unexpected result of the brightness tests than the assumption +that both of these animals had learned to discriminate by brightness +difference instead of by color. + + + +TABLE 26 + +GREEN-RED TESTS + +Brightnesses Different for Human Eye + + No. 2 No. 5 + +SERIES DATE BRIGHTNESS RIGHT WRONG RIGHT WRONG + VALUES (GREEN) (RED) (GREEN) (RED) + + 1 May 7 Green + Red 1800 c.m. 10 10 12 8 + 2 8 Same 12 8 11 9 + 3 9 Same 15 5 14 6 + 4 10 Same 18 2 12 8 + 5 11 Same 18 2 14 6 + 6 12 Same 19 1 16 4 + 7 13 Same 19 1 18 2 + 8 14 Same 20 0 20 0 + + +Brightness tests without colors were now given to determine whether the +mice had been choosing the brighter or the darker instead of the green. + + +TABLE 26--CONTINUED + + + NO. 2 NO. 5 + + SERIES DATE BRIGHTNESS VALUES RIGHT WRONG RIGHT WRONG + (GREEN) (RED) (GREEN) (RED) + + 8a 14 Brighter 74 c.m. 0[1] 10[2] 2[1] 8[2] + Darker 3 c.m. + 9 15 3 c.m. on left + 1800 c.m. on right 8 12 16 4 + 10 16 4 c.m. on left + 36 c.m. on right 5 5 7 3 + 11 16 Green 4 c.m. + Red 36 c.m. 9 1 8 2 + 12 17 11 c.m. on left + 1800 c.m. on right 7 3 6 4 + 13 17 Green 11 c.m. + Red 1800 c.m. 9 1 8 2 + 14 18 Mixed values + 3 to 1800 c.m. 7 3 8 2 + 15 19 Same 7 3 7 3 + 16 20 Same 7 3 7 3 + 17 21 Same 7 3 9 1 + 18 22 Same 9 1 8 2 + 19 23 Same 7 3 9 1 + 20 24 Same 10 0 8 2 + 21 25 Same 10 0 9 1 + 22 26 Same 9 1 10 0 + + +[Footnote 1: Brighter] +[Footnote 2: Darker] + + + +Immediately after the brightness series, the influence of making first one +color, then the other, the brighter was studied. Throughout series 9 the +brightness value of the left box remained 3 candle meters, that of the +right side 1800 candle meters. Number 2 was so badly confused by this +change that his mistakes in this series numbered 12; No. 5 made only 4 +incorrect choices. Then series after series was given under widely +differing conditions of illumination. The expression "mixed values," which +occurs in Table 26 in connection with series 14 to 22 inclusive, means +that the brightnesses of the green and the red boxes were changed from +test to test in much the way indicated by the sample series of Table 25. +In view of the results of these 22 series, 320 tests for each of two mice, +it is evident that the dancer is able to discriminate visually by some +other factor than brightness. What this factor is I am not prepared to +say. It may be something akin to our color experience, it may be distance +effect. No other possibilities occur to me. + +Table 26 shows that discrimination was relatively easy for Nos. 2 and 5 +with green at 3 candle meters and red at 1800. That their discrimination +was made on the basis of the greater brightness of the red, instead of on +the basis of color, is indicated by the results of the brightness check +series 8a. Increase in the brightness of the green rendered discrimination +difficult for a time, but it soon improved, and by no changes in the +relative brightness of the two colors was it possible to prevent correct +choice. + +In addition to giving point to the statement that red appears darker to +the dancer than to us, the above experiment shows that the animals depend +upon brightness when they can, and that their ability to discriminate +color differences is extremely poor, so poor indeed that it is doubtful +whether their records are better than those of a totally color blind +person would be under similar conditions. Surely in view of such results +it is unsafe to claim that the dancer possesses color vision similar to +ours. + +Perfectly trained as they were, by their prolonged green-red tests, to +choose the green, or what in mouse experience corresponds to our green, +Nos. 2 and 5 offered an excellent opportunity for further tests of blue- +green discrimination. For in view of their previous training there should +be no question of preference for the blue or of a tendency to depend upon +brightness in the series whose results constitute Table 27. + + + TABLE 27 + BLUE-GREEN TESTS + + NO. 2 NO. 5 + +SERIES DATE BRIGHTNESS VALUES RIGHT WRONG RIGHT WRONG + (BLUE) (GREEN) (BLUE) (GREEN) + + 1 June 1 Blue 74 c.m. + Green 36 c.m. 3 7 3 7 + 2 2 Same 5 5 4 6 + 3 3 Same 5 5 6 4 + 4 4 Same 6 4 3 7 + 5 5 Same 6 4 5 5 + 6 6 Blue 21 c.m. + Green 21 c.m. 6 4 7 3 + 7 7 Same 2 8 3 7 + 8 8 Same 5 5 4 6 + 9 9 Same 3 7 6 4 + 10 10 Same 2 8 4 6 + 11 12 Same 6 4 3 7 + 12 13 Blue 36 c.m. + Green 21 c.m. 3 7 4 6 + 13 14 Same 5 5 + 14 15 Blue 62 c.m. + Green 21 c.m. 4 6 + 15 16 Same 5 5 + 16 17 Same 5 5 + 17 18 Same 6 4 + + + + +Now, as a final test, blue and green glasses were placed over the +electric-boxes, the brightness of the two was equalized for the human eye, +and the tests of series 18 and 19 were given to No. 2:-- + + +TABLE 27--CONTINUED + + NO. 2 +SERIES DATE BRIGHTNESS VALUES + RIGHT WRONG + (Blue) (Green) + + 18 18 Blue 62 c.m. + Green 21 c.m 4 6 + 19 19 Same 6 4 + 20 20 Blue 21 c.m. + Green 88 c.m. 2 8 + + +The green was now made much the brighter. + + + 21 21 Blue 21 c.m. + Green 18 c.m. 7 3 + 22 23 Same 8 2 + + +To begin with, the blue and the green were made quite bright for the human +subject, blue 74 candle meters, green 36. Later the brightness of both was +first decreased, then increased, in order to ascertain whether +discrimination was conditioned by the absolute strength of illumination. +No evidence of discrimination was obtained with any of the several +conditions of illumination in seventeen series of ten tests each. + +On the supposition that the animals were blinded by the brightness of the +light which had been used in some of the tests, similar tests were made +with weaker light. The results were the same. I am therefore convinced +that the animals did justice to their visual ability in these experiments. + +Finally, it seemed possible that looking directly at the source of light +might be an unfavorable condition for color discrimination, and that a +chamber flooded with colored light from above and from one end would prove +more satisfactory. To test this conjecture two thicknesses of blue glass +were placed over one electric-box, two plates of green glass over the +other; the incandescent lamps were then fixed in such positions that the +blue and the green within the two boxes appeared to the experimenter, as +he viewed them from the position at which the mouse made its choice, of +the same brightness. + +Mouse No. 2 was given two series of tests, series 18 and 19, under these +conditions, with the result that he showed absolutely no ability to tell +the blue box from the green box. The opportunity was now taken to +determine how quickly No. 2 would avail himself of any possibility of +discriminating by means of brightness. With the blue at 21 candle meters, +the green was increased to about 1800. Immediately discrimination +appeared, and in the second series (22 of Table 27) there were only two +mistakes. + +The results of the blue-green experiments with light transmitted from in +front of the animal and from above it are in entire agreement with those +of the experiments in which reflected light was used. Since the range of +intensities of illumination was sufficiently great to exclude the +possibility of blinding and of under illumination, it is necessary to +conclude that the dancer does not possess blue-green vision. + +Again I must call attention to the fact that the behavior of the mice in +these experiments is even more significant of their lack of discriminating +ability than are the numerical results of the tables. After almost every +series of tests, whether or not it came out numerically in favor of +discrimination, I was forced to add the comment, "No satisfactory evidence +of discrimination." + +We have now examined the results of green-red, green-blue, and blue-green +tests. One other important combination of the colors which were used in +these experiments is possible, namely, blue-red. This is the most +important of all the combinations in view of the results already +described, for these colors represent the extremes of the visible +spectrum, and might therefore be discriminable, even though those which +are nearer together in the spectral series were not. + + +TABLE 28 +BLUE-RED TESTS + + + + No. 2 No. 205 +SERIES DATE BRIGHTNESS VALUES + RIGHT WRONG RIGHT WRONG + (BLUE) (RED) (BLUE) (RED) + +1 July 31 1800 c.m. on left + 24 c.m. on right 5 5 6 4 +2 Aug. 1 21 c.m. on left + 1800 c.m. on right 6 4 6 4 +3 2 1800 c.m. on left + 21 c.m. on right 8 2 6 4 +4 3 19 c.m. on left + 1800 c.m. on right 9 1 6 4 +5 4 1800 c.m. on left + 7 c.m. on right 7 3 5 5 +6 5 6 c.m. on left + 1800 c.m. on right 10 0 7 3 +7 6 18 c.m. on left + 74 c.m. on right 10 0 9 1 +8 7 1800 c.m. on left + 7 c.m. on right 8 2 8 2 +9 8 7 c.m. on left + 1800 c.m. on right 7 3 8 2 +10 9 Mixed values + 6 to 1800 c.m. 8 2 9 1 +11 10 Blue 3 c.m. + Red 1800 c.m. 7 3 6 4 + + +Brightness tests were now made, without the use of colors. + +11a 10 4 6 5 5 + +12 10 Blue 3 c.m. + Red 8 c.m. 4 6 6 4 +13 11 Blue 3 c.m. + Red 7200 c.m. 8 2 5 5 +14 13 Mixed values + 3 to 7200 c.m. 7 3 7 3 +15 13 Same 7 3 9 1 +16 14 Blue 3 to 6 c.m. + Red 112 to 3650 c.m. 10 0 10 0 + + +Series were now given to test the assumption that red appears dark to the +dancer. + + + +17 14 Darkness on one side + Red 3 c.m. 5 5 7 3 +18 14 Blue 3 to 3650 c.m. + Red 3 to 3650 c.m. 10 0 10 0 +19 15 Darkness on one side + Red 3 c.m. 5 5 4 6 +20 15 Blue 3 to 3650 c.m. + Red 3 to 3650 c.m. 10 0 9 1 +21 16 Darkess on one side + Red 72 c.m. 5 5 7 3 +22 16 Darkness on one side + Red 1800 c.m. 6 4 10 0 + + +As is shown by the results in Table 28, no combination of brightnesses +rendered correct choice impossible in the case of the blue-red tests which +are now to be described. Choice was extremely difficult at times, even +more so perhaps than the table would lead one to suppose, and it is quite +possible that color played no part in the discrimination. But that +brightness difference in the colors was not responsible for whatever +success these mice attained in selecting the right box is proved by the +brightness-without-color series which follows series II of the table. +Neither No. 2 nor No. 205 showed preference for the lighter or the darker +box. At the end of the sixteenth blue-red series, I was convinced that one +of two conclusions must be drawn from the experiment: either the dancers +possess a kind of blue-red vision, or red is of such a value for them that +no brightness of visible green or blue precisely matches it. + +The latter possibility was further tested by an experiment whose results +appear in series 17 to 22 inclusive, of Table 28. The conditions of series +17 were a brightness value of 0 in one box (darkness) and in the other red +of a brightness of 3 candle meters. Despite the fact that they had been +perfectly trained in _blue-red tests_ to avoid the red, neither of the +mice seemed able to discriminate the red from the darkness and to avoid +it. This was followed by a series in which the brightness of both the blue +and the red was varied between 3 and 3650 candle meters, with the striking +result that neither mouse made any mistakes. In series 19 red was used +with darkness as in series 17, and again there was a total lack of +discrimination. Series 20 was a repetition of series 18, with practically +the same result. I then attempted to find out, by increasing the +brightness of the red, how great must be its value in order that the +dancers should distinguish it readily from darkness. For the tests of +series 21 it was made 72 candle meters, but discrimination did not clearly +appear. At 1800 candle meters, as is shown in series 22, the red was +sufficiently different in appearance from total darkness to enable No. 205 +to discriminate perfectly between the two electric-boxes. For No. 2 +discrimination was more difficult, but there was no doubt about his +ability. It would appear from these tests that the dancers had not learned +to avoid red. Therefore we are still confronted with the question, can +they see colors? + + + +TABLE 29 + +VISUAL CHECK TESTS +With the Electric-boxes Precisely Alike Visually + + + No. 151 No. 152 +SERIES DATE + RIGHT WRONG RIGHT WRONG + +1 Sept. 29 6 4 4 6 +2 30 5 5 6 6 +3 Oct. 1 3 7 4 6 +4 2 5 5 3 7 +5 3 3 7 5 5 +6 4 6 4 5 5 +7 5 5 5 5 5 +8 6 -- -- 3 7 +9 7 -- -- 6 4 +10 8 -- -- 4 6 + +Averages 4.7 5.3 4.5 5.5 + + + +The account of my color vision experiments is finished. If it be objected +that other than visual conditions may account for whatever measure of +discriminating ability, apart from brightness discrimination, appears in +some of the series, the results of the series of Table 29, in which all +conceivable visual means of discrimination were purposely excluded, and +those of the several check tests which have been described from time to +time in the foregoing account, should furnish a satisfactory and definite +answer. I am satisfied that whatever discrimination occurred was due to +vision; whether we are justified in calling it color vision is quite +another question. + +I conclude from my experimental study of vision that although the dancer +does not possess a color sense like ours, it probably discriminates the +colors of the red end of the spectrum from those of other regions by +difference in the stimulating value of light of different wave lengths, +that such specific stimulating value is radically different in nature from +the value of different wave lengths for the human eye, and that the red of +the spectrum has a very low stimulating value for the dancer. In the light +of these experiments we may safely conclude that many, if not most, of the +tests of color vision in animals which have been made heretofore by other +investigators have failed to touch the real problem because the +possibility of brightness discrimination was not excluded. + +Under the direction of Professor G. H. Parker, Doctor Karl Waugh has +examined the structure of the retina of the dancing mouse for me, with the +result that only a single type of retinal element was discovered. +Apparently the animals possess rod-like cells, but nothing closely similar +to the cones of the typical mammalian retina. This is of peculiar interest +and importance in connection with the results which I have reported in the +foregoing pages, because the rods are supposed to have to do with +brightness or luminosity vision and the cones with color vision. In fact, +it is usually supposed that the absence of cones in the mammalian retina +indicates the lack of color vision. That this inference of functional +facts from structural conditions is correct I am by no means certain, but +at any rate all of the experiments which I have made to determine the +visual ability of the dancer go to show that color vision, if it exists at +all, is extremely poor. It is gratifying indeed to learn, after such a +study of behavior as has just been described, that the structural +conditions, so far as we are able to judge at present, justify the +conclusions which have been drawn. + + + + +CHAPTER XI + +THE ROLE OF SIGHT IN THE DAILY LIFE OF THE DANCER + +Darting hither and thither in its cage, whirling rapidly, now to the left, +now to the right, running in circles, passing through holes in the nest +box quickly and neatly, the dancer, it would seem, must have excellent +sight. But careful observation of its behavior modifies this inference. +For it appears that a pair of mice dancing together, or near one another, +sometimes collide, and that it is only those holes with which the animal +is familiar that are entered skillfully. In fact, the longer one observes +the behavior of the dancer under natural conditions, the more he comes to +believe in the importance of touch, and motor tendencies. Sight, which at +first appears to be the chief guiding sense, comes to take a secondary +place. In this chapter it is my purpose to show by means of simple +experiments what part sight plays in the dancer's life of habit formation. + +The evidence on this subject has been obtained from four sources: (1) +observation of the behavior of dancers in their cages; (2) observation of +their behavior when blinded; (3) observation of their behavior in a great +variety of discrimination experiments, many of which have already been +described; and (4) observation of their behavior in labyrinth experiments +which were especially planned to exhibit the importance of the several +kinds of vision which the dancer might be supposed to possess. The +evidence from the first three of these sources may be presented summarily, +for much of it has already appeared in earlier chapters. That from the +fourth source will constitute the bulk of the material of this chapter. + +My observation of the behavior of the mice has furnished conclusive +evidence of their ability to see moving objects. But that they do not see +very distinctly, and that they do not have accurate perception of the form +of objects, are conclusions which are supported by observations that I +have made under both natural and experimental conditions. In Chapters VII, +VIII, IX, and X, I have presented an abundance of evidence of brightness +vision and, in addition, indications of a specific sensitiveness to wave +length which may be said to correspond to our color vision. It is +noteworthy, however, that all of the experimental proofs of visual ability +were obtained as the result of long periods of training. Seldom, indeed, +in my experience with them, have the dancers under natural conditions +exhibited forms of activity which were unquestionably guided by vision. + +It is claimed by those who have experimented with blinded dancers that the +loss of sight decreases the amount and rapidity of movement, and the +ability of the animals to avoid obstacles. + +By means of the discrimination method previously used in the preliminary +experiments on color vision, a full description of which may be found in +Chapter IX, p. 133, the dancers' ability to perceive form was tested. +Immediately after the two males _A_ and _B_ had been given the "food-box" +tests, whose results appear in Table 15, they were tested in the same +apparatus and by the same method for their ability to discriminate a +rectangular food-box from a round one. In the case of the color +discrimination tests, it will be remembered that the circular tin boxes 5 +cm. in diameter by 1.5 cm. in depth, one of which was covered with blue +paper, the other with orange, were used. For the form discrimination tests +I used instead one of the circular boxes of the dimensions given above and +a rectangular box 8.5 cm. long, 5.5 cm. wide and 2.5 cm. deep. "Force" was +placed in the circular box. The tests were given, in series of 20, daily. + + + +TABLE 30 + +VISUAL FORM TESTS + +SERIES DATE MOUSE A MOUSE B + RIGHT WRONG RIGHT WRONG + (CIRCULAR (RECTANGU- (CIRCULAR (RECTANGU- + BOX) LAR BOX) BOX) LAR BOX) + 1 Jan. 5 10 10 9 11 + 2 7 12 8 13 7 + 3 10 6 14 10 10 + 4 11 7 13 10 10 + 5 12 9 11 10 10 + 6 13 11 9 11 9 + 7 14 13 7 9 11 + 8 15 10 10 11 9 + 9 16 10 10 11 9 + 10 17 11 9 9 11 + 11 18 11 9 12 8 + 12 19 12 8 10 10 + 13 20 10 10 12 8 + 14 21 10 10 8 12 + 15 22 10 10 10 10 + + Totals 152 148 155 145 + + + +The results of 15 series of these tests, as may be seen by the examination +of Table 30, are about as definitely negative, so far as form +discrimination is in question, as they possibly could be. From the first +series to the last there is not one which justifies the inference that +either of the dancers depended upon the form of the boxes in making its +choice. In view of the general criticisms I have made concerning the use +of hunger as a motive in experiments on animal behavior, and in view of +the particular criticisms of this very method of testing the +discriminating powers of the mouse, it may seem strange that space should +be given to a report of these tests. I sympathize with the feeling, if any +one has it, but, at the same time, I wish to call attention to the fact +that almost any mammal which is capable of profiting by experience, and +which, under the same conditions, could distinguish the rectangular box +from the circular one, would have chosen the right box with increasing +accuracy as the result of such experience. The results are important in my +opinion, not because they either prove or disprove the ability of the +dancer to discriminate these particular forms, the discrimination of which +might fairly be expected of any animal with an image-forming eye, but +because they demonstrate an important characteristic of the dancing mouse, +namely, its indifference to the straightforward or direct way of doing +things. + +Most mammals which have been experimentally studied have proved their +eagerness and ability to learn the shortest, quickest, and simplest route +to food without the additional spur of punishment for wandering. With the +dancer it is different. It is content to be moving; whether the movement +carries it directly towards the food is of secondary importance. On its +way to the food-box, no matter whether the box be slightly or strikingly +different from its companion box, the dancer may go by way of the wrong +box, may take a few turns, cut some figure-eights, or even spin like a top +for seconds almost within vibrissa-reach of the food-box, and all this +even though it be very hungry. Activity is pre-eminently important in the +dancer's life. + +In passing I may emphasize the importance of the fact that at no time did +the brightness or color discrimination tests furnish evidence of attempts +on the part of the dancers to choose by means of slight differences in the +form of the cardboards or the cardboard carriers. Several times form +differences, which were easily perceivable by the human subject, were +introduced in order to discover whether the mice would detect them and +learn to discriminate thereby instead of by the visual conditions of +brightness or color. As these experiments failed to furnish evidences of +form discrimination, the following special test in the discrimination box +was devised. + +[Illustration: FIGURE 22.--Cards used for tests of form discrimination.] + +The color discrimination box of Chapter X was arranged so that the light +at the entrance to each electric-box had a value of 20 candle meters, less +the diminution caused by a piece of ground glass which was placed over the +end of the electric-boxes to diffuse the light. The windows through which +the light entered the electric-boxes were covered with pieces of black +cardboard; in one of these cardboards I had cut a circular opening 4 cm. +in diameter, and in the other an opening of the same area but markedly +different shape. These openings are shown in Figure 22. As the mouse +approached the entrance to the electric-boxes, it was confronted by these +two equally illuminated areas, whose chief difference was one of form. +Difference in the amount of light within the boxes was excluded so far as +possible. The question which I asked was, can the dancer discriminate by +means of this difference in visual form? + +For the purpose of settling this point and of gaining additional knowledge +of the role of vision, two individuals were tested in the discrimination +box under the conditions which have just been described. During the first +ten days of the experiment each of these mice, Nos. 420 and 425, was given +a series of ten tests daily. At the end of this period experimentation +with No. 425 had to be discontinued, and the number of daily tests given +to No. 420 was increased to twenty. + +Instead of taking space for the presentation of the daily records, I may +state the general results of the tests. Neither of the mice learned to +choose the right box by means of form discrimination. In fact, there was +absolutely no sign of discrimination at any time during the tests. This +result is as surprising as it is interesting. I could not at first believe +that the mice were unable to perceive the difference in the lighted areas, +but assumed that they were prevented from getting the outlines of the +areas by the blinding effect of the light. However, decreasing the +intensity of the illumination did not alter the result. According to the +indications of this experiment, the dancer's ability to perceive visual +form is extremely poor. + +Thus far the purpose of our experiments has been to ascertain what the +dancer is enabled to do by sight. Suppose we now approach the problem of +the role of this sense by trying to find out what it can do without sight. + +[Illustration: FIGURE 23.--Labyrinth B. _I_, entrance; _O_, exit; 1, 2, 3, +doorways between alleys.] + +For the investigation of this matter the labyrinth method seemed eminently +suitable. The first form of labyrinth which was used in these visual tests +appears in ground plan in Figure 23. It was made of 1-1/2 cm. boards. The +length was 52 cm., the width 17 cm., the depth 10 cm. Each of the +doorways, _I_ (the entrance), 1, 2, 3, and _O_ (the exit), was 5 by 5 cm. +The alleys were 2-1/2 cm. wide. For this width the necessity is obvious +from what has already been said of the animal's propensity to whirl on all +occasions. As the mice almost never tried to climb up the walls, no cover +for the labyrinth was needed. The direct route is indicated by the symbols +_I_-1-2-3-_O_. If an error be defined as a choice of the wrong path as the +animal progressed toward the exit, five mistakes were possible in the +forward course: the first by turning to the left at the entrance; the +second by failing to pass through doorway 1; the third by turning to the +right after passing through doorway 1; the fourth by failing to pass +through doorway 3, and the fifth by turning to the left after passing +through 3. In case the mouse retraced its course, any mistakes made as it +again progressed towards _O_ were counted, as at first, no matter how many +times it went over the same ground. Thus an individual might make the same +mistake several times in the course of a single test in the labyrinth. + +With this labyrinth Nos. 7, 998, 15, 16, 151, and 152 were tested. At +first a record was kept of the time which elapsed from the instant the +animal entered _I_ to the instant it emerged at _O_, of the path which it +followed, and of the number of errors which it made; but later only the +number of errors was recorded. + + +TABLE 31 + +THE ROLE OF SIGHT + +Labyrinth-B Experiments + + + NO. 7 NO. 998 + +TEST DATE TIME ERRORS TIME ERRORS + 1 June 16 66" 8 127" 19 + 2 16 11 0 94 12 + 3 16 15 2 18 3 + 4 16 7 0 13 2 + 5 16 5 0 10 1 + 6 18 61 15 12 3 + 7 18 13 3 14 4 + 8 18 14 5 8 1 + 9 18 24 9 16 2 + 10 18 10 1 9 1 + 11 19 36 13 80 17 + 12 19 8 3 10 1 + 13 19 6 1 7 1 + 14 19 9 1 8 0 + 15 19 12 2 7 0 + 16 20 14 1 25 0 + 17 20 28 3 + 18 20 No efforts No efforts + to escape to escape + + + + + + TABLE 32 + + LABYRINTH-B EXPERIMENTS + + with + +Electric Shock given as Punishment for Mistakes + + No. 7 No. 998 +TEST DATE CONDITION ERRORS CONDITION ERRORS + + +1 June 29 Light 4 Light 9 +2 29 Light 1 Light 3 +3 29 Light 1 Light 2 +4 29 Light 0 Light 0 +5 29 Light 0 Light 0 +6 29 Light 0 Light 0 +7 29 Light 1 Light 0 +8 29 Light 0 Light 0 +9 29 Light 1 Darkness 0 +10 29 Light 1 Light 0 +11 29 Light 1 Darkness 0 +12 29 Light 0 Light 0 +13 29 Light 0 Light 0 +14 29 Light 0 Light 0 +15 29 Light 0 Light 0 +16 29 Light 0 Light 0 +17 29 Darkness 2 Darkness 0 +18 29 Light 2 Light 0 + with paper +19 29 Light 0 Light 0 +20 29 Darkness 0 Light 0 + with paper +21 29 Light 0 Light 0 +22 29 Light 0 Darkness 0 +23 29 Light 0 Odorless 0 +24 June 29 Light 0 Darkness 0 +25 29 Light 0 +26 29 Darkness 4 +27 29 Light with + paper 1 +28 29 Light 0 +29 29 Light with + paper 1 +30 29 Darkness 0 +31 29 Odorless 2 +32 29 Darkness 4 + + + + +As the results in Table 31 show, the time and number of errors rapidly +diminished. Number 7, for example, made no errors in the second test. The +chiefly significant fact which appeared in these preliminary experiments, +however, was that the mice soon ceased to care whether they got out of the +labyrinth or not. After they knew the path perfectly, they would enter the +wrong passages repeatedly and wander about indefinitely. It was obvious, +therefore, that the labyrinth could not be used to reveal the role of +sight unless some sufficiently strong motive for continuous effort to +escape from it could be discovered. Naturally I looked to the electric +shock for aid. + +The labyrinth of Figure 23, which for convenience in distinguishing it +from several other forms to be described later I have designated as +labyrinth B, was placed upon a board 90 cm. long and 30 cm. wide about +which had been wound two pieces of phosphor bronze wire after the manner +described on p. 94. At _O_, Figure 24, there was an opening closed by a +swinging door which led into a box 40 by 24 cm. In one corner of this box +was a small nest-box. The significance of this rearrangement of the +labyrinth is apparent. As in the preliminary tests, the dancer was started +at I, but instead of being allowed to wander about without any other +result than delay in escape, it was given a shock each time it made an +error. The satisfaction of escaping from the narrow bounds of the +labyrinth's passages, which alone was not strong enough to impel a dancer +constantly to do its best to escape, was thus supplemented by the powerful +and all-controlling tendency to avoid the disagreeable stimulus which +resulted from entering certain of the passages. The result of this +modification of method is strikingly exhibited by the data of Table 32. + +[Illustration: Figure 24.--Labyrinth B on an interrupted circuit board. +_I_-1-2-3-_O_, labyrinth path; _B_, nest-box; _N_, nest; _EW_, board wound +with phosphor bronze wire; _IC_, induction apparatus; _C_ electric cell; +_K_, key.] + +This table was constructed for the purpose of exhibiting the principal +features of the results obtained with labyrinth _B_ in certain preliminary +experiments in which the conditions were changed in various ways. Chief +among the important facts which appear in the illustrative data (for Nos. +7 and 998) which are presented, are the following. The dancers readily +learn the path of labyrinth B so that they can follow it quickly and with +perfect accuracy. After familiarity with the direct path from entrance to +exit has been gained, they become indifferent about escaping and tend to +wander aimlessly. The introduction of the electric shock as punishment for +the choice of the wrong passage impels them to do their best to avoid +errors. The path once learned can be followed in total darkness with few +or no errors. Table 32 indicates marked differences in the behavior of No. +7 and No. 998. The latter learned the path readily and was little +disturbed by any of the changes in conditions. In total darkness he +followed the path rapidly and accurately, as was indicated by the time of +the trip and the path that he left on a sheet of smoked paper that had +been placed on the floor of the labyrinth as a means of obtaining a record +of the errors made. The presence of the smoked paper did not seem to +interfere at all with his behavior, nor did the thorough washing of the +labyrinth and the resultant removal of its odors. In the case of No. 7 the +opposite was true. She did not learn the path readily, was confused by any +change in conditions, had great difficulty in finding her way in darkness, +made errors when the smoked paper was placed on the floor and after the +odors of the labyrinth had been removed by washing. Of the six dancers +which were observed in these preliminary tests, No. 7 alone gave +convincing evidence of the importance of sight. + +I think we may say in the light of the results of the table that such +errors as appear in the darkness tests are due rather to the disturbing +influence of a change in the conditions of the experiment than to the +exclusion of visual data, for as many or more errors were sometimes caused +simply by changing the position of the labyrinth, placing smoked paper on +the floor, or by introducing a new odor at some point. The exclusion of +the possibility of guidance by smell and touch did not seriously interfere +with the animal's ability to follow the path. + +The results which have just been considered seemed to be of sufficient +interest and importance to justify the further use of the labyrinth method +in the investigation of the role of vision. A series of experiments with +labyrinth B was therefore planned so that the importance of sight, touch, +and smell in connection with this form of habit should be more +satisfactorily exhibited. Does the dancer follow the path by sight, touch, +smell, by all, or by no one of them? + +This series of tests with labyrinth B, whose several purposes may best be +explained in connection with the various kinds of tests enumerated below, +consisted of: + +I. A preliminary test in which the dancer was permitted to wander about in +the labyrinth, without being shocked, until it finally escaped to the +nest-box by way of the exit. Thus the animal was given an opportunity to +discover that escape from the maze was possible. + +II. This was immediately followed by a series of tests at the rate of +about one per minute, with an electric shock as punishment for every +mistake. This was continued without interruption until the path had been +followed without error five times in succession. + +III. The labyrinth was now moved about 3 cm. to one side so that it +covered a new floor area, and a test was given for the purpose of +ascertaining whether the mouse had been following a trail on the floor. + +IV. Tests with smoked paper on the floor were now alternated with tests in +which the floor was plain. The alternation was rendered necessary by the +fact that the paper was laid over the electric wires and therefore +prevented the punishment of mistakes. The purpose of these tests was to +discover whether the smoked paper, which was an essential condition for +the next test, was itself a disturbing condition. These tests were +continued until the animal had followed the path correctly, despite the +smoked paper, twice in succession. + +V. The electric lights were now turned out and tests were given in total +darkness, with smoked paper on the floor as a means of obtaining a record +of the number of errors. These tests were continued until the path had +been followed once correctly. + +VI. The labyrinth was now thoroughly washed with warm water, to which a +little kerosene had been added, and quickly dried over a steam radiator. +This usually necessitated a delay of about five minutes. As soon as the +labyrinth was dry, tests were given to discover whether the odors of the +various passages had been serving as important guiding conditions. These +tests were continued until the path had been followed once without error. + +VII. A final test in darkness completed the series. + +As it was not possible for the observer to watch the animal and thus to +count the number of mistakes which it made in total darkness, the simple +method of placing a piece of smoked paper on the floor of the labyrinth +was used. The mouse left a graphic record of its path on the paper and +from this the number of errors could be ascertained. In the tests now to +be described the smoked paper was placed upon the electric wires, but +later a form of electric labyrinth was devised in which it was underneath +and therefore did not interfere with the electric shock. + +The above series of tests was given under the same external conditions in +a dark-room to six pairs of dancers. In all cases, two individuals, a male +and a female, which had been kept in the same cage, were experimented with +at the same time, _i.e._ one was permitted to rest in the nest-box while +the other was being put through a test. This was done in order that the +comparison of the results for males and females should be perfectly fair. + +The detailed results of this long series of tests may be presented for +only two individuals, Nos. 210 and 215, Table 33. In this table lines +separate the results of the seven different kinds of tests. + + +TABLE 33 + +THE ROLE OF SIGHT, TOUCH, AND SMELL IN LABYRINTH EXPERIMENTS + + + No. 210 No. 215 + + TEST CONDITION ERRORS CONDITION ERRORS + + I. 1 No shock 9 I. No shock 2 + +II. 2 Shock 5 II. Shock 3 + 3 Shock 4 Shock 1 + 4 Shock 2 Shock 0 + 5 Shock 3 Shock 0 + 6 Shock 0 Shock 0 + 7 Shock 0 Shock 0 + 8 Shock 0 Shock 0 + + 9 Shock 0 III. Labyrinth 0 + moved + + 10 Shock 0 IV. Paper on floor 4 + +III. 11 Labyrinth 0 No paper (shock) 0 + moved + +IV. 12 Paper on 0 0 + floor + 13 No paper 0 No paper 0 + (shock) + 14 Paper 1 Paper 1 + 15 No paper 0 No paper 0 + 16 Paper 7 Paper 4 + 17 No paper 0 No paper 0 + 18 Paper 0 Paper 0 + 19 No paper 0 No paper 0 + 20 Paper 4 Paper 0 + 21 No paper 0 No paper 0 + 22 Paper 2 V. Darkness 0 + 23 No paper 2 VI. Labyrinth 2 + 24 Paper 1 washed 0 + 25 No paper 0 VII. Darkness 2 + 26 Paper 0 + 27 No paper 0 + 28 Paper 0 + 29 No paper 0 + V. 30 Darkness 0 + VI. 31 Labyrinth 2 + washed + 32 0 +VII. 33 Darkness 0 + + + + +The average results for the twelve individuals (six of each sex) which +were subjected to the tests, I have arranged in Table 34. The Roman +numerals at the top of the table designate the seven groups of tests, and +the figures under each, the numerical results of the tests. I may explain +and comment upon the averages of the several columns of this table in +turn. + +Column I gives the number of errors made in the preliminary test. +Curiously enough, the males made many more errors than the females. + +For the second group of tests (II) two results have been tabulated: the +number of the first correct test, and the total number of tests before the +path was followed correctly five times in succession. The first correct +trip came usually after not more than five or six tests, but five +successive correct trips demanded on the average at least fourteen +training tests. + +Destruction of the floor path by movement of the labyrinth to one side, +without changing its relations to the points of the compass, disturbed the +mice very little. Only four of the twelve individuals made any mistakes as +a result of the change in the tactual conditions, and the average error as +it appears in Column III is only .3. + + +TABLE 34 + +ROLE OF SIGHT, TOUCH, AND SMELL IN LABYRINTH EXPERIMENTS + + + II. IV. + TRAINING TESTS SMOKED + I. NO OF TESTS BEFORE III. PAPER ON +MALES PRELIMINARY CORRECT LABYRINTH FLOOR + TEST. _____________________ MOVED. NO OF TIMES + ERRORS FIRST TIME FIVE TIMES ERRORS BEFORE COR- + RECT TWICE + + 210 9 5 9 0 9 + 212 2 3 8 1 3 + 214 6 10 28 0 22 + 220 25 4 8 0 14 + 410 11 6 20 0 10 + 420 14 6 14 1 7 + +AVERAGES 11.2 5.7 14.5 .3 10.8 + +FEMALES + + 211 16 6 10 1 5 + 213 7 5 14 1 21 + 215 2 3 7 0 6 + 225 14 6 18 0 14 + 415 6 6 13 0 3 + 425 10 7 13 0 8 + +AVERAGES 9.2 5.5 12.5 .3 9.5 + + + + V. + DARKNESS VI. + MALES LABYRINTH VII. + ERRORS IN NO. OF TESTS WASHED. DARKNESS. + FIRST TEST BEFORE COR'CT ERRORS ERRORS + + 210 0 1 2 0 + 212 2 2 0 0 + 214 0 1 -- 0 + 220 2 4 2 0 + 410 1 3 2 1 + 420 2 4 1 4 + +Averages 1.2 2.5 1.2 0.8 + +FEMALES + + 211 2 2 0 0 + 213 2 2 -- 3 + 215 0 1 2 2 + 225 3 2 0 0 + 415 1 3 2 1 + 425 1 7 0 0 + +Averages 1.5 2.8 0.7 1.0 + + + + +That covering the floor with smoked paper forced the mice to relearn the +path, in large measure, is evident from the results of Column IV. An +average of ten tests was necessary to enable the mice to follow the path +correctly. It is almost certain, however, that the interference with the +perfectly formed labyrinth habit which this change in the condition of the +floor caused was not due to the removal of important tactual sense data. + +As Column V shows, the number of errors in total darkness is very small. +Some individuals gave no sign of being disturbed by the absence of visual +guidance, others at first seemed confused. I have given in the table the +number of errors in the first darkness test and the number of the first +test in which no mistakes occurred. + +No more disturbance of the dancer's ability to follow the path which it +had learned resulted from washing the labyrinth thoroughly than from +darkening the room. Indeed it is clear from Column VI that the path was +not followed by the use of smell. However, the test in darkness, after the +odor of the box had been removed, proved conclusively that in most cases +the mice could follow the path correctly without visual or olfactory +guidance. + +The behavior of 18 individuals as it was observed in labyrinth B makes +perfectly evident three important facts, (1) In following the path which +it has learned, the dancer in most instances is not guided to any +considerable extent by a trail (odor or touch) which has been formed by +its previous journeys over the route; (2) sight is quite unnecessary for +the easy and perfect execution of the labyrinth habit, for even those +individuals which are at first confused by the darkening of the experiment +room are able after a few tests to follow the path correctly; (3) and, +finally, smell, which according to current opinion is the chiefly +important sense of mice and rats, is not needful for the performance of +this habitual act. + +At this point we may very fittingly ask, what sense data are necessary for +the guidance of the series of acts which constitutes the labyrinth habit? +I answer, probably none. A habit once formed, the senses have done their +part; henceforth it is a motor process, whose initiation is conditioned by +the activity of a receptive organ (at times a sense receptor), but whose +form is not necessarily dependent upon immediate impressions from eye, +nose, vibrissae, or even from internal receptors. These are statements of +my opinion; whether they express the truth, either wholly or in part, only +further experimentation can decide. + +In considering the results of these labyrinth tests it is important that +we distinguish clearly those which have to do with the conditions of habit +formation from those which instead have to do with the conditions of habit +performance. Sense data which are absolutely necessary for the learning of +a labyrinth path may be of little or no importance for the execution of +the act of following the path after the learning process has been +completed. Thus far in connection with the labyrinth tests we have +discussed only the relations of sight, touch, and smell to what I have +called habit performance. We may now ask what part these senses play in +the formation of a labyrinth habit. + +A very definite answer to this question is furnished by observation of the +behavior of the dancers in the tests. Most of them continuously made use +of their eyes, their noses, and their vibrissae. Some individuals used one +form of receptive organ almost exclusively. I frequently noticed that +those individuals which touched and smelled of the labyrinth passages most +carefully gave least evidence of the use of sight. It is safe to say, +then, that under ordinary conditions habit formation in the dancer is +conditioned by the use of sight, touch, and smell, but that these senses +are of extremely different degrees of importance in different individuals. +And further, that, although in the case of some individuals the loss of +sight would not noticeably delay habit formation, in the case of others it +would seriously interfere with the process. When deprived of one sense, +the dancer depends upon its remaining channels of communication with +environment. Indeed there are many reasons for inferring that if deprived +of sight, touch, and smell it would still be able to learn a labyrinth +path; and there are reasonable grounds for the belief that a habit once +formed can be executed in the absence of all special sense data. +Apparently the various receptive organs of the body furnish the dancer +with impressions which serve as guides to action and facilitate habit +formation, although they are not necessary for habit performance. + +The reader may wonder why I have not carried out systematic experiments to +determine accurately and quantitatively the part which each sense plays in +the formation of a labyrinth habit instead of basing my inferences upon +incidental observation of the behavior of the dancers. The reason is +simply this: the number and variety of experiments which were suggested by +the several directions in which this investigation developed rendered the +performance of all of them impossible. I have chosen to devote my time to +other lines of experimentation because a very thorough study of the +conditions of habit formation has recently been made by Doctor Watson.[1] + +[Footnote 1: Watson, J. B., _Psychological Review_, Monograph Supplement, +Vol. 8, No. 2, 1907.] + +What is the role of sight in the dancing mouse? How shall we answer the +question? The evidence which has been obtained in the course of my study +of the animal indicates that brightness vision is fairly acute, that color +vision is poor, that although form is not clearly perceived, movement is +readily perceived. My observations under natural conditions justify the +conclusion that sight is not of very great importance in the daily life of +the dancer, and my observations under experimental conditions strongly +suggest the further conclusion that movement and changes in brightness are +the only visual conditions which to any considerable extent control the +activity of the animal. + + + + +CHAPTER XII + +EDUCABILITY: METHODS OF LEARNING + +Nearly all of the experiments described in earlier chapters have revealed +facts concerning the educability of the dancer. In order to supplement the +knowledge of this subject thus incidentally gained and to discover the +principles of educability, the specially devised experiments whose results +appear in this and succeeding chapters were arranged and carried out with +a large number of mice. In the work on the modifiability of behavior I +have attempted to determine (1) by what methods the dancer is capable of +profiting by experience, (2) the degree of rapidity of learning, (3) the +permanency of changes wrought in behavior, (4) the effect of one kind of +training upon others, (5) the relation of re-training to training, and (6) +the relation of all these matters to age, sex, and individuality. + +As it is obvious that knowledge of these subjects is a necessary condition +for the intelligent appreciation of the capacities of an animal, as well +as of the choice of methods by which it may be trained advantageously, +perhaps it is not too much to expect that this investigation of the nature +and conditions of educability in the dancing mouse may give us some new +insight into the significance of certain aspects of human education and +may serve to suggest ways in which we may measure and increase the +efficiency of our educational methods. + +Merely for the sake of convenience of description I shall classify the +methods which have been employed as problem methods, labyrinth methods, +and discrimination methods. That these names are not wholly appropriate is +suggested by the fact that discrimination necessarily occurs in connection +with each of them. As problem methods we may designate those tests of +initiative and modifiability which involve the opening of doors by pushing +or pulling them, and the climbing of an inclined ladder. An example of the +labyrinth method has been presented in Chapter XI. The name discrimination +method I have applied to those tests which involve the choice of one of +two visual, tactual, or olfactory conditions. The white-black +discrimination tests, for example, served to reveal the rapidity and +permanency of learning as well as the presence of brightness vision. + +In the case of most mammals whose educability has been studied +experimentally, problem methods have proved to be excellent tests of +docility and initiative. The cat, the raccoon, the monkey, in their +attempts to obtain food, learn to pull strings, turn buttons, press +latches, slide bolts, pull plugs, step on levers. The dancer does none of +these things readily. Are we therefore to infer that it is less +intelligent, that it is less docile, than the cat, the raccoon, or the +monkey? Not necessarily, for it is possible that these methods do not suit +the capacity of the animal. As a matter of fact, all of the tests which +are now to be described in their relation to the educability of the dancer +bear witness to the importance of the selection of methods in the light of +the motor equipment and the habits of the animal which is to be tested. +Judged by ordinary standards, on the basis of results which it yields in +problem and labyrinth tests, the dancer is extremely stupid. But that this +conclusion is not justified is apparent when it is judged in the light of +tests which are especially adapted to its peculiarities. + +Problems which are easy for other mammals because of their energetic and +persistent efforts to secure food in any way which their motor capacity +makes possible are useless as tests of the dancer's abilities, because it +is not accustomed to obtain its food as the result of strenuous and varied +activities. There are problems and problems; a condition or situation +which presents a problem to one organism may utterly lack interest for an +organism of different structure and behavior. What is a problem test in +the case of the cat or even of the common mouse, is not necessarily a +problem for the dancer. Similarly, in connection with the labyrinth +method, it is clear that the value of the test depends upon the desire of +the organism to escape from the maze. The cat, the rat, the tortoise do +their best to escape; the dancer is indifferent. Clearly, then, methods of +training should be chosen on the basis of a knowledge of the +characteristics of the animal whose educability is to be investigated. + +The simplest possible test of the intelligence of the dancer which I could +devise was the following. Beside the cage in which the mice were kept I +placed a wooden box 26 cm. long, 23 cm. wide, and 12 cm. deep. Neither +this box nor the cage was covered, for the animals did not attempt to +climb out. As a way of passing from one of these boxes to the other I +arranged a ladder made of wire fly-screen netting. This ladder was about 8 +cm. broad and it extended from the middle of one side of the wooden box +upward at an angle of about 30° to the edge of the box and then descended +at the same angle into the cage. + +A dancer when taken from the nest-box and placed in the wooden box could +return to its cage and thus find warmth, food, and company by climbing the +ladder. It was my aim to determine, by means of this apparatus, whether +the dancers can learn such a simple way of escape and whether they learn +by watching one another. As it turned out, a third value belonged to the +tests, in that they were used also to test the influence of putting the +mice through the act. + +In the first experiment three dancers, Nos. 1000, 2, and 6, were together +placed in the wooden box. At the end of 15 minutes not one of them had +succeeded in returning to the cage. They were then driven to the bottom of +the ladder and started upward by the experimenter; with this assistance +all escaped to the nest-box. At the expiration of 5 minutes they were +again placed in the wooden box, whence the chilly temperature (about 60° +F.) and the lack of food made them eager to return to their cage. No +attempt to climb up the ladder was made by any of them within 15 minutes, +so the experimenter directed them to the ladder and started them upward as +in the first test. This completed the experiment for the day. The +following day two tests were given in the same way. In the second of these +tests, that is, on its fourth trial, No. 1000 climbed over of his own +initiative in 5 minutes. The others had to be assisted as formerly. On the +third day No. 1000 found his way back to the nest-box quickly and fairly +directly, but neither No. 2 or No. 6 climbed of its own initiative in the +first test. When their movements were restricted to the region of the box +about the base of the ladder, both of them returned to the cage quickly. +And on the second test of the third day all the mice climbed the ladder +directly. + +In Table 35 I have given the time required for escape in the case of 40 +tests which were given to these 3 individuals at the rate of 2 tests per +day. + +When the time exceeded 15 minutes the mice were helped out by the +experimenter; a record of 15 minutes, therefore, indicates failure. +Naturally enough the motives for escape were not sufficiently strong or +constant to bring about the most rapid learning of which the dancer is +capable. Sometimes they would remain in the wooden box washing themselves +for several minutes before attempting to find a way of escape. On this +account I made it a rule to begin the time record with the appearance of +active running about. The daily average time of escape as indicated in the +table does not decrease regularly and rapidly. On the fourth day, which +was the first on which all three of the dancers returned to the cage by +way of the ladder of their own initiative in both tests, the average is +214 seconds. In contrast with this, on the twentieth day the time was only +5 seconds. It is quite evident that the dancers had learned to climb the +ladder. + +At the end of the twentieth day the experiment was discontinued with Nos. +2 and 6, and after two weeks they were given memory tests, which showed +that they remembered perfectly the ladder-climbing act, for when placed in +the wooden box, with Nos. 4 and 5 as controls, they returned to the cage +by way of the ladder immediately and directly. + + + +TABLE 35 + +LADDER CLIMBING TEST + +Time in Minutes and Seconds + + +No. of Date No. 1000 No. 2 No. 6 Average Daily Av. + Exp. 1905 For All For All + + 1 Nov. 14 15' 15' 15' -- -- + 2 15' 15' 15' -- -- + + 3 15 15' 15' 15' -- -- + 4 300" 15' 15' -- -- + + 5 16 480" 15' 15' -- -- + 6 180" 300" 420" 300" 300" + + 7 17 450" 240" 540" 410" + 8 20" 15" 18" 18" 214" + + 9 18 90" 180" 135" 135" + 10 135" 105" 165" 135" 135" + + 11 19 480" 240" 330" 350" + 12 30" 120" 90" 80" 143" + + 13 20 360" 75" 120" 185" + 14 5" 6" 8" 6" 95" + + 15 21 105" 450" 120" 192" + 16 8" 80" 20" 54" 123" + + 17 22 255" 300" 180" 245" + 18 10" 30" 270" 103" 174" + + 19 23 300" 660" 450" 470" + 20 90" 120" 150" 120" 295" + + 21 24 240" 125" 225" 197" + 22 4" 6" 168" 59" 128" + + 23 Nov. 25 305" 85" 130" 173" + 24 5" 6" 118" 43" 108" + + 25 26 3" 8" 44" 18" + 26 19" 1" 176" 98" 58" + + 27 27 150" 79" 269" 166" + 28 26" 3" 31" 20" 93" + + 29 28 214" 18" 267" 166" + 30 40" 3" 4" 16" 91" + + 31 29 130" 45" 250" 142" + 32 12" 3" 25" 13" 77" + + 33 Dec. 2 61" 35" 44" 47" + 34 50" 5" 24" 26" 36" + + 35 3 66" 18" 2" 29" + 36 8" 5" 10" 8" 19" + + 37 4 9" 4" 3" 5" + 38 10" 5" 6" 7" 6" + + 39 5 5" 3" 5" 4" + 40 10" 4" 3" 6" 5" + + + + +One of the most interesting and important features of the behavior of the +dancer in the ladder experiment was a halt at a certain point on the +ladder. It occurred just at the edge of the wooden box at the point where +the ladder took a horizontal position, and led over into the cage. Every +individual from the first test to the last made this halt. Although from +the point of view of the experimenter the act was valueless, it may have +originated as an attempt to find a way to escape from the uncomfortable +position in which the animal found itself on reaching the top of the +ladder. Its persistence after a way of escape had been found is an +indication of the nature of habit. Day after day the halt became shorter +until finally it was little more than a pause and a turn of the head +toward one side of the ladder. I think we may say that in this act we have +evidence of the persistence of a particular resolution of physiological +states which is neither advantageous nor disadvantageous to the organism. +Had the act resulted in any gain, it would have become more marked and +elaborate; had it resulted in injury or discomfort, it would have +disappeared entirely. I have observed the same kind of behavior in the +frog and in other animals. What the animal begins to do it persists in +unless the act is positively harmful or conflicts with some beneficial +activity. The only explanation of certain features of behavior is to be +found in the conditions of their original occurrence. They persist by +sheer force of conservatism. They have value only in the light of the +circumstances under which they first appeared. Although this is merely a +fact of habit formation, it suggests that many of the problems which have +puzzled students of behavior for ages may be solved by a study of the +history of activity. + +That there are marked individual differences in intelligence in the +dancing mice is apparent from the results of the ladder-climbing +experiment. No. 1000 learned to climb quickly, and largely by his own +initiative; Nos. 2 and 6, on the contrary, learned only by reason of +tuition (being put through the required act by the experimenter). It +occurred to me that this experiment, since it was difficult for some +individuals and easy for others, might be used to advantage as a test of +imitation. If a dancer which knows how to escape to the cage by way of the +ladder be placed in the wooden box with one which, despite abundant +opportunity, has proved unable to form the habit on his own initiative, +will the latter profit by the activity of the former and thus learn the +method of escape? + +On November 20, Nos. 4 and 5 were placed in the wooden box and left there +for half an hour. As they had failed to escape at the end of this +interval, they were taken out of the box by the experimenter and returned +to the nest-box. November 21 and 22 this test of their ability to learn to +climb the ladder was repeated with the same result. On November 23 they +were placed in the box with the three mice which had previously been +trained to climb the ladder. The latter escaped at once. Apparently the +attention of Nos. 4 and 5 was drawn to the ladder by the disappearance of +their companions, for they approached its foot and No. 5 climbed up a +short distance. Neither succeeded in escaping, however, and they made no +further efforts that day. On the 24th, and daily thereafter until the +29th, these two dancers were placed in the box for half an hour, with +negative results. At the end of the half hour on the 29th, Nos. 2 and 6 +were placed in the box and permitted to go back and forth from one box to +the other repeatedly within sight of Nos. 4 and 5. The latter made no +attempts to follow them, although at times they seemed to be watching +their movements as they ascended the ladder. + +To render the results of this test of imitation still more conclusive No. +5 was given further opportunity to learn from No. 1000. Beginning December +2, the following method of experimentation was employed with these two +individuals. They were placed in the wooden box together. No. 1000 usually +climbed out almost immediately. Sometimes No. 5 apparently saw him +disappear up the ladder; sometimes she paid no attention whatever either +to the presence or absence of her companion. After he had been in the +nest-box for a few seconds, No. 1000 was returned to the wooden box by the +experimenter and again permitted to climb out for the benefit of No. 5. +This mode of procedure was kept up until No. 1000 had made from three to +ten trips. No. 5 was left in the box for half an hour each day. This test +was repeated on 18 days within a period of 3 weeks. No. 5 showed no signs +of an imitative tendency, and she did not learn to climb the ladder. + +To this evidence of a lack of an imitative tendency in the dancer I may +here add the results of my observations in other experiments. In the +discrimination tests and in the labyrinth tests I purposely so arranged +conditions, in certain instances, that one individual should have an +opportunity to imitate another. In no case did this occur. Seldom indeed +did the animals so much as follow one another with any considerable degree +of persistence. They did not profit by one another's acts. + +Excellent evidence in support of this conclusion was furnished by the +behavior of the mice in the discrimination experiments. Some individuals +learned to pull as well as to push the swinging wire doors of the +apparatus and were thus enabled to pass through the doorways in either +direction; other individuals learned only to pass through in the direction +in which the doors could be pushed open. Naturally I was interested to +discover whether those which knew only the trick of opening the doors by +pushing would learn to pull the doors or would be stimulated to try by +seeing other individuals do so. At first I arranged special tests of +imitation in the discrimination box; later I observed the influence of the +behavior of one mouse upon that of its companion in connection with visual +discrimination experiments. This was made possible by the fact that +usually a pair of individuals was placed in the discrimination box and the +tests given alternately to the male and to the female. Both individuals +had the freedom of the nest-box and each frequently saw the other pass +through the doorway between the nest-box, _A_, and the entrance chamber, +_B_ (Figure 14), either from _A_ to _B_ by pushing the swing door or from +_B_ to _A_ by pulling the door. + +Although abundant opportunity for imitation in connection with the opening +of the doors in the discrimination box was given to twenty-five +individuals, I obtained no evidence of ability to learn by imitation. The +dancers did not watch the acts which were performed by their companions, +and in most instances they did not attempt to follow a mate from nest-box +to entrance chamber. + +These problem tests, simple as they are, have revealed two important facts +concerning the educability of the dancer. First, that it does not learn by +imitation to any considerable extent, and, second, that it is aided by +being put through an act. Our general conclusion from the results of the +experiments which have been described in this chapter, if any general +conclusion is to be drawn thus prematurely, must be that the dancing mouse +in its methods of learning differs markedly from other mice and from rats. + + + + +CHAPTER XIII + +HABIT FORMATION: THE LABYRINTH HABIT + +The problem method, of which the ladder and door-opening tests of the +preceding chapter are examples, has yielded interesting results concerning +the individual initiative, ingenuity, motor ability, and ways of learning +of the dancer; but it has not furnished us with accurate measurements of +the rapidity of learning or of the permanency of the effects of training. +In this chapter I shall therefore present the results of labyrinth +experiments which were planned as means of measuring the intelligence of +the dancer. + +The four labyrinths which have been used in the investigation may be +designated as _A, B, C,_ and _D_. They differ from one another in the +character of their errors, as well as in the number of wrong choices of a +path which the animal might make on its way from entrance to exit. In the +use of the labyrinth method, as in the case of the discrimination method +of earlier chapters, the steps by which a satisfactory form of labyrinth +for testing the dancer was discovered are quite as interesting and +important for those who have an intelligent appreciation of the problems +and methods of animal psychology as are the particular results which were +obtained. For this reason, I shall describe the various forms of labyrinth +in the order in which they were used, whether they proved satisfactory or +not. At the outset of this part of my investigation, it was my purpose to +compare directly the capacity for habit formation in the dancer with that +of the common mouse. This proved impracticable because the same labyrinth +is not suited to the motor tendencies of both kinds of mice. + +[Illustration: FIGURE 25.--Labyrinth A. _I_. entrance; _O_, exit; 1, 2, 3, +4, blind alleys.] + +The first of the four labyrinths, A, appears in ground plan in Figure 25. +It was constructed of wood, as were the other labyrinths also, and +measured 60 cm. in length and width, and 10 cm. in depth. The outside +alleys were 5 cm. wide. In the figure, _I_ marks the starting point or +entrance to the maze, and _O_ the exit through which the mouse was +permitted to pass into its nest-box. Any turn in the wrong direction which +the animal made in its progress from entrance to exit was recorded as an +error. The four errors, exclusive of the mistake of turning back, which +were possible in this labyrinth, are indicated in the figure by the +numerals 1, 2, 3, and 4. By retracing its steps a mouse might repeat any +one or all of these errors, and add to them the error of turning back. + +In the experiments a mouse was permitted to enter the maze from a small +box which had been placed by the experimenter at _I_, and an accurate +record was kept of the number of errors which it made in finding its way +from entrance to exit, and of the time occupied. Each of five dancers was +given 31 tests in this labyrinth. The number of tests per day varied, as +is indicated in Table 36, from 1 to 4. The results of the tests, so far as +errors and times are in question, appear in the table. _T_ at the head of +a column is an abbreviation for time, _E_ for errors. + +The dancers did not learn to escape from this labyrinth easily and +quickly. In fact, the average time of the thirty-first test (198") is +considerably longer than that of the first (130"). The number of errors +decreased, it is true, but even for the last test it was 6.6 as compared +with only a little more than twice that number for the first test. The +last column of the table furnishes convincing proof of the truth of the +statement that the animals did not acquire a perfect labyrinth-A habit. +Was this due to inability to learn so complex a path, or to the fact that +the method is not adapted to their nature? Observation of the behavior of +the mice in the experiments enables me to say with certainty that there +was no motive for escape sufficiently strong to establish a habit of +following the direct path. Often, especially after a few experiences in +the maze, a dancer would wander back and forth in the alleys and central +courts, dancing much of the time and apparently exploring its surroundings +instead of persistently trying to escape. This behavior, and the time and +error results of the accompanying table, lead me to conclude that the +labyrinth method, as it has been employed in the study of the intelligence +of several other mammals, is not a satisfactory test of the ability of the +dancer to profit by experience. That the fault is not in the labyrinth +itself is proved by the results which I obtained with common mice. + + + +TABLE 36 + +RESULTS OF LABYRINTH A TESTS WITH DANCERS + + + AVERAGE +TEST DATE No. 1000 No. 2 No. 6 No. 4 No. 5 FOR ALL + 1905 + T E T E T E T E T E T E + + 1 Nov 23 130" 14 100" 8 170" 13 60" 6 190" 26 130" 13.4 + 2 24 140 19 78 7 60 8 149 6 211 25 128 13.0 + 3 25 392 31 87 1 98 5 185 13 120 9 176 11.8 + 4 26 448 38 38 3 47 2 50 3 121 12 141 11.3 + 5 27 142 8 21 2 27 3 27 2 17 1 47 3.2 + 6 28 45 2 61 7 63 5 102 8 33 4 61 5.2 + 7 29 303 17 64 7 36 3 42 2 57 4 100 6.6 + 8 30 222 15 26 2 37 5 42 3 7 0 67 5.0 + 9 Dec 1 185 9 36 5 48 3 63 3 94 8 85 5.6 + 10 2 52 2 71 4 19 0 196 5 95 11 87 4.4 + 11 3 180 8 32 2 107 4 52 3 38 4 82 4.2 + 12 4 310 10 133 11 65 3 242 6 125 6 175 7.2 + 13 4 153 9 335 55 130 10 195 15 154 18 193 21.4 + 14 5 330 7 69 2 42 2 201 6 130 10 154 5.4 + 15 5 287 7 34 4 61 4 136 7 25 2 109 4.8 + 16 5 455 15 65 4 25 0 110 8 160 15 183 8.4 + 17 6 120 15 280 9 33 0 168 4 39 2 128 6.0 + 18 6 120 4 164 10 81 4 101 5 85 4 110 5.4 + 19 6 132 12 78 7 110 6 40 2 151 12 102 7.8 + 20 7 258 10 223 16 33 1 92 5 37 1 129 6.6 + 21 7 110 7 23 3 44 4 20 4 305 23 100 8.2 + 22 7 100 4 60 8 167 15 44 7 58 4 86 7.6 + 23 8 43 1 179 7 356 6 34 3 65 3 135 4.0 + 24 8 92 5 56 5 42 3 17 1 23 1 46 3.0 + 25 9 85 5 114 3 62 3 129 8 31 0 84 3.8 + 26 9 30 2 36 4 109 15 12 1 34 2 44 4.8 + 27 9 69 5 40 4 85 6 36 3 16 1 49 3.8 + 28 10 169 7 80 3 28 0 142 5 35 2 89 3.4 + 29 10 155 5 266 8 91 5 27 0 37 2 115 4.0 + 30 10 29 1 25 2 124 14 83 6 111 12 74 7.0 + 31 10 465 6 208 8 95 3 65 3 159 13 198 6.6 + + + + +On the basis of two tests per day, two common mice, a white one and a gray +one, quickly learned to escape from labyrinth _A_ by the shortest path. +The time of escape for the gray individual (Table 37) decreased from 180" +in the first test to 21" in the tenth, and the number of errors from 6 to +1. Similarly in the case of the white individual, the time decreased from +122" to 8", and the errors from 5 to 1. A fraction of the number of tests +to which the dancer had been subjected sufficed to establish a habit of +escape in the common mouse. It is evident, therefore, that the dancer +differs radically from the common mouse in its behavior in a maze, and it +is also clear that the labyrinth method, if it is to be used to advantage, +must be adapted to the motor tendencies of the animal which is to be +tested. + + + +TABLE 37 + +RESULTS OF LABYRINTH A TESTS WITH COMMON MICE + + GREY MOUSE WHITE MOUSE +TEST T E T E + + 1 180" 6 122" 5 + 2 26 2 80 6 + 3 37 1 56 4 + 4 18 0 27 1 + 5 68 2 33 2 + 6 10 1 19 1 + 7 11 1 17 1 + 8 13 1 17 1 + 9 10 0 8 1 + 10 21 1 8 1 + + + +The behavior of the dancer made obvious two defects in labyrinth A. Its +passages are so large that the mouse is constantly tempted to dance, and +it lacks the basis for a strong and constant motive of escape by the +direct path. To obviate these shortcomings labyrinth B was constructed, as +is shown in Figures 23 and 24, with very narrow passages, and a floor +which was covered with the wires of an interrupted electric circuit so +that errors might be punished. The length of this labyrinth was 52 cm. and +the passages were 2.5 cm. wide and 10 cm. deep. Dancing in these narrow +alleys was practically impossible, for the mice could barely turn around +in them. In the case of all except the common mice and two dancers, a +depth of 10 cm. was sufficient to keep the animals in the maze without the +use of a cover. + +As an account of the behavior of the dancer in labyrinth B has already +been given in Chapter XI, I may now state the general results of the +experiments. In all, thirty individuals were trained in this labyrinth. +Each individual was given tests at the rate of one per minute until it had +succeeded in following the correct path five times in succession. The weak +electric shock, which was given as a punishment for mistakes, provided an +activity-impelling motive for escape to the nest-box. + +An idea of the extreme individual difference in the rapidity with which +the labyrinth-B path was learned by these dancers may be obtained by an +examination of Table 38, from which it appears that the smallest number of +training tests necessary for a successful or errorless trip through the +maze was one and the largest number fourteen. It is to be remembered that +each mouse was given an opportunity to pass through the labyrinth once +without punishment for errors, and thus to discover, before the training +tests were begun, that a way of escape existed. This first test we may +designate as the preliminary trial. Table 38 further indicates that the +females acquired the labyrinth habit more quickly than did the males. + + + +TABLE 38 + +RESULTS OF LABYRINTH-B EXPERIMENTS, WITH TWENTY DANCERS + + + MALES FEMALES + +NO. OF NO. OF FIRST NO. OF LAST OF NO. OF NO. OF FIRST NO. OF LAST OF + MOUSE CORRECT FIVE CORRECT MOUSE CORRECT FIVE CORRECT + TEST TESTS TEST TESTS + + 76 8 14 75 4 15 + 78 5 20 77 7 11 + 86 13 22 87 12 22 + 58 2 14 49 1 5 + 50 6 23 57 3 20 + 60 13 37 59 14 28 + 410 6 20 415 4 13 + 220 4 8 225 6 18 + 212 3 7 211 6 10 + 214 10 28 213 5 14 + + AV. 7.0 19.3 AV. 6.2 15.6 + + + + +A graphic representation of certain of the important features of the +process of formation of the labyrinth-B habit is furnished by Figure 26 in +which the solid line is the curve of learning for the ten males of Table +38, and the broken line for the ten females. These two curves were plotted +from the number of errors made in the preliminary trial (P in the figure) +and in each of the subsequent tests up to the sixteenth. In the case of +both the males and the females, for example, the average number of errors +in the preliminary trial was 11.3, as is indicated by the fact that the +curves start at a point whose value is given in the left margin as 11.3. +In the second training test the number of errors fell to 3.3 for the males +and 2.7 for the females. The number of the test is to be found on the base +line; the number of errors in the left margin. If these two curves of +learning were carried to their completion, that for the males would end +with the thirty-seventh test, and that for the females with the twenty- +eighth. + +[Illustration: FIGURE 26.--Curves of habit formation, plotted from the +data of labyrinth-B tests with ten males and ten females. The figures in +the left margin indicate the number of errors; those below the base line +the number of the test. _P_ designates the preliminary test. Males +____[solid line]; Females ----[broken line].] + +Time records are not reported for these and subsequent labyrinth tests +because they proved to be almost valueless as measures of the rapidity of +habit formation. At any point in its progress through a labyrinth, the +dancer may suddenly stop to wash its face, look about or otherwise examine +its surroundings; if a shock be given to hurry it along it may be +surprised into an error. It is my experience, and this is true of other +animals as well as of the dancing mouse, that a long trip, as measured in +time units, does not necessarily indicate the lack of ability to follow +the labyrinth path correctly and rapidly. Hence, whenever it is possible +(and the experimenter can always plan his tests so that it shall be +possible), the number of errors should be given first importance and the +time of the test second place. I have presented in Table 38 the number of +the first correct test, and the number of the last of five successive +correct tests. Space cannot be spared for records of the errors made in +the several tests by each individual. + +In general, labyrinth B proved very satisfactory as a means of testing the +ability of the dancer to learn a simple path. The narrow passages +effectively prevented dancing, and the introduction of the electric shock +as a punishment for mistakes developed a motive for escape which was +uniform, constant, and so strong that the animals clearly did their best +to escape from the labyrinth quickly and without errors. This maze was so +simple that it did not tend to discourage them as did the one which is +next to be described. It must be admitted, however, that, though labyrinth +B is perfectly satisfactory as a test of the dancer's ability to learn to +follow a simple path, it is not an ideal means of measuring the rapidity +of habit formation. This is due to the fact that the preliminary trial and +the first training test play extremely different roles in the case of +different individuals. A dancer which happens to follow the correct path +from entrance to exit in the preliminary trial may continue to do so, with +only an occasional error, during several of the early training tests, and +it may therefore fail for a considerable time to discover that there are +errors which should be avoided. The learning process is delayed by its +accidental success. On the other hand, an individual which happens to make +many mistakes to begin with immediately attempts to avoid the points in +the maze at which it receives the electric shock. I was led to conclude, +as a result of the labyrinth-B experiments, that the path was too easy, +and that a more complex labyrinth would, in all probability, furnish a +more satisfactory means of measuring the rapidity of habit formation. + +[Illustration: FIGURE 27--A record sheet, showing the plan of labyrinth C +(as made on the sheet by means of a rubber stamp) on which the +experimenter recorded the path followed by the mouse. This sample sheet +presents the path records for the first, fifth, tenth, and eleventh tests +of No. 2 in labyrinth C. 1, 2, 3, 4, 5 designate the several errors of the +labyrinth.] + +On the basis of the supposition that a maze whose path was so complex that +the animal would not be likely to follow it correctly in the early trials +would be more to the purpose than either A or B, labyrinth C was devised. +As is shown in the plan of this maze, Figure 27, five mistakes in choice +of path were possible on the forward trip. These errors, as a rule, were +more difficult for the dancers to avoid than those of labyrinths A and B. +Those which are designated by the numerals 2, 3, and 4 were especially +difficult. Error 4 was much more troublesome for left whirlers than for +right whirlers because, after turning around abruptly at the entrance to +the blind alley, the former type of dancer almost always followed the side +wall of the maze so far that it missed the correct path. Undoubtedly the +various errors are not of the same value for different individuals; but it +would be extremely difficult, if not impossible, to devise a maze which +should be equally difficult for several normal individuals. + +In order that records of the path followed by a mouse in test after test +might be kept with ease and accuracy by the experimenter, the plan of this +labyrinth, and also that of labyrinth D, were cast in rubber. The outlines +of labyrinths C and D which appear in Figures 27 and 28 respectively were +made with the rubber stamps which were thus obtained. Figure 27 is the +reproduction of a record sheet which presents the results of the first, +the fifth, the tenth, and the eleventh tests of No. 2 in labyrinth C. The +path followed by this individual in the first test was far too complex to +be traced accurately on the record sheet. The record therefore represents +merely the number of errors which was made in each region of the maze. For +the fifth test, and again for the tenth and the eleventh, the path was +recorded accurately. This simple device for making record blanks which can +readily be filled in at the time of the experiment should recommend itself +to all students of animal behavior. + +In labyrinth C ten pairs of dancers were given continuous training tests +at the rate of one test per minute until they were able to follow the +direct path correctly. Because of the difficulty in learning this maze +perfectly, it was not demanded of the mice that they should follow the +path correctly several times in succession, but instead the training was +terminated after the first successful trip. + + + +TABLE 39 + +RESULTS OF LABYRINTH-C EXPERIMENTS, WITH TWENTY DANCERS + + + MALES FEMALES + + NO. OF NO. OF FIRST NO. OF NO. OF FIRST + MOUSE CORRECT TEST MOUSE CORRECT TEST + + 2 11 29 15 + 30 33 49 34 + 50 49 57 15 + 52 22 59 15 + 58 16 215 10 + 60 17 415 10 + 76 3 75 8 + 78 6 77 11 + 86 5 87 9 + 88 25 85 11 + + AV. 18.7 AV. 13.8 + + + + +The results of the experiments with this labyrinth as they are presented +in Table 39 indicate that its path is considerably more difficult for the +dancer to learn than that of labyrinth B, that the females learn more +quickly than the males, and finally, that individual differences are just +as marked as they were in the case of the simpler forms of labyrinth. It +therefore appears that increasing the complexity of a labyrinth does not, +as I had supposed it might, diminish the variability of the results. +Certain of the individual differences which appear in Table 39 are due, +however, to the fact that in some cases training in labyrinth B had +preceded training in labyrinth C, whereas in the other cases C was the +first labyrinth in which the animals were tested. But even this does not +serve to account for the wide divergence of the results given by No. 2 and +No. 50, for the latter had been trained in B previous to his training in +C, and the former had not been so trained. Yet, despite the advantage +which previous labyrinth experience gave No. 50, he did not learn the path +of C as well in fifty tests as No. 2 did in eleven. The facts concerning +the value of training in one form of labyrinth for the learning of +another, as they were revealed by these experiments, may more fittingly be +discussed in a later chapter in connection with the facts of memory and +re-learning. + +[Illustration: FIGURE 28.--Plan of Labyrinth _D_, as reproduced from a +print made with a rubber stamp. _I_, entrance; _O_, exit; numerals 1 to +13, errors.] + +Labyrinth C is a type of maze which might properly be described as +irregular, since the several possible errors are extremely different in +nature. In view of the results which this labyrinth yielded, it seemed +important that the dancer be tested in a perfectly regular maze of the +labyrinth-D type. The plan which I designed as a regular labyrinth has +been reproduced, from a rubber stamp print, in Figure 28. As is true also +of the mazes previously described, it provides four kinds of possible +mistakes: namely, by turning to the left (errors 1, 5, 9, and 13), by +turning to the right (errors 3, 7, and 11), by moving straight ahead +(errors 2, 4, 6, 8, 10, and 12), and by turning back and retracing the +path just followed. The formula for the correct path of _D_ is simple in +the extreme, in spite of the large number of mistakes which are possible, +for it is merely "a turn to the right at the entrance, to the left at the +first doorway, and thereafter alternately to the right and to the left +until the exit is reached." This concise description would enable a man to +find his way out of such a maze with ease. Labyrinth D had been +constructed with an exit at 10 so that it might be used as a nine-error +maze if the experimenter saw fit, or as a thirteen-error maze by the +closing of the opening at 10. In the experiments which are now to be +described only the latter form was used. + +Can the dancer learn a regular labyrinth path more quickly than an +irregular one? Again, I may give only a brief statement of results. Each +of the twenty dancers, of Table 40, which were trained in labyrinth D had +previously been given opportunity to learn the path of C, and most of them +had been trained also in labyrinth B. All of them learned this regular +path with surprising rapidity. The numerical results of the tests with +labyrinths B, C, and D, as well as the behavior of the mice in these +several mazes, prove conclusively that the nature of the errors is far +more important than their number. Labyrinth D with its thirteen chances of +error on the forward trip was not nearly as difficult for the dancer to +learn to escape from as labyrinth C with its five errors. That the +facility with which the twenty individuals whose records are given in +Table 40 learned the path of D was not due to their previous labyrinth +experience rather than to the regularity of the maze is proved by the +results which I obtained by testing in D individuals which were new to +labyrinth experiments. Even in this case, the number of tests necessary +for a successful trip was seldom greater than ten. If further evidence of +the ease with which a regular labyrinth path may be followed by the dancer +were desired, it might be obtained by observation of the behavior of an +individual in labyrinths C and D. In the former, even after it has learned +the path perfectly, the mouse hesitates at the doorways from time to time +as if uncertain whether to turn to one side or go forward; in the latter +there is seldom any hesitation at the turning points. The irregular +labyrinth is followed carefully, as by choice of the path from point to +point; the regular labyrinth is followed in machine fashion,--once +started, the animal dashes through it. + + + +TABLE 40 + +RESULTS OF LABYRINTH-D EXPERIMENTS, WITH TWENTY DANCERS + + + MALES FEMALES + + NO. OF NO. OF FIRST NO. OF LAST OF NO. OF NO. OF FIRST NO. OF LAST OF + MOUSE CORRECT TWO CORRECT MOUSE CORRECT TWO CORRECT + TEST TESTS TEST TESTS + + 2 3 7 29 10 11 + 58 7 10 49 7 8 + 30 9 10 57 3 6 + 60 10 14 215 6 10 + 402 10 11 415 7 8 + 76 4 7 75 4 13 + 78 4 5 77 11 12 + 86 3 9 87 4 9 + 88 4 8 85 3 4 + 90 7 8 83 4 7 + + Av. 6.1 8.9 Av. 5.9 8.8 + + + + +From the results of these labyrinth experiments with dancers I am led to +conclude that a standard maze for testing the modifiability of behavior of +different kinds of animals should be constructed in conformity with the +following suggestions. Errors by turning to the right, to the left, and by +moving forward should occur with equal frequency, and in such order that +no particular kind of error occurs repeatedly in succession. If we should +designate these three types of mistake by the letters _r, l_, and _s_ +respectively, the error series of labyrinth C would read _l-l-r-s-l_. It +therefore violates the rule of construction which I have just formulated. +In the case of labyrinth D the series would read _l-s-r-s-l-s-r-s-l-s-r-s- +l_. This also fails to conform with the requirement, for there are three +errors of the first type, four of the second, and six of the third. Again, +in a standard maze, the blind alleys should all be of the same length, and +care should be taken to provide a sufficiently strong and uniform motive +for escape. In the case of one animal the desire to escape from +confinement may prove a satisfactory motive; in the case of another, the +desire for food may conveniently supplement the dislike of confinement; +and in still other cases it may appear that some form of punishment for +errors is the only satisfactory basis of a motive for escape. Readers of +this account of the behavior of the dancing mouse must not infer from my +experimental results that the electric shock as a means of forcing +discrimination will prove satisfactory in work with other animals or even +with all other mammals. As a matter of fact it has already been proved by +Doctor G. van T. Hamilton that the use of an electric shock may so +intimidate a dog that experimentation is rendered difficult and of little +value. And finally, in connection with this discussion of a standard +Labyrinth, I wish to emphasize the importance of so recording the results +of experiments that they may be interpreted in terms of an animal's +tendency to turn to the right or to the left. My work with the dancer has +clearly shown that the avoidance of a particular error may be extremely +difficult for left whirlers and very easy for right whirlers. + +I hope I have succeeded in making clear by the foregoing account of my +experiments that the labyrinth method is more satisfactory in general than +the problem method as a means of measuring the rapidity of habit formation +in the dancer, and I hope that I have made equally clear the fact that it +is very valuable as a means of discovering the roles of the various senses +in the acquirement of a habit (Chapter XI). From my own experience in the +use of the labyrinth with the dancer and with other animals, I am forced +to conclude that its chief value lies in the fact that it enables the +experimenter so to control the factors of a complex situation that he may +readily determine the importance of a given kind of sense data for the +formation or the execution of a particular habit. As a means of measuring +the intelligence of an animal, of determining the facility with which it +is capable of adjusting itself to new environmental conditions, and of +measuring the permanency of modifications which are wrought in its +behavior by experimental conditions, I value the labyrinth method much +less highly now than I did previous to my study of the dancer. It is +necessarily too complex for the convenient and reasonably certain +interpretation of results. Precisely what is meant by this statement will +be evident in the light of the results of the application of the +discrimination method to the dancer, which are to be presented in the next +chapter. The labyrinth method is an admirable means of getting certain +kinds of qualitative results; it is almost ideal as a revealer of the role +of the senses, and it may be used to advantage in certain instances for +the quantitative study of habit formation and memory. Nevertheless, I +think it may safely be said that the problem method and the discrimination +method are likely to do more to advance our knowledge of animal behavior +than the labyrinth method. + + + + +CHAPTER XIV + +HABIT FORMATION: THE DISCRIMINATION METHOD + +Discrimination is demanded of an animal in almost all forms of the problem +and labyrinth methods, as well as in what I have chosen to call the +discrimination method. In the latter, however, discrimination as the basis +of a correct choice of an electric-box is so obviously important that it +has seemed appropriate to distinguish this particular method of measuring +the intelligence of the dancer from the others which have been used, by +naming it the discrimination method. + +It has been shown that neither the problem nor the labyrinth method proves +wholly satisfactory as a means of measuring the rapidity of learning, or +the duration of the effects of training, in the case of the dancer. The +former type of test serves to reveal to the experimenter the general +nature of the animal's capacity for profiting by experience; the latter +serves equally well to indicate the parts which various receptors (some of +which are sense organs) play in the formation and execution of habits. But +neither of them is sufficiently simple, easy of control, uniform as to +conditions which constitute bases for activity, and productive of +interpretable quantitative results to render it satisfactory. The problem +method is distinctly a qualitative method, and, in the case of the dancing +mouse, my experiments have proved that the labyrinth method also yields +results which are more valuable qualitatively than quantitatively. I had +anticipated that various forms of the labyrinth method would enable me to +measure the modifiability of behavior in the dancer with great accuracy, +but, as will now be made apparent, the discrimination method proved to be +a far more accurate method for this purpose. + +Once more I should emphasize the fact that my statements concerning the +value of methods apply especially to the dancing mouse. Certain of the +tests which have proved to be almost ideal in my study of this peculiar +little rodent would be useless in the study of many other mammals. An +experimenter must work out his methods step by step in the light of the +daily results of patient and intelligent observation of the motor +capacity, habits, instincts, temperament, imitative tendency, +intelligence, hardihood, and life-span of the animal which he is studying. +The fact that punishment has proved to be more satisfactory than reward in +experiments with the dancer does not justify the inference that it is more +satisfactory in the case of the rat, cat, dog, or monkey. Methods which +yielded me only qualitative results, if applied to other mammals might +give accurate quantitative results; and, on the other hand, the +discrimination method, which has proved invaluable for my quantitative +work, might yield only qualitative results when applied to another kind of +animal. + +The form of the discrimination method whose results are to be presented in +this chapter has already been described as white-black discrimination. In +the discrimination box (Figures 14 and 15, p. 92) the two electric-boxes +which were otherwise exactly alike in appearance were rendered +discriminable for the mouse by the presence of white cardboards in one and +black cardboards in the other. In order to escape from the narrow space +before the entrances to the two electric-boxes, the dancer was required to +enter the white box. If it entered the black box a weak electric shock was +experienced. After two series of ten tests each, during which the animal +was permitted to choose either the white or the black box without shock or +hindrance, the training was begun. These two preliminary series serve to +indicate the natural preference of the animal for white or black previous +to the training. An individual which very strongly preferred the white +might enter, from the first, the box thus distinguished, whereas another +individual whose preference was for the black might persistently enter the +black box in spite of the disagreeable shocks. First of all, therefore, +the preliminary tests furnish a basis for the evaluation of the results of +the subsequent training tests. On the day succeeding the last series of +preliminary tests, and daily thereafter until the animal had acquired a +perfect habit of choosing the white box, a series of training tests was +given. These experiments were usually made in the morning between nine and +twelve o'clock, in a room with south-east windows. The entrances to the +electric-boxes faced the windows, consequently the mouse did not have to +look toward the light when it was trying to discriminate white from black. +All the conditions of the experiment, including the strength of the +current for the shock, were kept as constant as possible. + +Choice by position was effectively prevented, as a rule, by shifting the +cardboards so that now the left now the right box was white. The order of +these shifts for the white-black series whose results are quantitatively +valuable appear in Table 12 (p. III). That the order of these changes in +position may be criticised in the light of the results which the tests +gave, I propose to show hereafter in connection with certain other facts. +The significant point is that the defects which are indicated by the +averages of thousands of tests could not have been predicted with +certainty even by the most experienced investigator in this field. + +In Table 41 are to be found the average number of errors in each series of +ten white-black discrimination tests for five males and for five females +which were trained by being given ten tests per day, and similarly for the +same number of individuals of each sex, trained by being given twenty +tests per day. Since the results for these two conditions of training are +very similar, the averages for the twenty individuals are presented in the +last column of the table. For the present we may neglect the interesting +individual, sex, and age differences which these experiments revealed and +examine the significant features of the general averages, and of the +white-black discrimination curve (Figure 29). + + + + TABLE 41 + +WHITE BLACK DISCRIMINATION TESTS. NUMBER OF ERRORS IN + THE VARIOUS SERIES + + MALES FEMALES + + AVERAGES AVERAGES GENERAL AVERAGES AVERAGES GENERAL AVERAGES +SERIES FOR 5, FOR 5, AVERAGES FOR 5, FOR 5, AVERAGES FOR ALL + 10 TESTS 20 TESTS FOR 10 10 TESTS 20 TESTS FOR 10 (20) MALES + PER DAY PER DAY PER DAY PER DAY AND FEMALES + + A 5.8 6.0 5.9 5.8 5.8 5.8 5.85 + B 5.6 6.2 5.9 5.8 5.6 5.7 5.8 + 1 5.0 5.0 5.0 5.6 4.6 5.1 5.05 + 2 2.6 4.6 3.6 4.4 5.0 4.7 4.15 + 3 3.0 3.4 3.2 3.4 3.4 3.4 3.3 + 4 2.6 3.8 3.2 2.4 2.2 2.3 2.75 + 5 2.4 2.0 2.2 2.6 1.8 2.2 2.2 + 6 1.6 1.6 1.6 1.0 2.2 1.6 1.6 + 7 1.0 1.4 1.2 2.0 0.4 1.2 1.2 + 8 0.2 0.6 .4 1.4 1.6 1.5 .95 + 9 0.2 1.0 .6 0.6 0.8 .7 .65 + 10 0 .8 .4 1.0 0.8 .9 .65 + 11 0 .8 .4 0.8 0 .4 .40 + 12 0 .6 .3 0.4 0 .2 .25 + 13 0 0 0 0 0 0 0 + 14 0 0 0 0 0 0 + 15 0 0 0 0 0 0 + + + + +[Illustration: FIGURE 29.--Error curve plotted from the data given by +twenty dancers in white-black discrimination tests. The figures in the +left margin indicate the number of errors; those below the base line, the +number of the series. _A_ and _B_ designate the preference series.] + + The preference series, _A_ and _B_, reveal a constant tendency to choose +the black box, whose strength as compared with the tendency to choose the +white box is as 5.8 is to 4.2. In other words, the dancer on the average +chooses the black box almost six times in ten. The first series of +training tests reduced this preference for black to zero, and succeeding +series brought about a rapid and fairly regular decrease in the number of +errors, until, in the thirteenth series, the white was chosen every time. +Since I arbitrarily define a perfect habit of discrimination as the +ability to choose the right box in three successive series of ten tests +each, the tests ended with the fifteenth series. + +The discrimination curve, Figure 29, is a graphic representation of the +general averages of Table 41.--It is an error curve, therefore. Starting +at 5.85 for the first preliminary series, it descends to 5.8 for the +second series, and thence abruptly to 5.05 for the first training series. +This series of ten tests therefore served to reduce the black preference +very considerably. The curve continues to descend constantly until the +tenth series, for which the number of errors was the same as for the +preceding series, .65. This irregularity in the curve, indicative, as it +would appear, of a sudden cessation in the learning process, demands an +explanation. My first thought was that an error in computation on my part +might account for the shape of the curve. The error did not exist, but in +my search for it I discovered what I now believe to be the cause of the +interruption in the fall of the error curve. In all of the training series +up to the tenth the white cardboard had been on the right and the left +alternately or on one side two or three times in succession, whereas in +the tenth series, as may be seen by referring to Table 12 (p.111), it was +on the left for the first four tests, then on the right four times, and, +finally, on the left for the ninth test and on the right for the tenth. +This series was therefore a decidedly more severe test of the animal's +ability to discriminate white from black and to choose the white box +without error than were any that had preceded it. If my interpretation of +the results is correct, it was so much more severe than the ninth series +that the process of habit formation was obscured. It would not be fair to +say that the mouse temporarily ceased to profit by its experience; instead +it profited even more than usually, in all probability, but the +unavoidably abrupt increase in the difficultness of the tests was just +sufficient to hide the improvement. + +As I have suggested, the plan of experimentation may be criticised +adversely in the light of this irregularity in the error curve. Had the +conditions been perfectly satisfactory the curve would not have taken this +form. I admit this, but at the same time I am glad that I chose that +series of shifts in the position of the cardboards which, as it happens, +served to exhibit an important aspect of quantitative measures of the +modifiability of behavior that otherwise would not have been revealed. Our +mistakes in method often teach us more than our successes. I have taken +pains, therefore, to describe the unsatisfactory as well as the +satisfactory steps in my study of the dancer. + +[Illustration: FIGURE 30.--Error curve plotted from the data given by +thirty dancers, of different ages and under different conditions of +training, in white-black discrimination tests.] + +The form of the white-black discrimination curve of Figure 29 is more +surprising than disappointing to me, for I had anticipated many more +irregularities than appear. What I had expected, as the result of training +five or even ten pairs of mice, was the kind of curve which is presented, +for contrast with the one already discussed, in Figure 30. This also is an +error curve, but, unlike the previous one, it is based upon results which +were got from individuals of different ages which were trained according +to the following different methods. Ten of these individuals were given +two or five tests daily, ten were given ten tests daily, and ten were +given twenty tests daily. The form of the curve serves to call attention +to the importance of uniform conditions of training, in case the results +are to be used as accurate measures of the rapidity of learning. + +Examination of the detailed results of the white-black discrimination +tests as they appear in the tables of Chapter VII will reveal the fact +that some individuals succeeded in choosing correctly in a series of ten +tests after not more than five series, whereas others required at least +twice as many tests as the basis of a perfect series. In very few +instances, however, was a perfect habit of discrimination established by +fewer than one hundred tests. As the averages just presented in Table 41 +indicate, fifteen series, or one hundred and fifty tests, were required +for the completion of the experiment. One might search a long time, +possibly, for another mammal whose curve of error in a simple +discrimination test would fall as gradually as that of the dancer. It is +fair to say that this animal learns very slowly as compared with most +mammals which have been carefully studied. It is to be remembered, +however, that quantitative results such as are here presented for the +dancer are available for few if any other animals except the white rat. +Neither in the form of the curve of learning nor in the behavior of the +animal as it makes its choice of an electric-box is there evidence of +anything which might be described as a sudden understanding of the +situation. The dancer apparently learns by rote. It exhibits neither +intelligent insight into an experimental situation nor ability to profit +by the experience of its companions. That the selection of the white box +occurs in various ways in different individuals, and even in the same +individual at different periods in the training process, is the only +indication of anything suggestive of implicit reasoning. Naturally enough +comparison of the two boxes is the first method of selection. It takes the +dancers a surprisingly long time to reach the point of making this +comparison as soon as they are confronted by the entrances to the two +electric-boxes. The habit of running from entrance to entrance repeatedly +before either is entered, once having been acquired, is retained often +throughout the training experiments. But in other cases, an individual +finally comes to the point of choosing by what appears to be the immediate +recognition of the right or the wrong box. In the former case the mouse +enters the white box immediately; in the latter, it rushes from the black +box into the white one without hesitation. So much evidence the +discrimination tests furnish of forms of behavior which in our fellow-men +we should interpret as rational. + +[Illustration: FIGURE 31.--Curve of habit formation, plotted from the data +of labyrinth-D tests with ten males and ten females.] + +Comparison of the error curves for the labyrinth tests (Figures 26 and 31) +with those for the discrimination tests (Figures 29 and 30) reveals +several interesting points of difference. The former fall very abruptly at +first, then with decreasing rapidity, to the base line; the latter, on the +contrary, fall gradually throughout their course. Evidently the labyrinth +habit is more readily acquired by the dancer than is the visual +discrimination habit. Certain motor tendencies can be established quickly, +it would seem, whereas others, and especially those which depend for their +guidance upon visual stimuli, are acquired with extreme slowness. From +this it might be inferred that the labyrinth method is naturally far +better suited to the nature of the dancer than is any form of the +discrimination method. I believe that this inference is correct, but at +the same time I am of the opinion that the discrimination method is of +even greater value than the labyrinth method as a means of discovering the +capacity of the animal for modification of behavior. + +Inasmuch as my first purpose in the repetition of white-black +discrimination tests with a number of individuals was to obtain +quantitative results which should accurately indicate individual, age, and +sex differences in the rapidity of learning, it is important to consider +the reliability of the averages with which we have been dealing. Possibly +two groups of five male dancers each, chosen at random, would yield very +different results in discrimination tests. This would almost certainly be +true if the animals were selected from different lots, or were kept before +and during the tests under different environmental conditions. But from my +experiments it has become apparent that the average of the results given +by five individuals of the same sex, age, and condition of health, when +kept in the same environment and subjected to the same experimental tests, +is sufficiently constant from group to group to warrant its use as an +index of modifiability for the race. This expression, index of +modifiability, is a convenient mode of designating the average number of +tests necessary for the establishment of a perfect habit of white-black +discrimination. Hereafter I shall use it instead of a more lengthy +descriptive phrase. + +As an indication of the degree of accuracy of measurements of the rapidity +of learning which are obtained by the use of 5 individuals I may offer the +following figures. For one of two directly comparable groups of 5 male +dancers which were chosen from 16 individuals which had been trained, the +number of tests which resulted in a perfect habit of white-black +discrimination was 92; for the other group it was 96. These indices for +strictly comparable groups of 5 individuals each differ from one another +by less than 5 per cent. Similarly, in the case of two groups of females, +the indices of modifiability were 94 and 104. These figures designate the +number of tests up to the point at which errors ceased for at least three +successive series (30 tests). + +The determination of the probable error of the index of modifiability +further aids us in judging of the reliability of the measure of the +rapidity of learning which is obtained by averaging the results for 5 +individuals. For a group of 5 males (Table 43, p. 243) the index was 72 ± +3.5; and for a group of 5 females of the same age as the males and +strictly comparable with respect to conditions of white-black training, it +was 104 ± 2.9. A probable error of ± 3.5 indicates the reliability of the +first of these indices of modifiability; one of ± 2.9, that of the second. + +I do not doubt that 10 individuals would furnish a more reliable average +than 5, but I do doubt whether the purposes of my experiments would have +justified the great increase in work which the use of averages based upon +so large a group would have necessitated. + +Further discussion of the index of modifiability may be postponed until +the several indices which serve as measures of the efficiency of different +methods of training have been presented in the next chapter. + +From the data which constitute the materials of the present chapter it is +apparent that the results of the discrimination method are amenable to +much more accurate quantitative treatment than are those of the problem +method or the labyrinth method. But I have done little more as yet than +describe the method by which it is possible to measure certain dimensions +of the intelligence of the dancer, and to state some general results of +its application. In the remaining chapters it will be our task to discover +the value of this method and of the results which it has yielded. + + + + +CHAPTER XV + + +THE EFFICIENCY OF TRAINING METHODS + +The nature of the modifications which are wrought in the behavior of an +organism varies with the method of training. This fact is recognized by +human educators, as well as by students of animal behavior (makers of the +science of comparative pedagogy), but unfortunately accurate measurements +of the efficiency of our educational methods are rare. + +Whatever the subject of investigation, there are two preeminently +important aspects of the educative process which may be taken as +indications of the value of the method of training by which it was +initiated and stimulated. I refer to the rapidity of the learning process +and its degree of permanency, or, in terms of habit formation, to the +rapidity with which a habit is acquired, and to its duration. Of these two +easily measurable aspects of the modifications in which training results, +I have chosen the first as a means to the special study of the efficiency +of the training to which the dancing mouse has been subjected in my +experiments. + +The reader who has followed my account of the behavior of the dancer up to +this point will recall that in practically all of the discrimination +experiments the number of tests in a series was ten. Some readers +doubtless have wondered why ten rather than five or twenty tests was +selected as the number in each continuous series. I shall now attempt to +answer the question. It was simply because the efficiency of that number +of tests, given daily, when taken in connection with the amount of time +which the conduct of the experiments required, rendered it the most +satisfactory number. But this statement demands elaboration and +explanation. + +Very early in my study of the dancer, I learned that a single experience +in a given experiment day after day had so little effect upon the animal +that a perfect habit could not be established short of several weeks or +months. Similarly, experiments in which two tests per day were given +proved that even a simple discrimination habit cannot be acquired by the +animal under this condition of training with sufficient rapidity to enable +the experimenter to study the formation of the habit advantageously. Next, +ten tests in succession each day were given. The results proved +satisfactory, consequently I proceeded to carry out my investigation on +the basis of a ten-test series. After this method had been thoroughly +tried, I decided to investigate the efficiency of other methods for the +purpose of instituting comparisons of efficiency and discovering the +number of tests per day whose efficiency, as measured by the rapidity of +the formation of a white-black discrimination habit, is highest. + +For this purpose I carefully selected five pairs of dancers of the same +age, descent, and previous experience, and gave them white-black tests in +series of two tests per day (after the twentieth day the number was +increased to five) until they had acquired a perfect habit of +discriminating. Similarly other dancers were trained by means of series of +ten tests, twenty tests, or one hundred tests per day. Since it was my aim +to make the results of these various tests strictly comparable, I spared +no pains in selecting the individuals, and in maintaining constancy of +experimental conditions. The order of the changes in the position of the +cardboards which was adhered to in these efficiency tests was that given +in Table 12. + +At the beginning of the two-test training I thought it possible that the +animals might acquire a perfect habit with only a few more days' training +than is required by the ten-test method. This did not prove to be the +case, for at the end of the twentieth day (after forty tests in all) the +average number of mistakes, as Table 42 shows, was 3.2 for the males and +3.0 for the females. Up to this time there had been clear evidence of the +formation of a habit of discriminating white from black, but, on the other +hand, the method had proved very unsatisfactory because the first test +each day usually appeared to be of very different value from the second. +On account of the imminent danger of the interruption of the experiment by +the rapid spread of an epidemic among my mice, I decided to increase the +number of tests in each series to five in order to complete the experiment +if possible before the disease could destroy the animals. On the twenty- +first day and thereafter, five-test series were given instead of two-test. +Unfortunately I was able to complete the experiment up to the point of +thirty successive correct tests with only six of the ten individuals whose +numbers appear at the top of Table 42. That the results of this table are +reliable, despite the fact that some of the individuals had to be taken +out of the experiment on account of bad condition, is indicated by the +fact that all the mice continued to do their best to discriminate so long +as they were used. Possibly the habit would have been acquired a little +more quickly by some of the individuals had they been stronger and more +active. + +It should be explained at this point that the results in all the +efficiency-of-training tables of this chapter are arranged, as in the +previous white-black discrimination tables, in tens, that is, each figure +in the tables indicates the number of errors in a series of ten tests. In +all cases _A_ and _B_ mark preliminary series of tests which were given at +the rate of ten tests per series. The numbers in the first column of these +tables designate groups of ten tests each, and not necessarily daily +series. In Table 42, for example, 1 includes the results of the first five +days of training, 2, of the next five days, and so on. The table shows +that No. 80 made seven wrong choices in the first five series of two tests +each. This method of grouping results serves to make the data for the +different methods directly comparable, and at the same time it saves space +at the sacrifice of very little valuable information concerning the nature +of the daily results. It is to be noted, with emphasis, that the two-five +tests per day training established a perfect habit after four weeks of +training. This method is therefore costly of the experimenter's time. + + + +TABLE 42 + +EFFICIENCY OF TRAINING. WHITE-BLACK TESTS AT THE RATE +OF 2 OR 5 PER DAY + + MALES FEMALES +SETS 80 82 84 86 88 AV. 73 79 83 85 89 AV. +OF 10 + + A 5 5 4 8 5 5.4 5 6 7 7 6 6.2 + B 5 3 6 5 6 5.0 7 5 7 6 7 6.4 + + 1 7 7 6 6 6 6.4 7 6 9 4 6 6.4 + 2 2 1 0 6 6 3.0 6 5 6 5 5 5.4 + 3 4 5 5 1 2 3.2 6 5 2 4 1 3.6 + 4 3 4 7 2 0 3.2 4 3 1 4 3 3.0 + 5 2 3 3 2 4 2.8 - 3 4 3 1 2.7 + 6 2 2 - 2 2 2.0 - 0 2 2 0 1.0 + 7 - 1 - 0 1 0.7 - 1 0 2 1 1.0 + 8 - - - 1 1 1.0 - 1 1 0 0 0.5 + 9 - - - 0 1 0.5 0 1 1 0 0 0.5 +10 - - - 0 0 0 - 0 0 0 0 0 +11 - - - 0 0 0 - 0 0 0 +12 - - - 0 0 - 0 0 0 + + + + +TABLE 43 + +EFFICIENCY OF TRAINING. WHITE-BLACK TESTS AT THE RATE OF +10 PER DAY + + MALES FEMALES +SETS 210 220 230 410 420 AV. 215 225 235 415 425 AV. +OF 10 + + A 6 5 6 6 6 5.8 8 4 4 8 5 5.8 + B 6 8 8 5 1 5.6 8 7 6 6 2 5.8 + + 1 6 7 6 2 4 5.0 7 6 5 6 4 5.6 + 2 4 3 1 2 3 2.6 5 6 4 2 5 4.4 + 3 3 1 4 3 4 3.0 3 3 4 2 5 4.4 + 4 5 0 3 3 2 3.2 2 1 3 3 3 2.4 + 5 3 0 4 1 4 2.4 1 3 3 3 3 2.6 + 6 2 1 4 0 1 1.6 2 1 1 1 0 1.0 + 7 1 0 3 1 0 1.0 1 1 2 3 3 2.0 + 8 0 0 1 0 0 0.2 0 0 2 2 3 2.0 + 9 0 0 0 1 0 0.2 1 0 0 1 1 0.6 +10 0 0 0 0 0 2 1 0 2 1.0 +11 0 0 0 0 3 0 1 0 0.8 +12 0 0 0 0 0 2 0 0.4 +13 0 0 0 0 0 +14 0 0 0 +15 0 0 + + + + +TABLE 44 + +EFFICIENCY OF TRAINING. WHITE-BLACK TESTS AT THE RATE OF +20 PER DAY + + MALES FEMALES +SETS 72 74 208 240 402 AV. 217 230 245 403 407 AV. +OF 10 + + A 4 6 7 7 6 6.0 5 4 7 7 6 5.8 + B 6 4 6 8 7 6.2 7 3 5 8 5 5.6 + + 1 3 5 7 5 5 5.0 3 6 4 4 6 4.6 + 2 4 3 7 5 4 4.6 7 3 5 4 6 5.0 + 3 3 3 3 5 3 3.4 4 3 3 2 5 3.4 + 4 6 3 1 4 5 3.8 5 0 1 2 3 2.2 + 5 4 1 0 2 3 2.0 6 0 0 1 2 1.8 + 6 3 1 0 2 2 1.6 4 1 1 0 6 2.2 + 7 3 2 0 1 1 1.4 1 0 0 0 1 0.4 + 8 2 0 1 1 0.6 0 3 3 0 2 1.6 + 9 2 1 1 1 1.0 1 0 0 3 0.8 +10 1 2 1 0 0.8 0 1 1 2 0.8 +11 3 1 0 0 0.8 0 0 0 0 0 +12 1 2 0 0 0.6 0 0 0 0 0 +13 0 0 0 0 0 0 0 0 +14 0 0 0 +15 0 0 0 + + +The results of the ten-test training as they appear in Table 43 need no +special comment, for quite similar data have already been examined in +other connections. In the case of this table it is to be remembered that +each figure represents the number of errors for a single day as well as +for a series of ten successive tests. The results of Table 44, on the +other hand, appear as subdivided series, since each daily series was +constituted by two series of ten tests, or in all twenty tests. + +Finally, in Table 45 I have arranged the results of what may fairly be +called the continuous training method. In connection with several of the +labyrinth experiments of Chapter XIII continuous training proved very +satisfactory. It therefore seemed worth while to ascertain whether the +same method would not be more efficient than any other for the +establishment of a white-black discrimination habit. That this method was +not applied to ten individuals as were the two-five-test, the ten-test, +and the twenty-test methods is due to the fact that it proved practically +inadvisable to continue the tests long enough to complete the experiment. +I have usually designated the method as one hundred or more tests daily. I +applied this training method first to individuals Nos. 51 and 60. At the +end of one hundred and twenty tests with each of these individuals I was +forced to discontinue the experiment for the day because of the approach +of darkness. In the table the end of a series for the day is indicated by +a heavy line. The following day Nos. 51 and 60 succeeded in acquiring a +perfect habit after a few more tests. + + + +TABLE 45 + +EFFICIENCY OF TRAINING. WHITE-BLACK TESTS AT THE RATE OF +100 OR MORE PER DAY + + SETS 51[1] 60 87 Av. +OF 10 + +A 5 5 6 5.3 +B 5 3 7 5.0 + +1 6 6 5 5.7 +2 3 2 5 3.3 +3 5 4 7 5.3 +4 7 4 5 5.3 +5 6 2 3 3.7 +6 1 1 3 1.7 +7 4 2 3 3.0 +8 3 3 0 2.0 +9 2 2 3 2.3 +10 5 0 2 2.3 +11 1 2 2 1.7 +12 2 1 1 1.3 + +13 4 1 2 2.3 +14 1 2 1 1.3 +15 3 1 5 3.0 +16 3 3 2 2.7 +17 1 0 1 0.7 +18 2 0 1 1.0 +19 0 0 2 0.7 +20 0 0 0 +21 0 1 0.3 +22 - +23 - +24 - + +[Footnote 1: Age of No. 51, 22 weeks. Age of No. 60, 17 weeks. Age of No. +87, 8 weeks.] + +The results of the continuous training method for these two mice were so +strikingly different from those yielded by the other methods that I at +once suspected the influence of some factor other than that of the number +of tests per day. The ages of Nos. 51 and 60 at the time of their tests +were twenty-two and seventeen weeks, respectively, whereas all the +individuals used in connection with the other efficiency tests were four +weeks of age. It seemed possible that the slow habit formation exhibited +in the continuous training experiments might be due to the greater age of +the mice. I therefore selected a healthy active female which was only +eight weeks old, and tried to train her by the continuous training method. +With this individual, No. 87, the results were even more discouraging than +those previously obtained, for she was still imperfect in her +discrimination at the end of two hundred and ten tests. At that point the +experiment was interrupted, and it seemed scarcely worth while to continue +it further at a later date. The evidence of the extremely low efficiency +of the continuous method in comparison with the other methods which we +have been considering is so conclusive that further comment seems +superfluous. + +We are now in a position to compare the results of the several methods of +training which have been applied to the dancer, and to attempt to get +satisfactory quantitative expressions of the efficiency of each method. I +have arranged in Table 46 the general averages yielded by the four +methods. Although these general results hide certain important facts which +will be exhibited later, they clearly indicate that an increase in the +number of tests per day does not necessarily result in an increase in the +rapidity of habit formation. Should we attempt, on superficial +examination, to interpret the figures of this table, we would doubtless +say that in efficiency the two-five-test method stands first, the +continuous-test method last, while the ten-test and twenty-test methods +occupy intermediate positions. + + + +TABLE 46 + +EFFICIENCY OF TRAINING + +Number of Errors in White-Black Series for Different Methods of +Training + +SETS OF 10 2 OR 5 TESTS 10 TESTS 20 TESTS 100 OR MORE + PER DAY PER DAY PER DAY TESTS PER DAY + +A 5.8 5.8 5.9 5.3 +B 5.7 5.7 5.9 5.0 + +1 6.4 5.3 4.8 5.7 +2 4.2 3.5 4.8 3.3 +3 3.4 3.2 3.4 5.3 +4 3.1 2.5 3.0 5.3 +5 2.7 2.5 1.9 3.7 +6 1.5 1.3 1.9 1.7 +7 0.9 1.5 0.9 3.0 +8 0.7 0.8 1.1 2.0 +9 0.5 0.4 0.9 2.3 +10 0 0.5 0.8 2.3 +11 0 0.4 0.4 1.7 +12 0 0.2 0.3 1.3 +13 0 0 2.3 +14 0 0 1.3 +15 0 0 3.0 +16 2.7 +17 0.7 +18 1.0 +19 0.7 +20 0 + + + +We may now apply to the results of our efficiency-of-training tables the +method of measuring efficiency which was mentioned at the end of the +preceding chapter as the _index of modifiability (that number of tests +after which no errors occur for at least thirty tests)_. By taking the +average number of tests for the several individuals in each of the Tables +42, 43, 44, and 45 we obtain the following expressions of efficiency:-- + + + +METHOD INDEX OF MODIFIABILITY (EFFICIENCY) + +Two-five-test 81.7 ± 2.7 +Ten-test 88.0 ± 4.1 +Twenty-test 91.0 ± 5.3 +Continuous-test 170.0 ± 4.8 + + + +Since the difference between the indices for the ten-test and the twenty- +test methods lies within the limits of their probable errors (±4.1 and +±5.3) it is evident that it is not significant. Except for this, I think +these indices may be accepted as indications of real differences in the +value of the several methods of training. + +A somewhat different interpretation of our results is suggested by the +grouping of individuals according to sex. In Table 47 appear the general +averages for the males and the females which were tested by the several +methods. The most striking fact exhibited by this table is that of the +high efficiency of the twenty-test method for the females. Apparently they +profited much more quickly by this method than by the ten-test method, +whereas just the reverse is true of the males. I present the data of this +table merely to show that general averages may hide important facts. + + + +TABLE 47 + +EFFICIENCY OF TRAINING + +CONDITION MALES FEMALES + INDEX OF MODIFIABILITY INDEX OF MODIFIABILITY +2 or 5 tests per day 85.0 80.0 +10 tests per day 72.0 104.0 +20 tests per day 94.0 88.0 +100 or more tests per day 160.0 180.0 + + + +From all considerations that have been mentioned thus far the reader would +be justified in concluding that I made a mistake in selecting the ten-test +method for my study of the modifiability of the behavior of the dancer. +That this conclusion is not correct is due to the time factor in the +experiments. If the dancer could acquire a perfect habit as a result of +twelve days' training, no matter whether two, five, ten, or twenty tests +were given daily, it would, of course, be economical of time for the +experimenter to employ the two-test method. But if, on the contrary, the +two-test method required twice as many days' training as the five-test +method, it would be economical for him to use the five-test method despite +the fact that he would have to give a larger number of tests than the two- +test method would have demanded. In a word, the time which the work +requires depends upon the number of series which have to be given, as well +as upon the number of tests in each series. As it happens, the ten-test +method demands less of the experimenter's time than do methods with fewer +tests per day. The twenty-test method is even more economical of time, but +it has a fatal defect. It is at times too tiresome for both mouse and man. +These facts indicate that a balance should be struck between number of +tests and number of series. The fewer the tests per day, within the limits +of two and one hundred, the higher the efficiency of the method of +training, as measured in terms of the total number of tests necessary for +the establishment of a perfect habit, and the lower its efficiency as +measured in terms of the number of series given. The greater the number of +tests per day, on the other hand, the higher the efficiency of the method +in terms of the number of series, and the lower its efficiency in terms of +the total number of tests. By taking into account these facts, together +with the fact of fatigue, we are led to the conclusion that ten tests per +day is the most satisfactory number. + +If my time and attention had not been fully occupied with other problems, +I should have determined the efficiency of various methods of training in +terms of the duration of habit, as well as in terms of the rapidity of its +formation. As these two measures of efficiency might give contradictory +results, it is obvious that a training method cannot be fairly evaluated +without consideration of both the rapidity of habit formation and the +permanency of the habit. A _priori_ it seems not improbable that slowness +of learning should be directly correlated with a high degree of +permanency. By the further application of the method which I have used in +this study of the efficiency of training we may hope to get a definite +answer to this and many other questions concerning the nature of the +educative process and the conditions which influence it. + + + + +CHAPTER XVI + +THE DURATION OF HABITS: MEMORY AND RE-LEARNING + +The effects of training gradually disappear. Habits wane with disuse. In +the dancer, it is not possible to establish with certainty the existence +of memory in the introspective psychological sense; but it is possible to +measure the efficiency of the training to which the animal is subjected, +and the degree of permanency of habits. The materials which constitute +this chapter concern the persistence of unused habits, and the influence +of previous training on the re-acquisition of a habit which has been lost +or on the acquisition of a new habit. For convenience of description, I +shall refer to certain of the facts which are to be discussed as facts of +memory, with the clear understanding that consciousness is not necessarily +implied. By memory, wherever it occurs in this book, I mean the ability of +the dancer to retain the power of adaptive action which it has acquired +through training. + +I first discovered memory in the dancer, although there was previously no +reason for doubting its existence, in connection with the ladder-climbing +tests of Chapter XII. In this experiment two individuals which had +perfectly learned to escape from the experiment box to the nest-box by way +of the wire ladder, when tested after an interval of two weeks, during +which they had remained in the nest-box without opportunity to exercise +their newly acquired habit, demonstrated their memory of the method of +escape by returning to the nest-box by way of the ladder as soon as they +were given opportunity to do so. As it did not lend itself readily to +quantitative study, no attempts were made to measure the duration of this +particular habit. At best the climbing of a wire ladder is of very +uncertain value as an indication of the influence of training. + +Similarly, the persistence of habits has been forced upon my attention day +after day in my various experiments with the mice. It is obvious, then, +that the simple fact of memory is well established, and that we may turn +at once to an examination of the facts revealed by special memory and re- +learning experiments. + +The visual discrimination method, which proved invaluable as a means of +measuring the rapidity of habit formation, proved equally serviceable in +the measurement of the permanency or duration of habits. Memory tests for +discrimination habits were made as follows. After a dancer had been +trained in the discrimination box so that it could choose the correct +electric-box, white, red, blue, or green as it might be, in three +successive daily series of ten tests each, it was permitted to remain for +a certain length of time without training and without opportunity to +exercise its habit of visual discrimination and choice. At the expiration +of the rest interval, as we may designate the period during which the +habit was not in use, the mouse was placed in the discrimination box under +precisely the same conditions in which it had been trained and was given a +series of ten memory tests with the box to be chosen alternately on the +right and on the left. In order that the entire series of ten tests, and +sometimes two such series given on consecutive days, might be available as +indications of the duration of a habit, the mouse was permitted to enter +and pass through either of the electric-boxes without receiving a shock. +Had the shock been given as punishment for a wrong choice, it is obvious +that only the first test of the memory series would be of value as an +indication of the existence of a previously acquired habit. Even under the +conditions of no shock and no stop or hindrance the first test of each +memory series is of preeminent importance, for the mouse tends to persist +in choosing either the side or the visual condition (sometimes one, +sometimes the other) which it chooses in the first test. If the wrong box +is chosen to begin with, mistakes are likely to continue because of the +lack of punishment; in this case the animal discriminates, but there is no +evidence that it remembers the right box. Likewise, if the right electric- +box is chosen in the first test, correct choices may continue simply +because the animal has discovered that it can safely enter that particular +box; again, the animal discriminates without depending necessarily upon +its earlier experience. I have occasionally observed a series of ten +correct choices, made on the basis of an accidental right start, followed +by another series in which almost every choice was wrong, because the +animal happened to start wrong. + +As the results of my tests of memory are of such a nature that they cannot +advantageously be averaged, I have arranged in Table 48 a number of +typical measurements of the duration of visual discrimination habits. In +this table I have indicated the number and age of the individual tested, +the habit of discrimination which had been acquired, the length of the +rest interval, the result of the first test (right or wrong), and the +number of errors made in each series of ten memory tests. + + + +TABLE 48 + +MEASUREMENTS OF THE DURATION OF A HABIT + +Memory + + ERRORS +NO. AGE NAME OF TEST REST FIRST FIRST SECOND + INTERVAL CHOICE SERIES SERIES + +1000 25 weeks White-black 4 weeks Right 0 + 5 27 White-black 4 Right 5 7 + 210 15 White-black 8 Right 5 + 220 15 White-black 8 Right 4 + 230 15 White-black 8 Wrong 5 + 215 15 White-black 8 Right 5 + 225 15 White-black 8 Right 2 + 235 15 White-black 8 Right 7 + 410 15 White-black 8 Wrong 4 + 415 15 White-black 8 Wrong 6 + 420 15 White-black 8 Wrong 3 + 425 15 White-black 8 Right 3 + 2 28 Black-white 4 Wrong 9 + 7 17 Black-white 2 Wrong 1 + 7 21 Black-white 6 Right 1 + 7 27 Black-white 10 Right 1 6 + 998 18 Black-white 2 Wrong 3 + 998 22 Black-white 4 Right 0 + 998 28 Black-white 10 Right 5 5 + 13 10 Black-white 4 Right 3 + 14 10 Black-white 4 Right 3 + 15 10 Black-white 4 Right 2 + 16 10 Black-white 4 Right 4 +1000 25 Light blue-orange 4 Right 4 + 2 28 Light blue-orange 2 Wrong 5 + 5 28 Light blue-orange 6 Wrong 4 6 + 3 25 Light blue-orange 4 Wrong 8 + 10 24 Light blue-orange 2 Right 8 + 10 26 Light blue-orange 2 Right 5 + 11 25 Light blue-orange 2 Right 6 + 11 27 Light blue-orange 2 Wrong 5 + 151 13 Green-red 2 Right 1 0 + 152 13 Green-red 2 Right 5 1 + + + +This quantitative study of the duration of simple habits of choice showed +that in the majority of cases a perfectly acquired habit persists for at +least two weeks. To be perfectly fair to the animal I must restrict this +statement to visual conditions other than colors, for the dancer exhibited +little ability either to acquire or to retain a habit of distinguishing +spectral colors. Altogether, I made a large number of white-black and +black-white memory tests after rest intervals of four, six, eight, or ten +weeks. The results for the four-week interval show extreme individual +differences in memory. Number 1000, for example, was able to choose +correctly every time in a series of white-black tests after a rest +interval of four weeks, whereas No. 5 was wrong as often as she was right +after the same interval. I have placed the results for these two +individuals at the head of the table because they suggest the variations +which render averages undesirable. Number 1000 had a perfect habit at the +end of four weeks of disuse; No. 5 had no habit whatever. I shall reserve +further discussion of age, sex, and individual differences in the +permanency of habits for the next chapter. + +With Nos. 7 and 998 memory tests were made after three different rest +intervals. At the end of two weeks the black-white habit was present in +both individuals, although it was not perfect. After six and four weeks, +respectively (see Table 48), it still persisted; in fact, it apparently +had improved as the result of additional training after the earlier memory +tests. At the expiration of ten weeks it had wholly disappeared. In her +first series of memory tests after the ten-week interval No. 7 made only +one error, but a chance choice of the black (right) in the first test and +the subsequent choice of the box in which no shock had been received serve +to account for results which at first appear to be indicative of memory. +That this explanation is correct is proved by the fact that a second +memory series, in which the first choice happened to be wrong, resulted in +six mistakes. Evidently she had lost the habit. + +In no instance have memory tests definitely indicated the presence of a +habit after a rest interval of more than eight weeks. It is safe, +therefore, to conclude from the results which have been obtained that a +white-black or black-white discrimination habit may persist during an +interval of from two to eight weeks of disuse, but that such a habit is +seldom perfect after more than four weeks. + +The measurements of memory which were made in connection with color +discrimination experiments are markedly different from those which were +obtained in the brightness tests. As might have been anticipated (?), in +view of the extreme difficulty with which the dancer learns to +discriminate colors, the habit of discriminating between qualitatively +different visual conditions does not persist very long. I have never +obtained evidence of a perfect habit after an interval of more than two +weeks, and usually, as is apparent from Table 48, the tests indicated very +imperfect memory at the end of that interval. It seems probable that even +in these so-called color tests discrimination is partly by brightness +difference, and that the imperfection of the habit and its short duration +are due to the fact that the basis of discrimination is inadequate. This +is the only explanation which I have to offer for the difference which has +been demonstrated to exist between the duration of brightness +discrimination habits and color discrimination habits. + +The duration of a discrimination habit having been measured with a fair +degree of accuracy, I undertook the task of ascertaining whether training +whose results have wholly disappeared, so far as memory tests are in +question, influences the re-acquisition of the same habit. Can a habit be +re-acquired with greater facility than it was originally acquired? Is re- +learning easier than learning? To obtain an answer to the question which +may be asked in these different forms, ten individuals were experimented +with in accordance with a method whose chief features are now to be +stated. In each of these ten individuals a perfect white-black habit was +established by the use of the standard series of tests the order of which +is given in Table 12. At the expiration of a rest interval of eight weeks +precisely the same series of tests were repeated as memory and re-training +tests. In this repetition, the preliminary series, _A_ and _B_, served as +memory tests, and the subsequent training series, as re-training series. + +[Illustration: FIGURE 32.--Error curves plotted from the data given by ten +dancers in white-black discrimination tests. The solid line ([Symbol: +solid line]) is the error curve of the original learning process; the +broken line (------) is that of the re-learning process, after an interval +of eight weeks.] + +The striking results of this investigation of re-learning are exhibited in +the curves of learning and re-learning of Figure 32. These curves make it +appear that the mice re-acquired the white-black discrimination habit much +more readily than they had originally acquired it. But in addition to +furnishing the basis for some such statement as the foregoing, the curves +suggest a serious criticism of the experiment. + +In the original tests, the preliminary series indicated a strong +preference for black. In series _A_ it was chosen on the average 5.8 times +in 10, and in series _B_, 5.7 times. This preference was rapidly overcome +by the training series, and at the end of 130 tests discrimination was +perfect. All this appears in the curve of learning (solid line of figure). +On the other hand, these preliminary series when repeated as memory tests, +after a rest-interval of eight weeks, gave markedly different results. +Series _A_ indicated preference for white (5.6 times in 10) instead of +black, and series _B_ indicated only a slight preference for black. In +brief, series _A_ and _B_ show that the preference for black was +considerably stronger at the beginning of the training than at the +beginning of the re-training. + +In the light of these facts it is fair to claim that the effects of the +white-black training had not wholly disappeared as the result of eight +weeks of rest, and that the experiment therefore fails to furnish +satisfactory grounds for the statement that re-learning occurs more +rapidly than learning. I accept this criticism as pertinent, although not +necessarily valid, and at the same time I freely admit that the results +have a significance which I had not anticipated. But they are not less +interesting or valuable on that account. Granting, then, that at least +some of the ten individuals which took part in the experiment had not +completely lost the memory of their white-black training at the end of +eight weeks, it is still possible that an examination of the individual +results may justify some conclusion concerning the question which was +proposed at the outset of the investigation. Such an examination is made +possible by Tables 49 and 50, in which I have arranged separately the +results for the males and the females. + + + + +TABLE 49 + +WHITE-BLACK TRAINING. TEN TESTS PER DAY + + + Males + TRAINING RETRAINING + 210 220 230 410 420 AV. 210 220 230 410 420 AV. + + + A 6 5 6 6 6 5.8 5 4 5 4 3 4.2 + B 6 8 8 5 1 5.6 8 4 5 4 6 5.4 + + 1 6 7 6 2 4 5.0 3 3 4 7 3 4.0 + 2 4 3 1 2 3 2.6 2 4 2 5 3 3.2 + 3 3 1 4 3 4 3.0 1 4 1 4 1 2.2 + 4 5 0 3 3 2 2.6 0 1 0 1 2 0.8 + 5 3 0 4 1 4 2.4 0 2 0 2 0 0.8 + 6 2 1 4 0 1 1.6 0 1 0 0 2 0.6 + 7 1 0 3 1 0 1.0 0 0 0 0 + 8 0 0 1 0 0 0.2 0 0 1 0.2 + 9 0 0 0 1 0 0.2 0 0 0 +10 0 0 0 0 1 0.2 +11 0 0 0 0 0 +12 0 0 0 0 +13 0 0 +14 +15 + + + +Only three of the ten individuals failed to re-acquire the habit of white- +black discrimination more quickly than it had originally been acquired, +and, in the case of these exceptions, No. 220 required exactly the same +number of tests in each case, and No. 420 was placed at a slight +disadvantage in the re-learning series by an interruption of the training +between the seventh and the eighth series. Had his training been completed +by the sixth series he too would have had the same number of tests in +training and re-training. Moreover, and this is of preëminent importance +for a fair interpretation of the results, in several instances even those +individuals which exhibited as strong a preference for the black in the +memory series as in the preliminary series re-learned more quickly than +they had learned. Number 210, for example, although he gave no evidence of +memory, and, in fact, chose the black more frequently in the memory series +than he did in the preliminary series, re-acquired the discrimination +habit in less than half the number of tests which had been necessary for +the establishment of the habit originally. + + + + TABLE 50 + + WHITE-BLACK TRAINING. TEN TESTS PER DAY + + Females + + TRAINING RE-TRAINING + 215 225 235 415 425 Av. 215 225 235 415 425 Av. + A 8 4 4 8 5 5.8 5 2 7 6 3 4.6 + B 8 7 6 6 2 5.8 8 5 6 4 3 5.2 + 1 7 6 5 6 4 5.6 4 1 5 4 3 3.4 + 2 5 6 4 2 5 4.4 1 1 1 2 3 1.6 + 3 3 3 4 3 4 3.4 1 0 3 6 0 2.0 + 4 2 1 3 3 3 2.4 0 0 3 3 1 1.4 + 5 1 3 3 3 3 2.6 0 0 died 2 0 0.5 + 6 2 1 1 1 0 1.0 0 1 0 0.2 + 7 1 1 2 3 3 2.0 0 0 0 + 8 0 0 2 2 3 1.4 1 0.2 + 9 1 0 0 1 1 0.6 0 0 +10 0 2 1 0 2 1.0 0 0 +11 0 3 0 1 0 0.8 0 0 +12 0 0 0 2 0 0.4 +13 0 0 0 0 0 +14 0 0 0 +15 0 0 + + + +The facts which have been presented thus far become more significant when +the indices of modifiability for the learning and the re-learning +processes are compared. + + + INDICES OF MODIFIABILITY + + LEARNING RE-LEARNING + +Females . . . . . . . 104 42.5 +Males . . . . . . . . 72 54 + + +The behavior of the mice in the experiments, the detailed results of +Tables 49 and 50, and the indices of modifiability together justify the +following conclusions. Most of the ten dancers, at the end of a rest +interval of eight weeks, had so far lost the habit of white-black +discrimination that memory tests furnished no conclusive evidence of the +influence of previous training; a few individuals seemed to possess traces +of the habit after such an interval. In the case of each group of +individuals re-training brought about the establishment of a perfect habit +far more quickly than did the original training. This suggests the +existence of two kinds or aspects of organic modification in connection +with training; those which constitute the basis of a definite form of +motor activity, and those which constitute the bases or dispositions for +the acquirement of certain types of behavior. There are several +indications that further study of the modifiability of behavior will +furnish the facts which are necessary to render this suggestion +meaningful. + +Closely related to the facts which have been revealed by the re-training +experiments are certain results of the labyrinth experiments. For the +student of animal behavior, as for the human educator, it is of importance +to learn whether one kind of training increases the efficiency of similar +forms of training. Can a dancer learn a given labyrinth path the more +readily because it has previously had experience in another form of +labyrinth? + +The answer to this question, which my experimental results furnish, is +given in Table 51. In the upper half of the table have been arranged the +results for six individuals which were trained first in labyrinth B, then +in labyrinth C, and finally in labyrinth D. Below, in similar fashion, are +given the results for six individuals which were trained in the same three +labyrinths in the order C, B, D, instead of B, C, D. My purpose in giving +the training in these two orders was to ascertain whether labyrinth C, +which had proved to be rather difficult for most individuals, would be +more easily learned if the training in it were preceded by training in +labyrinth C. + + + +TABLE 51 + +THE INFLUENCE OF ONE LABYRINTH HABIT UPON THE FORMATION +OF ANOTHER + LABYRINTH B LABYRINTH C LABYRINTH D + NO. OF NO. OF NO. OF NO. OF NO. OF NO. OF +NO. FIRST COR- LAST OF FIRST COR- LAST OF FIRST COR- LAST OF + RECT TEST FIVE COR- RECT TEST FIVE COR- RECT TEST FIVE COR- + RECT TESTS RECT TESTS RECT TESTS + +76 8 14 3 19 4 7 +78 5 20 6 14 4 5 +86 13 22 5 12 3 9 +75 4 15 8 19 4 13 +77 7 11 11 29 11 12 +87 12 22 9 20 4 9 + +AV. 8.2 17.3 7.0 18.8 5.0 9.2 + + + LABYRINTH C LABYRINTH B LABYRINTH D + +58 16 -- 2 14 7 10 +60 17 -- 13 37 10 14 +88 25 35 9 22 4 8 +49 34 -- 1 5 7 8 +57 15 -- 3 20 3 6 +85 11 18 2 11 3 4 + +AV. 19.7 26.5 5.0 18.2 5.7 8.3 + + + +The results are sufficiently definite to warrant the conclusion that +experience in B rendered the learning of C easier than it would have been +had there been no previous labyrinth training. Those individuals whose +first labyrinth training was in C made their first correct trip as the +result of 19.7 trials, whereas those which had previously been trained in +labyrinth B were able to make a correct trip as the result of only 7.0 +trials. Similarly the table shows that training in C rendered the +subsequent learning of B easier. To master B when it was the first +labyrinth required 8.2 trials; to master it after C had been learned +required only 5 trials. In addition to proving that the acquisition of one +form of labyrinth habit may facilitate the acquisition of others, +comparison of the averages of Table 51 furnishes evidence of the truth of +the statement that no results of training can be properly interpreted in +the absence of knowledge of the previous experience of the organism. + + + + +CHAPTER XVII + + +INDIVIDUAL, AGE, AND SEX DIFFERENCES IN BEHAVIOR + +All dancers are alike in certain important respects, but to the trained +observer of animal behavior their individual peculiarities are quite as +evident, and even more interesting than their points of resemblance. +Omitting consideration of the structural marks of individuality, we shall +examine the individual, age, and sex differences in general behavior, +rapidity of learning, memory, and discrimination, which have been revealed +by my experiments. Observations which bear on the subject of differences +are scattered through the preceding chapters, but in no case have they +been given sufficient prominence to force them upon the attention of those +who are not especially interested in individual peculiarities. It has +seemed worth while, therefore, to assemble all the available material in +this chapter for systematic examination and interpretation. + +In the pages which follow, individual, age, and sex peculiarities are +discussed in turn. Within each of these three groups of differences I have +arranged in order what Royce has appropriately named the facts of +discriminating sensitiveness, docility, and initiative. Individuals of the +same age and sex no less than those which differ in sex or age exhibit +important differences in ability to discriminate among sense impressions +("discriminative sensitiveness"), in ability to profit by experience +("docility"), and in ability to try new kinds of behavior ("initiative"). + +Individual differences in sensitiveness to visual, auditory, tactual, and +olfactory stimuli have been revealed by many of my experiments. The +brightness discrimination tests conclusively proved that a degree of +difference in illumination which is easily detectable by one dancer may be +beyond the discriminating sensitiveness of another. Both the tests with +gray papers and those with the Weber's law apparatus furnished striking +evidence of individual differences in the kind of visual sensitiveness +which throughout this book has been called brightness vision. I suspect +that certain of the differences which were observed should be referred to +the experience of the individuals rather than to the capacity of the +visual organs, for training improves visual discrimination to a much +greater extent than would ordinarily be thought possible. To the truth of +this statement the results of the Weber's law experiments with No. 51 bear +witness. Likewise in color discrimination there are individual +differences, examples of which may be discovered by the examination of the +results given in Chapters IX and X. + +No differences in auditory sensitiveness appeared in my adult dancers, for +in none of them was there definite response to sounds, but among the young +individuals differences were prominent. I may call attention to the data +on this subject which Table 5, p. 89, contains. The mice in four out of +twelve litters gave no indications of hearing any sounds that I was able +to produce; the remaining individuals responded with varying degrees of +sensitiveness. I made no attempt to measure this sensitiveness, but it +obviously differed from mouse to mouse. I feel justified, therefore, in +stating that the young dancers exhibit extreme individual difference in +sensitiveness to sounds. + +My observations of differences in sensitiveness to other forms of +stimulation were made in connection with training tests, and although they +are not quantitative, I venture to call attention to them. Indeed, I am +led by the results of my study of various aspects of the dancer's behavior +to conclude that the race exhibits individual differences in +discriminating sensitiveness to a far greater extent than do most mammals, +not excepting man. The importance of this fact (for I am confident that +any one who carefully examines the detailed results of the various +experiments which are described in this book will agree that it is an +established fact) cannot be overlooked. It alters our interpretation of +the results of training, memory, heredity, and discrimination experiments, +and it leads us to suspect that the dancing race is exceedingly unstable. +I do not venture to make comparison of my own observations of the dancer's +sense equipment with those of Cyon, Rawitz, Zoth, and Kishi, for the +differences are too great in many instances to be thought of as other than +species or variety peculiarities. It has seemed fairer to compare only +individuals of the same breed, or, as I have done and shall continue to do +throughout this chapter, of two lines of descent. + +With respect to docility individual differences are prominent. We need +only turn to the various tables of results to discover that in +modifiability of behavior, in memory, in re-learning, not to mention other +aspects of docility, dancers of the same sex and age differed strikingly. +Let me by way of illustration cite a few cases of difference in docility. +Number 1000 learned to discriminate white from black more quickly and +retained his habit longer than any other dancer with which I have +experimented. I should characterize him as an exceptionally docile +individual. Table 44 offers several examples. Numbers 403 and 407, though +they were born in the same litter and were alike in appearance and in +conditions of life, acquired the white-black habit with a difference in +rapidity which is expressed by the indices of modifiability 50 and 100. In +other words, it took No. 407 twice as long to acquire this habit as it +took No. 403. Similarly the ladder-climbing tests revealed important +individual differences in ability to profit by experience. In the tables +of labyrinth tests (38, 39, 40) individual differences are too numerous to +mention. It required forty-nine tests to establish in No. 50 a labyrinth-C +habit which was approximately equal in degree of perfection to that which +resulted from twenty-two tests in the case of No. 52. The figures in this +and other instances do not exaggerate the facts, for repeatedly I have +tested individuals of the same litter, the same sex, and, so far as I +could judge, of the same stage of development, and obtained results which +differ as markedly as do those just cited. If space limits permitted, I +could present scores of similar differences in docility which the problem, +labyrinth, and discrimination methods have revealed. + +In examining the detailed individual results of the various tables for +differences of this sort, it is important to bear in mind that sex, age, +and descent should be taken into account, for with each of them, as will +be shown clearly later in this chapter, sensitiveness, docility, and +initiative vary. I have therefore based my statements concerning +individual differences in docility upon the results of comparison of mice +of the same litter, sex, and age. It is safe to say that human beings +similarly selected for comparison do not exhibit greater differences in +ability to profit by experience than did these dancing mice. + +The facts concerning individual differences in initiative which I have +discovered are not less definite than those of the preceding paragraphs. +From the beginning of my study of the dancer I observed that what one +individual would readily learn of his own initiative another never +learned. For example, in the ladder-climbing experiment No. 1000 +distinguished himself for his initiative, whereas Nos. 4 and 5 never +acquired the habit of escaping from confinement by using the ladder. I +noticed, in this test of the animal's ability to learn, that while one +individual would be scurrying about trying all ways of escape, +investigating its surroundings, looking, sniffing, and dancing by turns, +another would devote all its time to whirling, circling, or washing +itself. One in the course of its activity would happen upon the way of +escape, the other by reason of the limited scope of its activity, not the +lack of it, would fail hour after hour to discover even the simplest way +of getting back to its nest, to food, and to its companions. Hundreds of +times during the past three years I have noticed important individual +differences in initiative in connection with the discrimination +experiments. The swinging wire doors which one dancer learned to push open +before he had been in the box five minutes, another might not become +familiar with through his own initiative for hours or days. In fact, it +was not seldom that I had to teach an individual to pass from one +compartment to the other by gently pushing him against the door until it +opened sufficiently to allow him to squeeze through. Occasionally a mouse +learned to pull the doors open so that he could pass through the openings +in either direction with facility. This was a form of individual +initiative which I had not anticipated and did not especially desire, so I +did not encourage its development, but, nevertheless, at least one fourth +of the mice which I experimented with in the discrimination box learned +the trick. The other three fourths, although they were used in the box day +after day sometimes for weeks, never discovered that they might return to +the nest-box by pulling the swing-door through which they had just passed +as well as by entering one of the electric-boxes. + +Another indication of individual initiative in action appeared in the +tendency of certain mice to climb out of the experiment boxes or +labyrinths. It would have been extremely easy for any of the mice to +escape from the labyrinths by scaling the walls of the alleys, for they +were only 10 cm. in height, and when a dancer stood on its hind legs it +could easily reach the top with its nose. But, strange though it will seem +to any one who has not worked with the dancer, not more than one in ten of +the animals which I observed made any attempt to escape in this manner. +They lacked initiative. That it was not due to a lack of the power to +climb, I abundantly demonstrated by teaching a few individuals that a +scramble in one corner meant easy escape from the maze of paths. I do not +think any one of the mice was physically incapable of climbing, but I am +confident that they differed markedly, not only in the willingness to try +new modes of action, but in the readiness with which they could climb. I +have already said that individuals differ noticeably in the scope of their +activity. By this statement I mean that they try a varying number of kinds +of activity. As in the case of men, so in mice, one individual will do a +greater number of things in a few hours than another will in weeks or +months. The dancers differ in versatility, in individual initiative, as do +we, albeit not so markedly. + +Important differences which may with certainty be described as age +differences are not so obvious as are such marks of individuality as have +been set forth in the preceding pages. I have noted few changes in +discriminative sensitiveness, other than those with regard to auditory +sensitiveness, which could be correlated with age. In certain instances +adults appeared to be able to discriminate more accurately and more easily +than young mice, but it is difficult to say whether this change belongs +under sensitiveness or docility. I have not made an ontogenetic study of +the senses, and I am therefore unable to describe in detail the course of +their development and decline. Of one important fact I am certain, that +discriminative sensitiveness increases up to a certain point with age and +with training. + +Differences in docility which are obviously to be correlated with age +abound. In the prime of its life (from the second to the tenth month) the +dancer is active, full of energy, quick to learn; in its senility (during +the second year) it is inactive, but at times even more docile than during +the period of greatest physical development. Frequently I have noticed in +connection with labyrinth tests that individuals of the age of a year or +more learn much more quickly than do individuals of the age of two or +three months. But, on the other hand, I have contradictory observations, +for now and then I obtained just the opposite result in experiments to +test docility. Evidently this is a matter which demands systematic, +quantitative investigation. Casual observation may suggest conclusions, +but it will not justify them. + +Early in my investigation of the behavior of the dancer I conceived the +idea of determining the relation of modifiability of behavior (docility) +to age. The question which was foremost in my mind and for which I first +sought an answer may be stated thus: can the dancer acquire a given habit +with the same facility at different ages? Since the visual discrimination +experiment seemed to be well suited for the investigation of this problem +I planned to train, in the white-black discrimination experiment, five +pairs of dancers at the age of one month, and the same number for each of +the ages four, seven, ten, thirteen, sixteen, and nineteen months.[1] + +[Footnote 1: I have not been able thus far to determine the average length +of the dancer's life. The greatest age to which any of my individuals has +attained is nineteen months.] + +To test the same individuals month after month would be the ideal way of +obtaining an answer to our question, but I could devise no satisfactory +way of doing this. The effects of training last so long, as the results of +the previous chapter proved, and the uncertainty of their entire +disappearance is so serious, that the same training process cannot be used +at successive ages. The use of different methods of training is even more +unsatisfactory because it is extremely difficult to make accurate +quantitative comparison of their results. It was these considerations that +forced me to attempt to discover the relation of docility to age by +carrying out the same experiments with groups of individuals of different +ages. + +As my plan involved the execution of precisely the same set of tests with +at least seventy individuals whose age, history, and past experience were +accurately known, and of which some had to be kept for nineteen months +before they could be trained, the amount of labor and the risk of mishap +which it entailed were great. To make possible the completion of the +investigation within two years, I accumulated healthy individuals for +several months without training any of them. In March, 1907, I had +succeeded in completing the tests for the age of one month, and I had on +hand for the remaining tests almost a hundred individuals, whose ages +ranged from a few days to eighteen months. Had everything gone well, the +work would have been finished within six months. Suddenly, and without +discoverable external cause, my mice began to die of an intestinal +trouble, and despite all my efforts to check the disease by changing food +supply and environment, all except a single pair died within a few weeks. +Thus ended a number of experiments whose final results I had expected to +be able to present in this volume. However, the work which I have done is +still of value, for the single pair of survivors have made possible the +continuance of my tests with other individuals of the same line of descent +as those which perished, and I have to regret only the loss of time and +labor. + +As I have on hand results for ten individuals of the age of one month, and +for four individuals of the age of four months, it has seemed desirable to +state the problem, method, and incomplete results of this study of the +relation of modifiability to age. The indices of modifiability for these +two groups of dancers differ so strikingly that I feel justified in +persisting in my efforts to obtain comparable data for the seven ages +which have been mentioned. + + + + TABLE 52 + + PLASTICITY (RELATION or MODIFIABILITY TO AGE) + +Number of Errors in Successive Daily Series of Ten White-Black + Tests, with Dancers Four Months Old + + + + SERIES MALES FEMALES + + NO. 76 NO. 78 AV. NO. 75 NO. 77 AV. GENERAL AV. + + A 7 7 7.0 4 8 6.0 6.50 + B 8 6 7.0 6 5 5.5 6.25 + + 1 5 5 5.0 5 5 5.0 5.00 + 2 5 4 4.5 2 2 2.0 3.25 + 3 4 5 4.5 2 5 3.5 4.00 + 4 3 4 3.5 1 1 1.0 2.25 + 5 5 2 3.5 0 1 0.5 2.00 + 6 3 2 2.5 1 0 0.5 1.50 + 7 2 1 1.5 1 2 1.5 1.50 + 8 5 1 3.0 0 0 0 1.50 + 9 1 3 2.0 0 0 0 1.00 + 10 1 2 1.5 1 0 0.5 1.00 + 11 1 1 1.0 0 0 0.50 + 12 1 1 1.0 0 0 0.50 + 13 0 0 0 0 0 0 + 14 0 0 0 0 + 15 0 0 0 0 + + + +[Illustration: FIGURE 33.--Plasticity curves. In the left margin are given +the indices of modifiability (the number of tests necessary for the +establishment of a perfect habit). Below the base line the age of the +individuals is given in months. Curve for males, --•--•--•--; curve for +females, - - - -; curve for both males and females,----. When these three +plasticity curves are completed, they will represent the indices of +modifiability as determined for ten individuals at the age of 1 month, and +similarly for the same number of individuals at each of the ages, 4, 7, +10, 13, 16, and 19 months.] + +The detailed results for the one-month old individuals appear in Table 43; +those for the four-month individuals in Table 52. The general averages for +the former are to be found in the third column of Table 46, under the +heading "10 tests per day"; those for the latter in the last column of +Table 52. Mere inspection of these tables reveals the curious sex +difference which goes far towards justifying the presentation of this +uncompleted work. The index of modifiability for the ten one-month +individuals is 88 (that is, 88 tests were necessary for the establishment +of a habit); for the four-month individuals it is 102.5. The heavy solid +line of Figure 33 joins the points on the ordinates at which these values +are located. Apparently, then, the dancer acquires the white-black +discrimination habit less readily at the age of four months than at the +age of one month. + +Further analysis of the results proves that this statement is not true. +When the averages for the two sexes are compared, it appears that the +males learned much less quickly at four months than at one month, whereas +just the reverse is true of the females. The dash and dot line of the +figure extends from the index of modifiability of the one-month males (72) +to that of the four-month males (120); and the regularly interrupted line +similarly joins the indices of the one-month (104) and the four-month (85) +females. In seeking to discover age differences in docility or ability to +profit by experience we have stumbled upon what appears to be an important +sex difference. Perhaps I should add to this presentation of partial +results the following statement. Since there are only four individuals in +the four-month group, two of each sex, the indices are not very reliable, +and consequently too much stress should not be laid upon the age and sex +differences which are indicated. + +In view of this impressive instance of the way in which averages may +conceal facts and lead the observer to false inferences, I wish to remark +that my study of the dancer has convinced me of the profound truth of the +statement that the biologist, whether he be psychologist, anthropologist, +physiologist, or morphologist, should work with the organic individual and +should first of all deal with his results as individual results. Averages +have their place and value, but to mass data before their individual +significance has been carefully sought out is to conceal or distort their +meaning. Too many of us, in our eagerness for quantitative results and in +our desire to obtain averages which shall justify general statements, get +the cart before the horse. + +Figure 33 presents the beginning of what I propose to call plasticity +curves. When these three curves are completed on the basis of experiments +with five dancers of each sex for each of the ages indicated on the base +line of the figure, they will indicate what general changes in plasticity, +modifiability of behavior, or ability to learn (for all of these +expressions have been used to designate much the same capacity of the +organism) occur from the first month to the nineteenth in the male and the +female dancer, and in the race without respect to sex. So far as I know, +data for the construction of plasticity curves such as I hope in the near +future to be able to present for the dancing mouse have not been obtained +for any mammal. + +At present it would be hazardous for me to attempt to state any general +conclusion concerning the relation of docility to age. + +The initiative of the dancer certainly varies with its age. In scope the +action system rapidly increases during the first few months of life, and +if the animal be subjected to training tests, this increase may continue +well into old age. The appearance of noticeable quiescence does not +necessarily indicate diminished initiative. Frequently my oldest mice have +shown themselves preëminent in their ability to adjust their behavior to +new conditions. However, I have not studied individuals of more than +eighteen months in age. One would naturally expect initiative to decrease +in senility. All that I can say is that I have seen no indications of it. + +We may now briefly consider the principal sex differences which have been +revealed by the experiments. In sensitiveness I have discovered no +difference, but it should be stated that no special attention has been +given to the matter. In docility the males usually appeared to be superior +to the females. This was especially noticeable early in my visual +discrimination tests. The males almost invariably acquired a perfect habit +quicker than the females. I may cite the following typical instances. +Number 14 acquired the black-white habit with 40 tests; No. 13, with 60 +(Table 10, p. 109). Of the five pairs of individuals whose records in +white-black training appear in Table 43, not one contradicts the statement +which has just been made. It is to be noted, however, that under certain +conditions of training, for example, 20 tests per day, the female is at an +advantage. Recently I have with increasing frequency obtained measures of +docility which apparently favor the female. That this difference in the +results is due to a difference in age is probable. + +In labyrinth tests the female is as much superior to the male as the male +is to the female in discrimination tests. From the tables of Chapter XIII +I may take a few averages to indicate the quantitative nature of this +difference. A degree of proficiency in labyrinth B attained by the males +after 7.0 trials was equaled by the females after 6.2 trials. In labyrinth +C the males acquired a habit as a result of 18.7 trials; the females, as a +result of 13.8. And similarly in labyrinth D, 6.1 trials did no more for +the males than 5.9 did for the females. + +That at the age of about one month the male dancer should be able to +acquire a visual discrimination habit more rapidly than the female, +whereas the female can acquire a labyrinth habit more readily than the +male, suggests an important difference in the nature of their equipment +for habit formation. One might hazard the suggestion that the male depends +more largely upon discrimination of external conditions, whereas the +female depends to a greater extent than does the male upon the internal, +organic changes which are wrought by acts. At any rate the female seems to +follow a labyrinth path more mechanically, more accurately, more easily, +and with less evidence of sense discrimination than does the male. + +Finally, in concluding this chapter, I may add that in those aspects of +behavior which received attention in the early chapters of this volume the +dancers differ very markedly. Some climb readily on vertical or inclined +surfaces to which they can cling; others seldom venture from their +horizontally placed dance floor. Some balance themselves skillfully on +narrow bridges; others fall off almost immediately. My own observations, +as well as a comparison of the accounts of the behavior of the dancer +which have been given by Cyon, Zoth, and other investigators, lead me to +conclude that there are different kinds of dancing mice. This may be the +result of crosses with other species of mice, or it may be merely an +expression of the variability of an exceptionally unstable race. + +I can see no satisfactory grounds for considering the dancer either +abnormal or pathological. It is a well-established race, with certain +peculiarities to which it breeds true; and no pathological structural +conditions, so far as I have been able to learn, have been discovered. + +I have presented in this chapter on differences a program rather than a +completed study. To carry out fully the lines of work which have been +suggested by my observations and by the presentation of results would +occupy a skilled observer many months. I have not as yet succeeded in +accomplishing this, but my failure is not due to lack of interest or of +effort. + + + + +CHAPTER XVIII + +THE INHERITANCE OF FORMS OF BEHAVIOR + +In a general way those peculiarities of behavior which suggested the name +dancing mouse are inherited. Generation after generation of the mice run +in circles, whirl, and move the head restlessly and jerkily from side to +side. But these forms of behavior vary greatly. Some individuals whirl +infrequently and sporadically; others whirl frequently and persistently, +at certain hours of the day. Some are unable to climb a vertical surface; +others do so readily. Some respond to sounds; others give no indications +of ability to hear. I propose in this chapter to present certain facts +concerning the inheritance of individual peculiarities of behavior, and to +state the results of a series of experiments by which I had hoped to test +the inheritance of individually acquired forms of behavior. + +My study of the nature of the whirling tendency of the dancer has revealed +the fact that certain individuals whirl to the right almost uniformly, +others just as regularly to the left, and still others now in one +direction, now in the other. On the basis of this observation, the animals +have been classified as right, left, or mixed whirlers. Does the dancer +transmit to its offspring the tendency to whirl in a definite manner? + +Records of the direction of whirling of one hundred individuals have been +obtained. For twenty of these mice the determination was made by counting +the number of complete turns in five-minute intervals at six different +hours of the day. For the remaining eighty individuals the direction was +discovered by observation of the activity of the animals for a brief +interval at five different times. Naturally, the former results are the +more exact; in fact, they alone have any considerable quantitative value. +But for the problem under consideration all of the determinations are +sufficiently accurate to be satisfactory. + +The distribution of the individuals which were examined as to direction of +whirling is as follows. + + + + RIGHT WHIRLERS LEFT WHIRLERS MIXED WHIRLERS TOTAL + +Males 19 19 12 50 +Females 12 23 15 50 + + +The frequency of occurrence of left whirlers among the females is +unexpectedly high. Is this to be accounted for in terms of inheritance? In +my search for an answer to this question I followed the whirling tendency +from generation to generation in two lines of descent. These two groups of +mice have already been referred to as the 200 line and the 400 line. The +former were descended from Nos. 200 and 205, and the latter from Nos. 152 +and 151. Individuals which resulted from the crossing of these lines will +be referred to hereafter as of mixed descent. There were some striking +differences in the behavior of the mice of the two lines of descent. As a +rule the individuals of the 200 line climbed more readily, were more +active, danced less vigorously, whirled less rapidly and less +persistently, and were in several other respects much more like common +mice than were the individuals of the 400 line. It is also to be noted +(see Table 5) that few of the litters of the 200 line exhibited auditory +reactions, whereas almost all of the litters of the 400 line which were +tested gave unmistakable evidence of sensitiveness to certain sounds. +These differences at once suggest the importance of an examination of the +whirling tendency of each line of descent. + +The results for the several generations of each line which I had +opportunity to examine are unexpectedly decisive so far as the question in +point is concerned. + + + + INDIVIDUALS OF THE 200 LINE + + MALES FEMALES + +First generation No. 200, ? No. 205, ? +Second generation No. 210, Mixed whirler No. 215, Left whirler +Third generation No. 220, Mixed whirler No. 225, Mixed whirler +Fourth generation No. 230, Right whirler No. 235, Mixed whirler +Fifth generation No. 240, Right whirler No. 245, Left whirler + + INDIVIDUALS OF THE 400 LINE + + MALES FEMALES + +First generation No. 152, Left whirler No. 151, Left whirler +Second generation No. 410, Left whirler No. 415, Right whirler +Third generation No. 420, Left whirler No. 425, Left whirler + + + +One line of descent exhibited no pronounced whirling tendency; the other +exhibited a strong tendency to whirl to the left. Are these statements +true for the group of one hundred individuals whose distribution among the +three classes of whirlers has been given? In order to obtain an answer to +this question I have reclassified these individuals according to descent +and direction of whirling. + + + + INDIVIDUALS OF THE 200 LINE + + RIGHT WHIRLERS LEFT WHIRLERS MIXED WHIRLERS TOTAL + +Males 7 6 8 21 +Females 5 8 8 21 + 12 14 16 42 + + + + + INDIVIDUALS OF THE 400 LINE + + RIGHT WHIRLERS LEFT WHIRLERS MIXED WHIRLERS TOTAL + +Males 4 9 1 14 +Females 6 9 4 19 + 10 18 5 33 + + INDIVIDUALS OF MIXED DESCENT + + 9 10 6 25 + + +Three interesting facts are indicated by these results: first, the +inheritance of a tendency to whirl to the left in the 400 line of descent; +second, the lack of any definite whirling tendency in the 200 line; and +third, the occurrence of right and left whirlers with equal frequency as a +result of the crossing of these two lines of descent. + +It is quite possible, and I am inclined to consider it probable, that the +pure dancer regularly inherits a tendency to whirl to the left, and that +this is obscured in the case of the 200 line by the influences of a cross +with another variety of mouse. It is to be noted that the individuals of +the 200 line were predominantly mixed whirlers, and I may add that many of +them whirled so seldom that they might more appropriately be classed as +circlers. + + + +THE INHERITANCE OF INDIVIDUALLY ACQUIRED FORMS OF +BEHAVIOR + +The white-black discrimination experiments which were made in connection +with the study of vision and the modifiability of behavior were so planned +that they should furnish evidence of any possible tendency towards the +inheritance of modifications in behavior. The problem may be stated thus. +If a dancing mouse be thoroughly trained to avoid black, by being +subjected to a disagreeable experience every time it enters a black box, +will it transmit to its offspring a tendency to avoid black? + +Systematic training experiments were carried on with individuals of both +the 200 and 400 lines of descent. For each of these lines a male and a +female were trained at the age of four weeks to discriminate between the +white and the black electric-boxes and to choose the former. After they +had been thoroughly trained these individuals were mated, and in course of +time a male and female, chosen at random from their first litter, were +similarly trained. All the individuals were trained in the same way and +under as nearly the same conditions as could be maintained, and accurate +records were kept of the behavior of each animal and of the number of +errors of choice which it made in series after series of tests. What do +these records indicate concerning the influence of individually acquired +forms of behavior upon the behavior of the race? + + + +TABLE 53 + +THE INHERITANCE OF THE HABIT OF WHITE-BLACK DISCRIMINATION + +Number of Errors in Daily Series of Ten Tests + + + MALES FEMALES + +SERIES FIRST SECOND THIRD FOURTH FIRST SECOND THIRD FOURTH + GENERA- GENERA- GENERA- GENERA- GENERA- GENERA- GENERA- GENERA- + TION TION TION TION TION TION TION TION + + No. 210 No. 220 No. 230 No. 240 No. 215 No. 225 No. 235 No. 245 + + A 6 5 6 7 8 4 4 7 + B 6 8 8 8 8 7 6 5 + + 1 6 7 6 5 7 6 5 4 + 2 4 3 1 5 5 6 4 5 + 3 3 1 4 5 3 4 4 3 + 4 5 0 3 4 2 1 3 1 + 5 3 0 4 2 1 3 3 0 + 6 2 1 4 2 2 1 1 1 + 7 1 0 3 1 1 1 2 0 + 8 0 0 1 0 0 0 2 3 + 9 0 0 0 1 1 0 0 0 + 10 0 0 1 0 2 1 1 + 11 0 0 0 3 0 0 + 12 0 0 0 0 0 + 13 0 0 0 0 + 14 0 + + + +I have records for four generations in the 200 line and for three +generations in the 400 line.[1] As the results are practically the same +for each, I shall present the detailed records for the former group alone. +In Table 53 are to be found the number of errors made in successive series +of ten tests each by the various individuals of the 200 line which were +trained in this experiment. The most careful examination fails to reveal +any indication of the inheritance of a tendency to avoid the black box. +No. 240, in fact, chose the black box more frequently in the preference +series than did No. 210, and he required thirty more tests for the +establishment of a perfect habit than did No. 210. Apparently descent from +individuals which had thoroughly learned to avoid the black box gives the +dancer no advantage in the formation of a white-black discrimination +habit. There is absolutely no evidence of the inheritance of this +particular individually acquired form of behavior in the dancer. + +[Footnote 1: This experiment was interrupted by the death of the animals +of both lines of descent.] + + + + + +INDEX + + +Abnormal dancers. +Acquired forms of behavior. +Act, useless, repeated. +Activity, periods of. +Affirmation, choice by. +Age, peculiarities; + maximum age; + and intelligence. +Albino cat; + dog. +Alexander and Kreidl, young dancer; + behavior; + tracks of mice; + behavior in cyclostat; + behavior of white mouse and dancer; + structure of ear; + deafness. +Allen, G. M., drawing of dancer; + heredity in mice. +Alleys, width of, in labyrinths. +Amyl acetate for photometry. +Anatomy of dancer. +Animals, education of. +Appuun whistles. +Audition. _See_ Hearing. +Averages, dangers in. + + +Baginsky, B., model of ear of dancer. +Bateson, W., breeding experiments. +Behavior, of dancer; + inheritance of; + when blinded; + equilibration; + dizziness; + structural bases of; + of young; + changes in; + useless acts; + under experimental conditions; + in indiscriminable conditions; + value of sight; + in labyrinth experiments; + modifiability of; + history of; + explanations of; + individual differences in. +Blinded dancers, behavior of. +Blue-orange tests; + blue-red tests; + blue-green tests; + blue-green blindness. +Bradley papers. +Brain, structure of. +Breeding of dancers. +Brehm, A. E., "Tierleben". +Brightness vision; + preference; + check experiments; + relation to color vision. + + +Cages for dancers. +Candle meter. +Candle power. +Cardboards, for tests of vision; + positions of. +Care of dancer. +Castle, W. E., drawing of mouse; + cages. +Cat, albino; + training of. +Cerebellum of dancer. +Characters, acquired. +Check experiments. +China, dancers of. +Choice, exhibition of; + by affirmation; + by negation; + by comparison; + methods of. +Circling, a form of dance. +Circus course mice. +Cleghorn, A. G. +Climbing of dancer. +Cochlea, functions of. +Color blindness. +Color discrimination apparatus. +Colored glasses. +Colored papers. +Color patterns of dancers. +Color vision, problem; + methods of testing; + tests with colored papers, + tests with ray filters, + orange-blue tests, + yellow-red tests, + light blue-orange tests, + dark blue-red tests, + green-light blue tests, + violet-red tests, + green-blue tests, + green-red tests, + blue-green tests, + blue-red tests, + structure of the retina, + conclusions, + of different animals, +Comparative pedagogy, +Comparison, choice by, +Cones, lacking in eye of dancer, +Corti, organ of, in dancer, +Cotton mouse, +Curves, of habit formation, + irregularities of, + of labyrinth habit, + of discrimination habit, + of learning and re-learning, + of plasticity, +Cyclostat, behavior of dancer in, +Cyon, E. de, dancer pathological, + behavior, + behavior of blinded dancers, + varieties of dancer, + space perception, + individual differences, + anatomy of dancer, + hearing of dancer, + pain cries. + + +Dancers, occurrence among common mice, + varieties of, + hybrid, +Dancing, + forms of dance movement, + whirling, circling, figure-eights, manège movements, solo + dance, centre dance, + direction of, + periods of, + amount of, + causes of, + sex differences in, + individual differences in, +Darbishire, A. D., breeding experiments with dancers, +Deafness of dancer, + causes of, +Descent, lines of, +Development of young dancer, +Differences, individual, + sex, +Direction of movement, choice by, +Direction of whirling, +Discrimination, visual, box, + of brightness, + white-black and black-white, + of grays, + habits, + by odor, + by form, + method, + habit defined, +Diseases of dancer, +Dizziness, + visual, + rotational, +Docility, +Dog, albino, + training of, + fear of electric shock. + + +Ear, structure of, + structural types, + model of, + of rabbit, + functions of, + movements of, +Educability of dancer, +Education, human, + methods of, + of vision, +Efficiency of training, +Electric-box for visual tests, +Electric-labyrinth for habit experiments, +Electric-shock as punishment for mistakes, +Epidemic among dancers, +Equilibration in dancer, +Error curves, + form of, +Error records versus time records, +Errors, in labyrinths, + nature of, + types of, + value of, + number of, +Even numbers to designate males, +Excitability of dancer, +Experience, value of, + influence of, +Eyes, of dancer + opening of + retina of + + +Fear, in dancer +Females, + designated by odd numbers + dancing of + voice of + _See_ Sex +Fighting of dancers +Figure eight dance +Filters for obtaining colored light +Food of dancer +Form discrimination +Frog, + reactions of + repetition of act by +Functions of eye + + +Galton whistle +Gestation, period of, in dancer +Gray papers +Green blue tests +Green-red tests +Grouping for averages +Guaita, G von, + breeding experiments with dancers + + +Haacke, W, + description of dancer, + origin of dancer + breeding experiments +Habit, + of dancing, + discrimination, + useless + labyrinth, + duration of, + reacquisition of, + relations of, +Habit formation, + and the senses, + _versus_ habit performance, + in the dancer and in the common mouse, + curves of, + speed of +Habituation to sounds +Hacker, dancing shrews +Hair, appearance of +Hamilton, G V, experiments with dog +Hatai, S, the dancer +Head, shape of, in dancer +Hearing, + in dancer + in young + in adult + methods of testing, + in frog +Hefner unit of light +Heredity + _See_ Inheritance +Hering, E, colored papers +History, + of dancer + of acts +Hunger as motive in experiments +Hybrid dancers + + +Imitation in dancer +Index of modifiability +Individuality +Inheritance +Inhibition of an act +Initiative of dancer +Insight of dancer +Intelligence, + measures of, + comparisons +Interrupted circuit for experimental use +Irregular labyrinths + + +Janssen-Hoffman spectroscope +Japan, dancers in +Judgment in dancer + + +Kammerer, P, dancing wood mice, +Kishi, K, + dancer in Japan + origin of race, + equilibration, + blinded dancer, + structure of ear, + wax in ears, + tests of hearing +König tuning forks, + steel bars +Kreidl, A + _See_ Alexander + + +Labyrinth, + forms of, + labyrinth A, + errors in, + tests, + labyrinth B, + tests, + labyrinth C, + labyrinth D, + a standard labyrinth, + regular and irregular labyrinths +Labyrinth errors and individual tendencies +Labyrinth habits, +Labyrinth method, +Labyrinth path, formula, + method of recording, +Ladder climbing tests, +Landois, H, account of dancer, +Lathrop, A, dancers, +Learning, process, + methods of in dancer, + by being put through act, + by imitation, + by rote, + rapidity of, + permanency of, + learning and relearning, + curves of, +Left whirlers, +Life span of dancer, +Light, reflected, + transmitted, unit of measurement, + control of, +Litter, size of, in dancer, +Lummer-Brodhun photometer. + +Males, dancing of, + fighting and killing young, designation of, + voice of, _See_ Sex +Manège movements, +Mark, E L, cages, +Maze _See_ Labyrinth +Measurements, of light, + of rapidity of habit formation, + of intelligence, + of efficiency of training, +Memory, defined, + for ladder climbing, + tests of, + measurements of, + span of, + for brightness, + for color, +Method, of studying dance, + for testing hearing, + indirect, + for testing vision, + motives, + for brightness vision, + for color vision, + of shifting filters, + of testing form discrimination, + of testing Weber's law, + development of methods, + of choice, + food box, + labyrinth, + of recording errors, + of training + problem method, labyrinth method, discrimination method, + of recording labyrinth path, + qualitative versus quantitative, + of studying senses, + values of methods, + of measuring intelligence, + quantitative, + comparisons of, +Milne-Edwards, origin of dancer, +Mitsukun, K, the dancer in Japan, +Mixed whirlers, +Modifiability, of behavior, + of useless acts, + index of, +Motives, for activity, + for choice, + avoidance of discomfort, + in labyrinths, + desire to escape, to get food, to avoid pain, +Motor, tendencies, + ability, + capacity +Movements, + of ears, +_Mus musculus L_, +_Mus spiciosus L_, +_Mus sylvaticus L_. + +_Nankin nesumi_, name for dancer, +Negation, choice by, +Nendel, R, gray papers, +Nerve, eighth, +Nervous system, +Nest materials, +Noises, effects of, +Numbers, odd for females, even for males, + reference, _See_ Bibliographic List. + +Odors, discrimination by, +Old Fancier's description of dancer, +Olfactory sense _See_ Smell +Orange-blue tests, +Orientation of dancer, +Origin of dancer; + by selectional breeding; + by inheritance of an acquired character; + by mutation; + by pathological changes; + by natural selection. + + +Panse, R., + structure of ear; + explanation of deafness. +Papers, Nendel's grays; + Bradley's colored; + Hering's colored. +Parker, G. H. +Path in labyrinth, record of. +Pathological condition of dancer. +Pedagogy, comparative. +Perception, + of brightness; + of color; + of movement; + of form. +Peru, dancers in. +Petromyzon, semicircular canals of. +Photometer, Lummer-Brodhun. +Plasticity of dancer; + curves of. +Position choice by, + of cardboards. +Preference for brightness, + tests of. +Preliminary tests. +Probable error. +Problems, of structure; + of method. +Punishment versus reward. +Putting-through, training by. + + +Qualitative methods. +Quantitative methods. + + +Rabbit, ear of. +Rawitz, B., behavior of dancer; + structure of ear; + deafness of dancer; + hearing in young. +Ray filters. +Reactions, to sounds; + to disagreeable stimuli; + valueless. +Reasoning, implicit. +Reconstruction method. +Records, of markings of dancers; + of time; + of errors; + of path. +Red, stimulating value of; + vision. +Reference numbers to literature. + _See_ Literature on Dancer. +Reflected light. +Refrangibility and vision of dancer. +Regular labyrinth. +Re-learning, relation to learning; + curves of. +Reliability of averages. +Repetition of useless acts. +Rest-interval. +Restlessness, of dancer; + cause of. +Retina of dancer. +Retzius, ear of rabbit. +Reward, for performance of act; + versus punishment. +Right whirlers; + behavior in labyrinth; + occurrence of; + inheritance of tendency. +Rods of retina. +Rotational dizziness. +Rubber stamps of labyrinths. + + +Saint-Loup, R. +Schlumberger, C.; + wood carving with dancers. +Selenka, ear of rabbit. +Semicircular canals. +Sense organs. +Senses, and habit formation; + differences in. +Sensitiveness +Sex, recognition of, designation + of, peculiarities +Shellac to coat cards +Shrews, dancing +Sight, role of, _See_ Vision, + Brightness Vision, and Color Vision +Smell sense of, in labyrinth habits +Sniffing by dancer +Solutions as ray filters +_Sorex vulgaris L_ +Sound, reactions to +Space perception +Spectroscope +Spectrum, stimulating value of +Standard, candle, light, + labyrinth +Stine, W M, photometrical measurements +Strength of dancer +Structure, of brain, + of ear, of eye +Swinhoe, mice in China + +Temperament of animal +Temperature sense +Tests, visual, number of, + per day, +Threshold of discrimination +Time records +Touch, and labyrinth habits +Training conditions of, Weber's + law, methods of, + and retraining, in labyrinths, + efficiency of, two test, + ten-test, twenty-test, + continuous, relation to + methods, spread of +Transmitted light. + +Variability of dancer, +Variable light. +Varieties of dancer +Violet red tests +Vision, brightness + vision, color vision, + training of, importance + of, conclusions concerning +Visual dizziness +Voice of dancer + +Watson, J B, habit formation +Waugh, K, color vision apparatus, + retina of mouse +Wax, plugs of, in ear of mouse +Weber's law, tests of, apparatus +Weldon, W F R, breeding experiments, +Whirling of dancer + +Yellow Red tests +Young dancers, killing of, by male, + description of, development + of, hearing of, intelligence + of, size of + +Zoth, O, origin of dancer, size + of young mice, the senses of + dancer, behavior, dancing, + equilibration, climbing + dancers, individual differences, + tests of hearing, + vision + + + + + +End of the Project Gutenberg EBook of The Dancing Mouse, by Robert M. Yerkes + +*** END OF THE PROJECT GUTENBERG EBOOK THE DANCING MOUSE *** + +This file should be named 8729-8.txt or 8729-8.zip + +Produced by Juliet Sutherland, Michael Oltz, +Charles Franks and the Online Distributed Proofreading Team. + +Project Gutenberg eBooks are often created from several printed +editions, all of which are confirmed as Public Domain in the US +unless a copyright notice is included. Thus, we usually do not +keep eBooks in compliance with any particular paper edition. + +We are now trying to release all our eBooks one year in advance +of the official release dates, leaving time for better editing. +Please be encouraged to tell us about any error or corrections, +even years after the official publication date. + +Please note neither this listing nor its contents are final til +midnight of the last day of the month of any such announcement. +The official release date of all Project Gutenberg eBooks is at +Midnight, Central Time, of the last day of the stated month. 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