The Dancing Mouse Part 5

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 ca.n.a.ls 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 ca.n.a.ls. It is highly probable, therefore, that the ca.n.a.ls 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 cooperation 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 ca.n.a.ls, the ampullae, the utriculus, and the cristae acusticae of the ca.n.a.ls 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 ca.n.a.ls 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 ca.n.a.l 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 s.p.a.ce perception which Cyon (9) had constructed, largely on the basis of the demonstration by Rawitz that the dancers have only one normal ca.n.a.l, is totally destroyed by Panse, Baginsky, Alexander and Kreidl, and Kishi, for all of these observers found in the dancer three ca.n.a.ls 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 ca.n.a.l which Rawitz found to be most strikingly defective. This fact he connected with his observation that the fish Petromyzon, which possesses only two ca.n.a.ls, moves in only two spatial dimensions. The dancer with only one functional ca.n.a.l 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 s.p.a.ce depends upon the number and arrangement of the semicircular ca.n.a.ls, and that the dancer behaves as it does because it possesses ca.n.a.ls 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 s.p.a.ces 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 ca.n.a.ls into the cochlea as a.s.sumed 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.

[Ill.u.s.tration: FIGURE 12.--The inner ear of the dancer. Reproduced from Kishi's figure in the _Zeitschrift fur wissenschaftliche Zoologie_, Bd.

71. _c.c._ crus simplex; o.b. anterior vertical ca.n.a.l; _h.b._ posterior vertical ca.n.a.l; _a.b._ horizontal ca.n.a.l.]

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 ca.n.a.ls as Rawitz's figures and descriptions represent. Rawitz states that the anterior ca.n.a.l 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 ca.n.a.ls are normal in shape and that the usual connection between the anterior and the posterior ca.n.a.ls, 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 ca.n.a.ls 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 ca.n.a.ls 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 ca.n.a.ls 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 ca.n.a.ls, which frequently are clearly visible in the gray mouse, are lacking (21 p. 478).

[Ill.u.s.tration: 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 s.p.a.ce 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 ca.n.a.ls 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 ca.n.a.l 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 a.s.suming 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).