In the lower animals the analogue of the labyrinth is shrunk to a little vesicle filled with a liquid and containing tiny crystals, auditive stones, or otoliths, of greater specific gravity, suspended on minute hairs. These crystals appear physically well adapted for indicating both the direction of gravity and the direction of incipient movements. That they discharge the former function, Delage was the first to convince himself by experiments with lower animals which on the removal of the otoliths utterly lost their bearings and could no longer regain their normal position. Loeb also found that fishes without labyrinths swim now on their bellies and now on their backs. But the most remarkable, most beautiful, and most convincing experiment is that which Dr. Kreidl instituted with crustaceans. According to Hensen, certain Crustacea on sloughing spontaneously introduce fine grains of sand as auditive stones into their otolith vesicle. At the ingenious suggestion of S. Exner, Dr. Kreidl constrained some of these animals to put up with iron filings (ferrum limatum). If the pole of an electro-magnet be brought near the animal, it will at once turn its back away from the pole accompanying the movement with appropriate reflex motions of the eye the moment the current is closed, exactly as if gravity had been brought to bear upon the animal in the same direction as the magnetic force.[104] This, in fact, is what should be expected from the function ascribed to the otoliths. If the eyes be covered with asphalt varnish, and the auditive sacs removed, the crustaceans lose their sense of direction utterly, tumble head over heels, lie on their side or their back indifferently. This does not happen when the eyes only are covered. For vertebrates, Breuer has demonstrated by searching investigations that the otoliths, or better, statoliths, slide in three planes parallel to the planes of the semi-circular canals, and are consequently perfectly adapted for indicating changes both in the amount and the direction of the mass-acceleration.[105]
I have already remarked that not every function of orientation can be ascribed exclusively to the labyrinth. The deaf and dumb who have to be immersed in water, and the crustaceans who must have their eyes closed if they are to be perfectly disorientated, are proof of this fact. I saw a blind cat at Hering's laboratory which to one who was not a very attentive observer behaved exactly like a seeing cat. It played nimbly with objects rolling on the floor, stuck its head inquisitively into open drawers, sprang dexterously upon chairs, ran with perfect accuracy through open doors, and never bumped against closed ones. The visual sense had here been rapidly replaced by the tactual and auditive senses. And it appears from Ewald's investigations that even after the labyrinths have been removed, animals gradually learn to move about again quite in the normal fashion, presumably because the eliminated function of the labyrinth is now performed by some part of the brain. A certain peculiar weakness of the muscles alone is perceptible which Ewald ascribes to the absence of the stimulus which is otherwise constantly emitted by the labyrinth (the labyrinth-tonus). But if the part of the brain which discharges the deputed function be removed, the animals are again completely disorientated and absolutely helpless.
It may be said that the views enunciated by Breuer, Crum Brown and myself in 1873 and 1874, and which are substantially a fuller and richer development of Goltz's idea, have upon the whole been substantiated. At least they have exercised a helpful and stimulative influence. New problems have of course arisen in the course of the investigation which still await solution, and much work remains to be done. At the same time we see how fruitful the renewed co-operation of the various special departments of science may become after a period of isolation and invigorating labor apart.
I may be permitted, therefore, to consider the relation between hearing and orientation from another and more general point of view. What we call the auditive organ is in the lower animals simply a sac containing auditive stones. As we ascend the scale, 1, 2, 3 semi-circular canals gradually develop from them, whilst the structure of the otolith organ itself becomes more complicated. Finally, in the higher vertebrates, and particularly in the mammals, a part of the latter organ (the lagena) becomes the cochlea, which Helmholtz explained as the organ for sensations of tone. In the belief that the entire labyrinth was an auditive organ, Helmholtz, contrary to the results of his own masterly analysis, originally sought to interpret another part of the labyrinth as the organ of noises. I showed a long time ago (1873) that every tonal stimulus by shortening the duration of the excitation to a few vibrations, gradually loses its character of pitch and takes on that of a sharp, dry report or noise.[106] All the intervening stages between tones and noises can be exhibited. Such being the case, it will hardly be assumed that one organ is suddenly and at some given point replaced in function by another. On the basis of different experiments and reasonings S. Exner also regards the assumption of a special organ for the sensing of noises as unnecessary.
If we will but reflect how small a portion of the labyrinth of higher animals is apparently in the service of the sense of hearing, and how large, on the other hand, the portion is which very likely serves the purposes of orientation, how much the first anatomical beginnings of the auditive sac of lower animals resemble that part of the fully developed labyrinth which does not hear, the view is irresistibly suggested which Breuer and I (1874, 1875) expressed, that the auditive organ took its development from an organ for sensing movements by adaptation to weak periodic motional stimuli, and that many apparatuses in the lower animals which are held to be organs of hearing are not auditive organs at all.[107]
This view appears to be perceptibly gaining ground. Dr. Kreidl by skilfully-planned experiments has arrived at the conclusion that even fishes do not hear, whereas E. H. Weber, in his day, regarded the ossicles which unite the air-bladder of fishes with the labyrinth as organs expressly designed for conducting sound from the former to the latter.[108] Störensen has investigated the excitation of sounds by the air-bladder of fishes, as also the conduction of shocks through Weber's ossicles. He regards the air-bladder as particularly adapted for receiving the noises made by other fishes and conducting them to the labyrinth. He has heard the loud grunting tones of the fishes in South American rivers, and is of the opinion that they allure and find each other in this manner. According to these views certain fishes are neither deaf nor dumb.[109] The question here involved might be solved perhaps by sharply distinguishing between the sensation of hearing proper, and the perception of shocks. The first-mentioned sensation may, even in the case of many vertebrates, be extremely restricted, or perhaps even absolutely wanting. But besides the auditive function, Weber's ossicles may perfectly well discharge some other function. Although, as Moreau has shown, the air-bladder itself is not an organ of equilibrium in the simple physical sense of Borelli, yet doubtless some function of this character is still reserved for it. The union with the labyrinth favors this conception, and so a host of new problems rises here before us.
I should like to close with a reminiscence from the year 1863. Helmholtz's Sensations of Tone had just been published and the function of the cochlea now appeared clear to the whole world. In a private conversation which I had with a physician, the latter declared it to be an almost hopeless undertaking to seek to fathom the function of the other parts of the labyrinth, whereas I in youthful boldness maintained that the question could hardly fail to be solved, and that very soon, although of course I had then no glimmering of how it was to be done. Ten years later the question was substantially solved.
To-day, after having tried my powers frequently and in vain on many questions, I no longer believe that we can make short work of the problems of science. Nevertheless, I should not consider an "ignorabimus" as an expression of modesty, but rather as the opposite. That expression is a suitable one only with regard to problems which are wrongly formulated and which are therefore not problems at all. Every real problem can and will be solved in due course of time without supernatural divination, entirely by accurate observation and close, searching thought.