1911 Encyclopædia Britannica/Equilibrium
EQUILIBRIUM (from the Lat. aequus, equal, and libra, a balance), a condition of equal balance between opposite or counteracting forces. By the “sense of equilibrium” is meant the sense, or sensations, by which we have a feeling of security in standing, walking, and indeed in all the movements by which the body is carried through space. Such a feeling of security is necessary both for maintaining any posture, such as standing, or for performing any movement. If this feeling is absent or uncertain, or if there are contradictory sensations, then definite muscular movements are inefficiently or irregularly performed, and the body may stagger or fall. When we stand erect on a firm surface, like a floor, there is a feeling of resistance, due to nervous impulses reaching the brain from the soles of the feet and from the muscles of the limbs and trunk. In walking or running, these feelings of resistance seem to precede and guide the muscular movements necessary for the next step. If these are absent or perverted or deficient, as is the case in the disease known as locomotor ataxia, then, although there is no loss of the power of voluntary movement, the patient staggers in walking, especially if he is not allowed to look at his feet, or if he is blind-folded. He misses the guiding sensations that come from the limbs; and with a feeling that he is walking on a soft substance, offering little or no resistance, he staggers, and his muscular movements become irregular. Such a condition maybe artificially brought about by washing the soles of the feet with chloroform or ether. And it has been observed to exist partially after extensive destruction of the skin of the soles of the feet by burns or scalds. This shows that tactile impulses from the skin take a share in generating the guiding sensation. In the disease above mentioned, however, tactile impressions may be nearly normal, but the guiding sensation is weak and inefficient, owing to the absence of impulses from the muscles. The disease is known to depend on morbid changes in the posterior columns of the spinal cord, by which impulses are not freely transmitted upwards to the brain. These facts point to the existence of impulses coming from the muscles and tendons. It is now known that there exist peculiar spindles, in muscle, and rosettes or coils or loops of nerve fibres in close proximity to tendons. These are the end organs of the sense. The transmission of impulses gives rise to the muscular sense, and the guiding sensation which precedes co-ordinated muscular movements depends on these impulses. Thus from the limbs streams of nervous impulses pass to the sensorium from the skin and from muscles and tendons; these may or may not arouse consciousness, but they guide or evoke muscular movements of a co-ordinated character, more especially of the limbs.
In animals whose limbs are not adapted for delicate touch nor for the performance of complicated movements, such as some mammals and birds and fishes, the guiding sensations depend largely on the sense of vision. This sense in man, instead of assisting, sometimes disturbs the guiding sensation. It is true that in locomotor ataxia visual sensations may take the place of the tactile and muscular sensations that are inefficient, and the man can walk without staggering if he is allowed to look at the floor, and especially if he is guided by transverse straight lines. On the other hand, the acrobat on the wire-rope dare not trust his visual sensations in the maintenance of his equilibrium. He keeps his eyes fixed on one point instead of allowing them to wander to objects below him, and his muscular movements are regulated by the impulses that come from the skin and muscles of his limbs. The feeling of insecurity probably arises from a conception of height, and also from the knowledge that by no muscular movements can a man avoid a catastrophe if he should fall. A bird, on the other hand, depends largely on visual impressions, and it knows by experience that if launched into the air from a height it can fly. Here, probably, is an explanation of the large size of the eyes of birds. Cover the head, as in hooding a falcon, and the bird seems to be deprived of the power of voluntary movement. Little effect will be produced if we attempt to restrain the movements of a cat by covering its eyes. A fish also is deprived of the power of motion if its eyes are covered. But both in the bird and in the fish tactile and muscular impressions, especially the latter, come into play in the mechanism of equilibrium. In flight the large-winged birds, especially in soaring, can feel the most delicate wind-pressures, both as regards direction and force, and they adapt the position of their body so as to catch the pressure at the most efficient angle. The same is true of the fish, especially of the flat-fishes. In mammals the sense of equilibrium depends, then, on streams of tactile, muscular and visual impressions pouring in on the sensorium, and calling forth appropriate muscular movements. It has also been suggested that impulses coming from the abdominal viscera may take part in the mechanism. The presence in the mesentery of felines (cats, &c.) of large numbers of Pacinian corpuscles, which are believed to be modified tactile bodies, favours this supposition. Such animals are remarkable for the delicacy of such muscular movements, as balancing and leaping.
There is another channel by which nervous impulses reach the sensorium and play their part in the sense of equilibrium, namely, from the semicircular canals, a portion of the internal ear. It is pointed out in the article Hearing that the appreciation of sound is in reality an appreciation of variations of pressure. The labyrinth consists of the vestibule, the cochlea and the semicircular canals. The cochlea receives the sound-waves (variations of pressure) that constitute musical tones. This it accomplishes by the structures in the ductus cochlearis. In the vestibule we find two sacs, the saccule next to and communicating with the ductus cochlearis, and the utricle communicating with the semicircular canals. The base of the stapes communicates pressures to the utricle. The membranous portion of the semicircular canals consists of a tube, dilated at one end into a swelling or pouch, termed the ampulla, and each end communicates freely with the utricle. On the posterior wall of both the saccule and of the utricle there is a ridge, termed in each case the macula acustica, bearing a highly specialized epithelium. A similar structure exists in each ampulla. This would suggest that all three structures have to do with hearing; but, on the other hand, there is experimental evidence that the utricle and the canals may transmit impressions that have to do with equilibrium. Pressure of the base of the stapes is exerted on the utricle. This will compress the fluid in that cavity, and tend to drive the fluid into the semicircular canals that communicate with that cavity by five openings. Each canal is surrounded by a thin layer of perilymph, so that it may yield a little to this pressure, and exert a pull or pressure on the nerve-endings in each ampulla. Thus impulses may be generated in the nerves of the ampullae.
The three semicircular canals lie in the three directions in space, and it has been suggested that they have to do with our appreciation of the direction of sound. But our appreciation of sound is very inaccurate: we look with the eyes for the source of a sound, and instinctively direct the ears or the head, or both, in the direction from which the sound appears to proceed. But the relationship of the canals on the two sides must have a physiological significance. Thus (1) the six canals are parallel, two and two; or (2) the two horizontal canals are in the same plane, while the superior canal on one side is nearly parallel with the posterior canal of the other. These facts point to the two sets of canals and ampullae acting as one organ, in a manner analogous to the action of two retinae for single vision.
We have next to consider how the canals may possibly act in connexion with the sense of equilibrium. In 1820 J. Purkinje studied the vertigo that follows rapid rotation of the body in the erect position on a vertical axis. On stopping the rotation there is a sense of rotation in the opposite direction, and this may occur even when the eyes are closed. Purkinje noticed that the position of the imaginary axis of rotation depends on the axis around which the head revolves. In 1828 M. J. P. Flourens discovered that injury to the canals causes disturbance to the equilibrium and loss of co-ordination, and that sections of the canals produce a rotatory movement of a kind corresponding to the canal that had been divided. Thus division of a membranous canal causes rotatory movements round an axis at right angles to the plane of the divided canal. The body of the animal always moves in the direction of the cut canal. Many other observers have corroborated these experiments. F. Goltz was the first who formulated the conditions necessary for equilibration. He put the matter thus:—(1) A central co-ordinating organ—in the brain; (2) centripetal fibres, with their peripheral terminations—in the ampullae; and (3) centrifugal fibres, with their terminal organs—in the muscular mechanisms. A lesion of any one of these portions of the mechanism causes loss or impairment of balancing. Cyon also investigated the subject, and concluded:—(1) To maintain equilibrium, we must have an accurate notion of the position of the head in space; (2) the function of the semicircular canals is to communicate impressions that give a representation of this position—each canal having a relation to one of the dimensions of space; (3) disturbance of equilibrium follows section; (4) involuntary movements following section are due to abnormal excitations; (5) abnormal movements occurring a few days after the operation are caused by irritation of the cerebellum.
On theoretical considerations of a physical character, E. Mach, Crum-Brown and Breuer have advanced theories based on the idea of the canals being organs for sensations of acceleration of movement, or for the sense of rotation. Mach first pointed out that Purkinje’s phenomena, already alluded to, were in all probability related to the semicircular canals. “He showed that when the body is moved in space, in a straight line, we are not conscious of the velocity of motion, but of variations in this velocity. Similarly, if a body is rotated round a vertical axis, we perceive only angular acceleration and not angular velocity. The sensations produced by angular acceleration last longer than the acceleration itself, and the position of the head during the movements enables us to determine direction.” Both Mach and Goltz state that varying pressures of the fluid in the canals produced by angular rotation produce sensations of movement (always in a direction opposite to the rotation of the body), and that these, in turn, cause the vertigo of Purkinje and the phenomena of Flourens. Mach, Crum-Brown and Breuer advance hydrodynamical theories in which they assume that the fluids move in the canals. Goltz, on the other hand, supports a hydrostatical theory in which he assumes that the phenomena can be accounted for by varying pressures. Crum-Brown differs from Mach and Breuer as follows:—(1) In attributing movement or variation of pressure not merely to the endolymph, but also to the walls of the membranous canals and to the surrounding perilymph; and (2) in regarding the two labyrinths as one organ, all the six canals being required to form a true conception of the rotating motion of the head. He sums up the matter thus: “We have two ways in which a relative motion can occur between the endolymph and the walls of the cavity containing it—(1) When the head begins to move, here the walls leave the fluid behind; (2) when the head stops, here the fluid flows on. In both cases the sensation of rotation is felt. In the first this sensation corresponds to a real rotation, in the second it does not, but in both it corresponds to a real acceleration (positive or negative) of rotation, using the word acceleration in its technical kinematical sense.”
Cyon states that the semicircular canals only indirectly assist in giving a notion of spatial relations. “He holds that knowledge of the position of bodies in space depends on nervous impulses coming from the contracting ocular muscles; that the oculomotor centres are in intimate physiological relationship with the centres receiving impulses from the nerves of the semicircular canals; and that the oculomotor centres, thus excited, produce the movements of the eyeballs, which then determine our notions of spatial relations.” These views are supported by experiments of Lee on dog-fish. When the fish is rotated round different axes there are compensating movements of the eyes and fins. “It was observed that if the fish were rotated in the plane of one of the canals, exactly the same movements of the eyes and fins occurred as were produced by experimental operation and stimulation of the ampulla of that canal.” Sewall, in 1883, carried out experiments on young sharks and skates with negative results. Lee returned to the subject in 1894, and, after numerous experiments on dog-fish, in which the canals or the auditory nerves were divided, obtained evidence that the ampullae contain sense-organs connected with the sense of equilibrium.
It has been found by physicians and aurists that disease or injury of the canals, occurring rapidly, produces giddiness, staggering, nystagmus (a peculiar twitching movement of the muscles of the eyeballs), vomiting, noises in the ear and more or less deafness. It is said, however, that if pathological changes come on slowly, so that the canals and vestibule are converted into a solid mass, none of these symptoms may occur. On the whole, the evidence is in favour of the view that from the semicircular canals nervous impulses are transmitted, which, co-ordinated with impulses coming from the visual organs, from the muscles and from the skin, form the bases of these guiding sensations on which the sense of equilibrium depends. These impulses may not reach the level of consciousness, but they call into action co-ordinated mechanisms by which complicated muscular movements are effected.
Full bibliographical references are given in the article on “The Ear” by J. G. McKendrick, in Schäfer’s Textbook of Physiology, vol. ii. p. 1194. (J. G. M.)