The skin is supplied with nerves—naked fibrils—in the richest abundance. They are most easily demonstrated in the layer which covers the cornea, thanks to its transparency; in this, as shown in [Fig. 41], having branched on the front of the fibrous tissue of which the cornea is composed, the nerves pass towards the surface, forming connections with every one of its cells, or, at any rate, with every cell of the more superficial of the three or four layers of which the epithelium is made up. Ramified nerve-twigs of this type do not, under ordinary conditions, convey any sensations to consciousness. So long as the skin-cells with which they are connected are healthy, the nerve-twigs establish for them connections with the central nervous system by which their nutrition is regulated; but they carry no impulses to which we can direct attention. The movement of blinking is accompanied by no sensation until the edges of the eyelids come in contact. A pencil pressed against the lid evokes touch-sensations from the skin, but none from the cornea which underlies it. When a tiny beetle injures the surface of the cornea by scratching the epithelial cells with its horny wings and legs, the ruptured nerve-filaments convey to consciousness impulses, or, as we prefer to express it, an influence which is felt as pain. But even the pain caused by injury to the cornea is trifling as compared with that which originates in the under-sides of the lids, where not only is the epithelium supplied with branching nerve-twigs, but specialized organs of touch are present to localize the seat of injury. Everywhere the epithelium covering the surface of the body is so abundantly supplied that a successful staining of nerve-filaments induces one to think that every epithelial cell has its nervous affiliation. These are the nerves of common sensation, if we retain the term; but sensation so common, so obscure, so little differentiated that we know no more about it than we know about the air which envelops our hands and faces on a warm, windless day. Yet the air, when it moves, gives rise to a dim, broad, generalized sensation, which may be focussed into definiteness by a sensitive nerve.
Fig. 41.—Vertical Section of the Epithelium which covers the Surface of the Cornea,
and of a Small Portion of the Corneal Substance, highly Magnified.
The black lines are naked nerve-fibres (stained with chloride of gold), which are distributed amongst the cells of the more superficial strata of the epithelium in very great abundance. The corneal substance is composed of sheets of transparent fibres with intervening cells. As the fibres of the several sheets cross one another at various angles, they are cut, some transversely, others in the direction of their length.
An observer who has devoted himself for many years to the investigation of skin-sensations, and especially of the “referred pains” which are due to diseases of the viscera, recently caused the large cutaneous nerve which supplies the thumb side of the forearm and hand to be cut in his own arm, in order that he might study carefully the revival of sensations. He found that he never lost his ability to recognize displacements of the tissues beneath the skin. Pacinian bodies and other end-organs of deep-lying nerves recorded pressure and tension caused by pushing or rubbing with a blunt instrument. Seven weeks after the injury he began to recognize stimuli that do harm—hot things, cold things, pricking with a pin—although his power of localizing the spot injured was extremely vague. In seven weeks, that is to say, the protopathic nerves, which do not follow the same definite lines as the nerves of the special senses, but form open networks with many alternative paths, had re-established their skin connections. Only gradually and very slowly did critical sensations return—the ability to distinguish degrees of warmth, to recognize as separate two points of a pair of compasses, to feel a touch with cotton-wool.
According to a theory set forth in this book ([p. 312]), pain is not a set of sensations, but a condition of the central nervous system which renders it unduly excitable, or excitable in a particular manner, to impulses which have the same local origin as the nerve-current which sets up the condition of pain. When a nerve of the skin has been cut, the epithelial ramifications are renewed before any specialized tactile or other sense-organs have regained their nervous connections. When the area which has regained its surface ramifications, but has not regained its sense-organs, is injured, no localization of pain results. Indeed, the obscure sensations which are then experienced if the skin be injured can hardly be described as painful. The ramified nerves pour their agitation into the grey matter of the spinal cord; but it is not the agitation per se which causes pain. It is the passage of impulses through the agitated area that gives to them, when they reach consciousness, not only a topographical meaning, but also a distressful feeling. Until the specialized organs of the skin have been restored to working order, there are no impulses to pass through the agitated grey matter, and therefore no feelings of pain. According to this view there are two systems of afferent nerves, the protopathic and the specialized or critical. The former is very widely and very abundantly distributed to the surface of the body, the lungs, the alimentary canal, and other viscera. It has no end-organs, no defined tracts in the central nervous system, no definite connections with the cortex of the great brain. The currents which it conducts, if they originate in the visceral part of this system, have no direct effect in consciousness; but if they originate on the surface of the body, or in the alimentary canal at the lower end of the œsophagus, or in certain other situations, they co-operate with stimuli of heat, cold, or traction. The critical system works in a more definite way. Its impulses originate in sense-organs. Starting with a certain potential, they are transmitted by the discharge of a succession of linked neurones. When they reach the cortex their potential is sufficiently high to evoke consciousness. Their distribution in the cortex is as definite as their origin.
Specialized sense-organs are necessary for the origin of all sensations. Within the epithelium are certain cells which look as if they were specialized for sensory purposes. The deeper sheet, or derma, of the skin is abundantly provided with structures in which nerves end in the most elaborate and complicated ways ([Fig. 42]). They are found especially in the papillæ of connective tissue, which, set in rows, form the ridges that one can see at the finger-tips and in various other situations. All of these organs are made up of groups of epithelial cells which, displaced from the epidermis, have sunk into the derma, with the nerves connected with them. In their further development the nervous part of the apparatus is complicated by branching, the branches being thickened and usually flattened into ribbons, which lie on the external surfaces of the cells or between them. A more or less marked capsule is provided for the organ by condensation of connective tissue.
Anyone can convince himself that the skin is not uniformly sensitive. He may test it first for the minimal stimulus which excites a sensation of touch. With a hair of the head—it must not be a very fine one—cut across with scissors, and held between finger and thumb at the right distance from the cut end, the skin of the palm of the hand is prodded. Every here and there a spot is found which is insensitive to so slight a pressure. These spots are neither large nor very close together. If the hairless skin of the arm between the elbow and the armpit be investigated in the same way, much larger blank areas are met with—oval patches more than ¼ inch in diameter. When a hairy surface is tested, it is found that contact with a hair can always be felt; and when the hairs are shaved, the touch-spots are found to extend around or from the points at which hairs pierce the epidermis. Touchless areas lie between them. Hair-follicles receive tufts of nerve-filaments, and it appears that they are the chief organs of touch. “Touch-corpuscles,” which are found in great numbers in the papillæ of the skin of the fingers and elsewhere, may probably be regarded as, genetically, hair-follicles which have not developed hairs.