The Expression of the Emotions in Man and Animals (Evolutionary Theory). Чарльз Дарвин

nerve, which transmits its influence to certain nerve-cells, and these in their turn excite certain muscles or glands into action; and all this may take place without any sensation or consciousness on our part, though often thus accompanied. As many reflex actions are highly expressive, the subject must here be noticed at some little length. We shall also see that some of them graduate into, and can hardly be distinguished from actions which have arisen through habit?31 Coughing and sneezing are familiar instances of reflex actions. With infants the first act of respiration is often a sneeze, although this requires the co-ordinated movement of numerous muscles. Respiration is partly voluntary, but mainly reflex, and is performed in the most natural and best manner without the interference of the will. A vast number of complex movements are reflex. As good an instance as can be given is the often-quoted one of a decapitated frog, which cannot of course feel, and cannot consciously perform, any movement. Yet if a drop of acid be placed on the lower surface of the thigh of a frog in this state, it will rub off the drop with the upper surface of the foot of the same leg. If this foot be cut off, it cannot thus act. “After some fruitless efforts, therefore, it gives up trying in that way, seems restless, as though, says Pflüger, it was seeking some other way, and at last it makes use of the foot of the other leg and succeeds in rubbing off the acid. Notably we have here not merely contractions of muscles, but combined and harmonized contractions in due sequence for a special purpose. These are actions that have all the appearance of being guided by intelligence and instigated by will in an animal, the recognized organ of whose intelligence and will has been removed.”32

      We see the difference between reflex and voluntary movements in very young children not being able to perform, as I am informed by Sir Henry Holland, certain acts somewhat analogous to those of sneezing and coughing, namely, in their not being able to blow their noses (i.e. to compress the nose and blow violently through the passage), and in their not being able to clear their throats of phlegm. They have to learn to perform these acts, yet they are performed by us, when a little older, almost as easily as reflex actions. Sneezing and coughing, however, can be controlled by the will only partially or not at all; whilst the clearing the throat and blowing the nose are completely under our command.

      When we are conscious of the presence of an irritating particle in our nostrils or windpipe—that is, when the same sensory nerve-cells are excited, as in the case of sneezing and coughing—we can voluntarily expel the particle by forcibly driving air through these passages; but we cannot do this with nearly the same force, rapidity, and precision, as by a reflex action. In this latter case the sensory nerve-cells apparently excite the motor nerve-cells without any waste of power by first communicating with the cerebral hemispheres—the seat of our consciousness and volition. In all cases there seems to exist a profound antagonism between the same movements, as directed by the will and by a reflex stimulant, in the force with which they are performed and in the facility with which they are excited. As Claude Bernard asserts, “L’influence du cerveau tend donc à entraver les mouvements réflexes, à limiter leur force et leur étendue.”33

      The conscious wish to perform a reflex action sometimes stops or interrupts its performance, though the proper sensory nerves may be stimulated. For instance, many years ago I laid a small wager with a dozen young men that they would not sneeze if they took snuff, although they all declared that they invariably did so; accordingly they all took a pinch, but from wishing much to succeed, not one sneezed, though their eyes watered, and all, without exception, had to pay me the wager. Sir H. Holland remarks34 that attention paid to the act of swallowing interferes with the proper movements; from which it probably follows, at least in part, that some persons find it so difficult to swallow a pill.

      Another familiar instance of a reflex action is the involuntary closing of the eyelids when the surface of the eye is touched. A similar winking movement is caused when a blow is directed towards the face; but this is an habitual and not a strictly reflex action, as the stimulus is conveyed through the mind and not by the excitement of a peripheral nerve. The whole body and head are generally at the same time drawn suddenly backwards. These latter movements, however, can be prevented, if the danger does not appear to the imagination imminent; but our reason telling us that there is no danger does not suffice. I may mention a trifling fact, illustrating this point, and which at the time amused me. I put my face close to the thick glass-plate in front of a puff-adder in the Zoological Gardens, with the firm determination of not starting back if the snake struck at me; but, as soon as the blow was struck, my resolution went for nothing, and I jumped a yard or two backwards with astonishing rapidity. My will and reason were powerless against the imagination of a danger which had never been experienced.

      The violence of a start seems to depend partly on the vividness of the imagination, and partly on the condition, either habitual or temporary, of the nervous system. He who will attend to the starting of his horse, when tired and fresh, will perceive how perfect is the gradation from a mere glance at some unexpected object, with a momentary doubt whether it is dangerous, to a jump so rapid and violent, that the animal probably could not voluntarily whirl round in so rapid a manner. The nervous system of a fresh and highly-fed horse sends its order to the motory system so quickly, that no time is allowed for him to consider whether or not the danger is real. After one violent start, when he is excited and the blood flows freely through his brain, he is very apt to start again; and so it is, as I have noticed, with young infants.

      A start from a sudden noise, when the stimulus is conveyed through the auditory nerves, is always accompanied in grown-up persons by the winking of the eyelids.35 I observed, however, that though my infants started at sudden sounds, when under a fortnight old, they certainly did not always wink their eyes, and I believe never did so. The start of an older infant apparently represents a vague catching hold of something to prevent falling. I shook a pasteboard box close before the eyes of one of my infants, when 114 days old, and it did not in the least wink; but when I put a few comfits into the box, holding it in the same position as before, and rattled them, the child blinked its eyes violently every time, and started a little. It was obviously impossible that a carefully-guarded infant could have learnt by experience that a rattling sound near its eyes indicated danger to them. But such experience will have been slowly gained at a later age during a long series of generations; and from what we know of inheritance, there is nothing improbable in the transmission of a habit to the offspring at an earlier age than that at which it was first acquired by the parents.

      From the foregoing remarks it seems probable that some actions, which were at first performed consciously, have become through habit and association converted into reflex actions, and are now so firmly fixed and inherited, that they are performed, even when not of the least use,36 as often as the same causes arise, which originally excited them in us through the volition. In such cases the sensory nerve-cells excite the motor cells, without first communicating with those cells on which our consciousness and volition depend. It is probable that sneezing and coughing were originally acquired by the habit of expelling, as violently as possible, any irritating particle from the sensitive air-passages. As far as time is concerned, there has been more than enough for these habits to have become innate or converted into reflex actions; for they are common to most or all of the higher quadrupeds, and must therefore have been first acquired at a very remote period. Why the act of clearing the throat is not a reflex action, and has to be learnt by our children, I cannot pretend to say; but we can see why blowing the nose on a handkerchief has to be learnt.

      It is scarcely credible that the movements of a headless frog, when it wipes off a drop of acid or other object from its thigh, and which movements are so well coordinated for a special purpose, were not at first performed voluntarily, being afterwards rendered easy through long-continued habit so as at last to be performed unconsciously, or independently of the cerebral hemispheres.

      So again it appears probable that starting was originally acquired by the habit of jumping away as quickly as possible from danger, whenever any of our senses gave us warning. Starting, as we have seen, is accompanied by the blinking of the eyelids so as to protect the eyes, the most tender and sensitive organs of the body; and it is, I believe, always accompanied by a sudden and forcible inspiration, which is the natural preparation for any violent effort. But when a man or horse starts, his heart beats wildly against his ribs, and here it may be truly said we have an organ which has never been under the control of the will, partaking in the general reflex movements of