Fig. 1.--The connection from the back of the hand,
which is receiving a stimulus, and the arm muscle which makes the
response. The nerve center is indicated by the dotted lines.
The nervous system resembles a city telephone system. What passes along the nerve is akin to the electricity that {28} passes along the telephone wire; it is called the "nerve current", and is electrical and chemical in nature.
Fig. 2.--(From Martin's "Human Body.") General view of
the nervous system, showing brain, cord, and nerves.
All nerve connections, like the great majority of telephone connections, are effected through the centers, called "centrals" in {29} the case of the telephone. Telephone A is connected directly with the central, telephone B likewise, and A and B are indirectly connected, through the central switchboard. That is the way it is in the nervous system, with "nerve center" substituted for "central", and "sense organ" and "muscle or gland" for "telephones A and B."
Fig. 3.--Location of the cord, cerebrum and cerebellum.
The brain stem continues the cord upward into the skull cavity.
(Figure text: cerebrum, cerebellum, cord, tongue)
The advantage of the centralized system is that it is a system, affording connections between any part and any other, and unifying the whole complex organism.
The nerve centers are located in the brain and spinal cord. The brain lies in the skull and the cord extends from the brain down through a tube in the middle of the {30} backbone. Of the brain many parts can be named, but for the present it is enough to divide it into the "brain stem", a continuation of the spinal cord up along the base of the skull cavity, and the two great outgrowths of the brain stem, called "cerebrum" and "cerebellum". The spinal cord and brain stem contain the lower or reflex centers, while the cerebellum, and especially the cerebrum, contain the "higher centers". The lower centers are directly connected by nerves with the sense organs, glands and muscles, while the higher centers have direct connections with the lower and only through them with the sense organs, glands and muscles. In other words, the sensory nerves run into the cord or brain stem, and the motor nerves run out of these same, while interconnecting nerve strands extend between the lower centers in the cord and brain stem and the higher centers in the cerebrum and cerebellum.
The spinal cord contains the reflex centers for the limbs and part of the trunk, and is connected by sensory and motor nerves with the limbs and trunk. The brain stem contains the reflex centers for the head and also for part of the interior of the trunk, including the heart and lungs, and is connected with them by sensory and motor nerves. The nerve center that takes part in the flexion reflex of the foot is situated in the lower part of the cord, that for the similar reflex of the hand lies in the upper part of the cord, that for breathing lies in the lower or rear part of the brain stem, and that for winking lies further forward in the brain stem.
Big movements, such as the combined action of all four legs of an animal in walking, require cord and brain stem to work together, and throw into relief what is really true even of simpler reflexes, namely that a reflex is a coordinated movement, in the sense that different muscles cooperate in its execution.
Internal Construction of the Nerves and Nerve Centers
We shall understand nerve action better if we know something of the way in which the nervous system is built. A nerve is not to be thought of as a unit, nor are the brain and cord to be thought of as mere masses of some peculiar substance.
Fig. 4.--A motor nerve cell from the spinal cord,
highly magnified. (Figure text: dendrites, cell body, axon,
termination of axon in muscle)
A nerve is a bundle of many slender insulated threads, just as a telephone cable, running along the street, {32} is a bundle of many separate wires which are the real units of telephonic communication. A nerve center, like the switchboard in a telephone central, consists of many parts and connections.
The whole nervous system is essentially composed of neurones. A neurone is a nerve cell with its branches. Most nerve cells have two kinds of branches, called the axon and the dendrites.
The nerve cell is a microscopic speck of living matter. Its dendrites are short tree-like branches, while its axon is often several inches or even feet in length. The axon is the "slender thread", just spoken of as analogous to the single telephone wire. A nerve is composed of axons. [Footnote: The axon is always protected or insulated by a sheath, and axon and sheath, taken together, are often called a "nerve fiber".] The "white matter" of the brain and cord is composed of axons. Axons afford the means of communication between the nerve centers and the muscles and sense organs, and between one nerve center and another.
The axons which make up the motor nerves are branches of nerve cells situated in the cord and brain stem; they extend from the reflex center for any muscle out to and into that muscle and make very close connection with the muscle substance. A nerve current, starting from the nerve cells in the reflex center, runs rapidly along the axons to the muscle and arouses it to activity.
The axons which make up the optic nerve, or nerve of sight, are branches of nerve cells in the eye, and extend into the brain stem. Light striking the eye starts nerve currents, which run along these axons into the brain stem. Similarly, the axons of the nerve of smell are branches of cells in the nose.
The remainder of the sensory axons are branches of nerve cells that lie in little bunches close alongside the cord or {33} brain stem. These cells have no dendrites, but their axon, dividing, reaches in one direction out to a sense organ and in the other direction into the cord or brain stem, and thus connects the sense organ with its "lower center".
Fig. 5.--Sensory and motor axons, and their nerve
cells. The arrows indicate the direction of conduction. (Figure text:
eye, brain stem, skin, cord, muscle)
Where an axon terminates, it broadens out into a thin plate, or breaks up into a tuft of very fine branches ( the "end-brush"), and by this means makes close contact with the muscle, the sense organ, or the neurone with which it connects.
The Synapse
Now let us consider the mode of connection between one neurone and another in a nerve center. The axon of one neurone, through its end-brush, is in close contact with the dendrites of another neurone. There is contact, but no actual growing-together; the two neurones remain distinct, and this contact or junction of two neurones is called a "synapse". The synapse, then, is not a thing, but simply a junction between two neurones.
Fig. 6.--The synapse between the two neurones lies just
above the arrow.
The junction is good enough so that one of the two neurones, if itself active, can arouse the other to activity. The end-brush, when a nerve current reaches it from its own nerve cell, arouses the dendrites of the other neurone, and thus starts a nerve current running along those dendrites to their nerve cell and thence out along its axon.
Now here is a curious and significant fact: the dendrites are receiving organs, not transmitting; they pick up messages from the end-brushes across the synapse, but send out no messages to those end-brushes. Communication across a synapse is always in one direction, from end-brush to dendrites.
This, then, is the way in which a reflex is carried out, the pupillary reflex, for example. Light entering the eye starts a nerve current in the axons of the optic nerve; these axons terminate in the brain stem, where their end-brushes arouse the dendrites of motor nerve cells, and the axons of these {35} cells, extending out to the muscle of the pupil, cause it to contract, and narrow the pupil.
Or again, this is the way in which one nerve center arouses another to activity. The axons of the cells in the first center (or some of them) extend out of this center and through the white matter to the second center, where they terminate, their end-brushes forming synapses with the cells of the second center. Let the first center be thrown into activity, and immediately, through this connection, it arouses the second.
Fig. 7.--Different forms of synapse found in the
cerebellum, "a" is one of the large motor cells of the cerebellum (a
"Purkinje cell"), with its dendrites above and its axon below; and
"b," "c" and "d" show three forms of synapse made by other neurones
with this Purkinje cell. In "b," the arrow indicates a "climbing
axon," winding about the main limbs of the Purkinje cell. In "c," the
arrow points to a "basket"--an end-brush enveloping the cell body;
while "d" shows what might be called a "telegraph-wire synapse."
Imagine "d" superimposed upon "a": the axon of "d" rises among the
fine dendrites of "a," and then runs horizontally through them; and
there are many, many such axons strung among the dendrites. Thus the
Purkinje cell is stimulated at three points: cell body, trunks of the
dendrites, and twigs of the dendrites.
The "gray matter" comprises the nerve centers, lower and higher. It is made up of nerve cells and their dendrites, of the beginnings of axons issuing from these cells and of the terminations of incoming axons. The white matter, as was said before, consists of axons. An axon issues from the {36} gray matter at one point, traverses the white matter for a longer or shorter distance, and finally turns into the gray matter at another point, and thus nerve connection is maintained between these two points.
There are lots of nerve cells, billions of them. That ought to be plenty, and yet--well, perhaps sometimes they are not well developed, or their synapses are not close enough to make good connections.
Fig. 8.--A two-neurone reflex arc. (Figure text:
stimulus, skin, sensory axon, bit of the spinal cord, motor axon,
muscle)
Examined under the microscope, the nerve cell is seen to contain, besides the "nucleus" which is present in every living cell and is essential for maintaining its vitality and special characteristics, certain peculiar granules which appear to be stores of fuel to be consumed in the activity of the cell, and numerous very fine fibrils coursing through the cell and out into the axon and dendrites.
The reflex arc can now be described more precisely than before. Beginning in a sense organ, it extends along a sensory axon (really along a team of axons acting side by side) to its end-brush in a lower center, where it crosses a synapse and enters the dendrites of a motor neurone and so {37} reaches the cell body and axon of this neurone, which last extends out to the muscle (or gland). The simplest reflex arc consists then of a sensory neurone and a motor neurone, meeting at a synapse in a lower or reflex center. This would be a two-neurone arc.
Fig. 9.--A three-neurone arc, concerned in respiration.
This also illustrates how one nerve center influences another.
(Figure text: white matter, gray matter, lung, respiratory center in
the brain stem, diaphragm, motor center in cord for the diaphragm)
Very often, and possibly always, the reflex arc really consists of three neurones, a "central" neurone intervening between the sensory and motor neurones and being connected through synapses with each. The central neurone plays an important rôle in coördination.
COÖRDINATION
The internal structure of nerve centers helps us see how coördinated movement is produced. The question is, how {38} several muscles are made to work together harmoniously, and also how it is possible that a pin prick, directly affecting just a few sensory axons, causes a big movement of many muscles. Well, we find the sensory axon, as it enters the cord, sending off a number of side branches, each of which terminates in an end-brush in synaptic connection with the dendrites of a motor nerve cell.
Fig. 10.--Coördination brought about by the branching
of a sensory axon. (Figure text: cord, sensory neurone, motor neurone)
Thus the nerve current from a single sensory neurone is distributed to quite a number of motor neurones. Where there are central neurones in the arc, their branching axons aid in distributing the excitation; and so we get a big movement in response to a minute, though intense stimulus.
But the response is not simply big; it is definite, coordinated, representing team work on the part of the muscles as distinguished from indiscriminate mass action. That means selective distribution of the nerve current. The axons of the sensory and central neurones do not connect with any and every motor neurone indiscriminately, but link up with selected groups of motor neurones, and thus harness together teams that will work in definite ways, producing {39} flexion of a limb in the case of one such team, and extension in the case of another. Every reflex has its own team of motor neurones, harnessed together by its outfit of sensory and central neurones. The same motor neurone may however be harnessed into two or more such teams, as is seen from the fact that the same muscle may participate in different reflex movements; and for a similar reason we believe that the same sensory neurone may be utilized in more than one reflex arc.
Fig. 11.--Coördination brought about by the branching
of the axon of a central neurone. (Figure text: sensory, central,
motor)
The most distinctive part of any reflex arc is likely to be its central neurones, which are believed to play the chief part in coördination, and in determining the peculiarities of any given reflex, such as its speed and rhythm of action.
Reactions in General
Though the reflex is simple by comparison with voluntary movements, it is not the simplest animal reaction, for it is coördinated and depends on the nervous system, while the simplest animals, one-celled animals, have no nervous system, any more than they have muscles or organs of any {40} kind. Without possessing separate organs for the different vital functions, these little creatures do nevertheless take in and digest food, reproduce their kind, and move. Every animal shows at least two different motor reactions, a positive or approaching reaction, and a negative or avoiding reaction.
The general notion of a reaction is that of a response to a stimulus. The stimulus acts on the organism and the organism acts back. If I am struck by a wave and rolled over on the beach, that is passive motion and not my reaction; but if the wave stimulates me to maintain my footing, then I am active, I respond or react.
Now there is no such thing as wholly passive motion. Did not Newton teach that "action and reaction are equal"?--and he was thinking of stones and other inanimate objects. The motion of a stone or ball depends on its own weight and shape and elasticity as much as on the blow it receives. Even the stone counts for something in determining its own behavior.
A loaded gun counts for more than a stone, because of the stored energy of the powder that is set free by the blow of the hammer. The "reaction" of the gun is greater than the force acting on it, because of this stored energy that is discharged.
An animal reaction resembles the discharge of the gun, since there is stored energy in the animal, consisting in the chemical attraction between food absorbed and oxygen inspired, and some of this energy is utilized and converted into motion when the animal reacts. The stimulus, like the trigger of the gun, simply releases this stored energy.
The organism, animal or human, fully obeys the law of conservation of energy, all the energy it puts out being accounted for by stored energy it has taken in in food and oxygen. But at any one time, when the organism receives {41} a stimulus, the energy that it puts forth in reaction comes from inside itself.
There is another way in which the organism counts in determining its reaction. Not only does it supply the energy of the response, but its own internal arrangements determine how that energy shall be directed. That is to say, the organism does not blow up indiscriminately, like a charge of dynamite, but makes some definite movement. This is true even of the simplest animals, and the more elaborate the internal mechanism of the animal, the more the animal itself has to do with the kind of response it shall make to a stimulus. The nervous system of the higher animals, by the connections it provides between the stimulus and the stores of energy in the muscles, is of especial importance in determining the nature of the response.
Stimuli are necessary to arouse the activity of the organism. Without any stimulus whatever, it seems likely that the animal would relapse into total inactivity. It should be said, however, that stimuli, such as that of hunger, may arise within the organism itself. The stimulus may be external or internal, but some stimulus is necessary in order to release the stored energy.
In general, then, a reaction consists in the release by a stimulus of some of the stored energy of an animal, and the direction of that energy by the animal's own internal mechanism of nerves and muscles (and, we may add, bones and sinews) into the form of some definite response.
EXERCISES
1. Outline of the chapter, being at the same time a "completion test". Complete the following outline by filling in the blank spaces (usually a single word will fill the blank, but sometimes two words will be better):
A. Definition: A reaction is a response to a ___________.
The stimulus energy stored in the organism, and the __________
has a definite form determined by the organism's own machinery
of ________ and ______.
B. Among very prompt reactions are the reflex and the "simple
reaction". The reflex differs from the "simple reaction" in that:
(1) It usually takes less________.
(2) It requires no___________,
(3) The machinery for it is ________in the organism.
C. The machinery for a reflex consists of:
(1) a________ organ.
(2) a ________nerve.
(3) a nerve ________,
(4) a _________nerve.
(5) a muscle or _________.
D. The sensory and motor nerves consist of ________ which are
branches of ______. The cells for the motor nerves lie in the
________, and those for the sensory nerves lie in two cases in
the _________, and in all other cases in bunches located close
beside the _________or ________,
E. The neurone is the _______ of which the nervous ______ is
composed. It consists of a ________ and of two sorts of
branches, the ________ and the ________. Internally, the neurone
shows a peculiar structure of ________ and ________.
F. Communication from one neurone to another occurs across
a _____ called the synapse. The _________of an axon here comes
into close contact with the ______or with the _________of
another neurone. The communication takes place from the
________of the first neurone to the ___________ of the second.
G. The "nerve current" in a reflex therefore runs the following
course: from the sense organ into a ________ axon, along this to
its _________ in a nerve, and across a _________ there into the
_________ of a neurone, and thence {43} out along the _______of
this neurone to the ________or _________ that executes the
reflex. This is a two-neurone _________, but often there is a
third, ________neurone between the _________ and the
_____________.
H. Coördination is effected by the ________ of the axons of the
sensory and ________ neurones, by which means the nerve current
is ______ to a team of ________ and so to a team of _________.
2. Is the reaction time experiment, as described in the text, an
introspective or an objective experiment?
3. Mention two cases from common life that belong under the
"simple reaction", two that belong under "choice reaction", and two
that belong under the "associative reaction".
4. Arrange the reflexes mentioned in the text under the two heads
of "protective" and "regulative".
5. Draw diagrams of (a) the neurone, (b) a synapse, (c) a reflex
arc, and (d) a coördinated movement. Reduce each drawing to the
simplest possible form, and still retain everything that is
essential.
6. What part of the nervous system lies (a) in the forehead and
top of the head, (b) in the very back of the head, (c) along the
base of the skull, (d) within the backbone, (e) in the arm?
7. Using a watch to take the time, see how long it takes you to
name the letters in a line of print, reading them in reverse order
from the end of the line to the beginning. Compare with this time
the time required to respond to each letter by the letter following
it in the alphabet (saying "n" when you see m, and "t" when you see
s, etc.). Which of these two "stunts" is more like reflex action,
and how, nevertheless, does it differ from true reflex action?
8. The pupillary reflex. Describe the reaction of the pupil of the
eye to light suddenly shining into the eye. This response can best
be observed in another person, but you can observe it in yourself
by aid of a hand mirror. On another person you can also observe the
"crossed" pupillary reflex, by throwing the light into one eye only
while you watch the other eye. What sort of connection do you
suppose to exist between the two eyes, making this crossed reflex
possible?
9. The lid reflex, or wink reflex, (a) Bring your hand suddenly
close to another person's eye, and notice the response of the
eyelid, (b) See whether you can get a crossed reflex here, (c) See
whether your subject can voluntarily prevent (inhibit) the lid
reflex, (d) See whether the reflex occurs when he gives the
stimulus himself, by moving his own hand suddenly up to his eye.
(e) What other stimulus, besides the visual one that you have been
using, will arouse the same response?
REFERENCES
C. Judson Herrick, in his Introduction to Neurology, 2nd edition, 1918, gives a fuller and yet not too detailed account of the neurone in Chapter III, and of reflex action in Chapter IV.
Percy G. Stiles, in his Nervous System and Its Conservation, 1915, discusses these matters in Chapters II, III and IV.
Ladd and Woodworth's Elements of Physiological Psychology, 1911, has chapters on these topics.
CHAPTER III
REACTIONS OF DIFFERENT LEVELS
HOW SENSATIONS, PERCEPTIONS AND THOUGHTS MAY BE CONSIDERED AS FORMS OF INNER RESPONSE, AND HOW THESE HIGHER REACTIONS ARE RELATED IN THE NERVOUS SYSTEM TO THE SIMPLER RESPONSES OF THE REFLEX LEVEL.
Having defined a reaction as an act of the individual aroused by a stimulus, there is no reason why we should not include a great variety of mental processes under the general head of reactions. Any mental process is an activity of the organism, and it is aroused by some stimulus, external or internal; therefore, it is a reaction.
I hear a noise--now, while the noise, as a physical stimulus, comes to me, my hearing it is my own act, my sensory reaction to the stimulus. I recognize the noise as the whistle of a steamboat--this recognition is clearly my own doing, dependent on my own past experience, and may be called a perception or perceptive response. The boat's whistle reminds me of a vacation spent on an island--clearly a memory response. The memory arouses an agreeable feeling--an affective response, this may be called. In its turn, this may lead me to imagine how pleasant it would be to spend another vacation on that island, and to cast about for ways and means to accomplish this result--here we have imagination and reasoning, aroused by what preceded just as the sensation was aroused by the physical stimulus.
In speaking of any mental process as an act of the individual, we do not mean to imply that he is always conscious {46} of his activity. Sometimes he feels active, sometimes passive. He feels active in hard muscular work or hard thinking, while he feels passive in reflex action, in sensation, and in simply "being reminded" of anything without any effort on his own part. But he is active in everything he does, and he does everything that depends on his being alive. Life is activity, and every manifestation of life, such as reflex action or sensation, is a form of vital activity. The only way to be inactive is to be dead.
But vital activity is not "self-activity" in any absolute sense, for it is aroused by some stimulus. It does not issue from the individual as an isolated unit, but is his response to a stimulus. That is the sense of calling any mental process a reaction; it is something the individual does in response to a stimulus.
To call a sensation a form of reaction means, then, that the sensation is not something done to the person, nor passively received by him from outside, but something that he himself does when aroused to this particular form of activity. What comes from outside and is received by the individual is the stimulus, and the sensation is what he does in response to the stimulus. It represents the discharge of internal stored energy in a direction determined by his own inner mechanism. The sensation depends on his own make-up as well as on the nature of the stimulus, as is especially obvious when the sensation is abnormal or peculiar. Take the case of color blindness. The same stimulus that arouses in most people the sensation of red arouses in the color-blind individual the sensation of brown. Now what the color-blind individual receives, the light stimulus, is the same as what others receive, but he responds differently, i.e., with a different sensation, because his own sensory apparatus is peculiar.
The main point of this discussion is that all mental {47} phenomena, whether movements, sensations, emotions, impulses or thoughts, are a person's acts, but that every act is a response to some present stimulus. This rather obvious truth has not always seemed obvious. Some theorists, in emphasizing the spontaneity and "self-activity" of the individual, have pushed the stimulus away into the background; while others, fixing their attention on the stimulus, have treated the individual as the passive recipient of sensation and "experience" generally. Experience, however, is not received; it is lived, and that means done; only, it is done in response to stimuli. The concept of reaction covers the ground.
While speaking of sensations and thoughts as belonging under the general head of reactions, it is well, however, to bear in mind that all mental action tends to arouse and terminate in muscular and glandular activity. A thought or a feeling tends to "express itself" in words or (other) deeds. The motor response may be delayed, or inhibited altogether, but the tendency is always in that direction.
Different Sorts of Stimuli
To call all mental processes reactions means that it is always in order to ask for the stimulus. Typically, the stimulus is an external force or motion, such as light or sound, striking on a sense organ. There are also the internal stimuli, consisting of changes occurring within the body and acting on the sensory nerves that are distributed to the muscles, bones, lungs, stomach and most of the organs. The sensations of muscular strain and fatigue, and of hunger and thirst, are aroused by internal stimuli, and many reflexes are aroused in the same way.
Such internal stimuli as these are like the better known external stimuli in that they act upon sense organs; but it {48} seems necessary to recognize another sort of stimuli which act directly on the nerve centers in the brain. These may be called "central stimuli" and so contrasted with the "peripheral stimuli" that act on any sense organ, external or internal. To do this is to take considerable liberty with the plain meaning of "stimulus", and calls for justification. What is the excuse for thus expanding the notion of a stimulus?
The excuse is found in the frequent occurrence of mental processes that are not directly aroused by any peripheral stimulus, though they are plainly aroused by something else. Anything that arouses a thought or feeling can properly be called its stimulus. Now it often happens that a thought is aroused by another, just preceding thought; and it seems quite in order to call the first thought the stimulus and the second the response. A thought may arouse an emotion, as when the thought of my enemy, suddenly occurring to mind, makes me angry; the thought is then the stimulus arousing this emotional response.
If hearing you speak of Calcutta makes me think of India, your words are the stimulus and my thought the response. Well, then, if I think of Calcutta in the course of a train of thought, and next think of India, what else can we say than that the thought of Calcutta acts as a stimulus to arouse the thought of India as the response? In a long train of thought, where A reminds you of B and B of C and C of D, each of these items is, first, a response to the preceding, and, second, a stimulus to the one following.
There is no special difficulty with the notion of "central stimuli" from the physiological side. We have simply to think of one nerve center arousing another by means of the tract of axons connecting the two. Say the auditory center is aroused by hearing some one mention your friend's name, {49} and this promptly calls up a mental picture of your friend; here the auditory center has aroused the visual. What happens in a train of thought is that first one group of neurones is aroused to activity, and then this activity, spreading along the axons that extend from this group of neurones to another, arouses the second group to activity; and so on. The brain process may often be exceedingly complex, but this simple scheme gives the gist of it.
The way nerve currents must go shooting around the brain from one center or group of neurones to another, keeping it up for a long time without requiring any fresh peripheral stimulus, is remarkable. We have evidence of this sort of thing in a dream or fit of abstraction. Likely enough, the series of brain responses would peter out after awhile, in the absence of any fresh peripheral stimulus, and total inactivity ensue. But response of one brain center to nerve currents coming from another brain center, and not directly from any sense organ, must be the rule rather than the exception, since most of the brain neurones are not directly connected with any sense organ, but only with other parts of the brain itself. All the evidence we have would indicate that the brain is not "self-active", but only responsive; but, once thrown into activity at one point, it may successively become active at many other points, so that a long series of mental operations may follow upon a single sensory stimulus.
The Motor Centers, Lower and Higher
A "center" is a collection of nerve cells, located somewhere in the brain or cord, which gives off axons running to some other center or out to muscles or glands, while it also receives axons coming from other centers, or from sense organs. These incoming axons terminate in end-brushes and so form synapses with the dendrites of the local {50} nerve cells. The axons entering any center and terminating there arouse that center to activity, and this activity, when aroused, is transmitted out along the axons issuing from that center, and produces results where those axons terminate in their turn.
Fig. 12.--Side view of the left hemisphere of the brain, showing the
motor and sensory areas (for the olfactory area, see
Fig. 18). The visual area proper, or
"visuo-sensory area," lies just around the corner from the spot
marked "Visual," on the middle surface of the hemisphere, where it
adjoins the other hemisphere. (Figure text: frontal lobe, parietal lobe,
central fissure, occipital lobe, motor area, somesthetic area, auditory
area, fissure of Sylvius, temporal lobe, brain stem, cerebellum)
The lower motor centers, called also reflex centers, are located in the cord or brain stem, and their nerve cells give rise to the axons that form the motor nerves and connect with the muscles and glands. A muscle is thrown into action by nerve currents from its lower motor center.
The principal higher motor center is the "motor area" of the brain, located in the cortex or external layer of gray matter, in the cerebrum. More precisely, the motor area is a long, narrow strip of cortex, lying just forward of what is called the "central fissure" or "fissure of Rolando".
If you run your finger over the top of the head from one side to the other, about halfway back from the forehead, the motor areas of the two cerebral hemispheres will lie close under the path traced by your finger.
Fig. 13.--(After Cajal.) Type of the brain cells that
most directly control muscular movement.
(Figure text: Axon. Giant pyramid cell from the motor area of the cerebral cortex, magnified 35 diameters. Cell body of same farther magnified)
The motor area in the right hemisphere is connected with the left half of the cord and so with the muscles of the left half of the body; the motor area of the left hemisphere similarly affects {52} the right half of the body. Within the motor area are centers for the several limbs and other motor organs. Thus, at the top, near the middle line of the head (and just about where the phrenologists located their "bump of veneration"!), is the center for the legs; next below and to the side is the center for the trunk, next that for the arm, next that for head movements, and at the bottom, not far from the ears, is the center for tongue and mouth.
Fig. 14.--The nerve path by which the motor area of the
cortex influences the muscles. The upper part of this path, consisting of axons issuing from the giant pyramids of the motor area and extending down into the spinal cord, is the pyramidal tract. The lower part of the path consists of axons issuing from the motor cells of the cord and extending out to the muscles. The top of the figure represents a vertical cross-section of the brain, such as is given, on a larger scale, in Fig. 18. (Figure text: cortex, cord, muscles)
The largest nerve cells of all are found in the motor area, and are called, from their shape, the "giant pyramids". They have large dendrites and very long axons, which latter, {53} running in a thick bundle down from the cortex through the brain stem and cord, constitute the "pyramidal tract", the principal path of communication from the cerebrum to the lower centers. The motor area of the brain has no direct connection with any muscle, but acts through the pyramidal tract on the lower centers, which in turn act on the muscles.
How The Brain Produces Muscular Movements
The motor area is itself aroused to action by nerve currents entering it through axons coming from other parts of the cortex; and it is by way of the motor area that any other part of the cortex produces bodily movement. There are a few exceptions, as, for example, the movements of the eyes are produced generally by the "visual area" acting directly on the lower motor centers for the eye in the brain stem; but, in the main, any motor effect of brain action is exerted through the motor area. The motor area, as already mentioned, acts on the lower motor centers in the cord and brain stem, and these in turn on the muscles; but we must look into this matter a little more closely.
A lower motor center is a group of motor and central neurones, lying anywhere in the cord or brain stem, and capable of directly arousing a certain coördinated muscular movement. One such unit gives flexion of the leg, another gives extension of the leg, a third gives the rapid alternation of flexion and extension that we see in the scratching movement of the dog. Such a motor center can be aroused to activity by a sensory stimulus, and the resulting movement is then called a reflex.
The lower center can be aroused in quite another way, and that is by nerve currents coming from the brain, by way of the motor area and the pyramidal tract. Thus flexion of the leg can occur voluntarily as well as reflexly. The same {54} muscles, and the same motor neurones, do the job in either case. In the reflex, the lower center is aroused by a sensory nerve, and in the voluntary movement by the pyramidal tract.
The story is told of a stranger who was once dangling his legs over the edge of the station platform at a small backwoods town, when a native called out to him "Hist!" (hoist), pointing to the ground under the stranger's feet. He "histed" obediently, which is to say that he voluntarily threw into play the spinal center for leg flexion; and then, looking down, saw a rattler coiled just beneath where his feet had been hanging. Now even if he had spied the rattler first, the resulting flexion, though impulsive and involuntary, would still have been aroused by way of the motor area and the pyramidal tract, since the movement would have been a response to knowledge of what that object was and signified, and knowledge means action by the cerebral cortex, which we have seen to affect movement through the medium of the motor area. But if the snake had made the first move, the same leg movement on the man's part, made now in response to the painful sensory stimulus, would have been the flexion reflex.
Facilitation and Inhibition
Not only can the motor area call out essentially the same movements that are also produced reflexly, but it can prevent or inhibit the execution of a reflex in spite of the sensory stimulus for the reflex being present, and it can reinforce or facilitate the action of the sensory stimulus so as to assist in the production of the reflex. We see excellent examples of cerebral facilitation and inhibition in the case of the knee jerk. This sharp forward kick of the foot and lower leg is aroused by a tap on the tendon running in front {55} of the knee. Cross the knee to be stimulated over the other leg, and tap the tendon just below the knee cap, and the knee jerk appears. So purely reflex is this movement that it cannot be duplicated voluntarily; for, though the foot can of course be voluntarily kicked forward, this voluntary movement does not have the suddenness and quickness of the true reflex. For all that, the cerebrum can exert an influence on the knee jerk. Anxious attention to the knee jerk inhibits it; gritting the teeth or clenching the fist reinforces it. These are cerebral influences acting by way of the pyramidal tract upon the spinal center for the reflex.
Thus the cortex controls the reflexes. Other examples of such control are seen when you prevent for a time the natural regular winking of the eyes by voluntarily holding them wide open, or when, carrying a hot dish which you know you must not drop, you check the flexion reflex which would naturally pull the hand away from the painful stimulus. The young child learns to control the reflexes of evacuation, and gradually comes to have control over the breathing movements, so as to hold his breath or breathe rapidly or deeply at will, and to expire vigorously in order to blow out a match.
The coughing, sneezing and swallowing reflexes likewise come under voluntary control. In all such cases, the motor area facilitates or inhibits the action of the lower centers.
Super-motor Centers in the Cortex
Another important effect of the motor area upon the lower centers consists in combining their action so as to produce what we know as skilled movements. It will be remembered that the lower centers themselves give coördinated movements, such as flexion or extension of the whole limb; but still higher coördinations result from cerebral control. {56} When the two hands, though executing different movements, work together to produce a definite result, we have coördination controlled by the cortex. Examples of this are seen in handling an ax or bat, or in playing the piano or violin. A movement of a single hand, as in writing or buttoning a coat, may also represent a higher or cortical coördination.
Fig. 15.--(From Starr.) Axons connecting one part of
the cortex with another. The brain is seen from the side, as if in
section. At "A" are shown bundles of comparatively short axons,
connecting near-by portions of the cortex; while "B," "C," and "D"
show bundles of longer axons, connecting distant parts of the cortex
with one another. The "Corpus Callosum" is a great mass of axons
extending across from each cerebral hemisphere to the other, and
enabling both hemispheres to work together. "O. T." and "C. N." are
interior masses of gray matter, which can be seen also in
Fig. 18. "O. T." is the thalamus, about which more later.
Now it appears that the essential work in producing these higher coördinations of skilled movement is performed not by the motor area, but by neighboring parts of the cortex, which act on the motor area in much the same way as the motor area acts on the lower centers. Some of these {57} skilled-movement centers, or super-motor centers, are located in the cortex just forward of the motor area, in the adjacent parts of the frontal lobe. Destruction of the cortex there, through injury or disease, deprives the individual of some of his skilled movements, though not really paralyzing him. He can still make simple movements, but not the complex movements of writing or handling an instrument.
It is a curious fact that the left hemisphere, which exerts control over the movements of the right hand and right side of the body generally, also plays the leading part in skilled movements of either hand. This is true, at least, of right-handed persons; probably in the left-handed the right hemisphere dominates.
Motor power may be lost through injury at various points in the nervous system. Injury to the spinal cord, destroying the lower motor center for the legs, brings complete paralysis. Injury to the motor area or to the pyramidal tract does not destroy reflex movement, but cuts off all voluntary movement and cerebral control. Injury to the "super-motor centers" causes loss of skilled movement, and produces the condition of "apraxia", in which the subject, though knowing what he wants to do, and though still able to move his limbs, simply cannot get the combination for the skilled act that he has in mind.
Speech Centers
Similar to apraxia is "aphasia" or loss of ability to speak. It bears the same relation to true paralysis of the speech organs that hand apraxia bears to paralysis of the hand. Through brain injury it sometimes happens that a person loses his ability to speak words, though he can still make vocal sounds. The cases differ in severity, some retaining the ability to speak only one or two words which {58} from frequent use have become almost reflex (swear words, sometimes, or "yes" and "no"), while others are able to pronounce single words, but can no longer put them together fluently into the customary form of phrases and sentences, and still others can utter simple sentences, but not any connected speech.