drew diagrams and made models of the nervous system in which they
represented it as composed of conducting arcs. The arcs were superposed
to indicate that they were of various grades—spinal for ordinary
reflexes, bulbar for co-ordinated actions, through the grey matter
in the centre of the great brain for “ideo-motor” actions, through
the cortex of the great brain for voluntary acts. They spoke of
authority and responsibility, comparing the nervous system to an
army or a club. It is premature to attempt a theory of the nervous
system compatible with recent discoveries regarding its structure and
mode of working, but it is clear that the diagrams and metaphors to
which we have just referred were misleading. In place of attempting
to disarticulate the machine, we ought to emphasize its structural
unity. The results obtained by uniting heterologous nerves cannot be
explained by reference to a model made of wires and pieces of cork.
They do not fit in with any organization of human units or with any
postal system or telephonic apparatus for transmitting news. Probably
the lines of thought which will prove most fruitful are somewhat as
follows: (1) An efferent discharge occurs as the result of the opening
of a circuit from a muscle back to the muscle. Afferent impulses—call
them sensory, on the understanding that this does not imply that they
appear in consciousness—are ceaselessly flowing from receptors to
effectors in the muscle. A sensation—in the case of skeletal muscles
usually a skin sensation—reinforces them to discharging-point. If
the spinal cord has been severed from the brain, the up-and-down flow
does not reach beyond its grey matter. It is short-circuited. If the
brain is in normal connection with the spinal cord, sensory impulses
travel upwards to its cortex (without, save in exceptional instances,
arousing consciousness, or, as we should prefer to express it in this
connection, without attracting attention) to a degree which varies with
the several classes of receptor and with the animal. A monkey reduced
to the condition of a “spinal animal”—_i.e._, with its spinal cord
severed from its brain—is less competent than a dog, and a man is far
less competent than a monkey. In other words, a man habitually uses
his brain more than does a monkey, and a monkey more than a dog. The
proportion which brain-weight bears to body-weight roughly indicates
the part the brain plays in conducting the traffic of the body. (2)
Communication within the nervous system is almost unrestricted. If,
before the median nerve was divided at the wrist, receptor A usually
initiated a current which passed through the circuit to effector X, and
receptor B to effector Y, and if the new fibres which grew downwards
lost their way so that the one which used to receive messages from A
attached itself to B, and the one which used to transmit commands to
X attached itself to Y, A is not thereby cut off from X, or B from
Y. Such a mechanical association is restricted to our diagrams. It
does not enter into Nature’s plan. The spinal cord is not scored with
unchangeable paths. A messenger from A could always reach either X or
Y. It was not the path, but the struggle with competing messengers,
which directed him to X.
When we endeavour to picture the mechanism of the nervous system, we
find ourselves faced by phenomena which appear irreconcilable. One set
of observations leads to the conception of closed paths; another set
points to an open conductor. The experimental crossing of nerves to
which we have just alluded shows that the nervous system is adaptable,
to a degree which seems extraordinary to anyone who attempts to compare
it with any of Man’s devices for establishing communication. Paths
appear to make themselves. On the other hand, the more important, and
therefore dominant, reflex actions, such as swallowing, breathing,
the maintenance of position, are due to the union of receptors and
effectors by lines which are either reserved for their sole use, or, if
shared by other currents, it is on the understanding that they have a
first and altogether prepotent claim. No competing impulses can divert
them or block their way. All reflexes which in the history of the race
have established their right to dominance not only seize and hold a
route through the nervous system, to the exclusion of all competitors,
but, as we have already shown in the case of the swallowing impulse,
the traffic in neighbouring routes is suspended for their benefit.
At the other end of the scale we find reflexes which may be termed
“occasional,” in that, although of frequent occurrence, they exhibit
illimitable variability in form. Occasional reflexes require, as a
preliminary to their transmission, that the afferent impulses which
give rise to them should secure for a time the exclusive use of the
motor neurones by which they are carried out. The receptors bring the
motor neurones into tune with themselves, and while in tune they will
respond to impulses from no others. But the tuning lasts for a short
time only. Either receptor or neurone, or both, soon tire. There is no
danger of a particular reflex being prolonged to the detriment of the
organism as a whole. As an illustration of an occasional reflex, we may
cite the scratching movement of a dog. Its skin is punctured by a flea.
It scratches the place. A second flea bites it somewhere in the same
neighbourhood. The dog does not shift its hind-foot so as to scratch
midway between the two bites. It finishes out one scratch before paying
attention to its second tormentor. The exact position to which the
hind-foot is raised depends upon the position of the irritant; and
since this may be shifted over a very considerable surface, the form of
the reflex varies equally widely. Each of the very numerous receptors
in the skin tunes a slightly different group of motor neurones; and
since a second irritant may reinforce the first, instead of making an
alteration in the group of neurones which the reflex is discharging, it
is clear that there is no fixed path uniting receptor A with neurones
X, Y, Z and receptor B with neurones W, X, Y. If, however, the second
irritation occurs at a spot lying at a considerable distance from
A, in place of reinforcing the scratching movement which A has set
going, it weakens and shortens it. The receptor C, which is calling
for the discharge of a markedly different set of motor neurones, tends
to inhibit those which are already active. These results are tested
with precision upon a “spinal dog” and with the aid of an electric
needle, the other pole from the battery being a large flat plate
placed in contact with the animal’s body. The conception of definite
paths, to which the contemplation of permanent reflexes gives rise, is
inappropriate to occasional reflexes. The latter show so wide a range
of variability and adaptability as to prove that a given receptor
may bring any of a great variety of groups of motor neurones into
connection with itself; just as a given group of neurones may be played
upon by impulses from a great number of different receptors. We have
called it a tuning of the motor neurones. One metaphor is as good as
another. The physical process which in the brainless frog underlies the
preparation for discharging motor neurones in the spinal cord, on the
same side as the leg on which vinegar is placed, so long as that leg
is free, and on the opposite side, when that leg is fixed, is unknown.
We seem to catch a glimpse of a doubleness of action, receptors in
the muscles combining with receptors in the skin in determining the
paths along which impulses shall be reflected—the efficient muscles
sensitizing their own neurones to the tuning influence of impulses from
sounding cutaneous nerve-endings. But it is impossible to formulate a
working scheme in the present state of knowledge.
=Sense-Organs and Nerve-Centres.=—A vast amount of labour has been
devoted to the study of the external form of the central nervous
system and to unravelling its internal structure; to plotting out its
various groups of nerve-cells, to disentangling its innumerable tracts
of fibres. The surface of the brain and spinal cord has been mapped
and measured. Every millimetre of its substance has been cut into
sections on the micro-tome. Organs which, fifty years ago, appeared
too complicated for investigation have been described in the minutest
detail. An immense accumulation of data is available for purposes of
reference; yet anyone who submits the theory of the nervous system as
it is held at the present day to a general review must allow that the
results of anatomical research enter but little into its construction.
The reason for this is not far to seek. As knowledge has advanced, the
apparent, or rather the expected, complication of the system has given
place to ideas of unity and simplicity. Its external configuration and
the varied arrangement in “nuclei” of its nerve-cells may, without
impropriety, be described as accidental. The form of the body and the
consequent location of the clients of the nervous system determine
the disposition and degree of concentration of its various business
centres. It shows, when followed throughout the whole animal kingdom,
extreme variability of its constituent organs, with absolute uniformity
of plan. Indeed, from the physiological point of view the term “organ”
is scarce admissible. It implies diversity of function in too high a
degree. The several parts into which the central nervous system is
obviously divisible co-operate so intimately as to preclude us from
thinking of them as separate organs.
If the citadel of the central nervous system is to be captured, all
lines of approach must be tried. Its outward form must be studied,
its minute structure examined with the microscope, its modifications
in various animals compared, its development followed, its reactions
to artificial stimuli tested, its pathological deficiencies and
vagaries watched. Yet, of all the means which have been made use of in
attempting to penetrate its secrets, the study of its history, by the
methods of comparative anatomy and embryology, has probably contributed
most to the development of sound ideas regarding the manner of its
working. The first differentiation visible in the blastoderm—the
globe of cells into which the ovum divides and out of which the embryo
is built—has relation to the formation of the nervous system. If
the earliest stages of its growth are followed, and the different
phases through which it passes are compared with the forms which it
assumes permanently in lower animals, the plan or type upon which
it is constructed shows up distinctly. Looking down the line to the
earliest vertebrata, we can discern clearly the form of nervous system
possessed by their prototype. Not that this “ideal ancestor” ever
existed. Experience teaches that it is unlikely that any animal that
ever lived was absolutely regular and symmetrical in all its parts;
nevertheless, the type can be presented in a perfectly regular scheme.
The ideal ancestor of the vertebrata was segmented, like a caterpillar
or a worm. Its mouth was not at the anterior extremity of the body, but
two (or more) segments behind it. Every segment bore a sense-organ (at
one period two sense-organs) on either side. Beneath each sense-organ
there was a clump of “grey matter.” Each segment also contained
(although not at the earliest epoch) two clumps of nerve-cells and
neuropil in a more central situation. These “ganglia” were united by
longitudinal and transverse commissures. They received the axons of the
cells which lay in the clumps beneath the sense-organs. They gave axons
to various muscles. Such is the type out of which the modern nervous
system has developed: two separate sense-organs and a complete nervous
system for each segment, the sense-organs connected with the ganglion
of the same side, the ganglia of the two sides bound together across
the middle line, and each row of sense-organs and each row of ganglia
united by longitudinal commissures into a chain. From the nervous
system as we see it now the majority of these segmental sense-organs
have disappeared; but the mode of formation of the cerebro-spinal
ganglia shows that they are the clumps of nerve-cells which lay beneath
the vanished organs. In the nose and the eye the grey matter retains
its original situation in the immediate vicinity of the receiving
epithelial cells—as the olfactory bulb and the deeper (anterior)
layers of the retina. The ganglia of the auditory nerve lie within the
bones of the ear. Spinal ganglia are close to the spinal cord. Auditory
and spinal ganglia contain only the cell-bodies of the first collecting
neurones (sensory nerves) together with certain curious bracketing
cells already referred to (p. 324), all the other constituents of the
peripheral clumps of grey matter which are found in the olfactory bulb
and retina having been withdrawn from the spinal ganglia into the axis
of the brain and spinal cord.
The sense-organs in front of the mouth have had from the beginning
an immense advantage over the others as observing-stations. Whereas
the body-organs collected information regarding the things with which
the animal came in contact, and consequently specialized in touch,
pressure, temperature, and, in the case of fishes, sensitiveness to
the chemical constitution of the medium in which the animal lived,
the head-organs specialized in responsiveness to forces acting from a
distance—particles suspended in air, vibrations of light, pulsations
of sound. Sensitiveness to touch, if it is to be useful, must be
widely distributed. The body-organs therefore broke into scattered
groups of sense-cells. Touch-spots are scattered all over the surface,
although they are set much closer together in the areas of skin which
are usually the first to come into contact with external objects
than they are elsewhere. The efficiency of the sense-organs of the
head—nose, eye, and ear—depended upon their remaining compact.
Progress in animal life, as we understand it—the rise from lower to
higher forms—has depended upon increasing integration of the body and
co-ordination of its functions. The nervous system is the agent which
has accomplished this unification. Each step in advance has depended
upon the provision of more nerve-tissue for the lacing together of
the various parts. We have seen already (p. 329) how intimate is
the union of receptors and effectors of every kind via the spinal
cord and brain. The overwhelming predominance in the direction of
action of the nose, the eye, and the ear has led to the accumulation
in their vicinity of the ever-increasing grey matter. The cerebral
hemispheres, or “great brain,” are pouched outgrowths from the first
pair of ganglia directed towards the olfactory pits. The original eyes
bore a similar relation to the second pair of ganglia—the epithet
“original” implying that the eyes which we now use are not the organs
with which our prevertebrate ancestors saw. First one of the original
eyes disappeared, and then the other. The vestige of the second is
still to be seen in the “pineal body” which is found on the dorsal
side of the brain of every vertebrate animal—in a mammal deeply
hidden in the cleft between the cerebrum and cerebellum. In place of
the pineal eyes two other sense-organs have specialized as eyes. They
are constructed on a different plan, being, to put it shortly, pineal
eyes turned inside out; for whereas in the pineal eyes, as in most of
the eyes of invertebrate animals, the rods and cones, which are the
cells of the retina sensitive to light, are directed forwards towards
the lens, the rods and cones of our permanent eyes are directed away
from the source of light. This change has made it possible to provide
more abundantly for their nutrition, and hence a greater power of
discriminating separate points in space and of distinguishing colours
is conferred upon them. The substitution of other sense-organs for the
original eyes has complicated the pictures which are presented to us by
a brain in its successive stages of growth; but it does not prevent us
from recognizing the general plan. Probably the secondary eyes, like
their predecessors, belonged to a pre-oral segment. The sense-organs
of a segment behind the mouth developed into ears; and the ear was
in its earliest phases, and still is, something more than an organ of
hearing. Its semicircular canals give information of displacements
in space. Knowledge of the position of its body is, to a fish, of
far more importance than its ability to hear breakers on the rocks.
Three looped tunnels, opening at either end into a common chamber, are
hollowed in the bone which contains the ear (_cf._ Fig. 38). Placed
at right angles one to the other, they occupy all three dimensions of
space. Open a notebook until, one of its covers lying horizontally,
the other is vertical, and place a sheet of paper vertically against
the bottom of the pages. A curved line drawn on each of these three
surfaces will represent the three semicircular canals. Arrange another
notebook in the same way, and let the two rest on the table with the
two vertical covers inclining one to the other, anteriorly, at an
angle of 90 degrees. The six surfaces will be in the planes of the six
semicircular canals. Within each bony canal is a membranous tube, to
which nerves are distributed, filled with fluid. When the position of
the head is changed, the fluid within the membranous tubes slides on
their walls. It is left behind at the moment the movement commences. It
overtakes its receptacle when the movement stops. The stimulus received
by the nerve-endings is recognized as indicating an alteration in the
orientation of the head. If the movement of the fluid is violent,
as when one waltzes, the loss of the sense of position disconcerts
the brain to such an extent that giddiness results. For a time the
quiet assurance upon which so much depends, that one knows how the
body stands in relation to its surroundings, gives way to a chaos of
sensations. From the nature of the case, the information which the
semicircular canals afford relates to change. They give no help in
ascertaining the position of the head when it is at rest. This must be
the reason, although the connection is not very clear, for the waning
of the effect in consciousness when stimulation is prolonged, and also
for the very marked after-sensation. At the commencement of a voyage
attention may be unpleasantly attracted to the rolling of the ship.
After a few days it ceases to be noticeable; yet when the voyager,
the night after landing, wakes in the dark, he finds his bed-room as
unsteady as his cabin. Rising hurriedly, the attempt to adjust his
position to the heaving floor (we speak from personal experience)
may result in a heavy fall. Although this phenomenon must be classed
with other “after-sensations,” it is so prolonged as to suggest that
consciousness, having become accustomed to a world which causes a
backward and forward flow of endolymph, misinterprets the absence of
sensation as indicative of change.
Taste is, practically, a special kind of smell. A fish’s olfactory
membrane, taste-buds, and chemical organs “of the lateral line” serve
the same sense, although, no doubt, they are applicable to the analysis
of different forms of matter in solution.
Our ideal prevertebrate has now left its primitive undifferentiated
condition. In front of its mouth it bears organs with which it searches
the world. Close behind the mouth are its auditory and orienting
organs. The rest of the surface of the body is endowed with the
capacity of recognizing “taste,” temperature, and contact. Smell,
sight, and orientation determine the development of the brain.
The cerebrum which has eventually become, as the seat of consciousness,
and hence the apparatus of mind, the dominant factor in the nervous
system, was in the first instance the part of the brain concerned with
the distribution to the muscles of impulses generated in olfactory
organs. There is scarcely any indication in a fish’s brain of the
representation in the cerebral hemispheres of any other sense, even
that of vision.
A bird’s brain presents a striking contrast to the brain of a fish.
With the exception of the apteryx and other ground-birds of New
Zealand, all birds are apparently destitute of the sense of smell.
Vision is the sense upon which their activity depends. It has invaded
the cerebrum, converting it into an organ in which sensations of
sight are worked up into “mind-stuff.” The optic lobe connection is
restricted to the production of reflex actions in which vision is
immediately followed by movement.
All the senses are represented in the great brains of mammals.
The cerebrum, which owes its existence to its connection with the
favourably-situated sense-organ of the nose, and grew in importance
when vision invaded it, has now taken in the senses of hearing, taste,
and touch. Only what may be termed in general visceral sense, and the
sense of orientation, are excluded.
Looking back to the starting-point, we see a segmented animal; its
segments of equal value; its nervous reactions unisegmental, although
linked in functional sequence. If it starts to walk, owing to
stimulation of one of its sense-organs, the impulse to walk spreads
from segment to segment. Comparing the latest product of evolution
with the earliest, we find that nervous tissue has concentrated
at the anterior end of the body. The double chain of ganglia, now
condensed into the axis of the brain and the spinal cord, still
contain all the effector neurones by which muscles are called into
action. Sensory nerves still arborize in the axis, providing the
mechanism for actuating motor neurones. But the vast majority of
intermediate or intercalated neurones have been attracted to the two
huge brain-masses—the cerebellum and cerebrum. In the former all
sensations (not conscious) connected with tone, position, orientation
and equilibrium are worked into appropriate impulses for the regulation
of the muscular system. In the latter all sensations which convey
information regarding the relation of the environment, including the
body, to the ego—the _not-me_ to the _me_—are transformed into motor
discharges which set a-going the movements (and the thoughts) by means
of which the purposes of life are fulfilled; for in the cortex of the
great brain alone is the passage of nerve-currents accompanied by
consciousness. Concentration of nerve-tissue allows of the combination
of sensations. It also facilitates the no less important effect
of mutual influence, interference. Sensations are suppressed, and
therefore the multitude of reactions to which they would give rise are
inhibited, in the interests of restricted and sustained movement or
thought.
=The Cerebellum.=—Sharks and other swift-swimming fishes have large,
deeply fissured cerebella, for the cerebellum is the part of the brain
which has gathered into itself most of the grey matter associated
with balancing, attitude, posture. The cerebellum is in birds large
and deeply folded. Developed from the ganglia to which the auditory
nerve distributes impulses from the semicircular canals, it has
established connections with all the other nervous tissues concerned
with sensations of position, strain, or pressure, including the
eyes, which afford information regarding the position of our limbs
relatively to the trunk, and of the whole body relatively to external
objects. Morphologically it is a median growth. The adverb is one of
those qualifying terms, convenient in science, which direct thought
without confining it. As used above, it implies that anyone who passes
before his mind the cerebella of all animals from fishes to Man, and
in all stages of growth, from their earliest appearance in the embryo
to their condition in the adult, sees the organ as a median prominence
surmounting the medulla oblongata. The bulgings of its sides which,
in human anatomy, are termed hemispheres, do not disturb its central,
unpaired plan of structure. It has, it is true, a lateral appendage
on either side (the combined flocculus and paraflocculus of mammalian
anatomy), but this lobe, although of great historic interest, is
so small, as compared with the median growth, as not to affect our
general conception of the form of the organ. By transverse fissures the
cerebellum is divided into a series of lobes.
In appearance the cerebellum varies greatly in the different classes
and orders of Vertebrata. Yet underlying this variety there is
marked unity of plan. A sagittal section of the organ of a shark, of
a bird, of a kangaroo, of a dog, of a whale, of Man, shows that it
is divided, from before backwards, into the same number of lobes in
animals occupying every position from the bottom to the top of the
vertebrate scale. A very little effort to grasp the significance of
this mystic number, nine, convinces one of the hopelessness of any
attempt to correlate the form of the cerebellum with the muscular
development or sensory endowments of vertebrates as a sub-kingdom.
It is the same for animals with limbs and animals without; animals
with well-developed noses or eyes, and animals destitute of one or
other of these sense-organs. This uniformity is extremely significant,
when contrasted with the wide differences exhibited by the cerebral
hemispheres. It shows that, unlike the great brain which mediates
between the several senses and the muscular system, the little brain is
concerned in bringing about adjustments to the environment which are
equally important to all animals, no matter how far they may depart
from the common type. The cerebellum is crossed by deep fissures,
dividing it into narrow convolutions or folia. The folia are grouped
in nine lobes. If the reader has secured as an illustration the brain
of a sheep, he will notice that the lateral regions of the cerebellum
present a complicated appearance owing to the contortion of the folia,
which results from the unequal development on its sides of the several
lobes. In its total size the cerebellum keeps step with the cerebrum,
the right side of one organ being associated with the left side of the
other.
[Illustration: FIG. 23.—VERTICAL SECTION OF THE CORTEX OF THE
CEREBELLUM, CUT PARALLEL WITH THE LONG AXIS OF A FOLIUM.
A shows three cells of Purkinje, their espalier
systems of dendrites being seen in profile. A
“mossy fibre” enters the granular layer from the
white matter. About a dozen of the granules are
shown, each with four or five dendrites and a
single axon. The axon bifurcates in the molecular
layer, its two branches running for a considerable
distance to left and right along the folium. B
shows the other nervous elements which are found in
the cortex: a cell of Golgi with a ramified axon,
a climbing fibre, a basket-cell, of which the axon
divides into four branches, and a small stellate
cell.]
The grey matter which covers the surface of the cerebellum, its
cortex, is singularly regular in microscopic pattern (Fig. 23). It
is divided into three sheets: superficially, the molecular layer in
which the dendrites of the cells of Purkinje branch; beneath this, the
thin layer in which are situate the cell-bodies of these neurones;
thirdly, the layer of small cells, or granules. Cells of Purkinje and
granules have been already described (p. 303). To these must be added
the stellate, bracketing cells of the molecular layer, the axons of
which divide to form baskets about a number of Purkinje-cells, and the
cells of Golgi of the granular layer. These last are comparatively
large cells, which have thornless dendrites, and axons which branch
repeatedly in the granular layer, without passing into the white matter
which underlies the cortex. Two kinds of nerve-fibre bring impulses
to the cortex: (1) “Mossy” fibres, which bear rosettes of filaments
which distribute impulses to the granules; and (2) “climbing” fibres
or “tendril” fibres, which, passing through the granular layer, cling
like ivy to the trunk and principal boughs of the dendritic processes
of Purkinje-cells. The axons of the cells of Purkinje undoubtedly carry
impulses away from the cortex, but their destination is not certainly
known.
The uniformity of structure of the cerebellum suggests that it “acts
as a whole.” Anatomy gives no warrant for the expectation that work of
different kinds is done by its several lobes. Its simplicity leads one
to hope that its mechanism may some day be understood; but at present
there are so many gaps in our knowledge that it is difficult, perhaps
hardly profitable, to attempt to string together the few anatomical
facts of which we are sure.
By means of tracts of afferent fibres the cerebellum has a very
extensive connection with the grey matter of the cerebro-spinal axis
(including the optic thalamus) into which sensory impulses of all kinds
are poured. Experimental results indicate that the organ distributes
impulses to the whole length of the cerebro-spinal axis, from the level
of the neurones which govern the muscles which move the eyes to its far
hinder end. No nerve-roots enter it. Its afferent fibres are the axons
of cell-bodies which lie in the posterior horns of the grey matter of
the spinal cord and in the corresponding grey matter of the axis of the
brain, especially that part related to the nerve from the semicircular
canals. Another set of afferent fibres lies at the periphery of the
spinal cord, forming one of the best defined of the spinal tracts. It
is also one of the oldest, being found in the same situation in all
vertebrate animals. Its fibres, which are exceptionally large, are
the axons of cells which form a very definite column—the “vesicular
column of Clarke”—on the median side of the posterior horn. Further
than this we cannot go. We are ignorant of the nature of the sensory
impressions collected by the cells of Clarke. The cerebellum also
receives through its middle peduncle the axons of cells which lie in
the pons Varolii on the opposite side; which cells are discharged by
impulses descending from the cortex of the great brain. It is not
improbable that it gives to the great brain as many fibres as it
receives from it.
If we had no experimental evidence as to the part which the cerebellum
plays in the harmonious working of the whole nervous system, we
should infer from its structure and connections that it is somewhat
mechanical, a co-ordinator of the activities of other parts rather
than in itself a functionally independent organ. Pathological and
physiological observations very definitely justify this conclusion.
They show that the cerebellum is not essential to life. It may be
completely destroyed by disease or removed by operation without robbing
the individual of any single function or capacity. Disease of the
cerebellum does not diminish the patient’s sensitiveness to every kind
of stimulus, nor does it deprive him of the use of any single muscle;
but it reduces him to the condition of a person who in gait, but not
in mind, is habitually drunk. When he walks he staggers from side to
side; when he stretches out his hand it trembles. His movements are
jerky; his head shakes, his eyes oscillate; he suffers from a feeling
of giddiness; his speech comes haltingly. Cerebellar ataxia, which is a
rare disease, resembles in many respects the much commoner “locomotor
ataxia” produced by disease of the spinal ganglia and the parts of
the cord connected with posterior roots; but careful analysis of the
symptoms shows that they are due, not to want of the sensations which
guide movements, but to inability to regulate the force of muscular
contractions. A man suffering from locomotor ataxia falls when he
closes his eyes, because, not being able to feel with his feet, he is
dependent upon vision for information as to his attitude. When the
cerebellum is diseased, the patient is no less unsteady with his eyes
open than he is with them closed.
The results of cerebellar disease or injury bring home to us the
fact that a nice adjustment of movements is needed to maintain
equilibrium. A dog from which the cerebellum has been removed retains
all its natural enterprise, all its instincts, all its emotions; but
every action which requires it to maintain its centre of gravity in
an unstable position gives it trouble. Placed in water so that its
body is supported, it swims almost as well as a normal dog. It is,
however, easy to lay too much stress upon the balancing function of the
cerebellum. The disturbance of this function attracts our attention;
yet it is probably but the indirect result of the suppression of
activities of a more widespread character. No animal ventures such
liberties with its centre of gravity as the biped Man accomplishes,
without thinking, every time that he descends a flight of stairs. Yet
the cerebellum of the limbless whale, that lives in a medium which
decentralizes its gravity, so to speak, bears the same proportional
relation to the rest of the nervous system as that of Man. Strangely
enough, it is the only cerebellum in the animal kingdom which so
closely resembles Man’s that it might be passed off as belonging to
a human giant; another reminder of the difficulty of deducing the
functions of the several parts of the organ from a study of their
relative development. What have a man and a whale in common which
determines the identity in form of their cerebella? How has it come
about that two cerebra as widely unlike as a man’s and a whale’s should
be associated with a common form of cerebellum?
If we apply to grey matter the distinction between sensory and motor
nerve-tissue—having no exact terminology, it is difficult to avoid
these metaphorical expressions—the cerebellum is essentially a sensory
development. It grows from the very margin of the infolding groove,
which, when closed, becomes the central canal of the brain and spinal
cord, its elements being marshalled in intimate association with
sensory root-fibres. Its millions of loops formed by the axons of
granules and the collecting processes of Purkinje-cells, are by-paths
which tap the conductors of sensory impulses. From some—those,
for example, which originate in the muscles and tendons, and in
the semicircular canals—more of the impulse is diverted to the
cerebellum, from others less. The organ has no motor functions. It
does not discharge neurones which control skeletal muscles, or plain
muscle, or glands. Yet it influences the passage of impulses through
sensori-motor chains, and apparently its influence is universal. It
regulates tone, reflex action, voluntary action. There is no part of
the nervous system over which its control is not felt. By its action
on the apparatus which binds the infinity of receptors which the body
contains to its muscle-fibres and other effectors, it unifies the
body. The cerebrum, as we shall see, is the organ which unifies the
personality. In the progress of evolution two functions which were
originally combined have, for convenience of concentration, been
divorced. The great brain has been set free from the more mechanical
part of the work. That it can perform the functions of the cerebellum
as well as its own is proved in cases of congenital deficiency of
that organ. In several instances malformation, amounting to a very
considerable reduction in the size of the cerebellum, was not detected
until after death, there being no symptoms of a sufficiently pronounced
character to call attention to it during life.
=The Cerebrum.=—All observations made on the great brain prior to
1870 showed it as absolutely inexcitable. Surgeons and physiologists
agreed that cutting, burning, passing electric currents through its
substance, neither yielded evidence of sensation nor movement of any
part of the body. Concerning its structure little was known beyond the
fact that whereas the grey matter, or cortex, which covers its surface
contains nerve-cells, only fibres are to be found in the white matter
which constitutes the greater part of its bulk. It seemed a hopeless
task to attempt to make anything out of a mass of tissue so uniform
in constitution and so irresponsive to experiment. Removing portions
of it appeared to cause a general dulling of the intellect without
loss of any particular mental quality. Physiologists, therefore, spoke
of the cerebrum as “functioning as a whole.” Phrenologists, having
classified the various phases of mental activity as “faculties,”
discovered “bumps” on the surface of the skull which they correlated
with the possession of the several faculties in a marked degree. They
parcelled out the brain in organs concerned with different kinds
of thought; but their localization of function was anatomically as
baseless as their classification of the various aspects of mind,
viewed as a system of philosophy, was absurd. In 1870 it was announced
that electrical stimulation of certain areas of the cortex of the
cerebrum of an animal under the influence of an anæsthetic, and
therefore incapable of voluntary action, induces definite movements.
Although the surgical applications of this discovery have proved
immensely important, its physiological value, as affording a method of
investigating the functions of the brain, is extremely small. Yet the
discovery gave an impetus to the further study of the cortex, which
has been rewarded with many exact results. By the discovery of its
excitability to electric currents it was proved that the whole cortex
has not exactly the same work to do, or—perhaps this is the safer form
of statement—does not do its work in exactly the same way. As soon as
it was known that it is divisible into areas differing in function,
many methods by which the delimitation of the areas might be attempted
were devised. The converging efforts made during the past forty years
by comparative anatomists, histologists, physiologists, pathologists,
and physicians, have resulted in the acquisition of an accurate, if
very restricted, understanding of the construction and mode of working
of the apparatus of thought. Of some of the new data the psychologist
is able to make use; but so far as the physiologist is concerned, it is
the vehicle of mind which is the subject of study, not its contents.
A new subject has been created since 1870. There is therefore nothing
to be gained, so far as our present purpose is concerned, from the
consideration of views which were current before that date; and since,
as must always occur when a science is rapidly advancing, observations
which logically should have been the first to be made were not thought
of until it became necessary to devise methods of checking results
obtained in other ways, we will consider the various sources of our
information without regard to the chronological order in which they
were opened up.
The cerebral hemisphere contains two large central masses of grey
matter, the nucleus caudatus and the nucleus lenticularis, often
described as a single structure under the name “corpus striatum.” Their
functions are unknown. The nerve-fibres which connect the cerebral
hemispheres with the rest of the central nervous system form two thick
limbs or crura on the under side of the brain. Each crus turns upwards
into its hemisphere, between the nucleus caudatus and optic thalamus
(the latter belongs to the “between-brain”) on the inner side, and
the nucleus lenticularis on the outer. In this passage the compact
crus, which is somewhat flattened, is termed the “internal capsule.”
Immediately above the three grey masses the internal capsule disperses
as a fountain of fibres which go to all parts of the cortex. Mingled
with these radiating fibres are vast numbers of others, proper to the
hemispheres, which run tangentially. Some, crossing the median plane,
as the corpus callosum, bind the two hemispheres together. Others form
tracts which can be followed from one end or pole of the hemisphere to
the other. Groups of fibres, dipping but little below the cortex, unite
nearly adjacent spots or neighbouring convolutions.
The folding of the cortex beneath fissures is due to the necessity
of disposing of a certain bulk of grey matter without increasing its
thickness beyond the proper limit. Since the superficial area of a
sphere varies as the square of its radius, whereas its capacity varies
as the cube, it is possible for a fixed relation to be maintained
between the amount of cortex and the amount of white matter in the
brain, only by the folds increasing in depth as the size of the brain
increases. Fissuring is a response to a mechanical need. This does not
imply, however, that the lines along which it takes place are devoid of
morphological meaning. The similarity in pattern of the convolutions
and fissures in various animals, and the regular progress of their
development in each individual, prove the contrary. If they are not
absolutely trustworthy as boundaries of areas of separate function—and
further evidence will be needed before a decision can be pronounced
upon this disputed question—they are in the main satisfactory as
landmarks.
As the nervous system grows, the axons of its neurones acquire
their fatty (myelin) sheaths in the order in which they come into
functional activity. The passage through them of impulses is the
stimulus which leads to the deposition of fat. The study of the
progress of myelination enabled the anatomist Flechsig to ascertain
the situation within the brain of the tracts of fibres related to
the several senses, and hence the traffic of the areas of the cortex
to which they go. Glistening white streaks appear successively in
the pulpy yellowish-pink substance of the interior of the brain. At
the time of birth all the fibres which enter or leave the cerebral
hemispheres have acquired their myelin sheaths. In the baby’s brain the
sense-organs have established all their connections with the cortex. No
new fibres will appear in the nerves of the eye, the ear, or the other
sense-organs, nor will their end-stations in the cortex be further
multiplied. (The use of the expression “end-stations” is legitimate
so far as sensations are concerned; notwithstanding that all sensory
impulses are retransmitted by neurones in the cerebro-spinal axis.) But
the cortex is very far from having finished its growth. It contains
a large amount of embryonic tissue, which gradually spreads outwards
from the developed areas into the surrounding unoccupied zones. The
taking up of new territory, and the consequent increase in the size
of the brain, is continued into adult life. The study of progressive
myelination enabled Flechsig to divide the cortex into “sensory
centres,” and intervening “association-zones”; although, doubtless, the
difference in function between the portions which receive sensations
direct and the portions in which the products of sensation are worked
up is one of degree, and not of kind.
[Illustration: =Fig. 24.=—VERTICAL SECTIONS OF THE CORTEX OF THE
CEREBRUM—A, OF THE VISUAL SENSORY; B, OF THE VISUAL ASSOCIATION AREA.
Between the two sections are shown the principal
types of cell, at the levels at which they
are severally found: _a_, small pyramid; _b_,
medium-sized pyramid; _c_, large pyramid. The size
of a pyramid is an indication of the distance to
which its axon extends before branching; the longer
its traject, the more widespread, it would seem, is
its terminal arborization. The axon of _c_, one of
the very large pyramids found in this association
area, passes to the front of the cerebrum, where
it breaks up in an association area of the tactual
sense of the hand, or of sensations concerned
with the regulation of gait, or in a centre for
movements of the eyeball. _d_, a tangential cell
of the surface; _e_, a Golgi cell with ramified
axon; _f_, a polymorph cell, with its axon directed
towards the surface. In sensory areas, tangential
fibres and granules are more numerous; in
association areas, small and medium-sized pyramids.]
The structure of the cortex is not quite the same in sensory and
association areas; but it is everywhere so far from showing the
diagrammatic simplicity which characterizes the cortex of the
cerebellum as to make it difficult to summarize the modifications which
distinguish its various regions. To a considerable extent its elements
shade one into the other, differing in size and in orientation rather
than in form. Commonly it is described as divisible into five layers:
(1) A thin superficial layer, containing cells of various forms and
fibres derived from the cells of the deeper strata. Some of the cells
are pluripolar, possessing several axons which run parallel with the
surface. Their destination is unknown. They do not appear to form
baskets like the cells of the molecular layer of the cerebellum. The
dendrites of pyramidal cells extend into this layer. (2) The layer
of small pyramids; cells with a branching apical process, root-like
dendrites from the basal angles of the pyramid, and an axon which sinks
into the white matter. (3) Granules. Carmine or other nuclear stains
show that small cells are present in very large numbers, especially
in the sensory areas; but since they are not, like the granules of
the cerebellum, coloured by the chrome-silver method, their form
and the disposition of their axons are unknown. (4) Large pyramids
exactly similar in form to the small ones. Their apical processes
are very thorny. Their axons give off several collaterals. Pyramids
are the most conspicuous elements in the cortex. Properly speaking,
they do not occur in layers, but are scattered throughout its whole
thickness, although their cell-bodies are not seen in either its most
superficial or its deepest strata. The largest are those of which the
axons either descend into the spinal cord or pass to a very distant
region of the cortex. They are found singly or in small clusters in the
deeper levels. (5) Polymorphous cells, some of them pyramids lying on
their sides, or even directing their axons towards the surface; some
fusiform or irregular cells; some Golgi-cells (p. 340). The axons of
pyramids enter the white matter, and many fibres from the white matter
radiate towards the surface between the pyramids; but the way in which
afferent, sensory fibres are connected with the collecting processes,
dendrites, of the pyramids is not known. We have already referred to
thorns, and to the possible nerve-net (p. 301). Sheets of tangential
fibres also occur in the cortex. A particularly distinct sheet divides
the granules in the visual cortex into two strata. In sections of this
region the sheet of fibres appears as a white line, distinctly visible
without a lens.
The limits of the several areas can be determined by examining the
structure of the cortex; but the individual peculiarities of the
various regions are not so marked as to indicate that they have
different kinds of work to do; if by kinds of work we wish to imply
that one part is “sensory,” another “motor,” a third concerned with
“intellectual processes.” On the contrary, its relative uniformity
shows unmistakably that all parts are engaged in the same work.
Nevertheless, certain broad conclusions can be drawn with regard to
the form of the neurones more immediately concerned with sensation,
with motion—that is to say, with the discharge to the grey matter
of the cerebro-spinal axis of the impulses which call its neurones
into activity—and with the secondary processes, called collectively
“association,” which occur within the cortex. Granules, as everywhere
throughout the nervous system, are receivers and distributors of
sensory impulses; although a study of the cerebral cortex does
not justify the conclusion that they are necessary links in its
sensori-motor arcs. Large pyramids are occupied with the nutrition
of fibres which have a long traject through the system. Hence they
are “motor.” They constitute a marked feature of the area which
is susceptible to stimulation. They occur also in the visual area
and elsewhere. Small pyramids are associational; that is to say,
their axons do not leave the cerebral hemispheres. They distribute
impulses from sensory areas to association-zones, and from one part
of an association-zone to another. The layer of polymorphous cells
is relatively thicker in animals in which the cortex of the brain
exercises less control over action than in animals in which the cortex
is supreme—in a rabbit thicker than in a monkey; in a monkey thicker
than in Man. This layer is therefore said to be concerned with the
lower functions of the cortex, whatever this expression may mean. Since
the relative abundance of small pyramids is a test of the supremacy
of the cortex, we may speak of them vaguely as concerned with its
higher functions. But a surer test of the capacity of the cortex for
the elaboration of the raw materials of thought which sensory nerves
deliver to it is the relative abundance of the tissue which intervenes
between its cells. The number of cell-bodies to be counted in a square
millimetre of a section of a given thickness is smaller in Man than in
a monkey, in a monkey than in a dog, and in a dog than in a rabbit.
A comparison of the brains of various mammals in which particular
sense-organs are either deficient or exceptionally well developed
affords the clearest proof of the localization of sensory areas. This,
if it were possible to make satisfactory measurements, would be by
far the best class of evidence as to the part played by the several
senses in an animal’s mental life. Unfortunately, measurement appears
to be out of the question; but a glance at a rabbit’s brain, placed by
the side of a mole’s, shows that vision is localized in the occipital
region. All marine mammals are destitute of the sense of smell; the
brain of a dog, compared with that of a porpoise or a whale, shows
that the sphenoidal region (_cf._ Fig. 25) is associated with this
sense. The brain of an otter exhibits very clearly the area into which
impulses arising in the nerve-endings of the sensory bristles of the
cheek are poured.
“Nihil est in intellectu quod non prius in sensu fuerit.” The organ
of the intellect is the cortex of the great brain, a sheet of grey
matter which has developed in connection with the various sense-organs.
The cerebral hemisphere of an infant is merely an extension of the
nerve-tissue associated with its sense-organs. Such it remains in a
microcephalous idiot. In the lower animals its capacity of growth after
birth is very small. But in a normal child the inflow of impressions
through sense-organs, the experience acquired regarding itself and its
surroundings, education, whether accidental or directed, causes the
extension of nerve-tissue from the sensory areas into the expansible
intervening zones.
There is still some uncertainty as to the nature of the sensations
received in the excitable area. They may be termed “kinæsthetic”
(sensations connected with movement) without more exact definition.
Some physiologists consider that tactile sensations, as well as the
obscure sensations, originated in the nerve-endings in muscles, around
tendons, or on joint-surfaces, are distributed to the areas, which,
when stimulated, are shown to represent fingers, hand, arm, and other
parts of the body. Others have sought, though with doubtful success,
for a tactile area, independent of the kinæsthetic centres. When first
discovered, these centres were termed “motor,” and still this term
may be retained, on the understanding that it does not imply that the
exchanges which occur in the kinæsthetic centres are of a different
nature to those which take place elsewhere. The region which they
occupy has become the motor area of the cortex because voluntary
movement is possible only under the guidance of sensations of movement.
A sound or a retinal image may prompt the movement; but the part of the
temporal region, or of the occipital region in which the sound-movement
exchange or sight-movement exchange occurs must act through the motor
area by opening kinæsthetic-movement arcs. Destruction of a part of
the kinæsthetic cortex causes in Man and the higher apes permanent
paralysis for the movements directed by the spot destroyed. In lower
animals the definition of the movement centres is vague, and their
removal produces only temporary results. Their mastery over the muscles
is less complete than in the higher apes and Man.
Practically nothing is known with regard to localization of function in
the association-zones, with the exception of the localization of the
centres for words; but this exception is so remarkable as to suggest
that if there were any other faculties, interference with which caused
defects as distinct as those which characterize disorders of speech,
it would be found that the association-zones are made up of definite
centres. As the evidence stands with regard to the broadest continental
divisions, we can merely state that it points, although not very
clearly, to the connection of the frontal zone, the region in front of
the kinæsthetic area, with ideas of personality, of other zones with
ideas of environment. Injury to the frontal region has in certain cases
resulted in the victim’s losing his knowledge of himself, his name, and
his relation to his family. On the other hand, gunshot wounds and other
definite injuries have in a large number of cases destroyed portions
of the cortex behind the forehead without causing any recognizable
intellectual change. It is quite certain that this part of the brain
performs no functions which are of a different, or, as it is often
called, higher order than those of other association-zones. It has been
stated that disease of the zone which intervenes between the visual
and auditory areas is more likely to cause hallucinations, disease of
the frontal zone delusions. A patient fancies in the one case that
he sees things that are not there, or hears voices when no one is
speaking; in the other case he imagines himself a king; but evidence
connecting localized disease with mental derangement is very scanty.
The functional disturbance which causes lunacy is usually of a general
character; or, if local to begin with, it becomes general before the
death of the patient makes possible the examination of his brain.
[Illustration: FIG. 25.—THE SURFACE OF THE LEFT CEREBRAL HEMISPHERE,
CEREBELLUM, AND MEDULLA OBLONGATA.
Sensory areas are enclosed by broken lines; certain
centres in the association-zones are marked by
dots. The sensory area of smell is on the inner
aspect of the brain; so also is the area of vision
which borders the calcarine and retrocalcarine
fissures, and only rarely extends on to the
external surface, as shown in the diagram. The
sensory area of hearing is largely hidden within
the fossa of Sylvius, the opening into which
is indicated by the dark line above it. The
kinæsthetic-sensory areas for the various muscles
of the body occupy the territory between the
dotted line in front and the bottom of the fissure
of Rolando behind. They do not extend on to the
posterior wall of this fissure. It is impossible
at present to define the boundaries of any of the
centres in the association-zones.]
Derangements of =speech= throw a flood of light upon the organization
and manner of working of the association-zones; and, owing to the
accident of the continuation of the line of the carotid artery by
the middle cerebral artery, which supplies the speech centre, there
is no other spot in the cortex so likely to be thrown out of gear. A
little plasma coagulates on one of the cardiac valves, or about an
atheromatous spot in the aorta. Detached by the blood-stream, it is
shot into one of the branches of the middle cerebral artery, which
it plugs, causing apoplexy. A larger or smaller number of muscles on
the opposite side of the body are paralysed. If the plugging occurs
on the left side of the brain, it is accompanied by aphasia; but only
if it occurs on the left side, owing to the fact—perhaps the most
remarkable in connection with the localization of speech—that only on
the left side is the cortex trained to utter words. In course of time
the patient may recover the power of speaking, but not until he has,
with almost as much labour as in childhood, educated the right side to
do the work. There are four speech-centres, quite distinct one from
the other. Near the visual area is the centre for seeing words, or
rather the centre for seeing the meaning of words. If this centre be
diseased, a written word is merely a crooked line. Behind the auditory
area is the centre for recognizing the meaning of words heard. If it
is interfered with, the most endearing or commanding phrases produce
no more impression on the hearer than a bird’s song. In front of the
hand-area—its localization is less certain than that of the other
three—is the centre for writing. In it are associated words heard
or seen, with the movements necessary for the making of letters. In
the centre first referred to, as being the one most often thrown out
of gear, which lies in front of the area for the mouth and throat,
words heard or seen are translated into movements of the parts which
give them sound. No other actions illustrate so clearly the “law of
neural habit.” In the infant’s brain sounds of words are distinguished
from other sounds. They are associated with the objects which they
name. Movements of the mouth and throat, made at first ineffectively,
blunderingly, succeed after a time in securing the thing of which
they sound the name to the child’s satisfaction. Thus, two centres
are gradually established in his mind. Sounds and ideas of things are
associated in the one; words and ideas of the movements necessary to
their pronunciation in the other. Either of the four speech-centres
may be placed out of action without the others suffering. A man may
be able to write without being able to read what he has written. He
may read aloud, although apparently deaf to speech. He may be unable
to write or unable to speak, although understanding what he reads or
hears. Aphasia, when partial, illustrates still further the law of
neural habit. The ability to remember nouns, especially proper names,
is most easily lost. Few are the people who, as age advances, do not
suffer from this failing. Even the names which are most familiar elude
the memory. From one point of view this is strange. Nouns-substantive
are the words first learned. Of all words they have the most definite
objective association. But it is just their definiteness which makes
them difficult of approach when the apparatus of mind is working badly.
There are so few paths by which they can be reached. Their mental
associations are limited. A patient who is recovering from the effects
of a lesion which has rendered him partially aphasic may be able to
recall adjectives when he cannot recall nouns. He may say, “Give me
the black,” when he wants ink, and “Give me the white,” when he needs
paper. Or he may retain control of verbs. “Where is the—— what I put
on—what I think with?” may be the circumlocution for hat.
Psychologists explain the voluntary production of a movement as the
setting flowing of a sensori-motor current. Everyone agrees that it
is impossible to think of the impulses which produce movement as
originating without sensory antecedents. Hence psychologists picture
the nerve-current as originating on the sensory side. Kinæsthetic
images of the sensations which will result from the movement are
described as being called up in the mind by the agitation of the
part of the brain which, by association, is linked with the neurones
which discharge impulses to the appropriate spots in the grey
matter of the spinal cord. The idea of movement flows over to the
muscles. But this conception of the relation of mind to body assumes
too much. It postulates an existent mind in which the images of
movement-sensations—the memories, that is to say, of the sensations
which previously accompanied movement—are stored. The study of the
apparatus of mind does not warrant this assumption of an existent
mind. It finds nothing in the nervous system but apparatus. There is
no mind existent in the brain during sleep. It would appear to be
sufficient to describe the origination of a voluntary movement as
the opening of the channels which convey the afferent impulses which
are ceaselessly pouring into grey matter from nerve-endings in and
about muscles into efferent channels. Our conception of the number
of sensations which reach the realm of consciousness is ludicrously
restricted by our inability to pay attention to more than one sensation
at a time—a restriction, it is needless to remark, which is imperative
in the interests of consistency of behaviour. Two personalities paying
attention to different sequences of sensations would give incompatible
orders. One would command the muscles to cause the body to recline;
the other would direct them to make it stand up. From myriads of
sense-organs impulses are continuously rippling through the cortex
of the brain. The term “impulse” is too heavily weighted by its
association with the idea of currents which are strong enough to prove
effective without the intervention of consciousness; but no other is
available. They ring the bell of consciousness, however little may be
the attention which their summons secures. Attention cannot be directed
to two things simultaneously. It moves, as it were, on a succession of
points. On some it rests longer than on others. They make an impression
which can be recalled; the rest being passed by so rapidly that
they are not remembered, it is as if they had never been perceived.
They blend, as a succession of moving lights blend, in producing a
background to consciousness. Not recognizing their separateness, we
interpret them as fused. A good deal of misleading metaphor has been
used, as it seems to the writer, in accounting for the effect upon
the mind of impressions which make but a weak demand upon attention.
They are spoken of as “marginal” perceptions, from the analogy of
the ineffectiveness of impulses generated at the periphery of the
retina, as compared with those which give rise to direct vision. A
“subconscious,” or even “unconscious,” self is evoked. The self cannot
be less than conscious. Self is the passage of attention from sensation
to sensation. Its relation to the not-self is temporal, not spatial.
Every sensation which is called up into consciousness, though it
occupy attention for the shortest possible time, tends to give rise to
movement—is, indeed, in its very nature an impulse flowing through
a sensori-motor arc. The circuit for the voluntary execution of a
movement is represented as flowing through kinæsthetic-movement arcs.
This may be necessary for volitional actions, but it is not essential
for reflex actions. A spinal frog will remove an irritant from its back
with its hind-leg, after the roots of all the afferent nerves of the
hind-leg have been cut. In this case the reflex is direct, from injured
skin to muscles of the leg. It is not double—muscular sensations from
the leg, liberated into efferent leg-muscle-nerves by skin-sensations
originated simultaneously in a part anterior to the segments in which
the roots have been cut.
The unit of sensation to which attention can be directed has yet to
be defined. Like sensations—sensations which are correlated in
experience, that is to say—seem to fuse in consciousness. A sequence
of similar sensations appeals to attention. Unlike sensations interfere
one with another. The apparent fusion is not a composite neural effect
which consciousness views as a single unit. Not even identical images
simultaneously formed on the two retinæ produce a superimposed effect
upon a particular spot in the brain. Different brain-spots receive the
two separate images which the mind views as one. This raises a doubt
as to whether perceptions are, properly speaking, fused. It suggests
that they are separate points upon which attention rests in rapid
succession; but such a hypothesis does not preclude the conception
of the production of a composite sensation by impulses coming
simultaneously from the same sense-organ—_e.g._, a unified neural
effect as the result of several musical tones.
Every neural agitation which attracts attention has an effect upon
the growth of the nerve-strands in which it occurs. =Memory= is not
an existent. It is the repassage of the same strands. There is no
such _thing_ as memory. It is the neural apparatus which responds in
a similar way to a similar agitation. It is difficult to speak of
association and neural habit, the phenomena upon which not only all
mental life, but all co-ordinated activities, are based, without using
such expressions as “the broadening of the path” or “the thickening
of the conductor” by the impulses which pass through it. Apparently
these analogies may with safety be pressed curiously far. Chaotic
response to stimulation is unknown. Thanks to the nervous system,
action exhibits an ordered relation to stimulation. This relation is
determined by education, giving the term a connotation wide enough
to cover all experience. Nerve-tissue adjusts itself to experience;
and since the nerve-matter which takes the pattern is not labile,
the process of organization is consecutive and the result permanent.
One pattern is not destroyed as another is impressed. Hence temporal
associations are formed. What has been thought once will be thought
again, if the circumstances in which it was thought recur. What has
been done once will be done again under the influence of a similar
sequence of stimuli. The conductors are widened every time that they
are used. But, so far as concerns the mind, a reversed influence comes
into play. The wider the conductor, the less appeal to attention is
made by the impulses which pass through it. It is as if currents which
have to overcome resistance in a narrow path acquire a higher potential
than those which find an open road. And since the making of the road
depends upon attention, the limit of broadening is reached when a
volitional act becomes a habit. The first time that a piece of music is
played consciousness is alert. Marks on the page and movements of the
fingers are felt intensely. With each repetition the need for attention
subsides.
A skilled movement is impossible in the absence of guiding sensations.
I decide to button my coat. Sensation-paths from the muscles of the
forearm are opened into motor paths extending from the large pyramids
in the arm-centres of the kinæsthetic cortex. But it is not sufficient
that the action be started: it must be guided by the sensations which
movement produces. If my fingers are numb with cold, I cannot button
the coat. The muscles which move the fingers are warm enough beneath
the sleeve, but my attempts to will them to move are as futile as they
would be if the muscles belonged to some other person. The will has
no power over the muscles. It is essential that the sensations which
accompany the act of buttoning the coat flow through the same paths as
hitherto in the cortex of the brain. Flowing through the same paths,
they produce the same effect in consciousness, the same perceptions. In
ordinary parlance, one cannot perform any act unless one can remember
what it felt like to perform it on a previous occasion. It is almost
as sound physiology to describe the voluntary action of fastening a
button as commencing in the skin of the fingers as to describe is as
commencing in the brain. The act is due to the direction of attention
to impulses which flow from muscle to muscle, and from skin to muscle.
All skill in the use of muscles is acquired by the method of trial and
error. Familiar movements are tried, combined, modified with a view to
the production of a new result. A man accustomed to striking with the
right hand forwards endeavours to swing a golf-club with the left hand
backwards. For a long time the result is anything but a success. At
length the head of the club takes the right curve. It not only hits the
ball with its centre, but it carries it through in the right line. The
ball travels 120 yards or so towards the green. In golfing terminology,
a successful drive is always “an awful fluke”; but the fluke once
accomplished, nothing is easier for the golfer than to drive equally
well on all succeeding occasions. He need merely remember exactly what
it felt like to give the club a perfect swing, and exclude all other
sensations while he is passing these memories through his sensori-motor
arcs!
The fact that we can deliberately improve an action, fitting it to
the attainment of the object of desire, by suppressing wrong and
emphasizing right sensations, shows how large a part consciousness
plays in the affairs of the nervous system. This brings us to
the frontier of physiology. At this boundary the authority of
the physiologist ends. He cannot define consciousness; he cannot
investigate it. Yet he naturally asks whether the machine which he is
investigating is a machine and nothing more. When the possibilities of
reflex action were first recognized, thought tended to dethrone feeling
and Will in favour of automatism. If the actions of a spinal frog
exhibit so distinct a purposive character, why, it was asked, should
we assume that the frog with a brain is anything more than a reflex
machine? Light, heat, sound are playing upon its sense-organs; surely
these stimuli suffice to set going all the sensori-motor currents which
lead to the various movements which in their totality constitute the
frog’s behaviour! And why assign to a mammal a self-directing authority
which we deny to a frog? The increased complexity of its behaviour is
more than accounted for by the greater variety of its nervous arcs.
All animals, it was argued, including Man, are reflex machines. Their
thoughts and actions are the effects of the play upon their nervous
systems of forces from the outer world. Each inherits a nervous system
of a certain pattern. Its individual development is conditioned by
the sensations which pass through it. The sensations are impressed by
the environment. Therefore the individual is a puppet, his activities
the dance of circumstance. Consciousness is an “epiphenomenon.” Few
physiologists or students of animal behaviour take this material view
of life at the present day. The fact that it leads inevitably to the
conclusion that consciousness is an “epiphenomenon” (Huxley’s term) is
its _reductio ad absurdum_. It is not in harmony with the economy of
Nature that an animal should be endowed with the capacity of feeling
pain and pleasure, if such endowment is useless to it. It can be useful
only by directing activity towards the attainment of pleasure and
the avoidance of pain. This admitted, the mechanical theory falls to
the ground. There is an “It” which feels, selects feelings, chooses
those which have a pleasant tone, wills to perform the acts by which
they are attained. It follows that the value of consciousness lies in
the prerogative which it confers of adapting action, within certain
limits, to circumstance. An animal succeeds in life in proportion as
the nervous system which it inherits reacts satisfactorily to its
environment. A chick which, after being hatched in an incubator, has
been isolated for twelve hours without food, seizes a grain of corn the
instant that it sees it. Its brain contains ready-made sensori-motor
arcs connecting the spot in its cortex in which the visual impression
of the grain is perceived and the motor neurones which control the
pecking muscles. A sheep-dog is quickly broken to sheep, because its
ancestors have been selected by mankind from amongst dogs that readily
adapted themselves to this work. The breeder has selected a pattern of
brain with the same success with which, when appearance is the only
desideratum, he selects a pattern of coat. Beavers set to work at
constructing a dam at the only spot in a valley at which it is possible
to create an artificial lake, because for countless ages Nature has
ruled out the animals which constructed their dams in unsuitable
places. Man also inherits a brain-pattern; but, not being required to
shift for himself soon after birth, he goes through a long period of
infancy and tutelage, during which, by force of circumstance and his
own Will, the pattern is elaborated. His supreme success is due to his
capacity for adapting means to ends. He inherits very few instincts.
Except as regards organic functions, his spinal cord is subservient in
almost all respects to his brain. Most of the actions of an animal are
instinctive—a word which has been sadly misapplied. Its connotation
is negative rather than positive. Owing to the marked pattern of its
brain, an animal finds it difficult to avoid acting in a particular
way. As the nights grow longer and its hours for feeding are curtailed,
a swallow is impelled by its instinct to go South. It makes the same
use of its sensations during its migration, and is as completely
dependent upon them for its guidance as a man would be. The lower we
descend the scale, the more inevitable do an animal’s movements become;
but there can be no doubt but that consciousness is of value to an
animal, as to Man, in that it gives to its individuality the capacity,
within such limits as Nature has selected, of resisting or modifying
its ancestral instincts when they are not absolutely appropriate to the
occasion.
Sentience implies personality. “No system of philosophy can extrude
the ego.” The difference between the performance of the animal machine
as a physiologist studies it, and its behaviour when under the control
of its own driver, is the difference between reflex action and choice.
The ego interacts with physical forces. It does not come within the
province of the physiologist to explain the source of the force which
interferes with force. He finds no trace of it on either credit or
debit side when making up the body’s accounts. He is unable to enter,
“Item, to the development of consciousness ... so much.” He can form
no conception of this immaterial manifestent which hovers over the
infinitely numerous sensori-motor exchanges which are always occurring
in the cortex of the brain, giving to a particular group of agitations,
now here, now there, a special quality; but the manifestent is needed
to account for the potency of the reinforced agitations which enables
them to take possession of the nerve-paths by which muscles are reached.
It is for the psychologist to define the application of the terms
“consciousness,” “attention,” “will.” He cannot define the attributes
of the ego which these terms connote. The moralist must show the way
in which they determine, or should determine, conduct. Yet within the
plain limits of physiology, attention, using the word in its every-day
sense, modifies the responses of the nervous system in a degree which
cannot escape observation. It is astonishing to anyone accustomed to
hospital surgery (although even in this field singular exceptions are
met with) to see the grave operations which a veterinary surgeon may
perform, without the animal showing any evidence of pain, provided its
apprehension has not been aroused and its attention directed to what
is being done. A horse standing in front of a crib of oats, untied,
will hardly whisk its tail while the surgeon is making a great wound
in its flesh, and sawing off a bony excrescence. The knife does not
come within the experience of a horse. It has no anticipations, and
its skin, intensely sensitive to the tickling of a fly or the smart
of a whip, is relatively insensitive to a cut. An eminent surgeon of
the last generation (the writer, as a student, “dressed” for him in
his old age) was in the habit, having arranged that his patient could
not see what he was doing, of performing operations of a very painful
nature whilst assuring his patient, “I am merely making a thorough
examination, in order that I may be perfectly certain of the cuts that
I shall have to make to-morrow in the operating-theatre when you are
under chloroform.” We are not concerned with the ethics of his method;
but the assurance, “Now that’s all over; you will never need to have
that operation performed again,” saved many a sufferer from a night of
apprehension and a miserable “coming round.”
It was stated, during the South African War, that at Ladysmith the
bearer of a critical despatch, who was struck in the palm of the hand
by a bullet which traversed the whole length of his forearm, did
not discover that he was wounded until he saw the dripping blood,
after his errand was successfully accomplished. To deliberately cut
oneself with a razor is most painful, yet shaving in the morning, with
thoughts concentrated on the doings of the day, it is often the sight
of blood which directs attention to the fact that the skin is severed.
Of all evidences of self, the power of paying attention is the most
noteworthy. We can direct attention to certain sensations, which then
become perceptions, and we can deliberately ignore others, within
certain restricted limits.
The control of the nervous apparatus by the self is a truth which no
student of the physiology of human beings can ignore. Isolated from its
relation to all other scientific truths, it has been made the basis of
a nescience which, although positively merely foolish, is, negatively,
harmful—yet a form of folly which answers well to the needs of persons
of a certain category.
It may be objected that the picture of the relation of mind to brain
which is here presented—the one, activity, motion, the other a
labyrinth of conducting paths—makes all mental phenomena entirely
dependent upon current sensations. No results could happen if the
sensations were not there. It affords no ground for the explanation of
=mental images=, =hallucinations=, =dreams=. A few lines may be spared
to show that this objection does not hold. We cannot attempt to explain
the conscious control of thought. It is a part of the impenetrable
mystery to which we have just referred. But, granted that it obtains,
the direction by the ego of afferent nerve-currents through the same
strands which formerly vibrated to sensations which drew a picture, and
hence the revival of its image, is no more incomprehensible than the
liberation of afferent impulses from muscles into efferent channels.
Brain-chains are composed of many links. Their interconnection is
illimitable. When I recall the appearance of the house in which I lived
as a child, I throw into the chain impulses (from somewhere) which
traverse the final links, where passage implies consciousness. At the
edge of the lace-work of linked threads the impulses light up a pattern
which childhood’s experience worked into the apparatus of thought.
If we were to admire the perfection of any special aspect of the
brain’s functioning, the rarity of hallucinations might give us cause
for wonder. That impulses so seldom leave their own paths is more
astonishing than that occasionally, when the brain is excited and its
nutritive conditions deranged, the impulses which the ego can direct
into channels where they revive an image should sometimes, and with
far greater force, make their own way down well-worn paths, lighting
up a picture which deceives the ego. Dreams, by contrast, throw up
in a strong light the part played by attention in intelligent life.
The capacity for alertness is due to the favouring of one set of
impulses by suppressing others. The favoured impulses hold the road.
Concentration of attention is keeping thought to one line by resisting
all temptation to wander into by-paths. The waking condition is the
state in which all nerve-ways are closed, with the exception of those
which consciousness is using. The more severe the closure, the more
vivid is consciousness. In sleep all paths are open. In none is the
potential acquired by impulses in the process of overcoming resistance
high enough to evoke consciousness. A burst of impulses ascends from
the stomach, set a-flowing by undigested fragments of salmon and
cucumber, or mounts from the arm on which the sleeper has been lying
until its circulation has been arrested. They reverberate through the
open corridors of the brain. If they are sufficiently noisy to awaken
the sleeper, he, detecting them in this path and in that, supposes
them to be on the same errands as the impulses which commonly pass
thus. If dreams are analysed, it will be found that, although the
combinations of impressions may be uncommon and extremely bizarre, the
impressions are selected from the most familiar. The images of which
the dream is compounded, which may have lost all normal relations and
may have assumed impossible proportions, are those which the mind
most frequently conjures up. In the large majority of instances some
happening of the day preceding can be recognized as the prompting
cause. A remembered dream is the photograph taken by consciousness
of the sensations which have bombarded it into activity. Especially
if due to impulses originated by visceral discomfort, the dream may
have an unpleasant tone. This may take various forms, but the emotion
most commonly aroused is fear. The objects visualized may have
preposterous dimensions, or they may be not sufficiently distinct for
recognition—elusive imps; but most commonly distress is caused by
the want of harmony of sensations, due to the absence of kinæsthetic
elements. A man is lying on the railway-line; a train is approaching
with increasing speed; he cannot get up. He is in the pulpit, but
cannot speak. Dreams thus confirm the view set forth above as to the
cause of volitional action. Ability to perform an act depends upon the
flow through the kinæsthetic centres of the brain of impulses generated
in the muscles by which the act is to be, or is being, performed.
Kinæsthetic sensations do not under any circumstances play the same
part in mental life as sensations from the skin, the eye, or the ear;
when the body is passive in bed they are not flowing into the cortex.
The dream-photograph shows elements demanding movement, but affords no
evidence that movement is in progress.