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THE MECHANISM OF NERVOUS ACTION
THE MECHANISM OF
Nervous Action
ELECTRICAL STUDIES OF THE NEURONE
By
E. D. Adrian
M.D., D.SC., F.R.S., F.R.C.P.
Foul erton Research Professor oj the Royal Society
Philadelphia
U N I V E R S I T Y OF PENNSYLVANIA PRESS
Copyright 1932
UNIVERSITY OF PENNSYLVANIA PRESS
New Edition 1 9 5 9
Published in Great Britain, India, and Pakistan

by the Oxford University Press
London, Bombay, and Karachi
Printed and Bound in the United State* of America by

Book Craftamen Associate* Inc., New York
THE MECHANISM OF NERVOUS ACTION
PREFACE
On October 5th, 1 9 1 6 , Keith Lucas, my director of
studies at Trinity, was killed in an aeroplane accident on
Salisbury Plain. A biologist with the training of an
engineer, he had succeeded Gotch as the foremost authority
in Great Britain on the physiology of nervous conduction,
and in 1914 he was planning an attack on the problems of
the central nervous system. A few weeks before his death
he talked to me of the great possibilities which might lie in
the use of the thermionic valve for amplifying nerve action
currents. His forecast was justified in a few years by the
work of Forbes and Gasser: and these lectures deal with
the use of the method at Cambridge for the study of the
units of the nervous system.
Had Keith Lucas lived, it is certain that the present stage
would have been passed long ago; it is even possible that
the method would have already served its turn. As it is,
we are still collecting information and have scarcely begun
to build up hypotheses to explain what we have found.
Fortunately the main results can be seen at a glance from
photographic records which need very little discussion.
B y the aid of these I have tried to illustrate the chief lines
of work which have been carried out by a particular tech-
nique. I have not tried to review a particular field of
physiology—the list of references will be enough to show
how little I have said of any other line of enquiry. But this
is by no means a personal record, for much of the work
described in it has been due to my colleagues, particularly
to Β. H. C. Matthews, who has done so much to improve
the technique, and has turned his instruments to such good
account in his own work; and to Professor D. W. Bronk,
with whom I have spent so many strenuous and exciting
days in the laboratory and elsewhere. His kindness in
vii
MECHANISM OF NERVOUS ACTION
inviting me to give the second course of Eldridge Reeves

Johnson lectures is very poorly repaid in these pages, but
it would take a great deal to repay the debt which I owe to
him and to my friends at the University of Pennsylvania.
I have to thank the Royal Society and the Physiological
Society for permission to reproduce some of the figures
which have appeared in their publications.
E. D. A.
March 26, IQ32.
viii
CONTENTS
Page
PREFACE vii
I. THE DEVELOPMENT or ELECTROPHYSIOLOGY

Recording Instruments 2
Amplification of Electric Changes 5
T h e Activity of Peripheral Nerves 8
T h e Activity of Single Nerve Fibres 10
T h e Nature of the Nervous Messages 14
Gradation of Activity 16
T h e Membrane Hypothesis 19
II. THE ACTION OF T H E SENSE ORGANS
Adaptation 23
T h e Production of the Rhythmic Discharge. . . 27
Depolarisation in the Sense Organ 29
T h e Injury Discharge 30
Differences in the Sensory P a t h w a y : Touch and
Pressure 34
Wever and Bray's Work on the Auditory Nerve 37
III. PAIN
T h e Tactile Endings 43
Protopathic and Epicritic Fibres 44
Slow Impulses in the Nerves of the Frog 50
T h e Nerve Fibres Concerned with Pain 53
Specific Nerve Impulses 56
IV. DISCHARGES IN MOTOR NERVE FIBRES
T h e Motor Units 61
T h e Electromyogram 64
T h e Response of Single Units 65
Sympathetic Discharges 73
V. THE ACTIVITY OF N E R V E CELLS
Respiration—The Automatic R h y t h m 78
ix
Potential Changes in the Brain Stem 81

Potential Changes in Insect Ganglia 83
Periodic Discharges in Injured Nerve Fibres. . 86
The Effect of Potential Gradients 88
Synchronous Activity 89
REFERENCES 95
INDEX 101
χ
Chapter I
THE DEVELOPMENT OF ELECTRO-

PHYSIOLOGY
N London we have just celebrated the hundredth anni-

I versary of Faraday's discovery of electromagnetic in-
duction, thanks to which we have every day some new
comfort and some new machine to supersede the human
worker. No one would blame Faraday because our social
organisation cannot always keep pace with the progress of
electrical engineering, but his discovery came at a time
when the science of electrodynamics was well under way,
and if we are really to date the present era from any one
experiment a good case could be made for an observation
in medical physics in the year 1786. Galvani of Bologna
was investigating the effects of static and atmospheric
electricity on the muscles. He had prepared the bod" of a
frog and had fixed a brass hook in the vertebral canal
intending to hang it up on the terrace outside his house.
He placed the frog on an iron plate so that the hook touched
the iron, and when it did so the body moved. " En motus
in rana spontanei, varii, haud infrequentes!" and when the
movements ceased they could be revived by pressing the
hook more firmly against the iron. Galvani had produced
an electric current and had the wit to realise that he had
done so.
It was some time before the full meaning of this observa-
tion was understood, for the role of the frog's muscles was
uncertain. Galvani had made other experiments which
led him to believe that a muscle could generate electricity in
the absence of metals, and it was known that a fish, the
torpedo, could give a severe electric shock. For a few
1
years, until 1792, it seemed that Galvani's phenomena were

due to a form of electricity which was peculiar to living
tissues. Enthusiasts held that this 'Animal Electricity'
was the true life-force, that it would explain movement and
sensation and lead to the cure of all kinds of nervous
disease. Then Volta pricked the physiologists' bubble by
showing that when the two metals were used, the frog
acted merely as indicator and a source of moisture. The
electric current occurred in inanimate systems, life was
not an essential ingredient, and the science of electro-
dynamics became a branch of physics and not of medicine
or biology.
But Galvani's experiments and the controversy with
Volta laid the foundation of a considerable branch of
medical physics, for attention was focussed on the frog's
muscle as well as on the nature of the electric current.
After much discussion it became clear that Galvani was
right in supposing that muscles could produce electricity on
their own account, and before long the two salient facts of
electrophysiology had emerged, namely that a current will
make a muscle contract and that a contracting muscle
develops a current of its own. The two facts are true
not only of the activity of muscle, but of nerve as well.
Electric changes take place whenever a nerve fibre carries
a message, and it is this association of electric change with
nervous activity which forms the main topic we have to
discuss. Owing to recent developments in technique we
can now record a great deal of what takes place electrically
in the nerves, and we shall have to consider how far these
records can be made to reveal the working of the nervous
system.
Recording Instruments
The history of electrophysiology has been decided by the
history of electric recording instruments. Progress was
very rapid at first. In spite of Voita's denial of its special
character, the study of animal electricity seemed the
2
DEVELOPMENT OF ELECTROPHYSIOLOGY
shortest cut to an understanding of life. I t attracted men

of the stamp of du Bois R e y m o n d , Pflüger, and Helmholtz,
and by 1848 du Bois R e y m o n d had published two large
volumes on the subject, and the use of electric currents
for stimulating-—producing the active state—had become a
general method of research. I t would be hard to think
of any other method which has done so much to show us
how the body works, for it gives us a means of throwing a
muscle or a nerve into activity at will by an agency which
does no damage and can be precisely controlled. B u t the
chief interest centred round the converse effect, the produc-
tion of currents by active cells.
Here there were great technical difficulties. T h e currents
are small and as they follow every change of activity they
m a y fluctuate very rapidly; the early mirror galvanometers
could detect them but could not show how they changed
from moment to moment; indirect evidence had to be
sought and lively controversies took place. T h e next stage
came when Bernstein devised an ingenious analysing instru-
ment, the rheotome, which made it possible to follow out
the true form of the potential change which occurs when a
muscle contracts as the result of an electric shock. T h e
events following stimulation in muscle and nerve were
explored and evidence accumulated to show that the electric
response was a true index of the active state and could not
be dissociated from it. T h e capillary electrometer in a
form especially modified for physiological work confirmed
Bernstein's results. It was now possible to record the brief
potential change in a stimulated nerve trunk, to measure
its rate of conduction, and to compare it with that of the
w a v e of activity. Then in 1903 came Einthoven's string
galvanometer, an instrument far more sensitive than the
capillary and yet rapid enough to follow the potential
change in muscle. T h e knowledge which had seemed so
academic was at once put to practical use in medicine, for
3
with the string galvanometer it was possible to record the

electric currents produced by the heart-beat in man.
The use of the electrocardiogram for analysing the
working of the heart is a good illustration of the electrical
method in general. When any part of the heart muscle
becomes active, a potential gradient is established between
the active and inactive regions, and as the wave of activity
spreads, the gradients change their position. Consequently
the potential fluctuations between any two fixed points on
the heart muscle are determined by the route taken by the
wave of activity and by its rate of travel in different regions.
T o record these changes there is no need to expose the heart.
I t is conveniently placed in the thorax with the air-filled
lungs on either side, so that by making electrical connection
with the arm and leg of the subject we are in effect leading
from the base and apex of the heart. Thus the electro-
cardiogram shows at a glance how the heart is behaving,
although we do not directly observe the contraction of the
muscle. Now in a nerve we cannot directly observe the
change from rest to activity, because no visible change
takes place; but when the activity is produced by electric
stimulation there is the same kind of potential change as
in the heart muscle and we can use this in the same way
to explore the working of the fibre. If it is true then that
nerve activity, however produced, is always associated
with an electric change, it should be possible, given instru-
ments rapid and sensitive enough, to analyse the normal
working of the muscles and nerves as we can analyse the
working of the heart.
In the special field of the heart muscle the string galva-
nometer added an entirely new chapter to physiology, but
in the more general field its increased sensitivity was not
great enough to help much. It showed, however, that the
electric changes in nerves and muscles were not produced
only when artificial stimulation was used. When every
fibre in a nerve is thrown into action by an electric shock,
4
D E V E L O P M E N T OF ELECTROPHYSIOLOGY
t h e p o t e n t i a l c h a n g e is m u c h larger t h a n when the u n i t s

a c t i n d e p e n d e n t l y ; so the earlier work, with less sensitive
i n s t r u m e n t s , h a d dealt mainly with the effects of electric
s t i m u l a t i o n . W i t h the s t r i n g g a l v a n o m e t e r it was pos-
sible in selected cases to record electric changes in nerve
d u r i n g reflex a c t i v i t y . I t was rarely possible to say more
t h a n t h a t r a p i d p o t e n t i a l changes were t a k i n g place, b u t
this was e n o u g h to show w h a t m i g h t be expected with an
even m o r e sensitive recording system.
T h i s t a k e s us to t h e period immediately before t h e war
when K e i t h L u c a s was investigating t h e nervous impulse
with t h e capillary electrometer and Piper was using t h e
string g a l v a n o m e t e r to record muscle action c u r r e n t s d u r i n g
v o l u n t a r y c o n t r a c t i o n . B o t h i n s t r u m e n t s were still giving
valuable results, b u t t h e y were results of t h e kind needed
to consolidate a position already won, a n d there was n o t
m u c h h o p e of f u r t h e r a d v a n c e w i t h o u t a fresh d e v e l o p m e n t
on t h e i n s t r u m e n t a l side. F o r t u n a t e l y for us t h e fresh
d e v e l o p m e n t has now a p p e a r e d . T h e a d v e n t of triode
valve, or v a c u u m t u b e amplification has so altered t h e
whole position t h a t we can c o m p a r e ourselves to a micro-
scope worker who has been given a new objective with a
resolving power a t h o u s a n d times greater t h a n a n y t h i n g he
has h a d before. W e h a v e only to focus our i n s t r u m e n t on
t h e field t o find s o m e t h i n g new a n d interesting.
Amplification of Electric Changes

T h e v a c u u m t u b e amplifier is a device which will m a g n i f y
small changes of potential of a n y form. T h e o u t p u t change
is, within limits, a t r u e copy of t h e i n p u t , a n d as it is
d e r i v e d f r o m t h e batteries or mains which feed the amplifier,
t h e p o w e r available for driving t h e recording i n s t r u m e n t
m a y be m a d e very large indeed. T h e result of this is t h a t
t h e sensitivity of the recording i n s t r u m e n t has ceased to
be an i m p o r t a n t consideration, a n d our choice can be
d e t e r m i n e d entirely by t h e r a p i d i t y of the moving system.
5
For some experiments the ideal instrument is the inertialess

cathode ray oscillograph, but the moving iron oscillograph
designed by Matthews (1928) is rapid enough for most
purposes, and there are various instruments of the moving
coil or string type with much the same range. Another
advantage which comes from the increased power at our
disposal is that we are not restricted to optical recording
systems. The shifting beam of light from an oscillograph
can be photographed on a moving strip of film or viewed
with a revolving mirror, but we can often learn something
more if the amplified potential changes are made audible as
well as visible, and this can be done, if the changes are
rapid enough, by leading them into telephones or a loud
speaker.
The amount of amplification which can be obtained
nowadays is far greater than we are ever likely to need in
physiological work. For instance, there would be no
particular difficulty in demonstrating the potential change
in a frog's nerve fibre as an audible signal in the course of
a radio lecture. The power available in the input circuit
would be of the order of i o - M watts, and the transmitting
station might radiate 50 kilowatts or more. But there is a
definite limit at the other end of the scale. We can increase
the magnification of a microscope as much as we like by
using more and more powerful eyepieces, but the resolving
power is still determined by the objective. In the same
way the resolving power of an amplifier, the smallest
potential change which can be detected, depends on the
working of the first tube—the magnitude of the output
change is immaterial. At present we can detect an input
change of one or two microvolts in a circuit of high resist-
ance like a small nerve trunk, but we cannot deal with
anything less than this because fluctuations of half a
microvolt or so are constantly occurring quite apart from
any physiological effects. They may be due to the amplifier
itself, e.g., to unsteady emission in the first tube, and if so
6
there is some hope of reducing them, but there must

always be an irreducible minimum of disturbance due to
the thermal agitation of the electrons in the input circuit,
and it may be that we have already reached the limit set
by this. For some purposes greater sensitivity would be
an advantage, but we may have to remain content with
the great sensitivity we have already.
It will be seen that we have a method of detecting and
recording small electrical changes which is far in advance
of anything possible without amplification. We can record
far smaller changes and we can record very rapid changes
with far greater accuracy than before. I t would be a
sad confession of failure if with these new resources we had
been able to learn nothing fresh about the working of the
nervous system, but in fact they have already given us
direct evidence on many points and the field now open is
so large t h a t it will be a long time before the method has
exhausted its usefulness.
The work with which I shall deal is mainly concerned

with the behaviour of the units which make up the nervous
system. The units, the nerve cells and the threads of
protoplasm which run out from the cell body, are built
up into a complex central mass, the brain and spinal cord,
and a series of nerve trunks connecting the central nervous
system with the periphery. The nerve trunks have a
relatively simple structure, for they consist of bundles of
nerve fibres, grouped together for anatomical convenience
but each acting as an independent conducting unit. Now
it is best to begin with the simplest case and from the
point of view of electrical analysis it is much easier to
decide what is going on in the nerves than in the central
nervous system. All manner of potential changes can be
recorded by placing electrodes in contact with the brain
and cord, but it is often very hard to decide what structures
are responsible for them. No doubt these difficulties will
7
M E C H A N I S M OF NERVOUS ACTION
be overcome before long—some methods are already

available—but meanwhile the most definite results have
come from the peripheral nerves where no such difficulties
arise.
The Activity of Peripheral Nerves
A record of the electrical activity of a sensory nerve is
shown in Fig. i. The nerve is one of the small cutaneous
trunks in the cat's hind foot. The animal has been
F i c . ι . Electric changes in nerve from cat's toe joint on flexing the toe slowly,
rapidly and still more rapidly (Adrian and Umrath, 1929). Record made with
Matthews oscillograph and amplifier. T h e potential changes are of the order of
15 microvolts. The slow rise of the base line in the lowest record is due to a
movement of the nerve on the electrodes; it has no physiological significance.
decapitated under an anaesthetic and the body is kept

alive by artificial respiration. The nerve has been cut
through near the ankle, dissected out for an inch or so,
and placed on two electrodes which lead to an amplifier
and a recording system. All the terminal branches of the
nerve have been cut except one very small twig which
leads to a group of sense organs in the neighbourhood of
the toe joint; these sense organs have not been interfered
with in any way and they can be stimulated, as they
would be in the intact animal, by bending the toe. The
8
records show the potential changes which occur between

the two electrodes when the toe is bent slowly or rapidly.
T h e beam of light from the mirror of a M a t t h e w s oscillo-
graph is thrown on to a moving strip of bromide paper so
that any change of potential is recorded as a movement up
or down of the dividing line between light and shadow.
As long as the toe is at rest the line remains horizontal
apart from the slight unsteadiness always present with a
sensitive amplifying system; when the toe is bent a series
of rapid potential changes take place in the nerve, each
change consisting of a sudden deflection first above and
then below the zero line. T h e deflections are all of much
the same f o r m ; they v a r y in size over the same range in
all three records, but they are more closely crowded when
the toe is bent more rapidly.
There is no doubt about the interpretation of these
potential changes, for they are of the same form as the
much larger changes which are produced when a motor
nerve is stimulated electrically. T h e stimulus then sets up
in each nerve fibre a brief impulse or w a v e of activity
which travels rapidly down the fibre and produces a sudden
contraction in the muscle. A t each point on the nerve the
activity lasts for only a few thousandths of a second and
wherever it appears it involves a potential change, the
active region becoming negative to the inactive regions
beyond it. T h u s if the nerve rests on two electrodes a
potential change occurs as soon as the active region reaches
the first (Fig. 2) and when it reaches the second the po-
tential change is reversed.
T h e diphasic potential waves shown in Fig. 1 are the
exact counterpart of the diphasic w a v e which accompanies
the volley of nerve impulses produced by electrical stimu-
lation, though they are very much smaller, as they are
bound to be if they are due to single nerve fibres acting
independently. I t appears then that a succession of im-
pulses is passing up the nerve fibres from the sense organs
9
at the toe joint, and that the sensory message consists of

a series of brief active periods in each fibre separated by
pauses of varying length.
ι аяш ι
• д
I ι
DIPHASIC
POTENTIAL. WAVE
Fio. a. Production of a diphasic potential wave by the passage of an impulse

down a nerve fibre.
The Activity of Single Nene Fibres

When a nerve is stimulated electrically the potential
change in each fibre is of fixed magnitude and duration; it
depends only on the state of the fibre and not on the
strength of the stimulus which set it in motion. In the
record in Fig. ι the potential changes are not all the same,
for there are at least three different sizes. It is conceivable
that the waves set up by a sense organ can vary in size
although those set up by electric stimulation of the nerve
cannot, but it is much more likely that the different sized
waves are due to different sized nerve fibres, for the fibres
running to the toe joint are not all exactly alike. T o
decide this point and generally to learn what is happening
in each nerve fibre the preparation must be arranged so
that only one nerve fibre can come into action.
There are various methods by which this can be done
and they play an important, indeed an essential part, in
the analysis of nerve activity. The simplest method in
10
D E V E L O P M E N T OF ELECTROPHYSIOLOGY
theory and the most difficult in practice is to cut through

nearly all the fibres in the nerve in the hope of arriving at
a stage when only one active fibre remains intact. Pro-
fessor Bronk and I have spent many hours in this tedious
pursuit, for it is the only way of dealing with the motor
nerve fibres. With the sensory fibres it is usually much
better to leave the nerve alone and try to arrange matters
so that only one end organ will be thrown into action by
the stimulus. This cannot be done in a preparation of the
sort used in Fig. i, for there the end organs are of the type
known as Pacinian corpuscles and there is a bunch of
eight or more between the tendon and the joint capsule.
But in other situations and with other types of end organ
it is often fairly easy to stimulate one at a time. For
instance Matthews ( 1 9 3 1 ) has found that one of the small
toe muscles in the frog contains as a rule only one end
organ (a muscle spindle) which responds when the muscle
is stretched; in the skin it is often possible to stimulate a
single pressure organ or a single hair and so on.
When one of these preparations is used the sensory
message takes on a much simpler aspect. Fig. 3 is a
record made by stretching the frog's toe muscle and
... Τ
L ^\ L -1 LTL i ~- 1L LI LI TL kMkMi U U k b L L k L U ΙL ΙL ' k1 k k k L y
' IT 1 LI T I 1 11
Ι o·/ secoA/iУ .
FIG. 3 . Impulses in a single nerve fibre from a frog's muscle spindle stimulated
by stretching the muscle (Tsai, 1931)· Matthews' toe muscle preparation.
The potential waves are all alike; their frequency increases with the tension and
then declines slowly when the tension is constant.
recording the potential changes in the nerve. The potential

changes are now of uniform size; they form a regular series
with a frequency which increases to a maximum as the
muscle is extended and then declines again as the stretch
is maintained. The regular succession of waves is most
11
clearly marked in the discharge of sense organs which

become slowly adapted to the stimulus, for when adaptation
is rapid, as in many of the organs in the skin, the discharge
is too brief to show much regularity. But the potential
changes are again of uniform size, and an illustration of
this is given in Fig. 4, which shows the activity in a few
cutaneous nerve fibres of the frog, the skin being stimulated
by intermittent pressure.
FIG. 4. Impulses from sensory endings in the frog's skin stimulated by light
touch.
If these records give a true measure of the activity in

the sensory nerve fibres it is clear that they transmit their
messages to the central nervous system in a very simple
way. The message consists merely of a series of brief
impulses or waves of activity following one another more
or less closely. In any one fibre the waves are all of the
same form and the message can only be varied by changes
in the frequency and duration of the discharge. In fact
the sensory messages are scarcely more complex than a
succession of dots in the Morse Code.
The same kind of electrical activity is found in the
motor nerve fibres. When a message passes down a motor
fibre from the central nervous system to arouse a muscular
contraction we find again a rhythmic succession of potential
changes alike in size but varying in frequency. Fig. ς
gives a record from a single motor fibre supplying the
peroneus longus muscle in a spinal (decapitate) cat. The
record was taken at the beginning of a flexion reflex evoked
by pinching the foot; the movement developed slowly and
the development goes hand in hand with the increase in
frequency of the potential waves. Thus the effect produced
12
on the motor nerve fibres by the excited nerve cell in the

spinal cord is like t h a t produced on the sensory fibre by
the stretched muscle spindle ( F i g . 3 ) . It m a y be noted
that in F i g . 5 the recording instrument w a s a capillary
electrometer, not an oscillograph, and as the m o v e m e n t is
v e r y highly d a m p e d the true form of the potential w a v e s
is not shown, although it can be worked out by a fairly
simple analysis.
1*5'*«*-
FIG. 5. Impulses in a single motor nerve fibre supplying the peroneus longus
in the cat (spinal). The nerve has been cut down until only one active fibre
remains (Adrian and Bronk, 1929). A flexion reflex is produced by pinching the
foot and the frequency of the discharge increases as the contraction develops.
Record made with capillary electrometer and amplifier. The potential changes
are actually diphasic though the damping of the electrometer movement is too
great to show their true form.
• 4? ''J-Jee.
F i c . 6. Impulses in a motor nerve supplying the tail muscles in a caterpillar

(Adrian, 1930^). Record made at the end of a creeping movement showing the
declining frequency of the discharge. T h e nerve has not been cut down, but the
number of motor fibres is small enough to allow the individual impulses to be seen
(cf. p. 7 1 ) .
T o complete the series and to show t h a t the behaviour

of invertebrate nerve fibres agrees with t h a t of vertebrate,
two more records are given, this time from the nerves of
the caterpillar. Fig. 6 is a discharge f r o m one of the
motor nerves in the tail; the record w a s m a d e at the end
instead of the beginning of a muscular contraction so that
13
the frequency of the waves declines instead of rising as in

Fig. 5. Another difference is that more than one nerve
fibre is in action, for there are some very small potential
waves as well as the large waves of uniform size, but in
spite of this the general resemblance is clear enough.
Fig. 7 is also from the nervous system of the caterpillar,
but in this case the electrodes are placed not on a peripheral
FJG. 7. Impulses in central nerve cord of a caterpillar. A portion of the

central chain of ganglia has been isolated and one of the ganglion cells is discharging
spontaneously.
nerve but on the nerve cord which joins two of the ganglia
of the central nervous system. A message is passing from
one ganglion to the other and it is shown b y the usual
succession of diphasic waves.
The Nature of the Nervous Messages

According to these records nervous communication of all
kinds is carried out in the same w a y by a succession of brief
impulses. T h e impulses are not all exactly alike—those in
the sympathetic system and in the smallest sensory fibres
are conducted much more slowly than those in the large
motor and sensory fibres— but the records show no evidence
of any kind of activity other than that involved in the con-
duction of impulses. B u t can we assume that these records
of potential change really tell us w h a t is happening, or all
that is happening in the nerve? C a n there be nervous com-
munication without electrical change, or if there is always
an electrical change can we use it as a measure of w h a t is
going on in the fibre? These are questions which must be
faced before we go further, though they involve some
discussion of theory.
14
It is of course impossible to prove that electric changes

invariably occur when a nerve is in action. All we can
do is to point to the large number of experiments in which
they have been found, and to the close correspondence
between the message, as inferred from the electric changes
and as inferred from the effects it produces in the body.
In every case so far investigated whenever there has been
reason to suppose that a nerve is in action, the usual
potential changes have been detected, provided that the
conditions for recording them were favourable. This last
clause offers a refuge which is seldom needed, but it is
needed to explain the failure to record potential changes
from non-medullated sensory fibres in the mammalia.
This will be discussed further in connection with the
mechanism of pain, but meanwhile it is reasonable to
suppose that the failure is due not to the absence of electric
effects, but to their very small intensity under the condi-
tions of the experiment.
There are many instances where the record of electrical
activity has shown some unexpected feature which on
investigation is found to account for some special reflex or
central effect of the message. F o r instance Hoffmann
(1922) found some years ago that if the patellar tendon is
tapped whilst the thigh muscles are contracting, the
resulting knee jerk is followed by a brief pause in the
contraction. This 'silent period' has been explored by
Denny-Brown (1928) and shown to be a reflex effect of the
sensory discharge from the thigh muscles. Recently
Matthews (1931^) has recorded the impulses coming from
the muscle spindles and has found that a sudden contraction
of the muscle causes a brief pause in the sensory discharge.
This was unexpected but it evidently explains the silent
period after the knee jerk. Another example is that of
the persistent discharge of impulses in the sympathetic
nerves: as the discharge is concerned in maintaining the
tone of the blood vessels we supposed that the impulses
15
would be arranged in a fairly regular and uniform sequence,

but instead we found that they are often arranged in groups
occurring in phase with the movements of respiration.
This result, though quite unforeseen, is really quite in
accord with what we should have expected if we had looked
up the literature beforehand, for Fredericq showed in 1882
that the blood pressure may rise and fall with the frequency
of respiration, although mechanical effects on the blood
vessels have been excluded by opening the chest.
Unexpected agreements of this kind are the most satis-
factory vindication of the electric record as a true index of
the message in the nerve fibres, and so far there has been
no clear instance of lack of agreement. In fact the cor-
respondence has occurred so often that according to all the
rules we are justified in assuming that it is universal.
Gradation of Activity
There remains the second question, whether we can
assume that the potential changes are a true measure of
the impulses in the nerve fibre, whether the impulses can
v a r y in size without there being any corresponding variation
in the electric response. There is certainly no evidence
to suggest that the impulses are graded in size, for the fact
that sensory messages may produce a small or a large effect
according to the intensity of the stimulus is naturally
explained by the varying number of fibres in action and by
the varying frequency of the discharge in each fibre. B u t
the point is so important that it is worth trying for the
moment to take up the role of a hostile critic with a rooted
belief in the complexity of nature. Such a critic might
argue like this: " I f I look out on to a main road at night I
see a succession of bright lights travelling rapidly. T h e y
are all of much the same intensity, but would you ask me
to believe that they are all due to motor cars of the same
make? This is what you expect me to believe of the nerve
fibre when you argue from the potential wave to the
16
impulse. Surely each impulse in a message may have a

distinct individuality, and instead of being like a succession
of dots in the Morse Code the message may be like a
succession of numbers, or words, or sentences. And I am
not at all sure that the message is necessarily made up of
distinct impulses. To account for your potential changes
let us suppose that when any part of the nerve fibre
becomes active there is an immediate breakdown of a
polarised surface membrane, giving a potential gradient of
fixed intensity, that the breakdown is immediately repaired,
but that if the underlying activity persists the surface
breaks down again giving another potential wave, and so on.
On this view the potential change may be merely incidental:
the essential activity which the nerve fibre transmits may
be perfectly continuous, rising and falling with the excita-
tion of the sense organs or nerve cells and capable of much
more rapid and delicate fluctuations in intensity than are
revealed by the electric record. You have something
which you can measure and no doubt it is very closely
related to the activity of the nerve fibre. But changes
in lactic acid content seemed to be just as closely related
to the contraction of muscle, and we have had to give up
the belief that they tell the whole story. The potential
changes may show that a message is passing, but I see no
reason to believe that the whole content of the message is
revealed by them."
In answering this we have to be careful to define what
is meant by the impulse or the wave of activity in a nerve.
It must be taken to mean not the whole sequence of changes
which take place at each point in a nerve fibre after a
stimulus, but only the change which propagates itself down
the fibre and leads to the activity of other structures such
as muscles or glands. The passage of a single impulse
down a fibre may give rise to chemical and thermal changes
which are still in evidence five minutes later, five minutes
after the impulse has done all that it can in the way of
17
activating other regions. These chemical and thermal

changes are no doubt of graded intensity, since each new
impulse must add its own after-effects, and in this sense
the activity of a nerve fibre may be graded and persistent.
But the 'propagated disturbance,' the activity which
arouses other parts of the fibre, is a momentary affair and
its intensity cannot be altered except by changing the local
condition of the fibre.
It is true that the evidence must be taken from experi-
ments on motor nerves stimulated electrically, but there is
abundant proof that the impulses set up in this way are
identical with those arising from sense organs or nerve
cells. The impulse produced by a single electric shock is
certainly incapable of any kind of gradation which might
play a part in the working of the nervous system, for one
set up by a strong stimulus can do no more than one set
up by a weak. It travels no faster and no further (if
there are obstacles to its progress) and causes no greater
effects on the structures which are activated by it. Its
brevity is shown most clearly, I think, by an old experiment
of Boycott's (1899). If two stimuli are sent into a frog's
nerve at an interval of less than about .003 sec. the muscle
gives a twitch no greater than if only one stimulus is used.
When the interval is lengthened progressively the twitch
suddenly increases. Evidently the second stimulus has
now become able either to prolong or intensify the first
propagated disturbance or else to set up a fresh disturbance
of its own. Now if the nerve is cooled somewhere between
the stimulated region and the muscle, the twitch reverts
to its original height. This is easily explained if there are
two brief impulses but not if there is one long one. On
the former view the failure of a very early second stimulus
is due to its falling at a time when the nerve fibre is still
disorganised by the passage of the first disturbance, and
this refractory state will be lengthened in the region where
the nerve is cooled. On the other hand if the second
18
stimulus, when successful, produces not a second dis-

turbance but a prolongation of the first there is no reason
why it should cease to be effective when the nerve is cooled.
The Membrane Hypothesis
There are many other arguments of the same kind
depending on the phenomena of the refractory state, but
when all is said and done the best argument comes from
the whole body of evidence in favour of what is known as
the membrane hypothesis of nervous conduction. Ac-
cording to this hypothesis the resting nerve fibre is enclosed
by a polarised membrane, and this breaks down locally
under the action of the stimulus, establishing a potential
difference between the active and inactive regions. The
potential gradient (or the concentration gradients which
produce it) causes a differential movement of anions and
cations at the boundary of the depolarised region, and this
leads to a spread of the depolarisation and at the same
time to a restoration of the surface already depolarised
(Fig. 8). The restored surface is at first refractory and
therefore unable to break down again, and thus the area
REFRACTORY ACTIVE RESTING
DIRECTION OF TRAVEL >
FIG. 8. Spread of active (depolarised) area according to the membrane

hypothesis. The local currents cause a repair of the active region and a breakdown
of the resting surface beyond.
19
of breakdown shifts down the nerve fibre; the potential

change in each section acts as the stimulus to the next
section and the surface is automatically restored to the
polarised state after a brief period of activity. It follows
that the potential change is an essential link in the trans-
mission of the impulse: if it is found not to vary with the
stimulus there can be no possible variation in the impulse:
the duration of the period of surface breakdown must
coincide with that of the potential change, and the surface
breakdown is, in fact, the impulse, the disturbance which
propagates itself down the nerve fibre.
If we accept the membrane hypothesis in its general
outlines we need not hesitate to take the potential changes
as a faithful record of the way in which the nerve fibre
performs its function. There is no one experiment which
excludes all other explanations of nervous conduction but
there are very many phenomena of nerve and muscle
activity which can be most simply explained on this basis
and none which contradict it. It has been supported most
strikingly by R. S. Lillie's iron wire model—a non-living
system in which surface changes are rapidly transmitted
by the agency of the local currents set up at the margins
of the active region—and by the recent work of Α. V. Hill
(1932) which shows that only an extremely small amount
of heat is evolved during the passage of an impulse in
comparison with the amount evolved afterwards in restoring
the free energy of the fibre. Lillie's work shows that a
transmission of this kind is a physical possibility, that it
would be likely to happen in a structure such as a nerve
fibre and that the character of the transmission would
agree with that found in nerve. Hill's shows that the
main event in the conduction of an impulse is most probably
a physical change like the movement of ions postulated on
the membrane hypothesis. The hypothesis is in no sense
a complete picture of nerve activity but it is the only
satisfactory picture of the mechanism of conduction.
20
We come back then to our records of nervous messages

with a reasonable assurance t h a t they do tell us what the
message is like. It is a succession of brief waves of surface
breakdown, each allowing a momentary leakage of ions
from the nerve fibre. The waves can be set up so t h a t
they follow one another in rapid or in slow succession, and
this is the only form of gradation of which the message is
capable. Essentially the same kind of activity is found in
all sorts of nerve fibres from all sorts of animals and there
is no evidence to suggest that any other kind of nervous
transmission is possible. In fact we may conclude that
the electrical method can tell us how the nerve fibre
carries out its function as the conducting unit of the
nervous system, and that it does so by reactions of a
fairly simple type.
21
Chapter I I
THE ACTION OF THE SENSE ORGANS
Ε sensory discharge is something which can be

.neasured. It gives us our most direct information
about the working of the sense organs, for there is no
adequate measure of sensation, and reflex effects, though
measurable, are too far removed from the sensory appa-
ratus. A record of nerve impulses cannot tell us what
effect the message will have when it enters the central
nervous system, but it shows how the sense organ responds
to the stimulus.
Unfortunately we cannot always identify the sense organ
which gives rise to a particular series of impulses, for over
most of the skin the different types are so closely spaced
that it is impossible to say which of them are in action.
I t is for this reason that much of our information relates to
sense organs which have not been identified histologically,
to pressure receptors, temperature receptors, etc., and not
to Meissner's corpuscles or Golgi Mazzoni bodies. But
there are some receptors whose action can be identified,
e.g., the muscle spindles, the Pacinian corpuscles, the hair
endings, and certain types of ending in the viscera, and
no doubt the list could be extended without much difficulty.
The sense organs listed above are all stimulated mechani-
cally, and with all of them the effective change seems to be
a stretching of the terminal part of the sensory nerve fibre.
As with most sense organs, the ending of the fibre has
various accessory structures in close contact with it. These
may be intimately concerned with the process of excitation,
but their most obvious effect is to produce a given defor-
mation of the ending in response to a given stimulus.
22
ACTION OF T H E S E N S E ORGANS
With the hair organ a lateral movement of the tip of the

hair will distort the ring of endings round its base; with
the Pacinian corpuscle pressure in any direction will
elongate the central core and stretch the endings in it;
the muscle spindles will be stretched by a pull on the
tendon, and so on. T h e form of the accessory structures
will determine the direction in which the force must be
applied, and their rigidity will decide the amount of force
needed for a given extension of the endings. They will
act like a system of levers, springs, and shock absorbers,
and it is possible that this is their only function. I f so
the different kinds of corpuscles, etc., may serve merely to
fix the range of stimuli to which each ending will respond
and so to provide sense organs differing greatly in their
excitability to mechanical forces.
B u t the sense organs do not differ merely in excitability.
There are, of course, the receptors for thermal and chemical
stimuli which are not activated mechanically, but if we
confine ourselves to the receptors which are activated by
deformation we find great differences in the time course
of the discharge. With some of them the discharge only
takes place during the actual movement, with others it
persists as long as the deformation remains, though at a
gradually declining frequency.
Adaptation
T h e rapid failure in the exciting effect of a constant
stimulus is a well-known phenomenon in nerve physiology.
A constant current applied to a nerve gives as a rule only
one impulse, and a current which increases gradually from
zero may never excite at all. T h e excitation only occurs
whilst the conditions are actually changing, and the rate
of change must exceed a certain value. T h e persistent
stimulus does not give a persistent discharge because the
fibre becomes rapidly adapted to the new conditions.
23
Adaptation is perhaps a dangerous term to use, for it

seems to imply a positive reaction on the part of the nerve
fibre. There may be such a reaction, but for all we know
the rapid failure of the stimulus may be due to simple
physical changes such as diffusion or polarisation. The
change may not even take place in the nerve fibre at all,
for Bishop (1928) has suggested that polarisation in the
nerve sheath may be largely responsible. At any rate we
find the same rapid or slow decline in the effect of a stimulus
to a sense organ. Here we are justified by long usage in
speaking of adaptation, and it is unlikely that there is any
great difference in the process as it occurs in the sense
organ and in the nerve fibre.
The variations in adaptation rate can be seen most
clearly by recording the discharges of the sensory fibres in
a mixed nerve, or better in one of the dorsal nerve roots.
When one of these is placed on the electrodes it is found
that impulses are constantly passing up from sense organs
in the periphery. They come mainly from deep-seated
sense organs which adapt very slowly and signal the
posture of the limb. The slightest interference with the
limb produces a sudden but transient increase in the
discharge. As long as the movement continues, impulses
pass up in the nerve fibres from hairs and tactile endings,
but most of this extra discharge ceases when the movement
is over, for the endings adapt rapidly and are no longer
excited although the hairs remain in the bent position.
The steady discharge from the postural organs also increases
during the movement and may be left with an altered
frequency corresponding to the altered distribution of
muscle tension and pressure.
Almost any preparation containing a good many sense
organs will react in this way, with a persistent discharge
in some nerve fibres during an abnormal posture, and a
brief discharge, confined to the period of actual movement,
in others. In the skin the rapidly adapting endings are
24
in the majority, in mammalian muscles both are present

in varying proportions, and in frog's muscle nearly all the
endings adapt slowly. But the distinction is one of degree
only. It is worth making when we are discussing the
function of the different organs, but it does not mean that
there is any fundamental difference between the mechanism
of the muscle spindle and of the hair ending. They are
merely at opposite ends of the scale, and in fact some
muscle spindles adapt fairly rapidly and some hair endings
fairly slowly.
It should be pointed out that, as adaptation always
takes place to some extent, there is no sense organ in
which the frequency of the discharge can be said to depend
entirely on the intensity of the stimulus. We can say if
we like that with all sense organs the frequency is deter-
mined by the intensity of the excitation, though this is
merely defining what we mean by intensity of excitation.
But the stimulus, the mechanical change inflicted on the
sense organ, is always discounted more or less rapidly by
the adaptation which it produces, and the time factor is
quite as important as the intensity factor, even with the
organs which adapt very slowly. The slowly adapting or
postural end organs which have been investigated up to
the present are those responding to muscle stretch in the
frog and mammal, the Pacinian corpuscles and various
unidentified organs responding to pressure, the sensory
endings of the vagus in the lung root and those of the
cardiac depressor nerve and the nerve to the sinus caroticus.
With all of them the frequency of the discharge depends
on the rate of increase, the final intensity and the duration
of the mechanical change which acts as the stimulus, and
to obtain the maximum frequency the stimulus must not
only reach a great intensity but must reach it very quickly.
For instance Matthews (19310) finds that to make the
frog's muscle spindle discharge at the highest possible rate
the muscle must be suddenly pulled out by a spring, and
25
Bronk and Stella (1932) find that the rate of rise of the
arterial pressure is a very important factor in the discharge
of the sense organs in the aorta and sinus caroticus.
Records of the sensory discharge from the vagal endings in
the lung are given in Fig. 9. In the lowest record the lung
is inflated suddenly and the inflation is maintained. T h e
С
Fic. 9. Sensory impulses in the vagus from a single end organ in the lung.
Nerve cut down to show single fibre discharge.
A. Decerebrate cat breathing normally; the record shows the discharge during
a single movement of inspiration, maximum frequency 16/sec.
B. Another animal, hyperpnoea due to breathing into a closed space, maximum
frequency 72/sec.
C. Same preparation as B. Discharge produced by sudden, forcible inflation
of the lungs. The frequency rises to 150/sec. and has fallen to 1 1 5 after the first
second.
frequency of the discharge falls appreciably in the first

second, though the impulses are too closely crowded for
this to show well in the reproduction.
B u t provided we specify the whole evolution of the
stimulus we can often predict the response with very great
accuracy indeed. With a mammalian preparation it is
naturally difficult to keep all the conditions constant over
long periods, but with a frog's muscle spindle M a t t h e w s
could repeat a series of stimuli knowing that the response
to them would v a r y by less than 5 % in an experiment
lasting several hours—and the response is a complex affair
26
A C T I O N OF T H E SENSE ORGANS
rising and falling in the w a y shown in F i g . 3. T h i s machine-

like regularity of behaviour m a y be taken as a proof that
the working of the nervous s y s t e m ought not to be beyond
mechanical description. B u t those who dislike mechanism
might well retort that the sense organs would be of little
use to the body if they could not be trusted to give the
s a m e message for the same stimulus. T h e receptor and
effector a p p a r a t u s must respond with rigid precision if the
central nervous system is to be in full control, and it is
only in the central nervous system that our models m a y
be of questionable value.
The Production of the Rhythmic Discharge

When we come to the actual mechanism concerned in
the discharge of impulses we are on less certain ground.
T h e main evidence is that concerning the frequency and
regularity of the discharge. When a frog's muscle spindle
is stretched the discharge is regular, in the sense t h a t the
intervals between successive impulses lie on a smooth
c u r v e , as long as the frequency is above about 20 a second.
Below this there is much greater variation, though the
intervals are never shorter than one-twentieth of a second.
W i t h m a m m a l i a n sense organs the regularity persists at
much lower frequencies: the pressure receptors often dis-
charge regularly at 1 0 a second and the muscle spindles are
still regular at 5 a second.
T h e r e is a simple explanation of this behaviour, but
u n f o r t u n a t e l y it does not t a k e us v e r y far. A nerve fibre
itself can only be excited intermittently, for each impulse
sets up a r e f r a c t o r y state which passes a w a y gradually.
B u t if the end of the fibre were altered in such a w a y that
it could exert a continued stimulating effect on the rest
of the fibre it could not fail to produce a regular discharge.
T h e discharge would be regular because a f t e r each impulse
the normal fibre recovers along a smooth curve. An
impulse must be set up each time the excitability returns
27
to the level at which the stimulus is effective, and if the

stimulus and the rate of recovery do not change the
impulses will be evenly spaced. The regularity would
cease when the frequency falls below a limiting value; the
excitability does not go on rising indefinitely after each
impulse, and a necessary condition for the automatic
firing is that it should occur before recovery is complete.
An irregular response must have some other explanation,
but it is clear that any condition which can act as a per-
sistent stimulus to a nerve fibre would give a regular
series of impulses over a certain range of frequency.
The mere regularity of the discharge can only tell us
that a persistent stimulus is acting on a structure which
must respond intermittently, but in the sensory discharge
the particular range of frequency tells us a little more.
It shows that the structure in question is not the unaltered
nerve fibre, but is some transitional region between the
nerve fibre proper and the part of the ending which is
sensitive to the stimulus. A region of this kind is necessary
to account for the wide range over which the discharge
may vary without losing its regular character, for with a
mammalian nerve fibre the recovery is so rapid that a
persistent stimulus could not produce a regular discharge
at a frequency much lower than 150-100 a second.
When they were first examined it seemed that the whole
range of frequencies in the sensory discharge was lower
than it would be in a stimulated nerve fibre, but later
work has shown that there is no clear difference at the
upper end of the scale. With intense stimulation the
frequency from a sense organ can be made to rise to a
value very little below the maximum frequency for nerve
(Matthews, 1931л, Adrian, Cattell and Hoagland, 1931).
Thus the structure in which the discharge arises must have
an absolute refractory period scarcely longer than that of
the nerve fibre, but the later stages of recovery must be
slower. It is by no means certain that rapid and slow
28
ACTION OF THE SENSE ORGANS
discharges arise from the same point: there may be a

gradual transition from nerve fibre to nerve ending with
a gradual slowing of time relations, and an intense stimulus
may take effect at a point where recovery is rapid. But
in any case the intermittent activity, the succession of
impulses, must begin at some point in the nervous apparatus
of the sense organ and the recovery rate is a matter of
detail.
Our chief concern is with the persistent stimulus which
sets up the discharge. We must look for some kind of
change in the sense organ which would have this activating
tendency and would be brought about by such diverse
causes as mechanical deformation, temperature change,
and change in chemical environment.
Depolarisation in the Sense Organ

One simple possibility is that the change in the sense
organ is a surface depolarisation like that occurring in the
active nerve fibre, but persistent instead of momentary.
This is suggested by the recent work of R . S. Lillie on the
iron wire model of nervous conduction. Lillie's model is
an iron wire coated with a thin protective film, probably
an oxide, by immersion in strong nitric acid. If the film is
destroyed locally, a w a v e of surface disintegration travels
down the wire under the influence of the potential gradient
between the active and inactive surfaces. As in nerve,
the surface film is re-formed in the wake of the wave and
the activity at each point lasts only a short time. T h e
analogy with the nervous impulse is very close indeed, and
the analogy m a y be extended to the sense organ. If the
conditions are arranged so that one end of the wire is kept
permanently in the active state, this exerts a persistent
stimulating effect on the rest of the wire and a succession
of waves travel down it (Lillie, 1929). T h e stimulating
effect is due simply to the difference of potential between
the exposed or active iron at the end of the wire and the
29
passive film beyond it, in fact t h e active region sets u p a

train of waves for the same reason t h a t causes t h e spread
of the individual wave from one point to another. T h e
frequency of t h e waves depends on the rate of recovery of
the wire and on the size of the permanently active area;
it can be varied therefore by increasing or reducing this
area. In the same way persistent activity, or, in terms of
the m e m b r a n e hypothesis, persistent depolarisation in a
sensory nerve ending might account for the r h y t h m i c
impulse discharge. A partial or complete breakdown of a
polarised surface is an event which would be likely to
follow mechanical deformation; and if the extent of the
breakdown depends on the a m o u n t of deformation it
would determine the frequency of the rhythmic discharge
as in the iron wire model. T e m p e r a t u r e change or chemical
change might have the same destructive effect on the
surface of the endings which respond to these stimuli, and
again the area involved or the degree of depolarisation
would depend on the intensity of the stimulus.
The Injury Discharge

Whether or not depolarisation explains the discharge
from sense organs, there is no doubt t h a t i n j u r y , which
involves p e r m a n e n t depolarisation, can set up a rhythmic
discharge in a nerve fibre. These injury discharges are
abnormal effects, but the injured and the active states are
so closely allied t h a t it is worth spending a little time over
them. Both give the same potential change, for both
involve a breakdown of the polarised surface membrane,
though in the intact fibre the breakdown is promptly
repaired. Now we suppose t h a t an impulse travels down
a fibre because the depolarisation of one section causes a
depolarisation of the next. I t follows t h a t an injury
should s t a r t an impulse down the fibre, and if nothing
occurs to diminish its effect we should expect it to produce
fresh impulses whenever the fibre is ready to conduct them.
30
A C T I O N OF T H E S E N S E ORGANS
In many nerve fibres this repeated activity is not found.

A permanently depolarised area is formed whenever a
nerve is cut through, but the response may v a r y from a
single impulse to m a n y thousands: in fact the rate of
adaptation to the stimulus may v a r y as much in a nerve
fibre as it does in a sense organ. In the nerve of a frog it
is unusual to find much persistent activity due to nerve
section, and mammalian motor fibres give only a brief
discharge when they are cut through. B u t mammalian
sensory fibres are often much more active (Adrian, 1930a).
A medium-sized nerve trunk removed from the body and
kept in a warm, moist atmosphere is seldom free from
impulses. T h e y are due to the smaller nerve fibres and
arise from one or other of the cut ends. T h e discharge
m a y subside after a few minutes but it is often renewed by
slight changes in the condition of the nerve, e.g., by
irrigation, readjustment on the electrodes, or a draught of
cool or warm air. Another factor which affects the injury
discharge is the presence or absence of a thick sheath
round the nerve. If the sheath is stripped a w a y the
discharge ceases, and if there is a v e r y thin sheath (as in
the smallest nerves) there is usually v e r y little activity.
W h y the sheath should make so much difference is not
very clear, but the effect is not confined to the sensory
fibres, for the motor discharge is also longer when a nerve
is cut in a region where the sheath is well developed.
T h e simplest type of injury discharge is of high frequency;
it begins when the nerve is cut through and it can always
be produced in an intact nerve by placing on it a crystal
of calcium chloride or of any salt which will give a high
osmotic pressure. As this treatment gives a progressive
injury it allows us to study the beginning of the discharge
as well as its later phase. T h e sequence of events is then
as follows: at first an occasional impulse appears, then the
rate quickens, but the discharge in each fibre is still irregular
until it suddenly becomes regular at a frequency in the
31
neighbourhood of 150/sec. (at 35 0 C.). The frequency

rises to 500-800/sec. and the loud speaker record changes
from a low note to a high-pitched wail with more and more
fibres taking up the cry as they feel the effects of the
injury. Eventually the frequency in each fibre declines
again and there is the same abrupt change from the regular
to the irregular discharge below 150 impulses a second.
An example of the high frequency discharge in a single
nerve fibre is given in Fig. 10. In this case the discharge
was brought to an end by cooling the preparation slightly
and the records show the onset of the irregular phase.
FIG. 10. High frequency injury discharge. Dorsal cutaneous nerve of cat,
removed from the body and set up in a moist chamber. Continuous discharge
(300/sec.) changing to an irregular discharge of low frequency. The change was
produced by lowering the temperature about 5 0 C .
Discharges of this type show what happens when a

persistent stimulus acts on a nerve fibre rather than on the
transitional region which we have postulated in the sense
organ. The stimulus in this case is the injury and pre-
32
sumably the depolarisation which occurs at the injured

region, and the discharge resembles that from a sense
organ, though it occurs over a higher range of frequencies.
The frequency range is explained by the fact that a mam-
malian nerve fibre at body temperature takes about 1/150
sec. to recover its full excitability after an impulse. Conse-
quently the regular discharge cannot occur at a lower
frequency than 150/sec. and it must vary between this and
the upper limit set by the refractory period of the nerve
fibre. T o account for the irregular discharge is as difficult
here as it is with the sense organs; we must suppose that
there are irregular fluctuations in the threshold of the
nerve or in the strength of the stimulus, though this merely
amounts to the statement that something is irregular—
and this we knew already.
T o revert now to the sense organs; we have a reasonable
working hypothesis in the notion that the stimulus causes
a persistent depolarisation, or surface leakage, of variable
extent and that this sets up the intermittent waves of
depolarisation which spread down the nerve fibre. But
there is another possibility, though it can only be stated in
vague terms. It rests on the fact that an electric stimulus
appears to act on a nerve fibre by setting up some prelimi-
nary change which is not itself a depolarisation. Bishop
(1927) has shown this by recording the potential changes
in the region under the stimulating electrodes: an inade-
quate stimulus, though it increases the excitability of
the fibre, causes no potential change and therefore no
depolarisation, and a slowly increasing current causes no
depolarisation until the moment when the impulse is fired
off. This means that the excitability of the fibre, or, to
be more concrete, the instability of the surface membrane,
can be made to increase up to the point when the surface
breaks down, although until it does so there has been no
change in permeability. Thus the essential change in the
sense organ may be not a depolarisation but a development
33
of the condition which leads up to it. There would then

be no persistent depolarisation in any part of the ending
or fibre but merely the brief waves set up in the region
where the surface has become unstable. To decide the
point we should have to record the potential difference
between the sense organ and its nerve fibre and this,
though simple in theory, would be very difficult to carry
out experimentally. So for the present we must leave
this part of the picture in obscurity. It is very much
better to do so, for we have so little real knowledge of the
physics and chemistry of the cell surface that any picture
must be a crude likeness with half the features left out.
Differences in the Sensory Pathway: Touch and Pressure

Variations in excitability and in adaptation rate are the
chief differences which we can observe in the reactions of
the sense organs which respond to mechanical stimulation.
We shall see later that there are differences in the rate of
conduction of the impulses as well—the differences which
have been worked out by Gasser and Erlanger. But
apart from the peripheral apparatus we find that there are
also great variations in the time course of the central
events. These are revealed, I think, by the quality of
sensation which we experience when different receptors are
stimulated. Arguments which jump from the material
plane to the mental may land us in all kinds of error, but
this is the argument for what it is worth.
Firm pressure on the surface of a limb will stimulate
some of the slowly adapting endings, and these will give a
regular succession of impulses at a frequency depending on
the pressure. An example of such discharge is given in
Fig. I i . It was made from one of the sensory nerve roots
in the cat and shows the effect of pressure on the side of
the foot. A t the beginning, with very light pressure, the
frequency of the impulses averages only 9 a second. This
may be below the threshold of sensation, but sooner or
34
A C T I O N OF T H E SENSE ORGANS
later the frequency will become a d e q u a t e . Y e t pressure

applied to our own sense organs gives a sensation w h i c h is
perfectly continuous: it rises and falls along a c u r v e which
seems to run closely parallel to the c u r v e of rising and
falling impulse frequency and it never becomes i n t e r m i t t e n t .
A s a rule, no d o u b t , a good m a n y sense organs m a y be in
action and the composite discharge in several nerve fibres
would not h a v e any definite r h y t h m , but a restricted
stimulus m u s t occasionally t a k e effect on a single pressure
organ and there is no reason to think t h a t the sensation
would then be intermittent. It follows t h a t somewhere
on the p a t h w a y between the sensory nerve fibre and the
FIG. II. R e c o r d from one of the lumbar sensory rootlets in the c a t . Discharge
caused by pressure applied gradually to the side o f the foot. T h e upper record
shows the beginning o f the discharge. In the lower it has reached a f r e q u e n c y of
about ιοο/sec. and another nerve ending has begun to discharge at a lower frequency
(20/sec.). T i m e marker gives intervals of o . l j sec.
mind the intermittent message from the pressure organ

must be s m o o t h e d o u t into a s t e a d y a c t i v i t y . F o r this
to t a k e place each impulse m u s t p r o d u c e an excitation
which rises and declines so slowly t h a t successive impulses
can sum their effects. T h i s is not quite the same thing as
slow a d a p t a t i o n , but it implies a sluggish reaction in some
part of the central mechanism. Sherrington has given
m a n y examples of ' i n e r t i a and m o m e n t u m ' in reflex arcs
and the same conception m a y be applied here. N o w if we
turn to the receptors for light t o u c h we find an entirely
different s t a t e of affairs. T o produce a regular succession
of impulses at all c o m p a r a b l e w i t h those in F i g . 11 we
h a v e to use an intermittent stimulus r e p e a t e d at regular
35
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If you have exercised yourself in a manner worthy of the Olympic
Festival, if you have been guilty of no slothful or ignoble act, go on with
a good courage. You who have not so practised, go whither you will.[5]
The names of those who were to enter for the games were then written up
on a white board, and should a man withdraw after that, he was branded as a
coward. As soon as the competitor was finally enrolled, a boar was offered
in sacrifice to Zeus, and then he had to take a solemn oath that he was a full
Greek citizen, that he had fulfilled all the conditions necessary for the
games, that he would abide by the rules of the contest, and that he would
play fair, and such was the spirit of honour and fairness in which the games
were played, that in more than a thousand years there appear the names of
only six or seven competitors who were guilty of breaking their oath.
The first day of the festival was given up to sacrifices and processions.
The different states always sent official representatives to the Games, and
these would make public entrance in their chariots, richly arrayed and
bearing costly gifts to place in the treasury of the temple. The next three
days were devoted to the actual contests.
Long before the dawn on the first of these three days, every seat in the
stadium was occupied. It was situated at the foot of a hill, and every
available spot on the slope of this hill was used by the spectators. Should
anyone leave his place, even for an instant, it would be lost, and there the
spectators sat the whole day through, until the sun went down. What
refreshments they needed, they brought with them. The sun beat down on
their bare heads, for the Games were in honour of Zeus and he was looked
upon as present, and no one might enter the presence of the Father of Gods
and Men with covered head. Not until the setting sun gave the signal for the
end of the day's contests, did they hurriedly rush off to their tents and snatch
an hour or two of sleep before the coming of the dawn warned them to rise
and secure their seats for the next day's spectacle.
The contests probably took place in the following order: First, there were
the foot races: there were several of these varying in length from two
hundred yards to three miles. The shortest race of two hundred yards was for
a long time the race which brought greatest honour to the winner. Then
followed the pentathlon which consisted of five contests: throwing the
discus, throwing the spear, running, jumping and wrestling, and the winner
was required to have won three out of the five. In the pentathlon, in
particular, great importance was attached to the gracefulness of every
movement, and the jumping, discus and spear throwing were generally
accompanied by the music of the flute. Then came what was later regarded
as the greatest and most exciting race of all, the four-horse chariot race. This
was a race that poets loved to describe. Homer tells us how the charioteers
all together lifted the lash above their steeds, and smote them with the
reins and called on them eagerly with words: and they forthwith sped
swiftly over the plain; and beneath their breasts stood the rising dust like
a cloud or whirlwind, and their manes waved on the blowing wind. And
the chariots ran sometimes on the bounteous earth, and other whiles
would bound into the air. And the drivers stood in the cars, and the heart
of every man beat in desire of victory, and they called every man to his
horses, that flew amid their dust across the plain.[6]
The boxing and wrestling matches came last, and these were the roughest
and fiercest of all the contests.
On the last day of the festival the prizes were awarded. They were very
simple, but more highly valued than greater honours could have been. Each
prize consisted of a wreath of olive, which had been cut from a sacred olive
tree with a golden knife by a boy especially chosen for the purpose, and an
old tradition required that both his parents should be alive. These wreaths
used at one time to be placed on a tripod in the sight of all the people, later, a
beautiful table of gold and ivory was made for them. A herald announced the
name of the victor, his father's name and the city from which he came, and
then one of the judges placed the wreath on his head. This was the proudest
moment of his life, and though other rewards followed on his return home,
nothing ever quite equalled that glorious moment.
The last day of the festival was given up to sacrifices to Zeus, followed
by banquets and feasting which lasted late into the night. Every kind of
honour was shown the victors: poets wrote odes celebrating their victories,
and sculptors made models for statues of them, for to every athlete who had
won three victories was granted the honour of being allowed to have his
statue erected in the open space outside the temple of Zeus.
DISCOBOLUS OF MYRON.
5th Century B.C.
Vatican Rome.
The festival over, the victors and their friends and the great throng of
spectators returned to their homes. The victors were not only proud on
account of their own achievements, but for the glory they had brought to
their city. The news of the approaching arrival of a victor was sent on ahead,
and the day of his return to his native city was always honoured by a public
holiday. In some places it was an old custom to pull down a part of the city
wall and make a special entrance, in order that he who had brought the city
such glory might enter by a path never before trodden by other men. Songs
of triumph were sung to greet him, and he was led to his father's house along
a road strewn with flowers. Rich gifts were presented to him, and in every
way he was treated as a man whom the city delighted to respect and honour.
At Athens the returning victors were honoured by being allowed to dine
thenceforth at the public expense in the hall where the councillors and great
men of the city took their meals.
Pausanias, the traveller to whom we owe descriptions of so much in

ancient Greece that has now perished, visited Olympia, and he tells us that
many a wondrous sight may be seen, and not a few tales of wonder may
be heard in Greece; but there is nothing on which the blessing of God
rests in so full a measure as the rites of Eleusis and the Olympic Games....
and Pindar, the Greek poet who has most often sung of the Olympic Games,
summed up the feelings of every victor in the words: "He that overcometh
hath, because of the games, a sweet tranquillity throughout his life for
evermore."
[1] See p. 47.
[2] Pausanias.
[3] Pausanias.
[4] Dion Chrysostom.
[5] From E. N. Gardiner: Greek Athletic Sports and Festivals.
[6] Iliad, XXIII.
CHAPTER IV
THE GREEK CITY-STATE
Whenever men live together in communities, no matter how small they

may be, some form of law has to be observed, in order to maintain order, and
that there may be justice between man and his neighbours. The form that this
law takes in different places and in different communities is what is called
government.
The earliest form of government in Greece was, like all primitive

government, that of the family, and the word of the head of the family was
law to all those belonging to it. The land on which they lived belonged to the
family as a whole, not to separate individuals, and the dead were always
buried there, until in time the family claimed as their own that land, where
they had lived for generations, and where their ancestors were buried.
After a time it became more convenient for families to join together and
live in one community. By this means the labour of cultivating the land
could be more evenly distributed, and in times of attack from enemies, larger
and stronger forces could be used for defence. This grouping of families
together made a village and the strongest and most capable man in the
village would become its chief.
In time, just as families had found it more to their advantage to group
themselves together and form villages, so did the villages living in the same
neighbourhood find it a better thing to join together and form a still larger
community, which became known as a kingdom, because instead of having a
chief they were ruled by a king. At first the kings, like the chiefs, were
chosen because of their ability and power, later the office became hereditary
and was handed down from father to son.
Now because the Greek communities lived in the plains, separated from
each other by mountains, instead of forming one large kingdom, they formed
a great many small ones. There was in ancient times no King of Greece, but
Athens, Sparta, Corinth, Thebes and countless other cities had their own
independent forms of government, their own rulers, their own armies, their
own ships, and except that they were all Greek and were all bound together
by ties of language and religion, they were quite independent of each other.
All these independent cities became known in time as City-States, for to the
Greek the state meant the city, the territory immediately surrounding it was
included in the state, but the city was the most important part of it.
All communities are always governed in one of three ways: either by one
man, or by a few men, or by many men, and the Greeks tried all these ways,
until they found the one that answered best to their ideals of what a city-state
should be. All states did not develop in the same way, but one stands out
from the others as having most nearly reached the Greek ideal. That state
was Athens. Her story shall be told in its own place; in this chapter we will
see what the Greeks thought an ideal state should be, and what they believed
to be the duties of a good citizen.
The Greek philosopher Aristotle wrote a book in which these ideals were
set forth.[1] He believed that the end for which the State existed was that all
its citizens could lead what he called a "good life," and by that he meant the
life which best gives opportunities for man to develop his highest instincts,
and which makes it possible for every citizen to develop his own gifts
whatever they may be, in the highest and truest way. To realize such a life
there must be law and order in a state, and Aristotle considered that the first
thing necessary to ensure this was that the state must not be too large. He
believed that the greatness of a state was not determined by the size of its
territory or the number of its population, but that though a certain size and
certain numbers helped to make a state dignified and noble, unless these
were combined with good law and order, the state was not great. States, he
said, were like animals and plants or things made by human art which, if
they are too large, lose their true nature and are spoilt for use. But how is one
to know when the limit in size and population has been reached? Is there any
test by which it can be discovered whether a state has grown or is in danger
of growing too large?
Aristotle answered this question by saying that the state must be large
enough to include opportunities for all the variety and richness of what he
called the "good life," but not so large that the citizens could not see it or
think of it in their minds as one whole of which they knew all the parts. He
also thought it necessary that the character of all citizens should be well-
known, an impossibility in too large a community, but how else, he asked,
could men elect their magistrates wisely?
The duty of the State was, then, to ensure the possibility of a "good life"
to all its citizens. What was the Greek ideal of citizenship? First of all, every
citizen was expected to take a direct and personal share in all the affairs of
the State. To the Greek there was no separation between private and public
life, all things concerning the State were his affairs, and it was expected that
everyone should have an opinion of his own, that he should think clearly on
all matters of common interest and not allow himself to be swayed by his
feelings without honestly thinking the matter out, and to a Greek, thinking
meant straight thinking, the power to know right from wrong, to judge justly
without prejudices or passion, to separate the important from the
unimportant, and to follow undismayed wherever the truth might lead.
This belief in the duty of the citizen to be personally active in the affairs
of the State tended to keep the State small, for if every citizen was to attend
the meetings of the Assembly, the latter must be of such a size that everyone
could be heard if he desired to speak, and it was necessary that a very short
journey should bring the country-dweller into the city to attend to the State
business, for frequent journeys and long absences from his farm or his flocks
would be impossible for the countryman.
Further, the Greek believed that wealth was allowed to a man only as a
trust. Certain privileges and rights came to him because of its possession, but
they were privileges and rights that required of their owner distinct duties.
The more a man had, the more did the State require of him; he had to give
his time to the making of laws, his wealth built ships, bore the expense of
public festivals, adorned the city with beautiful buildings, it was spent not on
himself alone, but shared with his fellow-citizens, and given to that which
was their common interest. This resulted in a passionate devotion of every
Greek to his city, for every individual had a definite share in some way or
other in the making of it, and by the sacrifice of his life in times of danger,
he proved again and again that he was in very truth ready to die for it.
The ideal city demanded very high standards of her citizens, and no
Greek State attained these perfectly. But in their search for what they
conceived to be the highest perfection, the Greeks found out truths both
concerning government and the real meaning of citizenship that have
remained one of the priceless possessions of mankind.
[1] The Politics of Aristotle. See p. 384.
CHAPTER V
SPARTA[1]
Of all the city-states in Greece, two stand out from the others as having
played the leading part in Greek history. These two are Athens, which most
nearly approached the ideal city-state, and Sparta, the foremost military state
in Greece.
Sparta was the chief city in the peninsula in the south of Greece called the
Peloponnesus, or the Land of Pelops, one of the ancient mythical heroes of
that land. During the period of the Greek migrations, the Peloponnesus was
probably the last district to be settled, and the tribes which came down into it
were called the Dorians. They invaded Laconia, of which the chief city was
Sparta, and settled it and the surrounding country. Only the citizens of the
city itself were called Spartans; those who owed allegiance to the city, but
who lived in the country outside were known as Lacedaemonians.
Unlike most other Greek cities, which were well fortified and defended
by a citadel built on high ground, Sparta, "low-lying among the caverned
hills," had grown out of a group of villages and had no walls. But if she had
no outward signs of security from her foes, she had the mightiest warriors in
Greece ready to defend her to the utmost. Lycurgus, the wise man of Sparta,
was once consulted as to whether it would not be a wise thing to build a wall
round the city, and he answered those who came to him with the words: "The
city is well fortified which hath a wall of men instead of brick."
The Spartans were always afraid of attack from their slaves. These had
been the former inhabitants of the land and had been conquered and made
slaves by the Spartans. These slaves were called Helots; they were severely
and often cruelly treated by their masters, and were always ready to revolt
when opportunity came. This was one of the reasons that made the Spartans
spend more time than other Greeks in military training. This common danger
also had the result of drawing all Spartans very closely together, and of
making them subordinate all other interests to the supreme duty of protecting
the state.
I. THE GOVERNMENT OF SPARTA: LAWS OF LYCURGUS
The Spartans always held Lycurgus to have been their great lawgiver,
though they never agreed as to when he lived. It is probable that the laws
were not made by one single lawgiver, but that many wise men, whose
names were unknown to later generations, had helped to create the laws over
a long period of time. As Lycurgus was the wise man whose name was
known to the Spartans, they came to look back to him as their only lawgiver,
because it was by the government he was supposed to have established and
the laws he made that Sparta became so great a state. He was the brother of a
King of Sparta who died leaving a child as his heir. Everyone thought that
Lycurgus would take the opportunity to seize the throne and make himself
King, but he declared that he would only rule until his nephew should be old
enough to become King. Some people, however, would not believe this, and
plots were made against his life. This decided Lycurgus to leave Sparta for a
time and to visit some other countries with the intention of learning things
from them that might be of use to his own land. He went first to Crete,
where,
having considered their several forms of government, and got an
acquaintance with the principal men amongst them, some of their laws he
very much approved of and resolved to make use of them in his own
country. From Crete he sailed to Asia, with design, as is said, to examine
the difference betwixt the manners and rules of life of the Cretans who
were very sober and temperate, and those of the Ionians, a people of
sumptuous and delicate habits, and so to form a judgment. Lycurgus was
much missed at Sparta, and often sent for, "for kings indeed we have," the
people said, "who wear the masks and assume the titles of royalty, but as
for the qualities of their minds, they have nothing by which they are to be
distinguished from their subjects."
Lycurgus only, they said, knew both how to rule and how to make the people
obey him.
So Lycurgus returned, but on his way he went to Delphi to consult

Apollo, and the oracle called him "beloved of God," and said to him:
Lo, thou art come, O Lycurgus, to this rich shrine of my temple,
Beloved thou by Zeus and by all who possess the abodes of Olympus.
Whether to call thee a god, I doubt, in my voices prophetic, God or a
man, but rather a god I think, O Lycurgus.[2]
The oracle then went on to say "that his prayers were heard, that his laws
should be the best, and the commonwealth which observed them the most
famous in the world."
On his return to Sparta, the first thing Lycurgus did was to reform the
government. This was now to consist of three parts: the Kings, the Senate or
Council, and the Assembly. Sparta had always had two Kings at a time, who
succeeded each other from father to son. It was they who had the right of
commanding the army in war, they were always accompanied by a
bodyguard of specially picked men, at the public sacrifices and games they
had special seats of honour, and at all banquets and feasts double portions of
everything were served to them. When it was necessary for the State to
consult the Oracle, it was one of the Kings who decided on the messenger to
be sent.
The Senate or Council consisted of the two Kings who were members by
right of their birth, and of twenty-eight other men who were elected as
Senators for life. Every candidate had to be sixty years old, for Lycurgus
believed that until a man had reached that age, he was not wise enough nor
fit enough to be
entrusted with the supreme authority over the lives and highest interests
of all his countrymen. The manner of their election was as follows: the
people being called together, some selected persons were locked up in a
room near the place of election, so contrived that they could neither see
nor be seen, but could only hear the noise of the assembly without; for
they decided this, as most other affairs of moment, by the shouts of the
people. This done, the competitors were not brought in and presented all
together, but one after another by lot, and passed in order through the
assembly without speaking a word. Those who were locked up had
writing tables with them, in which they recorded and marked each shout
by its loudness, without knowing in favour of which candidate each of
them was made, but merely that they came first, second, third, and so
forth. He who was found to have the most and loudest acclamations was
declared senator duly elected. Upon this he had a garland set upon his
head, and went in procession to all the temples to give thanks to the gods.
The duties of the Senate were to prepare all the laws and matters of
public interest which were to be brought before the general Assembly; it
acted as a court of justice for criminal cases, and its deliberations carried
great weight.
Every Spartan citizen over thirty years of age was a member of the
Assembly. It was the duty of the Spartans in the Assembly to give or
withhold approval of all the matters brought before them by the Senate. It
was they who elected the Senators and all other magistrates, and they
declared war and made peace. In spite of this Assembly of citizens, the
government of Sparta was really in the hands of the Senate, for the members
of the Assembly might not discuss the laws submitted for their approval, but
only ratify or reject them. At no time were the ordinary citizens given much
opportunity to speak at length in public. The Spartans did not like long
speeches, and Lycurgus believed that no one should be allowed to talk much
unless he could say a great deal that was useful and to the point in a few
words. This way of talking became so characteristic of the men of Laconia,
that it was called by their name, and even today speech that is short and
sharp and to the point is called laconic. Many stories are told of this Spartan
manner of speech. King Leonidas said once to a man who was speaking
about some important matter, but at the wrong time and place, "Much to the
purpose, Sir, elsewhere"; and one who kept pressing for an answer to the
question "Who is the best man in Lacedaemon?" received as his answer:
"He, Sir, that is least like you."
Long after the death of Lycurgus, another special body of men was
elected to help in the government. These men were called Ephors, and there
were five of them. It was their business to watch the conduct of the Kings, to
see that the laws were all carried out and that order and discipline were
maintained in the state. Probably no other Greek state would have submitted
to such oversight, but the Spartans were well-disciplined and did not look
upon such an office as one that interfered in any way with their personal
freedom. Plato, writing long after of their authority, said that it was,
"exceedingly like that of a tyrant."
II. CUSTOMS IN SPARTA
Having established the government, Lycurgus next set himself to

introduce what he considered good customs amongst the Spartans. The first
thing he did was to re-distribute the land amongst the citizens. He found that
some were very rich and others poor, and he determined that they should all
live together sharing in everything alike. So he divided the land into lots and
distributed it equally amongst all the citizens. After this had been done, and
the time of harvest had come, Lycurgus, "seeing all the stacks standing equal
and alike, smiled, and said to those about him, 'methinks all Laconia looks
like one family estate just divided among a number of brothers.'"
During his travels in other parts of the world, Lycurgus had had
opportunities to compare rich states with poor ones, and he had concluded
that the richest were not always the best governed, and that wealth did not
always bring happiness. He was determined that the Spartans should become
good soldiers and that they should be great in war, for he believed that
simple habits and simplicity of living were more easily acquired by a nation
of warriors than by one devoted only to peace. To this end he wanted to have
neither rich nor poor in Sparta, but that all should be alike. Lycurgus,
however, was wise enough to know that some people would always manage
to make more money or to save more than others, so instead of dividing up
all the money in the state equally between everyone, he
commanded that all gold and silver coin should be called in, and that only
a sort of money made of iron should be current, a great weight and
quantity of which was but very little worth; so that to lay up twenty or
thirty pounds there was required a pretty large closet, and, to remove it,
nothing less than a yoke of oxen.
Not only did this prevent any one man from becoming too rich, but it
lessened the crime of theft in Sparta, for no one would want to steal what
was of no value.
This lack of money also resulted in doing away with luxuries; for
there was no more means of purchasing foreign goods; merchants sent
no shiploads into Laconian ports; no gold or silversmith, engraver or
jeweller set foot in a country which had no money, so that luxury wasted
to nothing and died away of itself.
Everything needed in their houses had to be made by the Spartans
themselves, with only the simplest tools, and the houses were roughly built.
The law required that the
ceilings of their houses should only be wrought by the axe, and their gates
and doors smoothed only by the saw, and as no man would furnish such
plain and common rooms with silver-footed couches and purple coverlets
and gold and silver plate,
all Spartans grew accustomed to the plainest and simplest surroundings.
It is reported that one of their Kings was so little used to the sight of
any other kind of work, that being entertained at Corinth in a stately
room, he was much surprised to see the timber and ceiling so finely
carved and panelled, and asked his host whether the trees grew so in his
country.
The last law made by Lycurgus to ensure simplicity of living was that all
Spartan men and youths should eat at common dining-tables, and they were
only allowed to eat such food as was permitted by the law. Each table seated
about fifteen men, who shared in providing the food; each of them was
"bound to bring in monthly a bushel of meal, eight gallons of wine, five
pounds of cheese, two and a half pounds of figs and some very small sum of
money to buy flesh and fish with." All the food was prepared in a very
simple manner, but "their most famous dish was the black broth, which was
so much valued that the elderly men fed only upon that, leaving what flesh
there was to the younger." Other Greeks, however, thought this black broth
very disagreeable.
It was the custom that at any one table, only those should sit who were
friendly to each other. When a newcomer wanted to join a certain table, all
those already seated at it voted as to whether they would have him or not. An
urn was passed round the table and everyone present dropped into it a small
ball of bread. Those who voted for the newcomer dropped their balls without
altering their shape, those who voted against him flattened the ball with their
fingers before placing it in the urn. One flat ball was enough to exclude a
man from the table.
When dinner was over, "every man went to his home without lights, for
the use of them was on all occasions forbidden, to the end that they might
accustom themselves to march boldly in the dark."
III. SPARTAN EDUCATION
Lycurgus was determined that every Spartan should be so trained that he

might become a good soldier, and some of his most important laws
concerned the education of children. As soon as a child was born, he was
carried to
the elders of the tribe to which he belonged; their business it was
carefully to view the infant, and if they found it stout and well-made, they
gave order for its rearing, but if they found it puny and ill-shaped, ordered
it to be taken to a cavern on Mount Taygetus, where it was left to perish,
for they thought it neither for the good of the child itself, nor for the
public interest that it should be brought up, if it did not, from the very
outset, appear made to be healthy and vigorous.
There was a belief in Sparta that wine was more strengthening than water for
a bath, and so the first bath a baby had was always in wine.
There was much care and art, too, used by the nurses; the children
grew up free and unconstrained in limb and form, and not dainty and
fanciful about their food; not afraid in the dark, or of being left alone;
without any peevishness or ill-humour. Upon this account, Spartan nurses
were often hired by people of other countries.
At the age of seven, Spartan boys left their homes and their mothers, and
the State took charge of the rest of their education.
As soon as they were seven years old they were to be enrolled in
certain companies and classes, where they all lived under the same order
and discipline, doing their exercises and taking their play together. Of
these, he who showed the most conduct and courage was made captain;
they had their eyes always upon him, obeyed his orders, and underwent
patiently whatsoever punishment he inflicted; so that the whole course of
their education was one continued exercise of a ready and perfect
obedience. Reading and writing they gave them, just enough to serve
their turn; their chief care was to make them good subjects, and to teach
them to endure pain and conquer in battle.
As they grew older, Spartan boys were taught to undergo all kinds of
hardships. They wore very little clothing, even in the cold of winter, and one
coat had to serve them for a year.
After they were twelve years old they lodged together in little bands
upon beds made of the rushes which grew by the banks of the river
Eurotas, which they were to break off with their hands without a knife.
The old men had an eye upon them at this time, coming often to the
grounds to hear and see them contend either in wit or strength with one
another, and this seriously and with much concern; so that there scarcely
was any time or place without someone present to put them in mind of
their duty, and punish them if they had neglected it.
Besides all this, there was always one of the best and honestest men in
the city appointed to undertake the charge and governance of them; he
again arranged them into their several bands, and set over each of them
for the captain the most temperate and boldest of those they called Irens,
who were usually twenty years old. This young man, therefore, was their
captain when they fought, and their master at home, using them for the
offices of his house; sending the oldest of them to fetch wood, and the
weaker and less able, to gather salads and herbs and these they must
either go without or steal; which they did by creeping into the gardens, or
conveying themselves cunningly and closely into the eating-houses; if
they were taken in the act, they were whipped without mercy, for thieving
so ill and awkwardly. They stole, too, all other meat they could lay their
hands on, looking out and watching all opportunities, when people were
asleep or more careless than usual.
If they were caught, they were not only punished with whipping, but hunger
too, for they were then reduced to their ordinary allowance, which was
purposely kept very small, in order to force them to use cunning and skill if
they wanted to add to it. "So seriously did the Lacedaemonian children go
about their stealing, that a youth having stolen a young fox, and hid it under
his coat, suffered it so to tear with its teeth and claws, that he died rather
than let it be seen."
In every way the Spartan youths as they grew up were severely

disciplined. Every year the older boys were whipped in public before the
altar of Artemis, in order to teach them to endure pain without crying out,
and it is said that some boys died under this whipping rather than utter a
complaint.
The Iren used to stay a little with them after supper, at which time he
would bid one of them sing a song, to another he put a question which
required an advised and deliberate answer; for example, Who was the
best man in the city? What he thought of such an action of such a man?
They accustomed them thus early to pass a right judgment upon persons
and things, and to inform themselves of the abilities or defects of their
countrymen. Besides this, they were to give a good reason for what they
said, and in as few words as might be; he that failed of this, or answered
not to the purpose, had his thumb bit by his master. Sometimes the Iren,
did this in the presence of the old men and magistrates, that they might
see whether he punished justly or not; and when he did amiss, they would
not reprove him before the boys, but, when they were gone, he was called
to account, and underwent correction, if he had run far into either of the
extremes of indulgence or severity.
Furthermore, in his desire firmly to implant in their youthful souls a

root of modesty Lycurgus imposed upon these bigger boys a special rule.
In the very streets they were to keep their two hands within the folds of
the cloak; they were to walk in silence and without turning their heads to
gaze, now here now there, but rather to keep their eyes fixed upon the
ground before them. And you might sooner expect a stone image to find a
voice than one of those Spartan youths; to divert the eyes of some bronze
statue were less difficult.[3]
Not very much time was spent by the boys in learning to read and write;
most of their education was given to their gymnastic training, to running,
jumping, boxing and wrestling, and to every kind of exercise that would fit
them to be brave and hardy soldiers. They learnt some music, chiefly
singing, but they only sang such songs as would put life and spirit into them,
and their battle songs were sung with great enthusiasm.
During a war, the Spartan young men were treated a little less severely
than when in training at home. They were allowed to curl and adorn their
hair, to have costly arms and fine clothes. They had better clothes, too, and
their officers were not so strict with them. They marched out to battle to the
sound of music. "It was at once a magnificent and terrible sight to see them
march on to the tune of their flutes, without any disorder in their ranks, any
discomposure in their minds or change in their countenance, calmly and
cheerfully moving with the music to the deadly fight."
Spartan discipline did not end when the boys and youths had become
men.
Their discipline continued still after they were grown men. No one
was allowed to live after his own fancy; but the city was a sort of camp,
in which every man had his share of provisions and business set out, and
looked upon himself not so much born to serve his own ends as the
interest of his country.
A GIRL RACER.
Early 5th Century B.C
Vatican, Rome.
The girls were educated at home, but, like the boys, they were given a
gymnastic training, and they learned to run and wrestle, to throw the quoit
and dart, and to be as strong and brave as their brothers. As the Spartan boys
were trained to become good soldiers, ready to die for Sparta, so were the
girls trained to become good wives and mothers of Spartan men, and if they
could not themselves die for their country, to be willing to sacrifice those
whom they loved the best. Every Spartan when he went to war, carried a
shield that was so heavy that if he fled from the enemy, he would have to
throw it away, so it was considered a great disgrace to return home without
one's shield; if he died in battle the Spartan was carried home upon it. The
Spartan mothers knew this, and when they said farewell to their sons who
were setting out to war, they bade them return home with their shields or
upon them.
The Spartans held their women in great honour; they listened to their
counsel and often acted upon it. A lady of another city once said to a
Spartan, "You are the only women in the world who can rule men." "With
good reason," was the answer, "for we are the only women who bring forth
men."
Having established all these laws and customs, Lycurgus forbade the
Spartans to travel, for he was afraid that contact with foreign people would
teach them bad habits and make them discontented with their simple way of
living. "He was as careful to save his city from the infection of foreign bad
habits, as men usually are to prevent the introduction of a pestilence."
At last the time came when Lycurgus felt that his laws and customs were
firmly established, and that they were all familiar to the people, but he was
afraid that after his death they might be changed. So he thought of a plan
whereby he might make them last forever. He called a special Assembly of
the people together and told them that everything was well-established, but
that there was still one matter on which he would like to consult the Oracle.
Before he departed on this journey, he made the two Kings, the Senate and
the whole Assembly take a solemn oath that they would observe his laws
without the least alteration until his return. "This done, he set out for Delphi,
and having first offered a sacrifice to Apollo, he asked the god whether his
laws were good and sufficient for the happiness and virtue of his people."
The Oracle answered that the laws were excellent, and that the state which
kept them should be greatly renowned. Lycurgus sent this oracle in writing
to Sparta, and then having once more offered a solemn sacrifice, he took
leave of his friends, and in order not to release the Spartans from the oath
they had taken, he put an end to his own life, thus binding them to keep his

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The Mechanism of Nervous Action:

Electrical Studies of the Neurone E. D.

The Integrative Action of the Autonomic Nervous System

Studies in the History of Science E. A. Speiser

Montfort History Early Research and Recent Studies of

Diseases of the Nervous System, 2nd Edition Harald

Glycobiology of the Nervous System 2nd Edition Cara-

Primary Mathematics 3A Hoerst

The Nervous Stage Nineteenth century Neuroscience and

Aids to the Examination of the Peripheral Nervous

Foul erton Research Professor oj the Royal Society

Published in Great Britain, India, and Pakistan

Printed and Bound in the United State* of America by

inviting me to give the second course of Eldridge Reeves

I. THE DEVELOPMENT or ELECTROPHYSIOLOGY

Potential Changes in the Brain Stem 81

THE DEVELOPMENT OF ELECTRO-

N London we have just celebrated the hundredth anni-

years, until 1792, it seemed that Galvani's phenomena were

shortest cut to an understanding of life. I t attracted men

with the string galvanometer it was possible to record the

t h e p o t e n t i a l c h a n g e is m u c h larger t h a n when the u n i t s

Amplification of Electric Changes

For some experiments the ideal instrument is the inertialess

there is some hope of reducing them, but there must

The work with which I shall deal is mainly concerned

be overcome before long—some methods are already

decapitated under an anaesthetic and the body is kept

records show the potential changes which occur between

at the toe joint, and that the sensory message consists of

Fio. a. Production of a diphasic potential wave by the passage of an impulse

The Activity of Single Nene Fibres

theory and the most difficult in practice is to cut through

recording the potential changes in the nerve. The potential

clearly marked in the discharge of sense organs which

If these records give a true measure of the activity in

on the motor nerve fibres by the excited nerve cell in the

F i c . 6. Impulses in a motor nerve supplying the tail muscles in a caterpillar

T o complete the series and to show t h a t the behaviour

the frequency of the waves declines instead of rising as in

FJG. 7. Impulses in central nerve cord of a caterpillar. A portion of the

The Nature of the Nervous Messages

It is of course impossible to prove that electric changes

would be arranged in a fairly regular and uniform sequence,

impulse. Surely each impulse in a message may have a

activating other regions. These chemical and thermal

stimulus, when successful, produces not a second dis-

REFRACTORY ACTIVE RESTING

DIRECTION OF TRAVEL >

FIG. 8. Spread of active (depolarised) area according to the membrane

of breakdown shifts down the nerve fibre; the potential

We come back then to our records of nervous messages

THE ACTION OF THE SENSE ORGANS

Ε sensory discharge is something which can be

With the hair organ a lateral movement of the tip of the

Adaptation is perhaps a dangerous term to use, for it

in the majority, in mammalian muscles both are present

frequency of the discharge falls appreciably in the first