ELECTROENCEPHALOGRAPHY AND CLINICAL NEUROPHYSIOLOGY
THE ELECTROENCEPHALOGRAM
OF ASCENDING INFLUENCES
OF THE
P. K. ANOKHIN
Sechenov Institute of Physiology of the First Moscow Medical Institute, Moscow (U.S.S.R.)
The character of the slow electrical activity of
the electroencephalogram is a problem that has
become increasingly acute in recent years. This
tendency to investigate the true physiological
sense of the slow electrical potentials of the cor-
tex is natural enough. The electrical indices of
cortical activity have been tacitly accepted as
direct reflections of cortical neuronal processes.
It seemed quite evident that everything occurring
in the cortical mass, in the interrelations between
its cellular components, was directly reflected
in the EEG. This premise is quite evidently a
result of the history of the development of our
conceptions of the functions of the central nerv-
ous system.
From the very beginning researchers have
attempted to analyze exterior symptoms of be-
haviour on an anatomical level, to discover what
nervous processes develop and where they occur.
The schematic conception of the reflex arc, first
published by Rend Descartes, is the first attempt
at projecting, on a nervous substratum, the
peculiarities of behaviour in man and animals.
Later, when the microscope had revealed a
whole world of diverse connections and cells in
the nervous system, the interpretation of nerv-
ous processes immediately rose to the level of
the finest neuro-anatomic relationships. It is
sufficient to recollect the enormous role of the
synapses which determine the whole process of
selective formation of specific activities at every
instant.
Here we come across a remarkable historical
paradox: the conception of nervous activity on a
fine neuronal level must somehow assimilate the
new technical achievement of electroencephalo-
graphic study of the brain. The fact that a direct
General summation of: The neurophysiological
bases of the EEG.
27
AS A RESULTANT
ON THE CELLS
CORTEX '
connection has been established between the
EEG and the conception of nervous activity on
the neuronal level must perforce be regarded as
quite natural.
The summary character and ext~'eme gener-
alization of the electrical phenomena compos-
ing the E E G have been revealed to, a growing
extent by concrete data. Research )~as made it
ever clearer that slow electrical activily cannot be
the reflection of an actual neurological process,
either in respect to its speed, or in respect to the
selective interaction of individual cells. Special
lines of research were thus defined in what 1 will
venture to call the critical period in the develop-
ment of electroencephalography (Ft,ssard, Jas-
per, Mountcastle, Anokhin and others).
The present meeting has very o,rrectly set
this problem for profound discu:~sion. The
measure must be regarded as quite timely, since
we are faced with the dangerofgrou~dlessopin-
ions on brain activity, distant from tl,c truth, as
a result of the broad use of EEG in physiological
laboratories and in the clinic. There can be
little doubt that the success and importance of
EEG in different lines of science depel~d directly
on the final conception of slow electrical activity
of the cerebral hemispheres and other areas of
the brain. In short, we must establish which part
of the general nervous integration tetermines
slow electrical activity, how this activiz...~is formed
in the brain tissue itself, and what its correla-
tions are with the selective processes :~t the cel-
lular level.
The origin of numerous new lit:tes of re-
search, which could be classified under t he general
term of correlative research, becomes evident in
the light of all that has been said. This research
usually establishes the correlations o" ordinary
EEG indices with the discharges of individual
Electroenceph. clin. Neurophysiol., 1964. 16:27-43
28
nerve cells caused by potentials, with pharma-
cological factors and, finally, with individual
phases of the conditioned reflex and general
behaviour.
The present meeting is, to some extent, a
summary of the various researches in modern
neurophysiology. The collection of experimental
material was founded on the following method-
ological principles: (a) correlation between the
E E G and discharges in individual neurons; (b)
correlation between the E E G and grouped cellu-
lar discharges recorded from the same electrode;
(c) correlation between the EEG and evoked
potentials, both primary and secondary; (d)cor-
relation between the E E G and specific chemical
and pharmacological factors; (e) correlation be-
tween the E E G and conditioned reflex behav-
tour, formed under qualitatively different bio-
logical conditions; ( f ) the ontogenetic investiga-
tion of the maturation of the electrical signs of
nervous activity.
This meeting has shown that we have now
achieved considerable success in applying correla-
tive methods of research. I should like to give a
short summary of the material of today's meeting
and some information on the research done
in our own laboratory in connection with this
important problem.
I. CORRELATION BETWEEN SLOW ELECTRICAL
ACTIVITY AND THE F U N C T I O N A L STATE
OF I N D I V I D U A L NEURONS
The first correlative research on comparison
between slow electrical activity of the cerebral
cortex and the discharges of individual neurons
was conducted in the laboratories of Fessard,
Jasper, Mountcastle and Amassian. Judging
from the latest results reported at this meeting
by Jasper, we may definitely maintain the non-
existence of direct, linear dependence between
slow electrical activity and individual cell dis-
charges. From a neurophysiological point of view
this means that the processes of individual neu-
ronal activity forming the base for the organ-
ization of large functional systems of the organ-
ism can occur to a certain extent independently
of slow electrical activity.
It could hardly be said that these two aspects
of the process have no connection whatever, but
the considerable results of the research conduct-
t'. K. ANOKHIN
ed by Jasper and other scientists are particu-
larly important because they raise the questions
of what is the exact participation of slow electri-
cal aclivity in the work of individual nervous
elements and whether this slo~ activity is not
a sort of epiphenomenon, i.e., a summary by-
product of activity that has alretLdy occurred on
a fine molecular level. Here we are faced with
one of nature's riddles, which dc:idedly impedes
our advance in the cognition of h e general laws
of nervous activity with the aid of EEG meth-
ods of research.
We also have results, to a considerable extent
coordinated, in the work of Morazzi and Baum-
garten. With some variations, th~ results of their
research lead to the same conclusion: that there
is no direct, so to speak linear, connection be-
tween the two forms of activity. A,t the same time
there is one detail that has been revealed in the
work of all researchers: that s o w oscillations
with a definite amplitude have an undoubted
influence on the condition of 1he neuron and
lead to the intensification of its spontaneous dis-
charges on the highest point of the slow wave.
What does this mean? Does it nean that slow
activity promotes the activatio~ of the cellular
discharges taking part in some rervous process,
or is this only an accessory result of the level of
electrical potentials, playing no mbstantial part
in real interactions on a netLrophysiological
level?
Our latest research on slov activity and
grouped cellular discharges retorded simulta-
neously from the same point in t~e cortex of the
cerebral hemispheres throws s(,ne light on the
problem. This technique can easiy be applied by
using certain forms of electronic filtration of the
incoming potentials.
With the aid of this simple :tevice we were
able to show that cell activity has no definite con-
nection with any form of slow el:ctrical activity.
For example, in largegroups ofcelt.~discharges may
occur at the very moment of de :ynchronization
of slow activity, developing in:o the type of
regular slow rhythm at 4-7 c/sec, described by us.
This rhythm is known to corresF~ond to the gen-
eral tension of the animal under unfavourable
biological conditions; for the rabbit we called
it "stress rhythm" (Fig. 1, 2). Tile very fact that
discharges of cellular activity can occur at E E G
Electroenceph. clin. Neurophysh~'~., 1964, 16:27-43
 ASCENDING INFLUENCESON CORTEX 29

"']---J,-~,lO
I '...~ TONE "2"00 DgF~'NSl .........

i
i

R.E

(o)
L.S.M. (b)

RSM

T.l.

TR

O.k.

OR.

N.VP.L.L.

N.VRLR.

EK.G.

2OOpv[Is,c

Respir,
Reticular stimulation 1OO C/see 3v
P
Fig. I
(a)Response of corticalelectricalactivityto the conditioned
defensive stimulus (200 c/sec). It is seen that slow
electrical activity immediately changes to desynchroniza-
tion in sensori-motor cortex (SMC) and to the regular
stress rhythm in the non-specific thalamic and reticular
formation areas. THL, thalamus left; THM, thalamus Fig. 2
midline; RF, reticular formation. .General scheme of the disposition of regular stress rhythm
(b) Appearance of the regular stress rhythm during direct m cortical and subcortical structures. Th, thalamus;
stimulation of the reticular formation. LSM, left sensori- Hyp, hippocampus; RF, reticular formation. + indi-
motor; RSM, right sensori-motor; TL, thalamus left; TR, cates stress rhythm, - - indicates areas for preferential
thalamus right; OL, occipital left; OR, occipital right; desynchronization.
NVPLL, nucleus ventralis postero-lateralis left; NVPLR,
nucleus ventralis postero-lateralis right: EKG, electro- Eleetroenceph. clin. Neurophysiol., 1964, 16:27-43
cardiogram.
30 P. K. ANOKHIN

Fig. 3
Comparison of EEG and mass cellular activity at the same point in the reticular formatior. It is evi-
dent that the increase in frequency of the regular rhythm accompanies the increase in cellul tr activity
(a). Conversely, the cellular activity ceases when the regularity of the rhythm is disturbed ! b).

indices of different quality shows convincingly
that this rhythms as a species of slow activity,
can have no direct relation to the origin of cellu-
lar discharges. Nevertheless we were able to sep-
arate one parameter of this slow activity which
is more closely connected with the origin of cel-
lular discharges. Fig. 3 shows that, while the
rhythm is very regular, cellular activity remains
ort a high level, but, as soon as the regularity is
disturbed, firing of the cells immediately ceases.
The cells can be active only while the slow
rhythm remains absolutely regular, that is, while
the amplitude of the waves remains constant.
The question naturally arises as to what pre-
cedes this? Is the regular slow rhythm the result of
extreme celhdar discharge, or, on the contrary,
is the cellular discharge facilitated by a preceding
slow rhythm?
A comparison of the results of diverse exper-
iments conducted in our laboratory shows that
cellular discharges can hardly be the primary
factor in the several cases of these joint changes.
Cellular discharges occur without interruption
during a rather long period while the regu-
larity of the tension rhythm forHed in response
to painful irritation of the ski:.-, is maintained.
Meanwhile, on the background ,~f this constant
discharge, slow oscillations ha, e a rhythmical
character, i.e., their electrical index is consider-
ably altered.
The wide spread of the ~ension rhythm
through different areas of the brain is a particu-
larly strong argument against ~, direct depend-
ence between regular slow rh~lhm and the ac-
tual neurophysiological proce.~ses selectively
developing at a neuronal level. For example, if
you touch the hind leg of a rabbit with the point
of a needle, the tension rhythm is immediately
manifested with remarkable precision in several
areas of the brain. It appears in the reticular
formation, the hypothalamu~, the medial
thalamus, the hippocampus, lhe parietal and
occipital fields of the cortex At the same
moment, desynchronization develops in other
Electroenceph. clin. Neurophysi~,l., 1964, 1 6 : 2 7 4 3
 ASCENDING INFLUENCES ON CORTEX
parts of the brain, for instance, in several parts
of the reticular formation, in the lateral thalamus
and in the sensori-motor field of the cortex. If we
study the condition of mass cellular activity in all
these areas, we shall find everywhere an inten-
sification of cellular discharges, independent of
the different indices and the quality of slow elec-
trical activity (Banzekina 1962).
Yet another consideration leads us to think
that slow activity develops in individual ways
and has its own place in the total sum of elec-
trical activity of the cortex. Looking at the wide
field over which regular slow rhythm is spread
(see Fig. 2) we involuntarily ask ourselves: how
can the same slow rhythm exist in these hetero-
geneous areas of the brain, where individualized
nervous interactions and integrations occur? On
the other hand, we cannot deny that this rhythm
does play some role, to us still incomprehens-
ible, in the interactions of the general work of the
brain.
It is interesting that, if we study the border
between the zone of desynchronization and the
zone of regular slow rhythm, or tension rhythm,
we can find a whole set of transitory forms of
electrical activity which can be separated into its
component parts with the aid of an analyser.
We have tried to analyse the nature of the
desynchronization of cortical electrical activity
occurring after painful irritation of an animal in
the waking state. If a rabbit is given a light ure-
thane narcosis, desynchronization of the sen-
sori-motor field will be replaced by the regular
rhythm which is characteristic for the fields de-
scribed above, but in this case cellular activity is
not so considerably altered. It is therefore ab-
solutely evident that the subcortical mechanism
forming the regular rhythm of individual areas
of the cortex escapes the influence of anaesthet-
ics, while the desynchronization mechanism of
the sensori-motor region can be blocked by
them.
In conclusion to this set of experiments we
may say that regular slow rhythm, which is one
of the forms of slow electrical activity of the cere-
bral cortex~ cannot be directly connected with
inter-neuronal activity. On the other hand this
rhythm, which occurs simultaneously in wide
expanses of the brain tissue, undoubtedly pro-
motes inter-neuronal activity to some extent and,
31
according to data on hand, has its own mecha-
nism of originating and spreading through the
mass of the brain.
We are especially inclined to advance this
hypothesis in view of the results of experiments
conducted in the laboratory of Moruzzi and
reported at this meeting.
11. E V O K E D P O T E N T I A L S OF T H E C E R E B R A L C O R T E X
A N D THEIR I M P O R T A N C E F O R THE D I S C L O S U R E O F
THE N E U R O P H Y S I O L O G I C A L BASIS OF THE EEG
The evoked: potential is perhaps the outstand-
ing electrical phenomenon of the central nerv-
ous system and especially of the cortex. Owing
to its constancy and rather exact dependence on
the intensity and quality of stimulation, this
phenomenon has been rightly used on a wide
scale to explain the nature of electrical poten-
tials in general and cortical electrical activity in
particular (Dempsey, Morison, Forbes, Bishop,
Grundfest, Purpura and others). On the other
hand, some of its peculiarities, described in
different recent works, arouse some scepticism
in regard to its participation in what we now
call "real nervous activity".
I shall enumerate the peculiarities o f the
evoked potential which cause some d~,ubts as to
its being an important component ~,f nervous
activity in general and of cortical ~tctivity in
particular.
(a) The evoked potential of the comex is well
defined when the animal is sunk in deep narcotic
sleep, that is when, according to our concep-
tions, the work of establishing relatic~ns of the
organism with the environment, which is most
characteristic of the cortex, ceases c,,mpletely.
(b) The amplitude of the evoked p~tential is
considerably reduced, and the poter~tial may
even completely disappear, at the moment of
activation of the cortex, i.e. at the very time
when the latter manifests its activit3 with the
greatest energy.
(c) The configuration of the evoked notential,
consisting principally of a complex o" positive
and negative oscillations, is extraordi~ arily ho-
mogeneous in extremely different aninlals and
even in nervous formations of different struc-
ture. The biphasic character of the superficial
evoked potential is found in amphibi~tns, birds
and lower mammals and it can be e~tablished
Electroenceph. c/in. Ne,rophysio/~. 1964. I6:27 43
32 P. Ko ANOKHIN
in the cortex, the hippocampus and hypothala-
mus.
All these physiological peculiarities of the
evoked potential unavoidably lead to the ques-
tion: what does the evoked potential actually
represent and what place does it occupy in the
mechanisms of formation of the summated
electrical activity of the cortex?
The ontogenetic view point, represented here
in the very interesting morpho-physiological
report of Purpura, is a very important way of
elucidating the neurophysiological foundations
of the evoked potential. The process of matura-
tion of electrical phenomena is especially con-
venient for the establishment of the nature of
the complicated phenomena of nervous activity
with which we have to deal in our study of the
adult organism.
The ontogenetic development of the nervous
system is governed by definite laws, which have
been well studied in our laboratory in the course
of the last thirty years. The first and most im-
portant of these laws, which governs the process
of maturation of all the vitally important func-
tions necessary to the newborn animal, is that
maturation of nervous and other structures is a
selective process, proceeding within the frame-
work of a large system of relations, the whole
forming a definite functional system which,
together with other systems, helps the newborn
organism to remain alive.
We have applied to this development, selec-
tive as regards time and structure, the term
s)'stemogenesis, since the final function of the
newborn is the directing factor in the founda-
tion, growth and consolidation of diverse struc-
tures in embryogeny (Anokhin 1933, 1937, 1948,
1958).
In the light of these concepts, no morpholog-
ical substratum, in particular the structures of
the brain, can ripen °'in general" in ontogenesis.
They will unavoidably reflect the degree of im-
portance and necessity of the individual struc-
tures and functions at different stages of the
post-natal development of the given species. In
other words, not individual organs or structural
units, but all the numerous fragments of the
given functional system develop in complete
coordination.
From this point of view Purpura's research
is of exceptional interest. Electron-microscopic
data have shown that between the maturation of
the pleximorphic layer and the layer of pyrami-
dal cells there is a definite interval: at the time
when the synaptic formations of the pleximor-
phic layer have already matured, the axosomatic
synapses, i.e., in the fourth layer, are still in the
stage of maturation. This heterochronic devel-
opment of the synaptic and ti)~rous structures
of different layers of the cortex has undoubtedly
a deep historical significance. If we take into
consideration Herrick's data (1948) on the struc-
ture of the brains of amphibia and the physio-
logical synthesis of these data ~ecently made by
Bishop (1958) and Grundfest (1959), the im-
portance of the morphological results obtained
in Purpura's research will become quite evident.
Thus, the fractional charm:ter and conse-
quences of the maturation process in individual
microscopic structures of the .:ortex can be a
very important means of studying separately the
cortical processes which, in an ~dult animal, are
already an organic part of eleclro-physiological
complexes.
Electro-physiological researca on evoked po-
tentials and slow electrical activity immediately
showed this heterochronic gr~,wth of cortical
structures. The research condtcted by Sherrer
and Von Economo has alread~ shown that the
negative phase of the evoked potential appears
first in ontogenesis. The gradutl growth of the
preceding positive deviation caanot be detected
for 6-8 days (Sherrer and 'Con Economo).
Unfortunately, no connection was drawn be-
tween this paradoxical fact ard the nature of
cortical electrical activity in gereral, or the pre-
vailing theory of evoked pote1~tials, elaborated
mainly by Eccles (1953).
The systematic work of out laboratory and
that of Purpura enables us to understand
the true physiological sense of superficial
cortical evoked potentials. Purpura has shown
that, directly after birth, the process of matura-
tion of the evoked potential proceeds in direct
connection with the growth ard maturation of
cortical synaptic formations in tifferent areas of
the cortex. Until the corresponding synapses are
mature and can, consequently, undertake exci-
tation and develop subsynaptic potentials, they
cannot produce the corresponding form of elec-
Electroenceph. clin. Neurophysiol., 1964, 16:27~,3
 ASCENDING INFLUENCES ON CORTEX
trical potential. These initial signs of evoked
potentials enable us to throw light not only on
the nature of the potential itself, but also on the
electrical activity of the cortex called "spon-
taneous".
Purpura's laboratory and our own have been
in sufficient accord in showing that the resolu-
tion in time of the different components of the
evoked potential occurs in ontogenesis, and that
the process of resolution depends directly on the
consecutive maturation of the synaptic organi-
zations of the pleximorphic and fourth layers of
the cortex.
This decisive conclusion, which can be
reached on the strength of factual data, enabled
us to develop the hypothesis of the dual neuro-
physiological nature of the evoked potential.
We believe that the initially maturing plexi-
morphic layer is the substratum forming the
negative phase of the evoked potential, while
the positive phase, which appears later, is the
product of post-synaptic potentials, formed on
the axosomatic synapses of the fourth layer of
the cortex. The substance of our conception is
that the evoked potential is formed on the basis
of a double message of ascending excitation.
The first message consists of excitations develop-
ing in a corLduction route that has matured ear-
lier and sends fine fibres directly to the pleximor-
phic layer and apical dendrites of the cortical
neurons. This substratum can he the analogue of
ancient structures of the anterior brain when it
had widely developed neuropi! structures. The
second message, which develops later in phyloge-
nesis and matures later in ontogenesis in a synap-
tic relation, is directed to the axosomatic synap-
ses of the fourth layer. These latter fibres are
thicker in diameter and are a typical example of
a specific thalamic radiation, conducting excita-
tion considerably quicker than the first system in
the pleximorphic layer. This is the explanation
which must be given for the fact that, approxima-
tely on the tenth day of post-natal life, the positi-
ve potential appears before the negative. After a
whole series of control experiments involving
chemical action and excitation by conjugate stim-
uli we have been still more strengthened in the
opinion that individual phases of the evoked
potential have different origins. It seems to me
that Purpura's excellent experiments, reported
33
250ms.
i
Fig. 4
This figure shows the importance of interxals for ascer-
taining the phases of paired evoked potentials of the
cerebral cortex in a 15 day old rabbit. (a! The interval
between two stimuli to sciatic nerve equals 130 reset.
The arrows show the points of single stimulation of
sciatic nerve (first and second). It is evident that the second
stimulus gives neither positive nor negative phase. (b) The
same conditions, but the interval between stimuli is
increased to 250 msec. It is evident that in response to the
second stimulus only the positive phase of the evoked
potential appeared. On the contrary, at an earlier stage
of ontogenesis, using the same pairing of stimulation,
only the negative phase of the evoked potential appeared.
here, also speak in favour of this hypothesis.
I can show several figures giving us further
confirmation of this concept (Fig. 4). This shows
that, by pairing two individual stimuli with a
definite interval between them, we can obtain a
result in which the negative potential only will
be absent in the second evoked potential, while
the positive will be present. These correlations
could not take place if both potentials had the
same neurophysiological origin (Anokhin 1958;
Sun Ven 1963). The extreme sensibility of the
negative component of the evoked potential
to various chemical influencesmust also be referred
to it. It is known from the results previously
published by usthaturethanenarcosis completely
blocks the negative components of the initially
evoked potential, leaving, however, little change
in the positive component (Ata-Muradova ! 960).
Experiments with local application ofgamma-
aminobutyric acid can be referred to the same
set of proofs of the special natur~' and spe-
cial origin of the negative componenl: of the in-
duced potential. It is known, from 1:he experi-
ments of Purpura and of my colleagues, that the
use of this substance causes the immediate re-
moval of the negative component, which is
another strong proofk>f the specific peculiarity
of these synaptic formations.
Eleetroeneeph. elin. Neurophysiol., 1964, 16:27-43
34 P . K . ANOKHIN

(b)

/ a 8
-d.

/,, !,:.. /;:

itlf
L
S S S

IVs Ns Us Ns % Ns 5 Ns ~ Ns
Fig. 5
A synthetic scheme showing the progressive stages of maturation of the synaptic organizations on the
cells of different layers of the cortex, a, b, c - - different degrees of appearing surfaces of e',,oked po-
tentials; P.L. - - plexiform layer; S,S - - specificascending pathways; Ns, Ns - - non-specific :~scending
pathways. (a) 3rd day after birth; (b) 10th day after birth. Progressive growth of synaptk: connec-
tions from primary epicentres in the plexiform layer is shown.

It is interesting to note that the growth of
the ascending and conducting paths and the
establishment by them of synaptic connections
in the pleximorphic layer proceed in a definite
sequence governed by a definite law. A very
narrow route, perhaps represented by a few
fibres, is first organized. On reaching the plexi-
morphic layer these fibres enter into synaptic
contact with a limited number of dendrites, this
being expressed in the very limited area from
which the evoked potential can be led off on the
first day after birth. Later on this zone widens,
which points to new dendritic synapses, related
to neighbouring cortical areas being involved
(Fig. 5).
The following experiment can be made. After
the limited nature of the evoked potential has
been ascertained at the point of maximum activ-
ity we shall see, by increasing the strength of the
stimulus, that neighbouring points, which until
then had not produced an evoked potential, be-
gin to produce a considerable potential.
On the strength of these coT~dderations and
the experiments presented by Parpura, we can
build the following universal scheme, reflecting
the laws of increasing spread ,_-,f ascending in-
fluences on elements of the p]cximorphic and
fourth layers of the cortex.
The cortical evoked potential, which pre-
viously seemed homogeneous, c resists of many
components, each of which or:~;;inates in sepa-
rate subcortical mechanisms. Th~s fact forces us
to re-examine, very seriously, ~,ur conceptions
of the variability with which electrical cortical
phenomena are manifested in geJ~eral.
If the configuration of the iJ~duced potential
depends on several ascending messages to the
cortex and if these messages or ginate in differ-
ent subcortical formations, wc may evidently
admit the extreme changeability of the interval
between these individual ascending messages.
This change may occur owing t,,~ the change of
externally acting agents, owing to the selective
influence of various chemical substances on indi-
Electroenceph. clin. NeurophysioL, 1964, 1 6 : 2 7 4 3
 ASCENDING INFLUENCES ON CORTEX
vidual subcortical centres and, finally, as we
shall see later, also owing to variable ascending
influences caused in the subcortex by humoral
agents.
As prototype of this possible explanation of the
configuration of electrical potentials in the cor-
tex, we can use the correlation in time of two
ascending currents of excitation, determining the
positive and negative phases of the evoked po-
tential (Fig. 6). As can be seen from this drawing,
~: b 20rnsec ~ a b
Fig. 6
Various types of primary evoked potentials which arise,
depending on intervals between two ascending volleys of
impulses--axo-somatic and axo-apical.
(A) The positive surface potential did not have enough
time for development of its complete amplitude. Volleys
of impulses along the other ascending pathways came to
the plexiform layer while the positive potential was yet
developing.
(B) The second volley of impulses to the apical dendrite
area in the plexiform layer was either blocked at the level
of the subcortical nuclei, or the initial message was not
strong enough. The positive phase bad sufficient time
for maximal development of its amplitude. The amplitude
of the negative phase is partly obscured.
(C) The volley in the specific pathways (axonal discharge)
is followed by the volley of impulses along the fine
ascending pathways. The positive phase of the surface
potential is not developed because of the small interval
between volleys,a: Arrival of volleys at the cerebral cortex
along the specific thalamic pathway, b: Arrival of axo-
apical excitation, possibly along the thalamo-reticular
pathways, a-b: Interval between development of post-
synaptic potentials from two ascending volleys.
the waves of excitation following ascending
routes as far as the pleximorphic layer may enter
the cortex at different intervals from the excita-
tion wave proceeding through the specific thal-
amus. Depending on the nature of this interval,
we can have a different expression of the positive
phase o f the evoked potential. If pleximorphic
35
excitation develops very quickly after the for-
mation of axosomatic potentials, the positive
potential of the cortex may not have time to de-
velop and we shall then have a well defined nega-
tive component only. If, on the contrary, the
excitations are somewhat delayed in comparison
with the specific excitations, the positive phase
may develop to the end and will then predomi-
nate in the general configuration of the evoked
potential.
That the extremely variable general configura-
tion of slow electrical activity in the t:ortex may,
to some extent, depend on the ascending subcor-
tical excitations changing in time is a very tempt-
ing idea, We must also consider the fact that
simple inter-relations, on the level of physical
electricity only, may determine the general pic-
ture of slow electrical activity of the cortex. We
have seen that one potential, as a physical phenom-
enon (positive component), may be cut off by
another physical phenomenon, having the op-
posite sign (negative component). The question
which naturally arises is: cart these interrela-
tions of electrical phenomena influc~ce in any
way the neurophysiological substratum which
caused them? This is quite evidently a fundamen-
tal question of the nature of slow elec~ rical activ-
ity of the cortex.
Recent experiments, conducted it, our labo-
ratory, in which polarizing direct current was
applied, have confronted us with ver~ paradoxi-
cal phenomena, which we shall certainly elab-
orate further. The evoked potential was regis-
tered from two points in the brain w'th the aid
of two separate electrodes. The first s'as at the
point of maximal activity of the evo~.ed poten-
tial in the sensori-motor field, the see,rod in the
specific thalamic nucleus through which passed
the stream of excitations caused by stirr, ttlation of
the sciatic nerve. As we know, well c~<fined, or-
dinary biphasic potentials are, in this t use, regis-
tered at both levels, in the specific thalamic
nucleus the recording bipolar electrode was in
a co-axial metal cover which served f,,r the po-
larization of this point of the brain a ith direct
current.
The experiment showed that, by applying pos-
itive direct current (anode)through the thalam-
ic electrode, we can observe the sharp growth
of the thalamic evoked potential whose ampli-
Electroenceph. c/in. Neurophysiol., 1964, 16:27-43
36
tude, especially in the anodic component, grows
two and a half times. But the cortical evoked po-
tential does not change in intensity. The oppo-
site experiment can be made by changing the
direction of the current and making the polari-
zation in the thalamic nucleus cathodic. In re-
sponse to this polarization, the evoked potential
of the thalamic nucleus reduces its amplitude
considerably. At the same time the amplitude of
the evoked potential in the cortex does not un-
dergo any noticeable change (Fig. 7).
N +
2oo,,,v
r
I'
~, i~ ,~/~--x [ 2oo~,v
~d
i
I 1 2oo,,,,v
IOMSE¢
Fig. 7
WiIh polarization of nucleus ventralis postero-lateralis
a n d c h a n g e of potentials in this nucleus, evoked poten-
tials of the cerebral cortex remained u n c h a n g e d . Below
are given inter-relations between areas of recording a n d
areas o f polarization. N - - evoked potentials before
polarization of nVPL. I: the first area o f s e n s o r i - m o t o r
cortex. 11 : the second area o f s e n s o r i - m o t o r cortex. 11I :
nucleus ventralis postero-lateralis; stimulation of the
radial nerve.
A paradoxical situation is thus created. It
would seem that the cortical evoked potential
is the direct result of the evoked potential in the
specific thalamus and so must its components be,
but the considerable changes of amplitude in
the thalamic evoked potential are in no way re-
flected in the amplitude of the cortical evoked
potential. One might think that the cortical po-
tential is determined by some collateral excita-
P. K. ANOKHIN
tion not subject to the influence of polarization.
But a control experiment excludes the possibility
of such a hypothesis. If we make the polarizing
electrode coagulating, thereby destroying the
brain tissue at the point of conduction, the cor-
tical evoked potential disappears; entirely.
All the experiments described above lead us to
think that the oscillations of the type of evoked
potentials can easily be neutralized by applying
polarizing currents of different sign, whereas the
truly neurophysiological process develops in the
thalamic nucleus and reaches tile cortex inde-
pendently of that change in the general aspect
of electric sign.
I am glad to note here that i~ the last two or
three years the study of evoked potentials, as the
most outstanding phenomenon ~n the electrical
activity of the cortex, has entered on a new phase,
in which the neurophysiological :mbstratum and
"IT the processes developing in it have begun to be
considered as the central factor, determining the
configuration and external expression of the
TIT
electrical indices or the central ~ervous system.
I11. ROLE OF BLOCKING AND EXCI! ING SUBSTANCES
IN ANALYSIS OF THE ELECTR{,?AL ACTIVITY
OF THE CORTEI~
A special neurophysiologica approach has
appeared in recent years in rese~irch on the elec-
trical activity of the cortex, naHely, the use of
pharmacological drugs in analys~s of the natule
of electric potentials. Reference.: to the works
of E. King (1956), Bradley (1958), Killam and
Killam (1958), Rothballer (1957) and other
authors is sufficient to understated the enormous
benefit which the analysis of c.)rtical electriea!
activity can derive from the use (,f this approach.
The principal reason for usitg various phar-
macological substances in nearophysiological
analysis is that all these subst~nces, especially
anaesthetics, have a specific selective influence on
definite nervous structures. Frc,m the point of
view of modern biology this selective faculty must
consist in the given substance, peculiar from a
chemical standpoint, finding in ..ome complicat-
ed protoplasmic complex a sensitive receptor
i'0r its influence. In short, in order to influence
points in some nervous formation, the given
chemical substance must ente~" into chemical
reaction with them.
Electroenceph. clin. Neurophysioi, 1964, 16:27-43
 ASCENDING INFLUENCES ON CORTEX 37

Thanks to this mechanism of diverse phar-
macological substances, we are now able to
affirm that there is an exceptional variety of
chemically specific metabolisms in the subcorti-
cal field, especially in the hypothalamus and
reticular formation. We may think, and there are
already experimental reasons for it, that classi-
fication of anaesthetics, for example, according
to the same criterion - - sleep, is founded on the
influence exercised by these different substances
on the structures of the reticular formation
which determine the state of wakefulness. From
that their influence spreads in the first place to
the subcortical formations with a generalized
action (activation, secondary discharges, recruit-
ing). This is the role of pharmacological sub-
stances in the analysis of the neurophysiological
basis of slow electrical activity of the cortex.
I have frequently had occasion to point out
that different narcotic substances, Nembutal and
urethane for instance, block in different ways the
subcortical mechanisms that exercise a constant
ascending influence on the cortex (Anokhin
1958). Just now 1 should like to examine two

a ~ I

2s0~vt__
5 s~c"

t ~ . ~ / k a ~ l ~ "

b~"

Fig. 8
(a) Action of a nociceptive stimulus under urethane anaesthesia. The stimulus evokes remarkable
generalized desynchronization of electrical activity of the cerebral cortex. (b) The same stimulus does
not cause desynchronization in the cortex after injection of chlorpromazine (Aminazine).

a neurophysiological point of view this means
that the given group of substances has the ca-
pacity of selectively suppressing the metabolism
of the nervous elements which constantly exer-
cise an activating influence on the cortex, thus
determining the state of wakefulness.
But each of these substances also has a sup-
plementary spectrum of chemical action on
other subcortical formations. Thus, the sum of
both these effects characterizes the given nar-
cotic as a unique chemical substance.
The use of substances belonging to the group
of narcotics and the group of tranquillizers shows
examples which convince us that the "narco-
tized" animal, sunk in deep sleep, is always found
to be, from a neurophysiological point of view,
a peculiar animal, depending on the type of narco-
tic which has been applied.
For example, urethane, given in a definite
dose, sends the animal to sleep, arid we know
that that sleep corresponds to narcotic sleep in
all its symptoms. But, if a painful stimulus is
applied to such a rabbit, the cortex will immedi-
ately respond to it by sharp desynchronization,
no weaker than in ordinary conditions of wake-
fulness (Fig. 8). On the contrary, under the same
Electroenceph. clin. Neurophysiol., 1964, 16:27-43
38 P . K . ANOKHIN

~1, (a)

(b)

0.2 mY -

60 m sec
Fig. 9A
After coagulation of the rostral part of the reticular formation the Forbes secondary discharge under
pentobarbitat disappears. In this example, urethane is given instead of pentobarbital, and one can
see that the marked "secondary urethane discharge" appears.

(a ) 1_ (c )

Fig. 9B
Dependence of the "secondary urethane discharge" (a and c) on the ascending influences of the
subthalamus and hypothalamus. Coagulation of hypothalamus (b and d) leads to elimination of the
"secondary urethane discharge" but fully preserves the primary discharge. (The negative component
of the primary discharge is known to be blocked by urethane.)
Electroenceph. clin. Neurophysiol.~ 1964, 16:27-43
 ASCENDING INFLUENCES ON CORTEX
conditions of sleep induced by another narcotic,
Nembutal for instance, there will be no desyn-
chronization in the cortex. In the case of a rab-
bit under the influence of urethane, desynchroni-
zation in response to painful irritation disap-
pears as soon as a small dose of chlorpromazine
is given to it.
In order to understand these different effects
of painful irritation on cortical electrical activ-
ity, we must imagine that individual complexes,
such as the complex of reaction of the whole or-
ganism to pain, have components that are indi-
vidually sensitive to different chemical substan-
ces. As we know, chlorpromazine, given in this
dose, does not send the animal to sleep, but con-
siderably reduces the animal's reaction to con-
ditioned painful stimulus.
After comparing these very limited facts, we
may therefore say that the final effect, i.e., the
effect on behaviour, of any substance depends on
its highly selective influence on the subcortical
structures. Every pharmacological substance of
a narcotic or tranquillizing nature exercises a
fractional dissociation in the subcortical nuclei,
letting some influences reach the cortex, while
it blocks other influences. From this point of
view every chemical substance of this kind is a
peculiar filter, which changes the complex of
ascending influences passing through it into the
cortex.
The selective action of this ascending influ-
ence and, consequently, the influence on the
general aspect of slow electrical activity of the
cortex, was noted very precisely in the experi-
ments of our colleague, Liu-Chang Hui. He
showed that, as a generalizing influence of the
subcortex on the cortex, the well known "For-
bes secondary discharge" is a direct result of the
narcotic substance applied in the given case.
This secondary discharge can manifest itself es-
pecially well under Nembutal narcosis. But if
urethane is used, this secondary discharge, pro-
duced by the rostral part of the reticular forma-
tion, disappears and a secondary discharge
appears in the region of the hypothalamus. It
is interesting that the Forbes secondary discharge
is eliminated by the destruction of the rostral
part of the reticular formation, while the "ure-
thane secondary discharge" is eliminated by the
destruction of the posterior wall o? *he hypo-
39
thalamus. Here is a clear indication that the use
of different narcotics is a decisive factor, giving
free passage to the cortex of the given complex
of ascending excitations and, on the contrary,
blocking the other set of excitations (Fig. 9).
Aside from the methodological importance
of these facts, warning us against an absolute
neuro-physiological estimation of the slow elec-
trical phenomena observed in the cortex, we may
point out that comparison of different anaesthet-
ics leads to a deeper understanding er the con-
stitution of slow electrical activity of ~he cortex.
For example, in response to stimulation of the
sciatic nerve under Nembutal narc,)sis, both
phases of the evoked potential are definitely
seen, i.e., the positive and the negatiw:. (Fig. 10).
On the contrary, under urethane narcosis, in the
same conditions of experiment, we get the posi-
tive phase only.
(a) (b)
[ . J
Fig. 10
T w o cortical surface evoked potentials ( , ; n s o r i - m o t o r
area) : (a) u n d e r N e m b u t a l anaesthesia, (b) after changing
N e m b u t a l to u r e t h a n e anaesthesia. It is see~ that in the
second case the negative c o m p o n e n t disappeared.
Imagine for a minute that the only narcotic
at our disposal is urethane; then the factual re-
sults of our research on the evoked potential of
the cortex would be completely different and so
would naturally be our conceptions ef the neuro-
physiological nature of the potential. All these
arguments make us realize the necesdty of spe-
cial research on the individual role c,f every nar-
cotic in the formation of cortical electrical activ-
ity.
IV. ASCENDING INFLUENCE OF HUMORAL ORIGIN
ON THE CORTEX
Until recently there were several examples of
humoral influences on the reticular formation,
leading to an activated condition. We must first
mention the role of adrenalin which, through a
Electroenceph. clin. Neurophysiol., 1964, 16:27--43
40 P. K. ANOKHIN
mechanism so far unknown, exercises an activat-
ing influence on the cortex through the blood
(Dell and Bonvallet 1954). Although Italian work-
ers have shown that, in this case, there can be
no question of a direct influence of adrenalin on
the activating substratum of the reticular for-
mation, it remains nevertheless certain that
adrenalin, injected into the blood, proves an
activating factor for cortical electrical activity.
Carbon dioxide is another natural factor caus-
ing the excitation of the activating mechanisms
of the reticular formation. It is known, also from
experiments conducted in Dell's laboratory, that
increased concentration of CO2 in the blood
leads to the activation of cortical electrical activ-
ity. Recent experiments in our laboratory, in
which highly concentrated CO2 was used on
newborn rabbits, showed that, at the moment of
birth, the reticular formation proves to be highly
sensitive to the influence of an increased content
of carbon dioxide in the air breathed by the
animal, which reacts to it by the appearance of
the stress rhythm already known to us. It is
interesting that this reaction to carbon dioxide
is much more strongly expressed than the reac-
tion to painful stimulation of the rabbit's hind
limb.
One form of natural humoral influence on the
activating mechanisms of the brain was discov-
ered in our laboratory in recent years. It was
shown, for example, that a cat which has received
no food for two days before the experiment mani-
fests a definite desynehronization of electrical
activity in the frontal areas of the cortex, even
under narcosis. All the remaining parts of the
brain remain in the condition of slow electrical
activity characteristic of sleep. It is quite evident
that we have here a case where the "starved
blood", exciting the hypothalamic structure
(Andersson 1959) of the feeding centre, trans-
fers this excitation to the activation of the frontal
section of the cortex.
This surmise should be controlled in special
experiments. We decided to satiate th'e animal
under narcosis and follow the evolution of elec-
trical activity of the cortex. Artificial feeding was
accomplished by two methods: by the intra-
venous injection of glucose and by introducing
milk through the natural tract, i.e., through the
mouth and oesophagus into the stomach. The
experiment showed that, after the "feeding" pro-
cess given under narcosis, desynchronization in
the frontal parts of the cortex is almost immedi-
ately transformed into the slow rhythm character-
istic of sleep. It is interesting t~lat maximal ef-
fect is produced only by the two t'actors together,
i.e., injection of glucose into the blood and milk
given through the mouth into the stomach (Suda-
kov 1961).
Analysis of this interesting paenomenon has
shown that desynchronization of the frontal
regions of the cortex can also be switched to
slow rhythm by cutting the spinal cord at the
level of the third thoracic seg1~aent, and both
vagus nerves. It was thus provec' that the desyn-
chronizing influence proceeds in 1his way through
the intestinal tract and, together ,~'ith the exciting
influence of starved blood, leacIs to the imme-
diate activation of the frontal section of the
cortex.
Perhaps the most interesting feature of these
experiments is the fact that the i afluences affect-
ing the composition of the electrical activity of
the cortex are not only ascendi~g and blocking
but, evidently, many of them ar~ initially due to
humoral factors circulating in tie blood of the
animal.
These experiments naturally r .rise very impor-
tant questions; in the first plat:e, the question
that the influence of the narcotic, and the narco-
tic condition of the animal itselF, are not at all
universal blocking factors for a l the ascending
excitations from the subcortical field to the cor-
tex. There is probably also a covsiderable quan-
tity of ascending activation tl~at escapes the
influence of the narcotic. We act lally have many
kinds of generalized effect whica can very well
be ascertained under narcosis (secondary dis-
charges, urethane, painful de~,ynchronization,
associative responses, "recruitil~g" etc.).
On the basis of this materitl, we may say
that study of the nature ofcortic;~ Lelectrical activ-
ity and its relations to the ne~rophysiological
substratum of nervous function must include, as
a most important line, classifica~~ion of ascending
influences from the point of view ~f their capacity
to "escape" the blocking influence of anaesthetics
and tranquillizers. It seems to me that this line
of research on electrical phenomena can also be
of great importance in wide clinical practice.
Electroeneeph. elin. Neurophysiol, 1964, 16:27-43
 ASCENDING INFLUENCES ON CORTEX
V. "REBOUND ACTIVATION" AS ONE OF THE ITEMS
OF SLOW ELECTRICAL ACTIVITY OF THE
CORTEX
As we have seen from the previous state-
ments, the cortex receives numerous ascending
influences and relays them in the postsynaptic
potentials of definite systems of connections,
where their final synthesis takes place. There is
one more form of influence, called by us "re-
bound activation", which can be included in the
number of these ascending influences.
Until now we have spoken of ascending in-
fluences that are initiated somewhere on the
periphery, transformed at the subcortical level
and spread through the cortex with some degree
of selection. At the same time, experimental work
has long ago shown the existence of a whole series
of cortico-fugal influences, i.e., influences from
the cortex to subcortical formations.
The usual analysis of these cortico-fugal in-
fluences stops at their significance for the condi-
tion of subcortical cells. These can be cells of the
hypothalamus, thalamus and reticular forma-
tion. But the study of the reverse influences of
these subcortical conditions has not been system-
atically conducted, as far as ! know.
I can only mention the case of reverberation
in thalamo-cortical interrelations, which involves
highly localized cyclic interactions of excitation,
and not generalized ascending influences.
! propose to set here the question of another
form of interrelation which, it seems to me,
contributes considerably to the general electrical
activity of the cortex. There is no doubt that, in
natural conditions, the activity of the cortex
is not limited to the reception of ascending influ-
ences. This refers to cases where the synthetic
process, for example, of recognition ends with
the inclusion of powerful emotional subcortical
complexes, ensuring entirety of behaviour. In
this case the process of excitation, in its very
substance highly selective, has a double direc-
tion: at first it is cortical, afterwards it excites, in
a cortico-fugal direction, a certain complex of the
subcortical formations and these produce a reverse
generalized ascending influence on the cortex.
It follows from these considerations that the
observable cortical electrical activity of a normal
animal is always a complicated interlacing of all
41
types of influences and interactions. This circum-
stance is a stumbling-block in the explanation of
the neurophysiological bases of cortical elec-
trical phenomena.
We have elaborated in our laboratory the
model of an experiment enabling us to under-
stand the degree of this type of reverse activa-
tion, or reverse influence on the cortex. By
application of paper soaked in strychnine on
a definite spot of the cortex, we ca~ evoke a
very interesting phenomenon. After 10-15 rain
we can observe throughout the corte:~ paroxys-
mal and epileptiform discharges, occurring with
the same latency independently of their distance
from the initial point on which strychnine was
used. These generalized influences are ~aotexclud-
ed by cutting the corpus callosum, although the
strychnine was applied on one hemisphere only.
Analysis of the phenomenon has convinced
us that strychnine excitation, at first highly local-
ized, produces powerful excitation of the cor-
tico-fugal type in the subcortical sy;tems, and
this excitation in turn exercises a generalized
reverse influence on the cortex. If this initial
strychnine point is refrigerated after a certain
time, for example, an hour after application of
strychnine, and all electrical activity is stopped
in that spot, paroxysmal discharges will still
occur in other parts of the brain (Shelikhov 1960).
We obtained the same phenome~on in an-
other experimental form. If 5 or 6 fine needles
are joined together and the cortex is slightly
pricked with this packet of needles, with a slight
abrasion of the cortical tissue, the ~hole cortex
will immediately be desynchronized As in the
first case, cutting the corpus callosum does not
exclude these generalized changes ol the cortex,
although irritation by means of the needles is
applied to one hemisphere only. The effect is elim-
inated on the non-irritated hemi,,phere and
retained on the irritated hemisphere ~tfter cutting
the commissural fibres in the region of non-
specific thalamic formations (Rogacheva 1962).
In comparing and estimating these two types
of reverse activation, we can note that these arti-
ficially obtained reverse influences o~1 the cortex
undoubtedly occur in natural conditions also, in
integration of the entire behaviour. It follows that
this "reverse" activation Ls, besides olhers, a com-
ponent of the slow electrical activity of the cortex.
Electroenceph, clin. Neurophysiol., 1964, 16:27-43
42 P. K. ANOKHIN
CONCLUSION
All the above material shows the great variety
of the components of the phenomenon which we
call electrical activity of the cortex. We are prob-
ably far from knowing all the forms of influence
jointly determining this extremely variable elec-
trical activity. The complication is so great that it
would seem as if it must lead us to a very pessi-
mistic opinion about the chances of success in its
study.
On the other hand the results of gradual ana-
tomical study of component parts of cortical
electrical activity give us reason to hope for suc-
cessful results in our research work.
if we take it that the cortical electrical activity
of an animal in a wakeful condition is the result
of a great number of ascending influences, the
question involuntarily arises as to the way in
which these influences are synthesized and where
the formation of the final resultant of all these
nervous influences takes place.
It seems to me that the overwhelming ma-
jority of the experimental results, especially
those presented at this meeting, are evidence that
the electrical activity observed by us in the cortex
cannot be the result of the summation of impulse
discharges in individual cells. There are many
more reasons for holding that the resting elec-
trical activity of the cortex is an integrated result
of electrotonic conditions. The latter can be due
not only to working nervous impulses but, which
is particularly important, to certain specific as-
cending influences which, because of their physio-
logical nature, are capable of changing the elec-
trotonic conditions of the whole mass of the brain.
We can point to the fact of generalized hy-
persynchronizations, which have been well elabo-
rated in the laboratory of Moruzzi, who has
proved the existence of special centres in the
lower part of the reticular formation.
Ontogenetic research shows likewise that, in
a definite stage of ontogenesis, directly after
birth, painful stimulation of the rabbit's hind
leg with hot water produces only large ampli-
tude slow activity instead of the desynchroni-
zatio~, usual for an adult animal. It follows that
the mechanism of high voltage slow activity
matures heterochronically in relation to the mech-
anism of desynchronization, which is proved
by its separate substratum (Ata-Muradova 1960).
The same fact is excellently proved in the report
made by Purpura (1959).
It is quite possible that we shall have to con-
sider the existence of a special mechanism in the
subcortical regions, which can include wide areas
of the brain in the same regime~ of slow electri-
cal activity, as we see in the exampleofthegener-
alized stress rhythm. The existe3~.ce of this mech-
anism is now almost proved, ~,o that it only
remains for us to solve the problem of the inter-
relation between it and the gerteral integrative
processes which determine inter-rteuronal and se-
lective interrelations.
The most difficult problem for lhe understand-
ing of slow electrical activity ol the cortex, ob-
served by us with the aid of the electroencepha-
lograph, is the evident interactim and mutual
suppression of individual electri,~:al phenomena,
independently of observable ir~eractions on a
neurophysiological level.
If we take it that the electrical ighenomenon of
cortical activity is always only an epiphenornenon,
i.e., an unavoidable physical rest~lt of the neuro-
chemical process already developed before it, we
are faced with a very sad prospect in the estima-
tion of the general role of slow ele,.'trical phenom-
ena in integral nervous activity.
These electrical epiphenomer~:., developing in
such volumetric electrolytic cot:ductors as the
brain mass, can enter in the m o , t diverse inter-
relations, whose final form depen Js only on their
electrical signs, formed as the re,~tdt of either sub-
traction, or addition, or compk'te mutual neu-
tralization.
Given this premise, the neJrophysiological
processes and real inter-relation ; between neu-
rons could never be adequately reflected in the
EEG seen by us. At least this is ti,e conclusion to
which we are led by the above f~ zts of the inde-
pendence of form of the potential; induced in the
cortex in cases of sudden chanb'~s of the inter-
mediate potentials in the speciti: nuclei of the
thalamus (Agafonov, Duriny'. n). However,
these problems are only just ap:)earing on the
horizon of our research inteests and the
immediate future must show just low correct are
all our suppositions regarding li'~e interrelatior~
between slow electrical activity a~-td the real pro-
cesses of nervous activity.
Electroenceph. c/in. Neurophysiol.. 1964, 16:27-43
 ASCENDING INFLUENCES ON CORTEX
REFERENCES
ANOKHIN, P. K. [Contemporary problems of embryogen-
esis of nervous activities.] Advanc. modern Biol.
(U.S.S.R.), 1933, 2: 40-69.
ANOKHIN, P. K. [Functional system as the basis of integra-
tion of nervous activities in embryogenesis.] Work o f
I/lth Congress o f Soviet Physiologists, Tbilisi, 1937.
ANOKmN, P. K. Systemogenesis as general regularity of
evolution. Bull. exp. Biol. Med., 1948, 10: 361-385.
ANOKHIN, P. K. [On morphological regularities of the
development of the functions in embryogenesis in
animals and man.] Session o f Anatomists o f the USSR,
1958, 1: 25-36.
ANOKHIN, P. K. [The electroeneephalographic analysis o f
the conditioned reflex. 3 Medgiz, Moscow, 1958.
ANOKHIN, P. K. The multiple ascending influences of the
subcortical centres on the cerebral cortex. In M. A. B.
BRAZIER (Editor), Brain and behavior. National Science
Foundation, Washington, 1961, l: 139-170.
ATA-MURADOVA, F. [On changes of the evoked potential
in the brain during the process of post-natal ontogeny.]
First Joint Conference on the Reticular Formation,
Moscow, 1960.
BANZEKINA, M. M. M. Sci. Thesis, Moscow, 1962.
BIsHoP, G. The place of cortexin a reticular stem. In
H. H. JASPER e t a l . (Editors), Reticular formation o f
the brain. Little, Brown and Co., Boston, 1958: 413.
BRADLEY, P. B. The central action of certain drugs in
relation to the reticular formation of the brain. In
H. H. JASPER et al. (Editors), Ret&ular formation o f
the brain. Little, Brown and Co., Boston, 1958: 123-
149.
BRAZIER, M. The action of anesthetics on the nervous
system with special reference to the brain stem reticular
formation. In E. D. ADRIAN et al. (Editors), Brain
mechanisms and consciousness. Blackwelt, Oxford,
1954:163-193.
DELL, P. et BONVALLET, M. Contr61e direct et r6flexe de
l'activit6 du syst~me r6ticul6 activateur ascendant du
tronc c6r6bral par l'oxyg~ne et le gaz carbonique du
sang. C. R. Soc. Biol. (Paris), 1954, 148: 855-858.
ECCLES, J. C. The neurophysiological basis o f mind. Claren-
don Press, Oxford, 1953.
Reference: ANOKHIN, P. K. The electroencephalogram as a resultant of ascending influences on the cells of the
cortex. Electroenceph. clin. Neurophysiol., 1964, 16: 2 7 4 3 .
43
GRUNDFEST, H. Evolution of conduction in the nervous
system. In A. D. BASS (Editor), Symposium on evolu-
tion o f nervous control from primitive organisms to man.
American Association for Advancement of Science,
Washington, D.C., 1959: 43-83.
HERRICK, C. J. The brain o f the tiger salamander. Uni-
versity of Chicago Press, Chicago, 1948, 409 p.
HERRICK, C. J. and BIsHoP, G. H. A compar;~ti 'e SUlvey
of the spinal lemniscus systems. In H. H. JASPER etal.
(Editors), Reticular formation o f the btain. Little,
Brown and Co., Boston, 1958: 353-361.
KILLAM, K. F. and KILLAM, E. K. Drug actium on path-
ways involving the reticular formation. In H . H .
JASPER et al. (Editors), Reticular formatiot~ ,~f the brain.
Little, Brown and Co., Boston, 1958:1 ] 1-123.
K1NG, E. E. Differential action of anesthetic~ and inter-
neuron depressants upon EEG arousal and recruiting
responses. J. Pharmacol. exp. Ther., 1956, 116: 404-
417.
KING, E. E. ,NAQUET, R. and MAGOUN, H. ~ ' Alterations
in somatic afferent transmission through the thalamus
by central mechanisms and barbiturates, d Pharmacol.
exp. Ther., 1957, 119: 48-63.
PURPURA, D. P. Nature of electrocortical potentials.
Int. Rev. Neurobiol., 1959, 1: 107-163.
ROGACHEVA, S. K. [About return generalizz~tion of the
excitation in the cortex after its traumatization.] First
Joint Conference on the Reticular Formatfi,n, Moscow,
1960,
ROTHBALLER, A. B. The effect of phenylephri~le, metham-
phetamine, cocaine, and serotonin upon the adrenalin-
sensitive component of the reticular actiw~ting system.
Electroenceph. clin. Neurophysiol., 1957. 9: 409~,17.
SHELmHOV, V. N. L'6tude 61ectrophysiologique du rfle des
formations sous-corticales dans la g6n6ralisation de
l'excitabilit6 de l'6corce des grandes h6misph6res.
Zh. Nevropat. Psikhiat., 1960, 60: 145-148.
StJDAKOV, K. V. [The experience of study of ascending
activating influences on the cortex by m~:ans of local
polarization of hypothalamus.] Fiziol. Z h Leningrad),
1963 (in the press).
SUN VEN. [Comparative electrophysiologica! analysis of
changes of lability of components of c~,, tical evoked
potential in postnatal ontogeny.] Fiziol Zh. (Lenin-
grad), 1963 fin the press).