Soviet Psychology

ISSN: 0038-5751 (Print) (Online) Journal homepage: http://www.tandfonline.com/loi/mrpo19

Advances in Brain Research

Petr Kuz'mich Anokhin

To cite this article: Petr Kuz'mich Anokhin (1966) Advances in Brain Research, Soviet
Psychology, 5:1, 14-23

To link to this article: http://dx.doi.org/10.2753/RPO1061-0405050114

Published online: 20 Dec 2014.

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 Brain Research

Nauka i chelovechestvo [Science and

humanity], Znaniye, 1965, 35-47
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Petr Kuz’mich Anokhin

(Sechenov Institute of Physiology, USSR Academy of Medical Sciences, Moscow)

ADVANCES IN BRAIN RESEARCH

Current study of the brain has reached a stage and help to map the route for further penetration
where the researcher in this field realizes that of the secrets of the brain. The need for gen-
he is only just beginning to unravel the secrets eralization and elaboration of the guiding prin-
of the brain. It is becoming increasingly clear ciples for conducting research is more and more
that even the most significant physiological prin- urgently felt.
ciples of nervous activity discovered s o far a r e The concept of the conditioned reflex was in
merely the portals of a hitherto little-known i t s time just such a generalizing factor in neuro-
world of molecular processes that play a deci- physiology. It is for this reason that the dis-
sive role in the working of the brain. covery and elaboration of conditioned reflexes
Each year brings in enormous quantities of played such a great historical role. The applica-
experimental information. How can this peculiar tion of the subtlest electrophysiological methods
crisis of overproduction of isolated facts be of research in the work of recent years has
avoided and given some sort of unifying scientific shown that the conditioned reflex provides not
bas is ? only the most practicable and effective means of
The view has gradually been accepted that one studying behavior, but also serves as a model
preventive would be to work out theoretical mod- to show the correlation of the separate aspects
els for research that would assist in rapid anal- of a given phenomenon to the whole. This must
y s i s of incoming facts and provide direction for be the explanation for the surprising revival of
the collection of new data. In other words, a interest in the conditioned reflex on the part of
universal “exploration chart” is needed that “pure” neurophysiologists.
would determine the exact location of every dis- At the same time the latest advances in cyber-
covery and its importance in the general scheme, netics and electronics have created new condi-

14
 VOL. V, NO. 1 15

tions for the further development of our concep- For the last twenty years numerous facts have
tions concerning the working of the brain, the been gathered from various scientific fields
conditioned reflex, and the general physiological which show that, from the physiological view-
regulation of the organism. The generalized point, a more general approach is needed for ex-
-
achievements of separate fields biology, phys- plaining the purposive nature of behavior as a
iology, physics, and even the social sciences - form of adaptation of an organism to the external
have made practicable the creation of a more world. It has become quite apparent that there
comprehensive theory embracing all the diver- a r e a number of complementary mechanisms
sity of the main mechanisms of higher nervous which expand the narrow reflex pattern and give
activity, and behavioral acts in particular. Con- it a complete physiological structure which ex-
currently, the conditions have also matured for plains problems that the reflex theory could not.
working out such a universal model of behavior These mechanisms help answer questions that
which would help in the orientation of separate must be resolved in order to understand the dif-
facts received through the most subtle electro- ferent phases of a behavioral act. What is the
physiological experiments by relating them to a true excitatory cause that creates the response?
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precise point within the model. In this way the What real physiological factors lead to the for-
interests of investigators of both fine neuro- mation of purpose, the decision to perform a
physiological processes and of behavioral acts specific behavioral act? How can the organism
would be organically combined. itself, out of millions of possibilities, perform
Our laboratory has worked out such a behav- precisely the actions that conform to the inten-
ioral model based on the principles of the func- tion? Lastly, and perhaps the most interesting
tional system [funktzional’naya sistema]. It as- problem: how can the organism and its nervous
sists both in setting up new research problems system “recognize” that the action has been suc-
and in interpreting the results obtained. -
cessfully completed whether it corresponds
We attempted a sort of reformulation of the to the set purpose or whether it deviates from
classical Cartesian “reflex arc” by introducing i t in some way and thus ceases to be an adaptive
a series of new physiological mechanisms which act?
broaden its capacity for explaining purposive For a long time these questions, essential to
behavior on a materialist basis. the understanding of behavioral acts, were not
According to Descartes’ theory, the “reflex systematically examined because they could not
arc” consists of three elements: the stimulus - be explained within the bounds of the traditional
an impulse which travels from the external en- “reflex arc.”
vironment to the brain by way of the sensory The present article will attempt to relate
nerve fibers; the translation within the brain of briefly the new principles, demonstrated by our
this stimulus into a motor reaction; the com- laboratory, that must be incorporated into the
mand conveyed by the motor nerves and executed reflex concept to create a comprehensive func-
by muscular action. This is how the reflex pat- tional system.
tern was understood for several centuries. We define a functional system as an aggregate
In time the concept of the “reflex arc” w a s ex- of processes and mechanisms dynamically united
tended, starting with the work of I. M. Sechenov. in terms of given situational demands, and neces-
But a particularly forceful impetus was given to sarily resulting in behavior adaptive for the or-
its development with I. P. Pavlov’s study of con- ganism in the given situation.
ditioned reflexes. And although Pavlov himself If human behavior were to be analyzed from
strove to remain within the theoretical frame- this viewpoint in any situation, it would appear
work of the “reflex arc,” h i s discovery of the that it can be subdivided into stages which a r e
conditioned reflex was of decisive importance themselves definite dynamic unities leading to
for the further elaboration of new concepts con- some final result. For example, you want to
cerning the essentials of the reflex theory. take a book from the top shelf of the bookcase:
 16 SOVIET PSYCHOLOGY

you stand on tiptoe, stretch out your hand, and action of the true stimulus. The excitations
take the book off the shelf. At the very moment from a conditioned stimulus, i.e., a stimulus for
when all your muscles were strained to help you some s o r t of reaction, combine in a synthetic
adopt the most comfortable position for getting unity with other excitations which have come
the book, a special, o r dynamic, functional sys- from preceding stimuli. The system of antece-
tem was formed, whose activity concluded in a dent excitations does not produce a conditioned
very definite result: you got the book. reaction, but rather determines its shape and
Thus it follows that the functional system does extent. A few years ago this hidden system of
not relate solely to the cerebral cortex or even excitations was entitled antecedent integration
to the brain as a whole. It is by its very es- [predpuskovaya integratziya], implying a system
sence a central-peripheral formation in which capable of forming a reaction as soon as the cor-
impulses circulate equally from the center to responding stimulus should take effect.
the periphery and from periphery to center, The dynamic stereotype, which was fully
which gives the central nervous system uninter- worked out in Pavlov’s laboratory, serves as a
rupted information about results achieved on the typical example for confirming this position. It
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periphery. is well known that if conditioned stimuli are re-

The formulation of the functional system goes peated in the same order over a long period,
beyond the bounds of the reflex theory, which af- eventually a stereotyped system of consecutive
firms that any external stimulus going to the excitations, independent of the external stimulus,
brain as a form of excitation is distributed is created in the animal’s brain.
throughout the central nervous system. In the experiment a dog is subjected to the
Numerous experiments conducted by col- action of 6 stimuli, each producing a definite
leagues in our laboratory have raised serious amount of saliva: an audible tone - 8 drops, a
doubts about the correctness of the classical light - 5 drops, a bell - 13 drops, a light - 5
position. It appears that excitation arising from drops, a bell- 13 drops, an audible tone - 8
the stimulation of some sensory organ, known in drops. The order of the stimuli was not changed
physiology as afferent, enters into subtle inter- f o r a year. A definite stereotype was worked
actions with other afferent excitations, and only out in the dog. Then the experimenter used only
the synthesis of all these afferent excitations the light as the stimulus. The dog reacted as
creates the conditions for the formation of pur- though all the other stimuli had been given as
poseful actions. In other words, the brain pro- well. He salivated at the same rate as in the
duces a highly specific synthesis of all external previous experiments.
signals which reach it through the various sen- Consequently, the conditioned stimulus loses
sory organs and which have different functional its essential concrete activity, and reflects a
significance. From this the concept of afferent situation previously created by a dynamic stereo-
synthesis arose. type. In other words, the new result is charac-
What is the significance of separate stimuli teristic of conditioned stimuli that have been
in this process? It consists of the following: used earlier, and not of the current, real stim-
any stimulation can either arouse of itself some ulus.
sort of reaction, in which case it is the true The direct nature of this interesting phenom-
stimulus for the appearance of this reaction; or enon w a s revealed by means of recording brain
it can serve as a factor preparatory to the ap- waves. The electroencephalogram shows that,
pearance of a given reaction. Experiments prior to the arrival of the excitation of the next
demonstrate that both kinds of stimulus form an stimulus, an anticipatory excitation occurs at a
organic unity, but that each contributes its own corresponding point in the cerebral cortex, re-
specific part to this unity. -the
flecting the course of external events (vide
-
The final adaptive result or effect, as phys- experiments of our colleague, A. D. Semenenko).
-
iologists say is not merely the result of the The heterogeneity of afferent excitations,
 VOL. V, NO. 1 17

which synthesize before the behavioral act is the animal’s behavior.

formulated, has been even more clearly demon- Ordinarily the dog had behaved quietly during
strated in the experiments of our colleague, I. I. the experiment; in the intervals between the
Laptev. He has connected one single conditioned conditioned stimuli it had sat in the middle of
stimulus (the bell) with two other non-conditioned the stand and, in response to the stimulus, which
stimuli. After the morning bell the dog was fed, was an initiatory stimulus to a given reaction,
at night the same bell was accompanied by an had run to the appropriate feeding-trough. This
electric shock. Ultimately a state was reached was the result of afferent synthesis by which the
where the same stimulus, the bell, gave highly animal’s behavior was coordinated with the ex-
different results according to the time it was perimental situation. The dissolution of the
used. In the morning, when the bell rang, the synthesis of afferent excitations, which occurred
animal clearly demonstrated a hunger reaction after the removal of the frontal sections of the
and salivated. In the evening, the same bell pro- cortex, changed the dog’s behavior. The animal
duced a distinct, defensive reaction. We are rushed from trough to trough, back and forth like
presented with the complex synthesis of an ini- a pendulum. These movements were a specific
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tiatory excitation (the bell), stimuli of the ex- reaction to the situation of alternate choice of
perimental conditions, and of the time at which trough that had taken place in the normal animal
the experiment occurs. Thus the final result is in the form of hidden antecedent [ predpuskovoy]
a product of the synthesis of different types of integration occurring after the presentation of
excitations: initiatory, and those pre-established the initiating [puskovoy] conditioned stimulus.
through earlier integration. We can conclude from the experiments de-
How is the process of afferent synthesis de- scribed that the cerebral cortex produces the
veloped? Which parts of the brain a r e involved, synthesis of numerous and functionally diverse
and what properties does each participating ele- afferent influences from which the adjusted be-
ment possess? havioral act ensues in conformity with a given
The mere fact that the perception of all these situation.
afferent influences cannot be evaluated as a lin- A t the present time there is a sound basis for
ear diffusion of excitations shows that we must speaking of four components that comprise the
seek a definite physiological framework to con- development of afferent synthesis.
solidate this synthesis. 1. Totality of situational stimuli, creating in
This was precisely the conclusion reached by each separate case an antecedent integration of
A. I. Shumilina’s systematic experiments, which excitations which, although hidden, can be im-
established that the concept of afferent synthesis mediately revealed as soon as the releasing
is an independent phase in the formation of the stimulus is activated.
behavioral act. By studying the physiology of 2. Initiating stimulus. The physiological
the frontal sections of the cortex she showed sense of this is to make the totality of hidden
that it is precisely here that the ability to store excitations, i.e., the antecedent integration,
different kinds of sensory influences in some come into play at the moment that is most ad-
synthetic unity is retained. vantageous for the success of the behavioral act.
The experiments consisted of the following: a 3. Motivating excitation. Without it there can
dog was put in a stand with two feeding-troughs be no behavioral act, for the motive to act al-
which were alternately filled with food as soon ways expresses an internal need of the organism.
as the conditioned stimulus was presented. A The behavioral act is called upon to satisfy this
conditioned reflex w a s built up in the dog to dis- need by creating the most beneficial conditions
tinguish between the two sides of the stand in re- for the existence of the organism in its given
sponse to two different conditioned food stimuli. state. The following example shows how im-
The dog then had its frontal lobes removed. A s portant the participation of motivating excitations
a result of the operation, changes occurred in is in afferent synthesis. If a dog is fed, the con-
 18 SOVIET PSYCHOLOGY

ditioned stimulus of food will not arouse hunger even in cases where they have acted on the re-
reaction, Why? Because there is no motive; ceptor (sensory) apparatuses through a very
the dog is satisfied, There is no ascending, ac- limited stimulus (e.g., a flash of light). A sim-
tivating excitation from the subcortical sections ilar stimulus on reaching the subcortex is widely
(the hypothalamus) in the dog’s brain to maintain diffused by arousing all the main subcortical
the hunger reaction with its energy. formations. Only after this do the excitations
4. Past experience. Afferent synthesis would that have been aroused enter the cerebral cortex
not be possible if the totality of situational and as separate currents as well as in a generalized
initiating stimuli was not connected with past ex- form. In practice almost every cell of the cor-
perience stored in the memory apparatuses. tex, through its synaptic organization, is brought
These fragments of past experience a r e set in into action by the excitations arriving from the
motion by these special mechanisms, enrich the subcortex and becomes a component part of the
present behavioral act, and give it the maximum whole system of excitations. Of course not all
of precision. the cells of the cortex are equally excited. On
These a r e the components by which afferent the contrary, there are ample grounds for be-
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synthesis is realized. However, the synthesis lieving that in different sections of the cortex
of information flowing to the brain could not be systems of numerous and unequal excitations oc-
completed if it were not accompanied by an ac- cur which also form the final result, the be-
tive orienting investigatory reaction. This re- havioral act.
action always arouses the reticular formation Thus contemporary study of the brain has
and the hypothalamus which, through their en- been enriched by new presentations of the inter-
ergy, tone up the cerebral cortex. Only then do action of excitations. It has become clear that
the unification of afferent excitations and the no single initial “focus of excitation” plays a
formation of a decision, corresponding to the re- leading part, but there is instead an extensive
quirements of a given situation and to the goal system of varying excitations, differing in
of behavior, become possible. strength and in the degree of diffusion, which
From our point of view the secret of forming enter into specific interaction between them-
a goal for the attainment of any result lies in the selves. How exactly does this take place in the
development of afferent synthesis. For this very brain? A few years ago we embarked upon an
reason detailed neurophysiological analysis of attempt to unveil this mysterious mechanism.
its component parts acquires such special im- Experiments conducted by our colleagues, V.
portance. For the last decade we have made a G . Agafonov, A. I. Shumilina, V. Gavlichek, and
thorough study of this problem, drawing upon the others, have shown that the excitations received
achievements of contemporary electronics in by the cerebral cortex from the subcortex are
our research. by no means uniform as was previously sup-
Study of the processes of the interaction of posed but are, on the contrary, completely
different afferent excitations on the synaptic varied, ascending excitations. These excitations
(contact) organization of one and the same cell have already acquired their specific biological
has now become exceedingly important f o r under- character while being formed on the level of the
standing the working of the brain. Suffice it to subcortical apparatuses, which determines the
mention some famous neurophysiological labora- part they play in the processes that go on at the
tories of various countries - that of A. Fessard cortical level. It is characteristic that the
(France), R. Jung (Western Germany), V. Mount- synaptic organization on the body of the cor-
castle (USA), H. H. Jasper (Canada), G . Moruzzi tical cell is drawn into these processes se-
(Italy), and many others. lectively and strictly in accordance with the
A s a result of numerous experiments con- specific biological character of the ascending,
ducted here and abroad, the striking variety of activating excitation. This law plays an enor-
ascending excitations has been clearly shown, mous part in the synthetic activity of the cere-
 VOL. V, NO. 1 19

bra1 cortex. be evaluated accordingly.

It is well known that each cortical cell absorbs A long s e r i e s of experiments have shown that
into its body and dendrites, by means of numer- ascending impulses of different biological qual-
ous synapses, several thousand different im- ity, on reaching the brain, activate synaptic links
pulses that come from thousands of other cortical possessing different biochemical natures. Thus
cells. The colossal number of contacts between the selective blocking of the cholinergic synapses
cortical cells creates unlimited possibilities f o r with amisil completely excludes the participation
interaction patterns. If the varied world of sub- of these synapses in the hunger reaction of the
cortical ascending excitations, which we have cortex. A s the electric charges demonstrate,
already analyzed and which enter the same cor- the cortical cells do not react to the ascending
tical cells, is added to this, the sum total com- excitations of hunger. However, these same
prises a true arena in which the process of af- cells a r e fully receptive to ascending excitations
ferent synthesis can be carried out. of a defensive, basically painful nature. If the
Imagine the chaos that would arise in our ac- sciatic nerve is stimulated when the hunger ac-
tivity and behavior if all the diverse connections tivation is eliminated from the cortex by means
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between the nerve cells were aroused uniformly of amisil, then the cortical cells under the same
and unspecifically throughout the cortex ! An effect freely pass into a state of desynchroniza-
amazing selectivity exists in the diffusion of ac- tion, as does the whole cortex.
tivating influences. Just as a river in flooding So, by pure chemical influence on the cortical
f i r s t sends its waters into channels, ravines, and cells it appears to be quite possible to obtain a
over low-lying ground, s o the ascending, activat- dissociation of the synaptic organization con-
ing excitations selectively attract those nervous nected with the ascending excitations of different
elements of the cortex and the related synapses biological nature since they have a different
which have been connected historically with a chemical structure. Thus a new aspect has been
given biological state. added by which the work of the cerebral cortex
Experiments in our laboratory have shown can be studied, especially the specific chemical
that the subcortical apparatuses of the brain, character of those synaptic structures that in-
which produce innate and acquired reactions of corporate all forms of ascending excitations and
different biological qualities, possess different realize the final stage of afferent synthesis, the
chemical properties closely linked with the pe- decision to act.
culiarities of their interchange (metabolism). It Why does the cerebral cortex unite two reac-
is now possible to speak of the specific chemistry tions of the organism which a r e opposed in their
of certain conditions, at least of the “chemistry biological relationship: hunger and defense?
of f e a r ” and the “chemistry of pleasure.” It is We cannot answer this question yet. But the
quite legitimate to pose the question of what the answer may radically change our conceptions of
chemical essence of all these different ascending the most obscure nature of the working of the
impulses is at the cortical level, i.e., where brain. It is now quite clear that we have reached
the process of afferent synthesis itself takes the point where the purely functional and spatial
place. Do these excitations preserve the spe- properties of excitations are fixed in established
cific character of their subcortical chemistry, systems of molecular reciprocity. The chem-
o r does their molecular nature change? ical variety of reactions, processes, and mech-
A method of applying various chemical sub- anisms that a r e of decisive importance in af-
stances, whose effect on definite cortical cells f erent synthesis dictate this conclusion.
was known, helped us answer these questions. The deduction is clear: all these diverse
The application took place a t a moment when processes, directed toward the assessment of
these cells were producing a definite, known the subtlest properties of external afferent in-
electric charge. The changes in this charge fluences, a r e a completely independent and qual-
showed definite chemical alterations and could itatively delineated stage in the development of
 20 SOVIET PSYCHOLOGY

the behavioral act. At the same time all this in the normal manner the stimulus of the exter-
shows that the brain receives a multitude of ex- nal appearance of the biscuits must enter the
citations from every situation that differ both in central nervous system and establish contact
their qualitative meaning and in their functional with the collating apparatus which had been
purpose (consecutive, situational, initiating, etc.). formed immediately after the original action of
Only the complex work of afferent synthesis the conditioned stimulus.
permits the organic consolidation of the thou- Of course, the external appearance of the meat
sands of ascending excitations and their distribu- also stimulated the gustatory, visual, and olfac-
tion f o r the formation of a concrete aim in be- tory receptors of the dog. However, the excita-
havior. tions s o formed were not identical to those that
A s soon as the goal of the action has been arose in evaluating the excitations from the bis-
formed, very interesting and highly important cuits. Thus a “discordance” [rassoglasovaniye]
mechanisms come into play, whose physiological was produced between the stored complex of af-
purpose consists in assisting in the evaluation of ferent excitations and the real complex of excita-
the information which has yet to enter the cen- tions entering the central nervous system. At
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t r a l nervous system from the future results of the time these experiments were made (in 1933),
the action. This is a genuine apparatus for eval- the idea of “discordance” w a s not popular in
uating and collating results with an established physiology, and therefore the concept of “stored
goal in mind. The idea of its existence w a s sug- excitation” [zagotovlennoye vozbuzhdeniye] was
gested to us by some experiments with condi- introduced.
tioned reflexes. Here is one of them. Gradually we reached the hypothesis that there
For a number of years experiments were car- must be a special mechanism, formed before
ried out with the conditioned hunger reflex in a the action is completed and the result has ap-
dog accompanied by a ration of dog biscuits. Ac- peared, which simultaneously contains all the
cording to the standard ideas on the hunger reac- attributes of the future result. This apparatus
tion, the animal should have made no distinction has been called the action acceptor [aktzeptor
between biscuits and meat since both a r e non- deystviya] (from the Latin word “acceptare,”
conditioned stimuli. However, the experiment meaning to accept, to approve; “acceptor” -
showed that the substitution of meat for bread accepting, approving). This apparatus is in-
led to a marked orienting-investigatory reaction tended f o r interpreting information of an ob-
and a temporary refusal of food. We were inter- tained result and comparing it with the charac-
ested in the mechanisms of origin of this orient- teristics of the results which had already been
ing-investigatory reaction. The conditioned formed a t the actual moment of the action of the
stimulus was normal, a ration of food. The gen- conditioned stimulus. Further experiments have
eral reaction of the animal in the form of a se- aimed at discovering the concrete neurophysio-
cretory reaction remained the same. What was logical mechanisms which provide a basis for
lacking to make the animal react in the usual the formation of this anticipatory apparatus.
manner, i.e., to eat the food at once? What fac- A dynamic stereotype was established in a
t o r s provoked s o violent an orienting-investiga- human subject, using three stimuli (a siren, a
tory reaction in the animal? bell, and a light). A recording of brain waves
The analysis of this question convinced us that was made simultaneously. The electroencephalo-
use of the conditioned stimulus not only pro- gram showed that the f i r s t two stimuli did not
motes the formation of a conditioned hunger re- change the indices of electric activity of the
flex, but also a simultaneous formation of a com- brain. But the light produced a desynchroniza-
plex of excitations whose properties correspond tion of electric activity in the frontal sections
to a complex of signs and features of the impend- of the brain. Then the experimenter changed the
ing result (in this experiment, to the eating of order of the stimuli. When in place of the light
the biscuits). For the reaction to be completed there was a sudden ring of the bell, a quite
 VOL. V, NO. 1 21

striking phenomenon occurred; in spite of the stimulus. Each time, as soon as the current
absence of the light action, a desynchronization was turned off, the r a t began to p r e s s on the
took place as before in the visual area of the pedal more strongly to get the pleasant sensa-
cortex, as though the light had been given and tion.
not the bell. Analysis of the electroencephalo- Here we find the same law as in the cases al-
gram showed that desynchronization started a ready described. Each of the rat’s movements
little before the bell was rung. It follows, there- to the pedal contains a form of action acceptor
fore, that an excitation reached the visual area anticipating a pleasant emotional state from
before any external stimulus could get there. stimulation of the brain. When it fails to get
It is clear that, in constructing a chain of such a sensation, the r a t begins to strive for it
stimuli connected with conditioned-reflex laws, by applying more frequent and stronger pressure
the excitation is dispersed through the brain to the pedal.
from point to point far more rapidly than the ac- This experiment shows primarily that the ac-
tual external stimuli. The excitation forestalls tion acceptor is invariably formed at the very
the real stimulus which has yet to act upon the beginning of the formation of the behavioral act.
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central nervous system. So the idea of a special This is the only way in which the immediate ap-
kind of excitation arose that was called ”antici- pearance of compensatory devices can be ex-
patory excitation” [operezhayushcheye vozbuzh- plained if the information on results does not
deniye]. coincide with the properties of the excitation
The capacity of nervous tissue to organize a which were established when the aim and action
chain of processes by means of anticipatory ex- acceptor were formed. The sudden cutting off
citation is an elementary basis f o r the develop- of the current and the rat’s consequent non-re-
ment of the conditioned reflex and of any predic- ception of a positive emotional state at once
tion or, as it is expressed in cybernetics, “prog- create conditions for “discordance” between the
nosis’’ of future phenomena. This formation of expected results in the form of pleasant sensa-
a complex of excitations, in which the properties tion and its sudden absence (Yu. Makarenko’s
of future results are coded, ensures that the re- experiments).
sults obtained a r e compared with the reality Special studies of the respiratory center con-
posed by the conditioned stimulus o r totality of ducted by our laboratory colleague, V. A. Pol-
situational stimuli. The action acceptor is uni- yantzev, have shown that it has the same s o r t
versally present, and scarcely any behavioral of functional organization.
-
act is possible - even the simplest without By means of electronic devices it was con-
the preliminary formation of this apparatus. trived that the order given by the respiratory
We have studied the mechanism of prediction center to the respiratory muscles to take in a
under conditions of so-called self-stimulation. specific quantity of air was changed en route
The American scientist J. Olds worked out the and the lung received only a part of the neces-
well-known method by which a rat can stimulate sary air. This situation corresponded physio-
itself through electrodes introduced into the re- logically to the occasion when the dog got meat
gion of the hypothalamus. The experiment instead of biscuits. Naturally the lung receptors
showed that if the electrodes were situated in the signaled to the respiratory center that an insuf-
zone that forms a biologically positive state of ficient quantity of air had been inhaled. This
the “pleasure” type, the r a t would stimulate it- information of the actual result w a s clearly in
self uninterruptedly, sometimes pressing the “discordance” with that predicted by the respi-
button as often as 14,000 times an hour. ratory center. Therefore it immediately in-
We have somewhat modified this method s o creased the number of nerve impulses directed
that the experimenter can at any moment turn to the respiratory muscles. A s a result the
off the electric current and by the same means lungs took in far more air than was needed to
eliminate the positive emotional effect of the compensate for the insufficiency in the previous
 22 SOVIET PSYCHOLOGY

breath, Consequently the respiratory center, in solution to the problem of purposiveness in hu-
sending a command to the periphery to take in a man and animal behavior. In fact the very ad-
specific quantity of air, simultaneously formed aptation of the ‘reflex arc” as the central model
an apparatus for comparing the supposed result for explaining behavior for many years excluded
with the one received. all possibility of treating results as a motivating
Yu. Fel’dsherov and A. Karpukhina of our factor in the formation of variability of behavior.
laboratory adapted the microelectrode technique There was simply no place for results in the re-
to determine the formation of the processes of flex model and so their physiological character
prediction. They discovered those neurons of or, more precisely, their decisive role in form-
the respiratory center which receive information ing the functional systems of the organism re-
of the results of action. This experiment enabled mained outside physiologists’ field of vision.
u s to reveal the details of the formation of the When speaking of physiological apparatuses
action acceptor. that form the probable model for future results
Other observations have also been made of the which have not yet been obtained, we have fre-
action acceptor apparatus in human subjects. quently mentioned that the properties of the re-
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They all confirm that a universal law exists in sults obtained a r e the decisive factor determin-
the working of the brain. It could be formulated ing the further formation of the behavioral act.
as follows: whenever the brain sends an excita- Naturally the question arises of the central ner-
tion to the peripheral functioning organs, some vous system’s information concerning these
nervous model is formed at the same time as properties.
the ’command,” a model capable of anticipating We have called such information sanctioning,
the characteristics of the impending result and o r inverse afferentation [sanktzioniruyushchaya
collating them with the result actually obtained o r obratnaya afferentatziya]. Sanctioning af-
on completion of the action. ferentation is information about results that
This law is of particular importance in human fully coincide with an intended aim and, conse-
behavior, which can pursue the most varied quently, with the physiological properties of the
aims. And here the action acceptor, formed at action acceptor as well. Introduction of the con-
the moment of decision-taking, subsequently also cept of inverse afferentation has “closed” the
defines the degree of coincidence between the classical “reflex arc.”
intended and the actual result. Thus the cybernetic ‘closed” circuit with an
It seems strange that for s o many years the inverse connection was exhaustively formulated
results of action were never the subject of spe- and worked out in detail in physiology long be-
cial physiological analysis, since they a r e the fore the appearance of Norbert Wiener’s works
vital connecting link between the different stages on cybernetics. As early as the thirties, inverse
of the behavioral act. This is even stranger if afferentation from properties of results had ac-
we think of the true nature of behavior. As a quired a special meaning in the mechanisms of
matter of fact, men and animals are always in- the functional system. It enabled the expansion,
terested in the results of action. It is only be- on a true physiological basis, of the structure of
cause of them that often long chains of behavioral the ‘reflex arc” and the construction of a physio-
acts are embarked upon, for only failure to ob- logical framework in which the main mecha-
tain the desired results acts as the stimulus for nisms of all processes are connected in a single
further acts until what is attained corresponds causal system.
in some way to what is desired. In such a functional system inverse afferenta-
In the classical “reflex arc” there is no room tion, i.e., information about properties of ob-
for the evaluation of results. This is the most tained results, on being channeled to the appara-
outstanding failing of the reflex theory, which tus of the action acceptor, completes the whole
showed the dualism of its creator and for a long logical pattern of an individual behavioral act.
time distracted physiologists from a materialist This concluding phenomenon is the process of
 VOL. V, NO. 1 23

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comparison (collation) of the result properties mation. It is in this sense that the expression
of a completed act with properties which were “sanctioning afferentation,” which was used as
formed in the action acceptor during the process early as 1935, must be understood.
of afferent synthesis. The mechanism of this It is clear that this new model for analyzing
collation constitutes the most obscure aspects all the main mechanisms involved in the behav-
of our deliberate behavior. ioral act requires considerable additional elab-
How does the action acceptor apparatus work; oration and definition. However, the general
how is the model of a future result created within logical system of the behavioral act, from the
it? At the present time we a r e directly approach- motivating stimulus to the evaluation of results
ing the solution of this problem, one of the cen- obtained, is already sufficiently clear and I do
.
tr a1 problems of neur ophysiology not now envisage any other possibility for under-
From what has been said we can conclude that standing the entire behavioral act that would
the behavioral act is a process which develops change the model in principle.
in two stages. The first is the moment when af- It would, of course, be wrong to consider that
ferent synthesis has been completed, the aim of the search for new and more perfect models has
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behavior and the program of afferent action and come to an end because a model outlining the
the action acceptor have been formed. However, principles of behavior and its main mechanisms
the action itself, as a combined state of working has been constructed. Just as the functional
organs at the periphery, and, consequently, the system of behavior is itself an expansion and
result of the action as well, have not yet been transformation of the classical “reflex arc,” so
formed. the further elaboration of the functional system
In the second stage the action has already oc- itself may lead to the discovery of new proper-
curred. It leads directly to the obtaining of re- ties. The model of the behavioral act set forth
sults and to the inevitable influence of their af- above has made our work of studying the de-
ferent properties on the already-existing model tailed neurophysiological processes of the brain
-
of results the action acceptor. Inverse af- much easier and more comprehensible. It has
ferentation connects the results of the action become, metaphorically speaking, the “universal
with the physiological complex of the action ac- chart” of exploration by which the location and
ceptor which, in its entirety, is the probable af- significance of any experiment can easily be de-
ferent model for future results. Inverse af- termined.
ferentation is the passive consequence of re- Thus all our experiments and theoretical argu-
sults obtained at a given moment. Its role is to ments show how far a physiologist can go in his
give accurate information on the results. The investigations in this area of the minutest neuro-
action acceptor provides the corrective mecha- physiological processes with the general model
nism that must give final sanction to this infor- of the behavioral act at his disposal.