JPRS ID: 9217 TRANSLATION NEUROPHYSIOLOGY STUDY OF SYSTEMIC MECHANISMS OF BEHAVIOR BY V.B. SHUYRKOV

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~~'~TE1~ I ~ MECH~tI~ I ~~5 ~F ~EH~~' I
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JPRS L/9217
- 24 July 1980 -
Translation
NEUROPHYSIOLOGICAL STUDY OF
- .
,
Y SYSTEMIC MECHANiSMS OF BEHAVIO.R
: By
_ V:B. Shuyrkov
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JFRS i/9217 ~
24 July 1980
NEUROPHYSIOLOGICA~ STUDY OF SYSTEMIC MECHANISMS OF BEHAVIOR -
Moscow ~iEYROFIZIOLOGICHESKOYE IZUCHENIYE SISTENINYKH MEKH,ANIZMOV -
. POVEDENIYA in Russian 1978 signed to press 31 Aug 78 pp 2-240
[Book by V.B. Shuyrkov, Izdatel'stvo "Nauka", 3,150 copies]
CONTENTS ~
Annotation 1
Introduction 2
Chapter 1. Systemic Description of the Behavioral Act =
Qualitative Distincticn of Behavior From Elementary
Physiological Processes 7
Goal Orientation of the Behavioral Act 11
Isolation of the BehavioraZ Act in the Continuum of Behavior 20 -
Organization of Physiological Functions in the Behavioral ~~ct 25 ~
, ~perational Architectonics of the Functional System in the -
Elementary Behavioral Act 29
Chapter 2. Electrophysiological Correlates of Systemic Processes
in the Elementary Behavioral Act
Electrical Activity of the Brain in Behavior 34
, Synchronism and Similarity of Configuration of EP of Various
Struc~tures in Behavior 37
Link Between EP and Time of Behavioral Act 43
"Endogeny" of EP in Behavior ~ 45
Link Between EP ~:~d Future Events 47
EP Components--Correlates of Systemic Processes of the
Behavioral Act 56 _
-a- LI -USSR-CFOUO]
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Chapter 3. Systemic Organization of Neuronal Activity in Behavior
Link Between Overall Activity and Neuronal Tmpulsation 74
Link Between Neuronal Activity and EP 75
Synchronism and Similarity of Neuronal Discharge Patterns in
Various Brain Structures ~g
Determination of Neuronal Discharge Pattern by Pretriggering
Integration 90
Involvement of N e~ons in Systemic Mechanisms of the
Behavioral Act 95 `
Chapter 4. Mechanisms of Transformation of External Information Into
- Organization of Processes in the ~unctional System of a
Beha.vioral Act
Relationship Between Prior Experience, Motivation and Information
= About the Current ~tate of the EEAVironment in Determination of _
Goal-Directed Activity 118
Organization of Memory 12~
Use of Exogenous Info`-mation to Organize Purposeful.Neuronal
_ Activity in the Behavioral Act 133
Role of Goal in Organization Processes 149
Involvemen~ of Different Regions of the Brain in the
Functional System of the Behavioral Act 164
Chapter 5. Mechanisms of Involvement of a Single Neuron in the
Functional System of the Behavioral Act
MPChanisms of G~neration of a Goal-Directed Pattern 176
~ Correlation Between Functional Synaptic Fields in
Pretrigg~ring Integration lgg
Chapter 6. Functional System Theory and the Psychophysiological
- Problem
Impossibility of Direct Correlation of Mental and Neurophysiological
Processes 197
The Problem of Correlation of' Systemic and Mental Processes 199
Correlation of Systemic and Neurophysiological Processes 204
Conclusion
Bibliography 209
- b -
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PUBLICATION DATA
English title . Neurophysiological Study of Systemic
- Mechanisms of Behavior
Russian title : Neyrofisiologicheskoye izucheniye
~ sistemnykh mekhanizmov povedeniya _
Author . V. B. Shvyrkov
Editor . K. V. Sudakov, corresponding me~aber
- of the USSR Academy of Medical Sciences
Publishing house . Nauka
Place of publication . Moscow
Date of publication . 1978
Signed to press . 31 August 1978
Copies . 3150
COPYRIGHT . Izdatel~stvo "Nauka", 1978
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ANNOTATION
This study deals with analysis of behavioral mechanisms from the stand-
point of functional system theory created by P. K. Anokhin, and it sub-
stantiates the need for the systemic approach to investigation thereof.
From the point of view of P. K. Anokhin's theory, the elementary beha-
vioral act is considered as a cycle of "exchange of information" between
the environment and the organism. Mechanisms of involvement of an
individual neuron in the system of the behavioral act are examined. -
There is discussion of correlations between mental, systemic and neuro-
physiological processes in behavior. _
. 1
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INTRODUCTION
The behavior of living organisms is the subject of investigation of many
disciplines, in each of which special aspects of behavior are studied. '
This circumstance, as noted by R. Hinde (1975) in the preface to his
book, does not allow us to define the concept of "behavior." However, ;
for many areas of research, including neurophysiology and psychology,
behavior in the most general sense can be defined as the relations of ~
an organism and the environment. For this reason, the study of behavior
should include analysis of both the environment and processes within an
= organism, and interaction between the organism and environment. The
, concept of "behavior" should pertain to all forms of interrelations '
between the organism and environment, including those that are reflected
in the psychological aspect of the organism.
At the present time hardly anyone will deny the role of the psychological
factor in behavior. Yet, it is obvious that behavior is based on physio-
logical functional processes of specific sorphological structures of the
organism. The correlation between mental and physiological processes
constitutes the so-called psychophysiological problem.
The materialistic thesis of unity of behavior and the psyche rules out the
possibility of gaining full knowledge about the mechanisms of behavior ~
without determining the role of inental processes~in behavior. Any
behavior theory that "throws out" or excludes mental processes is not,
in our opinion, consistent with reality, since it is expressly through
mental processes, through informational correlations that the external ~
environment determines behavior, as reflected in the theses of reflecting ,
and regulatory role of the psyche in beb.avior. At the same time, the unity
of behavior and the psyche implies that it is impossible for mental pro-
cesses to occur apart from behavior, i..e., certain physiological processes. ~
Thus, a given solution to the problem of inechanisms of behavior of ~
- necessity leads to a given solution of the psychophysiological problem ,
as well. '
- The psychological problem cannot be solved solely on a physiological or
solely on a psycnological basis; consequently, neither physic,logy nor
psychology can offer a complete description of behavior. Nor can this
be achieved by means of direct correlation of inental and physiological
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processes. As validly noted by A. N. Le~nt~yev (1975, p 7), "the fact of
the matter is that no direct correlation of inental and cerebral physiolo-
gical processes solves the problem. The theoretical alternatlves that
arise with such direct comparison are weli-known: there is either a
hypothesis of parallelism which inevitably leads to interpretation of
the psyche as an epiphenomenon, or a thesis of naive physiological deter-
minism with the ensuing reduction of psychology to physiology, or else,
finally, there is a dualistic hypothesis of psychophysiological inter-
action, which assumes. that the nonmaterial psyche affects tangible
processes occurring zn the brain. There is simply no other solution for
metaphysical thinking, only the terms change to refer to the same alterna-
tives."
It is presentiy bzcoming obvious that synthesis of psychol~gy and physio logy
_ to describe bel:yvior is possible only on some higher basir~ common to both
disciplines. The systemic approach is such a basis, and it is now being
developed in many areas of knowledge (Anokhin, 1�73a; Ke3rov, 1974;
_ Kuz'min, 1976; Lomov, 1975, and others). Of the many variants of the
systemic approach, functional system thPOry, which was dev~loped by
Academician F. K. Anokhin (1935-1974) aF~pears to us to be the most adequate
to the problems of physiology and psychalogy and the task of their synthesis
in the description of b~~havior. This theory proceeds from the most general
- biological theary, theory of ec:,lution, to explain behavior.
L~~ Indeed, unlike many variants of the systemic approach in biology, which
_ propose to study the properties of systems on formal models (Mesarovich,
1970), functional system theory is er.tirely based on biological facts, and it
uses the concept of survival, or useful adagtive result, as the foundation
for the method of isolating the system. Like all fundamental.initial con-
cepts (Kedrov, 1962), the concepts of system and result are definad
in functional system theory through the relationship between them. T'ne
result is a state of tti~? environment that allows the system to survive.
The system is an aggregate of elements so organized as to achiev~ this
result. Survival is the main result that is ultiimately reached by bio-
logical systems. Hence, the behavior of biological systems is goal-
oriented, and any behavior occurs to reach some useful adaptire result
that ultimately leads to survival.
Of course, there are very diverse forms of interactions between an organism
_ and the environment; behavior can be defined as interaction, in which both
the organism and environment are whole. Then behavior wi11 appear as a
two-way exchange of organization or information between the environment and
organism, which can occur only by means of informational or specifically
_ systemic processes th~t cannot be reduced to separate physiological pro-
cesses or separate effects of the environment.
Systemic processes describe the state of both the organism and environment;
for this reason, a neurophysiological or psychological description of
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- behavior is a particular description of the same systea~ic processes of
' correlation between the integral organism and environment.
From this point of view, the dESCription of the correlation between the
organism and environment in terms of systemic processes should constitute
the special subject of "syst~mology." Correlations between systemic and
elementary neurophysiological processes are the relations between informa-
tion and its material carrier, since systemic processes are distinctive
informational processes (Ferster, 1964; Gorskiy, 1974). But the correla-
tions between mental and systemic processes ar~� the relations between -
internal and external ifnormation. External lnformation is the organiza-
tional attribute of environmental elements, ~while internal information
- is that of organization of elements of the organism. Thus, one can com-
_ pare neurophysiological and mental processes only through qualitatively
unique systemic processes, which exist in the organism as processes of
organization of va~ious elements into a s~ngle whole, a functional system.
- Since systemic processes, one asp~ct of which is the psyche, are represented
in the organism by processes of expressly organization of physiological
functions, this view avoids equating mental and physiological processes.
. It also avoids psychophysiolog~.cal parallelism, since sysLemic processes
are processes of organization of expressly physiological functions and the
psyche is the product of the brain. Since internal organization is de-
, termined by organization of the environment, i.e., its object structure,
the psyche cannot be excl.uded from analysis of inechanisms of behavior.
Finally, since systemic processes "consist" exclusively of physiological
processes and a new quality is attained exclusively as a result of their
organization, physiological and psychological dei:ermination of behavior
is inseparabl~ united and the two do not exist without one another, which
precludes any psychophysiological interactions.
Evidently, this point of view is consistent with conceptions of correla-
tion between the psyche and brain as information and its carrier, which ~
are being developed from the philosophical point of view (Ponomarev, 1967;
Dubrovskiy, 1971, 1976). Thus, functional theory system serves as the
basis on which one can find an experimental solution to both the problem _
of inechanisms of behavior and the psychophysiological problem. From
the standpoint of this theory, neurophysiology of behavior and the psyche
' can be interpreted as the study of systemic processes of exchange of or-
ganization between *_he integr.al organism and objective environment using ;
_ neurophysiological nethods.
In research on behavior, the problem of elementary phenomenon was always ~
considered the key one, whi.ch determined all subsequent theoretical
developments and direction of research. Since the times of Descartes,
the organism's reaction to some environmental agent was alwais considered
to be an elementary manifestation of behavior. There is a certain inter-
val between a"stimulus" and the "reaction" to it, and in difierent
- aspects. it is referred to as "delay," "reaction time," "reflex time," etc.
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The poler~ics on the subject of processes that occur in this interval
gathered, or concentrated, all of the contractions in. psychology,
physiology and other disciplines that deal with th~ brain and behavior.
- The problem of determination of behavior, the psychophysiological prob- -
lem, the problem of localization of functions, cybernetic proolems of
information coding and regulation of the organism's relations with
the environment, and all other general biological problems of behavior
and the psyche are related in sone way or other to determination of the
~ mechanisms of the elementary behavioral act.
- From time immemorial, such mental processes as perception, comparison
[ccallation], remembering, etc., have been attributed to this interval
also. Measurement of this interval in different experimental modifica-
tions is widely used to describe the most diverse mental processes and
states; and it is even believed that "the method of ineasuring reaction
- time is the best method for studying higher functions, and it has a.
great future" (Shoshol', 1966, p 316).
In spite of the complexity and diversity of processes that one relates to -
_ elementary behavioral acts, for a long time the neuraphysiological inter-
pretation of processes occurring between the "stimulus" and "reaction"
amounted to conducti.on of stimulation from receptors to effectors, as
was dictated ~y reflex theory.
- The conception of the behavioral act as a reflex was not based on direct -
studies of neuronal mechanisms of behavior and not on physiological facts
or even anatomical conceptions, but exclusively on the ideas of inechanistic
determinisin. In his "Answer of a Physiologist to Psvchologists," I. P.
Pavlov wrote: "It is generally accepted that the concept of reflex
originates with Descartes; but what was known about the detailed construc- -
tion of the central nervous system or about its activity in the times of .
Descartes? Physiological and anatomical separation of sensory nerves from
_ motor nerves occurred only in the early lOth century. Obviously, the
idea of determinism was for Descartes the essence of the concept of
reflex, hence his conception of the animal organism as a machine. This
was the interpretation of reflexes of all subsequent physiologists, who
_ related different activities of the organism to ditferent stir~~:li, gradu-
ally isolating elements of neural structures in tfie form af various affer-
ent and efferent nerves, and in the form of special pathways and centers -
of the central nervous system, and finally also gathering the typic,al
dynamic features of the last mentioned system" (1949, p/~95).
Even at the moment of its inception, the idea of reflexes "made the
first breakthrough in the strong wall of mystical and religious concep-
~ tions that separated the researcher from real facts" (Anokhin, 1945, p 6).
_ The principle of determinism, contained in the reflex concept was not
only used to fight against interpretation of behavior from the te].eolo-
gical positians of idealism, but served as a natural methodological basis
for experimental research on the nervous syste~. The contemporary
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- advances in neurophysiology became possible only on the basis of the ana-
lytical approach, which enabled neurophysiologists working with local
processes or substrates to use the same approach that had been used and ~
_ glo~~ingly justified itself in mechanics.
At the present time, the principle of "mechanistic determinism" (also -
qualified as "linear" and "naive physiological") as applied to interpret
biological processes and behavior is being critized from the most varied
positions, including philosophical (Dubrovskiy, 1971; Serzhantov, 1974), -
- cybernetic (Menitskiy, 1975; Svinitskiy, 1976), psychological (Lomov, 1975),
biological (Oparin, 1964) and neurophysiological (B urns, 1969; Relenkov, -
- 1975, 1976; John, 1973; Sudakov, 1976, and others).
Although it was obvious to many, rather long ago, that the reflex inter-
pretation of the elementary behavioral act was unsatisfactory, for a long
time more constructive solutions of this problem were delayed by the
fact that considera~le revision of the entire system of conceptions that
had been formed would be necessary to reject ;.entury-old reflex traditions
in physiology. As noted by B. Burns with reference to one of the earliest
and most striking critics of the reflex, "Iashle~ s argumentation was weak
because Lashley quantitatively tested the reflex or telephone
theory of behavior and found it to be invalid, but did not offer any other =
' promising system of concepts" (1~09, p 19). _
Functional system theory created by P. K. Anokhin provides such a system
of concepts. V. F~ Serzhantuv believes that "acceptance of this concep-
tian leads to certain consequences for the entire theoretical system of
- biology and psychology: the principle of functional system permits
deeper interpretation of biological and psychological concepts formulated
in science to this time; hence the need to reorganize the entire concep-
_ tual structure of these areas of science" (1974, p 74).
Application of the conceptual apparatus of functional system theory to
- problems pertaining to the elementary behavioral act alters radically
the very methodology of research. For this reason, analysis of the neuro- _
physiological mechanisms of the elementary behavioral act from the
positions of functional system theory requires preliminary consideration -
- of behavior in the terms of this theory.
We shall make such an examination as compared to known and customary
conceptions of reflexes; however, our main goal will not be to compare -
_ the two approaches, but to define the subject of investigation and
formulate concrete problems, for which experimental neuraphysiological
solutions must be found.
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CHAPTER 1. SYSTEMIC DESCRIPTION OF THE BEHAVIORAL ACT
_ Qualitative Distinction of Behavior From Elementary Physiological Processes _
~ The psychological description of relations between an organism and the _
environment includes such concepts as memory, motivation, percepti~~n,
action, emotion, etc., i.e., concepts that characterize the organism as
- a whole, as a subject. The environment is also described in psychology
as "objective" [objectt-related], and the correlation of an integral or-
ganism and objective environment emerges as the correlation between a
subject and object. As validly observed by L. M. Vekker, "the ultimate, .
final characteristics of any mental prucess in the general case can only
_ be described in terms of properties and relations of external objects.
Thus, perception or a conception cannot be described in other than
terms of shape, size, consistency, etc., of the perceived or imagined
object. Thought can be described only in the terms of the features of _
objects, the relations between which it discloses; emotion can be des- ~
cribed in terms of attitude toward events, objects or individuals that
induce it, while voluntary decisions or a volitional act cannot be
expressed in other than terms of the events in relation to which the _
action or deed is performed" (1974, p 11). �
Thus, psychology describes the relations between the organism and environ-
ment in terms of properties and relations of expressly environmental -
elements. This is an extremely important aspact of behavior; however, -
psychological concepts do not describe internal processes at all, i.e.,
processes that take place in the organism, since "phenomena of subjective
reality constitute information given to the person so to speak in 'pure
form (Dubrovskiq, 1976, p 54).
Internal processes have always been referred to the area of physiology, -
which has its own conceptual system. From the very beginning, physiology
developed as an experimental and analytical science. Neurophysiological
concepts., such as stiznulation;~ excitation or inhibition, afferentation or
efferentation, etc., were created to describe processes occurring in
diffP:.ent, morphologically distinguishable organs or nerves. For many
reasons, for a long time experimentation was possible only on preparations
of animals, in which, o.f course, there is no behavior.
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In undertaking development of physiology of behavior, I. T'. Pavlov intro-
~ duced into the practice of physiological experimentation w~.~rk with integ-
ral organisms. It is expressly to I. P. Pavlov that we o*:e formulation
of tne task of physiological study of behavior, which l:as not losr its
meaning to this day. Already in 1903, he o~+~erved; " The tremendous com-
_ plexity of higher, as well as lower, organisms continues to exist as a
' whole only so long as all of its elements are finely and precisely
linked, balanced with one another and with the surroundings. Analysis -
of this balancing of the system con~titutes the first and foremost task -
_ and goal of physiological research as purely objective research" (1949,
p 337).
_ However, having formulated the task of studying inte:gral behavior, neverthe- `
_ less the Pavlovian school first concentrated its expprimentation on the
study of actual function of a single salivary gland, rather than mechanisms
of integral behavior, and this was of decisive significance to developsent
~f the entire conceptual system of the teaching on higher nervous activity.
= Having concentrated its efforts expressly on analysis of brain function,
the teaching on higher nervous activity used the analytical concept of
"reflex," which already existed in physiology and was developed to des-
cribe processes demonstrated in preparations, i.e., expressly beyond _
integral behavior, as the foundation for conceptions of the mechanisms of ,
the integral behavioral act. For this reason, the descriptions of
behavior of an integral organism and processes occurring in preparations
turned out to be identical. ,
~ With reference to adaptive behavior, I. P. Pavlov wro te: adaptation
is based on a simple ref lex act, which is initiated by certain exogenous
= conditions that affect only a specific kind of endings of centripetal
nerves, from which stimulation passes over a specific nervous pathway to
- the center, from their to a gland, also over a specific pathway, thus
causing specific function in it" (1949, p 334). Application of the ana-
- lytical concept of "ref lex" to analysis of inechanisms of integral behavior -
resulted in setting aside from the main line of physiological research '
the qualitative specifics of expi�essly integral behavior.
Confusion of concepts ~.escr~'~ing the function of disconnected physiological
mechanisms and the integral organism made it impossibl e, for a long time,
to see the actual problem of integrity, since "excitation of neurons" -
= ultimately produced "excitation of a center" and even "excitation of the
brain," while "inhibition of a reflex" was attributed to "inhibition of ~
neurons" of the corresponding centers. This "energet ic" description of ,
processes occurring in the organism and implementing behavior also required
an "energetic" description of the environment as an aggregate of different
"stimuli" or "irritants." The correlation between the organism and
environment with reference to behavior was actually reduced to conformity
between elements of stimulation and elements of reaction. '
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Applicatio:~ of the analytical concept of "reflex~~ to descx'iptian of integral
behavior appeared to disclose ~he po~s9.bility of describing ~:nternal
processes of behavior in traditiQnal physiological concepts describing the
state of different organs and ti,ssues. However, this approach also closed
the way for relating "properties and relations of external objects" to
processes within the organism. Tndeed, if, as is assumed in reflex theory,
relations between the organism and environment consist of element-by-
- element conformity between stimuli and reactions, some sort of special
concepts would be utterly superfluous to describe "properties and
relations of external objects" and processes of correlation between
- expressly the integral organism and objectiv~ environment. As we know,
= this circumstance had dramatic consequences with re~ard to the contact
_ bztween physiology and psychology, and made it impossible to develop a
_ conceptual system comnion to these two disciplines that describe behavior.
Development of the ideas of I. P. Pavlov concerning the systemic nature
of higher nervous activity led to creation of functional system theory
- (Anokhin, 1935-1974), which reflected the qualitative uniqueness of inechan-
isms of integral behavior, as compared to reflex mechanisms of spinal
preparations and anesthetized animals. As observed by V. F. Serzhantov,
11functional system theory grew from reflex theory in its Pavlovian inter-
- pretation, it is a continuation of the latter, but at the same time it is
- also a negation thereof in a certain sense. However, this is dialectical
negation" (1974, p 70). -
P. K. Anokhin expounded functional system theory on the basis of physiolo-
- gical facts that disclosed the Qualitative specificity of processes of
integration of different physiological processes into a single whole, the
functional system of integral behavior. This disclosed an absolutely new
- type of processes in the integral organism, a type of systemic processes
- or "processes of organization of physiological processes."
Discovery of systemic processes in the organism automatically leads to -
a certain interpretatian of both the environment and correlation between
the organism and envir~nment. According to functional system th2ory, unlike
material-energetic relations between a local stimulus and local reflex
reaction occurr~ng in anesthetized or spinal preparations, beha~~ior is -
a means of two-way informational correlation between the organism and
environment.
Aighly organized organisms exist in an organized environment; in the course
of evolution they had to adapt to such environmental factors as behavior of
the prey.or predator, availability of mater3.a1 to build a nest, behavior
of sexual partner, etc. Al1 these adaptations required integral evaluation
~ of different material-energetic factors and attitude toward their specific
- organization as to a whole, i.e., an object.
_ The environment affects different receptors of the organism in the form of
different, separate energies; Che object, i.e., organized aggregate of
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_ environmental elements, may even find passive reflection only in organization _
_ of activity of many receptors, when the organism emerges as a whole. There
- is reflection in hehavior of the environment, not only ob~ectively, but -
actively; the organism constantly searches for and obtains the information
it needs, unlike a preparation whtch is indeed "submitted to the effect of
a stimulus." '
= Processes within the organism referable to behavior can also not be reduced
_ to energy processes of "excitation and inhibition." Any aggregate of ~
excited elements per se does not create the phenomenon of behavior. It is ~
expressly processes of coordination of specific elements and organization .
thereof in a singlz who]e, in which everything is "finely and precisely
related, balanced within itself and with external conditions," expressly
_ these processes of organization constitute the essence of internal mechan-
isms of behavior of the integral organization, and not "stimulation of .
cells af a functional organ" per se.
_ The systemic approach compels one to consider behavior as the correlation
between organization of the environment and organization of processes
within the organism. And determination of behavior by the external environ-
ment emerges as determination of organization of processes within the
_ organism expressly by organization of the environment. ~
Just as "life is characterized by a special, specific combination of
- properties, rather than any particular properties" (Oparin, 1924, p 36), so i
_ behavior is not referable to some special processes, but to specific "
organization of processes on the physiological level. Processes of
organization are qualitatively specific and bilateral: environmental ~
organization determines organization of processes in the organism, ;
which in turn leads to organized influences of the organism on the
environment and new organization of the environment, etc. This entire, ;
continuous cyclic process is designated in systemic terminology by
the general term, "behavior." ~
~
- As we know, biological existence of an organism is implemented by absorp-
tion of organization, or "negentropy" of chemical bonds (Schroedinger, 1947). -
This principle is common to all living things, regardless of complexity
of the organism. The behavior of multicel?.ular and particularly highly
organized organisms can be viewPd as development of this capacity and use
by the organism, to maintain its integrity and organization, not only ;
of organization of chemical bonds, but other, higher forms of organization
of the environment. In this regard, "adaptation of organisms to it ~
acquires a qualitatively new form, which is related to reflection of ~
_ objectively object-related reality" [Leont'yev, 1972, p 49).
Thus, behavior as a qualitatively specif ic form of adaptation of the integ-
ral organism to the objective environment is based on systemic mechanisms
of organization of different physiological processes into a single whole,
a functional system. ~
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Functional system theory provides the hasis for descrihing hehavior in
_ terms of systemic processes proper, i.e., processes o� cprrelation between
external and internal organization. Exchange of organization between the
organism and envixonment can only he descrihed by systemic categories,
which characterize the environment azd processes in the organism from the
standpoint of correlation [compariscn]. The environment must be charac-
terized not oniy by a specific orgar~ization of its elements in time and
= space, but existence in the organisn studied of the capacity to make use,
in some way or other, of this organization of environmental elements in
behavior. In turn, processes in the organism must characterize not only
a specific organization of elements of the organism, but the link between
these organization and certain exogenous events. Therefore, the concepts
of functional system theory, such as "goal" or "result," "memory" or
"motivation," refer both to specific organization of the environment and
specific organization of elements within the organism.
At the present time, actually only the "skeleton" of the system ic conceptual
apparatus has been created, and different concepts will be constantly
defined; however, such definition must be made on the basis of concrete
factual material. Description of these processes in terms of "properties
= and relations in objective-object-related reality" is the subject of
psychology; their description in terms of activity of endogenous elements
of the organism is the subject of behavioral neurophysiology.
As noted by K. Lashley, reflex theory "has the advantage of simplicity,
which explains its popularity as a slogan" (1933, p 188). Systemic
_ categories do not have this aclvantage. They are not referable to
traditional or intuitively obvious categories; nevertheless, for the
methodological considerations stated above, the objectives of neurophysio-
logical studies of behavioral mechanisms should ensue expressly from a _
systemic description of behavior as exchange of organization betw~en the
organism and environment. ~
Goal Orientation of the Behavioral Act
The purposefulness of behavior of living organisms has actually never been
completely denied, since even mechanicism, which considered a"reaction"
to be the immediate consequence of a"stimulus," was also compelled to
recognize at least "seeming" purposefulness of behavior. This recognition
ensued from the adaptational nature of behavior directed toward survival
of the organism. While rejecting the concept of "goal" to interpret a
specific behavioral act, not a single biologist could deny that "all life
is the pursuit of one goal, namely of preserving life itself" (Pavlov, 1951,
p 33).
The conviction taken from mechanics, that the only scientific interpre-
tation is interpretation in terms of linearly related "causes" and "effects," _
and at the same time the obvious orientation of behavior of organisms
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toward reaching the goal of "surviving" generated numerous attempts ~
at explaining the purposefulness of behavior without using the concept
of "goal." This situation was cleverly described by the expression:
"Teleology is a lady, without whom not a single biologist can live,
but he is ashamed to appear in public with her" (quoted in: Mesarovich,
197Q).
. ;
The law of Thorndike's effect, which postulates that there is a retro- ~
_ active effect of the result of action on the "stimulus" and ~'reaction"
link, the conceptions of the Pavlovian school about "copying" an un-
conditioned (already adaptive) reaction to a conditioned signal, as well
as the inborn permissive [oz resolving] mechanism of ethologists--all
these are attempts to explain purposeful behavior through cause and effect
relations between "stimulus" and "reaction." This desire is attributable
to exclusively general philosophical considerations, since any behavior ~
is a continuum of behavioral acts, and in reality it is much more
convenient to classify natural behavior according to "actions" and "re-
sults," as is done by zoologists (Chauvin,1972), rather than according
- to "stimuli" and "reactions." The independence of reflex formulatian of
the problem of behavior from the subject of research proper can be very
graphically seen in the book by R. Hinde (1975).
While indicating the considerable advantages of describing behavior ~
according to the results attained and noting that "a description
according to consequences is often absolutely necessary for a complete
description of behavior"(p 21), R. Ninde nevertY~~eless views the problem
of behavior as "establishing a link between the phenomena studied and
events and conditions that immediately preceded them. Such analysis
is usually called 'causative analysis'"(p 12). ,
Both the natural forms of behavior, such as "food searching," "nest
building," "sexual" and "instrumental" behavior observed in experiments,
and such facts as the relationship between chemistry of saliva and
composition of future food, demonstrated in the classical experiments
of I. P. Pavlov--all these observations were a direct indication that
both the integral behavioral act and any behl~~ioral reaction are '
directly governed by future, rather than prior, events.
The obvious link between a given form of beha~vior and future events, or ~
results, also failed to serve as the theoretical basis for analysis of ;
behavior exclusively for general, philosophical considerations, since
it required an utterly different methodological approach. ~
The critical comments directed toward reflex theory and "causative`
analysis" of behavior, which became particularly numerous in the last ;
d~cade, made it absolutely obvious that mechanistic determinism (also
called linear, naively physiological, etc.) could not explain the behavior _
of living organisms. However, as noted by P. K. Anokhin (1962b), rejection
of inechanistic determinism led to teleological concepticns, which were
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found to be idealistic, since in the history of science, as a rule the
recognition of purposefulness ~.n living nature was set against materialistic
determinism. -
- The conceptians of purposeful behavior fe11 i.nto the stream of philosophical '
systems that extended the concept of "goal" to nature as a whole, which led
- to "vitalism," finalism," recognition of "entelechy," etc.
One can find a critique of these conceptions in recent philosophical
works (Volkova,et al., 1971; Ukraintsev, 1973, 1976, and others). Demon-
strating the inapplicability of philosophical te~.eology, the authors
- arrive at the conclusion that the goal-oriented approach to analysis of
biological phenomena is justified and absolutely mandatory.
~ Behavioristic theories such as "~~ymbol what it designates" (Tolman, 1951},
- as well as conceptions of "extrapolation reflexes" (Krushinskiy, 1967), of _
behavior guided by images (Beritov, 1961), "TOTE unit" type of concepts
(Millar, Galanter, Pribram, 1965), cybernetic behavior theories (Ashby,
1962; George, 1963), made a significant contribution to interpretation of
determination of future.by future events, but all of these conceptions
recognized.purposefulness of behavior, along with the reflex, which appeared
to be a satisfactory explanatory principle, at any rate, at least on the
level of some physiological mechanisms.
The "firmness" of the reflex in physiology was in contradiction to purpose-
- fulness of behavior as a whole, which led some to maintain that it was ~
"premature to physiologize" to interpret behavior (Tolman, 1951) and others -
to use reflexology even to explain human behavior.
At the present time, there are apparently few who would question the pur-
posefulness of human behavior, although there have been both philosophical
and physiological attempts to interpret human behavior from successively
reflex positions, a summary and critique of which can be found in the
book by Ye, A. Budilova (1972).
At present, it is imperative to have the goal category in an explanation of
human behavior (Leont'yev, 1975; Gal'perin, 1976; Bekhtereya, 1974). How-
ever, for some authors,. an obstacle arises when the conclusion of purposeful-
ness of behavior is extended to animals, which consists of consciousness
of human activity, the "humanitarian nature" of the concept of "goal," or
"anthropomorphism" of such extrapolation. These obstacles appear contrived,
since the concept of "goal" fn its application to analys~s of animal
behavior can be used without the adjective "conscious," ~~ing this term to
refer to the future, for which the behavior occurs. In this interpretation,
the concept of "goal" can apparently be used with equal significance to
describe both the behavior of a man who goes shopping in a store, and
of the earthworm who crawls up to the earth's surface for leaves and grass.
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- The systemic "goal" category refexs. to an eventfor which behav~.or occurs.
- Like all systemic categories, "goal" characterizes the relati.onships between
an integral organism and organized environment; ~or this reason, this _
category also refers to a specific organization of the environment, as -
well as specific organization of elements of the organism. -
- Functional system theory extended the principle of gurposefulness to all
levels of analysis of behavior and all physiological mechanisms, on which
behavior is based. Successive application of the principle of purpose-
fulness permits solving a number of "paradoxes" and creating a unified
and orderly system of concepts to explain both integral activity of
the organism i~? behavior and elementary neurophysiological processes con-
- tained in behavior.
The purposefulness of all biological processes is related to the very
~ history of appearan.ce of Iife on earth. Observing that "the analogy drawn
by mechanists between organisms and machines cannot by any means explain
the very thing it is called upon to explain, the 'purposefulness' of
organization of living beings," A. I. Oparin stresses that differentiation ~
of integral multimolecular systems from the primeval sottp could only
occur by virtue of the fact that the association of several molecules
enabled this structure to interact with the environment as a whole and
to preserve integrity. "By virtue of their differentiation, the emergence _
of such systems does not represent anything unique: at first these were
simply isolated regions in the primeval soup." And "any, even scattered !
~ chemical processes taking place in a drop, let alone some combination
or other thereof, were not indifferent to its subsequent fate"(1964, p 27).
Some of them aided in, while others prevented retention of the integrity
of multimolecular systems. "This is the route, already at the early
stage of evolution of coacervates, along which a form of selection
arose of the primitive syst:ems, according to the feature of conformity
y of their organization with the objective of preserving a biven drop
under conditions of its continuous interaction with the environment.
It is expressly on the basis of this new pattern, which emerged in the
very process of inception of life, that there was formation of the
metabolism inherent in all living things, a combination of different
reactions that, as an aggregate, was 'purposeful,' for constant self-
preservation and self-reproduction of living systems under prevailing
environmenta~ conditions" (1964, p 28).
The purposeful organization of different chemical processes constituting
- metabolism became enriched during evolution with more and more, also~ ~
purposeful additions. This is how D. Kenyon and G. Steinman describe how
metabolism became more complex in evolution: "There had to be a tim~
when the most easily assimilated nutrients (A) would be entirely used up;
then the eobionts (primitive prototypes af living cells), which were
capable of producing A from other available compounds (B) gained the _
_ advantage. When, in turn, the amount of secondary nutrients (B)
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diminished, it became necessary for A and B to furm from C, and so on.
Acquisition of the appropriate catalysts to accelerate these reactions
determinec? the degree of complication of this process" (1972, p 269).
All subsequent evol~tior~ and all, even qualitative, comglication of organiza- -
tion of biological systems and those derived from them were thereby -
guided by the same "system-forming factor" (P. K. Anokhin), the result that -
increased their chances of survival. This "patenc" significance of the
result to determination of purposeful behavior of systems with different
levels of organization was constantly stressed by P. K. Anokhin: "The
very appearance of stable systems with elements of self-regulation hecame _
, possible only because the first result of such seif~regulation emerged in
the form of stability itself, capable of withstanding exogenous factors.
Consequently, the regulatory role of the result of the system was the
first moving factor of development of systems, which accompanied all
stages of prebiological, biological and social development of matter"
(1975, p 339).
In any concrete study, we find contemporary organisms at a certain phase of
evolutionary development, when their structure reflects the entire history
of their survival. Since only purposeful forms of activity of organisms
were selected and structurally fixed in the course of evolution, the genetic
memory of organisms could only contain potentially purposeful behavioral
acts whichy under any conditions, led ultimately to survival of the organ-
- ism. Individually acquired behavioral acts were superimposed over innate
ones in accordance with the same evolutionary principle of survival.
The aggregate of all innate and acquired acts constitutes the general
stock of adaptive behavior of animals, which differs in different species
and specimens. This stock is life experience, or memory of the organism.
The systemic "memory" category refers to the aggregate of specific organi-
zations of elements of the organism that corresponded.~in the past to
some behavioral acts.
Aside from learning processes, adaptive behavior can be gleaned only from -
the store of inemory. For this reason, behavioral acts cannot in principle
be other than purposeful. According to functional system theory, the
selection of some particular goal out of all the material in memory and
of one behavioral act conforming with this goal occur under the influence
_ of several factors, designated as "motivation," "situation" and "trigger-
_ ing stimulus." The interaction of these factors is referred to as -
_ "afferent synthesis."
On the leuel of highly organized animals, the main goal of life, to
survive through the demands of tiss~rlar metabolism and homeostatic
mechanisms, is man~.fested in the form of motivations of behavior. Adapta-
tional behavior cannflt be unmotivated. Functional system theory makes
full use of the idea, vviced by I. M. Sechenov: "Vital needs generate
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desire, and then it leads to action; desire will then be a motive or goal,
while movement will be the action or means of reaching the goal. Movement
would be senseless in general without desire as a motive or impulse (1952,
p 516).
Motivation as a systemic category is concrete definttion of the goal to
survive. "Motivation retrieves all (behavioral) acts from memory that
- had at one time been related to satisfying this motivation" (Anokhin,
1974b, p 23). Since the same motivation (for example, the motivation of
hunger) can be satisfied by means of reaching different, even more eon-
crete,goals in the form, for example, of a specific type of food, further
reduction of potentially attainable goals and potentially feasible
behavioral acts occur under the influence of the situation, which permits
only the behavioral acts whose goals are attainable only in thi.s situation.
This state preceding the triggering stimulus was named preliminary
["pretriggering"] integration. These conceptions are illustrated in Figure l.
S i t u a t i o n
Possible acts~at ~ giden mbment
3 .
~ -
1 U! rd
Fulaire acts ~ o
a~~
s~o ~
oti uxi
atznn a~ v.~ m
- W U i~
, ~ �rl i.~" 4-I U
aa~~~c
Key: S) survival ' S '
Figure 1. Correlation between motivation and situation in pre- ~
liminary integration. The circles refer to behavioral i
~ acts constituting the organism's life experience. The
links between them reflect their position in the hierarchy of goals. �
Arrows show the direction of influences of motivation and situation
determining the priority (.circled numbers) of behavioral acts in a ~
state of preliminary integration. The circles without numbers refer
to "reduced degrees of freedom." ;
In highly organized animals, attainment of the main goal of survival is '
mediated by many hierarchically organized intermediate goals. Separate, ;
interrelated events serve as these goals, and the successive occurrence
thereof can lead the animal to satisfying its movitvation. These events
form the "tree of goals" of a specific motivation in the entire logic
net [system] of life experience. In different situations, the same goal
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_ can be reached by different actions; at the same time, under different con-
ditions, the same action is used to reach different goals. The final choice
of one goal and one behavioral act out of the many possible ones in a given
situation is made at the time when an event occurs in the environment that
favors orie of the goals already chosen by motivation and the ~ituati~n.
This event is called the triggering stimulus.
Only events included in tl-~e hierarchy of goals are actively "pursued" or
"expected" by the organ~sm, can guide the animal in choosing one concrete
goal out of all those that could be attained with the motivation at hand
- in a given situation, and lead to survival. In actuality, trigger stimuli
appear only as a result of prior behavioral acts in the continuum of be-
havior. Any unexpected events immediately interrupt goal-directed
behavior and induce an orienting-exploring reaction. The process of
selection of one goal and one action otit of all the material in memory,
= under the influence of alI ~lements of afferent synthesis, is referred to
as "decision making." Separation of afferent synthesis and decision
making only means that there is isolation of determinants, on the one hand,
and output functions of a single process, on the other, which translates
the orderliness of the environment into an orderliness of elementary
physiological processes in the functional system of behavior.
One goal, selected in the process of afferent synthesis and decision making,
is referred to as the "acceptor of action results." The model of this
goal,.which exists as a certain organization of elements extracted from
memory, in turn determines the organization of actuating mechanisms of
the behavioral act, i.e., organization of physical influences.of the organism
- on the environment. This organizatior of actuating mechanisms is referred
to by the term "program of action," while the organized influences on the
- environment are referred to by the term "action."
Action is a means of altering the correlation between the organism and the
environment, a means of "translating" the expected event--"goal" into a
real event--"result"; for this reason, action is totally determined by
= the model of a future event, rather than the txigger stimulus that directly
precedes the behavioral act. Determination of action by the goal, i.e.,
"anticipatory reflection of reality" (P. K. Anokhin) matees 3t possible
to elimin~te the so-called t~me-related paradox, which arises in reflex
interpr.etation of the behavioral act, in which the orientation of action
toward reaching future events does indeed appear paradoxical, since the
stimulus that directly preceded action is believed to be the cause of
action.
At present, conceptions of purposefulness [goal orientation] of behavior
' are becoming widely recognized. Eve~ such a strong proponent of reflex
. theory as E. A. Asratyan devoted part of his paper at the 21st Inter-
national Psychological Congress to a discussion of "neurophysiological
mechanisms of goal-directed nature of motivational motor acts" (1976, p 18).
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One car~ find a discussion of some conceptions of goal orientation (E. S.
Russell, W. Thorpe, G. A~ Deutsch and others) in a special chapter of -
the book by R. Hinde (1975), in which objections to these conceptions
- are also cited.
R. Hinde believes that "behavioral activity directed toward a certain goal
will attenuate as the goal situation is approached," and he sees an ob-
jection to this thes is in the fact that "rats run fastest when coming
close to the goal" (p 669). In actuality, these view is wrong in its
first premise. Sinc e"approaching the goal situation" is possible only
as the systematic attainment of more important goals in the hierarchy,
any "attenuation" of behavieral activity half way to the ultimate goal
is not justified ~.n any way.
Another objection is due to the fact that R. Hinde relates the goal
orientation of behavior to its determination by an "error signal," i.e., -
discrepancy between the real and "goal" situations, and he concludes:
- there :is no conclusive evidence that, for example, the reaction of
a wasp to damage to its nest is a reaction to the difference between
the damaged and intact nest, rather than a reaction to the edges of the
hole" (p 669-670). However, according to functional system theory,
a discrepancy between a real and goal situation by r~o means serves as
the cause of goal-directed behavior. A discrepancy between the goal and
- a real event can only induce a general orie nting-exploring reaction. But ~
goal-directed behavior is determined by the goals themselves, i.e., models
of future situations extracted from memory; these models precede actions -
- and determine them.
= A concre~e model is compared to the result only after the action, i.e., '
a comparison is made of the informationally equivalent real situation
- to the model of this same (and not future) situation. For this reason, ;
~ from the standpoint of functional sy~tem theory, in the example discussed _
by R. Hinde, the goal included in the hierarchy and ultimately leading to
; survival is the intact nest, and behavior is directed expressly toward
this goal, rather than the "difference between damaged and intact nest."
With such interpretation of the mechanisms of goal-directed behavior
there is in general none of the contradiction mentioned by R. Hinde, -
_ since elimination of the hole serves ae the more concrete goal of
an intermediate behavioral act in the sequence of acts of nest-building
behavior.
= Thus, the general conclusion nf R. Hinde, that the goal approach "should
_ be limited to cases, in which behavior includes reactions to inconsistency
between the existina and goal situations" (p 675), cannot be deemed
- warranted. Inconsistency between the existing and goal situations is
_ demonstrable only as an inadequate result, and it elicits "orienting"
behavior which disrup ts the ongoing goal-directed behavior. The goal '
approach is mandatory for all cases of behavior, since there simply are
- no unpurposef~al acts in the animal's prior experience. ~
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The other objections are probably based on pure mi5understandings. For
example, R. Hinde Frrites: "It is important to stress here that hehavior
may be goal-directed on one level of integration and not on another. At -
the early stages of nest-building behavior, the typical movements of the
weaver are directed and coordinated so as to obtain a finished "stitch,"
but these stitches are not directed so as to lead to building the nest.
The lowest levels are probably goal-directed, but behavior is not directed
toward completing building of the nest" (pp 672-673). R. Hinde does not
e~lain which mechanisms are involved in appearance of individual goals--
"stitches," and how precisely a nest and not something else is created in
integral nest-building behavior.
From the standpoint of functional system theory, any concrete goal may be
formed exclusively as concretization of a more general goal and ultimately
the main goal of survival, which is an inalienable property of everything
living.
We should also mention the instances of obviously useless and'"senseless"
animal behavior under inadequate environmental conditions as a frequently
raised objection to goal-directed behavior. Such behavior is inherent
even in mammals, For example, a fox or sled dog "buries" uneaten food by
scratching a wooden f~oor with its paws and, of course, does not produce
a resul~. (V. Fishel', 1973). In our opinion, these findings a~e in
contradiction to the purposefulness and success of behavior, rather than -
goal orientation, and they are apparently attributed to the fact that
both the goals and actions to reach ~hem can be retrieved only from memory,
while the store of inemory cannot be adequately used in an inadequate situ-
ation. If, however, training is included in such situations, which
broadens the store of prior experience, behavior may change radically~and
become quite purposeful. . ~
As a rule, examples of unpurposeful animal behavior are cited to illustrate
its difference from human behavior. However, all elements of human be-
havior are also extracted from the store of inemory, and man's behavior in
psychological experiments demonstrating the conservatism of thinking -
have much in common with the behavior of a fox that is placed in a cage.
The experiments of Lachins, for example, offer a striking demonstration
of this (see Liper, 1963, pp 301-302). _
Our description of the thesis in function~l system theory of the
- goal-orientation of behavior is only a scheme, and does not presume to
have offered a11 of the arguments in favor of goal-directed behavior, let
alone complete discussion of the mechanisms of goal-directed behavior. It
was necessary only to show that recognition of purposefulness of behavior
is not in contradiction to the principle of causation in explaining
- behavior, but develops it. Indeed, func:tional system theory considers
the immediate cause of action to be ? boal,� the:~taformation model of a
future event retrieved from memory. Alhhough the goal serves as a model
of the future, it already exists in the form of specific ne~YVOUS act~ivity
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even prior to action and, consequently, has the main property of a cause: �
it precedes its effect, i.e., action. It also has another property of
- a cause: it consistently induces its effects-�-actions, which were -
related in the past with reaching it.
At the same time, functional system theory discloses the causes and
mechanisms of goal formation proper. The goal emerges as a consistent
consequence of processes of choice and formation fr~m all elements in
memory of the model of only one event required for survival, and this
choice is made under the influence of both endogenous (motivation) and
exogenous (situation) factors. The trigger stimulus is considered to
be the result of prior behavior and one of the exogenous factors of
choosing the goal. For expressly this reason, the link between a stimulus
and action that follows it is stochastic, and the occurrence of the
same action following the same stimulus is merely a special case. _
Thus, functional system theory removes the seemingly unresolved contra- !
diction between the principles of causation and goal orientation in
explaining behavior.
Appearance of the result, i.e., a new event in the environment, leads to
a situation of conformity between the result of action and "acceptor of
result of action," thereby indicating the end of one cycle of exchange
in orderliness between the organism and the environment, and the start
of the next one.
Isolation of the Behavioral Act in the Continuum of Behavior
In order to investigate the neurophysiological mechanisms of systemic
processes in the behavioral act, it is absolutely mandatory to be able
to isolate the behavioral act in the continuum of behavior. _
In the reflex interpretation of behavior, it is assumed that any behavior
is made up of different reflexes. The unit of behavior is a single reaction
to a single stimulus. R. Chauvin calls this the "atomistic" approach and
_ observes that reactions "are never isolated; separation thereof leads to
impossibility of any interpretation of either these reactions or behavior
, as a whole. For ethologists, the concept of reflex in the na.rrow sense
is, we are not afraid to state, senseless" (1972, p 11). Successive
behavior in the reflex interpretation is viewed as a"chain reflex," i.e.,
a successive series of reactions to corresponding stimuli, which appeared
as a result of prior actions. P. Milner observes that "in many reactions,
the same movement is performed several times, but in each instance it is
followed by different movement; then the problem arises as to how two
different movements can be induced by the conditioned reflex mechanism,
by the same feedback signal. Of course, we can follow seve~al routes to
bypass this difficulty, but the simple basic theory does not ;~old up under
the burden of the required additions and changes" (1973, p 121).
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Because of the methodological convenience of delivering stimuli and
recording reactions, the usual artificial sequence of events in physiolo-
gical experiments (stimulation, delay, change in observed parameter) was
taken as the natural course of events. Orze can indeed detect such a
sequence in preparations (anesthetized animals, with severing of the brain -
and spinal cord, or with the use of curare), i.e., in cases where we are
- dealing with "machine-like" factors and there is no adaptive behavior. -
In a waking anim-al, the concepts of "stimulus" and "reaction" do not enable
us to unequivocally isolate a behavioral act. The presence of behavioral
acts without obvious exogenous stimuli ("false starts," "motivational
reflexes," "intersignal reactions," "reactions to time," etc.) indicates
that the processes determining action appear long before the stimulus, and
- this complicates signif~cantly deterrnination of the actual moment that
the processes referable to a given behavioral act begin. As for the
"reaction," this is not so much a reference to some definite, qualitative
reality of behavior, as it is a synonym for the philosophic al category of _
"effect," which is meaningless apart from its relation to the stimulus.
In some studies of behavior, electromyographic activity or some autonomic
parameters or other are taken as the rea~tion, in others movement of
some part of the body, in others yet, an event such as making contact -
by means of a telegraph key, etc.
_ Thus, the concepts of "stimulus" and "reaction" are philosophical, and _
they may include any changes in the environment (stimuli) and any changes
in the organism that follow the stimulus (reactions). This diffuseness
of the concepts of stimulus and reaction does not permit isolation of
expressly one behavioral act. Indeed, to describe elements of behavior
one must resort to such terms as, for example, "swallowing reaction,"
"running reaction," etc. On the one hand, they reflect the methodological
principle of inechanistic determinism, according to which the behavioral
act is a"reaction"; on the other hand, isolation of the behavioral act -
is accomplished with actual disregard of this principle.
Indeed, "the run" characterizes a segment of behavior only from its
phenomenological aspect, regardless of whether some stimulus is present.
Introduction of a stimulus still does not lead to unequivocal isolation of
a"stimulus-reaction" pair, since one ean consider the turn of the
animal`s head in the direction of the feeder and running toward it, as
well as taking feed and salivation, change in respiration and change in
cardiac activity, etc., as a reaction to a conditioned alimentary signal,
for example. At the same time, the run may be interpreted as the reaction =
to turning the head and taking feed as a reaction to the run, etc. It is -
_ only the proposed "neural link" between specific anatomical structures
(for exanple, between the "eye and salivary gland'~) that ~*ould permit ~
isolation of some reflex, but then a vicious logic circle is formed: for
we cannot study the mechanisms of a phenomenon isolated solely on the
basis of a hypothetical mechanism.
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~ '
_ Thus, there a~e ob~ections to the reflex approach, which separates behavior
- into reflexes induced by separate stimuli, not only when compared to the
continuum of behavior; ~,sol.at~,on of a behavioral act to study the neuro-
physiological mechanisms of behavior cannot Iae performed unequivocally
- when it is considered as a reaction to some stimulus or other, since '
neither the stimulus nor the reaction can be unequivocally defined in the
continuum of behavior.
J F'rom the standpoint of functional system theory, an individual behavioral
act is directed toward achieving a certain result, and it~ can be isolated
expressly by its result, i.e., by the event that it.causes in the environ-
- ment and to achieve which it is performed, The result has very specific
properties and meaning in the functional system, which are isolated
according to different criteria (Anokhin, 1968; Serzhanfiov, 1974) . Here,
we shall discuss only those of . its features, according to which it can
- be defined by external oliservation of behavior. ~ i
;
Since we are dealing witn behavior which, in the broadest sense, can be
defined as the "balance between the organism and environment" (I. P.
Pavlov) , the first property of the result of expressly a behavioral act '
is that the result is .a specific correlation between the organism and
environment, i.e., event-. Any behavioral act elicits numerous changes
in the environment, which may be indifferent and occasionally even
~ harmful to the organism. In accordance with the terminology of functional
system theory, let us call the result of the behavioral act expressly
- events, i.e:, organized aggregates of environmental elements that can be
- used in behavior. Let us call all incidental changes in the environment
the effects; ~e shall not discuss them in this work. The fact that
~ "events" characterize expressly the correlations between the enva.ronment '
and organism, and that any concrete special goal is. included in the -
_ hierarchy of goals and, consequently, in the structure of inemory or lif e
experience of the organism, automatically render the concept of "result"
_ applicable only to "familiary'' organized sets of environmental elements,
All so-called meaningful stimuli may be a result: inborn "releasers" or
acquired conditioned signals. ~
Since, according to functional system theory, behavior is goal directed -
and all actions are determined by th,~ goal, which is "translated01 by
acti~n into a result, the second decisive property of a r.esult is also
the fact that a result is an event that stops actior:s directed toward
reaching it. '
Since goals are hierarchically organized in the structure of expe~ience and
any exogenous factors become goala only if they bring the organism closer
to reaching the goal of survival, i;T the continuum of behavior, achieve-
ment of any result enables the animal to move toward achievement of the
next goal. For this reason, the third essential property of a result is
that it initiates the next behavioral act, which is determiaed by the next
goal in the hierarchy of goals leading to satisfaction~ oi a concre~te - -
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motivation and achi vement of the goal to survive. For expressly this
reasvn, a stimulus, i.e., result of prior behavior, is only a trigger
of subsequent behavior, whereas the specifics of the latter -�-a determined
by the model of a future event, i.e., goal.
These three properties of a result enable us to define, quite unequi- �
vocally, some of the results of behavior and, consequently, to single out
individual acts in the continuum of behavior. In our opinion, this idea
was very well expressed by V. F. Serzhantov: "A performed functional act
- that ends with a specific result causes the organ3.sm to move to other
similar acts. Thus, each separate act is qualitatively circumscribed in
- time, being separated from both preceding and subsequent phenomena of
vital functions. As life moves from one result to another, there is
distinctive expression of its rhythms on the level of the organism"
, (1974, p 73).
y Thus, according to functional system theory, the behavioral act can be .
isolated as a segment of the behavioral continuum from one result to
another.
These acts, which take place successively in time, do not form a"chain,"
but an hierarchy, since the goals are hierarchically organized in .
accordance with the general goal of "survival" and any result turns out
to be made up of more concrete results, and itself is part of a more
general result. In the~above~example of behavior, occurring after use
of a conditioned alimentary signal, the portion of food serves as a.
rather major result, to the achievement of which all of the behavior
discussed is directed. It is achieved, in turn, through the successive
achievement of more special results, which refer both to the change
in situation when the head is turned and change in po~ition of the animal
- in space as it runs.
At the same time, the portion of food serves only as a special result of
behavior directed toward satisfying the hunger motivation. Description
of the real hierarchy of goals would require knowledge about the entire _
subjective life experience of the animal; however, as.an example, we can
confine ourselves to listing only some of the obvious events included in
the hierarchy of goals and results: "to be satiated;'--"to eat a portion
of foot"--"to be near the feeder"--"to see the feeder"--"to receive the
conditioned signal." The goal "to eat a portion of food" contains all
of the preceding goals and is itself contained as an element of the
system with the more general goal "to be satiated."
~ According to this arbitrary hypotheti~al hierarchy, the result (for
examp~e, "to be near the feeder") is reached by all prior behavior,
including turning the head, which leads to a special result, �'to see
the feeder."
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Thus, by introducing the conception of hierarchic organization o~ goals -
in the stxucture of the qrgani:sm~s experience, ~unctional system theory
makes it possible to isolate a behavioral act no matter how minute,
in accordance with the result it achi.eyes, without removing it from the
continuum of behavior and without breaking behavior down into separate
"atoms." ~
At the same time, functional system theory permits retention of all the
methodologi.cal conveniences of delivering stimuli and recording reac~ions.
Indeed, in the case of integral behavior, a stimulus is, from the ~
_ standpoint of functional system theory, the result of prior behavior, ~ i-
sin~e it even has all of its external features: it is familiar, it ~
stops activity preceding it and causes the action to start that is
determined by the next goal in the str'ucture of experience on the road
- toward satisfaction of motivation. �
It is not the next, but prior behavior, which led to appearance of a ,
stimulus, that is informationally related to this stimulus, which exists
be�orehand as a goal, in the form of a certain "anticipatory reflection" '
(Anokhin, 1962). All those who work with animals by the conditioned
reflex method know that dogs literally require a conditioned signal facing
the experimenter and barking. Training alters this behavior, since
"stimulus-result" are attainable by means of "passive anticipation," '
which the experimenter specially develops by reinforcement with a ;
conditioned "calm background" signal.
As we have already stated, a stimulus serves only as one of the guide-
lines of future behavior, which permits the selection of one goal ~ ~
out of the many possible ones according to motivation and situation, by
means of a model thereof in the etructure of experience. Since this ~
goal can be reached by different means, depending on other conditions,
for exdmple, initial position, it is understandable that different
actions may follow the same "stimulus."
_ Since, by virtue of the complexity of organization of experience, the
same goal can be selected under various exogenous conditions, it is
understandable that action may be performed in the absence of a given
stimulus, whereas other conditions make it possible to reach this goal
(evaluation of this possibility by the animal may also be wrong). The~ -
distinct appearance of the same action following the same stimulus is a ;
special. case of goal-directed behavior, when a constant goal in a !
specific situation can be reached by the same ~eans and only in the' '
presence of the same prior "result--stimulus."
This situation is the most convenient for the study of neurophysiological
mec anisms of systemic processes, since in the case of a constant
, stimulus--result of prior action and stimulus--result of next action it
is easy to isolate the interval between the two results, in which all.of ~
activity is directed toward reaching only one goa1, i.e., we can isolate '
a single behavioral act.
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Organization of Physiological Functions in the Behavioral Act
Functional systein theory also alters conceptians of organization _f differ-
ent physiological functions in behavior, in accordance with conceptions of ~
determination of behavior by the goal.
In physiology, the concept of "function" was related to a specific sCructure
for a long time. A reflection of this approaca is seen in such concepts as
"spinal functions," "cortical functions," "function5 of the liver" or
"salivary gland." At the present time, the limitation of such an approach
for analysis of integral activity of the organism is obvious (Anokhin, 1940;
Luriya, 1962; Menitskiy, 1975, and others).
- Conceptions of reflex mechanisms of behavior were closely linked with con-
ceptions of reflex mechanisms of different physiological functions, down to
- the functions of a single neuran. Synthesis of "little" reflexes could
not yield anything but a"big" reflex: some receptor nerve unit is,
hit by some agent of the outside world or inner world of the organism.
This hit is transformed into a neural process, into the phenomenon of
neural excitation. Excitation trsv els over nerve fibers, as if they were
wires, to the central nervous system and from there, by virtue of estab- ~
lished links, over other wires to the functional organ, in turn changing
into a specific process in the cells of this organ. Thus, any agent _
consistently is related to some activity of the organism, like cause and
effect" (Pavlov, 1949, p 553).
Although this conception of the reflex has occasionally been labeled
as oversimplified or even "caricatured," it has not become enriched by
any basic changes in the last 70 years. As validly observed by D. N.
Menitskiy, "in spite of the enormous advances of natural sciences and
modern technology, as well as psychology and neurophysiology, the basic
tenets of conditioned reflex theory remained without appreciable change
until recent years.... The categorial structure, i.e., set of problems,
principles and concepts of the classical direction of physiology of
higher nervous activiLy remained the same" (1975, p.71).
Nor could these conceptions change, remaining reflex-oriented, since
the above quotation of I. P. Pavlov serves as an excellent definition of
the physiological concept of "reflex," reflecting real physiological
processes in spinal preparations and anesthetized animals. We believe
that authors who object to this definition of reflex do not actually
uphold successively reflex positions to interpret integral behavior,
and they put some other content into the distinct physiological con- -
cept of "reflex."
The conceptions of reflex mechanisms of physiological functions were
based on factual data, which continue to be submitted to this day.
They appear absolutely reliable with the use of modern investigative
methods. For example, the ares of spinal reflexes can presently be
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descrihed with exhaustive accuracy and thoroughness (Eccles, 1959; Kostyuk,
1971). However, all these data were obtained esclusively on preparations
(spinal, pretrigeminal, anesthesia, muscle relaxants, etc.), i.e., ex-
pressly in states that preclude goal-directed behavior.
This circumstance, i.e., demonstration of reflexes in the absence of
inte~ral behavior, was noted by I. P. Pavlov as far back as 1904, at the
very inception of conditioned reflex theory: amazingly, after
transection of all sensory nerves of the tongue, most substances that
reach the mouth when eating or forced in lead to absolutely the same
salivation as before they were severed. One has to resort to. more
- radical measures, to give a toxic ager~t to the animal, remove the ~
- higher branches of the central nervous system, to become conv~.nced of
the fact that there is not only a mental, but purely physiological link ,
between substances that stimulate the mouth and the salivary glands"
(1949, p 348). This link is also demonstrable in clinically important
reflexes.
In states that preclude goal-directed behavior, the.effects of stimulation
do indeed appear "automatically" ["machine-like"], since they are caused
by stable and in essence "dead" morphology, although it is purposeful,
_ which the experimenter actuates with stimulation. Under such conditions, -
stimulation does indeed serve as the cause of all processes occurring in
the preparation. The assumption that the animal uses certain morphological ~
elements in behavior just as they are used by the experiment in a pre-
paration was accepted without proof, since there simply was no methodo-
- logical possibility, for a long time, to examine the activity of the
nervous system in behavior.
In the case of integral behavior, in the presence of "spontaneous" nervous
system activity, even the primary nature of afferent processes in relation
to efferent ones is found to be related to ~.nterpretation of the behavioral
act as a reaction to a stimulus. The constant flows of impulses in both
directions make it possible to consider either direction as the first -
(Bernshteyn, 1966) or render such a choice generally impossible, since one ~
cannot single out the moment when there would be only afferent or only
efferent activity. The fact that behavior is a continuum of constant cyclic _
correlations between the organism and the environment relegates the ques-
tion of whicti is first, afferentation or efferentation, to problems of
the "egg and chicken" type. !
The conception of action as efferent activity and specific processes in ;
"functional organ" cells appears to be just as unjustified. As repeatedly ;
stressed by P. K. Anokhin, ",the conception that any exogenous stimulus
can produce a'reflex on a muscle,' 'reflex on a gland' or 'reflex on '
the heart' is more an expression of the technique used to evaluate
reactions than of our knowledge. about the mechanisms of reactions" (1975,
p 148). Even as a phenomenon, the behavioral act exists when and only
when there is organization of various processes into a single whole. P. K.
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Anokhin observed that only the deepest b~as could enable one to see re-
flexes iii the behavioral act. "Consider a kitten," he wrote," that
- performs rhythmic scratching movements to remove some irritant from the
. ear. This is not only a commonplace 'scratching reflex,' th~.s is
consolidation, in the true meaning of the word, of all parts of the
system for a result. Indeed, it is not only the paw that extends toward
the head, in this case, i.e., to the point of irritation, but the head
extends toward the paw. The cervical muscles on the irritated side ar�e
selectively tense, as a result of which the entire head is bent toward
the paw. The trunk is also curved in such a manner as to make free
manipulations with the paw easier. And even the three paws that are not
directly involved in scratching are so placed as to assure the success
- of scratching, from the standpoint of position of the body and center of
gravity. As we see, the entire body is turned toward the focus of the
result; consequen~ly, not a single muscle of the body remains uninvolved
in reaching a useful result. We are dealing, in the true sense of the
word, with a system of relations that is entirely subordinated to the
achievement of a result that is useful to the organism at a given time"
(1975, F 3..5).
This integration of activities of anatomically different structures and -
subordination of any physiological process contained in the behavioral � ~
act to the~general result rules out the possibility of performing any
- physiological function included in behavior as an independent "reaction"
to some separate factor, and this can be observed on preparations. It
is only organization as a whole that determines the form of activity of
- each structure, and "the components referable to some anatomical system
or other are mobilized and involved in the system only to the extent
that they aid in obtaining the programmed result "(Anokhin, 1973a, p 35).
Functional system theory makes it possible to extend the concept of
purpose�ulness to all levels of organization of physiological functions,
which leads to a revision of the content of the concept of function
itself. According to functional system theory, goal-directed behavior of
- the entire organism is organized from also goal-directed activities of
its elements, and the result of the entire int~gral behavior is achieved
by reaching the more elementary special results. Consequently, it is
possible to make any division of activity of the integral organism into
parts, i.e., into separate functions, only in accordance with the hierarchy
of the results. Achievement of some result in the organism is a function,
i.e., part of the general [overall] work, while the organized aggregate of
activities leading to attainment of this result is a f.unctional system.
"We interpret functional system as a combination of processes and mechan-
isms which, being formed dynamically in accordance with a given situation,
necessarily leads to an ultimate adaptive effect that is beneficial
_ to the organism in this very situation" (Anokhin, 1962b, p 77).
From this systemic thesis, not only any function is multistructural, but
any structure is multifunctional, since it is not one function, but all
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those that could take place w~,th the use o~ this structure that
are fixed in the structuxal distinctions. For example, such a result as
moistening food in the mouth is achieved by an entire functional system,
including the activity of many neural, muscular, vascular, glandular and
other morphological elements.
At the same time, the same process of salivation and activity of the same
structure, the salivary gland (for example, in the dog), can be used to
achieve different results: not only to moisten food and submit it to
primary enzymatic treatment, but for heat regulation, 1 icking a wound, etc. -
One could use the term "function of structure" to designate all these func-
tions, since t11e entire set of functions, in which the salivary gland can
be used in general, with only part of these possibilities being used in each
individual functional system, is fixed in the structural distinctions of '
the salivary gland.
Thus, according to functional system theory, all functions contained in the
functional system of the integral behavioral act are, in turn, organized
as functional systems of a lower order of complexity.
- Functional systems on different hierarchic levels were analyzed in detail
in the school of P. K. Anokhin. For example, many studies dealt wiCh func-
tional systems of regulation of respiration (Golubeva, 1971; Polyantsev,
1969; Yumatov, 1976), position (Shumilina, 1949; Agayan, 1970), arterial
pressure (Anokhin, 1947; Shumilina, 1961), autbnomic el ements of behavior ~
(Shidlovskiy, 1969), integral food-obtaining behavior (Sudakov, 1971; -
- Shuleykina, 1971; Khayutin and Umitriyeva, 1976) and many others, as can -
be seen, if only from the bibliography compilec~ by D. G. Shevchenko (1972).
Functional systems on the lowest level of complexity are functional elements
of more complex functional systems. The behavioral act is performed as the
immense hierarchy of functional systems on different levels of complexity:
"Of course, the correlation between actin and actomyosin constitutes a ~
well-circumscribed functional system, with regard to it s operational archi- ;
tectonics, which ends with a positive result that can be formulated as
the contraction of a muscular fibril. But such a func tional system is ,
merely an intermediate system between even finer molecular correlations ;
- of muscle cell protoplasm and between movement, for example, of a hunter
in the forest in search of game, since this movement is also ultimately ~
performed by means of actin and actomyosin. How wide the range is,
which contains numerous functional systems making up this immense hierarchy
of systems!" (Anokhin, 1973a, p 37).
An enromous number of various means of organizing elements is possible in
this hierarchy. However, it does not contain all possible combinations,
and is limited only to inborn and acquired integrations, since the very
- formation of some organization or other in philogenesis or through learning
is possible only under the system-farming influence of the result, and it is ;
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already the "purposefulness" of the morphological structure o� the organism
that reflects the restrict~on o~ "degrees of freedom" of different combina-
tions. Still, even the variants of organization,�which had, in principle,
at some time led to an adaptive result on some level of cotnplexity or
other and therefore were fixed in the inborn or acquired experience of
the organism, are present in sufficient number to make the necessary
- selection and organization of elements on each hierarchic level to rzach
an individual result.
Thus, from the standpoint of functional system theory, performance of any
function is related to organization. of specific activities, rather than
activity or a substrate per se. This link was determined already in the
course of inception of life. Since the main goal of biological systems,
to survive, is actually the goal of preserving integrity and organiza-
tion of inetabolic processes, the entire hierarchy of goals of highly
. organized animals is a hierarchy of organization of physiological processes
ultimately leading to preservation of integrity and organization of ineta- -
bolism within the enttre organism. And only those of the more elementary
functional systems, the results of which form the result of a larger system, _
are involved in some large functional system. Tlius,,the inter~e~at~,ons
of elements~in the system.are subordinated to the result of the entire
system. "The term 'system' can be used only for a set of selectively -
involved elements, in which interaction and interrelations acquire the
nature of interaction of elements to achieve a focused useful result"
- (Anokhin, 1975, p 37). ~
Consequently, the neurophysiological study of systemic processes in the
behavioral act is the study of processes of organization in behavior of the
activities of separate brain structures and separate neurons.
Operational Architectonics of the Funczional System in an Elementary
Behavioral Act
The orderliness of the environment, both present and past, wh3.ch makes up
the memory of an organism, is used to put in order the relations between
elements in the functional system of a single behavioral act. The correla-
tion between this order of environmental elements and processes of organiza- -
tion of elements of the organism is implemented through the operational
architectonics of the functional system of a behavioral act. According to -
the theory of P. K. Anokhin, the structure, or operational architectonics,
of a functional system of any degree of complexity is comprised of systemic
mechanisms, or stages, of afferent synthesis and decision ma.king, and
then the acceptor of results or ~oals of action and action program; per-
formance of action; achievement of results and comparison of feedback
from the parameters of the results to the acceptor of action results
(Figure 2).
In an elementary behavioral act, these systemic processes, i.e., processes
of interrelation between current and past information,and organization
t -
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of the system, are directly superimposed o~~er the time structure of the
behavioral act, and they can be precisely determined in time. There are
various systemic processes involved in a behavioral act, singled out as
- a segment of the behavioral continuum from one result to another, at dif-
ferent phases of its development: afferent synthesis and decision making
become involved between the result of the preceding act and start of
_ actuating mechanisms of the next one; the start of the actuating mechan-
isms of a behavioral act already coincides with implementation of the
- program of action and acceptor of results of action, while achievement of
. the result marks the time of occurrence of feedback and comparison
thereof to the acceptor of results of action (Anokhin, 1973b).
All these processes, or stages, of organization of elements into a system
exist in functional systems of all levels of complexity; however, they
present a number of distinctive features in an integral behavioral act,
which are related to the fact that behavior "equilibrates" expressly the
entire organism with the object-related [objective] environment.
Situational en~ ~ r
afferentation
. . ~ ' PR
Trigger` eci-
sion ~
stimulus ~ makin pA
.
' , ctio
Situationa~. Dominant
- afferentation otivatio
Afferen~
synthesis
Key: ARA) acceptor of action result PR) parameters of result
PA) program of action RA) result of action
Figure 2. Operational architectonics of a functional system
after P. K. Anokhin (1973a)
Even the first living systems were open (Anokhin, 1975, p 333) and
included interaction with the environment. The' result, in this sense,
is part of the system brought out into the environment, or part of
the environment contained in the system. Organization of the system can
be maintained only by means of organization of the environment (Ferster,
1964), and the very first living things had to utilize "negentropy"
from the environment (Schroedinger, 1947). For this reason, the results
on the level of biochemical systems were specific chemical substances,
organization of the relations of which was used to maintain metabolism.
On the level of highly organized organisms, an event in the environment
that became a result could also consist of only a.specific organization
of the environment. This organization of the environment, or information
flowing and already fixed in memory,.ultimately determines the selection ,
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and organization of elements and physiolog~.ca1 processes on all levels of
complexity in the functional system of the integral behavioral act.
In order to form an hierarchy, the "operational architectonics" of systemic
. processes must be basically invariant on all hierarchic levels of the systems
(Anokhi.n, 1973a). The functional system of the integral behavioral act 3s
made up of subsystems on the physiological level, each of which undergoes
a stage of afferent spnthesis and decision making, and includes its own
= acceptor of results of action and program of action. Of course, on the
level of physiological subsystems, all these processes take less time than
processes of organization of the entire system of the behavioral act. This
is related both to the lower volume of elements in physiological systems
and partial morphological fixing of some organizations "refined" in phylo-
genesis or ontogenesis.
At the same time, an individual behavioral act is always only one of the sub-
systems on the behavioral Ievel in the functional system whose goal is to
satisfy motivation, in which each systemic process may involve man~ elemen-
tary behavioral acts. Thus, the functional system of an integral (and, at
the same time, elementary) behavioral act must result in an even, i.e.,
correlation with the environment of the entire organism, and consists of
subsystems of only the physi.ological level, the results of which are certain
changes within and without the organism, constituting part of the events,
but not correlating the environment and organism as a whole.
We have already noted that, according to functional s;stem theory, the
choice of one goal and one behavioral act out of the entire store of
memory take place with the involvement of motivation and situation. It
actually signifies a choice of an enormous amount of subsystems on all
hierarchic levels and organization from them of a specific integration,
- or even an entire hierarchy of integrations of physiological processes.
The purposeful coordination of functions of different elements into an
integral system Lakes place by means of eliminating "superfluous" degrees
of freedom from the elements (Anokhin, 1973a, 1974a), related to Che
possibility of using the same element in diFferent systems.
Since exchange of orderliness between the organism and envixonment takes
place constantly, at any g~ven moment motivation and situatian make it
y possible to implement only a small number (probably about seven) of
behavioral acts (~huprikova, 1978). Motivation and situation reduce the
degrees of freedom of all subsystems used in behavior, so that in the
presence of one motivation and in a specific situation only limited
sets of elements can unite into the functional system of the behavioral
act. This preliminary selective organization of elements is what
constitutes "preliminary [pretrigger] integration" (Anokhin, 1968). The
latter concept ~.s referable to the next act, and in the continuum of
behavior preliminary integrations of future behavioral acts are formed
and change during current behavior, which is the expression of one of
the preceding preliminary integrations.
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= The process of translation of preliminary integrat~on into a behavioral
: act, i.e., final el,imination of all supexfluous degrees of freedom of ;
all subsystems of the physiological level and organization thereof into -
a single,purposeful functional system of tt~e integral behavioral act,
= occurs with appearance in the environment of some result of prior beha- ~
vior, on which depends the choi ce of a conerete goal and a r~eans of
reaching it. ,
Since there is no information in the environment as to expressly which -
subsystem organization will lead to satisfaction of motivation, while
memory of the organism consists entirely of such information, exogenous
infermation in the course of aff erent synthesis and decision making is
used expressly for selection from memory of specific information, from
which a concrete goal is set (acceptor of results of action), which is
reached through a single act and adequate motivation and situation.
_ These processes of organization of elements into'a system take up the '
interval between the�result of the preceding behavioral act (stimulus)
and start of purposeful action (reaction).
The acceptor of results of action, which appears after decision making,
can be theoretically related only to the programs of action that had
led to achievement of expressly this result in the past, and thi.,
determines the purposefulness of any action. Since actuating mec:'.ianisms �
of the behavioral act are determined by the acceptor of results ot
action and retrieved from memory, where they are c~ordinated beforehand,
_ the program of action arises immediately after decision making as an ~
"efferent integral" (Anokhin, 1968).
Reverse organization of system elements into a new order of environmental
elements occurs already by means of the systemic process of action, when
the organized work of actuating physiological subsystems is performed and
real results of integral behavior are achieved. Action is now mani-
fested as the coordinated function of selected subsy~tems, and until the
result is achieved coordination occurs only on the subsystem level,
whereas the correlations between the integral organism and organization
= of the environment are predetermined until the next result is achieved.
Upon completion of action and achievement of results, information about
the parameters of the real resul ts is compared to the.information of
the acceptor of results of action and, in the event they coincide, the
organism is able to move to the next purposeful behavioral act on the
road toward satisfying motivatio n; whereas in rlte case of noncoincidence, ~
this induces a universal orienting-exploring behavioral act.
Thus, the integral elementary behavioral act is an elementary cycle of "
_ correlating organization of the integral organism to the objective
enviranment. We can examine this cycle starting, for example, with
action: action leads to a result, i.e., specific organization of elements
of the environment which, along with motivation and situation, is used
to organize elements of the organism in processes of afferent synthesis
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and decision making; the oxganrzation formed after deci~ion znaking conforms
with the acceptor of resclts of action and related program of action.
Decision making is a transitional factor, after which all combina-
tions of stimulai acquire an actuating, efferent natu.re immediately
after making a decision there is formation of the integral of efferent
stimuli, which must first implement a peripheral action and then achieve-
ment of the results of action. There~is a precise, i~e., equivalent,
informational link between all these stages of formatio:~ of the act proper....
If we were to examine the results of action as consequences of organized
centrifugal flows of stimuli, these deterministic relations can be con-
tinued further, in the direction of information about the results obtained"
' (Anokhin, 1968, p 233). -
Now that we have discussed this elementary cycle in terms of systemic pro-
cesses, we can undertake the neurophysiological study of systemic pro-
cesses of the elementary behavioral act. .
Our objective should be to try to disclose the neurophysiological content -
of such systemic processes as afferene synthesis and de~ision making,
acceptor of results of action and program of action. All these concepts
are related to the concepts of "organization" and "information," which
cannot be unequivocally defined at the present time (see, for example,
Abramova, 1976; Kremyanskiy, 1976).
Perhap s, as the relevant neurophysiological data are accumulated, it will
be possible to define these concepts both in philosophical and cybernetic
terms. However, this wi11 require that the systemic principle be the
~ guiding one in neurophysiological studies of behavior. When it is dis-
regarded, a situation is formed in neurophysiology that is quite vividly
described by G. Somyen: when it is a matter of the central nervous
system, we are rich in facts but poor in theory. Data are accumulating
with incredible speed, but they form an amorphous~mass, rather than an
� organized structure. Advancement is inevitably retarded and the route _
becomes confused whenever there is an abundance of facts but not enough =
guiding principles (1975, p 235).
We should like to end this chap*er and, at the same time, give warning
in the words of K. Lashley: "The point of view of nervous activity
described here apparently does not give us the simple and clear explana-
tions that are possible if we~recognize the reflex hypothesis. But this
~ clarity was attained at the price of disto,.ting the truth, and we prefer
to admit our ignorance and be accused of vagueness, instead of shutting
our eyes to the most important problems (1933y p 196).
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~ CHAPTER 2. ELECTROPHYSIOLOGICAL CORRELATES OF SYSTEMIC PROCESSES IN THE
ELEMENTARY BEHAVIORAL ACT
Electrical Activity of the Brain in Behavior
Electrical activity of different brain structures and neurons is the most
accessible and widely used parameter of processes occurring in the central
nervous system in performing behavior. Since the functional system of the
goal-directed behavioral a.ct is formed by the coordinated activity of many
struc tures and neurons, it is equally important to neurophysi.ological
_ studies of systemic processes to determine the time characteristics of
neurophysiological processes in each separate structure or neuron and the
correlation between these processes.
- Of course, the systemic significance of time and space characteristics of
elect rical activity of different brain structures can be disclosed by
comparing them to the time intervals of the behavioral act and systemic
processes occurring in these intervals.
As we have alrea~ly noted, there must be general systemic processes of
afferent synthesis and decision making in the elementary behavioral act,
in the interval between the stimulus and start of action, i.e., processes
of coordination of activities of many elements on the scale of the
~ entire organism; there must also be genezal systemic pr.ocesses of
_ implementation of the acceptor of results and program of action between
- the s tart of action~to achievement of the result, when activity of the ;
organism as a whole is already coordinated and goal-directed, while coordi-
_ nation processes take place only on the level of physiological subsystems
of the integral behavioral act.
_ Systemic processes can be determined only in time, and they cannot be :
local ized in some structure, since systemic processes are processes of
interaction between many constantly functioning afferent and efferer.t
central and peripheral structures that are coordinated in a specific
way to achieve a concrete, adaptive result.
At the same time, it is apparent that local processes in separate struc-
tures, which perform different functions, must be related to expressly '
these specific functions. Electrophysiological phenomena are usually
compared to specific functions, since they are always recorded in some
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concrete local structures. For example, witn derivation from the visual
- cortex, all electrophysiological parameters are compared to the pxoper--
ties of visual stimuli and assessed as the correlates of visua~ informa-
tion processing, whereas with derivation of potentials from motor
structures they are compared to movement and considered as correlates of
specific motor functions.
When analyzing the significance of electrophysiological phenomena from the
positions of functional system theory, the question arises as to how
special and systemic processes are related and to what extent they are
ref lected in electrophysiological phenomena. This question is also
closely lintced with the problem of origin of electrophysiological pheno.-
mena. Do they reflect processes specific to the morphological structure
and relations of a specific structure, or processes of coordination of
activities of elements situated in different structures?
If electrophysiological parameters are correlates of processes related to
specific structure and physiological functions of specific brain structures,
they must be peculiar to each structure. But if electrophysiological
indicators are related to systemic processes and reflect coordination of
activities of elements "referable to different anatomical systems," these
parameters must be similar for different structures, but specific to a
specific behavior.
According to current data, the activity recorded with a macroelectrode _
in some point of rhe brain represents the sum of many processes occurring
in adjacent tissue. Overall electrica]. activity reflects both synaptic
potentials (Jasper, Stefanis, 1965; Frost, Gol, 1966) and dendritic ones _
_ (Purpura, 1963; Klee et al., 1965) and, perhaps, glial ones (Roytbak,
19Es5), as well as circulatory and tissular metabolic processes
(Aladzhalova, 1962); and the active elements do not remain constant,
so that the overall effects owe their origin to different neurons in
different time segments (Elul, 1972).
= The complexity of electrogenesis of total activity, which is also in-
- creased by anisotropism of brain tissue and presence of dipole relations
- in oriented structures, does not enable us to relate the charac~eristics
of overall activity to the activity of some specific structural elements
of brain tissue. However, total electrical activity of specific brain
structures can serve as an indicator of the state of these structures
and dynamics of processes in macrostructures.
It is for this purpose that one usually records overall electrical activity
of different brain structures in relation to behavior. At the present
time, there are very many works dealing with analysis of the EEG of _
activity in behavior.. As noted by V. I. Gusel'nikov, at the first stage
"there w~s a great desire to see, in the dynamics of the overall EEG, a
ref lection of the classical conceptions of the main patterns of brain
function, which led at best to repetition of the general schemes already -
proposed for them by I. P. Pavlov" (1976, p 8).
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However, already visual evaluat~on o~ overall act~,vity according to the _
change in frequency and amplitude o~ oscil,lations tnade it possible to ,
determine that during performance of behavioral acts, in response to
- a conditioned signal, activation is ohserved in many structures o~ the
brain (Gasteau, Roget, 1962), and there is selective involvement in
activation of vzrious structures with various forms of behavior
(Shumilina, 1959, 1961b). These facts already warranted reference to
systemic organization of processes in the brain during performance of _
behavioral acts (Shumilina, 1965; Naumova, 1968).
With the appearance of a possibility to asses s more precisely the frequency
and time characteristics cf overall activity, it was found that the oscilla-
tions of potentials become synchronous in var ious structures during
behavioral acts (Livanov, 1962, 1972). Synchrany is also observed in
selectively related structures, rather than all of �them (Anokhin et al.,
1973), and the set thereof changes with change in form of behavior
(Ioshii et al., 1969). `
_ Special experiments conducted in the laborato ry of M. N. Livanov (1972)
~ revealed that synchronous activation of different structures is closely
linked with behavior. On the one hand, a correlation was established ;
between the probability of moveiuent of a rabbit in response to a flash '
and level of spatial synchronization of the cortical EEG (Luchkova, 1971);
on the other hand, there was a link between spontaneous movements and
Ievel of EEG synchronization (Trush, Korol'kova, 1974). A correlation ~
was also established between human reaction t ime and level of spatial
synchronization of cortical activity (Vasil'yev, Trush, 1975) . '
Thus, studies of overall electrical activity revealed that, in perform- I-
ing behavior, there is,activation of an entir e system of structure, the
_ composition of which depends on the form of behavior; electrical processes
are synchronous in many structures; synchroni zation of processes is ~
referable to a specific set of structures, and it is necessary to
behavior. '
All these data already indicate that, in behavior, cerebral processes ~
= have systemic, rather than linear, organization. However, the EEG
describes processes occu rring in a particular structure in only the
~ most general features; moreover, evaluation of changes in the overall
EEG requires rather long time segments, and changes in the overall EEG
may be referable to Iarge segments of the behavioral continuum.
Separate oscillations of overall electrical activity, which are referred
to as "evoked potentials" (EP), "generated po tentials," "premotor poten-
tials," "motor potentials," as well as "wave of anticipation" or "condi-
= tioned negativeness," in relation to exogenous event.s or movements,
are more suitable for comparison to systemic mechanisms of the elementary ;
behavioral act.
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Like overall activity, EP constitute a complex phenomenon reflecting the
state of many different elements at the point of derivation. The configu-
ration of EP recorded in a given point probably also depends on both
synaptic (Purpura, 1963) and dendritic (Kullanda, 1964) potentials, and
perhaps glial ones as well (Roytb~k, 1965), and the location of the macro-
electrode in relation to orientation of dipoles alsa p lays a role in
polarity and amplitude of components (Guseltnikav, 1976). EP are used in
behavioral experiments and clinical examinations as a parameter of the I
dynamics of rather rapid processes in local points of derivation.
Synchronism and Similarity of Conf iguration of EP o~ Various Structures
in Bt~havior
_ For ri long time, EP were studied on animals ar.esthetized with barbiturates. -
Under such conditions, the oscillations of potential. in response to affer-
en~ stimulation were recorded from relatively local "focuses of maximum
activity," and they were stable in amplitude and conf iguration (Chang, 1959).
_ This circumstance caused wid~ use of the EP phenomenon in studies of the
morphology of relations in the central nervous system and publicati~n of
numerous studies dealing with a search for the pathways and structures
through which particular oscillations are "conducted." Reflex or "commutator"
conceptions of the mechanisms of behavior determined the same approach ta
the study ~of EP in integral behavior as well. However, already the use -
of chloralose (Buser et al., 1959) and muscle relaxants (Buser, Borenstein,
1959) revealed that, in response to the same stimulus, EP can be demon-
strated in many structures of the brain, while responses to different
stimuli can be recorded in the same structure. -
= EP were found to be very generalized (Kogan, 1965; Shul'ga, 1965) and
_ unstable in both localization and configuration in waking animals and man, -
which made it necessary to use the averaging procedure. Destruction of
different brain structures in waking animals did not el iminate EP in -
others (Chow et al., 1966; Cohn, 1969), while the observed EP changes were `
brief and could not be unequivocally explained (Cherkes, Lukhanina, 1972).
Nevertheless, debates continue concerning the links between EP components
and conduc~ion of afferent excitation over some "projection" and "non-
specific" pathways or other.
- Already in the early studies invulving recording of EP in response to
stimuli requiring a behavioral reaction it was found that there is very
early oscillation of potentials in cortical regions that were unrelated
to the stimulated analyzer (Artem'yev, 1956, 1959). A11 researchers
working ~n development of conditioned reflexes observed a phase of gerierali-
zation, when EP were demonstrable in virtually all leads (Shumilina, 1965;
- Naumova, 1968). E. R. John and his coworkers compared, in an entire .
series of experiments, the time parameters and configuration of EP in
many different brain structures, and demons~rated in many of them syn-
chronous EP, similar in configuration, in response to a stimulus that
induced behavior (John, 1969; John, 1972). In these studies, a comparison
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was made of configuration of EP averaged for many delivexies of a stimulus,
which permitted elimination of EP instab:ility i.n any structure, which is
common in waking animals. Analysis of the acfiivity af neuxonal "ensembles," -
- recorded with the same "quasimicroelectrodes" used for EP revealed that
- impulse activity of neurons of many anatomically and functionally differ-
ent structures was similarly organized in time when submitted to statis-
tical evaluation (John, Morgades, 1969). These data served as one of
the bases for "statistical configuration theory" (John, 1973).
The synchronism and similarity of.EP configurations in response to a
stimulus that evokes behavior do not extend to all structures of the ~
brain. In addition to the fact that EP are specific in:configuration and
distribution for different behavioral acts (for example, food-searching
and defense behavior), they are also variable and individual for each
animal (Myshkin et al., 1968). The question of individuality of EP has been
best studied in man. It was demonstrated, for example that, other condi-
- tions being equal, EP of twins have marked similarity (Dustman, Beck, 1465);
some of the individua~. characteristics of EP are beginning to be used in
differential psychophysiology (Rutman, 1974; Rusalov, 1974, 1975). ~
All of these distinctive features of EP do not.make it possible to intro-
duce a nomenclature of EP components that would apply equally to all condi- i
tions (Rutman, 1974) or to outline the typical topography of theix deriva-
tion. Under some conditions, marked EP may be recorded even from~"extra- j-
cranial" structures (Prichard et al., 1965), under others they are depressed i
a.nd not demonstrable at all (Coquery et al., 1972).
i
In our laboratory, we also observed synchronism of EP in different brain '
structures in response to a flash of light, in the case wher~e the flash
- triggered the rabbit's run to the feeder.
Experiments were conducted (with S. S. Trofimov) on five rabbits in a
special chamber (Figure 3). A flash of~light from the flash lamp
of a Soneclat stimulator (0.3 J, 50 us) was delivered from the ceiling of
the chamber (?0 cm above the floor). There was a 1-s interval between
the flash and automatic delivery of a feeder with 10-30 g of cabbage or ~
carrots.
The EEG of the right and left visual, right and left sensorimotor, right ;
auditory cortex, hippocampus, hypothalamus and reticular formation of the I
mesencephalon, was derived monopolarly by means of implanted electr~des. ~
The silent electrode was placed over the frontal sinus. A Polygraph-YVII, -
with concurrent recording on a Magnetor XIV, was used to record the EEG,
as well as EMG of cervical muscles and relevant marks. The bandpass '
constituted 0.3-200 Hz for the EEG channels. The EP were averaged
by reproducing the tape for 25 runs on an NTA-512B analyzer (bandwidth
2 ms, period ["epoch"] of analysis 512 ms or 1024 ms).
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_ . . , t ~ i
~ o 0
1, ~ 6
_...._:...;~s..~
. ;
~ ~
} Y 1
~ , '
- +,GYyN.
~Y4
, I + *
~a
i
ti:..:.
Figure 3. Genera]_ view of experimental chamber
- 1) flash lamp
2) feeder with automatic delivery of 10-20 g cabbage
3) contacts
= Figure 4 illustrates the tracing of total electrical activity of different
regions of the brain in a single behavioral act. It was demonstrated that
- EMG acti~ity marking the start of functioning of actuating mechanisms
appears on the rear front of the negative component of EP. On the left
are averaged evoked potentials (AEP) in different structures of the
brain corresponding to 25 such acts.
Figures 5 and 6 illustrate AEP of different regions of the brain in res-
ponse to a light, which triggers orienting~exploring behavior in one
gituation (a) and purposeful movement toward the feeder in another (b) -
for two different rabbits (in Figure 5, the tracings were obtained for
rabbit No 2 and in Figure 6, for rabbit No 4). These.same figures
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illustrate histograms o~ latency periods o~ II~G actiyation during the
corresponding acts.
GOs
, Cod
Cms '
Cm -
Cacs
r..~..~-....
Hfh
. Fr
,EMG I0~ '~`~w~1~`~''1~'`'~
_ ~
-E- 800 ms -j-
Zigfit Food -
Act -
Figure 4. Rabbit EEG during elementary behavioral act: In response
to the flash of light, the ra}abit heads for the feeder, in
which carrots appear after 800 ms. Top to bottom, leads: left and
right visual cortex, left and right motor9 left aiiditory cortex,
hypothalamus, reticular formation of the mesencephalon, EMG of
cervical muscles, stimulation marks, actogram (shows rabbit nearing i
the feeder)
Analysis of these tracings revealed that when the flash of light tri.ggered -
food-obtaining behavior, AEP are synchronous and similar in configuration
in several structures of the brain; for example, in Figure Sb, there are
very similar AEP in the sensorimotor and auditory cortex, and in Figure 6b, '
this applies to the right visual and sensorimotor cortex, as well as
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' reticular formation and hippocampus. If the similarity is assessed solely
according to time organization of processes without consideration of ampli-
tude of different elements and the initial level of constant potential, we -
can consider AEP to be similar in all leads, with the exception of the
hippocampus, in rabbit No 2(Figure 5b), and in all leads with the exception ~
of the hypothalamus and left visual cortex in rabbtt No 4(Figure 6b). The
_ difference in amplitude of different AEP components could be related to
the location of electrodes in relation to active tissular elements and
- different conditions of derivation of electrical activity for differently
localized electrodes.
a b
~ ~ Figure 5.
C.v.s Averaged evoked potentials in response
c.o.f. to flash of light triggering orienting
behavior (a) and running toward feeder
c.s-m.s (b) in rabbit No 2. -
ts-r'~.
The time of the flash is shown by the
~A~~ arrow. Leads, top to bottom: left and
yth.,.~,V,~,~,.�. right visual cortex, left and right
sensorimotor cortex, right auditory
~~f ' cortex, hypothalamus, reticular formation,
hippocampus. At the bottom of (b):
~p~ histogram of distribution of latency
' periods of EMG of cervical muscles in
25 averaged combinations.
~0o ms
w
In some structures, ALP have opposite pol.arity of all or some components.
This renders the AEP in such structures as the hippocampus in rabbit No2
and hypothalamus in rabbit No 4 dissimilar to AEP of other derivations; but
if we judge only the dynamics of processes and consider that the amplitude
and sign of components in each given structure are related to the location
_ of electrodes in relation to active elements of brain tissue, on the
basis of AEP configuration, we can conclude that ~n these structures also
- the dynam3cs of the processes have similar organization in time.
_ Thus, during a behavioral act, the processes in some.functionally and mor-
phoZogically different brain structures are synchronous and present similar
time organization.
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Getting somewhat ahead of our presPn-
tation, 1et us mention that we
b obtained the same results in experi-
1 a ~ ments with defense behavior, and
/ with recording o� activity of
individual neurons in different
c.0.s. parts of the brain during behavior.
G.O. d.
We believe that the same time organi- '
Cs-m,r zation of processes in functionally
~ cs-m.d and morphologically different struc-
tures, such as, for example, the
C.A.d. visual and sensorimotor cortex, ;
precludes interpretation of AEP as
yth. correlates of coding of some speci-
fic information, in this case !
R�F f"'~"'''ti" visual. Since physiological func-
Y ,~,i~~ tions of different structures
~'P~� ~
n evidently remain different in
~oa ms behavior, it must be accepted that
W common features of organization of ~
i-
activity are created by processes
Figure 6. that are common to many structures, ~
Averaged evoked potentials for rabbi.t and they occur only during behavior,
No 4. Designations are the same as~ but not under anesthesia, when EP
in Figure 5. are recorded in limited points.
The synchronism and similarity of
organization of processes in different structures also rules out the possi- j~
bility of "conduction of oscillations" from one structure to another,
and dascription of processes whose correlates are EP in terms of "con-
duction pathways" is generally inadequate.
If we were to compile a general nomenclature of all EP components demon-. _
strable in at least one lead a~d t~~.ynthesize"~an artificial "common" EP -
from them, indicating only the time interval.s taken up by specific compo-
nents, regardless of their sign and amplitude, we could gain an idea about ~
the time organization of processes in all derivations. A comparison of ;
EP in each specific lead to the "common" EP would show us the form of _
involvement of a specific structure in general~processes. And we learn
that some structures have a complete set of components: primary positive ~
oscillation.followed by great negativ3.ty, then positivity and slow ~
- ne~ative deviation. In other structures, there is not a complete set of
components, but the existing oscillatians are synchronous with some ~
components of the "general" [common] EP. This shows that the EP of
a separate structure, even if dissimilar in general configuration to EP
_ of other derivations, may reflect involvement of this structure in some
phases of the general process.
;
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A comparison of EP in two different behavioral situations, or3enfiing and
food-obtaining behavior (Figure Sa and b; Figure 6a and b), indicates that
although the configuration of EP in~the same structures varies widely,
the "general" EP consists of the same main components, each of which
spreads differently in structures and is complicated by various subcompo-
nents in different behavioral acts. -
The same phenomenon was demonstrated when a comparison was made of EP of
different rabbits with the same behavior (Figures 5 and 6): the distribu-
tion, configuration and even polarity of differen.t components could be
quite different; however, the four-component structure of the "general".
EP is apparently mandatory. We observed this four-component EP structure -
in virtually all of our experiments, and we sha11 adhere to the following ~
nomenclature of these components hereafter: primary, negativlty and late
positivity, which may be followed by a slow negative deviation under
certain conditions.
Link Between EP and Time of Behavioral Act
In the same experiments, we measured the latency periods of electrical ~
activation of cervical muscles, and plotted histograms of distribution of
these latency periods. The earliest EMG reactions appeared with a latency
period of about 50 ms; mean latency time ranged from 100 to 400 ms in .
different rabbits. A comparison of histograms of distribution of these
latency periods to time of development of EP revealed that only primary =
and negative components of EP develop in the latency period of the EMG
= reaction, whereas late positivity corresponded already to the start of
muscular contraction and, consequently, start of function of actuating
mechanisms of the behavioral act (Figures 5 and 6).
Many studies have been devoted to EP changes related to different reaction -
- times (Donchin, Lindsley, 1966; Bostock, Jarvin, 1970, and others). However,
researchers concentrated mainly on the correlation between reaction time
and amplitude of difFerent EP components. The question of correlation
between time characteristics of EP and time of behavioral act was not
posed, probably because of the prevailing view that EP are related to
"afferent processes." ~Still, R. Eason et al. (1967) demonstrated a link
between reaction time and latency of various EP components.
If we cempare the literature, according to which human reaction time to ~
different stimuli constitutes 100-300 ms (Shoshol', 1966), while EP in _
response to the same stimuli constitute 300-400 ms (Rutman, 1974), it is _
' easy to see that EP correspond to all processes in the behavioral act, _
rather than only "analysis of the stimulus."
In recent times, direct evidence has also appeared of the fact that the
start of motor activity coincides in time with the rear front of negati-
vity or anterior front of the positive component (Peymer, 1971; Ikeda,
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1973). A correlation was demonstrated: in all cases of increase in reac- �
tion time, regardless of the cause of this increase, there ~.s increase and
even dovbling of the negative ~omponent (k'eymer, 1971; Tkeda, 1973; Ritter
et al., 1972).
The ~act that EP conform with processes of the entir.e behavioral act makes
it un~ustified to classify potentials as sensory and motor, since they only
differ in means of isolation from the general, overall EEG. Indeed, if we
average the electrical activity of some structure due to a stimulus, we
obtain a sensory EP, whereas if we "reverse average" the same activity from
the start of the EMG, we obtain a m,otor potential. In view of the variabi-
- lity of the latency period of the EMG reaction, the configuration of motor
- potentials may differ somewhat from the configuration of the evoked
potential; however, the general composition of components rema.ins the same. ~
Tracings of the motor potential related to voluntary movement of the
foot, submitted, for example, in the work of L. Gilden et al. t1966), .
conform entirely with the late components observed in the behavioral act
and triggered by some stimulus: first a small positive component, then
negativ ity,' which reaches a maximum about 100 ms after its start,
followed by strong positivity. EMG activity begins together with the
posterior front of negativ~ity.
L. Decke et al. (1969) described a very similar sequence of components,
associated with finger movement: against the background of "potential
readiness" 86 ms before the EMG or. 117 ms before deviations on the
mechanogram, "premotor positiv~ity" was recorded, which changed 56 ms .
before the start of EMG activity into a"negative motor potential," the
~ posterior front of which corresponded to the start of EMG activity. The
' maximum level of this activity coincided with the next positive companent.
The same sequence of processes has been described with other forms of ~
motion and eye movement (Becker et al., 197~).
_ All these facts convince us that in all cases the summa~_�ed potentials
_ associated with a behavioral act correspond to all processes of organiza-
tion of this act.
- Neverthelecs, EP rec~rded in response to a sensory stimulus are usually
analyzed as correlates of only sensory processes, whereas the potentials
- isolated by "reverse averaging" are analyzed as correlates of only motor ,
pr~cesses. If we agree that both aualysis of the environment and
~rganization of actuating mechanisms are required to perform a behav~oral
act in any case and, as we have tried to demonstrate in the preceding.
section, that processes in sensory and motor structures have the same
_ time organization, it becomes apparent that EP reflect very unique and
qualj.tatively specific processes that occur during performance of
integral behavior. The use of anatomical and physiological categories
of "afferent--efferent," or "sensory--motor," is not adequate for evalua-
tion of processes, the correlates of which are EP.
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~'Endcgeny" of EP in Behavior
Evoked potentials were used in an enormous amount of studies of the most
diverse problems.
These studies made it possible to accumulate some very important facts
dealing with the dependence of diverse EP characteristics, such as configu-
ration, latency periods, amplitude and polarity of components, on the most
varied experimental conditions. Particularly many works dealt with deter-
mination of the dependence of EP on intensity of stimulation. Some authors
found a link between intensity of stimulus and primary EP component
(Schmidt, 1968); however, most studies demonstrated that laCer components
depend Dn intensity of stimulation (Beck, Rosner, 1968; Wicke et al., 1964).
EP were also found to be related to the content of stimuli, such as
slides or words (Lifshitz, 1966; John et al., 1967), as we11 as informa-
tion contained in the stimulus (Sutton et a1., 1967; Buchsbaum, Fedio, 1969)
and meaning of the stimulus for the sub~ect (Kostandov, 1977; Jennes, 1972)..
Many studies dealt with dependence of EP on level of attention (Garcia-
' Austt et al., 1964; Mackworth, 1969), and it was found that relevant, or
meaningful, stimuli that the subjAct had to count or to which he had to
respond always induced more marked EP than irrelevant ones. Here, the
link betw~en EP and the entire behavioral act�is particularly dist:inct.
In extreme cases of distraction of attention, EP are not recarded at all,
as had already been demonstrated by R. Hernandez-Peon (1960, 1961). ~
In our experiments, in response to presentation of rhythmic flashes of light
as a conditioned signal reinforced by electrocutaneo.us stimulation (ECS),
a complete EP developed only to the first flash in a series, after which
defense behavior began and the next flashes were "unmeaningful" (Figure 7)
(Shvqrkov, Velichkina, 1970). We obtainad similar data with regard to
food-re~a*.ed behavior (Shvyrkov, Grinchenko, 1972), and they have also
been d~;:=ribed by many other authors. A correlation was also demonstrated
between EE and nature of future motor response (.Spinelli, Pribram, 1970),
time and probability structure of presentation of s~imuli (Jenness, 1972a,
b; Boddy, 1973; Poon et al., 1976), etc., in o:'ther words, all factors
determining integral behavior.
At the same time, corx'elations were demonstrated b~t~aeen EP and physiolo-
gical parameters: the configuration of EP changed when stimuli were de-
livered at different phases of respiration or the cardiac cycle (Callaway,
- M. Buchsbaum, 1965), with change in state of the thyroid (Shagass,~1975),
with adm~niscration of pharmaceutical agents, etc.
Interpretations o� these facts are ~ust as diverse as the data themselves;
- however, they can be divided into three groups. Some authors prefer to
interpret L~P chang~es in psychological terms, such as "perception,"
"attentian," "recognition," etc. The second group of explanations refers
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- to physiological discussion of the sources and pathways of conduction of a
given component. The third group uses the terminology of information
processes: "evaluation of signals," "information processing," etc., etc.
_ a , '
b
c
Figure 7. Evoked p~tentials in response to rhythmic flashes of
light, in somatosensory cortex of the rabbit with
development of conditioned reflex
a) before development of reflex
b) lst to 20th combinations, delivery of~electrocutaneous stimulation
~ S00 ms after the 6th flash
c) 21st-40th combinations -
- All these interpretations make some use or other of. the link between EP _
and parameters of the stimulus that induces them, while changes in EP con- ,
figuration are related to the modulating influence of either attention, emo-
tions, etc., or nonspecific structures, or informational meaning. A. M.
Ivanitskiy believes that "the possibility of recording a response in the. _
absence of stimulus is an objection to conceptions of the exclusively
- modulating action of nonspecific influences on late waves of the response
(1976, p 73), and he cites extensive facts to support this possibility.
- Iiowever, A. M. Ivanitskiy believes that this is the only ob~ection.
We believe that the list of objectians must also include the resu~*_~ of
experiments involving recording of "motor" potentials, as well as data that
_ there are no EP in the presence of tiie stimuli described in the preceding
section. But the decisive objections which, we believe, compel us to '
abandon these conceptions, were obtained from systematic experiments j
in the laboratory of E. R. John, which demonstrated the "endogeny" of
all EP components after 40 ms (John, 1972) or even 25 ms (John, Morgades,
1969). The main experiment of E. R. John consists of delivery of flashes _
at a frequency of 3 Hz to cats trained to depress one lever in response
to flashes at a frequency of 2 Hz and another lever, in response to ~
- flashes at 4 Hz. This was associated with "generalization," and the cats ~
sometimes went to one lever and sometimes to the othez. The EP configura-
tion in response to such a"generalizing" signal corresponded expressly
to the light that was a"signal" for the ~ever to whicfi the cat went. . ~
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- The conditions of these experiments were changed in all sorts of ways.
The "generalizing" flash was replaced with a sound or electric stimula-
tion of brain structures; depression uf the lever was reinforced with
either food or elimination of a possible nociceptive stimulus. In addi-
tion to EP, aetivity of neuronal "ensembles" was also recorded, and a
computer was used to define the configuration of EP and "population
activity" corresponding to specific behavioral acts.
Al1 these modifications revealed that in response to any stimulus there
was reproduction of the EP configuration corresponding to specific beha-
vior, and it was unrelated to the parameters of the stimulus. In order
_ tu stress the independence of EP configuration from the parameters of ~
exogenous stimu~ation, E. R. John called the components "endogenous~~
after 25 ms or 40 ms, i.e., reflecting internal activity of the brain
read out of inemory (John, 1973). _
The fact that the early components were present with any form of behavior
served as grounds to consider these components (up to 25 or 40 ms)
"exogenous," related to input in the brain of ''external information."
The fact that EP reflect the "memory of prior experience" (John, 1973,
p 209), i.e., activity retrieved from the organismts memory, makes it
possible to explain the dependence of EP on a11 factors in the exogenous -
and endogenous environment. According to functional system theory,
retrieval of a given behavioral act from memory depends on both endogenous
= factors (motivation) and exogenou~ ones (situat~on). This shows that a
stimulus is exclusively an impetus, or trigger factor, which does not
determin~ endogenous brain processes, but only triggers them.
- This is also confirmed by experiments, in which an exogenous acoustic
stimulus was replaced with electrical stimulation of the auditory cortex
(Miller et al., 1969) or photic stimulus was replaced by stimulation of
the external geniculate body or visual cortex (M. t. Glickstein, 1972). -
The latency period of the motor response diminished with electrical stimuli
by exactly the magnitude of the interval occupied by the primary component.
These utterly artificial electrical stimuli 3pparently replace entirely
the exogerious trigger signal, although, of course, it is unlikely that
they carry "information about the physical properties of the stimulus." _
Since, according to our hypothesis, EP reflect processes of coordination
of elements of different structures into a single system, "endogeny" of EP _
signifies that the trigger stimulus reproduces processes of concordance
of the elements that previou~ly formed a functional system of the corres-
- ponding behavioral act.
Link Between EP and Future Events
According to function.al system theory, the activity of different brain
structures in the behavioral act is not only "endogenous," i:e., retrieved -
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from memory, but goal-directed. In other words, the choice of a specific
activity from memory is determined by hierarchically organized goals or
anticipated future events.
The link between EP configuration in response to any trigger stimulus
and future events can already be seen in the fact that EP to a
conditioned stimulus followed by reinforcement is significantly different
from the EP in response to an "indifferent" stimulus, as has been
demonstrated in a vast number of studies.
The experiments of A. I. Shumilina (1965) revealed that the configuration
of EP to a conditioned stimulus depends on the quality of reinforcement.
EP in response to the same flash of light, with respect to its physical
parameters, present different conf igurations when the flash serves as a
conditioned signal of future food or defense reinforcement. The above-
cited data of E. R. John can be interpreted as confirmation of another
aspect of dependence of EP on future events. While in the experiments of
A. I. Shumilina the same stimulus served as a signal of different future ~
events, in the experiments of E. R. John different stimuli, which triggered
the same behavioral act, were signals about the same future event.
There was direct demonstration of the possibility of purposeful transforma- :
tion of EP when a specific EP conf iguration leads to reinforcement in the
experiments of S. Fox, A. Ruddel (1970) and J. Rosenfeld, R. Owen (1972).
All these data warrant th~ assumption that the configuration of EP reflects
org~nization of processes that leads to a specific future event. Of
course, event isnot a purely physiological concept. We have already
_ stated that, as an organized aggregate of elements in the environment, it
can be compared only to a specific organization of physiological processes.
- For this reason, the link between configuration of EP in response to some
triggering stimulus and a future event can be demonstrated only by
comparing the configuration of EP corresponding to two consistently
successive events.
We found that with recording of EP in response.to light and EC.S in
the somatosensory cortex, in the conditoned defense reflex, that these EP
become amazingly similar (Shvyrkov, Velichkina, 1970). At that time we
evaluated this phenomenon as the correlate of "anticipatory reflection"
and manifestation of the model of future ECS, according to conditioned
si.gnal in expressly the somatosensory cortex. However, this conclusion '
was derived without considering the fact that EP reflect general cerebral
systemic processes, rather than special functions of the somatosensory
cortex.
The objective of the next series of experiments was to compare the EP ~
~ configuration with conditioned and unconditioned stimuli in several
structures of the brain performing di�ferent special functions. It is
~
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generally believed that the visual and somatosensory cortex perform different
functions in the conditioned defense reflex to light, reinforced by
electrocutaneous noxious stimulation: the conditioned stimulus is analyzed
in the former and the unconditioned, in the latter.*
It was assumed that comparisonfof EP configuration in response to light
and EC S would yield information about the extent of dependence of configu-
ration of EP on light and configuration of EP to ECS, and thereby about
reflection in EP configuration to light of the future event--electro-
cutaneous stimulatioa. A comparison of EP configuration in the visuzl and
somatosensory regions would ~grmit differentiation of components related
to general cerebral, systemic processes from components rela;.ed only to
special functions of one structure.
- Although there is no analogue in nature of constant combinations of light ~
and electrocutaneous stimulation, this experimental model is methodologi-
cally very convenient for the study of elementary behavior. One can `
arbitrarily consider reduction of the deleterious eff ect of electrocu-
taneous stimulation as the goal of this behavior (Laptev, 1949; Ivanova,
1970). " -
We conducted our experiments on nine adult rabbits whose paws~w~re
immobilized on a stand. The conditioned defense reflex and differentiation
_ were developed in one session, during which the rabbit received about
300 combined and separate stimuli. Three flashes of light, synchronized
with clicks delivered at 700-ms intervals, served as the conditioned
stimulus; 700 ms after the last flash we delivered reinforcing ECS,
square-wave pulse lasting 1-500 ms, with amplitude of 40-120 W, and
intersignal intervals of 30-90 s.
*In using the terms "conditioned reflex," "conditioned stimulus," etc.,
- we are merely following the physiological tradition of their referring
to certain experimental procedures, but by no means do we impart in these
terms their ori~inal conceptual meaning. In this book, we shall not ~
specially discuss the problem of formation of new behavioral acts; let
~ us merely indicate that, with the systemic approach to analysis of the
mechanism of learning, the very formulation of the problem changes: if
conditioned and unconditioned behavioral acts are not organized like
"ares" of corresponding reflexes, but as functional systems, there is
no physiological meaning to the question of bridging of a connection
between them. The organism does indeed detect a lYnk 3etween two events
and organizes a conditioned behavioral act, with due consideration of
future reinforcement, as established by I. P. Pavlqv. However, the
conditioned behavioral act is not a copy of an unconditioned reflex, but
new integration, a new functional system organized to achieve a certain
- result, which plays the part of a system-forming factor. -
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Light flashes attenuated with a yellow filter, also synchronized with
clicks, served as the differentiation signal. Since the rabbit retina -
contains only rods, differentiation apparently occurred according to
brightness. Cond:Ltioned (reinforced) and differentiation stimuli were
delivered in series of 25, which was motivated by the convenience of
subsequent processing of evoked potentials.
The electrical activity derived from the muscles of the front leg served ~
- as a criterion of development of the conditioned ref lex. Electrical I
activity of the visual and somatosensory cortex was derived with needle
_ electrodes inserted in the cranial bone and immobilized with dental
~ cement. After amplification by means of a Biophase universal unit,
along with recoridng on an ink recorder, the EP and EMG were recorded
on tape on a multichannel recorder ~"magnettor"], using frequency modula-
tion, and then they were averaged on a Mnemograph ac cumulator unit.
_ The bandwidth of all of the equipment constituted 1.2-500 Hz. Time of '
analysis of evoked poten tials constituted 400 or 800 ms, and averaging -
of 25 runs was perfor~ed. We analyzed the responses to electrical
stimuli and the~first in a series of flashes, since preceding experiments
convinced us that, under such conditions, the EMG reaction appears
already after the first conditioned signal flash, and the responses to
expressly the first f lash unde.rgo the main changes related to developed
- of the condi*_ioned reflex; the responses to other flashes are depressed,
and they con;ain only the primary complex, as illustrated in Figure 7.
Our obj ective made �it necessary to analyze expressly the configuration,
i.e., the time pzrameters and component composition of an evoked poten-
tial, rather than amplitude.
Before combining the flashes with ECS, the responses to the former ~
varied significantly in t?~,a visual cortex of different animals, and they ~
containEd a dissimilar nu:~ber of components (compare Figures 8 and 11) .
The responses to white (future conditioned) and yellow (future differentia- ;
tion) light presented the same configuration, but the latency period of
the response to yetlow light was usually several milliseconds longer. ~
It is a known fact that there is a correlation between latency period of
EP and brightness of flashes (Shevelev, 1971).
Light also induced some response or cther in the somatosensory region, ~
and such responses were virtually absent in only three rabbits (Figure 8).
In the other three rabbits, tr~e evoked potentials even contained the early ~
negative components first described by K. M. Kullanda (1964), with a ~
latency period of 15-20 ms (F~gure 11). ~
As can be seen in Figures 8 and 11, before development of the conditioned
reflex, the responses to light could vary, not only in the visual and
somatosensory regions, but even in the left and right visual areas -
(Figure 8), which we had already observed in freely behaving rabbits.
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Right Left Left ~
visual visual , soma.to?ens.
_ Lig ECS Ligh ECS Ligh ECS
J
_ ,
3 ~
~ -
S
6
7
8
200m . ~ -
a b .c~:. ,d ' e ~
Figure 8. EP to light and ECS in right and ~eft visual and left ~
somatosensory cortex during development of conditioned -
reflex ~
Averaging of 25 runs at a time:
1) before combinatiorLs 6) continuation, 76th-100th combina- ?
2~ lst-25th combinationc tions
- 3) 26th-50th combinations 7~ 51st-75th delivery of differenti- -
_ 4) 51st-75 combinations ated light
S) 1sC-15th delivery of differenti- 8) former differentiation yellow ~
ated yellow light light reinforced by ECS, lst- -
25th combinations ~
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Figure 9. .
Comparison of evoked potentials to one
a ~ another and to averaged EMG reaction.
~ '
Same experiment as in Figure 8:
a) response to ECS in somatosensory
cortex after development of
reflex (frame '~f" in Figure 8)
~
- b b) response to light in somatosensory '
~ cortex after development (frame '
e-6) '
- c) response to light in left visual
cortex (frame.c-6)
- d d) averaged conditioned EMG reaction,
~
I�-- 200ms--?I 76th-100th combinations
The responses to ECS were also
a b individual. They differed only
~ in the first combinations in the
. somatosensory and visual cortex. ~
Already after 25 combinations, the ~
;
_ 1 early components of re~ponses to
ECS in the somatosen~~ry and visual
cortex were the sam~ an~ had a
latency period of 10-2G ms ,
- (Figure 11). The responses
usually presented initial positivity, _
~I but the main typical component of
_ the response to ECS was negativity
-~-zoams in all areas, with a latency period
~ of 20-40 ms. Its duration was ~
Figure 10. very indiviuual, ranging from 40 to
Comparison of responses to conditioned 200 ms in different animals. Nega- _
(a) and differentiated (b) light in tivity was followed by a late
visual (I; (frames c-6 and c-7 in positive component. '
Figure 8) and somatosensory (II) ~
(frames e-6 and e-7) cortex. Same The dynamicg of responses to ECS
experiment as in Figure 8. during development of conditioned !
reflexes consisted of simplification ~
of configuration, and they were similar in ~ll of the examined parts of
the cortex (Figure 8) . ,
Responses to light, which had become a conditioned signal, were completely
_ transformed in both the visual and somatosensory regions after 25-50 com-
binations; at the same time, a stable conditioned EMG reaction appeared. ;
A comparison of configurations o~ EP to conditioned light in the visual
and somatosensory cortex showed them to be very similar (Figures 8, 9 and ;
52 -
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- 11); there was coincidenc~ of latency periods and duration of phases,
particularly the negative one, and often subcomponents as well.
Light ECS Conditioned Differ.
i
2 i
3 .
-r _
Figure 11. Comparison of responses in left visual (1) and somato- -
~ sensory left (2) and right (3) cortex to light before
development of reflex, to ECS, conditioned light (25th-
50th combinations) and to differentiation yellow light
(25th-50th deliveries). Averages for 25 runs, 400 ms
, frames
In other words, in defense behavior induced�by a conditioned or uncondi-
tioned stimulus, the evoked potentials in different parts of the cortex
were also synchronous and similar in configuration, as in the previously -
discussed food-related behavior. It is very important to note that, in
oiir experiments, the responses to conditioned light in the somatosensory
~
- cortex always contained a short, early component, which was usually
positive (~'igures 8 and 10). In two experiments and before development
= of the conditioned reflex, light induced an early negative component in
the soma.tosensory cortex, which persisted even after it was developed
(Figure 11). The early components had a latency period of 15-20 ms, which
was the same as the latency period of responses in the visual cortex -
(Figures 9 and 11). Appearance of such early oscillations of potential in
response to a conditioned signal, at the "point of reinforcement" had
already been reported in the literature (Artem'yev, 1959). The early
component was followed by negativity, which lasted 40 to 200 ms in
different ra~bits and was complicatea :ry a different number of subcompo-
nents, and late positive oscillation. -
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A comparison of the responses to conditioned light and ECS in somatosensory
regions confirmed our previous data (Shvyrkov, Velichkina, 1970) indicating
_ the similarity of their configurations. The responses to light and ECS in
the visual cortex after development of reflexes were also similar
(Figure 11). If the response to ECS changed, similarity was observed .
between the response a given rabbit presented before the conditioned ~
reflex, rather than the one ~hat was transformed as a result of develop-
ment thereof (Figures 8 and 9). This transformation of EP in response to
- ECS was studied by us in separate experiments (Shvyrkov, 1969), and we
shall not discuss it specially here. The similarity of configurations of '
responses to conditioned light and ECS was graphically demonstrable with
any individual configuration of EP (Figures 10 and 11).
The responses to differentiated light in the visual cortex differed in
most experiments from the conditioned evoked potential, in that they con- -
tained an additional negative oscillation and had no late positivity,
or else the latter was shifted in time (Figures l0.and 11). In the somato-
sensory cortex, the responses to differentiation light were always less
marked than before the combinations; they did not resemble the response
to ECS or conditioned signal and, what is important to note, they contained
no early components (Figures 8, 10 and 11). With the reinforcement of
- differentiation light, the responses immediately acquired all of the
features of a conditioned one, in both the visual and somatosensory regions ~
~ (Figure 8). ;
The conditioned EMG reaction appeared after 25-50 combinations, and it
had a relatively stable latency period for each rabbit, from 50 to 300 ms.
The EMG reaction began at the time of the posterior front of negativity
and late positive oscillation (Figure 9). According to the EMG reaction, ~
differentiation reached a 70-80% level after 25-75 separate presentations ~
of differentiation light.
These data indicate that the phenomenon that accompanies development of
a conditioned reflex does not consist merely of generalization or trans-
formation of ~he evoked potential to light, but the responses to conditioned
signals in the visual and somatosensory cortex become synchronous and
identical in configuration; and the responses to reinforcing ECS were
. also the same as before the combinations. This pattern was demonstrable in
all animals, with any individual configuration of EP.
The fact that, before and after development of the conditioned reflex, the
same stimulus could induce responses of utterly different configuration,
while different stimuli, light and cu~rent, cou13 evoke the same responses
after development (in different regions) demonstrates, once more, that
the ~echanisms that determined EP configuration in behavior are qualita-
tively different from the mechanisu:s that detErmined EP configuration in
anesthetized animals, in which case there is a distinct link between
configuration and distribution of EP, on the one hand, and anatomical
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porjections of receptor surfaces, on the other hand. This serves as
additional confirmation 6f the "endogeny" of processes, the correlates
of which are EP.
Our data indicate that the primary response, at least in
the somatosensory cortex, is also "endogenous," since presence or abaence
thereof are directly related to reinforcement and, consequently, like
the later components, the primary response also corresponds to prior
experience stored in memory. ~ _
The link between configuration of EP in response to light and to current
also indicates that retrieval of a certain organization of physiological
processes from memory takes place in accordance w ith a future event. The
_ order of processes may be as follows. According to functional system
tY:eory, any stimulus that appears in the environment finds preliminary
integration of elements prepared, and determined by the future event, the
_ appearance of which is predicted by motivatiqn and~situation.
In our case, this event, which generated preliminary [pretrigger] integra-
tion, was ECS. The result, i.e., attenuation of the deleterious effect of
ECS, w~as achieved by means of a functional system that contained specific
elements in different structures, including the visual and somatosensory
- cortex. Coordination of activity of expressly these elements was re-
flected in a specific EP configuration in response tc ECS. Since motiva-
tion (defense) and situation (constant) do not predict any future event
other than ECS, the preliminary integration in our experiments corresponds
mainly to one future event and one goal: to reduce the injurious effect
of ECS. The similarity of EP in response to light and ECS can be explained _
by the fact that the light flashed in the presence of preliminary integ-
ration created by ECS, and after it there was coordination of activity of
mainly the same elements that were involved in the functional system of
the unconditioned behavioral act.
We tested thi~s hypothesis in special experiments, where we recorded
neuronal impulsation activity, which is discussed in Chapter 4. Here,
let us merely mention that, since EP in response to differentiation
flashes differ from EP to both conditioned and indifferent flashes, it
- is imperative to assume that development of differentiation consists of
formation of a separate behavioral act, the functional system of which
is formed of different elements than the functional system of the other
acts studied.
Although delivery of combinations and differentiation flashes in our
experiments was performed in blocks of 25 presentations which, of course,
_ led to a change in preliminary integrations already after the first
flashes in a successive block, we can still assume that both preliminary
integrations exist from the time of introduction of differentiation
. light in any intersignal intervals.
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Triggering of a given integration is determined by the parameters of de-
livered light. Sin~:e, in our experiments, the EP to a conditioned light
differed from EP to differentiation light in tha t there was already a
primary response with a latency period of 15-20 ms in the somatosensory
cortex, it should have been assumed that not only "identification of
- physical properties," but "determination of signal meaning" of the light _
occur within the latency period of cortical EP. This conclusion. seems
paradoxical. However, one should apparently seek the cause of the para-
- dox in the conceptions of identification of physical properties and
determinatien of signal meaning of a stimulus as real processes,
EP Components--Correlates of Systemic Processes of the Behavioral Act
It appears to us that the EP distinctions demons trated in behavioral
experiments compel us to change the view of EP as a physiological pheno-
menon. Synchronism and similarity of EP configurations in different
structures do not offer grounds to maintain that there are some uni-
directional "afferent messages" cr "flows of excitation" spreading from
one structure to another. Rather, we can conceive of multilateral ex-
change of influences be~tween elements of many structures, which occurs at
- each phase of an evoked potential.
This hypothesis can also be extended to EP demons t rable under anesthesia,
the only difference being that elements involved in interaction processes ;
under anesthesia are limited to constant "narcoti c" preliminary integra-
tion, which could refer to the correlation between intact functional
links between struc~ures and links impaired by a specific anesthetic. As
we know, with the use of different types of anesthesia, stimulation of
- the same nerve elicits an evoked potential with d ifferent localization
and configuration (Nabil', 1969).
Thus, the possibility of recording EP under anesthesia, without integral
- behavior, is not in contradiction with the concep tion that EP are linked
with systemic processes of organization of the int egral behavioral act:
- EP is a phenomenon that reflects processes of any interaction of many
elements. In behavior, this interaction is determined by the goal and -
goal-directed organization of preliminary integra t ion; under anesthesia,
this interaction is due to the stable state creat ed by anesthesia.
The distinctions of EP in integral behavior are an expression of these ;
differences: synchronism and similarity in functionally different struc- ~
tures; dependence of configuration on future event s and relative indepen-
dence of parameters, modality and even presence of a trigger stimulus; ~
distinct link with time intervals of the behaviora 1 act. _
In the co~ltinu~~m of behavior, a single behavioral act--single organiza-
tion of activity of elements--replaces another behavioral act--another
- organization. Transitional processes triggered by a stimulus, the result _
of a prior act, are reflected in EP. ~
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According to functional system theory, there must be processes of afferent
synthesis and decision making in an elementary behavioral act, between
the stimulus and start of action: start of muscular contraction already
indicates implementation of the acceptor of action results and program
of action.
The rather stable correlation between EP components in different behavioral
acts and start of the EMG reaction warrants the assumption that the
_ primary response and negativity of EP correspond to processes of afferent ~
synthesis and decision making; late positivity already coincides with the
start of actuating mechanisms of the behavioral act, which are integrated
in processes of the acceptor of action results and program of action.
In order to determine more precisely the meaning of different EP components
as correlates of processes in the functional system of behavior, we con-
" ducted several series of experiments.
~
In the first series, we studied a behavioral act contained in the continuum
of behavior in order to track the dynamics of processes corresponding to
the moment of transition from one behavioral act to another.
According to functional system theory, motivation and situation retrieve
preliminary integration from memory, which corresponds to the goal of
entire behavior. This goal is hierarchically organized, and preliminary
integration includes all elements of future behavior. Performance of the
- first behavioral act and achievement of the first result out of the _
entire hierarchy, which ].eads tn achievement of the ultimate goal, must be
associated with the following successive processes: comparison of para-
meters of achieved result to the acceptor of results of action of this
act, afferent synthesis and decision making of the second act; then
there is formation of the acceptor of action results and program of
action for the second act, which determine action until the results of
the second act are reached, etc.
We simulated this segment of the continuum of behavior in the model of
instrumental behavior, in which rabbits turned on a flash of light and
headed for a feeder by pulling a ring with their teeth over a specific
distance. Both behavioral acts monitored by the experimenter (pulling
the ring and approaching the feeder) are contained in the general func-
tional system of food-obtaining behavior, but each of them is a functional
system with its own interim result. We can describe this segment
of the behavioral continuum schematically (Figure 12).
The first objective of our experiments was to have expressly the flash
of light serve as an interim result, i.e., the goal of tugging and
triggE~ stimulus to run toward the feeder. Experiments were conducted on
16 rabbits in a specially outfitted cage. Flashes of light were
delivered from the top at a distance of 70 cm.from the floor of the cage
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(flash energy 0.3 J and duration 50 us) from a Soneclat stimuZator lamp.
In experiments on 8 rabbits, we used a series of six flashes at 600-rms
intervals; in the experiments on the other 8 rabbits, we delivered
series of 3 flashes at 700-ms intervals.
Tric~ger , i
Trig er A'ffer- c ion stim. Affer,~ Action ~
stimu~us ent ^esult synth. resul~t '
s nth. acceptrs (light) accap
~r~g~ � (light) (c~bage~
~
, ci- ig Deci- ~ a9e
sion result sion result
aram. . param.. .
~
i. ' ~
- Action Action '
pmgram rogram
( u~l.n Aoin~n ,
~g t 5
xing) Resnlt eeder Result~ I
Figure 12. Schematic rendition of segment of behavioral continuum
In all of the experiments, activity of the visual and sensorimotor cortex
was derived monopolarly using implanted electrodes. The silent electrode
- was over the frontal sinuses. We used special stainless steel pins to
- derive electrical 3ctivity of cervical muscles, whir_h were inserted in the
_ skin on both sides of the neck; the EEG and EMG were recorded on a
Polygraph XVII electroencephalograph; in addition, a tape recording ,
was made of the experiments on the 8 rabbits to whom three flashes of ~
light were presented.
In these experiments, electrical activity of the cortex and EMG were '
recorded, after amplification on a universal Biophase unit, on magnetic
- tape. For reproduction from the tape, electrical activity was averaged ~
_ using a Mnemograph accumulator unit. ~
In the first experiments, the animals developed a classical conditioned
reflex: the flash was reinforced by automatic presentation of a feeder '
with 10-20 g cabbage. The control pulse for presentation of the feeder
as 50Q ms away from the last flash. Thus, the first fTash was more than
2 s ahead of presentation of the feeder. The conditioned reaction was
recorded as electrical activity of cervical muscles (Figure 14). Already
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after 3-5 combinations, the rabbits turned their head and went toward
the feeder in response to the conditioned signal. After 10-30 combinations,
running to the feeder was triggered by a flash in almost 100% of the cases.
During the second to fourth experiments, i.e., after 100-150 combinations,
we began to develop the instrumental behavior of tugging the ring, in
which we put a cabbage leaf for the first 3-5 times. A string connected
the ring to three contacts, which were so located that to bridge the
� first one the ring had to be pulled over 3 cm, for the second 8 cm and
the third 14 cm (Figure 13). By tugging at the ring, the rabbit could
successively bridge all contacts; however, the flash and then the feeder
- were presented only after bridging of the contact that the experimenter
connected to the stimulator.
.
- . . . I z~ ~ ' ~ -+~r.
II
.
,T
. 1 p 1 ~
y tirt , .;F~
! A V~I ~
- y~;~....
w~: .f.
� ~
~'!r ' r . '~q 7~
~I ~
~ ~7+~
f; ~f t f x"~
~ { �~'v~''~' ~ ~;ade?'~
~
~
J,
S'
Figure 13. Experimental cage. By tugging the ring with its teeth,
the rabbit moves a lever and bridges contacts I, II, III,
- one of which turns the light on
Interestingly enough, already in the first tests many rabbits, after
reaching for the cabbage leaf in the ring and unintentionally making
contact, dropped the leaf when the flash appeared and headed for the
feeder, which did not yet contain any cabbage.
At first, the first contact wa~r effectively produced. But when the
rabbits learned to tug the empty ring and this skill became f ixed (usually
after 50-70 times), each of the 3 contacts became alternately effective.
We tried to change contacts in random order; the procedure for this
change consisted of silent movement of the switch on a console 3 m from
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the cage. Thus, the rabbit did not receive a signal that the contact was
- changed, let alone which contact would be effective when it next pulled
the ring. ~
a b. c d . ~
i+1~-~~4'1,~I'h' +�.K+-~�"^M'1, ~1~.....+-avw~r ww~w.,.+.,-.,.w+~,.---�-�
1~I~N`yMN~ ~aht~~1~J'y,q1~M4~+~tWti?~.vv``"4'~N~+~~Ir~'+~+~~VM44~+1~A~Mikh~rM11+~
--~--~Irw~?'
- '-T-1~-'(- TT~"
I I ~11n111 ~ --Y
1~- .
Figure 14. Classical conditioned reflex (a) and instrumental
behavior (b, c, d) of rabbit. _
- Top to bottom: EEG of sensorimotor and visual cortex; EMG of cervical
muscles; marks for 3 flashes and presentation of feeder; marks for
making contact--in b, c, d, a--conditioned EMG activation,correspond-
ing to the rabbit's turning toward the feeder, begins after the
first flash.
Nevertheless, the experiments showed that all rabbits related quite
~ accurately pulling the ring to appearance of light: if they saw the.
light after pulling the ring over 3 cm (Figure 14d), they immediately
released the ring and headed for the.empty feeder; if,~ however, the -
_ experimenter rendered the third contact effective, the rabbits pulled
the ring to the maximum dietance. They did not always succeed in so doing
- at the first try; however, the rabbits did not stop trying and. did not -
head toward the feeder until the light was flashed (Figure 14b). When
they failed, they often "stood up" and sniffed the lamp. One of the -
rabbits, who could not pull the ring over a distance of 14 cm by moving
only its head, had to first tug the ring down with its paws, then grab
the string from the contacts in its teeth and addYtionally pu11 it out by ~
moving the head. As soon as the light appeared, the rabbit dropped the
string and ring, and headed for the feeder. ' ~
Since no signal was delivered as to the distance over which the ring had
to be pulled and the rabbits pulled it out each time over a different
_ distance, it must be conceded that, in our experiments, the scope and
discontinuation of pulling were not determined by the conditioned signal,
but by the goal. e
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The cage itself or the ring may be considered the signal to trigger
- pulling; however, these factors were constant, while the rabbits pulled
the ring over different distances each time, for which purpose they
performed different actions, including some to which they were not
specially trained. Since the volume of movements varied and could not
be determined by any stimulus prior to the start of movement, the event
that stopped pulling the ring had to appear as a result of movement.
_ Evidently, merely the position could not be such an event, since the
rabbits pulled the ring over different distances, and kinesthetic
aff erentation arising when the ring was pulled out for 3 cm was associ- _
ated with termination of action in some cases and continuation in others.
Evidently, the result of action could not be the feeder itself, since
the rabbits released the ring with appearance of the first flash.
- The light was expressly the result that stopped pull.ing at the ring
_ and it was necessary to stop pulling, whatever the mode of act~on.
By altering the effective contacts, the experimenter could always
predict which action would be performed and when it would stop.
Thus, our experiments showed once more that it is expressly the model
of the result, rather than any conditioned signal, that determines the
= range and mode of action performed to achieve it.
~ Whenever the ring was pulled there were many consequences: appearance of
the sound of movement of the lever, change in position of the ring, posi-
tion of the rabbit, etc. However, only the light~had the property of
= stopping the pulling. The distinction of light from the other conse-
quences is expressly that it emerges as a foreseeable and necessary
event, i. e. , as the goal of pulling.
_ ,
Of course, light acquired this property as a result of prior development
of signal-related link with the feeder, which was the more distant goal
of the entire food-obtaining behavioral cycle: approaching the ring--
pulling--obtaining light--approaching feeder--receiving cabbage. Thus,
we added light to the general hierarchy of results of food-obtaining
behavior and, consequently, it merely served as an interim, but immediate
goal, which was reached by pulling the light, and it was con~tained in the ~
hierarchically organized goal of the entire food-obtaining behavior.
We observed a rather interesting form of behavior in some experiments.
Af ter becoming satiated by the end of an experimental session, a
rabbit began to tug the ring often and regularly, relating the pulling
distance to appearance of light. It did not pay attention to the _
automatically presented feeder with cabbage, and could tug the ring even
at the moment the feeder was presented (Figure 15). After pulling the
ring over the required distance, it waited for the end of the series of
flashes, then pulled the ring again.
~
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~a,~,, ~ Thus, light added to the hierarchy -
_ of goals acquires inclependent
meaning as r~inforcea.:~ent, although -
the feeder, of which it is a signal,
temporarily lases this meaning
- ' due to elimination of the motiva- -
. tion of. hunger.
' ~ In our opinion, these findings also
corroborate the conclusion that
the flash of light is the immedi-
r~ ate goal of pulling the ring. Evi-
dently, achievement of this goal -
_ elicits some positive emotional
~ state, similar to the state of
~ satisfaction that appears when
the ultimate, biologically useful
effect is reached. It may be
Figure 15. assumed that, in this instance,
Satiated rabbit regularly pulls ring behavior is no longer guided by
and obtains light, but ignores the hunger, but by newly acquired,
feeder completely. Top to bottom: secondary (Miller, 1960) motivation
EEG of sensorimotor and visual cor.tex; that causes game behavior.
EMG of cervical muscles; marks of 3~
flashes and presentation of feeder; The results indicate that the
marks of closing contacts first objective of this series
was reached, that we developed a
method, with which any stimulus can be made the goal ot a behavioral act,
or method of "enrichment " of the acceptor of action results with
additional events.
The proposed modification of instrumental behavior does not differ
essentially from methods that are already known (Skinner, 1938; Beritov,
1961; Konorski, 1970, and others). However, addition of the procedure
of "variable action controlled by the resultt� enabled us to become con-
vinced that pulling the ring was indeed performed to obtain expressly
the light.
In the same experiments, we tried to demonstrate the electrographic cor-
relates of systemic'processes and, particularly, formation of the interim
acceptor of action results, i.e., prediction of light. For this purpose, .
_ we analyzed the electrical activity of the visual cortex at the time
- preceding pulling the ring and appearance of light. According to data
in the literature concerning the possibility of reproduction of the
rhythm of "marked" unconditioned reactions to a conditioned signal (John, ,
1966; Ruchkin, John, 1966), we could have expected appearance at this =
time of oscillations in the rhythm of future light in the visual cortex. '
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-
_ I ~.oo uv ~
~oo ~ -
~
Figure 16. 158th pull of the ring. No EEG oscillations in
- flash rhythm either before pulling or during the
action of light. Top to bottom: EEG of sotnatosensory and visual
cortex; EMG of cervical muscles; marks of flashes and presentation
- oi feeder; mark of making c~ntact
In additiore to visual appraisal of the ink tracings (Figure 16), we
performed reverse averaging using a mnemograph in the series of experiments
_ on the 8 rabbits exposed to three flashes of light. This procedure
consists of reproducing the tape by moving it in the reverse direction.
We averaged 25 runs at a time; the mark of the first flash served as
the trigger signal, and analysis time was 2 s. Our analysis revealed that,
when the rabbit pulls the ring, there is development of negative oscilla-
. tion in the visual cortex, which precedes appearance of the first flash
(FYgure 17a, 2, 3, 4). During the first pulls, it starts 1100 ms b~fore
appearance of the light (Figure 17a, 2) and upon fixing of instrumental
- behavior it starts 850 ms before (Figure 17a, 4). A comparison of this `
negativity to the start of Pulling at the ring was, unfortunately, not
feasible, since the moment of movement toward the ring was not fixed in
our experiments, and pulling itself lasted a different period of time -
in each instance. However, it is apparent that this oscillation also
existed during pulling, since the latter stopped after the flash.
The fact that the described negative oscillation increases with strengthen-
ing of the skill renders it similar to an anticipatory wave or conditioned
negativity (G. Walter, 1965). In our experiments, light served as the
expected stimulus, and therefore it may be assumed that the dynamics of
negative oscillation reflect the dynamics of formation of the interim
acceptor of action. Perhaps, the so-called readiness potentials that
precede voluntary movements of man (Deecke et al., I969), which are -
- similar to the above-described negativity in the rabbit, also reflect
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for~ation of the parameters of the result of voluntary movement. The
- schemes of the experiments are very similar, since in most studies the
subjects were instructed to achieve a"good~' movement, monitoring their
own EMG on the oscillograph screen. At the same time, this slow negative
_ oscillation must also conform with performance of the program of action,
since it develops while the rabbit is pulling and stops when it stops '
pulling.
~ -
a b ~
. -
i i-
Z ,
~ ' ~
~
- ~ 100 uV -
OOOms 2000 ms ' BOO.~ns I
_ Figure 17, Averaged activity of rabbit's visual cortex during !
classical conditioned reflex (1) and at different stages
- of instrumental behavior (2, 3, 4) ~
a) activity preceding presentation of light
b) evoked potentials in response to all 3 flashes; time of flashes
is narked by arrows at the bottom
c) response to first flash in the same combinations but a different
. time scale; in each case, 25 runs are averaged
1) classical conditioned reflex
2) instrumental behavior, 14th to 39th pull of the ring ~
_ 3) 76th to 100th pull; in (c), the arrow shows additional negativity
4~ l~lst to 176th pull; additional negativity also shown by arrow
It must be noted that we failed to demonstrate oscillations of rhythm of
, future light in the visual cortex, either in averaging,or analysis '
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of each separate tracing in the period before pulling the ring and during
the latter. Moreover, already upon formation of the clasaical conditioned
food reflex, we found that evoked potentials to the first and subsequent
flashes diff.ered in amplitude and configuration, as in defense behavior,
in accordance with previously described findings. The conditioned EMG
reaction started in our experiments after the first flash (see Figure 14a), ~
and cortical responses to the first flash were always the most marked
(Figure 17c, 1), whereas responses to subsequent flashes were deformed
and disappeared. The structure of EP to the first f lash before addition
of tugs did not differ from that described before.
In the situation of instrumental behavior, the rabbits stopped pulling
right after the first flash. The response to the f irs t flash remained
the most marked, although its configuration changed (Figure 17, 2, 3, 4).
- The chan~es in configuration consisted chiefly of appearance of an addi-
- ti~nal late negative oscillation with amplitude of about 100 uV, which
appeared at the site of ~he former late positive peak. Figure 11, 3, 4,
where this negative oscillation is shown by arrows, indicates that
late positivity does not disappear, but is shifted in time by about 100 ms.
Appearance of additional negativity did not cause an increase in latency -
period of the late positive oscillation in any of the rabbits, since this
_ negativity lasted only 30 ms.
As the skill in pulling the ring became more fixed, the additional negative
oscillation increased in amplitude. Figure 1Z shows that the increase in
additional negativity of EP is concurrent with increase in ~~egativity - -
preceding the flash. At Lirst, we assumed that the additionai negative
_ oscillation could be somehow related to comparison pro cesses (Shvyrkov, _
Grinchenko, 1972). However, the fact that similar additional negativity
appears in response to differentiation light in def ense behavior, as well
as that comparison processes occur also when the flash is delivered
without pulling, compelled us to re~ect this hypothes is. -
Since "light-goal" differs from "trigger light," not only in that the
former coincides with a specific moment in the rabbit's behavior, but
also in that it stops active pulling, whereas the triggering light stops
only passive anticipation, the hypothesis was expounded that the addi-
tional negativity is related to processes of discontinuing active pulling
(Trofimov, Grinchenko, 1975). However, control experiments proved this
to be wrong.
In the control experiments, which were conducted on 2 rabbits, only one _
flash was delivered, and a lag of 200 ms was produced between closing
contact and delivery of the flash. This resulted in a change in rabbit
behavior: after pulling the ring over some distance, the rabbit waited
for the light either after dropping the ring or holding it in its teeth.
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In this case, when we compared the EP in response to "triggering" light
and "light-goal," we also observed additional negativity (Figure 18). On
the basis of these experiments it was concluded that appearance of addi-
tional negativity was not related to the preceding acti, i.e., pulling,
but to the one for which the flash was a trigger, i.e., the act of ~
heading for the feeder.
Such doubling of negative oscilla-
tion has been described in experi- '
� ments on man with increase in number
of alternatives, out of which the ~
a sub~ect must choose one. For '
example, in the experiments of Ya.
A. Peymer (1971), additional "
oscillations appeared when the ~
b subject had to determine one out of '
- several possible positions of a
pointer on a briefly displayed
dial or in response to a flash
which was a signal for the reac-
tion of choice among four alterna-
tives. The possibility of alter-
500 ms ~ natives means that elements
. are involved in "pretrigger integ-
_ Figure 18. ration" in activity required to
Doubling of negative oscillation and perform all possible behavioral acts;
appearance of P-300 in EP to light the decision making mechanism, which
200 ms after making contact; arrow is triggered after the stimulus, -
indicates time of appearance of flash; chooses only one of them,~i.e.,
n=-25 it reduces the superfluous degrees
a) potential in response to of freedom.
trigger flash -
b) evoked potential to flash of It may be assumed that, in our experi- '
light--result of pulling ments, "t~~gger light" coincided
with only one formed pretrigger
integration in the simple conditioned reflex, corresponding to the run
toward the feeder. Implementation thereof corresponds to EP with one -
negativity. When pulling on the ring, the "light-goal" always was asso-
ciated with at least two mutually exclusive integrations corresponding to
conti.nuation and repetition of pulls, as well as running to the feeder. ~
This made decision making difficult, i.e., processes of implementing the -
one integration that corresponded to running to the feeder. Probably,
doubling of negativity was a reflection of this increase in superfluous
degrees of freedom and more difficult decision making.
It appears to us that this hypothesis can be extended to other instances
of doubling of EP negativity. In experiments involving a choice among
several alternatives, there was apparently no doubt as to the presence of -
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pretrigger integrations corresponding to several acts (Chuprilcov, 1978).
Addition of a differentiation stimulu~ in our experiments should also
have been associated with expansion of pretrigger integration.
Thus, double negativity can be interpreted as a reflection of "two-cycle"
reduction of superfluous degrees of freedom, i.e., a reflection of
processes of afferent synthesis and decision making that occurred twice. ~
Assuming that EP negativity corresponds, in any case, to concurrent
systemic processes of afferent synthesis and decision making, we can
observe that there remain only the latency period and primary response
for processes of comparison of par3meters of the result to acceptor of
results.
Since we had demonstrated in preceding experiments that appearance of a
primary response in the somatosensory cortex can be induced by a conditioned
signal and not by a differentiation one, one must assume that the appear-
ance of the primary EP component in nonspecific regions is attributable
already to the result of comparison and coincidence of parameters of the
light with some model thereof or other. This again leads us to the
_ assumption that such comparison is made during the latency period of cor-
tical EP.
The constancy of the primary EP component in projection regions, in rela-
tion to the modality of the stimulus, in different experimental situations
� l~d many authors to the conclusion that it is related to a reflection of ~
the "physical properties of the stimulus" (Ivanitskiy, 1976). However, the
"endogeny" of the primary component in nonprojection regions warrants the -
- a~sumption that it already reflects processes of implementation of pre-
- trigger integration, i.e., retrieval of activity of specific elements from
memory.
Thus, the primary component can be interpreted as the correlate of processes
of comparison of the real result--trigger stimulus and its model--acceptor
of action results.
We thus have some arguments for identifying the "anticipation wave," latency
period, primary component and negativity of EP with specific systemic
_ mechanisms of the eiementary behavioral act. -
The late positive component of EP, which may have different latency periods
under different experimental conditions and in different rabbits, coincides
~ with the start of EMG activity and changes to a slow negative oscillation, _
which we related to the function of the acceptor of action results and '
program of action. Consequently, it is contained between processes of
decision making and function of actuating mechanisms of the behavioral act.
The correspondence of this potential to the start of EMG activity led us
to analyze these correlations. As we have already mentioned, according to -
functional system theory, the behavioral act is implemented as organized
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_ activity of many elements, and any muscle could be involved in the actuating
_ mechanisms only to the extent of its contribution to achieving the result.
- The coordinated involvement of muscles is implemented by the "program of
action" or "effector integral," which can be determined from the order
of involvement of different muscle groups in actuating mechanisms.
Visual
cortex
Right .
motor
Lef t
motor
_ /
_ Z ~
3' ~-r ~
~
5
~ EMG
TAR
RT 100`ms
Figure 19. Correlation between EEG activity, latency periods (LP) ~f
EMG activation of different muscle groups, time of achieve-
ment of result (TAR) in one act of approaching feeder; RT--reaction
time.~ Top to bottom: EEG activity of right visual, right and
left motor cortex; 1, 2, 3, 4--EMG activation of right and left
groups of cervical muscles and posterior groups of brachial muscles '
of the right and left front legs, respectively. Below this, mark
for delivery of flash and feeder; 5--actogram
In experiments conducted with A. Kh. Pashina on ~five r.abbits involving the
simple conditioned reflex, where the light triggered going to the feeder,
we recorded activity of cervical muscles on the right and left, and
- activity of posterior muscle groups of the front legs, also on the right
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- and left. This activity was compared to the parameters of the late positive
component of EP recorded in the visual and sensorimotor cortex (Figure 19).
The experiments revealed that the order of involvement of different muscle
- groups is not constant, even after 500 runs to the feeder. One muscle
group, then another was first to be active (Figure 20), and the latency _
period of the very first EMG reaction constituted a mean of about 100 ms
(Figure 20).
In general, the distribution of
~s latency periods of involvement of
i different muscle groups coincided.
, When we compared EP to histograms
_ ~ of latency periods of EMG reactions
of any muscle, we only found that
~ , the earliest EMG reaction corres-
ponded to the posterior front of
negativity and anterior front of ~
S00 , late positivity, as was also
~ demonstrable when recording the
~ activity of one muscle.
i~ '
~n~ ,1 ~ A comparison of late positivity to
Z' ~ ' n latency periods of EMG activation _
, ~ , ,
� ~ ~ i l~'`_ ~ .,f of all muscles studied revealed
' ~ ` y~ ~ that all EMG activations begin
!00 ~ yi within the range cf the late posi-
tive component.
Zpp Combinations ZZg
Figure 21 illustrates the AEP in
Figure 20. the visual cortex, as compared to -
Correlation between latency periods of time of involvement of all muscles
activation of different muscle groups; in these 25 acts. A comparison of
x-axis, sequential number of behavi- ~ the top and bottom parts of this .
- oral ac~ y-axis, latency period, ms figure indicates that the form of
1) time of achieving result the late positive component of AEP
2) latency period of activation of corresponds to the compcsition of
right front leg muscles the time segments between involve-
3) left front leg ment of the firs;. and last muscles
4) right cervical muscles in acts that, were "avers~ed." It
_ 5) left cervical muscles is therefo,re possible that the
variabilil.y of configuration of
late posi;tivity in unaveraged EP
is correlated with the instability of time of in-~olvement of various somatic
and autonomic components in actuating mechanisms ~f single behavioral acts?
- By measuring the time between triggering of the first and last EMG reaction,
we can obtain information about the time of existing of the initial "efferent
integral." Subsequently, in the course of performing action, the constantly
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_ . Figure 21. ~
~ Correlation between late positivity '
- of averaged evoked potential (in '
` visual cortex) and time of involve-
r ment of all recorded muscles in ~
, ~ actuating mechanisms of the be-
havioral act. Averaged EP in
. 26th-50th acts (top) and lOlst- ~
125th acts (bottom). Under them
are the time segments in which ;
' ~ all muscles became involved.
~ The starting point on a line seg-
~ ment corresponds to the time of ~
S-~ involvement of the first muscle
~
and the last point, to.that of the
last muscle recorded. The top
� segment of the line corrresponds'to
the first run to the feeder and the
~ bottom segment to the 25th. The '
dot refers to combinations when
all muscles were involved simul-
- taneously and the arrow shows ~
~ ~ time of delivery of flash.
~
~
-
-
� - i---~
100 ms
~
incoming feedback, of course, corrects significantly and read~usts the ~
initial program of action within the framework of the precoordinated sub-
- systems on the physiological level.
According to functional system theory, the program of action is formed con-
currently with the acceptor of action results which actually determines !
the entire possible set of inechanisms included in the program. Conse-
quently, it is more correct to interpret the late positive component
as a correlate of the process of mobilization of actuating mechanisms of ~
the behavioral act, which trigger both the acceptor of action results and -
program of action. � � '
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The latency period of late positivity of AEP [averaged EP] in our experi-
ments could const~tute only 100 ms (according to maximum), but could also
= increa5e signif~cantly, particularly in tl~e experiments involving tugging
at the ring. For example, in Figure 17 this late positivity has a latency
period of about 300 ms. -
As we know, late positivity with a latency period of 300 ms, or so-called
P-300, has attracted the special attention of psychologists, since this
component appears in human EP in situations of "elimination of
uncertainty" (Sutton et al., 1965, 1967; Debecher, Desmedt, 1974; Ruchkin
et al,, 1975).
- Evidently, the latency period of the late positive component depends on
the duration of prior EP components. As we have already mentioned, the -
number of prior negative oscillations is related to the number of "cycles"
of afferent synthesis and decision making which, in turn, are determined
by the number of competing behavioral acts represented in general pre-
triggering integration. ~
Thus, late positivity does indeed appear at the ti~e when excessive
degrees of freedom, present in preliminary integration, are eliminated, '
but this "elimination of uncertainty" apparently occurs earlier, during -
the negative component.
From the point of view we are developing, P-300 does not differ in meaning
from the late positive component, which has a shorter latency period in ~
simple situations. In both instances, the late positive component cor-
responds to the process of mobilization of actu2ting mechanisms of the
_ behavioral act. The presence of P-300 only in response to "relevant" -
stimuli is probably related to the fact that it is only after such stimuli
that the corresponding actuating mechanisms become involved. The link
_ between P-300 and complexity of the situation can be explained by the
fact that it is only in such situations that additionll negative oscilla-
_ tions appear, which defer late positivity to a later interval; in simple
situations, the actuating mechanisms of the behavioral act become active
sooner, and one observes earlier positivity, and not P-300.
Thus, the general scheme of conformity of EP components with systemic
mechanisms of the behavioral act acquires the following appearance: in
the latency period and at the tine of the primary response there is com-
parison of parameters of the stimulus to its model; the negative component
corresponds to simultaneous processes of afferent synthesis and decision
making; late positivity is a correlate of simultaneous processes of forma-
tion of the acceptor of action results and program of action; "conditioned
negativity" serves as the correlate of actuating mechanisms of the beha-
vioral act--acceptor of results of action and program of action; the -
"stimulus-result" of a given behavioral act triggers the next cycle,
starting with the comparison procpss, etc. (Figure 80).
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All of the above-listed systemic pracesses are processes that relate the
organism to external events, both those existing before action and
those appearir_g during and after action.
It is convenient to begin our discussion of the entire cycle, in terms
characterizing organization of elements of the organism itself, with the
preceding behavioral act, when organization of real activity of elements
is related expressly to this prior behav~oral act. At this time, pre-
liminary integration, which corresponds to a possible subsequent behavioral
act, without being realized, increasingly loses "excessive degrees of
freedom" as the preceding act is performed. Upon achieving the result
of the preceding behavioral act, the "stimulus-result" initiates, like a
trigger, processes of reorganization of the activity of many elements; EP
are a reflection of this transitional process.
A change in organization probably does not occur during the latency period
of the primary component, and in this sense it is indeed a"latent" period.
The primary response corresponds to processes.of partial realization of '
preliminary integration, i.e., establishment of interaction between only
the elements whose "degrees of freedom" were coordinated at the time of
appearance of the "stimulus-result."
- tlegativity is a correlate of a comp.lete change from one form of organiza-
tion of elements to another. During zegativity, integratiun corresponding
to precerling behavior "falls apart," there is elimination of "excessive
degrees of freedom" of all elements contained in pretrigger integration,
- and only one form of organization of elements is left. In the ca~se of ~
competing organizations corresponding to different behavioral acts, within
the framework of a single preliminary integration, this process may be .
repeated several tiwes.
Late positivity corresponds to the process of involvement of all necessary
elements, i.e., implementation of a single integration, formed during
- negativity, and start of "maturation" of organization of a future behavioral
act.
Finally, slow negativity, referred to as "conditioned negativity," "wave
- of anticipation," "potential of readiness," etc., corresponds to processes
of implementation of actuating mechanisms of a current behavioral act and
"maturation" of preliminary integration for the next one, i.e., it corres- i
ponds to processes of organized function of physiological functional
systems contained in the hierarchy of the functional system of a given
integral behavioral act.
The characteristics of systemic processes from the standpoint of physiolo-
_ gical mechanisms are characteristics of processes of organizing the activity
of various elements into a single whole;for this reason, none of the
- EP components, according to the view presented, reflects only afferent or
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_ only efferent processes, and it is not related to stimulation of some
individual morphological, isolated pathways or structures. EP ar e
related to reorganization of activity of elements and relations within
the entire brain.
Of course, the above-presented conceptions concerning the correlation
between EP and systemic processes are speculative to a significant extent,
because of the complexity of the link between summated activity arid that
of single elements. In order to define the actual mechanisms invo lved
in processes of organizatLon, it is necessary to study the activi ty of ele-
ments, i.e., impulsation activity of single neurons.
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CHAPTER 3. SYSTLMIC ORGANIZATION OF NEURONAL ACTIVITY IN BEHAVTOR
Link Between Overall Activity and Neuronal Impulsation
For a long time, impulsation of single neurons, like overall electrical
- activity, was studied in accordance with reflex conceptions and related
conceptions of localization of functions. Neuronal activity appeared to ~
be a very obvious reaction to a stimulus, which came to the neuron under
study over specific pathways. According to the reflex conception,
excitation appearing in receptors activates chains of~neurons situated in
successively connected structures, which serve their own special functions
up to the effectors.
- These conceptions appeared to be so obvious and firm that the question
of inechanisms of the behavioral act was simply not posed in studies of
neur~onal impulsation. All efforts were concentrated on two different
ar~u unrelated directions; one consisted of examining neuronal impulsation
in different structures in order to determine the mechanisms of "sight,"
"hearing," "movement," etc.; the other concentrated on determination of
the neuronal mechanisms of learning, which was interpreted as formation of
a new "conditioned reflex," i.e., as "bridging of a new arc" between
receptors and other effectors than before.
From the standpoint of functional system theory, the main question that
should have been posed in studies of neuronal impulsation related to ~
behavior is the question of inechanisms of organization of activity of
single neurons into a single whole, into the functional system of the _
behavioral act. Since ttie :esearcher usually deals wi.th the activity
of only one neuron in his experiments, the question of organization of
activity of many neurons can be technically divided into two: first to
deternine organization of activity of single neurons in time and then,
after comparing the time organization of discharges of different neurons,
to obtain information about organization of neuronal activity in different
structures for the behavioral act. It could be of substantial help to
compare impulsation discharges of single neurons to the activity recorded
with a macroelectrode, since overall activity reflects processes of
interaction of many elements.
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There is a long history to the problem of correlation between impulsation
of single neurons and overall potentials (see, for example,
V. I. Gusel'nikov, 1975), and it is closely linked with the problem of -
electro genesis of summated activity.
The link between impulsation discharges and "spontaneous" summated poten-
tials turned out to be quite complex (Frost, A. Gol, 1966; Livanov, 1972;
Lebedev, Lutskiy, 1972; Elul, 1972). At the same time, it was shown that -
the oscillations of inembrane potentials of single neurons recorded intra-
cellularly correlated with macroactivity (Klee et al., 1965; Jasper, -
- Stefanis, 1965; Elul, 1964, 1972). Since changes in membrane potential
are related to entrance of synaptic influences in neurons, macroactivity
can be used to evaluate overall organization of synaptic influx in a
given structure as a function of time.
The reactions of single neurons to some stimulus or other were evaluated
in early studies only on the basis of impulse frequency, and they were
described as excitation and inhibition without consideration oP organiza- _
tion of impulsation in time. A more comprehensive analysis revealed that
neuronal reactions usually consist of alternating phases of activation -
and inhibition, which made it necessary to search for new criteria to
classify the entire neuronal reaction, reserving excitation and inhibition
only fo r evaluation of different phases.
A comparison of impulsation to evoked potentials opened up some utterly
new opportuni~ies for analysis of the time and space organization of
processes in the nervous system. Tt was found that the phases of neuronal
excitation and inhibtion often coincided with specific phases of EP
(Polyanskiy, 1965; Kondrat'yeva, 1967). At the present time, the link
= between single neuror.al discharges and some components or other of EP
has been demonstrated in virtually all parts of the brain, for example,
the retina (Fokiny Fomin, 1969), visual (Creutzfeldt et al., 1969), sensori- _
motor (Vasilevskiy, Soroko, 1970; Storozhuk, 1970) and other parts of the
cortex (Thompson et al., 1969), in the cerebellum (Bratus' et al., 1971),
_ hippocampus (Dubrovinskaya, 1971), activating structures (Shevchenko,
1975a), etc.
Since, as we strived to demonstrate in the preceding section, EP serves as
a correlate of general cerebral processes of organization of activity of
different elements into the functional system of the behavioral act, the
correlation between EP and impulsation of different neurons is of special
interest to us. For this reason, we shall discuss in detail the relation-
_ ship between EP and neuronal discharges.
Link Between Neuronal Activity and EP
_ When using anesthetized preparations, discharges of single neurons in
pro~ection regions, in relation to the stimulus, were demonstrable chiefly -
during the period of superficially positive EP oscillations. However,
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G. Fromm and G. Glass (1970) demonstrated that the form of correlation with
some EP component or other could be related to the constant cortical
potential, which changes with different doses of anestt?etic.
In experiments on waking animals, with the use of neutral stimuli, neuronal
discharges also appeared chiefly during superficially positive EP components,
which served as grounds to assume that there was inhibition of neurons ~
during negative EP waves. The few cells (2-3%} that presented a discharge
during negativity were interpreted as special inhibitory neurons. S. N. -
Khayutin demonstrated that, in the presence of natural, increased food
_ motivation and stimulation of the "hunger center" of the hypothalamus,
the number of neurons responding to neutral flashes of light with a dis-
charge in negativity ~f the evoked potential increased to 22% (1971, 1973).
He critized the conception of inhibitory pause and inhibitary neurons, -
and he concluded that the form of link between the neuronal response and
EP components was not fixed (Khayutin, 1973; Loseva et al:, 1970). ~
Neuronal discharges are observed in all EP phases, in response to stimuli
that trigger a given form of behavior (John, 1972; Shvyrkov, 1974 and
others), and the quantitative correlations between neurons presenting
discharges in the presence of different components of EP vary in different
~ behavioral acts and different structures (Aleksandrov, 1975; Shevch~nko, .
1975; Shevchenko, Aleksandrov, 1978).
We shall discuss the forms of correlations between impulsation and EP on
- the example of neurons of the somatosensory cortex with the use of electro-
cutaneous stimulation (ECS), which induces integral defensive behavior.
In our experiments, we examined the activity of 182 neurons of the somato-
sensory cortex on waking rabbits, stereotactically immobilized, with the
use of novocain alone. We used glass microelectrod es with tip diameter _
of about a micron, which were filled with 3 molar solution of KC1. EP
were derived from the surface of the somatosensory cortex with a silver
electrode immersed in agar, with which the trephination opening was filled.
ECS which consisted of square-wave pulses varying in duration and
intensity and delivered from a Physiovar stimulator by means
of needle electrodes, inserced subcutaneously in the assumed receptive _
field of the neuron under study, which was found in advance by testing
different parts of the body. -
We recorded impulsation and EP on tape, using the Ampex recorder, and
processed it on an A14096 analyzer. -
Under these experimental conditions, we observed the most diverse forms
of relations between neuronal discharges and EP components (figure 22);
_ and the same neuron could present discharges that coincided with several
or even all EP components. In other cases, a discharge appeared only -
during the anterior or posterior fron.t of one of the components (Figure 22). -
The phases of activation of single neurons could be more "divided" than the
EP components, but in general the pattern of activity of a single neuron
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could be described by the EP component:,, which coincide with the phases
of its activation. The discharges of only some neurons correspond to
specific EP components. In our experiments, 84 (46.4%) out of 182 neurons
presented discharges corresponding to some phase or o:.her of EP, and we
succeeded in demonstrating a primary response in 29 neurons (16%), dis-
charges during negativity in 45 units (24.8%) and late activation, which _
began during the late positive EP component or later, in 31 units (17.2~').
~ 30 _
24 LP 1Q ms Figure 22.
~0 Poststimulus histograms of 6
- neurons of the somatosensory cor-
tex as related to averaged evoked
Z~ potential (top)
LP 10 ms
� In these and all subsequent histo-
` grams the x-axis shows time in ms
- 30 an3 y-axis the nusber of impulses
Z~ LP 20 ms in the channel; n= 25, channel
!0 ~ width 5 ms. Averaging was done -
~ fm m the time of delivery of ECS;
30 - LP--latency period af the first
Zp ~ LP 30 ms phase of activation
IO I~ I y~ ~~~~~n~uf6Jl
30 LP 10 ms
20
!0 -
~-~1~1, wt ~ ~ia?~,~t
,O `~y11W~ ul~lll~~
50 I00 200 9pp ms -
_ �
_ In these same experiments, we examined the correlation between the pattern
_ of neuronal responses and ECS parameters, and we found that the neuronal
pattern could change entirely with different intensity and loca].ization
of stimulation. For this reason, the figures cited above characterize only
the number of neurons that served as the material of our study, and they do
not characterize activity of neurons of the somatosensory cortex in
any single behavioral act or ECS of specific parameters.
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Syzchronism and Similarity of Neuronal Discharge Patterns in Various Brain
Structures
The link between pattern-compone~~ts in a single neuren and EP components -
l~ads us to pose the question of correlation between time organization
of neuronal jischarges in different brain structures. This is a critical
question to reflex conceptions of t?:A ~behavioral act. Indeed, according
to reflex theory, the latency period of a~ehavioral reaction is defined
_ as the tine of conduction of excitation over the are of the corresponding
- ref lex. This conception is based chiefly on the idea of localization of
functions and the common sense of the temptingly understandable reflex
= scheme of stimulus--reaction. For example, processing of visual informa-
tion is viewed as the function of the visual analyzer, "from receptors to
the cortex," and the "output" of the visual analyzer then proceeds t~ _
motor structures that issue a"command" to actuating organs.
Thus, under the influence of a given conditioned or unconditioned stimulus,
excitation travels over a specific route, forcing some structures or
o*_her to perform their inherent functions. In this case, different
structures should discharge successively, and the time structure of neu-
ronal discharges in each structure should be related to the specific func- ~
= tion of this structure.
According to functional system theory, in the interval between st:imulus
_ and action there are processes of coordination of activity of different
elements into a single system. Of course, the coordination processes
- must be si~ilar and simultaneous in structures to be coordinated.
Quite a long time ago it was demonstrated that neurons of the same struc-
ture could response to stimuli of different modalities. At first, this -
property was believed to be specific for neurons of the reticular forma- ~
tion and other activatin~ structures on which the collaterals of specif ic
- or classical afferent pathways converge (Rossi, Zanchetti, 1960; Magoun,
1960}. Soon, however, "conzrargent properties" were aZso found in the
_ neurons of cortical projection regions (Jung et al., 1963; Buser, Imbert,
1964; Murata et al., 1965) and in all brain structures in general (Dubner,
_ 15G7; Bakl.avadzhan et al., 1971; Kazakov, Izmest'yev, 1972).
These data ~_ndicate that, after a stimulus, neurons situated in many parts
_ of the brain are activated somehow or other. A mere comparison of latency
~ periods of neuronal responses recorded in different structure after
arf erent stimuli shows tnat neurons of different structures can be
' stimulated simultaneously. For example, a click in an interval of up to -
, 30 ms elicits responses or alters activity not only of the auditory
analyzer, but neurons in other parts of the cortex (Voronin, Ezrokhi,
' 1973), hypothalamus (Baklava3zhan et al., 1971), hippoc2mpus
(Dubrovinskaya, 1971; Lidsky et al., 1974), cerebellum ~Khanbabyan, 1971),
a- and Y-motoneurons of the spinal cord (Buchwald et al., 1961), primary
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cutaneous afferents (Banno et al., 1972), optic nerve fibers (Spinelli,
M. Weingarten, 1966), etc.
J. Olds et al. (].972) obtained direct da*_a on time of activation of
neurons of different structures in the same behavioral act with the use
of numerous implanted electrodes. Tn these experiments, a click caused
_ rats to run to the feeder. A total of 64 different brain structures _
was examined. The experiments revealed that there are neurons that
fire in the interval of 0-20 ms. 4_n interval of 20 ms is the maximum
resolution capacity of the method.
Continuing these studies, J. Disterhoft and J. Olds (1972) demonstrated
that neurons that present discharges with the same latency period are
present in different structures in a different percentile ratio. In all
of these experiments, thick microelectrodes were used, the tip of which
was 62.5 um in diameter, which made it possible to describe the activity
of expressl y neuronal ensemliles, although one could also isolate thn ~
activity of single celis using a computer.
As we have already mentioned, in all structures the time organization of
discharges of single neurons corresponds to some components or other of
EP derived from this structure. Since the EP in different structures ~
become synchronous in response to stimuli that trigger a given form of
bet~avior, it is understandable that neuronal discharges with the same
latency period are demonstrable in many regions of the brain. E. R.
John et aI. (1969, 1972, 1974) also used the technique of recording
activity of neuronal ensembles, and they demonstrated that the activity of
ensembles is synchronous and similar in different structures. _
The objective of our studies was to coinpare the discharge patterns of
expressly single nPurons of different structures in the same behavioral
act. Experiments were conducted using the same method: rabbits immobi-
lized stereotactically, with anesthetization of the sites of fixation,
developed conditioned reflexes to a flash of light reinforced by ECS -
after 600 ms. We analyzed the evoked potentials and ~�ssponses of neurons
- in the visual and somatosensory regions of the cortex and reticular forma-
tion of the mesencephalon. Unlike the experiments of J. Olds and E. John,
in ours we used glass microelectrodes filled with 3 M KC1, with tip
diameter of about one micron, which enabled us to reliably isolate the
activity of expressly a single neuron.
Since our objective was to demonstrate the possible patterns of neuronal
discharges in the structures under study, ECS was delivered to different
points of the body surface so that we could assess the discharge patterns
in the cour se of various pretrigger integrations. . ~
Impulsation, EP and electrical activity of cervical muscles, which served
as a contro 1 of development of the conditioned reflex, were recorded on
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tape and processed on AI-256 and NTA-512 analyzers. The results, i.e.,
, averaged EP and poststimulus histograms of neuronal discharges, were
recorded on a two-coordinate automatic recorder, or photographed from
the analyzer oscilloscope.
We recorded 35 neurons in the visual cortex; 7 of them remained "areactive"
to light whatever the ECS parameters; 1 was always inhibited and 27
showed some pattern or other corresponding to EP. Since the pattern of
the response to light could change with changes in ECS parameters and the
discharges could correspond to several different EP components, we shall
classify the phases of neuronal activation independently of the number of
phases for a single neuron. -
We observed a discharge during the first EP component in response to light
with variois ECS parameters in seven neurons. In this phase, there were
usually only 1-3 impulses, which appeared with a latency period of 18 to
26 n~s. During the negative EP component, 5 neurons presented activation
with a latency period of 28 to 88 ms, and 26 neurons presented late
= activation with latency period of 100 to 500 ms. ~
Late activation was observed both in neurons that did not response or
- were inhibited during preceding phases (15 cells) and neurons that were
- activated in preceding phases.
In response to ECS, 3 out of 35 neurons showed a primary response; dis-
charges in negativity were observed in 13 cells; late activation was
found in 19 neu.rons, both among those that fired discharges in the
preceding EP phases (8 neurons) and those inhibited or that did not ~
respond during the early phases (11 neurons).
Data for the somatosensory cortex were obtained in experiments conducted
under the very same conditions also on 12 rabbits. We analyzed the
- activity of 83 neurons. Of this number, 33 cells responded to the
conditioned signal in accordance with the EP phases: we succeeded in
demonstrating a primary response with latency period of ~1-30 ms in 10
neurons, discharges in negativity (latency period of 30-86 ms) in 17 .
and late activation in 19, 6 of which did not present early phases.
~ Phasic activity was observed in 49 neurons out of 83 in response to ECS:
a primary response in 12, discharges in negativity in 29 and late activation
in 21, 8 of which did not present discharges in the early phases.
In analyzing the activity of neurons of the reticular formation, we com-
pared the phases of activation to EP derived from the surface of the
visual cortex, rather than reticular formation. Althoug'~ we also -
recorded EP from the reticular formation in these experiments, as the
microelectrode was introduced its position in relation to different
= tissular elements of the reticular formation changed, and this led to a
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change in configuration and even inversion of polarity of different EP
_ components.
The studies of A. Ramos et al. (1975, 1976) revealed that there could be
independent change in "focal EP" and impulsation derived with the same
microelectrode. This is probably related to the plastic geometric
localization of microelectrodes in tissue. When recording EP from the
cortical surface with a macroelectrode, the derivation conditions remain
constant, which enables us to compare the phases of activation of differ-
ent neurons, demonstrated even during different "passages" of the micro-
electrode, to the same components. _
We observed phasic reactions in response to a conditioned stimulus in 31
out of 68 neurons of the reticular formation. A,total of 20 cells fired
discharges~during the primary reaponse, 11 did so during EP negativity,
and late activation was demonstrated ir.~ 24 neurons, 8 of which did not
have early phases of activation. Phasic reactions to ECS were observed
in 41 neurons: primary resgonse in 25, discharges in negativity in 7 and
late activation in 26, 9 of the latter presenting only late activation.
A comparison of all these data leads us to conclude that, in all of the
structures examined, the time organization of neuronal discharge^ is
similar and that each of them contained neurons that presented identical
discharge patterns in response to a conditioned signal or ECS.
- Figure 23 illustrates poststimulus histograms of responses of neurons
_ of different structures to a conditioned signal, which contained the
- main components of the pattern.
The responses to ECS were also similar and had components corresponding to
- the phases of synchronous EP in different parts of the brain. For the
sake of comparison, Figure 24 illustrates the poststimulus histogram of
a reticular formation neuron, and Figure 25 illustrates poststimulus
histograms of a neuron of the somatosensory cortex, which show discharges
during the negative EP component, both in the conditioned and unconditioned
responses.
For direct comparison of time characteristics of neuronal activity in the -
visual and somatosensory cortex, we (with Yu. I. Aleksandrov) conducted
special experiments, in which we recorded neurorial activity in both
regions using two microelectrodes at the same time. The rest of the .
experimental conditions were analogous to the preceding ones.
We recorded 61 pairs of neurons: 48 neurons of the visual cortex and 53 of
the somatosensory. These proportions are attributable to t}ie fact that
we sometimes were able to record the activity of one neuror.~ in one
region "in a pair," with two or three successively demonstrable neurons in
another region. -
81
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SSC VC RF
~
VC ~
~ RF~
'0 10 ID
~ S 5 S
0 '~II'~-',..1.�'_-
0 4
VC ~
RF ~
~ !0
~ , 0^
I ~ ' ~
VC~ '
RF~
~~~~*1~ """T ~ r-'*~.w ~o .
10 5 S ~ -
5 ~ ~ ~
l~ ' � ~
0 "~f-T-'r- ~ , _
SO 150 Z50 350 50 150 Z30 350 60 150 230 350
Light Light Light
Figure 23. Identical types of neuronal responses to flash of
light, in somatosensory (SSC) and visual (VC) cortex,
and in mesencephalic reticular formation (RF).
Photograph of NTA-512B screen, n= 25; channel width 4 ms.
Averaged potentials in corresponding regions shown above the
histograms. T'he top row of histograms refers to neurons firing
discharges at the time of the primary EP response; the middle row .
is at the time of negativity and the bottom row, late activation
In some cases, we were able to directly observe neuronal discharges in
both regions that were synchronous and coinc~.ded~with the same EP components.
Figure 26 illustrates a vivid example of discharges of two neurons coincid-
ing with the primary components of EP, in response to both the conditioned
stimulus and ECS.
In this series, we tried to assess the order of involvement in activation
of neurons in different cortical regions after delivery of conditioned
signal. For this purpose, poststimulus histograms were plotted with
channel width of 2 and 4 ms. The latenc~ period of the neuronal response
was defined as the time between the stimulus and first maximum on the
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histogram. We analyzed neurons
with maximums within 100 ma after
the stimulus. We found 24 such
neurons in the visual cortex and
28 in the somatosensory cortex.
RF
25 Figure 27 illustrates histograms
!5 of distribution of neurons of the
5 visual and somatosensory cortex
according to latency periods. In
Light E S both regions, neurons become active
simultaneously and the maximums of
Figure 24. probability of their responses are
Responses of reticular formation in the range of 20 to 40 ms. These
neuron to conditioned stimulus probabilities constitute.0.58 for
(light) and electrocutaneous the visual cortex and 0.57 for the -
stimulation (ECS). Top: averaged EP somatosensory cortex.
of visual cortex and reticular for-
mation; bottom: poststimulus histo- In order to compare the dynamics
gram of reticular formation neuron of processes in these regions
responses. Channel width 3 ms; according to the parameter of
n= 25 (Slst to 75th combinations)y number of activated neurons, we -
650-ms interval between light and ECS calculated the latency periods
not only for the first, but all _
phases of activation of each neuron.
From these data we plotted histo-
~ grams of distribution of activation
phases of 34 neurons of the visual
imp� ' cortex and 40 of the somatosensory
~ Z4 , cortex (the rest of the neurons
Z~ presented no phasic activation).
!6 These "activity profiles" are
~~Z illustrated in Figure 28. Although
B they differ somewhat from one
4 another, due to the difference in
Z //2 22y 336 44B S60 1~Z 224 336 ms number of neurons in different '
C$ US~ regions that are active within a
Eigure 25. given interv~l, it is obvious that
- Poststimulus histogram of sornatosen- there is no question of any
sory cortical neuron firing dis- . successive [systematicJ involvement
charges in EP negativity. Bottom: of these regions.
histagram; channel width 4 ms; n= 25; ~us, after a stimulus that triggers
photograph from X=Y recorder. Arrows a behavioral act, the neurons of
show time of stimulation
each region fire discharges through-
out all phases of EP. There are
neurons that fire discharges synchronously in different structures at
any given time. -
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, i~,Y~`~"`~
2 ~ -
~
, ~
;~r~,-~+~.,.
~ -
4
I I.~l--l~-~
,
~ ~+~~I~ ~~1~ ~ ^r~l ~ ~
. ~fi' I :~~F,.
5
imp.
- a
imp.
b
~0
i ~ ~ ~ ~ ~ ,
CS /00 Z00 300 ~i!!0 500 US ~00
Figure 26. Synchronism of neuronal discharges in visual and
somatosensory cortex in response to conditioned light
flash (CS) and electrocutaneous reinforcement (US [unconditioned ~
stimulus]). Bottom: averaged EP and peristimulus histogram of
neurons of visual cortex (a) and somatosensory cortex (b). Channel
width 8 ms; n= 25 ~
1, 3) EEG of visual and samatosensory cortex, respectively _
2, 4) impulsation activity of neurons
5 ) EMG
6) mark.s of stimulation
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- n It is not always the same neurons
- 4 that fire synchronous discharges,
n.Zg � since the latency periods of
3 ~ activ3tion phases of the same
a Z neuron vary significantly, Figure
~ 29, tor example, illus.trates the
, dynamics of latency perioda of two
3 n�t~ simultaneously recorded neurons in
b Z the visual and somatosensory cortex
~ in response to a con~itioned signal.
~ When the stimulating electrodes
, were moved from the front leg to the
~ s ~ contralateral hir~d leg, the response
4 ; ~ patterns of these neurons to the
~ 3 ~ conditioned signal changed signifi-
Z ; , cantly, although the physical pro-
I ; ~ perties of the conditioned stimulus
~ ' remaiiied constant. This change in
20 ~io 60 BO !00 ms patterns was reflected in the change _
Figure 27. in dynamics of latency periods of
responses.
Histograms of neuronal distribution -
according to latency periods of re- Since there are neurons that are
actions to conditioned light stimulus. stimulated during any EP componenC
X-axis, latency period, ms; y-axis, in all regions, it is obvious that
number of neurons
- a) somatosensory cortex the discharges of some neuron in
b) visual cortex some part of the cortex are syn-
c) somatosensory (solid line) and chronous with discharges of some
visual (dash line) cortex neurons or other in other brain
structures. -
It appears to us that all of the above data warrant the conclusion that
there are simultaneously functioning neurons in all structures in the
.behavioral act. Although a different number o� neurons is activated in
each structure during different phases of EP, the overall time structure
of processes in different regions is identical, and it corresponds to the
time structure of "synchronous" or "general" EP.
We believe that synchronism and idei:tical nature of time organization of
both EP and neuronal discharges in diiferent structures rule out successive
performance of any functions by separate structures, and cannot conform
with conceptions of "conduction of stimulation" over the "reflex arc."
The neurons in each structure do not become active for a specific time -
as excitation advances from receptors to effectors, but they are involved
in all processes that participate in formation and implementation of the _
behavioral act.
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~p '
B
E ~
a ~
.2
6
b ~ '
2
CS 100 200 300 400 J00 jj~ !00 200 300 900 S00 600
Figure 28. Histograms of distribution of activation phases for
neurons of somatosensory and visual cortex, according
to latency periods of responses to conditioned light flash (CS) and
electrocutaneous stimulation (US). X-axis, latency period, ms;
y-axis, number of activation phases with the indicated latency period
a) in somatosensory cortex b) in visual cortex
It is also apparent that the similarity of EP configuration and neuronal
discharge p atterns in different structures precludea consicierati~n of EP
, configuration and neuronal discharge pattern as the expression of only
_ some specif~c function of a structure; the question of time pattern as a
means of co ding expressly specific information is also eliminated.
All of the submitted facts indicate, in our opinion, that processes in
different s tructures of tne brain acquire common features of organiza-
_ tion in performance of~a behavioral act and only during this act.
Since EP and neuronal discharge patterns corresponding to EP phases are
demonstrable locally, they reflect local physiological proceases; buC
since they are synchronous and common to different structures, it must be
- agreed that the same processes develop in many structures. Since both
physiological functions and links between various structures are different,
only process es of interacCion between elements of different structures can
have Che same dynamics. In other words, in the course of a behavioral
act, organization of physiological processes in time is the same for
different brain structures, and it is determined by tt~ie time structure
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of systemic processes of the behavioral act that are common to the entire
brain and organism, specific processes of organization of an integral
functional system out of special physiological mechanisms, rather than
the function of specific, for examp le, "visual," mechanisms in the visual
analyzer.
ms It is difficult to reconcile the
! 92~ a ~ fact that there is synchronism
Zy i and similarity of pattern configura-
~s tions in the responses of neurons
of different structures to the same
32 b stimulus, as well as the possibility
2~t of obtaining similar responses by
~s the same neuron to different sti-
iso Isz I,~4 ~ss ~S8 cos:'~ muli, with the analytical data
c' obtained on preparations as to
~Z~ dete~mination of the ~atency period
of a given discharge by a fixed
lOS number of synaptic arrests on the
way from a receptor to the recorded
74 neuron.
5B
Tndeed, the specific and constant
82 . d anatomical organization of links
66 in different parts ~f the brain
30 cannot explain the similar configu-
_ ,~y I ration of responses to the condi-
f tioned stimulus and ECS in, for
~B example, the somatosensory cortex,
~e0 ~ Z 1B~ 1B6 ,iBB where this has been described b
~ombinattons y
many authors (Shul'gina, 1967,
Figure 29. 1969; Vasilevskiy et al., 1972), or
Dynamics of latency periods of reac- similar patterns in reaponse to
tions of two simultaneously recorded the same conditioned stimulus by
neurons. X-axis, sequential number neurons of ~he visual and somato-
of combinations; y-axis, latency sensory cortex. Evidently, mor-
per~',ods of responses to 3.ight, ms. phological links rewain constant
Reinforcement ECS of 45 V was deli- and implement all types of time
' vered to the front (a, b) or hind organization of processes in all
(c, d) leg regions and with ali stimuli. How-
a, c) latency periods of responses ever, the link between the discharge
of somatosensory cortical of some neuron and arrival of
neurons to flash excitation over any specific path-
bt d) same for neurons of vieual ways can be established only on
cortex anesthetized preparations, in
which behavior is impossible.
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- Even the eariiest neuronal responses, with a latency period of 10-20 ms,
could be related to delivery to it of simultaneous influences from many
sources. The very possibility of such influences (for example, on neurons
of the somatosensory cortex) was demonatrated through etimulation of
various brain structures in anesthetized preparations (Storozhuk, 1974).
Some of these influences may be subliminal, with the use of anestheaia,
for onset of a spike response, while others may be so strong that they
induce EPSP [excitatory postsynaptic potentials] and spikes.
For example, stimulation of the amygdaloid complex of cats given nembutal
elicited a response in 11 out of 194 neurons of the somatosensory cortex,
in 5 of which an impulse appeared with a latency period of 1-2.2 ms,
and in another the "laten~y period of the response to stimulation of the
amygdaloid complex was even shorter than in response to stimulation of
VPL [expansion not known]: 2.2�0.2 and 2.4�0.17 ms, respectively, although
in both cases the response consisted of one impulse, and the probability -
of a response was the same, P= 1" [Storozhuk, 1974, p 149). In response -
to stimulation of the posterior hypothalamus, 15 out of 132 neurons res-
ponded, 4 of which had a latency period of 1.9-3.9 ms (p 150). Upon
stimulation of the pyramidal tract, orthodromal spikes appeared in 13 out
of 21 neurons, with latency periods of 2.6 to 7.5 ms.
The collateral influences from pyramidal cell axons could be addressed to
different neurons. In the opinion of V. M. Storozhuk, "in the case of
spontaneous activity this could cause a distinctive chain reaction of
dissemination of excitation in the somatic cortex" (p 151).
~ If we consider that fibers from the most diverse regions of the brain
come to the somatosensory cortex and that these structures have a"tonic"
� effect on neurons of the somatosensory cortex (Li, 1956; Tori et al., 1965),
even if stimulation Lhereof does not induce spikes in somatosensory
neurons, we arrive at the somewhat trite conclusion that the neurons of
the somatosensory cortex ar~~ under the inlfuence of all structures of the -
brain.
As we have already mentioned, a stimulus in the behavioral act induces
neuronal responses in many structures with short latency period and, in
particular, ECS induces responses with short latency periods by elements of
the optic nerve (Spinelli, Weingarten, 1966), reticular format3on
(Shevcher~ico, 1975a), hypothalamus (Bakl~,vadzhan et al., 1971), hippocampus
(Dubrovinskaya, 1971), etc. For this reason there are no grounds ta
questi~n the fact that, even with delivery of ECS that is "specifi^." for
the somatic analyzer, even the early responses of the pro~ection neurons
of the somatosensory cortex may be due to aynchronous comrergence of many
influences, rather than afferentiation over lemniscal pathways alone.
A. S. Batuyev er al. arrived at the same conclusion earlier, with reference
to the primary evoked potential: "The experiments convinced us that
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the primary response has a rather complex structure and contains components
that differ, not only in source in subcortical structures, but in distinc-
tions of their expression on the cortical level.... The main debatable ques- -
tion is whether the primary response is the result of one afferent volley in
a specific thalamocortical systew or whether it is the product of integra-
tion on cortical neurons of many afferent messages from various subcortical
structures, and we uphold the latter view" (1971, p 29).
These considerations are even more valid with regard to the early responses
of nonprojection regions and all late activations.
The presence of simultaneously discharging neurons in many structures and
the unquestionable links between them invalidate the question of how
expressly one structure affected another. From the point of view that we
are developing, all processes in all structures reflect interaction of
neurons in all structures, in accordance with the morphological links
between them. For example, while we know from morphological studies that -
_ the somatosensory cortex has direct communication with VPL, nonspecific
thalamic nuclei, reticular formation, other cortical regions, etc., it
is obvious that the discharges of neurons in the somatosensory cortex
are caused by influences that are carried over a certain number of fibers
from all these structures.
_ At the same t~me, the question of which exact fibers deliver influences -
to the somatosensory cortex, for example, from VPL or visual cortex, can
be answered on the level of VPL neurons or neurons of the visual cortex,
the activity of which, in turn, ~epends on all influences converging on
the VPL ar visual cortex neurons. Ultimately, each spike of each neuron -
is caused by the integr.ative activity of the entire brain. `
Thus, as applied to a waking organism, the explanation that any phase of
nesronal activation occurs by conduction of exeitation over some isolated _
pathways or chains of neurons is unjustifiably simple, since these chains,
by virtue of the specificity of morphological communications o~'each
neuron, must be specific and show quite diverse phases of activation in
different neuronv. It is only organization of processes in the entire
network of neurons that can cause similar patterns for different neurons -
that have a different place in this network. Consequently, it is only
organization of all processes that causes appearance of each spike.
It is known that quite a large number of synaptic influences should
reach a n euron simultaneously for at least one spike to appear (Kostyuk,
1971). The time and space organization of influences on a specific
neuron should, in turn, be a funcL-ion of coordination of integrative
processes occurring in many neurons. The appearance of phases of
neuronal reactions can apparently be attributed to the presence of sta- -
tistical maximums and minimums in the dynamics of number of selectively
- related and mutually coordinated elements. Expressly such dynamics of
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processes of coordination of activity of different elements must probably
be [he same and common to d ifferent parts of the brain involved in the
functional system of the behavioral act.
The fact that both neuronal discharges and EP, which are a reflection of
membrane potentials of many neurons, has the same time organization in
different structures indicates that different structures receive and
send influences simultaneously in the behavioral act. If we consider
the synaptic influx f rom dif ferent sources into each individual structure,
by virtue of synchronism of discharges in the other structures this
influx will have the same time organization as everywt~ere else, and it
will determine neuronal discharges also in accordance with the general time
structure of processes in all other parts of the brain. In this sense, we
can refer to systemic general cerebral time organization of synaptic
influx to any neuron. The similar EP in different structures are a
reflection of this general cerebral synaptic influx.
The identical time organization of processes in various structures indi-
cates that it is only the sp ecifi~ity of afferent and efferent coummunica-
tions that determines the difference in significance of spikes appearing
simultaneously in two different neurons in two different parts of the -
brain. According to functio nal system theory, any exoge:~ous stimulus
finds pretrigger integration ready, i.e., a dynamic system of interrelations
of elements prepared in advance, which is what determines the "spatial"
~:omposition of synaptic influx to each element.
Determination of Neuronal Discharge Pattern by Pretrigger Integration
When the activity of neurons was compared to EP in response to an insigni-
ficant stimulus, it was already found that the patterns of activity of
different neurons could chan ge (Kogan, Klepach, 1967). Moreover, in
some cases, the poststimulus histogram summated for many responses did
indeed refl.ect different patterns present in different responses, as
illustrated in Figure 30, taken Lrom the work of T. N. Loseva, S. N.
Khayutin and V. B. Shvyrkov (1970).
- E. John (1972, 1974) even expounded the view that neuronal responses are
always extremely variable, and that it is only the pattern of activity of
neuronal ensembles that does indeed have a constant correlation with EP.
However, the experiments of Ramos et al. (1976a, b, c) and Schwartz et al.
(1976) in the laboratory of E. John revealed that the response pattern
of the same neuron may even be more const~nt than the EP configuration
with the same behavior, and that it is demonstrable for several days.
The same neuron yielded diff erent patterns in response to the same stimulus ~
that triggered different behavior. The stability of neuronal responses
with stability of stimulus of behavior was also observed in the studies of
other authors (Olds, Hirano, 1969; Hirano et al., 1970; Phillips, Olds,
1969).
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, t~~''
- - , ~ , , _
- i ~
i
' ~ dfTr~"~ I I__1 1_ _ I ~ "
"'~T~
`~'-~"^~-^-~V 1 IV�~~/`V~`1~r~/~,^"`.v,- -
s ~ ~ ~J~~ Light
~
~ a _
d ~ i
~~,'1,.r~-~-
-----~--T--r- ;
'9 ~ ~ t sin ~
_ , ~--f-t--r-t-F-~~1-,~I�I-~~-T+
,,,~~i �~~t,,,�,
Mean number of im ises ~
A Z per ms g ~ .
1
u_ "~t,.~,_ ~ r,t~ _
_ SD w0 SOO ^ ms
~ ~-~---~r-
~ ~ ~
;--i-
_ ~ ---.r-~-- r~-r'~n~.-~.-
' ~~n~
ntilb'~~~,~~ti''.-_-.._'..r....,`_
6 i ~
~ ~ ----,--r-F+-~t~'a~fi~-~-tY;- , b Light -
~ . I
; I ~ , f i ~200 uV
VY
r #~----;-;;:~;r~!^I-- [1 mV
I I
Mean impulses
per Sp ~
Light i
B 2 I (
1 ~
-
Z~ I i ms
A+B ~E i
~~`n ~ -
.f0 7S0 SOr'I 1 I I ms
Figure 30. Reaction of one reuron of the visual cortex to exposure
of the retina to 9 successive 500-ms flashes. On each
strip: top line is superficial ECC [electrocorticogram]; the n~nbers -
_ on the left are sequential flash numbers.
A) average of 5 neuronal responses, including a primary resnonse and
late activation; inhibitory pause corresponds to slow negative oscill.
B) average of 4 responses of same neuron, where activation phase corres-
ponds to slow negative oscillation
A+B) averaging of all reactions of same neuron
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~ ~ ~T GF
~~~~'E1~ I~ 1~E~N~1~ i~1~~ QF ~EHA~' I~1~

~4 ,~UL~,'' ~HU"~~~~',~' ~ ~
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- Thus, the phases of activation of single neurons under different experi-
- menta.l conditions may be related to diverse ways to EP components. How-
_ ever, these relations are not formed at random, and they depend on the
- behavior triggered by the stimulus. As validly observed by M. N. Livanov,
- "one should probably think about participation of each neuron in the
formed system of integration, rather than initially inherent capacity of
neurons to react only to a given stimulus and only in a specific wa.y.
_ It seems correct to refer to nature of reaction of cortical neurons,
rahter than to types of such neurons" (1971, p 7).
.7ust like EP, the discnarge pattern of a single neuron is "endogenous" in
behavior, i.e., it can be quite different in response to the same stimulus
that triggers different behavior. This has already been demonstrated iri
, numerous experiments involving development o� conditioned reflexes, which
- showed that the responses of neurons in different parts of the brain to -
- a lioht flash, which became. a conditioned stimulus, changed in configura-
tion (Kondrat'yeva et al., 1970; Sviderskaya, 1971; Vasilevskiy, 1971,
_ and others).
At the same time, diff erent stimuli that trigger similar behavioral acts
can trigger similar discharge patterns in the same neuron, and this was -
- demonstrated in a comparison of patterns of conditioned and unconditioned
responses of neurons in defensP behavior (Vasilevskiy, 1973; Shul'gina, ~
= I968; Shvyrkov, Aleksandrov, 1973, and others). Many a.uth~rs have re-
ported that, with change in behavior, there is change mainly in the late
phases of the patcern of neuronal responses (Travis, Sparks, 1967; Ramos -
et al., 1976, and others). On the bas~s of analysis of activity of ~
neuronal ensembles, E. R. John and Morgades (1969) also believe that the
= early components of the pattem are "exogenous," unlike the "endogenous"
- late ones.
_ ~
According to our point of view about the conformity of EP with systemic ~
~ processes, the greater dependence of late phases on behaviox could be
attribu~ed to the fact that the late phases correspond to processes of ~
- implementation of one specific behavioral act, which changes with the
slightest change in exog:enous conditions. The early EP components reflect
: the transition to actuating mechanisms of one act from pretrigger integra- -
- tion, corresponding to all possible behavioral acts in a given situation;
f or this reason, the early phases are less related to one specific
' behavioral act, and they should change with change in a11 pretriggering
integration. ,
_ We tested this hypothesis in special experiments, which Yu. I. Aleksandrov
_ (1975) conducted on rabbits with developed conditioned defense reflex.
- The ECS parameters were changed to alter pzetrigger integration, since =
_ ECS diffe.ring in duration and force I.nduce, of course, different degrees
of deiensive motivation (Leander, 1973). ECS of different localization
- also has different ecological and behavioral slgnificance (Rozhanskiy,
1953; Menitskiy, Trubachev, 1974).
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- Experiments were conducted on 12 stereotactically immobilized rabbits, -
with anesthetization of fixation sites. The conditioned stimulus, a -
flash af light synchronized with a click delivered from a Soneclat
- stimulator (0.3 J, 50 ~ts), remained unchanged throughaut the experiment;
~CS delivered through needle electrodes from a Physiovar stimulator was
~ altered after every 25-75 combinations. The light was 600 ms away from
ECS, and interstimulus intervals ranged from 10 to 2 min. We recorded
impulsation of neurons of the visual cortex, EP and EMG, which served as
a control of development of the conditioned reflex, on magnetic t,ape,
and the data were processed on AI-256 and NTA-512-B analyzers. The
results (averaged EP ~nd poststimulus histograms of neuronal discharges) -
were recorded on a two-coordinate recorder or photographed from the
~ analyzer oscilloscope. -
. Of the 30 neurons of the visual cortex that presented phasic responses to -
the light o�r ECS, we succeeded at least once in altering the parameters
= of reinforcement in 15 neurons. In 9 of the latter, there was a change in
- pattern of responses to a light flash that was unchanged in physical -
parameters. In one neuron, a change in pattern was observed when the
- stimulating electrodes were moved over just a few centimeters. In others,
we succeeded in inducing such change only by changing significantly the _
intensity of ECS or moving the electrodes to another leg. The changes
, in pattern could consist of either disappearance of one uf the phases,
= or appearance of new components, and they were observed in neurons that
had differer.t "base patterns."
These data are listed in Table l, which also shows that, even in neurons
- whose activation phases did not change the response to change in ECS
parameters changed quantitatively, i.e., it contained more or fewer
spikes in the same phases.
Table 1. Modification of reactions of visual cortex neurons to condi-
tioned flash of light with chang~ in ECS parameters -
Type of response ~~ber of Qualitative and quantitative Changes in
neurons chan es in reactions atter.n
Primary activation 6 6 4
Negative activation 3 3 1
Late activation 5 5 4
,
It must be noted that even the primary phases of the pattern were related -
to ECS parameters. In three cases, the primary response present with the
= initial parameters disappeared and in another it appeared (Figure 31).
~ The responses of neurons of the visual cortex to ECS itself also changed
- with ci~ange in its par~meters (Figure 32). These changes were less marked,
- but they also involved all phases of the pattern, as can be seen in _
Table 2.
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1
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= � -
Table 2. Modification of reactions of neurons of visual cortex to ECS
with change in its parameters
_ Type of response N~ber of Qualitative and quantitative Changes in
neurans chan es in reactions attern
- Primary activation 1 1 1
Negative activation 9 4 1
Late activation 1 1 1
Figure 31.
a~" Appearance of primary response by
- imp. neuron of visual cortex to condi-
~ tioned light flash with increase
in intensity of ECS '
5
r
p~;x �,~~,t~d},~~_. t- j~, -f a) inhibition of neuronal activity ~
- in response to flash, reinforced I
by ECS of contralateral front ~
paw, 30 V ;
b) appearance of primary response
b to conditioned flash with in-
crease in ECS to 60 V,
Above (a) and (b): averaged EP;
below: peristimulus histogram of '
= 5~ impulsation activity. Channel
_ p ~-~t"�=`,~..~''a`L~`i
�r"i~''='_=-''?--`:~,+' width 4 ms, n= 25
_ ~00 ~00 I S00 CS--conditioned stimulus
CS u~ US--unconrlitioned stimulus
These data indicate that all phases of the pattern can change with a change '
in pretrigger integration. Since primary discharges appear in different
neurons in response t~ the same conditioned signal that triggers different
preliminary integrations, it may be assumed that even in regions that
_ are projections in relation to the stimulus the response is "endogenous," ,
i.e., it corresponds to activity of expressly the elements that were ~
included in advance in pretriggering integration. Of caurse, "endogeny" of
the primary response in projection regions could not be demonstrated -
when recording only macroactivity. -
Comparable results were also obtained for neurons of the somatosensory
cortex (Shvyrkov, 1974) and reticular formation (Shevchenko, 1975a).
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Figure 32.
Disappearance of visual neuron res-
- ponse to conditioned light and ECS
~ with change iz localizatior of
a
- stimulating electrodes.
imp.
~a 1) averaged EP
' f~ ~ 2) peristimulus histogram; channel
Z p . ' o width 2 ms, n= 25
CS 'T- U a) late activation in response to
light and activa*_ion in
- ~ ne~ativity with affierdischarges -
in response to ECS of contra-
b � lateral hind paw, 30 V ~
l~P' b) disappearance of reactions with
_ Z o~ .~~ty;;t~, ;~y,~~~ stimulation of ipsilateral front
= CS ~oa ao~
3vs yao sao US;o~ en:~ ms Paw with same intensity of
current
Tt~zre was equally graphic demonstration of the dependence of pattern of
activity of vis~ial neurons in response to light on pretrigger integration
in experiments where a change in preliminary integration was produczd by
- changing reinforcement from food to defensive (Shvyrkova and Shvyrkov,
1974; Shevchenko, 1976). We shall consider these findings in connection
~
with other questions.
_ Involvement of Neurons in Systemic Mechanisms of the Behavioral Act
Cortical neurons: For a long time, ~he activity of neurons of different ~
- structures of the brain was traditionally analyzed solely as a parameter
of the specific function of a given structure. It appeared quite logical
to relate all types of activity (for example, of neurons of the visual
cortex) to the parameters of a visual stimulus and processing of informa-
tian about this stimulus. Hoaever, the very first data obtained on waking _
animals in learning studies revealed that the activity of visual cartex
_ neurons :.hanges in relation to chan~ge in beha~;ior (Ricci et al., 1957; .
- Jasper et al., 1962). The same experiments revealed that neuronal activity -
in other structures, in particular, somatosensory, motor and frontal cortex,
depended on behavior. -
Subsequently, all researchers concerned with development of conditioned
reflexes observed a change in neuronal activity in response to a stimulus
that had become conditioned (Vasilevskiy, 1968; Shul'gina, 1967, 196A;
- Rabinovich, 1975, 1976, and others).
At the present time, because of development of techniques for recording `
neuronal activity of freely behaving animals, an increasing number of
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- works is being published demonstrating a direct link between activity
of neurons of all brain structures and specific stages of integral behavior,
rather than a given stimulus. For example, V. Mountcastle et al. (1975),
wno studied neuronal activity of the associative cortex demonstrated a
link between the discharges of these neurons and situational d~stinctions
that cannot be formuiated in other than terms of and objectives of
behavior.
An entire series of studies conducted by H. Niki (1974a, b) demonstrated
a relationship between neuronal activity in the prefrontal cortex and
specific stages of behavior. _
I.Ranck (1973), who studied hippocampal neurons, found that it was possible, -
_ in general, to compare their activity to only specific behavioral acts.
He calls this approach "microphrenology." . Hawever, we believe that
the ter~ "microethology" would be more suitable, since we are dealing
here with the link between activity of specific neurons and specific
behavior, rather than specific behavior and activity of a localized
_ region of the brain. ~
The experiments of J. Olds et al. (1969a, b; 1972) ar.d E. R. John (1969,
1972, 1974) showed that a link with specific behavior is observed for
neuronal activity in many structures of the brain.
A link with expressly behavior was noted by many authors; for neurons of
the reticular formation (Sparks, Travis, 1968; Travis, Sparks, 1967), as
well as neurons of cortical projection regions (Shvyrkova,Shvyrkov, 1975;
Andrianov, Fadeyev, 1976; Miller et al., 1972, 1974). J. Mi1J_er et al.
_ (1972) maintain that "cellular activity in sensory systems is strictly
determined by the behavioral situation and objective" (1972).
- All of the cited data, as well as our experiments that demonstrated a ~ ;
correlation between neuronal activity and parameters of future reinforce- '
ment, confirm the validity of the thesis of functional system theory, to
the effect that "elements rererable to some anatomical system or. other are
_ involved in the system of a behavioral act only to the extent that they
aid in achieving a preplanned result" (Anokhin, 1973a, p 35).
Indeed, each neuron participates in the behavioral act in the form of ~
impulses which, as they spread over axonal collaterals, have a specific I
influence on all elements linked with this neuron and, ultimately, the ~
entire system. The presence of "superfluous" impulses, as well as
absence of "necessary" ones woulci create a discordance in the system '
and~make it difficult t.o achieve the useful adaptive result. In this
sense, any impulses have actuating output functions.
According to functional system theory, impulses can only appear in neurons
whose past activity led to achieveme.nt of a specific result; in this sense,
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we can speak of goal-directed activity of each element in the functional
system of the behavioral act. The link with behavior can be explained,
from the point of view we are developing, by the fact that neuronal
activity is not a"reaction" to a"stimulus," but is retrieved fram
memory, as it is necessary to achieve the result of the entire behavioral
act.
- Of course, this is possible only when the model of the result is already
represented in pretriggering integration.
- The link between activity of single neurons and the entire behavioral act
= enables us to raise the question of involvement of nFnrons in specific _
= systemic mechanisms of the behavioral act. At the same time, this is a
_ question of how pretriggering integration is implemented in the goal-
= directed activity of many different neurons.
- Since, as we tried to prove in Chapter 2, EP components serve as correlates
of specific systemic proces~e~ in the behavioral act, and the discharges
of single neurons coincide.. in time with some EP components or other and -
have the same properties, the next hypothesis logically arises. Neuronal =
discharges corresponding to the primary EP component reflect processes
of implementation of the part of preliminary integration that is the
most "prepared" for the triggering stimulus; the discharges during EP
negativity must correspond to processes of afferent synthesis and decision
making, i.e., total replacement of prior organized activity of the
� integral organism by coordinated activity corresponding to the next
- behavioral act. The discharges during the positive EP component must _
correspond to processes of mobilization of organizerl actuating mechanisms =
= and, finally, late act~va*_ion should correspond to actual implementation `
of the acceptor of acti~~n results and program o� action, i.e., coordinated =
purposeful activity of ele-merts that were unite,; in prior processes into
a single functional system of the behavioral act. At the same time,
there must be "further maturation" of pretrigger integration of the next
behavioral act directed toward achievement of ~he next goal in the entire _
hierarchy of behavioral acts that ultimately result in sarvival of the
organism.
Since the same neuron can be activated in different EP phases, it must be
assumed that the same element can participate in several or even all systemfc
processes of the behavioral act. Tk?e.fact .that�a neuron can present phasic
activation in different behavioral acts suggests that the same neuron may _
be involved in different integrations. Since neuronal function, i.e., its
~ possible: contribution to achievement of the result of a behavioral act, is
determine~d exclusively by the topography of its axonal collaterals,
diverse integrations can be related only to the disersity of sets of
neurons included in a given integration.
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We tested all these hypotheses in collaboration with Yu. V. Grinchenko
b~ means of special experiments. We used the same method as for the study
= of evoked patentials in the continuum of behavior. The modification
was that the rabbit was trained to pull on a ring upon delivery of the
� conditioned signal of a click. The click triggered approach to the ring 1
and pulling on it; the light flash stopped the pulling and triggered
movement to the feeder.
~
In these experiments, in addition to recording of EP in the visual cortex
by means of a silver electro~e immersed in agar, which filled a trephina-
tion opening, we recorded the impulsations of 68 neurons of the visual
cortex. These recordings were made using a merhod developed in aur
laboratory (Grinchenko, Shvyrkov, 1974). We used a micromanipulator
attached to the rabbit's skull; a field transistor operating in the mode
of a source follower and connected in the circuit (Rosetto, Vandercar,
1972) was also placed on the skull. This technique enabled us to record,
with virtually no artefacts, the activity of single~neurons in the course
of several behavioral cycles and, occasionally, several hours.
After amplification with a UBP 1-02 and Biophase, the EEG of the visual
cortex, impulsations, EMG of cervical muscles and ma.rks.for sound, pulling,
light and presentation of feeder were recorded on magnetic tape of a 14-
. channel magnettor. These tapes were then reproduced on paper using an
automatic ink recorder, with reduction of feed rate to one-eighth.
= Impulsation was processed by the histogram method, and 50% deviations
from the background were considered as a change in activity. We selected
- 39 of the 68 neurons of the visual cortex under observation in these .
experiments for comprehensive analysis, am~ng those demonstrated in at
least five behavioral cycles. Of these 39, 7 neurons did not change their
- activity in any of the phases of the behavioral cycle. The change in
activity of the other 32 neurons coincided with some behavioral act which,
in accordance with functional system theory, we singled out as a segment -
of the behavioral continuum from one result to another: from the click to
appearance of light, and from the light to the feeder. In the firsti
behavioral act, we observed a change in activity of 20 neurons, 7 of
= which contained activation phases in their response and 13 only inhibi-
tion. In the second behavioral act, activation was demonstrated in the
responses of 18 neurons and 12 cells were inhibited; in all 30 neurons
showed a change in activity in the second act.
As in the preceding experiments, we found that neuronal activity in the
visual cortex is obser~ed in all time intervals of the behavioral act
(Figure 33). The early (up to 200 ms) phases of activation were
clearly related in time to the prior stimulus--result (Figure 34), whereas
the late ones appeared with a variable latency period. This "late"
activation was observed in 4 neurons in the first behavioral act and 6 in
� the second. They were demonstrable much better by the method of "reverse"
averaging, i.e., by plotting the "preresult histograms" (Figure 35). ~
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Expressly these activations are related to the actuating mechanisms of
the bchavioral act directed toward achievement flf a specific result.
_ Appearance of "stimulus--result" stops these activations (Figure 35).
In our rabbits, we occasionally observed intersignal tugging at the ring,
i.e., tugging that started without the click and, of course, did not
lead to appearance of the flash. In su~~h cases, the link between late -
activations and a specific behavior was also manifested in the abse:nce -
of a stimulus (Figure 35), which does not warrant consideration of
these activations as reactions to some factor. Actually, expressly these
late activations reflect the coordinated purposeiul activit}- of elements
of the inte.gral organism in a specific behavioral act.
A ~omparison of late activation of nesrons of the visual cortex in two
successive behavioral acts revealed that two neurons participated in
_ both behavioral acts and the remaining eight in only one of them. Five
- neurons that showed late activation only in the second act were not
reactive or inhibited in the first (FigurF:, 36~, while three cells, which
- showed late activation in the first act ~~~ere areactive or inhibited in~
the second, but two of them presented prim
Z
3
b
~
5
5 ~mP L
50.ms Light ~ECS
Figure 8~. Depression of background activity and all phases of
neuronal activation under the influence of GABA
1) EEG of somatosensory cortex 5) poststimulus histograms, channel
2) neuronogram width 24 ms, n= 25
3) EMG a) before GABA
4) averaged EP (n = 25) b) against the background of GABA '
Analysis of sensitivity to the agents used of different elements of the
pattern only enable us to state that in each phase the functional synaptic
fields change under the influence of several agents. Thus, of the 9 neurons
which demonstrated a primary response, 7 showed a change after phoresis of
not only one, but two or even three agents which changed background activity
differently (Figure 82). The primary response of different neurons was
sensitive to different agents. Since synpatic influences are mediated by -
different mediators and changes in FSF are specific for different agents
administered phoretically, the change in the primary response under the
influence of different a gents is indicative of the multimediator nature
of synaptic activity at the time of the primary response.
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a . . .
a '
..~v"1r~--~--
imn
JO ~
b 5
~ b _
c
_ 5
l~1
d c
- 5
. ECS L~p ECS 9BI4~290 ms
~ 10 ms �
Figure 84. Figure 85.
Comparative effects of atropine, GABA Appearance of new phase in neuronal
and glutamate on ECS-evoked neuronal response with ionophoretic adminis-
re~ponse; n= 15, on histograms the tration of acetylcholine
channel width is 24 ms a) initial nauronal reaction--
a) initial neuronal reaction (primary primary response
response and late activation) b) the primary response is
b) elimination of response by atro- eliminated by acetylcholine,
pine (+25 nA, 8 min) but there is appearance of a
c) inhibition of response by GABA phase of late activation
(+10 nA, 7 min) c) control series of combinations, -
d) elimination of initial pliases
and appearance of new (negative) n= 10, channel width 24 ms '
phase with L-glutamate (-15 nA, 8 min)
If we were to concur with r_he popular view (Orlov, 1974; Sherstnev, 1971; ;
Marsden, 1973; Spehlman et al., 1974a, b) that neurons of different brain
structures produce and use for synaptic transmission different mediators,
it is logical to conlcude that not only the primary response of the entire
set of neurons but, possibly, the.primary response of a single unit is
generated upon convergence on it of stimuli from many different sources.
We had arrived at this c~nclusion earlier on the basis of other data. The
same applies to the discharges in negativity, as well as the phase of late -
activation.
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The fact that different phases of the pattern can differ in sensitivity to
the same agent is indicative of differences in neuronal SFS in the course
of various systemic processes according to the neurochemical criterion also.
Since a change in the state of a neuron under the influence of some agent
does not lead to a chaotic reaction, but to elimination or appearance of
_ additional phases, but always phases corresponding to some components of
EP, it can be concluded ~hat the state of the nevron per se does not lead
_ to spike generation, bui influences the output pattern merely by selectively -
alte~:�ing the effectiveness of synapses making up the dynamic functional
- synaptic field, whereas initiation of phases is implemenL�ed by the phasic
synaptic influx.
As we know, synaptic influences on a neuron induce two sorts of effects:
"integrator," which are related to a change in state of the neuron,
and "detonator," which induce spike generation. According to P. Andersen
and T. Lomo (1967), integrating influences go to the distal parts of den-
drites, while detonator ones come closer to the soma. This implies that
there is morphological. fixation of integrator and detonator links vetween
neurons. P. G. Kostyuk (1974) believes that tonic and phasic influences
are distinguished exclusively by their functional and dynamic features,
and there can be both detonator and integrator influences from different
sources on a given neuron.
Since influences that elicit a neuronal discharge also alter the state of
the neuron, like the iniluences that do not elicit a discharge, we believe
there is more justification for making a distinction between effects,
rather than influences, on the assu~ption that any influence perceived by
a neuron will have an integratxve effec~, while generation of spikes would
depend on the entire ~et of influences and other conditions.
. _ - -
Under normal conditions, a given state of a neuron and selective effective-
ness or "detonatory nature" of specific synpases are apparently related
to corresponding integrator effects of all influences on a single neuron by
_ other elements of the entire integration. In our experiments, the selective
effectivenessas detonators of only some synaptic inputs and, consequently,
the pati:ern of a neuronal response could be determined by constant
integrator influences created by :he motivation of fear and defense activa-
tion, which always appear in a situ~ztion of delivery of nociceptive stimuli.
Since motivational influences are se~.ective and elicit only neuronal
~ states that led to acheivement of an a~daptive result in the entire system
_ in prior experience (Anokhin, 1968, 1974a; Sudakov, 1971), only the syn-
- apses whose activation would lead to purpos2ful activation of a neuron,
- aiding in achievement of this adaptive result, turn out to be effective.
Al,though numerous exogenous influences converge on a neuron with electro-.
' cutaneous stimulation, a real pattern appears as the result of selective
activation of the neuron only through specific effective synapes, which
~ 183
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is what pred`termines the appearance of a"purposeful" pattern in the reac-
tion to current synaptic activation and involvement of the neuron in some -
systemic pro~_esses of the behavioral act. Thus, the neuron does not emerge -
as a summator, but as an "organizer" of influences coning to it: from the
- organization of incoming influences, which does not conform well with the
goal, it creates an organization of discharges in time that conforms
more fully with the goal.
There are two points of view that can be discussed to explain the inter-
action of different synaptic influences on a single neuron. According to
one of them, the only form of interaction of synaptic influences is their
summation on the neuronal membrane. According to the other view, integra-
tive neuronal activity is not limited to summation of inembrane potentials:
some synaptic processes induce specific chemical changes in subsynaptic
regions that are integrated in a change in metabolism of the entire neuron,
and through the metabolic change they have a specific influence on the
effectiveness of other syn~aptic inputs using different mediators.
This intersynaptic integration, which is a reflection of all interneuronal
integration ~n the level of a single neuron, lea~s in turn to intermolecular
integration, which is the object of fixation in molecular mechanisms of
memory (Anokhin, 1974; Matthies, 1973, 1974; Huttunen, 1973). _
Phoretic administraticn of agents, which blocks or alleviates neuronal
activation with regard to some inputs, of course elicits a very complex
change in the synaptic input, which could also be related to different pre-
synaptic effects and some of its influences on adjacent elements. Never-
theless, phoretically administered agents, such as glutamate and GABA,
always alter background activity unequivocally, which can be interpreted
as an indicator of change in neuronal excitability, regardless of which ~
specific mechanism is involved in obtaining this ch3nge. If ue accept
the summation hypothesis, we must conclude that changes in the background
and reaction must proceed in the same direction, and this is what is usually
observed when the neuronal membrane potential is changed by means of
polarization (Kabur~eyeva, 1971). However, as noted by many authors,
when biologically active substances are delivered to a neuron the change in
background activity often fails to be correlated with the change in reac-
tion (Kozhechkin, Zhadina, 1973; Schmidt et al., 1974; Hess, Murata, Y974).
zn our e:cperiments, glutamate, which depolarized the membrane (Krn~evic,
1970; Bernardi et al., 1972), increased background activity, but could
eliminate the entire response or one of the phases of the response,
while GABA, which hyperpolarized the membrane (Krn~evic, 1970; Altman et
al., 1973) and blocked background activity almost entirely, could
- alleviate significantly the neuron's evoked reaction (Figure 86). At the
same time, different agents, even those that changed background activity
in the same way, c~uld alter very differently the patterr? of neuronal
reaction. The difference in directions of changes in background activity
and the evoked reaction, as well as different phases of neuronal reaction
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under the influence of the same agent cannot, in ou~ opinion, be attributed
to simple summation of inembrane potentials evoked by phoretic and natural
synaptic activation.
- a b ~
� ,
imp imp imp
I5 /5 15
!0 IO IO
S S 5
' ECS ECS ECSs~~
Figure 86. Comparative effects of L-glutamate and GABA on neuronal
response evoked by ECS
a) initial neuronal response contains a negative phase
b} with L-glutamace an additional phase of late activation is added
to the initial phase
, c) with GABA, the initial reaction is alleviated, while background
activity is depressed. On the histograms, channel width is 24 ms,
n=20
Within the framework of the summation hypothesis, this difference in direc-
tions could be explained by means of additional hypotheses to the effect -
that different components of the synaptic influh change differently under
~~he infliaence of the same agent. However, these hypotheses appear con- -
trived to us, since the absence of reaction change under the influence
_ of glutamate, which depolar:izes the membrane and izicreases background
activity in all cases,uAUld have to be attributed to the fact that the
heightened excitability of the neuron is "compensated" proportionately
to the decrease in synaptic influx, whereas blocking of the reaction (as,
for example, ~n Figure 82) would have to be related to blocking of only
one group of synapses, and one that is not isolated by the morphological
~ criterion, but the functional time criterion. Analogously, under the
influence of GABA, for example, in the case i].lustrated in Figures 86 and
87, we would have to assume that there is unexplainable intensification _
of synaptic influx under the influence of a universal inhibitory agent,
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and this intelsification of synaptic influx is so great that it overcomes
even a significant decrease in excitability of the neuron. "
a
b ~
~
c --~".~/1/."V`'�,'
I
ECS E S
Figure 87. Effect of GABA on neuronal responses as a function of
parameters of ECS. -
Left--responses of ne~iron to ECS of contralateral front foot, 15 V, 1 ms:
~ a) initial neuronal r.:sponse consists of negative phase and phase
of late activation
_ b) GABA (+6 nA, 14 min, which depresses background activity, alleviates
negative phase and depresses phase of late activation -
c) control series
Right--responses of neuron to ECS of contralateral hind foot, 30 V, 1 ms
a) 3.~itial neuronal response consists of negative phase
b)"GABA (+6 nA, 14 min), which depresses background activity, also
depresses negative phase of response '
c) control series of combinations
- Channel width 24 ms, n= 20
\
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These hypotheses turn out to be unnecessary, if we assume that constant
phoretic activation or blocking of some synaptic inputs leads, in addition
to change in membrane potential, transsynaptically to an increase in
effectiveness of some synapes and concurrent decrea~e in effectiveness of
others, which are activated immediately after the stimulus.
_ Since different synaptic inputs can be implemented by various mediators
(Orlov, 1974; Sakharov, 1974), the difference in direction of the effect
of the same agent on effectiveness of these inputs could be attributed to
a change in neuronal sensitivity to different mediators. According to
the integrative hypothesis, this selective change in sensitivity must be
related to a change in general intraneuronal metabolism induced by the
inf luence of a phoretically delivered agent.
- Since different agents can change the pattern of neuronal reaction in
different ways, it can be concluded that the change in metabolism induced
by phoretic application of an agent is rather specif ic, i.e., that there
is a specific link between activation of a certain functional synaptic
fieZd, metabolic changes within a neuron and new functional synaptic field, _
i.e., new organization of effective synapses. "
This specific conformity is~apparently determined by molecular processes
in neuronal protoplasm and constitutes the "substrate of neuronal memory."
- Thus, the integrative state of a neuron is mediated by neurochemical
- mechanisms. The general hypothetical scheme of correlation of neuro-
- physiological processes on the level of a single neuron and of neuro-
chemical processes is conceived as follows: when a neuron is involved in -
pretriggeri~ig integration, different integrative synpatic influences _
induce specific chemical changes in subsynaptic regions, which are integ- _
rated into a change in metabolism of the entire neuron, and through the
c,hange in metabolism they have specific influences on the effectiveness of
synapses that use different mediators. Probably, the neuron "recognizes"
a specific int~bration of synaptic influx as organization of ined.iators.
- At the present time, there are already several hypotheses concerning the
link between synaptic activation, molecular processes in neuronal proto-
plasm and impulse output of the neuron (Anokhin, 1974; Matthies, 1973,
- 1975; Matthies, 1974). The concrete intermolecular mechani.sms of integra-
tion are beyond the area of competence of the neurophysiologist. We only
have to stress the fact that, on all levels of thishierarchy,selection of
different mechanisms into the functional system of an integral behavioral
act occurs in accordance with a single evolutionary principle, namely
the criterion of their cooperation in achievement of a useful adaptive
result and, ultimately, survival of the organism. This entire hierarchy of
inte~rations, c~hich could be continued in both directions (in the direction
of "molecular memory" and in the direction of "memory of the arganism"),
is established during formation of the functional system of a behavioral
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act during trial behavioral acts, and it is fixed by the useful adaptive
result.
In accordance with the general direction of integrative activity of the
nervous system toward reduction of "degrees of freedom" and selection of
one behavioral act out of the many possible ones, integrative activity of
a neuron consists of reducing the "degrees of freedom" referable to time
of appearance of discharges and choosing one pattern of responses out of
the many possible ones (Anokhin, 1974). On the basis of the facts w~ have ~
submitted and data in the literature, it can be assumed that reduction of
"degrees of f~eedom" of neuronal discharges is achieved on the basis of
a general "principle of conformity." This principle is already manifested
an the periphery, and it consists of the fact that neuronal responses
appear only when there is conformity of stimulus properties with the pro-
perties of the peripheral receptive field. In the experiments described
above, this principle was manifested by the fact that, although many
synaptic influences converge on the neuron after ECS, a real pattern
appears as the result of conformity between "endogenously'(through meta-
bolic mechanismsl effective synapses with those that are really activated.
As a result of all this, neurons, whose set of elementary functions
corresponds to the goal and real info~ation, become involved in the func-
tional system, and behavioral acts are retrieved from inemory that corres-
pond to motivation and situation.
Since motivational influences that determine pretriggering integration are
selective and induce only neuronal states that led to achievement of
a given adaptive result in prior experience, on].y the synapses and FSF
whose activation would lead to purposeful neuronal activity corresponding
to achie~aement of one of the adaptive results are endogenously effective.
Additional reduction of degrees of freedom of the neuron is related to
the influence of numerous situational afferentations through which the
change in all of pretriggering integration narrows even more the area of
neuronal FSF.
Thus, the role of pretriggering integration in generation of a purposeful
neuronal pattern consists of reducing the degrees ~f freedom of the neuron
- by means of formation of funetional synaptic fields out of selectively
effective synapses, the use of which in prior experience had already led
to a useful adaptive result in a given situation.
Correlation Between Functional Synaptic Fields in Pretriggering Integration
The final choice of one degree of freedom in the behavioral act, i.e., the
final organization of one system of orderly and purposeful relations, among
all neuronal interactions that are possi hle due to divergence and conver-
gence of their axonal collaterals within FSF selected in pretriggering -
- integration, is established only after making a decision, when the
actuating mechanisms of the behavioral act begin to function.
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According to the conceptions ;ae are
- ' ` developing, auring pretriggering -
integration the FSF corresponding
2 . to future events related to the
given motivation and situation
should be effective.
a We tested this hypothesis by com- -
paring neuronal activity in cozdi-
tioned and unconditioned behavioral
~ acts wi*_h phoretic delivery of
different agents. The logic of
1 such comparison consists of the
. ~~I~~ ~���~.~��+~r�~ following: as we have established
~ ~ in the preceding experiments,
b neuranal activity in a conditioned
_ 2 behavioral act is determined by a
model of a specific future event--
electrocutaneous stimulation with
specific parameters; the activity
in an unconditioned behavioral act -
- is determined by a model of some
other future event, apparently, -
disc~ntinuation of the nociceptive ~
effect of ECS. These two goals
are hierarchically related, the
Light ~CSi~~ latter being "larger" in the entire
50 ms hierarchy of goals constituting
rigure 88. defense motivation.
Effect of glutamate on identical two- By comparing the effects of the
phase patterns of neuronal responses same agent delivered constantly on -
_ evoked by conditioned and uncondi- neuronal activity in the two be-
tioned stimuli. Channel width 24 ms, havioral acts, we hoped to separate
n = 25
a) initial res~onses of neuron are FSF corresponding to different
repreaented by negative phase and hierarchically organized goals.
� phase of late activation Of the total of 70 neurons, 21 res-
b) glutamate alleviates negative ponded to conditioned light.
phases and depresses phases of Delivery of some agent to eight
late activation neurons that presented identi.cal
. 1) neuronogram
2) averaged EP patterns in the two acts altered -
3) poststimulus histogram these patterns in six cases, in
the same direction (Figure 88). In
two cases, these patterns changed
independently: with delivery of the same agent, neuronal activity was
depressed in the first act and enriched by additional components in the
second one (Figure 89). There was also independent change in reaction
in the two behavioral acts in seven neurons that origina.lly presented -
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~ ~ ~T ~F
,
~~'~TE~t I ~ ~tE~H~I~ I ~F ~EH~~ I
,~l~L~ ~H~~'~1~~'~ ~ ~F ~

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- _ different patterns (Figure 90); '
a the changes in reactions to light
- and ECS were induced by different -
_ imp agents. Conditioned responses ~
10 appeared in 5 out of 14 neurons -
~ that had responded only in the =
_ ~ second act af ter delivery of the _
agent (Figure 91). In six neurons
= that presented ph~sic responses -
- only to the conditioned stimulus
_ b and did not react or were inhibited
_ witn ECS, no reaction to the un-
- ~-mP conditioned stimulus appeared -
- under the inf luence of application
_ of the agent, although responses
5 to the conditioned stimulus could
- change (Figure 92). Delivery of -
- agents to 35 neurons that were
- Light ECS inhibited or did not react in both
SOuis acts elicited a response to the
Figure 89. conditioned stimulus by only one
Different changes in patterns of neu- neur~n and to an unconditioned
' ronal responses to light and ECS under stimulus by another onF..
- the influence of glutamate. Channel -
~ width 24 ms, n= 15 In 12 neurons, we succee~ed in
- a) initially, neuron responded with altering the parameters of ECS in -
:
_ negative phases t~ light f.iash at least one instance. In seven =
_ and ECS of tnem, this elicited a change in
b) L-glutamate eliminated negative pattern of reaction, not only to
phases in both responses and ECS, as yhown in Figure 87~, but to `
"created" a primary response and the conditioned stimulus, and this _
= late activation in response to was also associated with a change -
ECS in effects of phoretically delivered -
agents (Figures 93 and 94). Inter-
- estingly enough, with change in ECS parameters, the patterns of conditioned -
reactions and sensitivity to different agents also changed in three neurons
that did not react to ECS of any parameters before (Figure 94).
; Let us discuss these findings from the vantage point of the qiiestions we
have posed. As it appears to us, the independence of changes in conditioned
and unconditioned patterns with change in state of the neuron induced by '
- ianophoretic application of agents, as well as the possiblity of appearance -
_ of conditioned act~vation with retention of the pa~tern for ECS, ~ndicate -
_ that the pattern of conditioned activity of many neurons cannot be deter-
mined by generator mechanisms or the synaptic inputs that really activate
the neuron and induce the unconditioned pattern.
190 -
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a
- ,,,,(~-n
- imp �
- 30 a
_ 25 ~
_ ~a =
_
_ Light EC ~
. b
_ b -
= imp~I~ ~
Y
30 ~ ~ -
- 20 j -
/0 ~~'I ~ J~,a � ry ~l~,n ;"j y~ ~n . ' _
~ ~ U~ ~1 ~ ~ j J"~
~
= Light ECS S~
~S Li:ght ECS ~PL -
_ ,f0 ms
I'igure 90. Figure 91.
Effect of L-glutamate on different pat- Appearance of late activation zn =
_ terns of neuronal responses in condi- response to con~3itioned light
= tioned and unconditioned acts. Channel under the ir.fluence of acetyl- _
width 4 ms, n= 25 choline; on histogr~ms: channel =
_ a) initiial neuronal responses: late widti:24 ms, n= 15 . _
~ activation to light and primary a) befnre acetylcholine -
response to ECS b) with acetylcholin.~ -
- b) with T.-glutamate, late activation c) control spries
to light is elimin.ated, primary
response to ECS does not change -
Indeed, the independence of changes in pattern of activity in the first and _
secand acts with delivery of some agent means that the neuron is activa.ted
in these two acts through different synaptic inputs. If we assume that
_ the functional synpatic field of ~ neuron in the conditioned act is
- determined by "detonator" ~nfluences, which really activate the nF:uron after
ECS, the changes in these influences, which weie observed witn ionophoresis,
should have also altered the pattern of tne response to the conditioned
signal in the same direction. Appearance of spikes in some neurons in ~
response to a previously ineffective condition_ed signal, with unchanged -
- pattern after ECS, also indicates that tlie functional synaptic fields de- -
termining the response to light could ~:hange, in the presence of the same `
191 `
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= pattern. The dependence of parameters of future ECS or. "conditioned"
activity of even neurons that do not generally respond to ECS, as we ob- -
ser~�ed not only in neurons of the somatosensory co;-te~, but those of
the visual cortex, speaks in favor of this conclusion.
a
b Figure 92. '
- Effects of different agents on a
_ neuron that is inhibited in the
,J~P unconditioned acL.
_ I ^ a) initial activity
~ ~ ~v`~" b) with atropine
c) control series
_ d) with L-glutamate ~
e) control series
- ~ f) with GABA ;
_ d ` Poststimulus histograms, channel
,
_ width 24 ms; above histograms
. ~1 ~ are averaged evoked potentials, -
. l~'~~11,~1 ~ ~II n = 25 ~
U -
First arraw--light, second--ECS.
= e~,_~~ ~ Calibration: 5 impulses, 50 ms :
~i -
� I~1~~_I,~II ~ ~ ,
I ~i
- .~1.~.~.~ 1 -
- f ,~.~,t~r,,,._.... ;
~ ~
; ~ 1 ~~~"~r~~~~ ~ i
~ i
L -
192 -
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A
Figure 93. _
a Effect of L-glutamate on conditioned
~
y ~ 2 and unconditioned responses of the
+1.~~~...~ same neuron as a function o f p a r a- _
meters of reinforcemeat
A) electrocutaneous stimulation of
contralateral hind foot (30 V, _
1 ms) _
b a) initial activity _
~ U b) with L-glutamate, a response _
_ ~ appears to conditioned
stimulus
c) control series -
B) ECS of contr:ilateral hir_d foot -
~ (50 V, 1 ms~
a) initial activity
b) L-glutamate eliminates primary
response to u~iconditioned -
B stimulus
~ In both cases, the L-glutamate -
dosage was the same (-10 nA, 6 min).
a -
Channel width 24 ms, n= 10 ~
1~~ i -
_ b
Light ECS
5 imp~
50 ms `
- The state.of the neuron very definitely determines che conditioned pattErn,
- and it does so to such an extent that a change in. state of the neuran by
- ionopH.oresis could even create a new response in neurons that did not -
react to light previously. A change in state of a neuron alters the -
- 193
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_ response even of neurons that do not respond to ECS. Thus, we arrive at
- the conclusion that it is expressly a specific organization of internal
metabolic processes that determines the functional synaptic field and,
consequently, the pattern of the neuronal response in a conditioned
- behavi~ral act.
= imp -
- 15 ~
a ~
5 ~ ~
~ ~ ~ ~
- ~s
, I
b s~ I I~ ~ ~I ~ ~I~~~ ~ I ~I ~ ~ ~
~ '
' I ~ I ,
J5 ~ -
- ! �
_ c 3 .I I
. ? i , I~ -
~
- Li_ght CS Z
p Light E S ~ -
Figure 94. Effect of acetylcholine on conditioned re$ponse of the
same neuron as a fun~tion of parameters of reinforce- ~
ment. -
- a) initial activity with ECS of 40 V, I ms, coatralateral hind foot -
_ (on the left) and 30 V, 1 ms, contralateral front foot (on the right) -
= b) against the background of acetylcholine ~
~ cc ntro'. series
Channel width 24 ms, n= 15. In both cases the acetylcholine I
losage is the same ~+50 nA, 8 min) -
~
~ i~
_ Normally, a given state of a neuron determining its FSF is probably
produced by all influences going to this neuron from elements related to
it, which have an integrative effect. Organization of these influences is
_ determined by motivation and situation. As it changes constantly, this -
organization of interneuronal interactions leads to constant changes in
endogenous metabolic processes whi.ch, as they determine neuronal FSF for
- each subsequent mo~nent, in turn determine organization of interneuronal
interactions.
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In our experiments, it is logical to relate organization of integrator in-
- fluences to defense motivation, which always appears in the situation of
" delivering electrocutaneous stimulation. The change in effects of pho-
retically delivered agents with change in parameters of ECS (Figures 93 and
~ 94) and the fact that delivery of an agent to neurons showing different -
! patterns in the two acts usually also elicits different changes in these
patterns and, in general, could influence or~ly one of the patterns, -
indicate that organization of integrator influences of defense motivation
creates differezt FSF in many neurons, to be used in the system of the
conditioned and ur_conditio*~ed act. It may be assumed that, in such neurons,
- the FSF in the conditioned act are created by the "metabolic model of
electrocutaneous stimulation," whereas in the unconditioned act they
are created by the model of "discontir.uation of electrocutaneous stimula- -
tion." This is also indicated by the fact thaC, with change in parameters
of ECS,there is also a change in effects of the agents we used.
In those neurons that present sim.ilar patters in the two successive acts, ~
_ the functional synaptic fields in the two systems are probably created by ~
- motiv2tion as the hierarchy of all goals and future events, wnich is
= what determines the sinilar sensitivit~ of similar patterns to ionophoresis
of the agents.
. Neurons whose activity does not generally change (i.e., "areactive" ones)
and, perhaps, inhibitory neurons probably simply do not have "metabolic -
- models" and, consequently, no syna~tic fields referable to the given motiva-
tion. For this reason, any change in the state of such neurons by mean5 -
- of clectrophoresis cannot create metabolic changes that would be specific
- in relation to organization of synaptic influx. _
Thus, de~ermination of FSF ~y motil~ation emerges a^ an overall ch~nge in
metabolism, which ultimately ?eaus to sat::sfaction thi~ ~notivation.
- Determination of FSF by the model of a concrete event, which is possible
~ in a given situation, energes as definition [specification] of inetabolic -
changes bq integrator infl.u?nces from the situation. On the level of
metabolic mechan.isms of inemory, the same "law of conformity" probably
~ appl.ies: metabolic. processes that would implement syn.t.heG:.~ of specific �
- biochemical recegtors for specific mediators that could be received -
. at a future time are probably initiated only when there is a conformity
_ between the receptors present at this time and real mediators. `
Of course, all these hypotheses, which have been expo~snded on the basis
_ of studies of only "~utput" impulse activity of neurons, require verifica-
tion in special experiments that would enable us to monitor [control] meta-
bolic processes in a neuron d:uri:ng a behavioral act. _
The operational architectonic:.~ of systemic p~ocesses must be invariant on
- all levels of the hierar~hy systems (Anakhin, 197.3). In order to form
� an hierarchy, ~he operational architectonic>> of the functional system of -
a behavioral act and functional system of a neuronal discharge must be _
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functionally identical. To continue the analogy to organization of systemic
processes of a behavioral act, it can be assumed that a spike (or discharge
within a single phase of EP) is the realizatioti of an e.lementary program
of action of the corresponding functional system. According to functional =
- s,ystem theory, generatiar. of spikes is the product of intiegrative activity -
of the neuron, in which convergence of different synaptic influences can
be interpreted as elementary afferent synthesis, while e~tablishment of
"metabolic conlor~ity" between the integrative state and detonator activa-
tion, i.e., functional synaptic field, as elementary decision mal;ing. Since
the neuror_is under the constant inf.luence of other elements of the system
- and through its discharges influences the state of the entire system,
appearance of a spike in the collaterals of its axon shou~d alter the state
of the entire system. This change can be interpreted as a result elicited
by the spike; the rev~~rse influence of the system on the neuron emerges -
as "feedback."
- Aay "anticipatory reflection" is based on anticipatory change in metabolism
(Anokhin, 1962a); this anticipa~ory change in metabolism and preparation of
, subsynaptic membranes for feedback that it elicits can be interpreted as
- an elementary "acceptor of results of action." Thus, organization of
intraneuronaJ. intersynaptic integration allows for analysis of its
functional architectonics from the standpoint ot functional system.theory.
The "principle of conformity" also applies in the performance of a single
behavioral act; although pretriggering integration allows for achieve-
ment of the same result by different means, which corresponds on the
level of a single neuron to potentiation of several FSF, real afferenta-
- tion after a stimulus activates only one functional synaptic field in _
each neuron 3nd determines implementation of a single [only] means of
reaching the goal. In the course of different systemic processes, the
integrator influences organized by each systemic process successively
"narrow down" functional synaptic fields, rendering them more adequate
to the goal and situation.
Complete exclusion of "superfluous degrees of freedom" and determination
of the pattern of neuronal c~ctivity of the neuron in actuating~mechanisms
of the behavioral act by the only goal of this act and spe~ific environment
are achiPVed by the set of integrator influences created at the stage of
afferent synthesis and decision making of expressly this elementary
behavioral act. Evidently, these integrator ~nfluences, created by
~ neuronal discharges during negativity of EP, determine the purposeful and ~
selective sensitivity of neurons to synaptic influxes, wnich appear during -
- performance of action until the result of a given behavioral act is
achieved.
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CHAPTER 6. FUNCTIONAL SYSTEM THEORY, AND THE PSYCHOPHYSIOLOGICAL PROBLEM -
Impossibility of Direct Correlation of Mental and Neurophysiological -
~ Processes
The nature of inental processes and their material substrate have always
been the subject of enormous interest to mankind. And the present time,
"the study of psychosomatic correlations continues to be a most pressing ~
problem, proper work on which would be inconceivable without the first
_ and foremost involvement of neurophysiology"(Dubrovskiy, 1971, p 271). -
As he began to study the brain by objective methods, I. P. Pavlov abserved: -
_ "In essence, there i~ only one thir?g in life that interests us, our mental
content" (Pavlov, 1949, p 351j.
Associatior.ism in psychology and reflex physiological conceptions led to -
interpretation of neurophysiological mechar.isms of the psyche in ~he
teaching on higher nervous activity on the basis of the idea of sameness
of "an elementary mental phenomenon"--associat~on and "the purely -
physiological phenomenon"--�-the conditioned reflex. I. P. Pavlov believed
that "here there is total ft~sion, total absorption of one by the other,
identificatior." (1949, p 521). For a long time this was the idea that
guided the research of both physiologists and psychologists.
The neurophysiological mechanisms of the behavioral act, interpreted as a `
reflex, were limited in essence to conduction of excitation over a specific
route, and they could be described as a succession.of physiological nro-
cesses occurring in different parts of the brain. In the very same way, -
mental processes were directly compared to physiological ones, which
were studied (as we mentioned i.^. the first chapter) in the absence of .
behavior ar.d mental activity.
It turned out ttiat these process:~s could not be~ compa.red, by virtue of
absolutely objective properties of physiological piocesses that are ~
always very definite in time and space, as well as mental processes local- ~
ized only within the entire brain and organism, and within the time of the
en*_ire behavioral act. This circumstance led I. Y. Pavlov to the ne~d
~o exclude psychological concepts from analysis af inechanisms of behavior:
- "As shown by all of the experiments, the entire substance of studying the
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reflex mechanism, wh ich is the foundation of central nervous activity, _
- amounts to spatial relations, determinatian of the pathways over which _
stimulation spreads and collects. Then it is absolutely clear that the -
probability of learning everything on this subject only exists for the -
concepts in this fi eld that are characterized as spatial concepts. This -
is why it must be clear that one cannot use psychological concepts, -
~ which are essentially nonspatial, to delve into the mechanisms of these -
relations. It is necessary to point with a finger to the site of stimulation -
and where it traveled. If you can conceive of this vividly, you will =
, comprehend the entire force and truth of the teaching that we uphold and
are developing, i.e., the teaching on conditioned reflexes, which has
excluded entirely from its realm any psychological concepts, and wliich
always deals only c~ith objective facts, i.e., facts that exist in time and -
space" (1949, p 385).
- Another solution to the prohlem of impossibi~ity of comparing mental and
- reflex processes was offered by psychologists, and it refers to the fact -
- that since "direct r econstruction of perception, feeling or a thought...
from the material of standard nervous i~npulses or graduated bioe~lect*_-ic
potentials cannot be done, this impossibility of formuZating the
characteristics of inental processes in the language oz physiology of
endogenous changes in their substrate is the opposite side of the possi- -
bility of formulatin g them only on the language of properties and re~a- -
tions af their object" (Vekker, 1974, pp 14-15). This conclusion was ~
very logically made by L. M. Vekker on the basis of an analytir_al reflex
premis: any mental process, like any other act of human vital
functions, originates from some human organ" (p 11).
Thus, mental and ref Iex mechanisms cannot be compared, both from the
physiological and psychological points of view.
The organism always emerges as a whole in a behavioral act, and such psycho-
logical concepts a5 motivation, perception, memory or goal reflect
conceptions about processes that are referable not only to the entire
behavioral act, but to the entire organism tliat performs behayior, and they
characterize it exp ressly as a whole. I. P. Pavlov observed that "mental
processes are very c losely linked with physiological phe:~omena, determining -
integral function of an organ" (1949, p 348j. However, with the analytical
approach, integral mental processes could only be compared to local and
special physiological ~rocesses, since systemic processes uniting elements -
into a single whole nad not yet been discovered.
We believe that it is e~cpressly at this point that the "possibility"
appeared of ruling o ut the mental factor from analysis of behavior,
_ since the mental factor is something ovar and above the sum of purely _
= nervous functions and, consequently, it also appeared to be over and
above behavior. Efforts to compare integral mental and special r~euro- _
_ physiological processes also led to psychophysiological parallelism, or
even directly equating mental processes with physiological ones, and the
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intensity of a sub,jective mental experience was compared to the force of
excitation of the corresponding structure, while the content of this ex-
perience was compared to localization of excitation. For example, visual
_ sensations and perceptions were viewed aa excitation of the visual ana-
_ lyzer or as a process accompanying such excitation; motivation was inter- -
- prete3 as excitation or the "subjective aspect" of excitation of som e
hypothalamic center, etc. The concept of threshold made only a quanti- _
tative separation of purely physiological excitatory processes from the
same excitation acc.nmpanied by an experience, and it did not permit posing
the question of quantitative spECifics of the nervous processes at the
- basis of psychological phenomena. At the same time, it is obvious that
by no means any nervous activity is associated with mental experiences.
This circumstance led to a search for the "center of consciousness,"
which was subsequently ca].~ed "anatomization of abstraction" (Burns, 1969).
We believe that, by exclu~ing the psychological element from mechanisms
of behavior, reflex theory only created the illus~on of the possibility of
a purely physiological explanation of behavior. It appears to us that it
- appeared because the reflex was the basis of all conceptions of reflex -
theory, i.e., the phenomenon that occurs in spina' and anesthetized
preparations,in which, o� cours~ there is no integral adaptive behavior
and, consequently, there is indeed n. u~~nd. Physiological reflex theory
provided a"purely" physiological interpretation of the causes and
mechanisms of behavior, in which reflection of objective reality by the
- brain was limited to physiological processes. In the reflex scheme of
the behavioral act, which was an arc linking different effects to different
reactions of different organs, there was simply no need for informational
relations ~between the environment and the organism as a whole nor, conse-
quently, for the mind. Since, however, there were few who were willing
to negate the mind in general, the latter always emerged as an
_ "epiphenomenon," which was not mandatory for performance of behavior.
~ For this reason, efforts to reconcile physiological and psycliological des-
_ criptions of an elementary behavioral act were always within the framework
~ of psychophy5iological parallelism.
The Problem of Correlation af Systemic and Mental Processes
Analysis of the development of psychology and physiology from the systemic _
point of view led P. K. Anokhin to the conclusion that, in order to paint a
complete natural scientific picture of brain function, it was not necessary
to blend or equate physiological and psychological elements, but that
a"conceptual bridge" was needed that would permit comparison of the
concepts of these two disciplines and see physiological mechanisms -
behind psychological phenomena.
_ Since there are specific systemic processes of organization, qualitatively
different from elementary ones, and integral behavioral acts are linked
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expressly with systemic processes, mental processes are based on systemic
- processes of organization of different processes into a single whole,
rather than elementary physiological processes of excitation or inhibition.
A comparison of neurophysiological and mental processes is possible only
through processes on the systemic level.
- A comparison of the concepts of systemic and psychological processes no
longer involves the difficulties that arise if one directly compares mental
and physiological processes. Indeed, in addition to time and space charac-
- teristics in common in physiological and systemic processes, the latter
are also characterized by the parameters of integrity [wholeness] and organi-
zation. Like any processes of organization, systemic processes of the be-
- havioral act are distinctive information processes, for which the "physio-
logical level" emerges as the "material carrier." A comparison of these
informational parameters of systemic processes to mental ones is then made
using the same gage with regard to meani.ng, since both information and
psychological processes are systemic properties of the organism as a whole.
The systemic nature of organization of processes is comparab le to psycho-
~ logical processes in the sense of the latter's reflective functior.. Psy-
chologists have provided convincing proof of the active role of perceptive '
actions in apprehension (Zinchenko, Lomov, 1960; Zaporozhets et al., 1967),
and this compels us to question any physiological conceptions concerning
purely sensory organization of in.formation processing in analyzers, from
"receptor to cortex." Some correlation or other can be demonstrated
between reactions and a stimulus in any part of the brain and, consequently,
this phenom~~non cannot be interpreted as an indication of expressly
mental reflection of the properties of a stimulus in the activity of some
analyzer. Such properties of apprehension as activity, integrity and
objectivity [in the iiature of an object] cannot be compared to processes
within a single analyzer, and they require a systemic foundation. At the
same time, as we have tried to demonstrate, analyzers are also involved
in such systemic processes as the program of action. Experimeuts with
reverse masking (D~nchin, Lindsley, 1964; Kostandov, Shostakovich, 1970;
Massaro, Kahn, 1973) and direct comparison of evoked potentials to
the re~orts of subjects (Rosner, Goff, 1969; Libet et al., 1967; Fox et
al., 1973), revealed that about 100 ms are required, with presence of both
earl}r and late components of EP, for the appearance of subjective sensa-
tions.
= At the same time, physiological experiments did not confirm the conceptions
that are popular in psychology and based on the reflex princip le to the
- effect that expressly the motor elements of perceptive actions, likened
to the properties of an object, implement the creation of an adequate image
of this object. Indeed, in afferent synthesis of the percept ual act,
material in memory about movements can only be one of the components,
along with information about the entire sensory.situation (p ast, present
and future).
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Since information about any event is used, as we tried to show, during all
systemic processes and for organization of activity of elements of the
entire system, perception as a reflection of the properties of an exogenous
object is linked with many central and peripheral structures, and with all
the key mechanisms of the functional system of the perceptual behavioral
act. Thus, the reflective role of the mind is thus comparable to the pro--
cess of the organism's use of exogenous information (organization of en-
vironment) to build its own organization (organization of physiological
processes).
As for the regulatory role of the mind, it couid be compared to the systemi~ ~
process of action: the higher the degree of organization of processes within _
the system, ~ut more perfect the behavioral acts and the better the result
is attained. Here, the process of afferent synthesis and decision making
translates information (order [organization]) about the environment into
an order of physiological elementary processes in the system, while reverse =
translation of organizaticn of the system into orderliness of the environ-
ment occurs with the function of systemic mechanisms of the acceptor of
results and prograr.: of action, when the action is performed, i.e., ~
organized function of physiological actuating mechanisms, and real results -
of behavior. are attained, i.e., new organization of the environment. Thus,
the function of determination of behavior by the mind can be compared to
the organizational parameters of systemic processes.
Insofar as systemic processes consist exclusively of physiological processes -
and organization of these processes creates a new quality--informational
parameter of the system, comparable to the conception of the mind, physio- ~
logical and mental determination of behavior are inseparable, and they do -
not exist without one auother or without informational or systemic
determination.
Although the "mental" factor is an attribute only o� integration as a whole,
_ this does n~t preclude the existence of a specific structure in mental pro-
cesses. Since mental processes are based on integral behavioral acts,
functional system theory, which describes the structure of behavioral acts,
is also applicable for description of the structure of inental processes,
and each of the key mechanisms of the functional system has, as can be seen
from the submitted experimental results, very concrete neuronal implementa-
tion. -
The properties of inental processes (for example, perception), demonstrated -
in psychological experiments, are found to be~comparable to systemic
mechanisms and, conseuqently, a very definite form of activity of concrete
. neurons. The activity of perception and apperception can be the consequence
of presence in the functional system oF the perceptual act of the system
of the acczptor of action results, which actively reques~s the required
information from the exogneous environment. Perception a.s objective [ob~ect-
related] and integral [whole], because information retrieved simultaneously
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from the most varied regions o~ the brain and combined in pretriggering -
integration is involved in afferent synthesis; it is constant when the
- 3ensory situation changes, because material from memory is inputted jn
afferent synthesis, rather than the recoded state of receptors. Of
course, this is only an example of possible comparison of properties of
mental ~nd systemic processes. -
Functional system theory provides the same "operational architectpnics" `
_ for any behavioral act; at the same time, psychology makes a distinction
between several mental processes, such as perception, thinking, remembering, -
etc. In view of the fact that there is objective manifestation of the
_ mind only through behavior and that the concept of activity (behavior for
animals) is included in current conceptions of any mentai prQCesses, it may
be assumed that the specific mental processes singled outby psychology ~
- can be compared to the specific characteristics of the same systemic
processes corresponding to behavior, which is consistent with the thesis
_ of integrity [wholeness] of the mind. The structure of systemic processes
is, apparently, referable to the common features of "an;~ menta~. process
which constitute the basis for distinguishing between the mental and
nnnmental" (Vekker, 1974, p 10).
Functional system theory enables us to refer to the concept of quantity
_ of information in the system and raise the question of dependen~ce of
� properties of integration on informational and energy characteristic~ of
the stimulus, as well as the problem of quantitative correlations between
the properties of integration and se~.sation, to provide objective quanti-
_ tative characteristics of sensation. Perhaps, the solution to these -
problems will rela.te ~he stimulus and sensation in a f~rmula free of the
objections th~t the psychophysiological law is presently encountering
(Luce, Galanter, 1967; Pieron, 1966; Lomov, ~574). The approaches to
_ the psychological solution of the problem of signal detection (Zabrodin,
= 1973) and the problem of reaction time (Stepanskiy, 1972; Oshanin, Konopkin,
1973) presently requrie analysis of the entire behavioral act and the =
entire experimental situation.
At the present time, we do not know at all which systemic characteristics
- uri.ll be comparable to s~me mental chara~teristics or other; hcwever, the
possibilities that are ~resently emerging of calculating such systemic _
characteristics as complexity, orderliness, wholeness, volume, composition, -
- organizatioLZ of a system, etc. (Ferster, 1964; Gorskiy, 1974) gives us _
- hope that there will be a qu~ntitative verification of this hypo*_hesis. `
The idea of expres~ly integrative activity of the nervous system has a?r,ng
_ history (see,for example: Anokhin, 1968, pp 194-202). The conception of
= systemic organization of pt~ysiological mechanisms in behavior has been ~
widely accepted ir. modera physiology, and it is winning increasing
_ recognition by psychologists and pnilosophers. -
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Having proved the existence of specifically systemic processes of integra- ~
= tion, qualitatively different from elementary physiological procPSSes,
functional system theory removed the main obstacle i:o synthesis of psycho- _
logy and physiology, which consisted of the fact that, in ana~ytical
- neurophysiological experiments, the researcher always deals only with local _
' and special processes, whereas behavior and mental processes are related
= to function of the brain and the organism as a whole. Thereby, functional
system theory made it possible to synthesize physiology and psychology
= while confirming the qualltative specifics of their objects of investiga- `
- tion. The solution to the psychophysiological problem apparently
- consists of the fact that organization of physiological processes into -
a single system occurs by means of qualitatively unique systemic processes;
their substrate is physiology, while their informational content refers to
the properties and relations of exogenous objects. Interpenetration and -
- mutual enrichment of physiology and psychology are po~ssible expressly
on the level of consideration of systemic processes. -
Functional system theory, first formulated in general terms in physiology _
as far back as 1935 (Anokhin, 1935), is becoming the logical basis of
_ systemic conceptions that are presently also being developeci in psycl-~ology
(Lomov, 1975)� The use of the same methodological approach to problems
- of consciousness and the mind in these disciplines opens up new opportu-
_ nities for synthesis of a single natural scientific idea about the world. ~
- At the same time, the systemic approach also imposes certain requirements
r.ot only of physiological, but psychological research.
- At the present time, we can still encounter quite often interpretation of -
different mental processes as independent realities, which merely interact
- with one another. ror example, it is assumed that prccesses of perception _
and memory, attention and thinking are independent, and determination is . �
- made of their influence on one another. This analytical, or "atomistic" -
~ approach, which was in its day a necessary stage of development of psy- `
_ chology, is now in contradictien with conceptions of integrity [wholeness]
_ of the mind and unity of behavior and the mind. As noted by D. T. -
Dubrovskiy, this approach led to a situation where "t;~e term 'mental,'
which is one of the most widely used in modern scientific parlance, -
entails a variegated 'train' woven of different meanings and values.
- And in this fo3-m it appears as the cornerstone of psychology, reflecting its
- lack of theoretical organization" (1971, p 162). _
At the same time, the integrity of inental phenomena, the impossibility of
_ breaking them down into pieces, are usually mentioned as one of the
fundamental features of the mind. It appears to us that the theses `
- gained by psychology concerning the integrity of the mirid and unity of -
behavior and mind constitute a good foundation for the systemic approach.
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Correlation of Systemic and Neurophysiological Processes
~
This section is a summation of sorts of the conceptions that were di~cussed
on the preceding pages. -
_ We shall begin the comparison of systemic processes to processes on the
: neuronal level with consideration of inemory ("life experience"). On the
behavioral level, memory emerges as an hierarchy of goal-directed behavioral
- acts, which lead to survival of the organism under some conditions or
= other. Or. the neurophysiological level, different behavioral acts are the
integration of a selective set of neurons with functions that are deter-
mined by organization of associations of each neuron.
_ On the level of a single neuron, its "life experience" consists of a set -
of functional synaptic fields that are used in any behavioral acts. These -
functional synaptic fields are hierarchically organized, and they are based
_ on an hierarchy of inetabolic processes within the neuron. -
- Thus, the behavioral act stored in memory"is the possibility of coordina- =
tion of activities, functions, functional synaptic fields and metabolism
= of many elements, which leads to survival under specific conditions. -
~otivations are based on metabolic changes, which determine the appropriate -
- organization of functional synaptic fields and, consequently, possible -
organizations of interneuronal relations. ~ -
- The situation affec~s the hierarchy of life experience in the opposite ;
~ direction: a certain organization of exogenous factors has a corresponding
= influence on coordination c~f neuronal metabolism through synaptic influ- ~
ences on specific functional synaptic fields.
Al1 these correlations ~occur onlythrough systemic processes: local changes ;
- in tissular metabolism, for example, in the hypothalamus, become motivation
, only through interneuronal and intertissular coordination of inetabolism on
_ the scale of the entire braln and organism; exogenous factors are considered '
= as a situation only by comparing the effects to functional synaptic fields
= which, in turn, are determined by metabolism. ~
~ Since the situation.and motivation change constantly, the interneuronal
= integrations, functional synaptic fields and metabolism of different neurons j
are in constant dynamic conformity with both motivation and situation.
- Goal-directed behavior develops when it is necessary and possible to
alter this conformity in the direction of im~irovement. _
In the continuum of behavior, pretriggering integration is formed while ,
performing the preceding action and achieving interim results. At this ;
- time, impulsation is related to coordination of functions of the subsystems
of the preceding action; at the same time, through integrator effects,.it
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adjusts the functional synaptic fields to ihe properties of the future re-
sult of the entire behavioral act and thereby reduces the degrees of -
freedom of both individual neurons and the entire organism.
There is constant comparison of exogenous properties to the parameters of
generated FSF; howev~?-, in the process of performing an action, the
exogeno~s environment is c:ompared to the parameters of FSF generated by
- subsystems on the physiological level. Performance of action leads to
appearance in the environment of a result--event, which is then used to
coordinate the functions in the entire brain and organism. For this reason,
_ when a result appears in the environment, its parameters are compared to -
the prepared FSF in all analyzers, in accordance with the goal of a given
behavioral act, which we considered to be the "preceding" one. ~
- Comparison of the real multimodal parameters of the result to the parameters -
of the goal leads to appearance of a primary response that is synchronous _
_ in many structures of the brain. This response occurs only in the set of -
neurons whose FSF were prepared for the parameters of the result of action
by the time it appeared. By virtue of hierarchi,c organization of FSF,
- only the fields ir.cluded in the hierarchy of tHe entire motivated behavior
_ can be prepared, and only in neurons whose function had ever been used to
_ attain the future result.
Thus, at the moment of the primary t~esponse, pretriggering integr~~tion
tha~t contains the possibility of performing several acts becomes signifi-
cantly reduced, and there is activation only of neurons whose FSF~must
meet one of two conditiozs: 1) ever been used.to reach some goal under cir-
cumstan ces analogous to the state of the environment at the time of the
prim.ary respcnse; 2) ever been used to attain the needed result under any =
circumstances.
~ The discharges of units with such properties through integrator effects _
lead to expansion of FSF in the direction of coordination with both the
situation and the goal. This stage correspor.ds to aff erent synthesj.s _
and decision making, and in the evoked potential it corresponds to
ne~ative oscillation. -
At this time, disr_harges appear in the set of neurons whose FSF meet
both conditions simultaneously, i.e., they had been used at some time to
reach a given goal in expressly the given environment. -
There can still be several integrations that could lead to a given goal �
in a given environment; of course, concurrent implementation thereof would -
disarrange the system and lead to diverse mistakes.
= The final choice of one means of coordination of the activity of all elements
on the scale of the entire brain and organism occurs next: when neuronal
discharges corresponding to negative oscillation of evoked potentials -
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alter FSF through integrator effects in such a way that discharges can
' appear only in neurons whose coordinated activity led in the past to the
required result. This stage corresponds to complete reductioa of deg~ees -
of freedom of both single neurons and the entire organism; this is the
stage of the acceptor of results and program of action. From this time on,
- the different physiological subsystems that were coordinated in the prc-
_ ceding stages of farmation of the functional system of th? behavioral -
act begin to function in accordance with the hi.erarchy of results making
- up the acceptor of results of action in the behavioral act. F
' The real information about results in subsystems alters integration and -
prepares new pretriggering integration for appearance of the result of
the entire behavioral act, etc. .
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CONCLTJSION -
H11 of the problems dealt with in our study of neurophysiological mechanisms
of systemic processes require further development and definition. 1'his -
~ applies to both the conceptual system and hypotheses, as wel.l as
- conclusions.
Use of functional system theory in our approach to behavior opens up a wide
- range of problems that must be submitted to experimental and theoretical
studies in both the physiological and psychological aspects. .
We have barely touched upon the problem of hierarchy of behavioral acts
in complex behavior and the problem of automation of the behavioral act,
when it probably becones a sybsystem on the physiological level. The
very content of our dtscussion of systemic processes may change appreciably
= when adequate ~uantitative gages will be found to describe organization,
integrity, composition, size and other systemic parameters.
- The feasibility of comparing mental and physiological processes with
the use of systemic ones raises the question of direct identification of
- neurophysiological bases of inental processes and states. For example,
= it may be that the "quantity of motivation" can be measured by the number
of elements involved in integration and extent of expansion of ttieir
functional synaptic fields, and that the "quantity of perception" can be
measured by the number of degrees of freedom removed by some perceived
_ event from elements and the organism as a whole. These are all problems
- that can be solved directly through psychophysiological epxeriments from
the positions of functional system theory.
- The learning problem, i.e., formation of new functional systems under the
system-f~rming influence of goals and results, requires systemic analysis,
' and it is becoming a part of general problems of systemogenesis.
. Apparently, development of neurophysiology, psychology and other disciplines,
the correlations between which are becoming possible by virtue of the -
- sameness of the systemic approach in these fields, will very soon lead to
significant clarification and even radical change in the initial theses
of functional system theory, as is presently happening with Darwinism.
_ However, the significance of functional system theory, which was expounded
207 -
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by P. K. Anokhin, is not that it is growing rigid, like a dogma, but
t~'~at "a genuine idea is capable of attracting, like a magnet, only
'iron' facts from a pile of diverse facts."*
- i-
. ~
;
*SOVETSKIY SOYUZ [Soviet Union], No 11, 1972, p 37.
208
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240. Buchwald, I. S.; Beatty, D.; and Eldred, E. "Conditioned Responses
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246. Chang, H. T. "The Evoked Potentials," in "Handbook of Physiology,"
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248. Chow, K. L.; Randall, W.; and Morrell, F. "Effect of Brain Lesion
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249. Coguery, I.-M.; Coulmance, M.; and Leron, M. "Modification of
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250. Cohn, R. "Visual Evoked Responses in the Brain-Injured Monkey,"
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[1172-10,657] i=
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