Many branches of natural science apply P.K. Anokhin’s theory of functional systems in practice, which is evidence of its universality. The academician is considered a student of I.P. Pavlov; it was only during his student years that he was lucky enough to work under the strict guidance of V.M. Bekhterev. The influence of the fundamental views of these great scientists prompted P.K. Anokhin to create and substantiate the general theory of functional systems.

Historical background

Some of the results of Pavlov's research are still studied in educational institutions today. It should be noted that Darwin's theory has not been removed from the school curriculum, but concrete evidence of its truth has not been provided to the scientific community. It is taken “on faith.”

However, observations of the Earth's ecosystem confirm that it does not exist: plants share nutrients and moisture with each other, distributing everything evenly.

In the animal world, you can notice that individuals do not kill more than is necessary to ensure their livelihoods. Animals that disrupt the natural balance through abnormal behavior (for example, they begin to kill everyone), as sometimes happens with some representatives of the wolf pack, are exterminated by their own relatives.

Observations of primitive tribes that survived into the twentieth century, studying their culture and everyday life, we can conclude about a primitive person who felt, understood, knew that he was part of the environment. When he killed an animal for food, he left something of what he killed, not as a trophy, but as a reminder of someone’s life wasted to continue his own.

From this follows the conclusion that ancient people had a concept of community, dependence on various environmental factors.

Field of research of Petr Kuzmich

PK Anokhin’s theory, on the contrary, is built on the basis of an extensive experimental base and a clearly structured methodology. However, many years of observations, practice, experiments, and theoretical elaboration of the results led the academician to this concept. The results of experiments by Pavlov, Bekhterev, and Sechenov played an important role in the formation of a systematic approach to the problem of purposeful activity. At the same time, the concept of functional systems cannot be called a “copying” or “continuation” of the theories of the listed authors due to the difference in methodology and general structure.

Methodological approaches of Pavlov and Anokhin

Upon a detailed examination of the concepts, one can notice that the positions of the methodology are understood and explained by the authors in completely different ways.

Methodological principles used in the authors' concepts
P. K. Anokhin	I. P. Pavlov
The author does not support the concept of universal methodology for all exact sciences. Emphasizes the importance of the influence of exogenous and endogenous factors on mental processes.	The universality of the methodology for studying the subject of all exact sciences is the main postulate of the scientific nature of the study of mental processes (most likely, this is an attempt to bring the study of consciousness to the level of “scientific” through the mechanical transfer of study methods from other areas of science).
Distinguishes between the laws by which living matter and the inorganic world function. He justifies his position by the presence of an “internal focus on survival” in living organisms, which is not characteristic of inanimate objects.	Mental processes, according to Pavlov, are subject to compliance with the laws governing the development and functioning of the material world.
The concept of “integrity” means the mobilization of the internal forces of the body to achieve a specific goal.	“Integrity” (close relationship) manifests itself when the body is exposed to external factors.
The hierarchy of processes implies the presence of feedback, which implies an influence on the control center of the coordinated elements of the system. Based on these interactions, the stages of the hierarchical structure are distinguished: molecular; cellular; organ and tissue; organismic; population-species; ecosystem; biosphere.	The organism is considered as levels of organization located within each other. Hierarchy is considered as a vertical management organization or a pyramidal organization of control centers without the possibility of reverse influence from lower-lying components of the system.
The mechanisms for reflecting reality are dynamic, not static, and are formed due to various external factors and a programmed goal in a specific period of time. The body has the ability of advanced reflection.	Conditioned and unconditioned reflexes according to Pavlov manifest themselves independently of other reactions of the body and consist of two processes - inhibition and activation.
Consciousness cannot be reduced to physiological reactions, arising on the basis of their development.	Elementary thinking arises on the basis of a combination of individual reflexes caused by a specific sensation or symbol.
the creator of the theory of functional systems, is based on the postulate “the law of a thing is in the thing itself.” Therefore, all processes are governed by patterns inherent only to them. Consequently, the structure of world laws resembles the principle of a “matryoshka” rather than a “pyramid”. Since management occurs with the help of different laws, the methods of study must be different.	The concept is based on the postulate “the law of a thing is outside a thing,” which indicates the independence of the law from the controlled process. At the same time, a hierarchy of subordination of laws (pyramid) is built. Consequently, all processes are subject to universal laws with observance in living, inanimate nature, and mental formations.

The given basic methodological principles of the authors allow us to draw a conclusion about their “opposite”. The theory of functional systems of Pyotr Anokhin cannot be a logical continuation of the materialistic teachings of I. P. Pavlov.

The influence of the works of V. M. Bekhterev

A historical fact is the disagreement between the creator of Objective psychology and Pavlov. Thanks to the latter’s vindictiveness and pettiness, Bekhterev was not awarded the Nobel Prize.

The author of the theory of functional systems describes the functioning of Pavlov's school as voicing many hypotheses (taken on faith) against the background of one fundamental discovery (the conditioned reflex). Indeed, the works of the famous physiologist (these are several volumes of Pavlovian environments) are a discussion with employees of the main hypotheses and assumptions.

Pavlov’s scientific works received recognition from the world community and were, for their time, quite progressive, but “reflexology”, formalized by Bekhterev, had the objectivity that Pavlov’s theory lacked. She studied the influence of human physiology on his socialization and behavior.

It should be noted that after the mysterious death of Vladimir Mikhailovich, both “Reflexology” and “Objective Psychology”, as scientific movements, were “frozen”.

Studying the legacy of Bekhterev and Anokhin, one can notice some general principles in the methodology of studying the subject. It is also worthy of attention that the theoretical assumptions of both authors were always based on practical research and observations. While Pavlov allowed “devastating reviews” only because of personal hostility.

The emergence of the concept, its development

The foundations of the theory of functional systems were laid back in the thirties of the twentieth century based on the study of the interaction of central and peripheral nervous activity. Pyotr Kuzmich gained rich practical experience at the All-Union Institute of Experimental Medicine named after A. M. Gorky, which served as the basis for the creation of the USSR Academy of Medical Sciences and the Leningrad Institute of Experimental Medicine in the forties.

The academician was able to study nervous activity not only at the general biological level. The first steps were taken in research into the embryological aspects of the functioning of higher nervous activity. As a result, the structural and functional approaches in Anokhin’s systems theory are recognized as the most advanced. It highlights private mechanisms and their integration into a more complex system of a higher order.

Describing the structure of behavioral reactions, the academician came to the conclusion about the integration of private mechanisms into a holistic behavioral act. This principle was called the “functional system”. It is not a simple sum of reflexes, but rather their combination into complexes of a higher order, according to the theory of functional systems, that initiates human behavior.

Using the same principles, one can consider not only complex behavioral reactions, but also individual motor acts. Self-regulation is the main effective principle in Anokhin’s theory of the functional system. Achieving planned goals that benefit the body occurs through the interaction and self-regulation of smaller components of the system.

The publication of Anokhin’s book “Philosophical Aspects of the Theory of a Functional System” includes selected works covering issues of natural and artificial intelligence, physiology and cybernetics, as well as system-forming factors.

Systemogenesis as the basis of the theory

The definition describes a “functional system” as the production of a useful result through the interaction of elements of a broad, constantly transforming distributed system. The universality of the theory of the functional system of Anokhin P.K. lies in its application in relation to any purposeful action.

From a physiological point of view, functional systems are divided into two categories:

• The first of them is designed to maintain the constancy of the basic parameters of the body through self-regulation, for example, maintaining body temperature. In case of any deviations, self-regulation processes of the internal environment are launched.
• The second ensures adaptation to the environment due to its connection with it, which regulates behavior change. It is this system that underlies various behavioral reactions. Information about changes in the external environment is a natural incentive to adjust various behavioral forms.

The structure of the central system consists of successive stages:

• afferent synthesis (or “bringing” to an organ or nerve center);
• decision-making;
• acceptor of action results (or “acceptance” of action results);
• efferent synthesis (“outgoing”, transmitting impulses);
• formation of action;
• assessment of the achieved result.

Various kinds of motives and needs (vital (thirst, hunger), social (communication, recognition), ideal (spiritual and cultural self-realization)) stimulate and correct the form of behavior. However, in order to move to the stage of purposeful activity, the action of “trigger stimuli” is required, with the help of which the transition to the decision-making stage occurs.

This stage is implemented on the basis of programming the results of future actions through the involvement of a person’s individual memory in relation to surrounding objects and methods of action to achieve the goal.

Goal setting in theory

Isolating the purpose of behavior in Anokhin's functional system theory is a key point. Both positive and negative leading emotions are directly related to goal setting. They set the vector and help highlight the purpose of behavior, laying the foundations of morality from the position of the theory of functional systems. Situational emotions act as a regulator of behavior at this stage of goal achievement and can provoke a goal abandonment or a change in the plan to achieve the desired.

The principles of P.K. Anokhin’s theory of functional systems are based on the statement that it is impossible to equate the sequence of reflexes with goal-directed behavior. Behavior differs from a chain of reflexes in the presence of a systematized structure based on the programming of actions using a proactive reflection of reality. Comparison of the results of an action with the program and other related processes determine the purposefulness of behavior.

Functional system diagram

Academic theory and cybernetics

Cybernetics is the science of the laws of control processes in various systems. Cybernetics methods are used in cases where the collision of a system with the environment has caused certain changes (adjustments) in the way the system itself behaves.

It is easy to notice that there are certain areas of contact between cybernetics and Anokhin’s theory of functional systems. We should briefly describe Pyotr Kuzmich’s attitude to the then new science. He is rightly called a propagandist and developer of cybernetics issues. This is evidenced by the articles included in the collection “Philosophical Aspects of the Theory of a Functional System.”

The book “Selected Works” is interesting in this regard. Cybernetics of functional systems". It describes in detail the issues and problems of cybernetics and their possible solution using the theory of functional systems, which is cited as the basic principle of control among biological systems.

The role of P.K. Anokhin in the development of the systems approach is to substantiate the scientific theory with precise physiological argumentation, unlike his predecessors. Anokhin's theory is a universal model of the body's functioning, which has precise formulations. It is also impossible to ignore the functioning of the model based on self-regulation processes.

The universality of the theory of functional systems is expressed in the possibility of studying the activity of systems of any complexity, since it has a fairly well-developed structured model. With the help of numerous experiments, it was proven that the laws of cybernetics are characteristic of any functional systems included in living organisms.

Finally

The theory of Anokhin Pyotr Kuzmich, which has existed for more than fifty years, defines a person as a self-regulating system that is in unity with the surrounding world. On this basis, new theories about the occurrence of diseases and their treatment, as well as many psychological concepts, emerged.

The works of P.K. Anokhin and N.A. Bernstein represent a manifestation of a new line in human physiology, associated both with a holistic understanding of the body as an indivisible integrity, developing in continuous interaction with the environment, and with the cybernetic models of purposeful activity that emerged a little later, which they anticipated.

Using his own terminology and placing emphasis somewhat differently, P.K. Anokhin also built physiology on new foundations, placing at the center of study “systems that have the ability of internal self-organization, dynamically and adequately adapting the body to changes in the external environment.” The key concept of P.K. Anokhin’s theory is the concept of a functional system. “By a functional system we mean a combination of processes and mechanisms that, being formed dynamically depending on a given situation, certainly leads to a final adaptive effect that is beneficial for the body precisely in this particular situation.” It “represents a strictly defined group of processes and structures united to perform some specific qualitatively unique function of the organism or an act of its behavior.”

This is a branched morphophysiological apparatus, covering all levels and systems of the body, “a central-peripheral formation in which impulses circulate both from the center to the periphery and from the periphery to the center (“reverse afferentation”), which creates continuous information from the central nervous system about the achieved on the periphery results." At any given moment, only one functional system can actually exist, which gives rise to incompatibility of movements included in different functional systems.

A living organism, therefore, is a special case of a self-regulating system, the structure of which must include at least the following elements:

“1) an effector (motor), the operation of which is subject to regulation according to this parameter;

2) a master element that introduces the required value of the controlled parameter into the system in one way or another;

3) a receptor that perceives the actual current values ​​of the parameter and signals them in some way to the comparison device;

4) a comparison device that perceives the discrepancy between the actual and required values ​​with its magnitude and sign;

5) a device that re-encrypts the data of the comparison device into correction pulses fed via feedback to the controller;

6) a regulator that controls the functioning of the effector according to this parameter”...

Norbert Wiener - founder of cybernetics, who owns a number of works devoted to issues of philosophy and methodology of science, the role of scientific knowledge in society, the problem of the universe, analysis of the possible consequences of the scientific and technological revolution, as well as the ethics of a scientist. In ancient Greece, cybernetics was the name given to the art of navigation. ( "cybernetes" means "helmsman", "helmsman" in Greek.) CYBERNETICS (Greek - art of management) - the science of managing, receiving, transmitting and converting information in cybernetic systems.

Wiener's interest in philosophical issues is not accidental: it is known that at first he was going to devote himself to philosophy, receiving a doctorate at the age of 18, and only then, continuing to improve his education, under the influence of Russell, gave preference to mathematics. Nevertheless, Wiener, in his scientific work, repeatedly turned to philosophical topics both in the “pre-cybernetic” period and when developing a project for a new science “about control and communication in animals and machines.”

Cybernetics is the science of the general laws of control in nature, society, living organisms and machines, or the science of control, communication and information processing. The object of study is dynamic systems. The subject is information processes related to their management.

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(26.01.1898 - 05.03.1974)

PC. Anokhin is a student and follower of Academician I.P. Pavlova, Doctor of Medical Sciences, Professor, Academician of the USSR Academy of Medical Sciences and Academy of Medical Sciences, State Prize laureate. Headed the Department of Normal Physiology at the State Medical Institute (1930-1934). Founder of the Nizhny Novgorod physiological school.

Pyotr Kuzmich Anokhin was born on January 14 (27), 1898 in Tsaritsyn (now Volgograd), Saratov province, into the family of a railway worker, who lived in the poorest part of the city, called “Ravine,” in a log house.

After graduating from higher primary school, Pyotr Anokhin in 1914 passed an external exam for 6 classes of a real school and the following year entered the land surveying and agronomic school in Novocherkassk.

P. Anokhin's student years coincided with the most complex and striking political events in the history of our country. He accepted the revolution as a struggle for justice. The revolutionary whirlwind picked up the young man and plunged him headlong into the cycle of the Civil War. PC. Anokhin participated in the establishment of Soviet power in the Don, worked as a press commissioner in Novocherkassk, and as editor of the newspaper “Red Don”, but very soon he realized his true destiny.

In 1921 P.K. Anokhin entered the Petrograd State Institute of Medical Knowledge (formerly the Psychoneurological Institute, created by V.M. Bekhterev), from which he graduated in 1926.

It must be said that, as a first-year student P.K. Anokhin felt a craving for research activities. Professor V.M. Bekhterev forever instilled in the soul of the young scientist an interest in the problems of studying the human brain and psyche. As a result, he wrote a scientific work, “The influence of major and minor combinations of sounds on excitation and inhibition in the cerebral cortex,” but later chose a different path. He is interested in experiments with animals, with the brain.

Working for Academician I.P. provided such opportunities. Pavlova. Their meeting took place in 1922. In the laboratory of the Military Medical Academy under the leadership of I.P. Pavlov Anokhin performed a number of works on the physiology of higher nervous activity.

In 1926 P.K. Anokhin was elected by competition as a senior assistant at the Department of Physiology of the Leningrad Zootechnical Institute. There he received good methodological training from representatives of the physiological school N.E. Vvedensky professors F.E. Tura, Yu.M. Uflyanda, as a result, performed an interesting and original work on the salivary gland. In 1929, a young researcher at the same department received an independent docent course.

In 1930 P.K. Anokhin, on the recommendation of Academician I.P. Pavlova was admitted to the newly created Nizhny Novgorod Medical Institute. He is entrusted with the organization of the Department of Normal Physiology. Pyotr Kuzmich becomes the first head of this department.

Arriving at the institute, the young professor immediately became a popular person. His lectures, brilliant in style and deep in content, were easy to remember. The ability to sharply pose questions and discuss them in a businesslike manner, erudition combined with a lively temperament - all this literally ignited employees and students.

With the arrival of a new professor at our institute, the work of the scientific student circle has noticeably intensified. Soon a group of young people rallied around him, who formed the core of his future school and later became his associates in science.

At the first stage of his activity at the Gorky Medical Institute, P.K. Anokhin proposed a new secretory-motor technique. Among the works of this period, begun in 1931, were studies of the problem of the relationship between the center and the periphery of nervous activity. As a result, the scientist came to the conclusion that the nervous system carries out its integrative activity not only according to a structural, but also according to a specifically functional principle.

It should be noted that the central problem of the professor’s scientific research was the development of the theory of functional systems of the body, which he worked on since 1930.

The result of the work of the team headed by P.K. Anokhin, was the publication in 1935 of a collection of works “The Problem of Center and Periphery in the Physiology of Nervous Activity”, in which the concept of a functional system as a dynamic morphofunctional organization of interacting components providing an adaptive effect beneficial to the body was first formulated. The formulation of reverse afferentation essentially won the priority of our country in physiological cybernetics, which was far ahead of the birth of the cybernetic direction as a whole.

An extensive study of the functional systems of the body led Professor P.K. Anokhin to the formulation of original ideas about the systemic organization of integral adaptations of body functions. It was a further creative development of reflex therapy, allowing us to reveal the scheme of adaptive activity, introducing new concepts into it, such as the result of activity, acceptor of the result of action, reverse afferentation about the result, etc.

The study of the mechanisms of maturation of functional systems during the period of ontogenetic development of functions led the scientist to the formulation of a new principle of development - systemogenesis. The essence of the theory of systemogenesis is that by the time of birth of animals and humans, first of all, those functional systems that ensure the survival of the newborn immediately after birth mature heterochronically and selectively.

Subsequently, the use of the theory of the functional system in pathological conditions led P.K. Anokhin to the development of a theory of compensation for impaired functions. The basic eight principles of this theory are now widely used in practical medicine and physiology.

Based on the theory of the functional system, for the first time in physiology and pathology, the mechanisms of compensatory adaptations in the functional respiratory system during total resection of the lung in humans were revealed.

The Department of Normal Physiology already in 1932 became a branch of the All-Union Institute of Experimental Medicine (VIEM), where Professor P.K. Anokhin became the head of the department of general physiology of higher nervous activity.

In 1934, Professor Anokhin, together with the branch, was transferred to Moscow, where the scientist continued his research, but back in 1935 he came to Gorky, gave lectures, and took part in discussions of scientific issues.

In Moscow, the structure of VIEM was finally developed, consisting of 28 departments, not counting branches. Professor P.K. Anokhin became head of the department of general physiology of nervous activity.*

In 1938, together with N.N. Burdenko Anokhin organized and headed the physiological sector at the Institute of Neurosurgery, which he directed until 1945.

During the Great Patriotic War, Pyotr Kuzmich worked as a physiologist-neurosurgeon and was the scientific director of a number of large hospitals. He independently operated on wounded people with injuries to the peripheral nervous system and achieved such success in this direction that his operations were demonstrated as demonstration ones for the purpose of training surgeons to work correctly with neurotrauma.

Of great importance for practical medicine, and especially for neurosurgery, were his numerous wartime scientific works, such as “Nerve transplantation (on the replacement of nerve defects after injury)”, “Surgical treatment of large nerve defects”, “Plasty of nerves in military trauma of the peripheral nervous system” systems”, etc.

The very approach to the problem of transplantation and plastic surgery essentially determined the strategic directions for the development of surgery in the second half of the twentieth century.

In 1942 P.K. Anokhin was elected professor of the Faculty of Biology of Moscow State University named after M.V. Lomonosov.

In 1944, with his direct participation, the USSR Academy of Medical Sciences was organized. PC. Anokhin became its full member (1945). Since 1949, he has been director of the Institute of Physiology of the USSR Academy of Medical Sciences.

In the summer of 1950, the infamous Pavlovian session broke out (not scientific, but politically programmed), which set back the science and practice of biomedicine for decades, discrediting the name of the great I.P. throughout the world. Pavlova.

This event was played out according to the scenario of the “black” session of the VASKhNIL in 1948. This happened under the protectorate of the “father of nations” V.I. Stalin. There were corresponding nominated performers who then took a monopoly in science. I.P. Pavlov was presented by them as a bright and frozen face, a step to the right or left from his teaching is death. Everyone who really developed this teaching - P.K. Anokhin, L.A. Orbeli, I.S. Beritashvili, A.D. Speransky and many others were discredited, deprived of the opportunity for full-fledged research and teaching activities, and removed from their positions (at best). So, for example, P.K. Anokhin was removed from all positions and was able to find work only in Ryazan, taking the position of head of the department of normal physiology at a medical institute.

The political situation prompted Pyotr Kuzmich to publish in 1952 in a physiological journal an article “On the fundamental essence of my mistakes in the development of the teachings of I.P. Pavlov and ways to overcome them.” He later wrote that dialectical materialism is able to prevent a scientist from erroneously rejecting his work “insurmountable for us from an ideological point of view.” Thus, Pyotr Kuzmich wanted to live and work, and for this it was better to be a conformist, and not a murdered fighter.

The acceleration of physiology was not as destructive as genetics, and after the death of the leader, everything began to return to normal.

Already in 1953 P.K. Anokhin began to head the department of higher nervous activity at the Central Institute for Advanced Medical Studies, and in 1955 he headed the department of normal physiology at the First Moscow Medical Institute. THEM. Sechenov.

Since 1958 P.K. Anokhin simultaneously headed the department of neurophysiology at the Institute of Normal and Pathological Physiology of the USSR Academy of Medical Sciences, and in 1966 he was again elected as a full member of the USSR Academy of Sciences.

The scientist created a theory of the functional systems of the body as a closed cyclic formation with the presence of feedback information about the result of their action. He identified the key mechanisms of the functional system: afferent synthesis, decision making, acceptor of action results. Result of action, reverse afferentia.

PC. Anokhin suggested that emotions and motivation are obligatory components of a functional system, constituting, together with situational and triggering afferents, the basis for afferent synthesis (1966-1968).

The definition formulated by the scientist of a functional system as a closed self-regulating organization, all components of which interact to achieve a result useful for the body with constant signaling of the result of the action, gave our country priority in physiological cybernetics.

During a visit to the laboratory of P.K. Anokhin in 1960, the founder of cybernetics Norbert Wiener admitted that the team’s research in the field of physiological cybernetics was far ahead of the emergence of this scientific direction in other branches of science.

PC. Anokhin developed the secretory-motor method of conditioned reflexes (1932), introduced the study of the autonomic components of the conditioned reflex as a necessary criterion for the state of the animal, gave a new interpretation of the mechanism of formation of internal inhibition (1935-1958), and clarified the main function of the frontal lobes in goal-directed activity (1949). The scientist was one of the first in the Soviet Union to use electroencephalographic techniques to analyze conditioned reflex reactions and subtle electrophysiological techniques to analyze the patterns of propagation of excitations of various modalities across

afferent nerve. PC. Anokhin established the specific systemic nature of ascending activations of the cerebral cortex during reactions of various biological qualities (1956-1962), put forward a new idea about the nature and composition of the evoked potential (1960-1964).

Formulated by P.K. Anokhin, the principles of compensation for impaired functions are widely used in the clinic to manage neurorehabilitation processes in neurology and neurosurgery.

PC. Anokhin clarified the mechanisms of adaptation and resistance of the body in extreme conditions and clarified a number of issues regarding the pathogenesis of the neurogenic form of hypertension (1960).

In 1968, the scientist was awarded a gold medal named after I.P. Pavlov for a series of brilliant works on the physiology of the central nervous system, for the development of a new direction in neurophysiology - a systematic approach to the functional organization of the brain. In the USA, at a congress of physiologists, he gave a brilliant report “Convergence of excitations on a neuron as the basis of integrative brain activity.”

This idea of ​​convergence and integration of various reflex pathways is akin to A.A.'s law of dominance. Ukhtomsky.

Thus, P.K. Anokhin stood on the foundations of such giants as I.P. Pavlov, V.M. Bekhterev and A.A. Ukhtomsky.

It should also be noted that the scientific works of Academician P.K. Anokhin’s works are distinguished not only by the huge number of completely new original facts and research methods, but also by the boldness of the judgments expressed. He reacted sensitively to everything new, and the gift of foresight allowed him to predict the most fruitful paths for the development of the physiology of the central nervous system.

PC. Anokhin created a large scientific school, widely known both in our country and abroad. Under his leadership, 25 doctoral and 180 candidate dissertations were completed.

At different periods of his life, he was a member of the Scientific Medical Council of the People's Commissariat of Health, a member of the Presidium of the USSR Academy of Medical Sciences, editor of the physiological department of the Great Medical Encyclopedia, member of the editorial board of the Physiological Journal of the USSR named after I.M. Sechenova and others.

Academician Anokhin’s personality attracted colleagues and students to him, who were impressed by the scientist’s human qualities: honesty, kindness, attention to people. Snobbery and arrogance were alien to Pyotr Kuzmich. Of course, he did not allow everyone into his inner world, but those who received this honor were amazed at the spiritual wealth and subtlety of his nature.

Thus, according to the memoirs and stories of those who worked with him, Anokhin the scientist had an amazing sense of the whole, the ability to separate the essential from the secondary, he was characterized by a “disciplined imagination.” As befits a true scientist, he knew how to present ideas without feeble-minded fear for their priority, and never sought to put his name with the author of the work he supervised. PC. Anokhin was an excellent teacher. He delivered his lectures brightly and sweepingly, captivating the audience with logic and imagination.

The creative life of Pyotr Kuzmich can be described as a vivid example of the collaboration of theory with practice, experiment with the clinic. He also collaborated with N.N. Burdenko, studying injuries to the peripheral nervous system, and with A.V. Vishnevsky, developing the physiological aspects of anesthesia and postpulmonectomy syndrome.

PC. Anokhin was also friends with foreign scientists and widely quoted them back in those years when this was called sycophancy and cosmopolitanism.

Until the last days of his life, Anokhin worked until 10-12 pm, reduced summer vacations to a minimum and never used them fully.

Pyotr Kuzmich united people, built bridges between scientific schools in different cities. “He stood on Sharik,” said N.I. Vavilov on the behavior strategy of a real Scientist.

It is interesting to know that Academician P.K. Anokhin did not interrupt contacts with our city, in particular, with the medical institute: he sent telegrams on the institute’s anniversaries, held meetings with colleagues from the department of normal physiology (photos of these meetings are carefully preserved at the department of normal physiology and in the NSMA museum).

On June 29, 1978, a memorial plaque with a bas-relief of Academician of the USSR Academy of Sciences and Academy of Medical Sciences, Lenin Prize laureate, founder of the Department of Normal Physiology of the State Medical Institute, Pyotr Kuzmich Anokhin, was unveiled on the facade of the Main building of the Gorky Medical Institute.

One of the most characteristic features of modern physiology is the attempt to criticize the enormous empirical material obtained mainly through analytical research, and to create a synthetic picture of the complex dynamic processes occurring in the whole organism.

Instead of the existing crude mechanical schemes trying to explain the dynamics of the life process, a wide variety of synthetic concepts are put forward in world literature, in most cases having clearly vitalistic tendencies.

For example, the theory of organicism has recently been put forward (organismic theory), which makes an attempt to embrace the entire diversity of individual physiological manifestations from the point of view of the coordinating influences of the organism as a whole. An ardent defender and even, perhaps, the founder of this theory is Ritter. In his book "Organismic conception etc" he tries to analyze the influence of the organism as a higher synthesis on the particular manifestations of its function and comes to the conclusion that in fact we cannot find a single physiological statement that could be considered in isolation. All appearances are connected by the organism as a whole and occur under its constant control. In addition to this direction, recently a movement known as “holism” (from the English word whole) has begun to enjoy success in American literature.

Embracing broader problems of organic and inorganic processes, holism also seeks to replace the dominance of analyticism in natural science with the dominance of a holistic manifestation of all natural processes. Undoubtedly, there are positive aspects to these trends, consisting of a natural reaction against the once revolutionary mechanistic view of physiological processes. But in them there is such a big tilt towards the vitalistic understanding of integrity that the correct proportion between analysis and synthesis is already lost, and the synthesis itself grows into a kind of categorical imperative, approaching entelechy.

In the field of physiology of nervous activity, in recent years a number of points of view have also been put forward that are aimed at revising the hitherto dominant reflex theory and replacing it with more complex concepts. All these ideas about the field of nervous activity are, in fact, only varieties of the Gestalt theory, which is currently beginning to penetrate all types of research into biological phenomena. Proponents of this theory in the physiology of nervous activity are mainly American and German neurologists, and some of them are known to the reader from translations of their articles that recently appeared in our press (Goldstein, Bethe, Weizsacker). Their general influence on the fate of modern neurology has been noted repeatedly in a number of articles; in this essay, these works will be considered mainly in connection with the authors’ attempts to combine central and peripheral factors of nervous activity into a single complex.

These attempts help us go beyond the traditional theory of centers when explaining complex complexes of nervous activity and build a worldview based on the constant mutual connection of central and peripheral processes. In this new formulation of the problem, the perceptive peripheral apparatuses and working response organs, together with the central nervous system, constitute a dynamic unity, in which only in individual cases can we speak with certainty about the dominance of one or the other.

Naturally, by introducing the constant regulatory and integrating role of peripheral apparatus into the system of nervous activity, this new point of view largely breaks with the traditional recognition of the prerogative of the central nervous system in the regulation of nervous activity.

Such a restructuring of the point of view on nervous activity completely changes the position of researchers in all areas of neurology, changes the very prospects of research and requires broad familiarization with a number of boundary problems. That is why the abandonment of the theory of centers, which is largely simple and schematic, and therefore convenient for manipulation, occurs very slowly and reluctantly. The situation is greatly aggravated by the fact that, in fact, today we do not have any definite, universally accepted formulation of the concept of the center. This state of affairs is strange if we take into account that everyone begins to talk about nerve centers from their student days. Later, however, people become so accustomed to this word, which has no precise content, that rarely does anyone set out to establish its relationship to the accumulating facts of nervous activity.

The formulation of the problem of center and periphery, and at the same time a more specific formulation of these concepts, was first given several years ago by Bethe (1931). Skillfully selecting enormous experimental and clinical material obtained over half a century by different authors and in different directions, he brought it together into a coherent system, giving it a certain meaning, which the researchers themselves often did not see. The Soviet reader is already familiar with Bethe’s general principled position from his article published in one of the issues of the journal “Advances in Modern Biology.” In this article, I set myself the task of giving a more detailed view of this problem, deepening its physiological argumentation based mainly on the experiments of my colleagues and trying to understand this problem in the light of the onto- and phylogenetic development of the relationship between central and peripheral processes.

CENTER AND ITS CHARACTERISTICS

Before moving closer to those dynamic ideas that are now emerging under the influence of a number of recent experiments, we will try to give a more or less accurate decoding of what is usually understood by the “center” by supporters of the exact localization of nervous functions.

Speaking about centers, we primarily mean those nerve formations that are, in the full sense of the word, the brain ends of various peripheral nerve trunks. These nerve trunks are the direct connection of any peripheral organ, perceptive or effector, with ganglion elements, which may lie in various parts of the central nervous system.

As an example of such centers, the centers of individual muscles, the center of the vagus nerve, the centers of the spinal cord that perceive skin irritations, etc. can be given. These centers, so to speak, in the first and last instance, communicate the central nervous system with the periphery; they are the first take on all the effects of external and internal stimuli on the body (“representant” according to Uexkuhl) and they also take on in their final form every nerve impulse, no matter how complex it may be and no matter how complex the path it has previously passed (“final way" by Sherrington). As follows from the above formulations, these centers of the first degree are most dependent on peripheral organs and most constantly exert their influence on them.

But the described centers, being the entrance and exit gates for a variety of impulses circulating throughout the central nervous system, are interconnected through a number of intermediate nerve formations in which the circulation of the nerve impulse, due to their multilateral connections, appears to be especially complex. From the point of view of the theory of centers, these intermediate nervous formations, which include mainly the subcortical formations and the cortex, complicate the course of the nervous process and the nervous manifestations of the body only due to their overall functioning. At each individual moment and at different moments of functioning, no matter how complex the existing excitation complex may be, the individual centers included in this complex never lose their specificity. In the most schematic form, this point of view is expressed in the position that any complex nervous act represents the sum and mutual influence of individual reflex arcs. It is quite obvious that, despite the apparent variability in the behavior of animals and humans, the above point of view introduces some predetermination into the degree and nature of the participation of individual centers. The properties of these centers, no matter how complex their combinations are from the point of view of the theory of localization, are strictly anatomically fixed. The changeability and formation of a new type of functioning of the nervous system, as, for example, is manifested in the formation of a conditioned reflex, does not exclude the above provisions of the theory of centers, for it is explained only as a new closure between completely predetermined nerve centers.

From what has been said, it is clear that if the nerve centers always retain their Specific properties, then those association fibers that make connections between them largely reflect the same specificity. They always carry only the type of excitation that is formed in the centers they connect.

From these provisions it naturally follows that the entire variety of activities of the central nervous system is the result of the variety of connections, mutual exclusions, etc. of individual nerve centers and their connections without them losing their specificity.

The functional specificity of certain nerve formations is an initial property that does not change under any circumstances. These are the basic provisions that are necessarily adhered to, consciously or unconsciously, by every clinician and neurophysiologist who takes a strictly localization point of view.

In the future, we will try to evaluate this position from the point of view of modern data; now it is necessary to analyze the methods themselves, with the help of which the main provisions of the theory of centers were established.

In the middle of the last century, when the achievements of natural sciences opened up one after another new aspects of the body’s activity, when technical improvements made it possible to dissect any of the organic manifestations into more detailed details, at that time the methods of irritation and extirpation reached a special flourishing in relation to the brain. From the point of view of analytical tendencies, these methods seemed to give an accurate indication of the presence of one or another center in the central nervous system.

In fact, we know that irritation of a certain zone in the cortex produces a contraction of certain muscle groups, which reaches particular isolation in primates and humans, and extirpation of, say, the occipital region leads to the elimination of the ability to distinguish and even perceive visual influences. With the development of histological methods and electrophysiological techniques, it became possible to apply new identifying features to individual localized centers.

The method of studying myelogenesis has made it possible to accurately establish that certain complex nerve formations are myelinated simultaneously, jointly, and often in isolated islands. This method emphasizes a specific localization of nerve centers (Dogel, Langwoursy). Recent recording of electrical oscillations in the cerebral cortex using an oscilloscope seems to convince us that certain cortical areas have a very specific electrical oscillation curve.

Whenever any perceptive apparatus functions, the zone corresponding to it in the cortex exhibits “electrical disturbances,” the nature of which has precise boundaries corresponding to both the physiological and cytoarchitectonic properties of this zone (S. A. Sarkisov).

Recently, in the laboratory of Prof. A.G. Gurvich found another very interesting way to characterize the nervous process, and at the same time its localization. It turned out that a nerve impulse of a certain quality and a certain intensity, be it in a peripheral nerve or in the brain, has a very specific mitogenetic effect that is perceived by yeast cells. As you know, the laboratory of prof. A.G. Gurvich found a way, thanks to spectral analysis, to qualitatively characterize this mitogenetic effect. With regard to the central nervous system, it has been shown that stimulation of the optic nerve produces spectra of varying quality in n. Optik i, visual thalamus and in the optical zone of the cortex.

Thus, in the mitogenetic effect one can see a powerful means of characterizing the process of excitation, “leaving far behind the methods of modern biochemistry in its sensitivity and especially in its physiology” (A. G. Gurvich, 1934).

This is the arsenal of means and methods that modern natural science has at its disposal to establish the localization of nerve centers. Of course, one cannot deny the importance of these methods for detecting one or another pattern of the nervous process, but it is also undeniable that they should be regarded only to the extent that they can help resolve the problem of localization. Meanwhile, these methods are undoubtedly overestimated by supporters of the theory of nerve centers.

If, when we stimulate any area of ​​the cortex, we obtain one or another motor effect in the periphery, then this cannot in any way be interpreted in the sense that we have found a motor center coordinating complex motor acts. The pyramidal cell zone is the outlet for the nerve impulse to the motor segments, i.e., to some extent, to use Sherrington’s language, a “check to bearer,” although he uses this expression in relation to the spinal cord.

Therefore, the excitation of any group of cells can only be a manifestation of the results of a complex complex of nervous processes occurring before them, but in no way an indicator of the construction of an entire motor act. Undoubtedly, this direct connection of the motor zone with the effector motor apparatus greatly helps, for example, a neurologist to accurately diagnose the source of destruction, but still this is only the destruction of the FINAL LINK in the complex picture of the circulation of the nerve impulse and nothing more. That is why all diagnostics of organic disorders of the central nervous system use “inclusive of the final motor component (movement of the limbs, oculomotor function, swallowing, etc.).

In the same way, the method of extirpation gives no more than the method of irritation. If extirpation of, say, the visual area of ​​an animal leads to complete or partial loss of vision, then this does not mean at all that we have excluded the “visual center”. Bethe correctly notes that, despite the fact that when the optic nerve is cut, the entire visual function is eliminated, no one would think of asserting that the “visual center” is located in the optic nerve, and yet, in relation to the visual area of ​​the cortex, most physiologists make this conclusion .

By removing the visual area, we remove the first instance in the circulation of the perceived impulse through the central nervous system, and therefore it is natural that in the future we do not see any usual indicators of this circulation (movement, secretion, etc.).

Thus, in the case of the final effector area and in the case of the Initial perceiver, we interrupt the circulation of the Nerve impulse at its extreme links, closely connected with the peripheral organs - this is the only way that all experiments carried out using the stimulation method can be evaluated. and by the method of extirpation.

Based on these considerations, other localization indicators can be assessed accordingly, such as the electrocerebrogram, mitogenetic effect and myelination. If, when the retina is irritated in the visual area of ​​the cortex, electrical oscillations can be recorded that are different from those in other areas, then this only indicates that the visual area has a more intimate, closer relationship to the passage of the light impulse than other areas.

Of course, an oscillogram can help us differentiate the participation of individual parts of the brain in any function, outline more or less accurately the boundaries of individual zones, but this does not mean establishing the localization of a certain complex function.

In the same way, myelination, representing not only an indicator of the functioning of the system, but also partly the biological prospects of a given organism (overgrow according to Koghill), can rather speak of the preferential circulation of the nerve impulse at a given stage of ontogenetic development. In addition, although Langwoursy insists on a certain parallelism between the beginning of the function of nervous tissue and the process of its myelination, voices have recently been heard indicating that such parallelism is extremely relative and in no case can serve as any argument in favor of localization functions (Anguio-Gonsalez, 1932).

All of the above applies equally to the mitogenetic indicator: it serves as an indicator of what qualitative changes a nerve impulse undergoes as it circulates through individual parts of the central nervous system, but cannot help in any way in identifying the centers that localize and organize this or that complex function.

Summarizing all these arguments regarding the methods of studying nerve centers, we must say that basically they always deal either with an increase in function in the final link, or, most often, with an interruption in the course of excitation at any of the numerous points of the complex nerve impulse . That is why the closer to the final links of this complex chain, and therefore to the peripheral organs, the destruction is localized, the more demonstrative and less compensated are the resulting dysfunctions. And vice versa, if the destruction is localized somewhere in the middle of this entire complex intertwined system, the more difficult it is to detect the direct result of this destruction, and therefore to localize the corresponding center. If we take into account (for example, in experiments with extirpation) the powerful influence of the inevitably occurring diashisis (Monakov, 1912), then all this makes the result of extirpation completely unreliable.

TEACHING ABOUT CENTERS AND DIFFICULT BEHAVIOR

AFFERENT CENTERS OF THE LARGE HEMISPHERE CORTEX

If we take complex complex behavior as an indicator of the presence of nerve centers, then the doctrine of nerve centers turns out to be especially untenable and runs counter to the factual material of modern neurophysiology. Since, from the point of view of the classical doctrine of nerve centers and their localization, it must be assumed that each center involved in complex behavior has a completely definite and never repeated function, one would expect that the removal of one or another center would lead to uncompensated loss. In fact, in experiments with the elimination, restoration and reproduction of skills, just the opposite happens. The overwhelming majority of experiments in which, after certain extirpations and disconnections, a previously developed skill was either preserved or could be re-formed, convinces that the participation of one or another part of the central nervous system in a complex developed act of behavior cannot be considered irreplaceable and in any case decisive. From the point of view of general compensation, it is comparatively more difficult to compensate for the removal of certain afferent parts of the cortex and the lower parts of the central nervous system in general, but only if we talk about restoring this particular function. General adaptive behavior is restructured in all such cases quite effectively, and this ensures the existence of the individual. Recently, the Lashley school has carried out a systematic study and characterized the afferent function of the cerebral cortex. Although Lashley studied the visual receptive area exclusively, we are interested in dwelling on his conclusions because everything related to visual function is equally applicable to other afferent areas of the cortex. The general conclusion from his experiments, carried out using the method of extirpation and subsequent assessment of “visual” behavior, is that the visual part of the cortex itself, even in such a relatively simply organized animal as the white rat, seems to be more or less definitely localized. These conclusions, given in a number of works by Lashley, somewhat contradict the understanding of his experiments that we have in the Soviet neurological literature (Yushchenko, Kharitonov). The misunderstanding stems from the fact that the relationship between the two functions of cortical tissue: differentiated and equipotential, which Lashley insists on, is not entirely correctly understood.

Here is how Lashley (1931) himself speaks about this: “... the same zone can function sometimes as a highly differentiated system, and sometimes as a single mass.” In various cortical functions there is any degree of specialization from limited exact correspondence of cells to conditions of absolute nonspecificity.

“We do not make a choice between the theory of localization and the theory of decentralization, but we should develop a broader view that would recognize the care and mutual dependence of both types of integration.”

From these quotes it is clear that Lashley does not deny or exclude the special function of certain afferent sections of the cerebral cortex; he only emphasizes that one special function is not capable of explaining the dynamic effect of the cortical mass on all processes occurring in the underlying sections of the central nervous system. This has been shown in numerous experiments.

We will not now dwell in detail on the forms and techniques of his experiments - this will be discussed in a special review of Lashley's latest works. Here it is important to more precisely formulate this additional function of the center, which has recently been supported by a number of other American authors. Indications of this function stem mainly from experiments in which a skill was destroyed only depending on the degree of disruption of the cortical mass, and not on its location. Moreover, the degree of loss and impairment of labyrinthine skill is directly proportional only to the mass of destroyed cortical tissue, regardless of where this destruction occurred. Along with this, Lashley showed that if a rat develops a skill after being previously blinded, it still loses it when the visual cortex is removed, and almost to the same extent as if it had acquired it with the help of the visual apparatus. This and other similar experiments led Lashley to say that the visual zone, along with its specific visual function, also has a non-specific, characteristic of all other zones of cortical tissue, the “non-visual function of the visual zone.” This “facilitation” function represents that “equipotential” property of the brain; cells, which unites the cortex into a single whole.

“We do not know the prevalence of such “facilitating” activities, but our work suggests that the entire cerebral cortex, and perhaps even each part of the nervous system, may, in addition to its specific functions, exhibit such a general “facilitating” effect on other parts. This may to some extent explain the quantitative relationships that exist between the size of the lesion and the correctness of execution, the size of the “relief” depending only on the number of active cells” (Lashley, 1931).

As can be seen from the above, the entire theory of equipotentiality, developed by Lashley, inevitably presupposes the presence of a more or less precisely localized and specific function and it is almost impossible to separate it from the undifferentiated function that constantly accompanies it - facilitation. Later, Lashley clarified his point of view in a number of specially designed experiments and began to talk about the interchange and equipotentiality of the receptor elements of the cortex within certain zones.

“In contrast to the theory of mosaicism of the functional activity of specialized zones, there are indications that within a special zone all parts are equivalent in certain respects and for certain functions.”

“The work shows that, within very wide limits, the absolute properties of the stimulus are relatively unimportant for behavior, and responses are determined by the amount of excitation, which is equally effective no matter to which group of receptor cells within the system it is applied” (Lashley, 1933).

All these conclusions are supported mainly by those experiments by Lashley, in which, despite the removal of 50% of the visual cortex, in both hemispheres and regardless of the location of the destruction developed before, a complex visual skill is preserved. Assessing all these and the above results, he concludes: “they show that the idea of ​​simple pathways from the receptor to the cortex with the establishment of integration between individual neurons there does not at all correspond to the actual picture of the afferent mechanism” (my discharge. - P.A.).

It is not surprising, therefore, that recently in the works of both Lashley himself and a number of other authors, an attempt has been made to find a morphological correlate of the dynamic concepts that follow from the experiments described above. The main question that came under attack from purely morphological research was the question of whether certain points of the retina correspond to well-defined and constant areas of the visual cortex. The question is of fundamental importance, because it is known that the reverse image of any moving object can move across the retina, and yet a person or animal has a very definite and constant idea of ​​​​the shape, size and other properties of the object.

How to combine this fact with the doctrine of precise and fixed localization?
The most complete and systematic studies on the degeneration method were done by Poljak. His data, very detailed, published in 1933, showed that between the retina and the cortex there is an exact zonal correspondence, in some cases truly a point-to-point relationship. Along with this, it was shown that the boundaries of the optical zone in the cortex are clearly limited, so that from their point of view it is impossible to talk about the dispersion of visual elements in neighboring cortical zones. These purely histological observations, based on tracing certain degenerated pathways, seem to speak against the theory of “scattered” nervous elements, which was recently put forward by I.P. Pavlov. According to this theory, along with the nuclear, so to speak, formation of any receptor or effector zone of the cortex, there are also scattered elements with the same function, scattered throughout the cortex far beyond the boundaries of this nuclear zone. We will not now enter into a discussion of whether there are any morphological correlates to the phenomenon of compensation, which was stated in the laboratory of I.P. Pavlova solely based on the functional ghost. It is only important to point out that the contradiction between the theory of “scattered elements” and Poljak’s data can be eliminated on other grounds. Undoubtedly, when the main, let's say visual, nucleus is eliminated, the general generalization of the excitatory impulse coming to the intermediate nuclear formations (let's say, the thalamus) makes it accessible to all remaining parts of the cortical tissue. Whether this will produce a specific visual function of these new elements or whether they will, in a “facilifation” type, clarify and stimulate the work of the subcortical apparatus, as it seems to have been in Lashley’s experiments, this question is subject to further in-depth study. But it seems to me beyond doubt that any process of compensation during the destruction of cortical tissue occurs primarily through enhanced generalization of the impulse in the primary nuclei lying close to the peripheral receptor apparatus, and only through this are new cortical zones activated. Schematically, this can be imagined as shown in Fig. 1.

This is partly indicated by the experiments of our laboratory (P. Fedotov) with the destruction of individual parts of the thalamus. The latter are compensated much more slowly and more difficultly than much more significant destruction of cortical tissue, and this is undoubtedly due to their intermediate positions in relation to the passing afferent impulse. One interesting fact in the study of Poljak, Despite the fact that a certain, precisely adjusted relationship was stated between the retina and the visual cortex, the relationship between the retina and the geniculate bodies turned out to be more diffuse and could not be precisely localized (Poljak, 1933). This indication has great physiological meaning. In fact, how can one imagine that an impulse, starting from a certain point in the retina, corresponding to a certain zone of the cortex, in the intermediate nuclear formations is diffuse, not confined to any specific parts of these formations?

These relationships once again emphasize that when taking into account the effect of any stimulus on the cerebral cortex, one cannot help but take into account very interesting relationships and propagation of the impulse that arise already in the first nuclear instances along the path of its circulation. But more on that below. Setting out to investigate precisely this part of the problem, Lashley most recently published the seventh report on "Machanism of vision", in which he presented a number of histological evidence that the crossing and non-crossing fibers of the retina are localized more or less constantly in certain primary optical centers. For example, crossing fibers do not end in the lateral geniculate bodies, but pass into the superior colliculus, while non-crossing fibers, on the contrary, end preferentially in the geniculate bodies. These data seem to contradict the results of Poljak, but in essence they do not say anything about the communication of the impulse, but only state the different behavior of crossed and uncrossed fibers of the optic nerve (Lashley, 1934).

Thus, the neurophysiological characterization of afferent centers faces a big problem: how to combine morphological data, indicating a more or less precise fixation and specialization of the nerve elements of the cortical zone, with neurophysiological data, which speaks in favor of the fact that this specialization does not play a decisive role in the dynamics of the function the entire central nervous system. On the contrary, where we are talking about the interchange of individual parts of the visual zone in a relatively primitive function, they all turn out to be equivalent (equipotential). The same problem faces neuromorphologists - to find out whether there are any morphological correlates for a general neurodynamic function such as facilitation or whether it is performed by the same nervous elements that also carry a special function. And if this last circumstance is present, then this functional polymorphism of cortical cells should be fully studied. Undoubtedly, this problem is very difficult and can only be solved by very subtle methods, if it can be solved at all. In this regard, layer-by-layer extirpation methods become especially important, making it possible to remove individual morphological layers of the cortex (from above) using certain fixing agents (Dusser de Barrene, 1934).

Regardless of whether a morphological correlate of the facilitating function is found, we can assert that physiologically both of these processes in each individual manifestation of nervous function are so closely linked to each other that we can speak of their complete unity. That is why the destruction of certain areas of the cortex, seemingly not interested in the development of this skill, nevertheless affects the subtlety of the animal’s behavior and even to no less a degree than the destruction of a special zone. Thus, speaking about the afferent centers of the cortex, we must remember that essentially nothing corresponds to this concept. There is a preferential distribution of the impulse in the first instances of its circulation, associated with the closest relationship with the perceiving organ, but already in these same first instances it becomes generalized thanks to universal connections with the cortex of such intermediate formations as the thalamus. The recognition of “scattered” elements essentially leads to the same conclusion, for if each of the foci having a special function has its own elements in the remaining parts of the cortex, then, taking into account all the special formations, we will obviously have to accept that in any function the cortex behaves as a single whole. We must not forget, of course, that the complication of animal organization in the process of phylogenetic development significantly changes the relationship between the two functions of cortical tissue described above - specialized and equipotential. At the lower stages, which show very weak development of the primary cortical tissue (hupopallium), the main operating factor is undoubtedly the facilitating function of the cortical tissue, while in highly developed mammals, and especially in humans, the moment of specialization is obviously the leading factor. This is precisely what can explain all the enthusiasm for schematic ideas about the localization of functions in the cortex. Evidence of this point of view can be provided by experiments with irritation of the motor zone in a crocodile. A fairly large part of the cortex, lying closer to the frontal part, gives general diffuse movements of the whole body, but no hints of local isolated movements can be obtained ( Herrick, 1926). Everyone, of course, knows that this experiment is easily possible in higher vertebrates and especially in primates and humans.

Summarizing all the material and considerations on this matter, we must say that it hardly makes any physiological sense to talk about the afferent centers of the cerebral cortex. This conclusion does not exclude its special significance for some areas than others in any nervous process, but these special areas themselves can only be considered as certain paths and points of propagation of the afferent impulse, and the points, of course, are only initial and therefore carry within them the inevitable results of the specificity of the impulse and or other receptive surface. In addition, these foci and zones associated with various perceptive organs cannot function only specifically; they inevitably have a common equipotential function with other zones, facilitating the occurrence of subcortical processes. But what is especially important for a number of schematic constructions of the reflex theory is that the cortical afferent zones honor in the perception of external stimuli not according to the principle of linear propagation of an impulse in the form of a reflex arc, but special and neurodynamic formations, constantly influencing the indexing and external resolution of processes in complex subcortical formations. An external impulse entering the brain does not go linearly in one direction, but, as Sherrington put it in his recent lecture, “oscillates” between separate ones; key points, establishing peculiar local relationships between excitation processes (Sherrington, 1933). Adrian’s work with recording biocurrents in the cerebral cortex seems to indicate this.

It is very likely (and there are indications of this from the phylogeny of cortical tissue) that the afferent impulse flows into the cortex along collaterals, and, once entering the cortex, it subsequently makes numerous circular circulation between the cortex and subcortical formations until one or another occurs external manifestation.

Considering the speed of propagation of excitation and the small size of the neurons of the large brain in most animals, we must say that the periods of these circulations must be extremely short. Hence it is clear that any scheme representing a linear, one-way directed propagation of excitation is incorrect. In practice, we must imagine that in response to any stimulus, the brain turns into a complex dynamically intense system, in which the periods and sizes of nervous oscillations between individual points of the whole brain are so far beyond the limits of our modern technical capabilities (see below). Nevertheless, knowledge of the afferent nodes of the brain gives us the opportunity to talk about the preferential distribution and circulation of a nerve impulse caused by any external stimulus.

EFFERENT CENTERS

We have already touched briefly on the main effector formations, which have long been known as “motor centers.” They can be called centers only in a very conditional sense of the word. They do not concentrate in themselves the control of any of the motor manifestations, but serve only as a final field, which receives a nervous process that has undergone complex and analytical processing somewhere before the motor centers. From this point of view, it is very interesting to note the special relationship of the cortical motor area to motor acts of varying complexity. The general characteristic of the cortical motor area is that it is credited with concentrating all processes of developing motor skills. This means that under the influence of external stimuli, any acquired motor act must ultimately manifest itself through the cortical motor area.

It is in this that the statement that acquired reactions are a function of the cerebral cortex gains its meaning. It follows, finally, that if the entire cortical motor zone were removed, the implementation of acquired motor reactions would become impossible.

Data from Wagner (1905), V.M. Bekhterev (1915), Gierlich (1913), N; I. Krasnogorsky (1915), V.P. Protopopov (1925) show that the preliminary development of a motor skill is eliminated if the motor zone of the corresponding side is extirpated in animals. Similarly, Pike and Chappell (1930) showed that the motor training ability of the cat is significantly reduced with very slight impairment of the motor area. Lashleu explains the lack of motor function in all these experiments by the fact that the motor zone of one side was removed and, thus, the general motor adaptation could be successfully carried out at the expense of the remaining healthy limb, i.e. the stimulus for restoring the lost motor skill was not enough .

That these considerations may be important is indicated by the experiments of Trendelerberg (1915) and Oden and Fanz (1917). When the motor areas of both sides were removed or destroyed, the restoration of acquired motor skills was much faster.

Likewise, still old experiments Marigue (1885), Exner and Paneth(1889) showed that if a circular vertical incision around the cortical motor zone is used to separate it from the rest of the cortex, the same effect is obtained as is observed with complete extirpation of the motor zone. But along with these localization data, indications were received a long time ago that the motor zone of the cortex is by no means a decisive factor in the regulation of the restoration of an acquired motor skill. Already Golz (1881) in his experiments received a hint of the non-absolute significance of motor zones.

Recent experiments, carried out with the special purpose of testing the absolute importance of motor zones for the development and retention of acquired motor skills, have given just the opposite results. Graham Brown (1916), Trendelenberg (1915), Lashleu (1924) showed that extirpation of motor areas does not eliminate the possibility of developing a new motor skill and restoring an old one, although the sufficiency of removal was confirmed in some cases by histological studies. Lashleu (1924) developed a complex motor skill in monkeys and showed that after sequential extirpation of one and the other motor zone, the skill is eventually carried out without any special changes. In our laboratory, this problem was tested on active selection material on a machine with two opposing feeders. When analyzing the developed motor act, we encountered the need to establish to what extent the participation of the cortical motor area is decisive in the implementation of this skill. The experiment was carried out in the following sequence: first, in response to certain conditioned stimuli, the animal was accustomed to eating either on the right or on the left side. In each individual case, when a conditioned stimulus was given, the animal was placed under conditions of active preference for one side or another of the pen (P.K. Anokhin, 1932). Active choice was detected with a large number of trials, movements, and the animal looking at both sides of the pen during the action of the main stimulus. The simultaneous removal of motor zones on their hemispheres showed that several days after the operation, without any preliminary training, despite the relatively disordered motor skills, the correct choice of the corresponding side appeared from the very first test of the conditioned stimulus (experiments by N. Chernevsky).

These experiments demonstratively showed that the data of Lashleu and other authors indicate a very regular phenomenon that can be obtained in various situations of developing motor skills.

How to understand such results?
How can we understand the fact that the cortical motor area does not play a decisive role in the development of conditioned motor skills? Explanations for these results could come from three directions. Firstly, one might think that the motor function, which previously belonged to the motor zones of the hemispheres, transferred after their removal to the subcortical formations. The most likely is the transition of this function to sections that are phylogenetically close to the cortical formations.

Lashleu (1924), having received complete restoration of motor skills after extirpation of the cortical motor zone, then destroyed a significant part of the nucl. caudatus on both sides. Experience has shown that the restored skill was largely preserved. Thus, in this case the subcortex did not replace the cortex. However, the experiments of the same Lashle u, which proved the restoration of receptive visual function through the nearest subcortical formations (thalamus), convince us that the question of the possible substitution of cortical function through subcortical formations should undergo careful and multifaceted study. Nevertheless, regarding the substitution of motor function, it is obviously necessary to come to a negative conclusion.

The second explanation was proposed by Fanz (1907) and was especially fully developed by the school of Acad. I. P. Pavlova. Fanz believed that after removal of the motor zone, the initially lost function passes to the limittrophic formations of the cortical mass, which quite adequately restore the lost motor skill.

Laboratory of acad. I.P. Pavlova admits that the distribution of motor nerve elements is not limited to the so-called motor zone; on the contrary, they turn out to be scattered throughout the entire cortical mass, like the afferent cells that we have already mentioned. (As Academician I.P. Pavlov puts it, there is “mechanical immunity.”) This understanding of “scattered” elements, based on purely functional considerations, is fully consistent with cytoarchitectonic data on the structure of limittrophic formations. According to morphological specialists, the idea of ​​scattered motor elements may well be supported by cytoarchitectonic data (personal consultation of the author with Prof. L. Ya. Pines).

Recently, this problem, attacked from different sides, has received a slightly different light thanks to a number of special experiments. The question is posed in this way: does compensation occur due to scattered elements or does the entire remaining mass of the brain, no matter what elements it has, take part and achieve compensation for the lost function. It is quite obvious that here two fundamentally different points of view collide: the function is compensated either by elements with a certain ability fixed to them (scattered), or the entire central nervous system rearranges the remaining elements in such a way that, regardless of the function fixed to them, they are more or less less fully restore the lost abilities of the body, and in this second assumption it is assumed that the remaining undestroyed cortical mass can participate in the restoration of the effect through non-specialized activity.

To test Fanz's assumptions mentioned above, special experiments were carried out by Leyton and Sherrington. The experiment was set up in this way: first, a skill was formed, then the motor zone was extirpated, and when the motor function was restored, the areas of the cortex closest to the initially extirpated zone were again removed. Observations on animals showed that the restored motor functions were not eliminated in a more or less noticeable manner.

The results of these experiments force us to understand the idea of ​​scattered elements somewhat differently.
In fact, if we accept that the dispersion of cellular elements of a certain function occurs with a certain gradient from the main nucleus, then, obviously, the cortical zones closest to this nucleus should have taken over the main part of the restored function. This means that with secondary extirpation of neighboring zones, the restored motor function would have to be lost almost completely. In reality this was not the case.

All these data make us think that the restoration of this or that activity after extirpation of a specific zone cannot be explained by the participation of specific elements of the cortex, which are available, as it were, “in reserve.” Data Colz (1881), Lashley (1929), Bornstein (1932) and others convince us that there is some dynamic equipotentiality of the nervous elements of different zones, which can be involved in the compensation process, regardless of where, in what specific zone the violation occurred. This gives rise to a whole series of experiments carried out by various authors, which are generally aimed at resolving the question: is there any definite and regular correspondence between the destruction of the cerebral cortex and the loss of one or another function. The general conclusion from these experiments is stated by Lashley: “the significant result of these observations is that limited destruction does not reduce any one relevant function, leaving others intact, but reduces the capacity for all kinds of functions” (Lashley, 1933).

“Motor acts, once acquired (for example, opening the bolt of a drawer), can be produced directly by motor organs that were not associated with these acts during training” (Lashley, 1933). All the above and many other experiments omitted due to lack of space indicate that there is no way to explain complex motor coordination and the development of motor skills, keeping only within the limits of the old localization “centrist” positions. The idea of ​​strict specificity of nerve elements, which arose in connection with the development of our histological knowledge of the central nervous system, must be changed and supplemented from the point of view of those synthetic ideas that arose on the basis of neurodynamic experiment. The fact that in our Soviet neurological literature synthetic views, for example Lashley, are criticized on the basis of a completely correct idea of ​​​​the line of its development, should not, however, eliminate from our field of view sometimes very valuable factual material.

One might think that if the central relationships are multifaceted and easily compensate for any violations, then the pathways along which the ready-made impulse, formed in the central nervous system, goes should have been strictly specific. In fact, this is not the case either. Already the old experiments of Starlinger, Osawa, and Rothman with bilateral transection of the pyramidal tract and subsequent restoration of motor coordination speak against this. From the point of view of compensatory processes, the Osawa experiments deserve special attention. By making successive sections of the columns of the spinal cord, first on one side, then on the other, he received the restoration of completely adequate coordination each time.

It is quite obvious that in this case the “specificity” of the pathways and cellular formations changed with each new operation. Concluding his assessment of a whole series of experiments of this kind, Bethe says: “From this it is well understood that certain fiber connections serve one specific, if not exclusive, purpose, just as a steam locomotive runs along a very definite path between large stations. If this most convenient and most passable path is destroyed, then a new one will be formed more or less quickly from existing other paths, which, however, is less convenient and passable. Each point is connected to every other by many paths - direct and indirect, but there are no specific paths that would represent one single connection (Bethe, 1931).

Thus, as a general conclusion from all this material and its interpretation, we can say that in an adult, mature animal, with which physiologists mainly experimented, under normal conditions there is a certain predominance of one pathway and one center over the other. But this preference is not an indicator of the absolute specificity and immutability of nerve formations; it changes whenever the normal circulation of an impulse is disrupted: this predominance is dynamic in nature. From this it is clear that the neurodynamic view does not assert the naked equivalence of brain elements, as some claim.

From the denial of the absolute specificity of nerve formations from a dynamic point of view, we move on to the problem: how and thanks to what operating factors in the process of ontogenesis this preferential circulation of the impulse and the special characteristics of the nodal points of the central nervous system present in an adult animal developed.

How and due to what does function compensation occur when certain areas of the central nervous system are destroyed?

While the localization theory relies almost on predetermined pathways and centers, the dynamic theory of nervous activity raises the question of the historical determinism of the structure and function of the central nervous system. In short, the question of the ontogenetic development of nervous activity, starting from the very first moments of embryogenesis, is decisively raised. The last chapters of our review will be devoted to this issue.

THE PROBLEM OF SPECIFICITY OF LOWER CENTERS

If the general dynamic characteristics of the cerebral cortex and complex acts of animal behavior led us to the conclusion that there is no absolute specificity of nerve formations, then experiments with changes in the specific properties of lower nervous formations, such as, for example, the spinal cord and medulla oblongata, are all the more important and fundamental. Here, in these sections, the function belonging to certain nuclei has long been considered innate, hereditarily fixed and not subject to any change. In essence, each of the experiments in this area, aimed at testing the centers, in itself was capable of causing a revolution in our neurological ideas, but most of them were consigned to oblivion, and the rest, although they entered modern physiology, did not have any noticeable impact on the reflex concept provided. Meanwhile, their real significance lies precisely in the fact that they call into question the principle of reflex with all its characteristic features (P.K. Anokhin, 1933).

This case is not unique in the history of science. Many of her most valuable achievements remained in the archives until the appropriate situation matured, reaching the level of forgotten experiments. The reflex concept has become too firmly entrenched in the minds of physiologists, and some even striking results that speak against it simply went unnoticed. The basic principle of all the experiments in this series is that a certain nerve center, historically associated with certain peripheral organs, is artificially given other peripheral organs that are unusual for it. Technically, this is accomplished by suturing dissimilar nerve trunks. This method of testing the specificity of nerve centers is of great convenience: it allows you to pre-assign the desired artificial heterogeneous connection. Historically, this method was used mainly for surgical purposes, and only very few experimenters did it for the purpose of a physiological experiment. Experiments with cross-linking of nerve trunks have shown that if dissimilar nerves are connected to peripheral organs that are unusual for them, then in the end this leads to the fact that both the “center” and the “periphery” work together and enter into adequate relationships. This “triggering” certainly occurs with the loss of the initial specialization of the nerve centers involved in this process. It is regrettable that neurophysiology undeservedly ignored experiments of this kind, since the first of them, and perhaps the most important of all, was carried out by Flourens back in 1848.

By cross-linking the nerves that give extension to the rooster's wing with the flexor nerves, he ultimately achieved the restoration of the original function of the wing. This experiment, carried out, as Bethe puts it, with the “explicit intention of testing the centers,” was carried out in such a classical form in the Essence that all subsequent experiments with the crossing of the somatic trunks could not give anything better. Even Bethe’s own experiments, carried out in 1912 with the crossing of opposite sciatic nerves, essentially only show that a violation of the specificity of centers can occur not only between flexion and extension on one side, but that centers of the opposite value can also be replaced.

Experiments Beth (1912), Kennedu (1911), Osborn and Klivingston (1911), carried out in the same direction, showed that the process of replacing the specificity of the centers ends with the corresponding inclusion in the function of the motor zone - the cerebral cortex. If, after complete restoration of motor function in the area of ​​the nerve chiasm, the cortical area corresponding to the operated side is irritated, then contractions of the muscles that have received altered innervation are obtained. It is not entirely established whether, under these conditions, localization relationships are always preserved in the cortex corresponding to the newly innervated areas, but in any case, control of this area is always concentrated in the motor zone of the corresponding side (Osborn, Klivingston, 1911).

The above experiments are the main ones that established the possibility of changing the specificity of spinal centers. Of course, the matter is not limited to these experiments: a huge number of operations were performed in the surgical clinic, which, pursuing purely practical goals, actually rest on the same theoretical foundation (Bayer, Perce, etc.).

An important conclusion from these experiments is, first of all, that the spinal centers are not specific, with a function fixed once and for all, but can, within a very limited time, change it to the diametrically opposite one. This fact in itself speaks against the generally accepted point of view that the entire central nervous system can be divided into sections with individually varying activity and into sections with hereditarily fixed activity.

Based on the fact that, despite the variability of the spinal centers, with the final establishment of correct coordination, the cerebral cortex enters into old localization relationships, one might think that all this restructuring of the spinal centers occurs under the influence of the cerebral cortex. In essence, these are precisely the conclusions drawn by all those authors who tried to establish a connection between the reintegrated portion of the spinal cord and the cerebral cortex. We verified this fact in a more subtle experiment using the method of conditioned reflexes. The animal had a cross n. obturatorius and n. femoralis on both hind legs. As a result of this crossing, the hind limbs of the animal developed a new coordination connection with the entire locomotor apparatus for a relatively long time. This development of coordination connections by posture could be divided into four periods.
The first period - an animal deprived of four nerve trunks, with significant paralysis of the hind limbs, compensatoryly adapts its locomotion to this defect. Here we can only talk, perhaps, about the acceleration of the regenerative process due to constant central impulses that carry out the compensatory process, but the regeneration process itself is not yet completed.
The second period - the nerve fibers of the central segments reach the effector apparatus, i.e. the muscles. The establishment of an elementary connection between the centers of the spinal cord and muscles that have never been innervated by them begins (due to the decussation of the nerve trunks). Although the animal begins to stand slightly on its feet, its muscles definitely have asthenia and it cannot maintain this position for a long time.
In the next, third, period, the development of locomotor acts begins, and if at first, in the second period, these movements in the animal were carried out by synchronous movement of both hind limbs, now noticeable reciprocity begins to develop between the right and left limbs. The animal in this period is an extremely interesting object for study. During its stroke, the front and hind limbs work uncoordinated: the hind limbs move in their own rhythm, and the front legs move in their own rhythm. Thus, this period must be recognized as the period of establishing general coordination, i.e., the involvement of the reintegrative portion of the spinal cord in the general locomotor complex.
And finally, in the fourth period, all limbs correctly distribute function among themselves, the hind limbs work in full accordance with the general coordination setting and in a normal relationship with the forelimbs. It is interesting to note that any obstacle that is presented to the animal during this period during its movement, however, causes a chaotic, confused reaction and it cannot overcome this obstacle.

It was during this period that we made tests as to the extent to which the cerebral cortex took part in this reintegrative process, or, more precisely, to what extent the connection between the cortex and the corresponding muscle groups had already been established. To check, we proceeded as follows. We developed in the animal a conditioned defensive reflex to the right hind limb, i.e., to the limb in which the nerve trunks had previously been crossed. The unconditioned reaction of this limb to irritation by electric current was completely normal, i.e., at least outwardly, it did not differ from the usual unconditioned reaction: when irritated by electric current, the animal threw its leg high, bending it at the hip and knee, and during flexion at the hip, everyone took part those muscle groups that received cross innervation. Despite the correctness and sufficiency of the unconditioned reaction, we could not obtain a conditioned reflex even after 400 times of electric shock reinforcement. In general, the animal’s conditioned reaction to a conditioned defensive stimulus was developed very early - already by about the 15th application of the conditioned stimulus, the animal began to express anxiety, squeal and make general movements.

These observations indicate that a conditioned reaction as a reaction of the whole organism in an animal to a given situation was developed, but we could not develop a conditioned defensive withdrawal of the reintegrated limb by this time.

This experiment has largely convinced us that the cerebral cortex does not yet exercise fine local control over the reintegrated area. If these observations are compared with other facts that we have obtained, namely with the facts of extirpation of the motor zone in an animal after the limb with the decussation of the nerve trunks has apparently restored its function, it turns out that the removal of this zone causes paralysis of the area that included in the crossover zone.

These two experiments, which are opposite in meaning, pose a question to us: how can we understand that the cerebral cortex does not exercise fine localization control and, at the same time, its exclusion leads to paralysis of the muscles involved in the decussation? We have tried to explain this tempo by considerations that were developed by Lashley and his students (see above). If we assume that the cerebral cortex performs not only a specific, to some extent localizing function, but also a general, non-specific one, in the form of a dynamogenic effect of the cortex on all nearby centers, then it will become clear that this general effect, being more or less diffuse, manifests itself already in a very early period of the reintegrative process, precisely at the moment when the spinal centers received communication with peripheral organs, with muscles. But at this moment, the subtle local specific effect of the cortex on this organ has not yet materialized.

This analysis allows us to draw a number of conclusions about the form of participation of the cerebral cortex in the reintegrative process. From the study of Wachholder et al. we know that the rhythm with which the impulse comes from the cerebral cortex to the spinal segmental centers differs significantly from the rhythm with which the impulse passes to the effector apparatus, and thus the local segmental reflex (such as Eigenreflex) can occur to some extent independently of the cortical impulse. It is in this circumstance, as it seems to us, that the explanation of the paradoxical phenomenon described above lies. Obviously, after the crossings, an installation occurs, a mutual synchronization of those reflex relationships and those circulation of impulses that occur at the level of the segment. This is especially necessary because when the nerve trunks cross, we change the relationship between the center and the periphery.

Thus, a specific subtle influence from the cerebral cortex can be carried out only if these subtle isochronic relationships have already developed at the level of these segments of the spinal cord, while the general diffuse connection with the centers of the spinal cord in the cortex remains the same. This nonspecific influence, very likely, keeps the entire course of the regenerative process under its control and enters into a relationship with the muscles as soon as the regenerating fibers have grown to them. This is obviously what needs to be explained by the paradoxical fact that the cortex - the most mobile and labile organ - does not establish a conditional connection with the reintegrated portion of the spinal cord.

Along with these studies, we carried out a number of special experiments in which we tried to clarify the role of afferent signaling from response organs in the reintegrative process of the spinal cord. For this purpose, by making decussations of the nerve trunks on both limbs, we deafferented one of them. Experience has shown that the ataxia that forms in the hind limb as a result of its deafferentation remains very stable throughout the reintegrative process, while the normal limb is already establishing its normal relationships.

Thus, we came to the conclusion that the reintegration process depends to a much greater extent on afferent signaling from peripheral organs than on the influence of the cerebral cortex, at least in the first period of reintegration. This same fact is supported by other experiments in which, some time before the decussation of the nerve trunks, we completely removed the corresponding hemisphere of the brain. Below we will dwell on a detailed description of these relationships, when we talk about the neurodynamic understanding of the origin and implementation of the reflex, and now we will move on to those experiments that are especially important for us, since they revealed to us a number of patterns and the restructuring of nerve centers under conditions of anastomosis.

Here it would be appropriate to briefly dwell on the whole volume. what the previous physiology did in the area of ​​nervous anastomoses, or rather, why it did nothing. We have already said that each of the experiments with the crossing of nerve trunks in itself was capable of causing a revolution, but this did not happen, and... in particular, in the area of ​​autonomic nerve anastomoses, nothing was added to what we knew about the function of the autonomic nervous system. Only general conclusions were made that nerve centers can be restructured, but what factors lead to this, what intermediate processes take place in the central nervous system - this has escaped the attention of physiologists. It is the absence of such an analysis that can explain that all experiments claiming that autonomic centers can rearrange their function are absolutely incorrect. The main disadvantages of these experiments were, firstly, the incorrect statement of the very fact of the restructuring of the nerve center and, secondly, the absence of any specific, strong indicators that could be used throughout the entire reintegrative process, i.e., restructuring of the nerve center taken as indices of this restructuring. As for the first drawback, it is determined by a not entirely logical conclusion from the experiment. In fact, if you attach cut nerve trunks, then at the end of regeneration, as Langeu showed, the fibers of the central segment, no matter which nerve they belong to, will grow into the peripheral segment and into the corresponding peripheral apparatus. But one circumstance, determined by the laws of the regenerative process, in no way determines the participation of the central nervous system in this process or its own restructuring. The overwhelming majority of previous researchers, irritating the nerve above the site of anastomosis under these conditions and obtaining, say, muscle contraction, concluded that the new center now controls the new periphery.

We assert, based on a number of our experiments, that this conclusion is devoid of any logical meaning. There is a huge distance between the moment when the nerve grows into the peripheral apparatus and the moment when the center includes this peripheral apparatus in its active work, and for many nerve centers, as we will see below, this process is not feasible at all. Meanwhile, this incorrect conclusion in all previous studies and in Bethe’s summary review of these studies is a postulate that asserts the variability of nerve centers. A general assessment of the restructuring of nerve centers, which can be done to restore locomotor function, as Flourens, Bethe and we did in our studies cited above, characterizing only the end of the compensatory process, is insufficient. That is why we raise this question so acutely that a whole series of our studies devoted to the characteristics of the nerve scar formed during anastomosis have convinced us that this area, unnoticed by anyone, unstudied by anyone, takes an extremely active part in the entire process of restructuring. Where, for example, Bethe speaks of the obligatory restructuring of nerve centers (as, for example, in the case of regeneration of segments of the same nerve), thanks to the scar we have the restoration of old, pre-operative relationships without any rearrangement of the nerve centers. That is why, although the general locomotor function of the body is restored as a result of the decussation of the nerve trunks, in order to say that this restoration took place through a rearrangement of the fixed relations of the nerve centers, it is necessary to carry out a number of additional studies. In order to monitor the restructuring of the nerve center over a long period of time, it is necessary to have a certain indicator of its function in the periphery and have access from the periphery. In short, for a successful analysis of the reintegrative process we must have both afferent and efferent indicators of center function.

For these reasons, it seemed to us most convenient to study the nucleus of the vagus nerve, which has very specific functions and specific responses to afferent stimulation. The idea of ​​the experiment was to project the nucleus of the vagus nerve onto the muscular apparatus, accessible for recording, and the skin, accessible for irritation. Obviously, this idea could be realized by anastomosis of the vagus nerve with one of the mixed nerves of the forelimb.

We performed this operation in several modifications. First, we anastomosed the central segment of the vagus nerve with three or four roots of the brachial plexus, placing a nerve suture all the way to the plexus. At the same time, for a more subtle analysis, we began to sew the central segment of the vagus nerve to one, two or three nerves of the forelimb that had already gone beyond the plexus, i.e., where they had already received their specific meaning and distribution zone (P.K. Anokhin, A.G. Ivanov). We hoped that as a result of this operation, the fibers of the vagus nerve would grow into the peripheral segments and grow into the skin and muscles. Thus, when the skin is irritated, we could receive some responses from the nucleus of the vagus nerve, and the nucleus of the vagus nerve, for its part, could send impulses to the muscles receiving innervation from the vagus. Our calculation was completely confirmed. It turned out that after a certain period of time, after the regenerative process is over, scratching the skin in the shoulder area first causes a slight cough, and then an increasingly stronger cough, turning into vomiting. This phenomenon is, of course, an extremely important phenomenon in physiology; for us it was of particular value also because it was extremely constant in all our experiments. Despite numerous experiments with similar anastomoses, we have not had a single case where there was no cough in response to scratching the skin, and it was this consistency that was extremely convenient for us to take into account how the relations between heterogeneous periphery and center change over time. From the point of view of convenience of understanding all this material, it is more profitable for us to divide it into two parts - the characteristics of afferent phenomena and the characteristics of efferent phenomena. First of all, it turned out that the central nervous system, in this case the nucleus of the vagus nerve, responds extremely accurately to a certain type of stimulus. While cough always occurs in response to scratching the skin, visceral vomiting with obvious signs of nausea before it occurs only in response to kneading the muscles innervated by the vagus. These phenomena, constant in the sense of their occurrence in all operated animals, are distinguished, however, by a certain variability in connection with the regeneration process. They reach a maximum in the 4-5th month of the regenerative process and then gradually decline, so that for about a year, regeneration is achieved with great difficulty, and sometimes even no effort can cause either coughing or vomiting.

Undoubtedly, this process is deeply significant for understanding the adaptation of the center to the periphery, for it indicates that impulses that previously easily called certain working complexes to action now pass by them. That this process of regenerative changes does not permanently eliminate the previous forms of relationships can be seen from the fact that by artificially increasing the excitability of the medulla oblongata and the nucleus of the vagus nerve, we can again restore all the phenomena of coughing and vomiting. We did this in many ways, which we will not talk about here (see the works in the collection “The Problem of the Center and the Periphery in the Physiology of Nervous Activity”). Let's mention just one of them - applying a hot test tube to an area of ​​the skin where previously a cough could easily be caused. The resulting second-degree burn turned into a small ulcer, and after a few days, lightly scratching around this point could easily induce coughing and vomiting. One must think that this point-ulcer, which is a point of prolonged irritation of the endings of the vagus nerve, creates an increased tonic state in the nucleus of the latter, and now, against this background, the previous irritation again causes the same reaction.

It is extremely interesting, of course, to trace the internal mechanism of these phenomena. Are the remnants of previous relationships restored or are the relationships not eliminated at all, but only replaced by some new ones? This question is of fundamental importance, because it is associated with the characteristics of the first central synapse, which receives various forms of peripheral excitations. Is its restructuring taking place, a restructuring that would have a different functional sign, or is it not happening? We have reason to think that such a reworking of the synapse occurs due to the different nature of the impulses constantly flowing from the periphery. This is proven by the fact that the vagus nerve of the side on which the anastomosis was performed, after a considerable time after this operation (about 2 years), when irritated by an electric current, gives a different central effect than the formal vagus of the opposite side. While irritation of the normal vagus causes the usual slowing of respiratory contractions or their complete stop, irritation of the opposite vagus leads to a clearly painful reaction in the animal (squealing, general muscle contractions, etc.). The current strength remained the same.

Another proof is that secondary transection of the vagus at that stage, after the end of regeneration, does not lead to the resumption of vagal phenomena.

In the future, we will dwell on this issue and try to more deeply analyze the meaning of the synapse, and now we will move on to the characterization of those effector phenomena that are obtained under the conditions of our anastomosis.

Chronologically, muscle contraction first appears in response to direct irritation of the nerves, i.e., what was usually for previous authors evidence of a restructuring of the nerve center. But this fact has no evidentiary force and only states that the fibers of the vagus nerve have grown to the muscles. This is evident from the fact that during this period (a month and 10 days) the muscles do not take any part in the active function of the vagus nerve nucleus. Half a month or a month after this, we notice that the muscles that received innervation from the vagus begin to contract synchronously with the respiratory rhythm.

Each breath is precisely accompanied by muscle contraction, and the slightest change in the respiratory rhythm is similarly reflected in a number of muscle contractions. If, for example, you clamp the trachea or give anesthesia, or somehow influence the state of the respiratory center and respiratory movements, then the muscle that has received innervation from the vagus absolutely accurately reflects all changes in the respiratory rhythm (B. A. Matveev, A. G. . Ivanov, P.K. Anokhin, 1935).

We subjected this fact to the most subtle analysis with recording of tuning fork vibrations, and it turned out that muscle contraction does not always go exactly synchronously with the respiratory act, sometimes it advances the expansion of the chest, sometimes it lags behind it, and sometimes it exactly follows this rhythm.

We paid a lot of attention to the analysis of this fact and, as a result, developed a working hypothesis about those forms of distribution of excitation that exist within a certain functional system.

Along with these respiratory contractions of the muscle, it is also involved in the swallowing act. If, for example, a swallowing act is artificially induced, then at some stage in the development of this act the vagal muscle comes into action and performs either one or a series of contractions, and several stages can be traced in the development of the form of contraction itself.

For the first time, when the swallowing contraction of the vagal muscle just appears, it contracts singly, with a small subsequent dicrotic wave. According to observations of the swallowing act itself, this single contraction corresponds precisely to the moment when the pharyngeal ring contracts and the cartilaginous apparatus of the larynx moves. Subsequently, with the development of the inclusion of muscles in the active work of the nucleus of the vagus nerve, additional small waves begin to be added to this single wave, which correspond to some subsequent moment in the development of the swallowing act (B. A. Matveev, A. G. Ivanov, P. K. Anokhin, 1935). This circumstance is of particular interest. It convinces us that the inclusion of a heterogeneous peripheral organ in the system of active work of an unusual center occurs in stages and, obviously, in each individual stage there are its own factors that include one or another part of the muscle in the work. That this is exactly the case can be verified by directly stimulating the vagus nerve and observing muscle contraction. The contraction is always more extensive and more powerful than the contraction to the natural active impulses of the vagus nerve nucleus. This convinces us that there is a selective inclusion of muscles in the active work of this center. The nature of contractions of the muscles involved in the act of swallowing has a surprisingly identical form in a wide variety of experimental animals. This indicates that the impulse sent by the central nervous system to the muscle is always timed to coincide with a specific stage in the development of the swallowing act.

Continuing the development of our experiments, we set ourselves the task of testing a wide variety of perceptive surfaces with different receptor formations as indicators of the specific properties of excitation in the central nervous formations. The most suitable object in this regard was the cornea of ​​the eye. It has a certain limited localization, which is great for irritation. It is excited not only by mechanical and temperature factors, but quite clearly also by chemical ones, etc.

To connect the cornea with the nucleus of the vagus nerve, we performed an anastomosis of the central segment of the vagus nerve with the cut optic nerve. This operation presents great technical difficulties, since for a number of conditions it is necessary to preserve the nutrition of the retina. However, all these difficulties are well worth the results obtained. We will not dwell here in detail on the results of this operation and will refer the reader to the work of A.I. Shumilina (1935), where he will become familiar with all the details. Here we can say that such a perceptive surface as the cornea distributed the forms of stimuli in the same order as the skin, i.e. coughing and gagging were caused only by mechanical irritations in scratching the cornea, no other forms of excitation - neither chemicals nor temperature - did not cause this reaction. Only electric current, when sufficiently strong, sometimes caused coughing phenomena, but in this case we still do not know exactly to what extent mechanical irritation from the applied cotton electrodes is excluded. This means that scratching as a certain type of irritation, which obviously stimulates only a certain form of the receptor apparatus in the skin, is completely identical in nature to scratching the cornea, although the latter, as we know, does not have special receptor apparatuses. These observations, when assessed comparatively, lead us to a large generalization on the characteristics of tissues from the point of view of their ability to respond to one or another form of irritation. There is no doubt that the ontogenetic relationship of the skin and cornea is of great importance for this form of reaction.

It was of great interest to take a receptor surface that would have different receptors and a different origin. An excellent object from this point of view is the mucous membrane in general and the oral mucosa in particular. In order to connect the mucous membrane of the tongue, let's say, with the nucleus of the vagus nerve, we performed a series of operations of anastomosis of the vagus nerve with both lingual nerves. The result of this operation was very different from all our previous operations. First of all, we had completely different temporal relationships in the appearance of individual afferent phenomena. While with the anastomosis of the vagus with the cutaneous nerve, cough phenomena were always noted at the very beginning (in the form of a slight cough), with anastomosis with the lingual nerve, no scratching of the tongue produced cough phenomena, and vice versa, electrical irritation, which did not cause any reaction on the skin , in this case, when the tongue was irritated, it only caused vomiting. However, subsequently, after a few months, coughing phenomena could be caused.

Thus, in this case it turned out that the impulses sent by the receptor surface to the central nervous system vary significantly with the same type of stimulation. Whether this difference is the result of only physical constants (rhythm, amplitude) or is it the result of some other additional factors is currently difficult to decide without an oscillographic analysis of these phenomena. In any case, they seem to admit at this stage explanations in accordance with the general position of Adrian (see the next section of this work).

One of the most interesting phenomena during this anastomosis (A.G. Ivanov) is that the salivary gland, which enters the anastomotic system as an effector organ, secretes the nuclei of the vagus nerve in response to a number of conditions. So, for example, if you introduce meat broth into a dog’s stomach, that is, force it to actively secrete gastric juice, then in parallel with this, an increase in the secretion of the salivary gland occurs. As a result of the anastomosis, the salivary gland undoubtedly received fibers of the vagus nerve associated with the secretory apparatus of the stomach and intestines, and thus, its joint secretion with the stomach convinces us that the secretory impulse is differentiated throughout the entire intestinal tract not so much in its effector part, as well as afferent impulses. Each section of the intestine obviously has a certain form of receptors, which are anatomically directed to certain nerve formations, and since we have established that irradiation does not occur in the primary afferent centers, then, obviously, this contributes to localized spread to the effector part of the vagus nerve nucleus . The impulse itself in the efferent part is unlikely to differ from impulses to other glands. In any case, this problem requires further research.