CONDITIONED REFLEXES

From Big Medical Encyclopedia

CONDITIONED REFLEXES (I. P. Pavlov's article, is printed on 1 prod. BME)

the Conditioned reflex — led now the separate physiological term designating a certain nervous phenomenon, detailed studying to-rogo to formation of new department in zoophysiology — physiology of higher nervous activity (see) as chapter 1 of physiology of the highest department of the central nervous system. For a long time empirical and scientific observations collected that the bruise or a disease of a brain and specially big hemispheres caused disturbance of the highest, most difficult behavior of animals and the person who is ordinary called by mental activity. In a crust, time hardly which of persons with medical education calls in question situation that our neurosises and psychoses are connected with easing or disappearance normal fiziol. properties of a brain or with big or its smaller destruction. Then there is a persistent fundamental question: what communication between a brain and the highest activity of animals and ourselves and from what and how to begin studying of its activity? It would seem, time mental activity is result fiziol. activity of a certain mass of a brain, from physiology also there had to be its research just as now with success activity of all other parts of an organism is studied. But, however, it long did not occur.

Mental activity for a long time (not one millennium) became object of studying of special science — psychology. And the physiology amazingly recently, only from 70th year

19 century, received the first exact facts relatively a nek-swarm motive) fiziol. functions of big hemispheres by means of the usual method of artificial irritation; by means of another, too usual, a method of partial destruction were, additional data on communication of other parts of hemispheres with the main receptors of an organism are acquired: eye, ear, etc. It excited was hope of both physiologists, and psychologists concerning establishment of close connection of physiology with psychology. On the one hand, at psychologists began usage to begin the guides to psychology with a preliminary statement of the doctrine about c. N of page and specially about big hemispheres (sense bodys). On the other hand, physiologists, doing experiences with switching off of different parts of hemispheres, discussed results of animal experiments psychologically by analogy with what would occur in our inner world (e.g., munkovsky «sees, but does not understand»). But there came the disappointment in both camps soon. The physiology of hemispheres considerably stopped on these first experiences and did not move significantly further. And between psychologists after that as was earlier, it appeared many resolute people standing on perfect independence of a psychological research from physiological again.

Near it there were also other attempts to connect the triumphing natural sciences with psychology through a method of numerical measurement of the mental phenomena. At one time thought was to form in physiology special department of psychophysis thanks to a happy find E. Weber and G. Fechner of the law (see Feeling) a certain numerical communication between intensity of external irritation and power of feeling. But further this only law the new department did not go. More Vundt's attempt was successful (W. The m of Wundt), the former physiologist, then become the psychologist and the philosopher to apply an experiment with numerical measurement to the mental phenomena in the form of so-called experimental psychology; thus it was built and considerable material gathers. Someone, by Fekhner's example, calls mathematical processing of numerical material of experimental psychology psychophysis. But now to meet not a miracle also between psychologists, and especially between psychiatrists, much bitterly disappointed in the active help of experimental psychology.

So, what to do? However one more way for the solution of a fundamental question was felt, imagined and outlined. Whether it is impossible to find such elementary mental phenomenon, a cut entirely rightfully could be considered at the same time and pure fiziol. the phenomenon, and, having begun with it, studying strictly objectively (as well as all in physiology) conditions of its emergence, various complications and an ischezaniye to receive objective fiziol. a picture of all highest activity of animals, i.e. normal work

of the highest department of a brain instead of earlier the conducted all experiments of his artificial irritation and destruction? Fortunately, such phenomenon was before eyes of many long ago; many stopped on it attention and nek-ry even began to study it [especially it is necessary to mention E. L. Thorndike], but stopped for some reason right at the beginning and did not develop it in the main, essential method systematic fiziol. studying of the highest activity of an animal organism. This phenomenon also was that now designates the term «conditioned reflex»; vigorous studying it quite justified just mentioned hope. Let's put two simple experiences, to-rye all will work well. Let's pour in a mouth of a dog in moderate solution some to - you; it will cause ordinary defensive reaction of an animal:

solution will be thrown out by the vigorous movements of a mouth, outside and at the same time in a mouth (and then outside) the plentiful saliva diluting entered to - that and washing it from a mucous membrane of a mouth will pour down. Now we will put other experience. Several times any external agent, napr, a certain sound, we will work on a dog just before entering it into a mouth the same solution. And what? Then will be to repeat enough only one this sound, and at a dog the same reaction will be reproduced: same movements of a mouth and same expiration of saliva.

Both of these facts are equally exact and constant. Both of them shall be designated same fiziol. term «reflex». Both of them will disappear if to cut or motor nerves to oral muscles and secretory nerves to sialadens, i.e. efferent drives, or afferent drives from a mucous membrane of a mouth and from an ear or, at last, to destroy the central stations of transition of nervous current, i.e. moving process of nervous irritation, from afferent drives on efferent; for the first reflex it will be a myelencephalon, for the second — big hemispheres. No strict thought will find objections in view of these facts against it fiziol. conclusions. But at the same time also the difference between these reflexes is visible already. First, their central stations are various as soon as it is specified. Secondly, as clearly from statement of our experiences, the first reflex was reproduced without any preparation, without any condition, the second — was received at special reception. What did it mean? At the first — transition of nervous current from one drives to others happened directly, without special procedure: in the second — this transition required something preliminary. Everything it is more natural to imagine business so. In the first reflex there was a direct carrying out nervous current, in the second preliminary formation of a way had to be made for nervous current; such concept was in nervous physiology for a long time and was expressed by the German word Bahnung — a protoreniye.

Thus, in the central nervous system there are two different central offices: direct carrying out nervous current and device of its short circuit and disconnection. It would be strange to stop in some bewilderment before such conclusion. The nervous system on our planet is inexpressibly the most difficult and thinnest instrument of the intercourses, bonds of numerous parts of an organism among themselves and an organism as the most difficult system with infinite number of external influences. If now short circuit and disconnection of electric current are our ordinary technical adaptation, then really it is possible to object to implementation of the same principle in this amazing tool? On the basis of the stated, continuous communication of the external agent with activity of an organism, response on him, it is lawful to call an instinctive reflex, and temporary — a conditioned reflex. The animal organism as system exists among the surrounding nature only thanks to a continuous equilibration of this system with the environment, i.e. thanks to certain reactions of live system on falling on it from the outside of irritation that at the highest animals is carried out preferential by means of a nervous system in the form of reflexes.

The first ensuring steadiness and consequently, and integrity of a separate organism, as well as its look, make instinctive reflexes (see) as the simplest (e.g., cough at hit of foreign matters in a windpipe), and the most difficult, ordinary called by instincts (see) — food, defensive, sexual, etc. These reflexes are excited as the internal agents arising in the organism and external, as causes perfection of steadiness. But the steadiness reached by these reflexes would be the external environment made only at absolute constancy. And as external environment at the extreme variety at the same time is in continuous fluctuation, unconditional bonds as continuous communications are not enough and addition with their conditioned reflexes, temporary bonds is necessary. E.g., it is not enough to animal to take away in a mouth only the food which is before it, then it often would starve and died of starvation, and it is necessary to find it on different accidental and temporary signs, and it also is the conditioned (alarm) excitators exciting the movements of an animal towards food which are coming to an end in its introduction to a mouth, i.e. in general a conditioned food reflex. The same treats also everything that is necessary for welfare of an organism and a look, both in positive, and in negative sense, i.e. to what should be taken from the environment and what it is necessary to be saved from. Temporary nervous communication is the most universal fiziol. the phenomenon in fauna and in ourselves. At the same time it and mental — what psychologists call association see) whether there will be this formation of connections from various actions, impressions or from letters, words and thoughts. What basis to somehow distinguish, separate from each other what the physiologist calls temporary communication, and the psychologist — association would be? There is full slity, a full absorption by one another, an identification. It would seem that it admits also psychologists since they (or at least nek-ry of them) declared that experiences with At. rubles gave a solid support of associative psychology, i.e. the psychology considering association the base of mental activity. And it especially as by means of the developed conditioned excitator it is possible to form a new conditioned excitator, and is convincingly proved recently on an animal (dog), as two indifferent irritations repeated one after another communicate among themselves, call each other.

For physiology At. the ruble became the central phenomenon, using the Crimea it was possible to study more stoutly and more precisely as normal, and patol. activity of big hemispheres. In a crust, a statement this studying which delivered a huge number of the facts by the present moment, of course, can be reproduced only in the main lines. The main condition of education At. the ruble is in general coincidence in time one pl several times of indifferent irritations with unconditional. In total it is rather and at the smallest difficulties this education happens at direct precedence of the first irritations by the last as it is shown above on the example of a sound acid reflex.

At. the river is formed on the basis of all instinctive reflexes and of various agents of internal and external environment both in an elementary look, and in the most difficult complexes, but with one restriction: from everything, for perception of what there are receptor elements in big hemispheres. Before us the broadest synthesis which is carried out by this part of a brain! But it is not enough. Conditioned temporary connection at the same time specializes to the greatest complexity and to the smallest divisibility both conditioned excitators, and some deyatelnost of an organism, specially skeletal and verbal and motive. Before us the thinnest analysis as a product of the same big hemispheres! From here the huge width and depth of fitness, steadiness of an organism with the environment. Synthesis is, obviously, the phenomenon of nervous short circuit. What there is as the nervous phenomenon an analysis? Here several separate fiziol. phenomena. The first grounds to the analysis are given by the peripheral terminations of all afferent nervous conductors of an organism, from to-rykh everyone is arranged especially for transformation of a certain type of energy (both out of, and in an organism) in process of nervous irritation, to-ry is carried out then as in special, scantier in number, cells of the lowest departments of c. N of page, and in the most numerous special cells of big hemispheres. Here, however, ordinary come process of nervous irritation spreads, irradiates on different cells on bigger or smaller distance. That is why, when we developed, we will put, At. rubles on one some certain tone, not only other tones, but also many other sounds cause the same conditional reaction. It in physiology of higher nervous activity is called generalization At. river. Therefore, here the phenomena of short circuit and irradiation at the same time meet. But then irradiation gradually is more and more limited; irritable process concentrates in the smallest nervous point of hemispheres probably in group of corresponding special cells. Restriction most happens at means of other basic nervous process soon, to-ry is called braking. Business happens so. We at first have a conditioned generalized reflex on a certain tone. Now we will continue with it experience, constantly accompanying it with an instinctive reflex, supporting it with it; but near it we will apply also others, so to speak, self-appointedly operating tones, but without reinforcement. At the same time the last tones will gradually lose the action; and it happens, at last, and to the closest tone, napr, tone in 500 fluctuations a second will work, and tone in 498 fluctuations — is not present, otdifferentsirut I. These the tones which lost action are slowed down now. It is proved as follows. If directly after use of the slowed-down tone to try constantly supported conditional tone, he or does not act at all, or — less than usual is sharp. The braking which abolished foreign tones means, made itself felt and on it. But this short-term action: at a bigger interval after the abolished tones it is not observed any more. From this it is necessary to conclude that brake process also irradiates, as well as irritable. But than more often not supported tones repeat, that irradiation of braking becomes less, brake process more and more concentrates both in time, and in space. Therefore, the analysis begins with special operation of peripheral devices of afferent conductors and comes to the end in big hemispheres at means of brake process (see Braking).

The described case of braking is called differentiating inhibition. Let's give other cases of braking. Usually, to have the constant defined more or less At. rubles, action of a conditioned excitator continue certain time and then attach to it an unconditional irritant, support. Then the first seconds or minutes of irritation, depending on duration of the isolated use of a conditioned excitator, have no action because as premature as signals of an unconditional irritant slow down. It is the analysis of the different moments of the proceeding irritant. This braking is called braking of a delayed reflex. But the conditioned excitator as alarm korrigirutsya by braking and in itself, becoming gradually zero if in a certain span is not followed by a reinforcement. It is ugasatelny braking. This braking keeps a nek-swarm time and then by itself disappears. Recovery of the died-away irritant accelerates a reinforcement. Thus, we have the positive conditioned excitators i.e. causing irritable process in bark of hemispheres, and negative, causing brake process. In the given cases we havingm special braking of big hemispheres, cortical braking. It arises under certain conditions where it was not earlier, it practises, it

fluctuates in a size, it disappears under other conditions — and it differs in it from more or less continuous and permanent braking of the lowest departments of c. the N of page and therefore is called unlike the last (external) internal. The name would be more correct: the developed,

conditioned inhibition. Braking participates in work of big hemispheres also incessantly, difficult and thinly, as well as irritable process.

As coming to hemispheres from the outside of irritation contact there in one cases the certain points which are in a condition of irritation, so the same irritations can enter in other cases, on the basis of simultaneity, temporary communication with a brake condition of bark too if it is in that. It appears that such irritants have brake action, cause in itself in bark brake process, are conditioned negative excitators. In this case, as well as in stated above, we have transformation under certain conditions of irritable process in brake. And it can be made for ourselves to a nek-swarm of degree clear, having remembered that in peripheral devices of afferent conductors we have continuous transformation of different types of energy into irritable process. Why under certain conditions not to occur to energy conversion of irritable process in energy brake and vice versa?

As we just saw, both irritable and brake processes, having arisen in hemispheres, at first spread on them, gathering to starting point. It is one of fundamental laws of all of c. N of page, but here, in big hemispheres, he acts with inherent only it mobility and complexity. Between the conditions defining approach and the course an irradiirova-niya and concoction of processes it is necessary to consider force of these both processes on the first place. The material collected still allows to conclude that at weak irritable process there is an irradiation, at an average — concentration, at very strong — again irradiation. Perfectly the same is at brake process. Cases of irradiation at very strong processes met less often and therefore are investigated less, especially during the braking.

Irradiation of irritable process at its weak tension as a temporary phenomenon makes explicit an abeyance of irritation from other cash irritant (but weak for its detection) or from recently former or, at last, from often repeating and left behind the raised tone of a certain point. On the other hand, this irradiation eliminates a brake condition of other points of bark. This phenomenon is called a disinhibition when the irradiatsionny wave of a foreign weak irritant turns action of a certain, cash, negative, conditioned excitator in opposite, positive. At an average tension of irritable process it concentrates, concentrating in certain limited point, being expressed in a certain work. Irradiation at very strong irritation causes the highest tone of bark when against the background of this irritation and all other replaced irritations give the maximum effect.

Irradiation of brake process at its weak tension is what is called hypnosis (see), and at food conditioned reflexes is characteristic it is found in both components — secretory and dvigatelnokhm. When under the specified condition there is a braking (differentsirovochny and others), the most ordinary fact — approach of special conditions of big hemispheres. At first, against the rule more or less parallel it is normal of change of size of salivary effect of conditioned food reflexes according to physical intensity of irritants, all irritants are equalized in effect (a leveling phase). Further weak irritants give more saliva, than strong (paradoxical phasa). And, at last, the perversion of effects turns out: the conditioned positive excitator remains absolutely without effect, and negative causes hypersalivation (an ultraparadox a lny phase). The same acts also on motor reaction; so, when the food (i.e. natural conditioned excitators work) is offered to a dog, she turns away from it and when the food is taken away, carried away away, reaches for it. Besides, in hypnosis sometimes it is possible to see directly in case of food At. river gradual distribution of braking on motive area of bark. First of all language and masseters will be paralyzed, then braking of cervical muscles and, at last, all truncal joins. At further distribution of braking down the brain sometimes can notice a cataleptic state and, at last, there comes the full dream. The hypnotic state as brake very easily enters temporary conditioned connection on the basis of simultaneity with numerous external agents.

During the strengthening of brake process it concentrates. It serves to differentiation of points of bark with an irritable state from points with brake. And since in bark the mass of the diverse points, irritable and brake, belonging as to the outside world (visual, acoustical, etc.), and to internal (motive, etc.), bark represents a grandiose mosaic with the alternating points of different qualities and different rates of strain of irritable and brake states.

Thus, the vigorous working condition of an animal and the person is mobile and together the localized that more coarse, that smallest breaking of irritable and brake conditions of bark contrasting with somnolence when braking at height of its intensity and extensiveness evenly spreads on all mass of hemispheres and in depth, down on the known distance. However and now there can be sometimes in bark separate irritable points, sentry, on duty. Therefore, both processes in a vigorous state are in a constant mobile uravnovesheniye, as if in fight. If at once from bark the mass of irritations external or internal disappears, then braking over irritation takes sharp overweight. Nek-ry dogs with the destroyed peripheral main external receptors (visual, acoustical and olfactory) sleep 23 hours a day.

Near the law of irradiation and concentration of nervous processes also other fundamental law — the law of mutual induction consisting that the effect of a positive conditioned excitator becomes more when the last is applied now or soon after concentrated brake as well as the effect brake is more exact and deep after concentrated positive also constantly works. Mutual induction is found as in a circle of point of irritation or a torkhmozheniye, along with their action, and on point after the termination of processes. It is clear, that the law of irradiation and concentration and the law of mutual induction are closely connected with each other, mutually limiting, counterbalancing and strengthening each other and, thus, causing an exact ratio of activity of an organism with environmental conditions. Both of these laws are found in all departments of c. N of page, but in big hemispheres — on again formed points of irritation and braking, and in the lowest departments of c. N of page — on more or less constant.

Negative induction, i.e. emergence or strengthening of braking in a circle of point of irritation, earlier in the doctrine about At. the river decreased or disappeared at action on an animal of the foreign, accidental irritant causing on itself most often an orientative reflex (see. Approximate and research reaction). It also was an occasion to connect the cases of braking described above (ugasatel-ny, etc.), under the name of internal inhibition as occurring without intervention of foreign irritation.

Except these two various cases of braking, in big hemispheres there is also the third. When conditioned excitators are physically excessively strong, the rule of a feedforward of size of effect of these irritants and their physical intensity is violated:

their effect becomes no more, and it is less than effect of irritants of moderate force — so-called ultraboundary braking. Ultraboundary braking acts both at one very strong conditioned excitator, and in case of summation of irritants not really strong separately. It is more natural to refer ultraboundary braking of everything to a case of reflex inhibition. If more precisely to systematize cases of braking, then it — or continuous, unconditional braking (braking of negative induction and ultraboundary braking), or temporary, conditioned inhibition (ugasatel-ny, differentsirovochny and braking of delay). But there are bases all these types of braking in their physical. - a chemical basis to consider for the same process only arising under various conditions.

All installation and distribution of irritable and brake states on bark of the hemispheres which occurred during a certain period under the influence of external and internal irritations, at the uniform, repeating situation more and more are fixed, being made easier and easier and avtomatichny. Thus the dynamic stereotype (systemacity) which support makes the lesser nervous work turns out in bark; the stereotype becomes inert, often difficult changeable, difficult overcome by a new situation, new irritations (see. Dynamic stereotype). Any initial installation of a stereotype is, depending on complexity of system of irritations, considerable and often extraordinary work.

Studying At. river at the mass of dogs gradually put forward a question of different nervous systems of certain animals, and, at last, the bases to systematize nervous systems on a nek-eye to their main lines turned out. Such lines it appeared three: force of the basic nervous processes (irritable and brake), their steadiness among themselves and mobility of these processes. The valid combinations of these three lines were presented in the form of four more or less sharply expressed types of a nervous system (see Types of higher nervous activity). On force animals were divided on strong and weak, strong on steadiness of processes — on balanced and unbalanced and balanced strong — on mobile and inert. And it approximately matches classical systematization of temperaments (see). Thus, there are strong, but unbalanced animals with both strong processes, but with dominance of irritable trial of brake — excitable impetuous type (choleric persons, according to Hippocrates). Further, strong, quite balanced, besides inert animals — quiet sluggish type (according to Hippocrates, phlegmatic persons). Then strong, quite balanced, besides labile — very live, mobile type (according to Hippocrates, sanguine persons). And, at last, weak type of animals, all more suitable to gippokratovsky melancholiacs; the prevailing and their common feature — an easy tormozimost as owing to internal inhibition, constantly weak and easily irradiating, so, in particular, and external under the influence of all, even insignificant, foreign external irritations. For the rest it is less uniform type, than all others; these are those animals with both equally weak processes: preferential, with extremely weak brake, the fussy, continuously looking around, on the contrary, constantly stopping, as if stiffening animals. The basis of this not one-image-nosti, of course, that animals of weak type as well as animals of strong types, differ among themselves on other lines, except force of nervous processes. But the prevailing and extreme weakness one brake, both processes destroys vital value of variations on other lines. The constant and strong tormozimost does all these animals equally by disabled people.

So, the type is an inherent constitutional type of nervous activity of an animal — a genotype. But as the animal since birth is exposed to the most various influences of a surrounding situation, on to-rye it inevitably shall answer with the certain deyatelnost which are often fixed for the rest of life, final cash nervous activity of an animal is alloy from the lines of type and changes caused by external environment — a phenotype, character (see the Genotype).

Everything stated, obviously, represents indisputable fiziol. material, i.e. objectively reproduced normal fiziol. work of the highest department of c. N of page. But it is also necessary to begin with studying of normal work and really usually begins fiziol. studying of each part of an animal organism. It, however, does not prevent physiologists to consider still a nek-eye the reported facts not belonging to physiology. A frequent case of a routine in science!

Easily described fiziol. to bring work of the highest department of a brain of an animal into a natural and direct connection with the phenomena of our subjective world on its many points. Conditioned connection as it is already stated above, is, obviously, what we call in ourselves association on simultaneity. Generalization of conditioned connection answers that it is called association on similarity. Synthesis and the analysis At. ruble (associations) in effect same basic processes of our mental work. At the concentrated dumaniye, at interest in some business we do not see and we do not hear what about us occurs — explicit negative induction. Who would separate in instinctive most difficult reflexes (instincts) physiological, somatic from mental, i.e. with experiences of mighty emotions of hunger, sexual desire, anger etc.? Our feelings pleasant, unpleasant, ease, difficulty, joy, torture, a celebration, despair are etc. connected with transition of the strongest instincts and their irritations to the relevant effector acts, with their detention, with all variations either the easy, or complicated course of the nervous processes happening in big hemispheres as it is visible to solve nervous problems of different degrees of difficulty on the dogs who are solving or not able. Our contrast experiences are, of course, the phenomena of mutual induction. At irradiirovan-number excitement we speak and we do what in a quiet state would not be allowed. Obviously, the excitation wave turned braking of nek-ry points into positive process. Strong falling of memory of the present — an everyday occurrence at a normal old age — is age decrease in mobility of specially irritable process, its inertness.

In the developed fauna on a phase of the person there was an extraordinary increase to mechanisms of nervous activity. For an animal the reality is signaled almost only only razdra-zheniyakh\sh and their traces in big hemispheres which are directly coming to special cells of visual, acoustical and other receptors of an organism. This that, as we have in ourselves as impressions, feelings and representations from surrounding external environment as all-natural, and from our social, excepting the word heard and seen. It is the first alarm system of reality, the general at us with animals. But the word made the second, specially our, alarm system of reality, being a signal of the first signals. Numerous irritations, on the one hand, removed with the word us from reality and therefore we constantly shall remember it not to distort our attitudes towards reality. However is not subject to doubt that the fundamental laws established in work of the first alarm system shall manage also and the second because this work of the same nervous tissue (see. Alarm systems).

The brightest proof of that studying At. the river was delivered on the correct way by a research of the highest department of a brain and that at the same time, at last, functions of this department and the phenomenon of our subjective world combined, identified, further experiences with serve At. rubles on animals, in to-rykh are reproduced patol. conditions of a nervous system of the person — neurosises and nek-ry separate psychotic symptoms, and in many cases also rational deliberate return to norm, treatment, i.e. true scientific mastering a subject is reached.

The norm of nervous activity is balance of all described processes participating in this activity. Disturbance of this balance is patol. the state, a disease, and is frequent in the most so-called butto rma, therefore, more precisely, in relative norm there is already known disbalance. From here the probability of a nervous disease clearly contacts type of a nervous system. From dogs under the influence of difficult experimental conditions soon and easily the animals belonging to extreme types nervously get sick: excitable and weak. Of course, extremely strong, exclusive measures it is possible to break balance and at strong balanced types. The difficult conditions breaking chronically nervous balance, it: an overstrain of irritable process, an overstrain of brake process and direct collision of both opposite processes, in other words — an overstrain of mobility of these processes.

We have a dog with system U. river on irritants of different physical intensity, reflexes positive and negative, applied in the same way in the same order and with the same intervals. Applying it is extraordinary, exclusively strong conditioned excitators, very much extending duration of brake irritants or making very thin differentiation, or increasing number of brake irritants in system of reflexes, at last, forcing to follow directly one after another opposite processes or even working with at the same time opposite conditioned excitators, or at once changing a dynamic stereotype, i.e. turning the installed system of conditioned excitators into an opposite number of irritants — we see that in all these cases the specified extreme types especially quickly come to a chronic patol. the state which is expressed at these two types variously. At excitable type neurosis is expressed that its brake process, constantly and normal lagging behind on force from irritable, now very much weakens, almost disappears: developed, though not absolute, differentiations rastormazhivatsya quite, fading extremely drags on, the delayed reflex turns in korotkootstavlenny etc. The animal becomes extremely violent and nervous at experiences in the machine: behaves violently, what is much more rare, falls into somnolence that did not happen to it earlier. At weak type neurosis has almost exclusively depressive character. Uslovnoreflektorny activity becomes extremely chaotic, and most often absolutely disappears, the animal in the machine is almost entirely in a hypnotic state, representing its various phases (there are no conditioned reflexes any, the animal does not take even the food offered it).

Experimental neurosises mostly accept long character for months and for years (see Neurosises experimental). At long neurosises with success methods of treatment were experienced. For a long time during the studying At. the ruble was applied bromine when business went about animals, to-rye could not cope with problems of braking. Also it turned out that bromine it is essential helped this animals. Long and various lines of experiences with At. rubles on animals undoubtedly established that bromine has special relation not to irritable process, reducing it, as usual considered, and to brake, strengthening it, toning up it. It was the powerful regulator and a reducer of the broken nervous activity, but under an indispensable and most essential condition of corresponding and its exact dosage but to types and conditions of a nervous system. At strong type and at enough still a strong state it is necessary to use on dogs high doses — to 2 — 5 g a day, and at weak it is obligatory to go down to centigrams and even milligrams. Such booking within one or two weeks sometimes already happened enough for radical treatment of chronic, experimental neurosis. Lately the experiences showing even more expressed become to lay down. action, in especially hard cases, a combination of bromine with caffeine, but besides at the thinnest, now to a mutual dosage. Treatment of sick animals turned out sometimes, though not so quickly and fully as well at one long or short, but regular rest from laboratory work in general or from elimination only of difficult tasks in system U. river.

It is more natural to compare the described neurosises of dogs of everything with a neurasthenia of people especially as nek-ry neuropathologists insist on two forms of a neurasthenia (see) — excited and depressive. Then here nek-ry traumatic neurosises, as well as others reactive patol will approach. states. Recognition of two alarm systems of reality at the person it seems will lead specially to understanding of the mechanism of two human neurosises: hysteria (see) and psikhas

teniya (see). If people on the basis of dominance of one system over another can be divided into thinkers mainly and artists mainly, then in patol. cases at the general disbalance of a nervous system it will be clear that the first will be psikhastenik, and the second — hysterics. Except clarification of a .mekhanizm of neurosises, fiziol. studying of higher nervous activity furnishes the clue to understanding of the nek-ry parties and the phenomena in pictures of psychoses. First of all we will stop on nek-ry forms of nonsense, on a variation of a persecution complex, on what P. Jean calls «feeling of mastering», and on E. Krechmer's «inversion». The patient is pursued by what he especially wishes to avoid; he wants to have the secret thoughts, and invincibly it seems to it that they open constantly, recognized by others; he wants to be to one, and he is tormented by an obsession, at least it actually also was in the room one that in it nevertheless someone is, etc. — feeling of mastering, across Zhane. U E. To a rech.mer two girls, having come to a time of puberty and having received an inclination to certain men, however suppressed in themselves this inclination on a nek-eye to motives. Owing to this fact at them at first persistence developed; to their painful grief it seemed to them that on a face them sexual excitement is visible and all pay attention to it, and their sexual purity, inviolability was very expensive to them. And then at once persistently began to seem to one and it was even felt as it that in it is and moves, reaching a mouth, the sexual tempter — the dragon which tempted Eve in paradise, and another — that she is pregnant. E. Krechmer also calls this last phenomenon inversion. It concerning the mechanism, is obvious, identical with feeling of mastering. It patol. subjective experience can be understood without stretch as fiziol. phenomenon of ultraparadoxical phasa. Idea of sexual integrity as the strongest positive irritation against the background of brake, dejectedness, in Krom there were both girls, turned into so strong opposite, negative representation reaching even degree of feeling at one — in idea of stay in her body of the sexual temper, and at another — in idea of the pregnancy as result of the sexual intercourse. The same and at the patient with feeling of mastering. Strong positive representation «I one» turns under the same conditions in same opposite «about me always someone».

In experiences with At. river at different difficult and patol. conditions of a nervous system often it is necessary to observe that temporary braking leads to temporary improvement of these states; at one dog two times the bright catatonic state which caused sharp improvement of a chronic persistent nervous disease, almost return to norm for several next days are noted. In general, it is necessary to tell that at experimental diseases of a nervous system the separate phenomena of hypnosis almost constantly act, and it grants the right to accept that it is normal reception fiziol. fight against the disease-producing agent. Therefore the catatonic form, or the phase of schizophrenia (see) entirely consisting of hypnotic symptoms can be understood as the physiological guarding braking which is limiting or absolutely excluding work of a sick brain, to Krom owing to some action while unknown, harmful agent danger of serious violation or final destruction threatened. The medicine in case of almost all diseases well knows that the first therapeutic measure — rest of the body which underwent a disease. What is the understanding of the mechanism of a catatonia at schizophrenia answers reality convincingly is proved that only this form of schizophrenia represents quite considerable percent of return to norm, without watching sometimes on multiannual (20 years) continuation of a catatonic state (see. Catatonic syndrome). From this point of view all attempts to affect catatonics with exciting receptions and means are directly harmful. On the contrary, it is necessary to wait for very significant increase in a recovery rate if to fiziol. to rest by means of braking to attach deliberate external rest of such patients, but not to support them among continuous and strong irritations of a surrounding situation, among other more or less uneasy patients.

During the studying At. rubles, except the general disease of bark, were repeatedly observed extremely interesting cases also experimentally and functionally made disease of separate very fractional points of bark. Let there is a dog with system of various reflexes and between them At. river on different sounds: tone, noise, blows of a metronome, is ringing etc. — and only one of these conditioned excitators can be made disease-producing, and the others will remain healthy. Patol. the condition of the isolated point of bark is made by the same receptions, to-rye are described above as disease-producing. The disease is shown in various forms, in various degrees. The easiest change of this point is expressed in its chronic hypnotic state: on this point instead of normal communication of size of effect of irritation with physical force of irritation leveling and paradoxical phases appear. And it on the basis of the aforesaid could be interpreted as a physiological precautionary measure at a difficult condition of point. At further development of disease state the irritant does not give positive effect at all, and always causes only braking. It in one cases. In others — absolutely on the contrary. The positive irritant becomes extraordinary steady: it dies away more slowly, than normal irritants, gives in to consecutive braking from other brake conditioned excitators less, he often sharply acts in size among all other conditioned excitators that earlier, to a disease, was not. Irritable process of this point means became chronically painfully inert. Irritation patol. point remains indifferent for other conditioned excitators of bark, it is impossible to touch this point with its irritant without all system of reflexes did not fall apart anyway. There is a basis to accept that at a disease of the isolated points when in sick point prevails brake process, irritable, the mechanism of disease state consists in disturbance of balance between opposite processes: weakens considerably and preferential one, other process. In a case patol. inertness of irritable process there is a fact that the bromine (strengthening brake process) often with success eliminates it.

Hardly the following conclusion can be considered fantastic. If as obviously directly, the stereotypy, iteration and perseveration have the natural basis in patol. inertness of irritable process of different motive cells, and the mechanism of persuasive neurosis and paranoia shall be the same. Business goes only about other cells or groups them connected with our feelings and representations. Thus, only one number of the feelings and representations connected with sick cells becomes abnormally steady and does not come under inhibitory effect of other numerous feelings and representations which were more untrue thanks to a healthy condition of their cells.

The following fact, to-ry observed many times during the studying of pathology of conditioned reflexes and to-ry has explicit relation to human neurosises (see) and to psychoses (see. Mental diseases), are to a circus-lyarnost in nervous activity. The broken nervous activity was represented more or less correctly fluctuating. That there was a strip of extremely weakened activity (At. rubles were chaotic, often disappeared absolutely or were minimum), and then is as if spontaneous, without the visible reasons, after several weeks or months there came bigger or smaller, or even the perfect return to norm which was replaced then again by a strip patol. activity. That in циркулярное™ the periods of the weakened activity and which is abnormally increased alternated. It is necessary to see in these fluctuations of analogy to a cyclotymia and maniac-depressive psychosis (see). Everything it would be more natural to reduce this patol. frequency on disturbance of the normal relations between irritable and brake processes, as for their interaction. Since opposite processes did not limit each other in due course and in a due measure, and worked independently from each other and excessively, the result of their work reached excessively — and only then there came change by one another. Thus another turned out, extremely exaggerated, frequency, week and monthly, instead of short and therefore absolutely easy daily periodicity.

At last, it is necessary to mention the fact which was found still in exclusively strong form, however, only in one dog. It is an extraordinary explosibility of irritable process. Nek-ry separate or all conditioned excitators gave the swiftest and overshot effect (both motive, and secretory), but quickly breaking even during action of irritants; and the dog at a reinforcement of a food reflex of food did not take any more. Obviously, business went about severe pathology of lability of irritable process that corresponds to irritable weakness of human clinic. Cases of a weak form of this phenomenon are frequent at dogs under nek-ry conditions.

All described patol. nervous symptoms act under the corresponding conditions as at the normal, i.e. quickly not touched dogs, so (in particular nek-ry of them, napr, a tsirkulyarnost) and at the castrated animals, means on integrally patol. to the soil. Numerous experiences showed that the main line of nervous activity of eunuchs is very strong and preferential weakening of brake process, at strong type eventually, however, leveled is considerable.

In conclusion once again it is necessary to emphasize to what by comparison of ultraparadoxical phasa to feelings of mastering and inversions, and patol. inertness of irritable process with persuasive neurosis and paranoia mutually become covered and merge fiziol. the phenomena with experiences of the subjective world.

Bibliography: Pavlov I. P., Complete

collection of works, t. 1 — 5, M. — L., 1940 — 19 49. I. P. Pavlov.



CONDITIONED REFLEXES. Further development of the doctrine of I. Pavlov

the Present stage of studying At. the river is characterized, on the one hand, by attempts to find experimental neurophysiological confirmations of initial hypothetical representations of I. P. Pavlov on the mechanism of short circuit of temporary communication at formation of a conditioned reflex. On the other hand, searches new fiziol are constantly conducted. mechanisms of work of a brain, to-rye could explain patterns of formation At. the rubles who are objectively established and studied by I. P. Pavlov's school. Modern understanding to it-rofiziol. mechanisms of education At. the river is based first of all on the new facts received at researches of integrative activity of c. N of page by means of new methods. Among such methods it is possible to note registration of evoked potentials of a brain, the analysis of total bioelectric activity both bark of big hemispheres, and underlying structures of a brain (see Bioelectric potential, the Brain, Elektroentsefalografiya). The research of the last became possible thanks to the developed stereotaxic method (see) introductions to a brain of electrodes. Use of a microelectrode method of a research (see) separate nervous cells in an acute and chronic experiment expanded ideas of neurophysiologists of their functions. Appeared many experimental facts indicating a functional polymodality of nervous cells and plasticity of mezhneyronalny bonds. In this plan there were perspective researches of structure and function of synoptic formations of a brain (see the Synapse).

During the postpavlovsk period in various laboratories various modifications were described and investigated At. river. Developed at animals in various experimental situations using modern technical means U. rubles served further as convenient pilot models for the analysis of neurophysiological mechanisms of the organization of higher nervous activity (see). So, studying motive behavior of animals, E. Konorsky allocated At. river of the second type (tool conditioned reflexes). In P. S. Kupalov's laboratory situational conditioned reflexes were widely studied, to-rye it is possible to develop at animals on the same irritant located in different places of the experimental room and causing either positive, or negative conditional reaction. A kind of chain conditioned reflexes (see. A chain reflex) so-called serial and space and chain reflexes are, at to-rykh in a chain of consistently operating incentives everyone is supported or only the last incentive. Integration At. the river in more irregular shape was studied in laboratory E. A. Asratyana also received the name of uslovnoreflektorny switching. Ability of structures of a brain, mainly bark of big hemispheres, to «switching» of ways of implementation At. the river in different situations provides to an organism more exact and thin compliance of its reactions to the environment. Staff of laboratory E. And. Asratyan was studied by specific features of associations during the formation at the animal difficult behavioural acts which received the name of causal conditioned reflexes. Nevertheless, the original positions of the uslovnoreflektorny theory formulated by I. P. Pavlov continue to form a basis for new researches. His idea about formation of new functional linkages between nervous cells of the cortical center of an indifferent irritant at its combination to an instinctive reflex (see) found further development in E. A. Asratyan (1970) ideas of a so-called local conditioned reflex. Results of the conducted electrophysiologic researches showed that the combination of the indifferent and supporting irritants involves sharp increase in functional linkages between neurons of the cortical center of future prearranged signal. Assume that in an initial mode of formation of conditioned reflexes the similar mechanism promotes establishment of distantny contact between different cell populations of bark of big hemispheres. And between cortical structures of two combined centers of excitement there are bilateral (direct and the return) conditioned connections.

I. P. Pavlov's hypothesis of structurally functional bases of education At. the river which is based on a neurophysiological phenomenon of a banung or protorhenium of ways gained the development in researches A. And. Ukhtomsky considering the dominant center of excitement in c. N of page capable to attract to itself weak excitement from other centers of a brain (see the Dominant). The subsequent works

of V. S. Rusinov showed increase in excitability of the relevant cortical centers, synchronization of work of nervous cells, ability to sum up them excitement, creating a stationary dominant position in reflex activity of a brain. In particular, it was shown that if the dominant center is localized in the cortical center of any paw of an animal, then its movement can be caused by action of such indifferent irritants as light or a sound. The dominant center of excitement is capable to acquire a rhythm of naturally operating irritants or artificially imposed by direct electric irritation of the relevant cortical structure. Long preservation of this rhythm the dominant center demonstrates existence of mechanisms of fixing of vozbuzhdeniye in nervous elements that is an important point in the functional organization of other excited cortical centers. According to V. S. Rusinov (1972), the created dominant and the summatsion-ny reflex preceding it are the initial stage of formation of temporary communication which is characterized by a phase of generalization created At. river when it can be caused practically by any irritant. Only in the course of transformation of a dominant in At. river the last becomes specialized, localized and stable. Nek-roye looking alike a dominant has the phenomenon of post-that-ganichesky potentsiation for the first time studied by J. Ekpls and P. G. Co-styukom on spinal structures. After more or less long electric irritation of sensitive roots of a spinal cord in its structures long preservation of a hyperexcitability was observed. It had local character and was shown in strengthening of the corresponding spinal reflexes. In L. G. Voronin, A. I researches. Roytbak and other physiologists it is shown that the post-tetanic potentsiation is more shown in bark of big hemispheres, neurons a cut have ability to longer keep the trace phenomena in comparison with other structures of c. N of page. Numerous elektrofi-zdologichesky researches revealed that in cortical zones of representation of unconditional reaction the expressed changes of bioelectric activity are observed. In particular, in a cerebral cortex on an unconditional irritant at development At. the river occurs increase in the caused answer. At its fixing late negative components of evoked potentials are stabilized. The analysis of total bioelectric activity also indicates a condition of a hyperexcitability and domination of the cortical center of unconditional reaction.

Results of researches of change of level of excitability of cortical representation of a conditioned excitator at its combination to an unconditional irritant were less unambiguous. The observations which are saved up by I. P. Pavlov's school about gradual weakening of the orientative reflex caused in process of development At. rubles an indifferent irritant, led it to a conclusion about decrease in excitability of the cortical elements perceiving a prearranged signal. However the researches conducted on the person G. V. To a ruff, E. N. Sokolov, etc., showed increase in sensitivity to perception of sound, light and other indifferent irritants at their turning into a conditional incentive. In the cortical structures corresponding to these irritants the threshold of emergence of evoked potentials decreases. At hmikroelektrodny researches on animals it was shown that the impulsation of the cortical cells perceiving a conditioned excitator quite often increases at a combination of this incentive to an unconditional reinforcement. Thus, the numerous researches conducted for the purpose of the analysis of a functional condition of the centers of excitement in bark of big hemispheres and sent to search for estimated mechanisms and ways of formation of temporary communication are in a conflict with traditional provisions of the uslovnoreflektorny theory and forced researchers to reconsider approaches to the analysis of neurophysiological mechanisms of education At. river.

Recent trend of researches U. by river it was connected not only with wide use of an electroencephalography, but also with opening of the ascending activating influences of a reticular formation (see) mesencephalon, thalamus and other subcrustal structures on a cerebral cortex. A large number of researches of mechanisms of corticosubcortical relationship at education is carried out At. river. Until recently ideas of processes in a cerebral cortex were connected with classical projective pathways, on the Crimea excitement reaches a certain cortical field and from where begins to extend to all other areas. On etokhm the interpretation of irradiation and concentration of the basic nervous processes in bark of big hemispheres was based. Studying of activity reticular' formations promoted emergence of idea of the channels of afferent impact on a cerebral cortex which are not considered by former physiology of c. N of page. The analysis of the ascending activating influences of a reticular formation on a cerebral cortex, the carried-out J. Moruzzi and X. Megun (1949), allowed on - but-vomu to treat mechanisms of interaction and distribution of vozbuzhdeniye in bark of big hemispheres. Generalized excitement of a reticular formation supports a working tone of bark of cerebral hemispheres and by that provides a possibility of its associative activity. Results elektrofiziol. researches point to the leading role of a reticular formation in the phenomenon of generalization At. river at initial stages of its education. At the first stage of formation At. the river of the most characteristic is the reaction of desynchronization noted on the electroencephalogram and extending to all structures of a brain. In process of consolidation and specialization At. the river this generalization breaks up to unstable and various local changes of bioelectric activity in structures of a brain. The analysis of frequency and amplitude changes of bioelectric activity of bark and subcrustal formations of a brain at development At. the river, carried out by M. N. Livanov, showed that their localization also depends on a stage of formation and the nature of uslovnoreflektorny reaction. During the fixing At. river M. N. Livanov observed specific changes of bioelectric activity in structures of bark, thalamic kernels, a hypothalamus and a reticular formation of a mesencephalon. The conclusion was drawn that in process of fixing At. the river in background biopotentials of bark occurs weakening of those changes, to-rye appear at early stages of elaboration of temporary communication. According to many researchers, changes of bioelectric activity in a reticular formation appear earlier, than in a cerebral cortex, and are from-razheniyekhm the mechanism of short circuit of temporary communication. Only for the second time due to the ascending influences of a reticular formation At. the river becomes a cortical phenomenon. At the same time except a reticular formation of a mesencephalon and a thalamus function of short circuit of uslovnoreflektorny communication can be carried out by neurons of a rhinencephalon, striate body and associative bark. At the same time corticofugal influences constantly keep a functional condition of subcrustal structures under control. According to P. K. Anokhin, cortical control of integrative functions of subcrustal structures is exercised through approximate and research reaction to future prearranged signal. On the basis of the obtained data on mechanisms of corticosubcortical relationship of I. S. Beritashvi - whether, M. M. Hananashvili, N. Yu. Belenkov made a hypothesis that formation of temporary communication is made on the way: a cerebral cortex — subcrustal structures — bark. From this point of view short circuit At. the river occurs between the structures which are carrying out an orientative reflex and structures of the supporting instinctive reflex.

The instability of the supporting properties of an unconditional irritant found in I. P. Pavlov's laboratories induced scientists to investigate a role of motivational conditions of an organism in mechanisms of formation of conditioned reflexes (see Motivations). The main biol. motivations — hunger, thirst, fear, aggression, sexual desire, etc., called I. P. Pavlov the main inclinations of an organism, on external manifestation carry to compound instinctive reflexes, or instincts (see). On origins they are connected with deviations of various indicators of constancy of internal environment of an organism and transformation of the arisen requirement in motivational excitement of structures of a brain. Neurophysiological basis biol. motivations the ascending activating influences of subcrustal structures on bark of big hemispheres are. The motivational centers, the being hl. obr. in a hypothalamus (see), and also in limbic structures (see. Limbic system) and a reticular formation, are a source of the activating influences, to-rye on the electroencephalogram are shown preferential in the form of a desynchronization test of total bioelectric activity. The same picture on EEG at the ascending influences of subcrustal structures on a cerebral cortex gave the grounds to speak about a nespe-tsifichnost of these influences. The similar conclusion was drawn on the basis of direct irritation of subcrustal structures, in particular a reticular formation. Works of G1. To. Anokhina and it sotr. showed that at development At. rubles at animals in the experimental conditions coming to natural, the ascending activating influences from subcrustal educations on a cerebral cortex always have the specific character connected with a specific biol. reaction of an organism (food, sexual, defensive, etc.). Specific biol. the condition of an organism finds also reflection and in background pulse activity of neurons of various areas of bark of big hemispheres. Thus, motivational excitement is a basis of integration into c. N of page of all types of the afferent vozbuzhdeniye arising at action on an organism of various irritants. From positions of the theory of functional systems of P. K. Anokhin (see. Functional systems), such integration always takes place during the formation At. the river is also designated as a stage of afferent synthesis (see). At this stage any external or internal irritant can serve one of the factors preparing formation of a conditioned reflex (a so-called obstanovochny afferentation). E.g., on the same conditioned excitator the animal in one experimental situation shows defensive At. river, and in another — food. On the basis of the dominating motivation with involvement of mechanisms of memory (see) and taking into account an obstanovochny afferentation prestarting integration of vozbuzhdeniye in the central nervous system forms, edges is implemented in uslovnoreflektorny reaction at action of a conditional incentive (a starting afferentation).

System ideas of the organization of intracerebral processes during the formation of conditioned-reflex behavior of animals and the person are further creative development of original positions of the uslovnoreflektorny theory of I. P. Pavlov.

The present stage of studying of mechanisms of education At. the river is characterized by researches of of N of page at the cellular and synoptic levels. Neyrofiziol. mechanisms of short circuit of temporary communication began to consider as integration of vozbuzhdeniye at the level of separate neurons of various structures of a brain. Such formulation of the question was promoted not only by development of a microelectrode method of a research of nervous cells, a method of a microionophoresis, but also achievement of the general cytology, a submicroscopy, neurochemistry and molecular biology. Basis of processes of integration into c. the N of page is, as we know, the principle of convergence of vozbuzhdeniye for the first time formulated and studied by Ch. Cher a ringtone (1906) at operational analysis of a spinal cord. Ch. Sherrington also described emergence of occlusion and simplification as a result of steric and temporary interaction of the vozbuzhdeniye coming to one point of c. N of page. Universality of the principle of convergence for c. to N of page it was shown in works of pupils of I. P. Pavlov in the analysis of interaction in structures of a brain of conditional and unconditional vozbuzhdeniye. It was established that convergence of vozbuzhdeniye to separate neuron can be carried out practically in all structures of c. N of page. The multitouch convergence which is characterized by a meeting and interaction on neuron of two or more geterotopny or heterogeneous afferent signals of various touch modality is fullestly studied. Especially clearly multitouch convergence is shown in a pontomezentsefalichesky reticular formation, on neurons the cut is converged and interact excitement, arising at somatic, visceral, acoustical, visual, vestibular, cortical and cerebellar irritations. Ability of neurons to react on several shown irritants depends not only at most and a touch modality of irritants, but also on the level of a functional condition of all structures of a brain in general. Multitouch convergence takes place and in nonspecific kernels of a thalamus, the median center, a kernel having a tail, a hippocampus, structures of limbic system where neurons are found, to the Crimea converge excitement at somatic, visual, sound, olfactory, cortical irritations, and also at irritations of a reticular formation. These structures of the headsleg of a brain are as if the intermediate switching relays on the way of distribution of afferent excitement from receptive fields to a cerebral cortex. Especially numerous effects of multisen-weed convergence are observed on neurons of various areas of bark of big hemispheres. Researches of reactions of neurons of primary projective zones of bark (visual, acoustical, touch) showed that in these areas along with the specific neurons reacting only to an irritant of strictly certain touch modality there are neurons nonspecific, reacting to irritants of several touch modalities. Nonspecific neurons most often react multi-touch convergence, the shown reaction of neurons to several shown irritants (sound, light, tactile, etc.). A large number of multisensory neurons is found in associative areas of a cerebral cortex that reflects functional value of these areas in mechanisms of formation of temporary bonds. Depending on quantity and quality of the vozbuzhdeniye coming to neurons they are classified as bisensorny, trisen-weed and polytouch. In bark of big hemispheres also other types of convergence of vozbuzhdeniye on neurons, excellent from multitouch and characteristic only of a cerebral cortex were found. Studying of neural cortical responses during education At. showed river that to separate cortical neuron is converged excitement of a touch modality (conditioned excitator) and a biological modality (an unconditional irritant). Such convergence of vozbuzhdeniye was called by G. Dzhasper touch and biological. To separate neurons of a cerebral cortex can converge also ascending influences created by various subcrustal structures and having a specific individual biological modality (e.g., food and painful). This type of convergence is called by Yu. A. Fadeyev (1965) multibiological. To neurons of projective areas of bark can converge specific afferent excitement, and also excitement, extending on collaterals from efferent axons of cells of a pyramidal path during his antidromic irritation (afferent and efferent convergence). The established types of convergence of vozbuzhdeniye allow to present in a new way the mechanism of short circuit At. and to determine by river a role of separate cortical neurons in these mechanisms. Being the meeting place and interactions of conditional and unconditional vozbuzhdeniye, separate neurons inevitably become the arena of their interaction. At the level of subcrustal structures and a cerebral cortex the effect of interaction of two vozbuzhdeniye is implemented or in the simplification which is shown increase in quantity of impulses in the answer of neuron to irritation or in braking of activity of neuron in response to the second or subsequent afferent influence. The final effect of interaction of vozbuzhdeniye depends at most two irritants, temporary ratios between them, and also their touch and biol. modalities. In turn, reaction of neurons to several irritants of various modality significantly depends on the level of their excitability that as it was already noted, shown in change of background pulse activity. Therefore, neurons of c. N of page, especially bark of big hemispheres, being point of convergence and interaction of vozbuzhdeniye, it is possible to consider elementary base units of integration.

Big actual material according to the analysis of mechanisms

of integration of heterogeneous vozbuzhdeniye at the level of separate neurons of c is saved up. N of page at impact on an organism of irritants in the sequence standard for development At. river.

The researches conducted on the immobilized animals using narcotic substances and muscle relaxants showed that the most part of neurons practically of all cortical and subcrustal structures of a brain can give conditional reactions to earlier inefficient touch irritants in process of their combination to the effective supporting incentives. The greatest number of researches of reorganizations of pulse activity of neurons of a brain is carried out at development At. river at animals by a classical technique when earlier indifferent incentive was supported with an unconditional irritant. According to numerous researches, the amount of the mustache-lovno-reacting neurons in various formations of a brain fluctuates from 30 to 90% of all cells registered in an experiment. H. N. Vasilevsky (1968), G. I. Shulgin (1978) and Yu. I. Alexandrov (1976) was assumed that reaction of neuron to a conditional incentive repeats reaction of neuron to an unconditional irritant in the course of development At. river. According to other point of view stated by O'Brien and Fox (J. The N of O'Brien, S. S. Fox, 1969), true manifestation of uslovnoreflektorny reaction at the neural level is change of nature of reaction of neuron on opposite after an otkhmena of the supporting incentive. In experiments on the awake animals with registration of effector manifestations in the form of the electromyogram or local motor reaction it is offered to identify conditioned responses of neurons by the following criteria: test on a nonspecific sensitization; emergence of earlier not registered stable answers of neuron to a conditional incentive; change of a sign of conditional reaction of neuron at repeated combinations; reproduction on a prearranged signal of trace reaction to a reinforcement at its sudden cancellation. The most interesting and a little unexpected fact is increase in stage of latency of reactions of neurons of all studied cortical and subcrustal structures at the irritants getting in the course of development At. river alarm value. This fact, obviously, reflects a certain reorganization of processes in a brain, the essence and mechanisms a cut did not find rather accurate explanation yet. Besides, at different formation At. rubles responses of neurons to an alarm incentive are produced 'the corresponding behavioural act much earlier. According to numerous researches, conditional reactions of neurons as activating, and brake type appear within the first 10 — 15 combinations and reach significant level after 30 — 50 combinations of conditional and unconditional incentives while stable behavioural conditional reaction appears after 150 — 200 combinations. Thus, formation At. by river it is connected not only with change of a functional condition of separate neurons, but also with the advent of the new organization of interrelations between them. Even more contradictory and difficult analyzed were data obtained at registration of pulse activity of neurons of a brain at animals in the conditions of free behavior. In a similar situation at implementation by an animal strictly certain At. the river the studied population of neurons was heterogeneous on the nature of responses of each cell and on extent of its involvement in the carried-out behavioural act. Besides, at repeated performance of the stereotypic movement by an animal the same neuron gives considerable dispersion in reactions, i.e. has no stability. If to add the polyfunctionality of cells noted in many researches and selectivity of their involvement in response motor reaction to it, then an impression a nek-swarm of chaoticity and uncertainty in work of nervous cells is made. The offered G1. K. Anokhin (1968) a hypothesis of convergent short circuit At. the river helps to overcome the arisen contradictions formulation of the question about integrative activity of neuron. Systematic researches of the thin mechanisms providing simultaneous processing of numerous and diverse vozbuzhdeniye in a stage of afferent synthesis during the formation At. rubles, inevitably led to a conclusion that convergence of vozbuzhdeniye on neuron is the universal working principle of its integrative activity. The main mechanism of interaction of vozbuzhdeniye on neurons summation on a membrane of postsynaptic potentials with the subsequent influence on a generating zone of cells admits. However numerous cytochemical and microbio-chemical researches indicate the certain biochemical differences of various types of neurons depending on their localization and the carried-out function. The submicroscopy convincingly shows that process of excitement in neuron is not limited to a subsinaptiche-sky membrane, and actively develops in cytoplasm, involving various intracellular organellas. All these data inevitably lead to other explanation of the integrative activity of neuron constructed at association of vnutriney-ronny cytoplasmatic reactions and the postsynaptic processes which are continuation in dendrites and a neurocyton. It is taken for granted that communication between neuromediator effects on a post-snnapticheskoy to a membrane of neuron and intracellular metabolism is carried out by cyclic nucleotides, mainly cyclic adenosine-3', the 5th '-monofos-fathom (tsAMF). Schematically these processes can be presented as follows: action of a neurotransmitter on hemoretseitivny sites of a postsynaptic membrane —> activation of adenyl cyclase —> increase in synthesis of tsAMF —» activation of protein kinases —> changes a fosfori-lirovadiya of proteins conformational changes of nuclear or synoptic proteins.

The subsequent mezhneyronalny integration in scales of the whole brain during the formation At. the river can occur both on functional, and on a structural basis. Theoretical ideas of reverberant vozbuzhdeniye in neural networks of bark and subcrustal educations [Kendel (E. R. Kap-del, 1970); Yarvik (M. of E. Jarvik,

1972)], the idea about vnutrianalizatorny and mezhanalizatorny interaction on neurons (E. A. Asratyan, 1965; A.S. Batuyev, 1970), a hypothesis

of formation in c. N of page of nervous network on the basis of a functional linkage between neurons [Burns (V. O. of Burns, 1968); A. B. Kogan, 1973; John (E. R. John, 1973); G. I. Shulgin, 1978], assumptions of the organization of neurons in microsystems (M. M. Hananashvi - whether, 1972) are in a stage of development so far or are buttressed insufficiently up experimental by facts and cause dissatisfaction of many researchers. The close attention is drawn by the ultra-restructurings of the synoptic device revealed at electronic microscopic examinations on analog models U. river (see the Synapse). Increase in quantity of bubbles and their concentration at a presynaptic membrane of neuron, and also a thickening of postsynaptic membranes, especially aksodendritny synapses was observed. At the same time there can be conformational changes of synoptic proteins that leads to direct reorganizations

of the interneural bonds providing short-term fixing of effects of interaction of vozbuzhdeniye. The conformational changes of nuclear proteins arising as a result of postsynaptic activation of neuron can lead to activation of its genome with increase in protein synthesis and peptides. A part of proteins at the same time, especially specialized, is transported to rubbed-minalyam also to various sites som of neuron, providing stable changes of their properties and formation of the multineural systems forming a morfofunktsionalny basis of temporary bonds. The original hypothesis of structural bases of formation of vrekhmenny communication was made by A. I. Roytbak (1969, 1972). On its representations, any supporting irritant along with activation of the corresponding neurons causes

depolarization of membranes oligoden-

drotsit, strengthening their miyelinobra-zuyushchy activity. Shoots of oligodendrocytes, contacting

with deprived of a myelin aksonny presynaptic the term of l to a hole,

begin to wrap them, thereby increasing efficiency of transfer of vozbuzhdeniye and quantity of the released mediator. Repeated combinations of conditional and unconditional irritants lead to increase of layers of a myelin of a presynaptic bombway and consolidation of uslovnoreflektorny temporary communication.

Researches on reproduction are of great interest At. rubles at hematothermal animals after an injection of homogenates or bio-chemical the substrates of a brain taken from donor animals at to-rykh previously were developed by it certain forms of uslovnoreflektorny behavior. Cleaning and identification of the active proteinaceous beginning of extract of a brain was made, a cut received the name of a skotofo-bin (see the Neurochemistry). It is shown that at development at rats At. the river of avoiding of a dark box quantity of a skotofobin in a brain progressively accrues, reaching a maximum for


the 6th day of an experiment, and then its contents decreases. Intraperitoneal introduction of a skotofobin to kry-@-recipients caused in them a phobotaxis of darkness. Further expansion of researches on identification and protein synthesis of the extracts of a brain capable to cause reproduction in recipients of behavior of donors will allow to get more deeply into intimate mechanisms of education in c. N of page of uslovnoreflektorny temporary bonds. «True short circuit of uslovnoreflektorny communication —

P. K specified. Anokhin (1968), is finally always thin irotoplazma-tichesky process, and all anatomy, biophysics, power, transfer, distribution and interaction of vozbuzhdeniye are only means of bringing various information to those points of an organism where it can be fixed and „packed“ into strong chemical konstellyation».

Bibliography: Anokhin P. K. Biology and neurophysiology of a conditioned reflex, M., 1968, bibliogr.; it, Sketches on physiology of functional systems, M., 1975; it, System mechanisms of higher nervous activity, M., 1979; Asratyan E. A. Sketches on physiology of conditioned reflexes, M., 1970; it, Ivan Petrovich Pavlov, M., 1981; about N e, the Reflex theory of higher nervous activity, M., 1983; Batuyev A. With * the Highest integrative systems of a brain, JI., 1981; Belenkov N. Yu. The principle of integrity in activity of a brain, M., 1980; Konorski Yu. Integrative activity of a brain, the lane with English, M., 1970; To about t of l I r B. I., etc. Neural organization of uslovnoreflektorny behavior, M., 1983; Krylov O. A. Reproduction of uslovnoreflektorny activity by administration of biochemical substrates, Usp. fiziol. sciences, t. 5, No. 4, page 22, 1974; Livanov M. N. Space organization of processes of the heads * leg of a brain, M., 1972; Pavlov I. P., Complete collection of works, t. 1 — 5, M. — L., 1940 — 1949; Simonov P. V. Higher nervous activity of the person, M., 1975; Modern problems of physiology of higher nervous activity, under the editorship of, N. P. Bekhtereva, M., 1979; Vessels

of k of K. V. Biological motivations, M., 1971; Fadeyev Yu. A. Pulse activity of cortical neurons during the formation and implementation of purposeful behavior, Usp. fiziol., sciences, t. And, No. 3, page 12, 1980; X and N and -

nashvit M. M. Mechanisms of normal and pathological uslovnoreflektorny activity, JI., 1972; Shvyrkov V. B. Neurophysiological studying of system mechanisms of behavior, M., 1978; Electric activity of a brain at formation of simple forms of temporary communication, under the editorship of V. S. Rusinov, M., 1972; The Electrophysiologic research of stationary activity in a brain, under the editorship of M. N. Livanov, M., 1983; In user P. Higher functions of the nervous system, Ann. Rev. Physiol., v. 38, p. 217, 1976; Classical conditioning, ed. by A. H. Black a. W. F. Prokasy, N. Y., 1972; Sutherland N. S. a. Mackintosh N. J. Mechanisms of animal discrimination learning, N. Y., 1971. Yu. A. Fadeyev.

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