FUNCTIONAL SYSTEMS — dynamically developing units of integration of a complete organism selectively combining vocational central and peripheral educations and directed to achievement of results of adaptive activity.
The concept «Functional Systems» evolved from systematic researches of processes of compensation of the broken functions (e.g., imposing of a heterogeneous nervous anastomosis and overseeing by the course of recovery of functions, change of muscles for the purpose of giving of new functional value to them, their deafferentation etc.). Fiziol. an essence of compensatory adaptations (see. Compensatory processes) consists that each attempt of the person or an animal to correct available «defect» shall be estimated immediately by its result. It means that any following stage of compensation can come only when there was assessment of the previous stage. Thus, in each separate stage of compensatory process there is assessment of the received result and degree of its usefulness for an organism, and the chain of «positive takes» of compensation provides a complete recovery of the lost function
. Such system which is carrying out qualitatively outlined adaptive effect, all parts a cut enter the dynamic, urgently developing functional association on the basis of continuous return information on adaptive result, it was called P. K. Anokhin (1935) «functional system». This principle became central for an explanation of all those adaptive processes, to-rye gain lines of complete acts and come to an end with useful adaptive effect (see Adaptation). Everyone F. the page represents to a nek-swarm of degree a loop system thanks to a continuous communication with peripheral bodies and a constant afferentation from these bodies. Thus, everyone F. the page has a certain complex of afferent alarm systems, to-ry through an acceptor of action (see the Acceptor of results of action) directs performance of its function. Separate afferent impulses can proceed from the most various and remote one from another bodies. E.g., at the respiratory act afferent impulses go from a diaphragm, intercostal muscles, lungs, a trachea, however, despite their various origin, these impulses combine in c. and. page due to emergence of the thinnest temporary ratios between them.
Thus, activity of everyone F. the page is in close dependence on quality and quantity of afferent impulses, as from straight lines (i.e. being incentives to commission of action), and from the return affe-rentation (informing on results of perfect action). Experiments with section of back roots of a spinal cord at a frog and the subsequent testing of her motor functions (swimming, a jump, a turning from a back in normal situation) showed that intervention of the return afferentation happens only when standard functional relationship within whole F is broken. page (e.g., at increase in load of an extremity). Requirement F. the page in a certain quantity and quality of afferent impulsation depends on complexity of this system, and according to extent of automation and phylogenetic antiquity of system required quantity and quality of afferent impulsation variously.
The role of afferent functions is in a complete dependence from properties and from a final effect this F. page. In other words, T. the page as a unit directed to obtaining a certain adaptive result has an opportunity to dynamically redistribute participation of afferent impulsation, maintaining their activity on a fixed level. As complete education any F. the page has the properties, quite specific to it, giving it plasticity, mobility and a certain degree of independence of the ready, developed designs of various bonds as within c. N of page, and all organism.
Important nodal concepts of physiology are a homeostasis (see) and self-control of physiological functions (see). T. the page is the branched morpho-physiological device providing through a number of its own patterns effect both a homeostasis, and self-control. She uses various thin mechanisms of integration and directs the course of all intermediate processes, up to obtaining final adaptive effect and assessment of its sufficiency.
Central point F. the page is receptor education, a cut on the trigger (starting) properties (see. Trigger mechanisms) it is precisely adapted for physical or chemical parameters of this useful effect. The final adaptive effect of system and its receptor device make the interconnected complex. Receptor part F. the page is its most conservative education keeping often during all life of an organism constancy of useful effect. It means that situation K. Bernard about constancy of internal environment should be carried not to the most internal environment, tending to considerable fluctuations, and to receptor devices, to-rye perceive changes of this Wednesday and through c. N of page include or switch off various additional mechanisms maintaining constancy of separate components of internal environment of an organism (see). Thus, in structure F. pages enter at least two categories fiziol. mechanisms with very various properties: 1) the mechanisms having extreme conservatism (receptors of result) and relative conservatism (a final effect); 2) nodal mechanisms of system, namely means of achievement of adaptive result having high ductility and ability to mutual substitution.
456 FUNCTIONAL SYSTEMS
Universal model F. page — means of the studying of any integrative education supporting this or that useful effect or reaching it in the conditions of life activity of the whole organism. E.g., if to the place of a final effect to substitute the size of osmotic pressure of blood or level of blood pressure, then it will become clear that complete integration will function and give adaptive effect on the basis of the same fiziol. architecture. Universality of separate mechanisms F. the page demonstrates that vital process, once organized on the basis of most and regulatory devices, was created in the course of evolution F. page as device of the most difficult integrative devices.
Many such «finds of evolution» are known, to-rye were useful to progressive development of the organic nature in the course of natural selection and at the highest stages of its development were widely adopted. Molecule DNA can serve as an example (see. Deoxyribonucleic acid), edges, having been useful to transfer of ancestral features of an organism, gained universal value of substance of heredity for the most various representatives of the organic world, beginning from a virus and finishing the person. Can be the same example and the membrane of living cell, structure a cut essentially did not change during evolution, having remained identical with egg of a sea hedgehog and with a cell of bark of big cerebral hemispheres (see Membranes biological). Very perhaps, as T. page, providing the effect widely known under the general name «expediency», it was enriched at each stage of evolution with new contents, keeping, however, the basic architecture.
Structure F. the page is not defined by topographical proximity of structures or their belonging to any section of anatomic classification. V F. pages can be selectively involved as it is close, and the structures of an organism remotely located; it can involve fractional sections of any integral systems in the anatomic relation and even private details of separate bodies. The only factor defining selectivity of these connections is biol. and fiziol. architecture of function, in some cases even her mechanic (e.g., jump). The only criterion of full value of these associations is the final adaptive effect for the whole organism coming at expansion of processes in this F. page.
Everyone F. the village has the regulatory properties inherent in it as whole and absent at its parts. Regulatory properties F. pages are first of all that at any damage to one of parts F. the page leading to disturbance of useful effect there is a bystry reorganization of the processes making it. The clearest pattern of system activity is progressive elimination of afferent influences from the total amount of afferentation of this system as soon as it passes to stationary functioning (the principle of «narrowing of an afferentation»). The final result of narrowing of an afferentation always is preservation some residual, sometimes very limited, «the leading afferentation». Integrative character F. the page affects that at any disturbance of the leading afferent impulsation or at a deviation in a net result «reserve afferentation», i.e. afferent impulses eliminated earlier, owing to what F are instantly involved in action. the page as whole keeps the architecture, useful to an organism.
What specific mechanisms carries out F. page the regulatory properties, keeping reasonable character of adaptation at all levels of functioning? Researches show that everyone F. the page has the accurately outlined nodal mechanisms.
Afferent synthesis as specific mechanism of functional system. Idea of afferent synthesis (see) as about a necessary and universal stage of formation of any conditioned reflex or the behavioural act there was gradually also a hl. obr. on the basis of assessment of a relative role of a conditional incentive in formation of conditional reaction. I. I. Laptev and A. I. Shumilina's experiments (1949) showed ability of a brain to make very extensive synthesis of all those signals of the outside world, to-rye come to a brain through various sense bodys and have various functional value. It became clear that an external irritant, coming in the form of excitement to c. the N of page, extends on it by no means not linearly as it is postulated by the classical reflex theory (see). He by all means enters thin interactions with other afferent vozbuzhdeniye making other functional sense and only depending on synthesis of all these afferentation conditions for formation of purposeful action are created. Then this interaction is enriched with the new afferent irritations which are actively selected by means of an orientation response (see. Approximate and research reaction).
Value and the place of each of external irritants in the course of afferent synthesis can be expressed by the following general formula: fiziol. the sense of any of external and internal irritations consists that it can have or starting character, i.e. be a true incentive for emergence of any reaction, or it can be the peculiar factor preparing the integrated reaction, edges stays in the hidden look and does not come to light yet. All forms of such irritations make organic unity, but each of them contributes the specific share in this unity. So, usual training of an animal for experiments on conditioned reflexes (see) consists that the experimenter previously, before start-up of a conditioned excitator, makes a number of standard preparations (preparation of a forage, a sticker of a vessel for collecting saliva, closing of a door, etc.). It is accepted to think that the major and point in manifestation of a conditioned food reflex the conditioned excitator is decisive. However actually it not so. E.g., if from all the preparatory operations described above to pass only one — a podsypaniye of sukharny powder in a feeding trough, then at once it turns out that the usual, always well acting conditioned excitator is not capable to cause conditional secretory effect now. Why there was such depreciation of a conditional incentive if it is known that the listed above irritations make together only a complex of obstanovochny irritations, and each of them taken separately usually does not cause salivation? It is obvious that the final conditional effect is result of action not only a conditional incentive. Excitement from it enter synthetic unity with those prestarting vozbuzhdeniye, to-rye were prepared by set of the previous irritations. Prestarting excitement though also do not cause conditional reaction, but define its form and volume of identification. This hidden system of vozbuzhdeniye prepared by various previous conditions was called «prestarting integration», i.e. the system capable to create reaction as soon as the corresponding starting incentive works (P. K. Anokhin, 1949).
It is possible to give many such experiments where there is a similar transfer of ability of formation of conditional reaction from an incentive to a situation. To one of the most striking examples of it serve the phenomenon of a dynamic stereotypy (according to I. P. Pavlov, 1932). In experiences with
a dynamic stereotypy clearly came to light that in a conditioned excitator there are at least two independent parameters and that the conditioned excitator is included in some big system of the relations, under the known circumstances the prevailing value in definition of quality of conditional reaction can have edges.
Even more defiantly this heterogeneity of the afferent vozbuzhdeniye which are synthesized before formation of the behavioural act was shown in I. I. Laptev's experiences, to-ry connected the same irritant (call) with two various unconditional irritants. In the morning the call was supported with food; in the evening the same call was supported with influence of electric current. Experiments of this kind at an animal were resulted by a state, at Krom the same irritant (call) gave strictly various effects depending on when it was used. In the morning saliva was emitted for a call at an animal and it showed obviously feeding response while in the evening on the same call the animal gave clear defensive reaction (drew aside a paw). Here too difficult synthesis of starting excitement (call), obstanovochny irritation from all situation of an experiment is available and, at last, from time of statement of an experiment, a cut and creates domination of prestarting integration.
In these cases the qualitative party of a conditioned excitator as its alarm value in the form of an incentive of secretory or defensive reaction is defined by the synthetic education connected with character of earlier previous and fixed vozbuzhdeniye completely depreciates. Thus, and here, as well as in case of a dynamic stereotype, the final effect is result of integration of different types of afferent vozbuzhdeniye: starting and earlier created prestarting integration. Nek-ry researchers at the description of the facts concerning prestarting integration used expressions: «cortical switching» (E. Konorsky, 1936), «cortical adaptive domination» (L. S. Gambaryan, 1962), etc.
Prestarting integration, being the important mechanism in complete activity of a brain, defines variability and dynamism F. page. Thanks to prestarting integration relative dependence between a conditioned excitator and response to it is established. T. the page becomes variable and dynamic not only in a receptor part (concerning set and structure of afferent irritations), but also in an effector part (concerning motor categories of the central integration).
Hardly there can be a doubt that strict comparison of all changes of these various afferent signals represents complex synthetic process, in to-ry all parts of a brain are involved. A. I. Shumilina's experiments showed that frontal departments of a cerebral cortex have ability to hold different types of afferent influences in a certain synthetic association. Experience was as follows: having developed at a dog conditioned reflexes of distinguishing of two parties of the machine in response to two various conditioned excitator, A. I. Shumilina deleted frontal departments of bark of big hemispheres (the 6th and 8th fields of Brodmann) at an animal. After thatwhich operations the animal continuously passed from a feeding trough to a feeding trough, making the pendulum movements, but did not stop long at one of them. There was an impression that the animal ceased to distinguish qualitative features of starting and total obstanovochny influences.
Usually not operated animal in the conditions of this experiment behaves quietly, in intervals between uslovnsh irritants sits in the middle of the machine and runs up to a feeding trough only in response to action of a conditioned excitator — a sound of a starting incentive. In it afferent synthesis also affects, thanks to Krom the experimental animal strictly coordinates the behavioural opportunities according to this situation. Therefore, the changes of behavior which occurred after an extirpation of frontal departments of a cerebral cortex are result of disintegration of this extensive complex of afferent vozbuzhdeniye, each of to-rykh reflects this or that specific party of nervous activity. The sense of disturbances of this afferent synthesis consists that obstanovochny irritations, action to-rykh was usually detained before emergence of a starting conditioned excitator, after removal of frontal departments of a brain become starting incentives.
Thus, there are all bases to allocate four forms, or a fragment, afferentation, from to-rykh there is a stage of afferent synthesis:
1) the dominating motivation; 2) obstanovochny afferentation; 3) starting afferentation; 4) use of devices of memory.
1. The dominating motivation is a necessary component of any behavioural act because it is always directed to creation for an organism, proceeding from given it states, enough favorable conditions for existence. The behavioural act always satisfies any internal requirement of an organism (for the person and social). Participation in afferent synthesis of motivational vozbuzhdeniye is how important (see Motivations), it is possible to see at least that the conditioned food excitator is not capable to cause uslovnoreflektorny effect if the animal is densely fed i.e. if it has no ascending activating excitement going from a hypothalamus (see) and a reticular formation (see) a cut creates at the level of a cerebral cortex peculiar and always an electoral system of vozbuzhdeniye (P. K. Anokhin, 1962; K. V. Sudakov, 1963).
Motivational excitement plays especially important role in formation of a stage of afferent synthesis as in general it is difficult to imagine any behavioural act without the corresponding premises like motivation. Such motives can have various character, they can be caused as the exchange and hormonal processes happening in a body of an animal (or the person), and on higher, including and social, level (in the form of urgent needs to commission of specifically human behavioural acts). Practically any external information getting to c. the N of page, is inevitably compared and estimated on scales of the motivation dominating at present, edges is as if the filter which is selecting necessary and discarding unnecessary is more faithful inadequate, for initial motivational installation. In other words, at each this moment the importance of this information for the dominating motivational vozbuzhdeniye is defined.
From told it is visible that even active selection of the external information by means of approximate and research reaction can go over with the greatest success only because each fragment of this information is compared with the dominating excitement, a cut it is created by this motivation. This process of comparison is not simple. Using terms of cybernetics, one may say, that «search of information» shall go in the direction of the greatest suitability of the obtained information for implementation of this motivational installation. Thus, speaking about motivation as about a fragment of afferent synthesis, it is necessary to remember that it plays a preferential role in selection of information necessary for development of the decision to action and obtaining the corresponding adaptive effect.
2. The Obstanovochny afferentation combines not only a stationary situation, in a cut this or that behavioural act, but also a number of consecutive afferent influences is carried out. Set of these irritations creates in each separate case a peculiar prestarting integration of vozbuzhdeniye, edges though is in skrytokhm a state, can be immediately revealed as soon as works starting an irritant.
3. Starting afferentation. Fiziol. the sense of a starting afferenta-tion is that she dates and reveals set of the hidden vozbuzhdeniye by a certain moment, the most favorable from the point of view of efficiency of adaptation.
4. Use of devices of memory. Afferent synthesis would be impossible if set of obstanovochny and starting irritations was not closely connected with the last experience of an animal postponed in devices of his memory (see). On the basis of these mechanisms those fragments of last experience which are capable to enrich the behavioural act will be mobilized and to make it the most exact. All this makes one of intimate mechanisms of afferent synthesis.
The stated above mechanisms could not make, however, synthetic processing of information inflowing in a brain if insufficiently accurate information was not replenished with active process of approximate and research reaction all the time. Only at continuous toning of bark of big hemispheres from a reticular formation and a hypothalamus there is possible an association of not combining earlier afferent vozbuzhdeniye and formation of that «decision», a cut in the broadest sense conforms to requirements of a specific situation and the true purposes of behavior.
N eyrofizio logical substrate
of afferent synthesis. Studying of a stage of afferent synthesis gives the chance strictly scientific fiziol. explanations of those concepts and processes of «expediency», to-rye were a prerogative of idealistic psychology for a long time. Allocation of a stage of afferent synthesis made unnecessary any references to «spontaneous» processes in c. N of page. Thanks to afferent synthesis each adaptive reaction directed to obtaining specific result to the set purpose, can be considered only as equally effective various processes with various share of each of four ingredients of afferent synthesis.
From this point of view is of particular importance detailed fiziol. analysis of all mechanisms of afferent synthesis. On this way we meet exclusive complexity and plurality of interactions of specific processes at the cellular level. One is undoubted: afferent synthesis cannot take place without interaction of all those vozbuzhdeniye, to-rye are born in receptor devices, arise at the subcrustal level and then in various combinations rise to cells of bark of big hemispheres. Exactly here, at the level of a cerebral cortex, there is that fullest synthetic interaction of the afferent ascending vozbuzhdeniye, as a result to-rogo the purpose consisting in receiving these, but not other results forms.
After J. Moruzzi and X. By Megun (1949) the so-called nonspecific activating action of a reticular formation on a cerebral cortex was opened, it became clear that bark of big hemispheres receives very diverse on quality and localization of excitement. These excitement, being addressed to the same cells of a brain, create something new, the interactions of these vozbuzhdeniye which are an organic consequence at molecular level here. One of the most considerable achievements in this area is that amazing plurality of the ascending vozbuzhdeniye was accurately realized. And, if very limited irritation is even applied on receptor devices of a body (e.g., flash of light), then, reaching subcrustal devices, it undergoes in them broad dispersion, exciting the major subcrustal kernels. Only after such dispersion and specific processing of excitement come to bark of big hemispheres as the separate localized flows, and in a generalized look. Practically each cell of a cerebral cortex through certain synoptic organizations is involved into action by the vozbuzhdeniye coming from below and becomes the participant of big system of vozbuzhdeniye. At this moment bark is not diffuzno and evenly excited. Almost each site of a brain and each synoptic organization of bark take extremely individualized part in work. Therefore there are bases to think that the final result of these various ascending vozbuzhdeniye is a certain statistical system from the numerous uneven vozbuzhdeniye developing in various departments of a cerebral cortex.
Interacting excitement do not form any uniform and isolated «center of excitement», about the Crimea connected in the main concepts of cortical activity very much. Became obvious that the leading role is played by not some one initial «center of excitement», and extensive system of the interacting diverse vozbuzhdeniye having various specific weight. To imagine surprising variety of synthetic work of a cerebral cortex, it is necessary to remember that the majority of subcrustal educations (a reticular formation, a hypothalamus, thalamic kernels, a hippocampus, an amygdaloid kernel, a body having a tail, etc.) send the flows of vozbuzhdeniye converging often on the same cells of bark to a cerebral cortex.
Convergence of vozbuzhdeniye on the same neurons, apparently, is the central mechanism, without to-rogo there cannot be an afferent synthesis because convergence provides interaction, comparison and synthesis of vozbuzhdeniye in an axoplasm of nervous cells. How it occurs what cellular processes correspond to it and what excitement, coming from subcrustal devices, enter at the same time interactions?
Great attention is drawn to one of the important parties of afferent synthesis at the level of a cerebral cortex — to biol. modalities of those ascending vozbuzhdeniye, to-rye come into contact and synthetic interaction at the level of a cerebral cortex. If activation of bark is diffusion and nonspecific, then how obviously differentiated state in case of food, painful or defensive activation is created? The first observation was made on the animal who was under an urethane anesthesia. It turned out that at effect of this anesthetic suppressing the ascending activation of wakefulness are possible free carrying out excitement to the level of bark and activation of bark at pain stimulation. Thus, this experiment of V. G. Agafonov (1956) led to, apparently, paradoxical conclusion that there are at least two types of the ascending aktivation having various substrate from the point of view of their neurochemical specifics. Further experiments showed that the cerebral cortex receives not monotonous excitement, and, on the contrary, in each case is generalized is activated by the specific ascending vozbuzhdeniye. These excitement, forming at the level of subcrustal devices, already here get the biol. specifics, edges radically defines destiny of the ascending activating vozbuzhdeniye at the cortical level.
With discovery of specific character of the ascending vozbuzhdeniye there was obvious a process of selective involvement of synoptic educations on a body of a cortical cell according to biol. a modality of the ascending activating excitement that has a direct bearing on a problem of afferent synthesis. It is known that each cell of a cerebral cortex prishshat on the body and shoots in srednvkhm several thousands of various vozbuzhdeniye coming from thousands of other nervous cells (see the Synapse). Such cross supply with synoptic formations of cortical cells creates vast zero possible interactions. If to add to it not less diverse world of the subcrustal ascending vozbuzhdeniye, to-rye converge to the same cortical cells, then all this combined and will make that arena, on a cut process of afferent synthesis is played. What chaos would arise in activity of the person and his behavior if all these diverse bonds between nervous cells were excited is generalized, «diffuzno» and «nesiyetsifichesk»! However it occurs not so, and on the basis of full selectivity in distribution of the activating influences. The ascending activating influences selectively involve in the sphere of the action those nervous elements of bark and synapses on them, to-rye historically, i.e. in phylogeny and ontogenesis, are connected with data specific biol. state.
Experiences showed that the subcrustal devices of a brain mobilizing the inborn and acquired reactions various biol. qualities, have various biochemical features which are closely connected with specifics of their metabolism.
Saying that cross convergence of vozbuzhdeniye happens on various neurons of bark, it is necessary to recognize that the final stage of this synthesis consists in molecular reorganizations of those chemical ingredients of protoplasm, to-rye are inevitably involved in work as sub-sinaptichesky reactions. Therefore, in afferent synthesis as universal process it is necessary to distinguish both architectural and physiological patterns, and the molecular nature of processes which is its cornerstone.
The important role in implementation of afferent synthesis, undoubtedly, is played by also huge structural variety of cortical cellular elements (pyramidal cells of all sizes and layers, star-shaped cells, cells of a granular layer etc.). In implementation of afferent synthesis also the interkortikalny bonds combining the remote areas of a brain have serious value intra-.
The huge role in the mechanism of afferent synthesis is played by the approximate and research reaction arising against the background of and by means of the activating ascending influences from a reticular formation. Direct researches of influence of irritation of a reticular formation on cortical discrimination, i.e. sharpness of distinguishing, shows that at the activating influence of a reticular formation cortical processes proceed with bigger discrimination of external irritations, than without this stimulation. Besides, increase in the activating action from a reticular formation at the time of emergence of approximate and research reaction facilitates interaction between separate parts of bark through intra-and interkortikalny bonds. It is clear, that this second value of approximate and research reaction is of special interest as the main line of afferent synthesis is comparison and integration of the cortical processes remote one from another. The sense of approximate and research reaction at this stage consists in tokhm that in the course of continuous and active selection of external afferentation to make comparison of this under-birayekhmoy afferentation with the main dokhminiruyushchy motivation and with elements of last experience.
«Decision-making» as nodal mechanism of functional system. Odnikhm from the most remarkable moments in formation of the behavioural act is «decision-making» to commission of it, but not other action.
«Decision-making» is logical process F. page; in the same vrekhmya it — result quite certain fiziol. influences, to-rye shall be still studied. Tvkhm not less already formulation of the question about «the decision» is useful since it pokhmozht to find fiziol. an equivalent of this of a veskhm of the focused integration at the same time and to find out a role of nek-ry nervous elements with very specialized functions. The main objective sign of the specific state undergone by c. the N of page in this mokhment, consists that the organism inevitably shall make the choice of the unique possibility of behavior from numerous opportunities, to-rymi it has at each this moment. In relation to muscular system this process is called by A. A. Ukhtomsky (1945) «elimination of excess degrees of freedom». Process of «decision-making» inevitably is also the choice of one certain form of behavior. This choice can be made or on much the detained stageand afferent synthesis with inclusion of consciousness or to proceed instantly avtomatizir about in and other p at subjects.
Most clearly the stage «decision-making» is observed at use of a method of the active choice in specially designed two-sided machine with two feeding troughs. In nek-ry cases animals in response to a conditioned excitator sit in the middle of the machine for a long time. However on povoro-takhm the heads alternately in right, to the left side with obvious visual fixing of one and other feeding troughs it is possible to judge that there is an active selection of additional information and that the stage of afferent synthesis did not end. But in some mokhment of this expressly of approximate and research reaction the animal quickly rises from the place and Ikhmenno to that korkhmushka goes, edges it is signaled by this conditioned excitator, and already here waits for giving of a forage.
The same sort the phenomenon is probably developed in c. N of page of animal N at the moments which received the name «ideatsshg>, t*l of «a state like eureka». Eo all these cases active afferent selection of the maximum quantity of parameters of the given situation with the help of approximate and research reaction comes to an end with the adequate behavioural act. It is also turning point for formation of an efferent part of the behavioural act. The second essential objective sign of «decision-making» — a ratio between very large volume of the initial afferentation used in a stage of afferent synthesis and strictly certain limited quantity of the efferent vozbuzhdeniye joining after «decision-making» in formation of the behavioural act.
As typical model of such ratios the neuron, more precisely, a ratio of details on «entrance» and at «exit» of neuron can serve. Still Ch. Sherrington (1906) paid attention that the motor neuron represents «the general way» for numerous vozbuzhdeniye, however to his axon there is only one excitement quite specific in parameters of single irritation for this neuron. The same circumstance was emphasized by E. Adrian (1947), having called this transitional moment «homogenization of vozbuzhdeniye». Not surprisingly therefore that many neurophysiologists came to need to allow existence of a stage «decision-making» and therefore consider the described feature of neuron as demonstrative model. To the same neuron can converge the most various forms of vozbuzhdeniye. Already this one fact in itself assumes that in neuron there is some extremely complex work like afferent synthesis, to-ry comes to an end with «decision» on generating point: to send categories of such, but not other configuration to an axon.
So, «decision-making» represents critical point, in Krom there are a bystry release from excess degrees of freedom and the organization of a complex of efferent vozbuzhdeniye capable to provide quite certain action. Practically during the formation of the behavioural act of an event in the whole brain develop as well as in single neuron. But in all cases to «decision-making» there corresponds afferent synthesis because it in many respects defines formation of the subsequent stages of the behavioural act. Anyway, afferent synthesis on the scale of the whole brain inevitably comes to an end with «decision-making», i.e.
selective excitement of such complex of neurons, to-ry can create the only behavioural act adequate to results of this afferent synthesis on the periphery.
How universal character has «decision-making» in functions of an organism, it is possible to see not only under behavioural acts, but also on the example of purely vegetative functions. So, the level of lung ventilation (see), in particular amount of the air which is taken away by lungs, at each this moment is exact reflection of needs of an organism for receiving oxygen and release of carbon dioxide gas. Any change of this requirement immediately is implemented in reduction or increase in lung ventilation. It is obvious that final motor neurons of a respiratory center receive and give the «team» which is precisely reflecting requirements of an organism on the periphery. However this requirement difficult. It includes several components, to-rye shall be integrated, and only after that the final motor neuron of a respiratory center receives quite certain «decision»: «to take» 400 or 600 ml of air.
Results of action as independent physiological category. Immediately after «decision-making» the integral (sum) of efferent vozbuzhdeniye forms, to-ry provides peripheral action. The central integral of efferent vozbuzhdeniye (or «the program of action») corresponds the most exact image to flows of the efferent vozbuzhdeniye leaving to numerous peripheral bodies. These bodies can be in very big distance from each other that is in direct dependence on biomechanics of action.
Results, dependent on it, are just the same inevitable and natural consequence of the made action.
It seems strange that for many years results «actions never acted as an object special fiziol. the analysis, as a decisive link between a reflex effect and formation of the subsequent stages of the behavioural act. As a matter of fact the reflex, the reflex act or a reflex effect are of interest only to the researcher — the physiologist or the psychologist. Results of action are always interesting to an animal or the person. Only for the sake of them and concerning them often very long chains of behavioural acts are undertaken and results only received become an incentive for commission of new and new acts until reached comes to this or that compliance with желаемь^м.
Results of action only then will become the real fact allowing their scientific analysis when in each separate case all those parameters of results are listed most fully, to-rye in the set define afferent information on the received results.
Such understanding of business becomes especially necessary because parameters of results inform a brain on usefulness of perfect action and make in general the return afferentation, i.e. a peculiar afferent integral, afferent model of results. If action terminates in the fact that the person gave a hand to a subject, then results consist that the person took a subject in a hand. Here immediately the complex of afferent parameters also forms, to-ry represents afferent model of results (weight of a subject, tactile properties, temperature, a form).
Formulating the principle F. page as unit of integrative activity of an organism, first of all it is necessary to pay attention to information on results of action. Proceeding from purely fiziol. reasons, this information on results of action was called by P. K. Anokhin (1935) authorizing, or the return, afferentatsiy. Thus, cybernetic «closed path» with «feed-back» in physiology was more exhaustively formulated and developed in details long before emergence of the first works of N. Wiener on cybernetics (see). Already then among mechanisms F. page the return afferentation about parameters of results of action was of particular importance. It allowed on a real fiziol. to a basis to expand structure of «reflex arc» and to construct such fiziol. architecture, in a cut all nodal mechanisms are connected in system of the processes combined by the cause and effect relations.
The return afferentation as a form of afferent impact on an organism is of special interest as changes ideas of mechanisms of organized behavior of an animal. Being an analog of «feed-backs» in cybernetics (see the Feed-back), this type of an afferentation in physiology and medicine, naturally, draws close attention to itself. Pointed to a certain role of muscular afferent impulses still Ch. Bell (1824) and I. M. Sechenov (1863). However these first instructions connected generally with a muscular afferentation (see Pronriotseptora), did not create system of ideas of a role of the return afferentation, about results of action. These should explain the fact that in physiology the concept «reflex arc», a cut on the most fiziol still continued to dominate. the essence is not enough for an explanation of behavioural acts on the basis of the return information on results of action.
The sense of the return afferentation consists that in any fiziol. process or in the behavioural act of an animal the return afferentation informs on results of perfect action, giving the chance to an organism to estimate degree of success of the operation performed by it. So, e.g., the person on the basis of a number of objective processes has interactions of an organism and the environment intention to drink a glass to tea can ripen. He gives a hand to a glass with tea, takes it. Then tactile, temperature, weight, at last, visual stimulation from contact of a hand with a glass — all these afferent irritations in the sum give information
on as far as the result of action corresponds to initial intention. However at implementation of this action advance of a hand to a glass is continuously regulated by the proprioceptive alarm system testimonial of the correct and corresponding distribution of the reduced muscles, about rate of strain of a hand, about height of its situation in relation to the planned purpose etc. This last form of an afferentation, undoubtedly, is very important for implementation of the movement of a hand, but it on the most essence cannot give c. N of page of any information on results of this action, as only one position of a hand and consequently, only a proprioceptive afferentation whether does not inform on that, e.g., the necessary subject is taken in hand. This circumstance, unfortunately, is a little considered at assessment of the return afferentation in an organism and therefore often it is necessary to see that attempts of comparison of «feed-backs» in organisms and cars begin with the proprioceptive alarm system, and sometimes and come to an end with it.
Thus, the return afferentation arising at any motive act should be divided into two absolutely various categories: a) directing to the movement and b) a productive afferentation. While the first is presented only by proprioceptive impulses from the muscles which are carrying out the movement the second always in a complex and covers all afferent signs concerning the result of the undertaken movement.
In life of an organism, especially in human life, there are no actions, to-rye would not follow from the previous actions and would not cause the subsequent actions. Naturally therefore that the corresponding differentiations depending on with what return af-ferentation we deal shall be entered into a concept of the return afferentation: whether it concerns information on results of some intermediate action or informs on final performance of the initial intention which is logically completing the big behavioural act.
So, e.g., if the person, being at home, set a goal to make some purchase, then after emergence of this intention a number of separate actions is developed: the person puts on,
checks existence of money, opens doors, comes downstairs, crosses the street, opens a door of shop, chooses a product necessary to it etc.
Thus, intention of the person to buy something includes a number of buffer stages, each of to-rykh could not be carried out if the person ke received at each such stage of the return afferentation from its successful completion.
It follows from this that all category of the return productive afferentation should be subdivided into two separate forms: the step-by-step return afferentation, edges corresponds to implementation of a certain stage of this behavioural act, and the authorizing return afferentation, edges fixes the most successful integration of efferent vozbuzhdeniye and completes a logical functional unit of behavior (e.g., «I want to drink» — «got drunk»).
As criterion for such distribution of the return afferentation serves intention, a cut is result of afferent synthesis and a cut the person wants to carry out at present. The quantity of stages of performance of intention depends on the width of a task and on the nature of each action; in some cases the step-by-step afferentation can be and authorizing, and final.
Really, purchase something in shop is not the end of all activity of the person at all; it can be only a stage in implementation of longer chain of acts, napr, trips to other city.
Here it should be noted that the return afferentation on the most essence is an afferentation about results of action and, therefore, very close corresponds to that information, edges continuously arrives from an adjustable object through feedback channels in difficult cars with automatic regulation (see the Feed-back, Regulation).
Prediction and control of results of action. The formulation of the purpose of action represents critical point in development of the behavioural act. Only after that formation of a difficult complex of efferent vozbuzhdeniye shall begin, to-rye, being distributed on work vehicles, define obtaining the result which is precisely answering a goal. From the point of view of a being of this transitional moment it is important to emphasize that there is still obscure transformation of results of afferent synthesis in very adequate distributions of efferent vozbuzhdeniye on working bodies. In a phase of expansion of the behavioural act also others are put in action extremely the important mechanisms which are formed also immediately after the moment of «decision-making». As soon as the purpose of action and its program forms, at once, along with an exit of excitement to effector devices, the complex of vozbuzhdeniye, fiziol forms. the sense to-rogo consists that with its help assessment of that information is made, edges will come to c. N of page from future results of action. It is the original device of assessment and checking of results with a goal.
Tell experiments with substitution of an unconditional reinforcement about existence of such device. If at development of food conditioned reflexes the dog constantly received for a number of years a reinforcement in the form of 20 g of crackers, then substitution by their meat leads to emphasized approximate and research reaction and temporary failure from food. What origins of such approximate and research reaction? The analysis of this question convinces that it is about use of the conditioned excitator promoting formation * not only a complex of efferent vozbuzhdeniye on a basis to-rykh all forms a mustache a lovny feeding response, but along with it is a complex of the afferent vozbuzhdeniye corresponding in the parameters to a komplekv of signs and qualities, the forthcoming result, i.e. a look, a smell, tastes of crackers, etc. In order that the feeding response of an animal ended systematically and standardly, in c. N of page of an animal the irritation from outward of crackers shall arrive and come there into some contact with an afferent complex of checking, to-ry the nose le of the beginning of action of a conditioned excitator, but prior to the most conditional reaction was created at once. External signs of meat, of course, too irritate receptors of a dog. However the complex of the central vozbuzhdeniye created under the influence of meat was inadequate, not to the corresponding most probable vozbuzhdeniye from crackers. Occurred ^рассогласование» between the prepared complex of afferent vozbuzhdeniye and the real complex of vozbuzhdeniye which came to c. N of page.
On the basis of these experiments of a .voznikl the hypothesis of existence of the special afferent mechanism, to-ry forms earlier, than action will be made and there will be a result, but at the same time comprises all signs of these future results. This device which received the name of an acceptor of results of action (see) is intended for perception of information on the received result and its comparison with those parameters of results, to-rye developed even at the time of action of conditional irritation.
Special experiences with registration of EEG were put on the person: system from three irritants — a siren, light, it is ringing — it was repeatedly shown to the examinee to the person for several hours every day. From these three irritants only «light caused desynchronization of electric activity of a cerebral cortex (see Elektroentsefalografiya). It also was an episeme of what will occur in visual area of bark at substitution of irritants. The substitution of a call to the place of light made both in a stage of development of a stereotype, and in that stage when the stereotype from three irritants was already rather strong recorded was the second stage of experience. Experience yielded the following result: when instead of light it was rung, in visual area of bark there was still a desynchronization as though light was given, ?a is not ringing. The analysis of EEG showed that desynchronization occurred even slightly earlier, than was rung. It follows from this that excitement came to the visual area earlier, than the external irritant could come there.
It is obvious that at the organization of a chain of irritations excitement extends on a brain from point to point much quicker, than ^реальные consistently appearing external irritants. Excitement advances a real irritant, to-ry shall work only in the future on c. the N of page, also occupies those areas of bark, to-rye it in the future shall excite. These excitement, playing an adaptive role, received the name of «the advancing vozbuzhdeniye».
Ability of nervous substrate to organize a chain of processes with the advancing excitement is that elementary, but historically very ancient basis, on a cut both the conditioned reflex, and any prediction of future phenomena develop. Became obvious, as approximate and research reaction of an animal to substitution of bread meat could arise only because excitement from the conditioned excitator signaling about bread creates an afferent complex in those cells of bark where it shall be created only in the future, i.e. at the time of use of an unconditional irritant. Meanwhile afferent parameters of meat were incoincident with these «the prepared vozbuyaedeniye».
The principle of development of the advancing vozbuzhdeniye is a consequence of the constitutional properties of nervous tissue and therefore it takes place everywhere where there is a need of «precautionary reaction» (speaking I. P. Pavlov's language). Practically it «predicts» probable results of action at this solution and this purpose of action. At the same time a complex of vozbuzhdeniye, in Krom are coded properties of future results, completely provides comparison of the received results to what is set, or with set of signs of this situation. The device of an acceptor of results of action has universal distribution. Without preliminary formation of this device even the simplest behavioural act is hardly possible any.
Special researches of a respiratory center showed, as it has the same functional organization. Each efferent volley of nervous impulses which is coming out a respiratory center to respiratory muscles precisely reflects the current needs of an organism for lung ventilation (see Breath). Information on this requirement consists of all afferent indicators of breath (concentration of carbonic acid and oxygen in blood, an afferentation from vascular hemoretseptorny areas etc.), as makes a peculiar afferent synthesis respiratory F. page. On this basis the size of volley of efferent vozbuzhdeniye forms and always is prichinno connected with the current needs of an organism for breath. Ability of this volley to cause reduction of respiratory muscles of this or that force is exact reflection of results of afferent synthesis about the necessary level of lung ventilation. As soon as as a result of reduction of respiratory muscles lungs begin to be filled with air and alveoluses begin to extend, afferent signals from their mechanioreceptors immediately come to a respiratory center (see), precisely reflecting the amount of the taken air prompted by results of afferent synthesis. It is possible to prove that the respiratory center, having sent volley of efferent impulsation to respiratory muscles, at the same time creates also the supervisory body of information on the forthcoming results, i.e. an acceptor of results of action. This device on the most fiziol. essences are designed to receive afferent to them - a pulsation from the extending lung, from receptors of its alveoluses and to compare a volume equivalent of this to them - a pulsation to a volume ekvivalentokhm of the efferent team sent to the periphery.
Thus, in all cases of parcel by a brain of vozbuzhdeniye through final motor-neurons to peripheral work vehicles along with efferent «team» the afferent model capable to anticipate parameters of future results forms and to compare at the end of action this prediction to parameters of true results. The prediction of results of action is general function of a brain, warning any «mistakes», i.e. a fulfillment of the actions which are not answering the object set by an organism and the decision to action.
This pattern is of particular importance in difficult behavioural acts of the person, in the course to-rykh the most various objects of behavior — big and small can be set, and nevertheless the acceptor of results of action forming also at the time of decision-making also defines afterwards hit rate between conceived and received. It belongs also to the speech, to its neurophysiological structure. Here the analysis in the same direction is necessary. It is necessary to remember that «decision» to tell any phrase or to state judgment develops absolutely as well as any other «decision», i.e. after a stage of afferent synthesis. «Decision» to state any judgment, as well as in all other cases, creates an acceptor of results of action with all afferent signs inherent in it. Therefore, here not, as usual represent, formations of each word separately, only the acceptor of results of action on each phrase with the sequential arrangement of words, sometimes even with the advancing sense is created that is a sure sign of formation of an acceptor of results of action on the whole semantic system.
All nodal mechanisms of the behavioural act represent fiziol. unity, and any of these mechanisms it is impossible to consider separately, without imagining all architecture of the behavioural act, and, above all — a specific role of this mechanism in expansion of processes in F. page.
Functional systems of an organism. Functional systems — units of complete activity
of an organism. They represent the dynamic self-regulating organizations forming on a metabolic basis or under the influence of factors surrounding,
and at the person (in the first eyes
red) — the social environment. Everyone F. the page by means of nervous and humoral control selectively
combines various bodies and fabrics for ensuring adaptive results, useful to an organism. From positions of the theory F. the page is lost the traditional organ principle of creation fiziol. functions. The same bodies and fabrics combine various F. page for ensuring specific results of their activity.
The useful results, adaptive for a human body, creating F. pages of various level, are: a gomeosta
the tichesky results defining the course of various metabolic processes, optimum for normal life activity, in body tissues; the results of behavioural activity of animals and the person satisfying them the leading metabolic and biological requirements; results of gregarious (zoosocial) activity of animals; results of mental activity of the person; results of social activity of the person.
The variety of adaptive results, useful to an organism, indicates that number F. the pages making various parties of life activity of the whole organism it can be extremely big. One F. pages determine by the activity various indicators of internal environment of an organism (see the Homeostasis), others — behavioural activity and interaction with the environment. Pages are genetically determined by Nek-rye F., others develop in process of formation of separate requirements (especially behavioural) of the person.
Everyone F. the page of various level of the organization is based on the principle of self-control: deviation of result of activity F. the page from the level providing normal metabolism (life activity) of an organism itself is an incentive to mobilization of necessary elements of system for ensuring this result. It belongs both to behavioural, and to social F. page, activity to-rykh it is directed to achievement by the subject (or community) it is biological or socially important results in to the environment.
Any F. page, according to P. K representations. Anokhina, has essentially same organization and turns on the peripheral and central nodal mechanisms, following general universal for different systems: 1) useful adaptive result as leading link of functional system; 2) receptors of result; 3) the return afferentation going from receptors of a result in the central educations F. page; 4) the central architecture representing electoral association F. page of nervous elements of various levels; 5) the executive somatic, vegetative and endocrine components including organized purposeful behavior.
Essentially identical organization various F. the page in an organism characterizes their isomorphism.
Various F. villages for achievement of various adaptive results can use various or same internals. So, cardiac performance can be used as for maintenance of a fixed level of blood pressure, and for ensuring gas exchange etc. Work of a kidney, except secretory function, can be used for ensuring the pH levels of blood, osmotic pressure, body temperature etc., optimum for metabolism.
In everyone F. the page is also available a possibility of interchangeability, mutual compensation of their effector mechanisms. At failure of one or several executive components of everyone F. the page ensuring its net adaptive result can be carried out by other components entering it. These system processes it is extremely important to consider at compensation of the broken functions of an organism (see Kompensa-torn ye processes).
Unlike classical physiology, edge studies mechanisms of life activity of separate bodies and dynamics of their work, physiology F. the village studies an organism as set of the system organizations in their dynamics and interrelation.
At the same time inclusion of separate bodies in F. the page occurs by the principle of mutually assistance, to-ry is carried out for achievement of useful result, adaptive for an organism. Selective mutually assistance of fabrics clearly is traced, e.g., in mechanisms of inclusion of various hemadens in different F. page. In everyone the funktsiona to lny system is available the endocrine «arrangement» providing achievement of result of activity of system. In the whole organism mutually assistance various F. the page is under construction on the basis of the principles of their hierarchy and multicoherent, multiparametrical interaction of results of activity separate F. page.
The principle of hierarchy consists in tomg that in each this timepoint you define activity of an organism yag F. page, dominating in respect of survival or adaptation to the environment. In relation to dominating all others F. pages of an organism are built in a hierarchical order according to them biol. the importance and need for social activity of the person.
After satisfaction with the activity of an organism conducting requirements the requirement, next on the biological and social importance, creating dominating F takes control. page. In relation to this F. pages others are also built in a hierarchical order etc. Change dominating F. the page on another occurs constantly and reflects essence continuously of the events metabolism and continuous interaction of an organism with the environment. Hierarchical interaction various F. the page is under construction on the basis of the principle of a dominant opened in our country by A. A. Ukhtomsky (see). However the principle of a dominant, from the point of view of the theory F. the page, gains further development.
M at l t the SP and r and mt r iches to y p r intsshg interactions of various functional systems defines their generalized activity. As a rule, change of one indicator, result of activity any» functional system immediately affects results of activity of others F. page.
The principle multiparametrical» interactions clearly comes to light, e.g., in activity F. page, the blood defining optimum level of gas structure. In such F. the page is at the same time carried out regulation of several interconnected respiratory indicators (pH, rs02, r02).
For F. page of multicoherent interaction other principle of self-control is characteristic qualitatively»: the deviation from optimum level of this or that parameter of result is an incentive to the directed redistribution in certain ratios of values of all other parameters of result this F. page.
In uniform generalized F. pages can enter separate F. the pages providing various indicators of a homeostasis of internal environment of an organism among them: functional system of maintenance of optimum sizes of hemodynamic indicators (see the Hemodynamics), functional system of maintenance of optimum level of respiratory indicators of an organism (see Gas exchange, Breath), functional system of ensuring water-salt balance (see the Water salt metabolism), functional system of regulation of a blood coagulation (see. Coagulant system of blood), functional system of maintenance of optimum body temperature (see Thermal control) etc. Besides, generalized F. the page can develop from F. the page, one of to-rykh provide separate homeostatic indicators of internal environment of an organism, others are directed to achievement of this or that behavioural result satisfying biol. requirements of an organism.
Thus, from the point of view of the theory of functional systems, the complete organism represents hierarchy of a set F. page, acting by the principle of multicoherent interaction. Always it is available dominating F. pages and all others in relation to it are built in the coordinated order, at Krom the result of activity of the subordinated system is included into result of activity of system of more high level of domination. On the other hand, various F. pages work in a uniform organism by the principle of multiparametriche-sky interaction. It means that changes of result of activity of one of F. pages surely affect a number of results of activity of others.
The complete organism in each this timepoint represents harmonious interaction — integration (across and verticals) various F. page that defines the normal course of metabolic processes. Disturbance of this integration if it is not compensated by special mechanisms, means a disease and can lead to death of an organism.
Unlike a reflex as response of an organism to various irritations, T. pages have the closed self-regulating dynamic organization. All activity F. by page it is aimed at providing adaptive results, useful to an organism. Assessment of a net result of activity is constantly carried out by means of the return afferentation (see the Feed-back).
Representations about F. pages of an organism made the new orientation of thinking promoting understanding of patterns of the whole organism in normal, extreme and pathological conditions.
Discovery of patterns of the system organization of purposeful behavioural acts forced to recognize that the behavioural act is carried out not only by the principle of a reflex, i.e. from an incentive to action (see the Reflex), but also by the principle of self-control: deviation of this or that fiziol. an indicator in an organism from the level providing its normal life activity immediately puts the vigorous behavioural activity directed to recovery of optimum level of metabolic processes in action (see Self-control fiziologichesky functions).
Unlike a reflex, to-ry in any its form is reaction of an organism to this or that incentive, T. the villages participating in formation of a homeostasis and behavioural acts have a number of new properties. They not only react to external stimuluses, but also by the principle of a feed-back answer various shifts of the vital result controlled by them. Besides, in them the advancing valid events of reaction form, and also there is a checking (correction) of the achieved result to the current requirements of an organism.
Central organization F. page of various degree of complexity in comparison with the central mechanisms of a reflex arc is also more difficult and includes the stages which are absent in a reflex arc — afferent synthesis, decision-making, anticipation of potrebny result (an acceptor of result of action), efferent synthesis and multicomponent action. The reflex arc is only a component of the system organization fiziol. functions.
In development of the general theory F. pages are formulated new regulations
on system «quantization» of behavior, a sistemogeneza of behavioural acts, on the holographic principle of the system organization of the dominating motivations (K. V. Sudakov), an age and population sistemogenez (H. N. Kokina), about plasticity F. page depending on effectiveness of behavioural acts (A. V. Kotov).
Theory F. by page it began to be used widely in physiology of work and sport, in philosophy, medicine, pedagogics, psychology etc.
According to the general theory of functional systems all-Union seminars are regularly held: first seminar «Systems analysis of mechanisms of behavior» (Moscow, 1975); second seminar «Structural funktsiona-nye of pattern of a sistemogenez» (Moscow, 1976); the third seminar devoted to the 150 anniversary since the birth of the prof. A. I. Babukhin, «System properties of the fabric organizations» (Moscow, 1977); the fourth all-Union seminar devoted to the 80 anniversary since the birth of P. K. Anokhin, «System mechanisms of emotional reactions» (Moscow, 1978); fifth seminar «Integrative activity of neuron» (Moscow, 1979); sixth seminar «System mechanisms of motivation» (Suzdal, 1982).
On various problems of the theory F. pages are held also international symposiums and conferences: first Soviet-Yugoslavian symposium «Principles and mechanisms of formation of functional systems of an organism» (Novi Sad, Yugoslavia, 1977); second Soviet-Yugoslavian symposium «Principles and mechanisms of formation of functional systems of an organism» (Moscow, 1979); the French-Soviet-Bulgarian symposium on a yeyrofiziologiya devoted to P. K. Anokhin's memory — «The theory of physiological systems and its use in clinic» (the Tour, France, 1981); the international Soviet-American Pavlovsk conference devoted to P. K. Anokhin's memory, «Emotions and behavior: systems approach» (Moscow, 1984).
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