RETICULAR FORMATION [formatio reticularis (PNA, JNA, BNA); synonym: reticular substance, setevidny education] — set of the structures located in the central departments of a spinal cord and a brain trunk. For R. f. existence of a large amount of the nerve fibrils passing in the most various directions in this connection this education under a microscope reminds a reticulum is characteristic. It formed the basis for Deyters (O. F. Page of Deiters) to call it mesh education.
at the end of 19 — the beginning of 20 century were described outside department of R. f. a myelencephalon, the consisting preferential of gray matter (formatio reticularis grisea), internal department consisting preferential of white matter (formatio reticularis alba), lateral department (substantia reticularis lateralis), and also separate kernels of R. f. First systematic description of kernels of R. f. a brain trunk it was given in 1909 by H. Jacobsohn.
In Mésen and Olszewski's atlas (Meessen, J. Olszewski, 1949) described 32 kernels of a rhombencephalon of a rabbit. In the work of 1954 Olszewski describes 22 kernels of R. f. a myelencephalon, the bridge and a mesencephalon at the person.
Studying of a structure of separate kernels of R. f. and the neurons making them, and also R.'s bonds f. formed the basis for R.'s division f. on zones, parts or columns.
Comparative and anatomic, morphological and ontogenetic studying of R. f. allowed V. V. Amuntsu (1966, 1976, 1980) to group R.'s kernels f. in 3 departments: outside, internal and average — and to allocate transition zones between reticular and specific, and also between various reticular structures.
Characteristic of a neural structure of R. f. it was made Shaybel and Shaybel (A. B.Scheibel, M. E. Scheibel, 1962), G. P. Zhukova (1977), T. A. Leontovich (1978), Mannen (H. Mannen, 1966, 1975), Valverde (F. Valverde, 1961), N. S. Kositsyn (1976), etc.
Nervous bonds of R. f. were studied preferential by methods of a degeneration. Fibers P. f. divide on ra-diarny, directed ventrally, fibers concentric, crossing in the field of the centerline, and fibers longitudinal, grouped in bunches. These bunches make both afferent, and efferent pathways. R. given about bonds f. are generalized in Brodal (1957), Rossi and Tsanketti's works (G. F. Rossi, A. Zanchetti, 1957).
In the field of a brain trunk anatomic educations, excitement are had to-rykh exerts generalized tonic impact on front departments brain (see). This group of anatomic educations received the name of the ascending activating system of a reticular formation (fig. 1). The important part in maintenance of the awake state, and also in mechanisms of formation complete, and in particular uslovnoreflektorny reactions of an organism is assigned to it. Along with the ascending activating system allocate also descending reticulospinal systems exerting the controlling impact on reflex activity of a spinal cord. Activity of the systems both ascending, and descending is maintained by continuous inflow of the afferent impulses coming to R. f. on collateral fibers from touch conduction paths. An important role in maintenance of activity of reticular mechanisms is played by humoral irritants, in relation to the Crimea R. f. has high sensitivity that provides its participation in regulation of a number of vegetative functions. Along with this R. f. is the place of selective effect of many pharmacological means that finds broad application at treatment of a number of diseases of c. the N of page, and also causes new approach to studying of such major problems of medicine as, e.g., a problem of pain and anesthesia. In the area P. f. there is an extensive space coincidence and interaction of the impulses arriving from various peripheral receptor educations to the vozbuzhdeniye going from a cerebellum and bark of big hemispheres. Thanks to a large number of kortiko-reticular bonds the cerebral cortex exerts the controlling impact on activity of reticular mechanisms, regulating the level of their activity.
Reticulospinal ratios. Influence of a brain trunk on a physical activity of a spinal cord was for the first time shown by I. M. Sechenov in 1862 (see. Setschenow's inhibition ). To them it was shown that at irritation of a brain trunk both braking, and simplification of reflex reactions can be observed. However the mechanism and structures mediating these influences remained obscure. In 1944 X. Megun, applying direct electric irritation of a brain trunk, showed that irritation of certain sites of bulbar R. f. leads to dead stop of the movements caused as it is reflex, and irritation of motive sites of a cerebral cortex. This braking was the general and extended to all groups of muscles irrespective of their topographical ratios and physiological function. X. Megun suggested that the controlling R.'s influence f. it is carried out at the level of a spinal cord, but not at the level of a cerebral cortex. This assumption was confirmed to them in experiences on the decerebrated animals.
Concerning the mechanism of reticulospinal influences there is no consensus. While one researchers consider that R. f. can exert impact directly on motor-neurons of a spinal cord, others assume that these influences are transmitted to motor-neurons through some interneuronels, to-rykh the internuncial neurons participating in short circuit of segmented spinal reflex arcs can play a role.
Retikulokortikalny ratios and a problem of the ascending activating influence of reticular mechanisms on a cerebral cortex. Is known the fact that the main symptoms at defeat of certain sites of a brain trunk are the general decrease of the activity of the patient, an adynamia, drowsiness. Experiments showed that the similar phenomena can develop at destruction at an animal of subcrustal and trunk departments of a brain. These data formed for row R1ssledovateley the basis to believe that in the field of a brain trunk there are centers knowing the general activity of an organism, the centers of a dream and wakefulness (see. Centers of a nervous system ). Such assumption was proved by the fact that the direct irritation of certain departments of a brain trunk could cause a dream in an experimental animal or bring him out of this state. However original progress in this problem became possible only after began to apply to a research of a brain more and more widely electrophysiologic methods, in particular electroencephalography (see). Researches B. B. Pravdich-Neminsky (1925) showed that external irritations cause the characteristic changes of EEG consisting in change of the slow high-amplitude and low-frequency fluctuations characteristic of a condition of rest, bystry low-amplitude and high-frequency fluctuations. Such changes of EEG are observed at the person or an animal upon transition from a condition of a dream to wakefulness. In this regard the similar type of reaction received the name «reaction of activation of EEG», or «reaction of awakening».
Development of experimental methods of a research allowed to develop ways of thin irritation and destruction of separate subcrustal structures by means of the electrodes entered with a big accuracy into points, necessary for the experimenter (see. Stereotaxic method ). It allowed J. Mo - ruzz and X. To approach Megun in 1949 the solution of a question about tokhm what structures of a brain are responsible for emergence of «reaction of awakening». For the solution of this task of J. Moruzzi and X. Megun made a series of experiments, in the course to-rykh established that at irritation at animal certain points of a brain trunk change of the slow synchronous high-voltage fluctuations characteristic of a dream, with low-amplitude high-frequency activity is observed. These changes of EEG were diffusion, i.e. were observed on all cerebral cortex, but were better expressed in the hemisphere, of the same name in relation to the site of irritation. Changes of electric activity of bark of big polunga-riya were followed by external signs of awakening.
By further researches it is shown that the similar phenomena can be observed at irritation of various departments of R. f. a brain trunk — beginning from oblong and finishing a diencephalon. In the area myelencephalon (see) an excitable zone matches those sites of R. f., to-rye, according to X. Meguna and R.Rhines, exert also the descending impact on activity of a spinal cord. At the level of the bridge (see. Bridge of a brain ) and mesencephalon (see) this zone is located in the field of a tire, and at the level diencephalon (see) takes a subthalamic kernel and back hypothalamus (see), reaching medial thalamic kernels. These structures of a brain trunk also make the ascending activating reticular system, the cut is anatomic substrate a number of the ascending korotkoaksonny reticular ways. X. Megun and J. Moruzzi came to a conclusion that the changes of EEG observed by them were not result of antidromic carrying out impulses to bark of big hemispheres on the known corticofugal ways. These changes could not be explained also with carrying out excitement to bark on known classical sensitive (lemniskovy) to pathways as even after section of these ways the irritation of structures of a myelencephalon continued to cause clear changes of EEG.
J. Moruzzi's representations and X. Meguna about the activating system P. f. a brainstem gained further development and confirmation in the works performed in laboratories of many countries of the world. The main conclusions of J. Moruzzi and X were confirmed. Meguna is also specified topography of the structures carried to the ascending activating reticular system of a brain trunk.
The electrophysiologic experiments made by means of microelectrode technics showed that to the same neuron of R. f. impulses from various peripheral sources and bark of a great brain can meet. Interaction between these impulses also causes, apparently, variety of observed effects of external irritations.
By Dell's works (R. of Dell) and its schools it is shown that an important role in maintenance of activity of R. f. belongs to humoral factors, in particular adrenaline. It was established that the activating influence of adrenaline on a cerebral cortex is carried out through R. f. mesencephalon and bridge (fig. 2). After section of a brain trunk of a kpereda from a mesencephalon administration of adrenaline did not cause more than «reaction of awakening». Thus, rostral departments of a brain trunk have hypersensitivity to adrenaline. Moreover, Vogt's researches (M. of Vogt, 4954) and P. K. Anokhina (1956) in rostral departments of a brain trunk revealed existence of a large number of the elements containing adrenaline and noradrenaline. I. P. Anokhina (1956) it was revealed that the aminazine having ability to block alpha and adrenergic receptors prevents development of activation of EEG at pain stimulations. It gives the grounds to assume that the activating R.'s influence f. on a cerebral cortex at pain stimulations it is carried out due to involvement in activity and - adrenergic receptors of rostral departments of a brain trunk.
On what ways influence of reticular mechanisms on a cerebral cortex is carried out? X. Megun and J. Moruzz (1949) suggested that one of ways, through to-ry R. f. exerts impact on a cerebral cortex, the group of medial kernels is thalamus (see), making so-called nonspecific projective talamokortikalny system. nucleus reticularis, the item ventralis anterior, n. centralis medialis, n. lateralis, n. medialis, n. centrum medianum, n. parafascicularis and other kernels of an intralaminarny complex belong to this system (at a cat) (intra lamellar kernels of a thalamus, T.). At irritation of nonspecific kernels of a thalamus generalized changes of electric activity of a cerebral cortex are observed irrespective of the fact which from kernels is exposed to irritation; kernels of this system have among themselves a large number of bonds thanks to what at excitement this system reacts as uniform.
To studying of functional ratios between the ascending activating system P. f. a brain trunk and nonspecific projective (diffusion) thalamic system the large number of works is devoted. Many researchers just combine these systems in one, considering their close morphological unity. However J. Moruzzi (1958) believes that for such association there are no sufficient bases and it is more reasonable to carry nonspecific thalamic system to «spheres of influence» of R. f., especially as there are experimental data testimonial of reciprocal relationship between these educations. Detailed studying of the ways connecting R. f. a brain trunk with nonspecific kernels of a thalamus, carried out Peyps (J. W. Papez, 1956). According to its data, such ways three: reticular and thalamic, passing from R. f. a myelencephalon to a tsentromedialny kernel of a thalamus, and also to zadnemedialny and parafascicular kernels. On the lateral party of this way there pass sensitive ways (lemniskovy system), from to-rykh collaterals to kernels of a tire depart. From kernels of a tire the tag-mentno-thalamic way which is coming to an end in the central median kernel (centrum medianum) and cells adjoining to it begins. The most lateral on an arrangement is the tektotalamichesky way terminating in a boundary kernel of a thalamus. Thanks to these ways the nonspecific projective thalamic system becomes a link between the ascending activating reticular system of a brain trunk and a cerebral cortex.
Recognition of existence of two afferent systems (specific and nonspecific) having the different forms, ways and spheres of influence resulted in need of studying of features of the termination of fibers of both of these systems for a cerebral cortex. It was shown that nerve fibrils of these systems have the terminations in bark differing in a form and distribution in various bast layers. Specific afferent fibers terminate mainly in the 4th layer of bark, nonspecific fibers — in all layers of bark. Specific fibers terminate mainly on a body of a cell, nonspecific — on its dendrites. The Aksodendritichesky terminations of nonspecific fibers can create change of excitability of cortical neurons, facilitating or complicating synoptic transfer (see. Synapse ). These influences of a diffuzna are also changeable. The Aksosomatichesky terminations of specific fibers provide prompt and local replies. Interaction of both of these systems, according to Chiang (H. T. Chang, 1952), causes final reaction of cortical neurons.
It should be noted that if in 50th and 60th 20 century the look, according to R.'s Krom f prevailed. was considered as it is diffuzno organized - bark, rendering «nonspecific» the ascending and descending influence, in the 70th this opinion began to be reconsidered.
One of the first researchers who pointed that the activating R.'s influences f. always have certain biological «sign», there was P. K. Anokhin (1958). Progress of both physiological, and morphological methods of a research was the reason of radical review of views. First of all, it is necessary to refer improvement of the technology of the gistoflyuorestsent-ny analysis to the last that allowed to reveal the neurons containing monoamines (noradrenaline, serotonin and dopamine) in a brain trunk and to stand on hind legs of a branching of fibers of these neurons. It is noted that noradrena-Lowai and serotoninny systems contain typical reticular neurons. These data forced to reconsider morphological dogma, according to a cut all ways to a cerebral cortex switch in a thalamus. It was established that mezenets - falichesky serotonergic and noradrenalinergichesky neurons ascend directly from R.'s kernels f. in a cerebral cortex. In this regard there was a question of a neurochemical basis of cortical «reaction of awakening».
Significant contribution to R.'s physiology f. brought use microelectrode methods of a research (see). It was shown that one of important mechanisms the activating reticulothalamic and retikulo-cortical influences is suppression of brake interneyron, i.e. «braking of braking».
New data were obtained in the course of studying retikulomotor-ache coordination, i.e. in identification of the local groups of the cells which are specifically connected with control of certain forms of a physical activity, in particular in identification of neurons, regulating the movements of eyes, the neurons controlling mechanisms of a pose and locomotion, etc.
Opening in a number of R.'s structures f. high concentration of so-called opiate receptors, pointed to communication with the organization of painful sensitivity. Kernels of a median seam, also central gray matter around a silviyev of a water supply system, etc. belong to these structures.
Detection of functional specialization of various departments of R. f. led to revaluation of the relation to R. f. as to «nonspecific system». The special international symposium, materials was organized and carried out to the USA to-rogo are published in the monograph «Review of views of a reticular formation: specific function of nonspecific system» (1980).
However it should be noted that the researches which are summing up data on R. f., do not cancel a view of R. f. as on the system working in functional unity with analizatorny systems and exerting tonic impacts on below - and overlying departments of c. N of page.
It is necessary to consider that long before emergence of the early studies devoted to R. f., I. M. Sechenov showed that in the field of a brain trunk the structures controlling activity of a spinal cord are located. I. P. Pavlov also attached to subcrustal educations great value in maintenance of activity of bark of big hemispheres. He spoke about «the blind force» of a subcortex, about a subcortex as «an energy source for bark». Establishment of an important role of reticular mechanisms in activity of c. the N of page is a specification of theoretical representations of these ingenious Russian scientists.
Pathology of a reticular formation
Dysfunction of R. f. the hl develops owing to defeat of its kernels which are localized. obr., in a medulla, the bridge and a mesencephalon, and also afferent and efferent bonds at various levels.
Pathology of actually integrative functions P. f., as A. M. Vane (1974), Ilayesson showed (S. G. Eliasson, 1978), it can be shown in the form of frustration movements (see), disturbances consciousnesses (see), dream (see), vegetative dysfunction.
Motive frustration are caused by disturbance of phasic and tonic control of cross-striped muscles, to-ry is normal carried out by means of interaction of activating and braking R.'s influences f., transferred on alpha and gamma motor-neurons spinal cord (see) in reticulospinal and vestibulospinalny ways.
R.'s pathology f. a brain trunk, and also its afferent and efferent bonds can be followed as increase in a muscle tone and tendon jerks, and their decrease. The muscular hypertension and increase in tendon jerks arise owing to dominance of the activating influences of a reticular formation on and - and at - motor-neurons of a spinal cord, a cut is observed at defeat of a giant-cell reticular kernel at the level of a myelencephalon or the bridge, its afferent bonds with bark of a great brain and a kernel having a tail, and also at pathology of efferent pathways to motor-neurons of a spinal cord.
Diffusion defeat of R. f. at the level of a brain trunk can lead to falloff of a muscle tone and tendon jerks owing to lack of the activating influences on and - and 7 motor-neurons of a spinal cord.
Motive frustration at R.'s pathology f. touch not only cross-striped muscles of a trunk and extremities, but also the muscles innervated craniocereberal (cranial, T.) nerves.
Another a wedge, the syndrome observed at R.'s pathology f., disorder of consciousness up to emergence of a coma is. Coma (see) it is characterized by full loss of consciousness, lack of reaction to external irritants, a slow synchronized rhythm of EEG. Blockade by the ascending activating R. f is the cornerstone of a coma., responsible for processes of wakefulness, activation of attention to various touch incentives. Functional or structural disturbances of the ascending activating system at any its level, including oral departments of a brain trunk, a transparent partition, a hypothalamus, a thalamus and talamokortikalny bonds, can lead to disorders of consciousness. Comas most often develop at pathology of a brain trunk and mesencephalon or at the processes leading to their dislocation. But also other mechanism of a coma is possible, at Krom pathology of a cerebral cortex and disturbance of the descending influences of bark on R. f is its cornerstone., therefore the functional condition of R. f for the second time changes.
For R.'s defeat f. in the field of a tire of the bridge and a mesencephalon the syndrome of a pseudo-coma, or an akinetic mutism is characteristic. The syndrome of an akinetic mutism is characterized by loss of ability to react adequately to external stimuluses at safe consciousness or poorly expressed its disturbance. At the same time at the patient it is broken speech (see), the active movements, he does not remember the events which are taking place in this time term. Pupillary tests, tendon and periosteal jerks do not change. Strong pain and sound stimulations cause response motor reaction. Disturbance of the ascending activating system and its bonds with limbic structures of a brain is the cornerstone of a syndrome (see. Limbic system ), what results in lack of motives to action, difficulty of integration of motor functions, frustration memories (see).
Frequent symptom of defeat of R. f. and its bonds frustration of a dream is. Pathology of a dream (see) is caused or disturbance of functional reciprocal relationship between the ascending activating system and the gipnogenny synchronizing zones responsible for generation of a dream, or dysfunction of the gipnogenny zones located hl. obr. within a limbiko-reticular complex.
Frustration of a dream can be in the form of the increased drowsiness (hypersomnia) and various disturbances of a night dream (insomniya). Hypersomnias can be caused by hypofunction of the ascending activating system or hyperfunction of one of systems of mechanisms of regulation of a dream. The Gipersomnichesky states observed at organic lesion of mezentsefalno-diencephalic area of a brain are result of disturbance of functioning of the activating system P. f. Insomniya, characterized by sleeplessness, difficulties of backfilling, frequent prosypaniye, shortening of duration of a night dream, can be caused by rather increased functional activity of the ascending activating system, and also disturbance of functioning of the certain somnogen-ny areas of a brain responsible for generation of phases of a REM and slow sleep. Such gipnogenny zones are located in caudal department of a brain trunk (the so-called synchronizing J. Moruzzi's system), in kernels of a median seam, a hypothalamus. At their pathology disorganization of separate phases of a dream is observed. Disturbance of the cyclic organization of a dream without rough changes of separate phases is characteristic of pathology of the integrating devices regulating turning on of the synchronizing and desynchronizing mechanisms of a dream. Separate structures of a limbiko-reticular complex (a hypothalamus, a visual hillock, basal kernels of an end brain), and also the activating talamokortikalny system treat them.
At dysfunction of R. f. a brain trunk the syndrome of a neurangiosis can be observed. By data A. M. Vane, etc. (1981), vegetovascular frustration are noted at most of patients with pathology of a brain trunk. Vegetative disturbances are presented by cardiovascular, vasculomotor and respiratory frustration, to-rye can have a sympaticoadrenal or parasympathetic focus (see. Neurocirculatory dystonia ). At the heart of the vegetative disturbances arising at R.'s defeat f. a brain trunk, not only dysfunction of the specific vegetative centers (vasculomotor, respiratory), but also disturbance of the complete integrative function necessary for ensuring reasonable adaptive behavior lies. Therefore the respiratory, cardiovascular and vasculomotor frustration observed at R.'s pathology f. a brain trunk, are followed by changes of a muscle tone, motility and secretion of internals, endocrine disturbances, changes of mood, decrease in memory. There is a certain dependence a wedge, displays of vegetovascular dystonia from area of defeat of R. f. brain trunk. At disturbance of upper parts of a trunk vegetative frustration carry a sympathetic orientation, can be followed by easy neuroendocrinal disturbances. At patients with defeat of caudal departments of a trunk the parasympathetic orientation of a tone of the autonomic nervous system comes to light, vestibular disturbances are often observed. It is caused by existence of bonds of R. f. with kernels of a vagus nerve and vestibular nuclei.
Studying of physiology and R.'s pathophysiology f. allowed to deepen considerably ideas of mechanisms of development of many diseases of a nervous system. Pathology of the ascending activating system P. f. is the cornerstone of disturbances of consciousness and comas (at acute disorders of cerebral circulation, craniocereberal injuries, tumors, encephalitis, metabolic frustration). Blockade of the activating influences on bark of a great brain can be caused or directly center of defeat which is localized in a brain trunk, a mesencephalon or a hypothalamus or the hypostasis leading to a prelum, dislocation and secondary metabolic frustration in this area.
At the comas caused by disorder of metabolism (e.g., at a hypoglycemia) or intoxication medicines (barbiturates, tranquilizers, adrenolytic means), observe direct suppression of neurons of the activating reticular system or blockade of adrenergic receptors of synapses.
The increase in functional activity of system of wakefulness which is shown pathology of a dream in the form of insomniya is characteristic of neurosises. At patients with neurosises (see) also the syndrome of a neurangiosis, emotional frustration characteristic of dysfunction of a limbiko-reticular complex are often observed.
R.'s pathology f. plays a part and in development of a syndrome parkinsonism (see). Among typical morphological changes at this disease often find death of neurons of R. f., providing a condition of wakefulness. The increased drowsiness and an akineziya at parkinsonism depend not only on primary defeat of the activating system P. f., but also its blockade due to functional strengthening of brake influences of a kernel having a tail at the level of retiku-lokorkovy bonds.
Pathology of descending R.'s influences f. plays a role in formation of the central paralyzes and paresis, extrapyramidal rigidity, myoclonias.
Specification of function P. f. and clarification of its role in development patol. disturbances became possible on the basis of wide-ranging pilot and clinical trials. For studying of function P. f. a method of implantation of electrodes with definition of activity of cell populations, the elektroentsefalografichesky analysis, morphological researches are applied using submicroscopy (see), methods histochemistry (see) and biochemistry (see), including studying of a neurochemistry mediators (see). In a wedge, practice the printing methods of a research including at the same time are widely used electroencephalography (see), elektrookulografiya (see), electromyography (see), electrocardiography (see), with the help to-rykh it is possible to determine differentially the level of defeat of a nervous system, a functional condition of the ascending and descending systems P. f. and to reveal features of their response to use of various pharmacological means.
Is available a wide arsenal of the pharmacological drugs influencing function P. f. and its bonds with other formations of a brain. The selection action on the activating reticular system barbiturates possess, to-rye block the ascending impulses to a cerebral cortex. This mechanism is the cornerstone of their narcotic and anticonvulsant effects. Drugs of bromine, drugs of a fenotiazinovy row (aminazine, etc.), nek-ry tranquilizers (chlordiazepoxide, diazepam, oxazepam, nitrazepam) have direct overwhelming effect on the ascending activating system what their calming, anticonvulsant and easy somnolent effect is connected with. The ascending reticular system adrenergic mediators (adrenaline, noradrenaline), their predecessor activate L-DOFA, and also indirect adrenomimetik (caffeine, Nialamidum, an imipramine, amitriptyline, Phenaminum, meridil, Sydnocarbum, etc.). These means use in complex therapy of patients in coma, at the increased drowsiness, depressions, an adynamy. Cholinergic synapses of a reticular formation are blocked by the central cholinomimetics (Scopolaminum, amizyl, meta-lizil) that leads to decrease in parasympathetic influences of R. f. a brain trunk on internals. Reduction of a flow of sympathetic impulses by the periphery can be reached by use of sympatholytics (Reserpinum, Methyldopa), to-rye break formation of catecholamines and activate brake structures of R. f. There are means having selective effect on serotonin-ergichesky structures of R. f. (L-tryptophane, dizerit). These drugs use in clinic for the purpose of normalization of a dream. The effect of braking of reticular macrophages of a caudal brain trunk as liorezat, to midokala, diazepam, is used by such drugs at treatment of patients with increase in a muscle tone.
Correction of syndromologic disturbances of R. f. is a part of complex therapy of diseases of a nervous system, in a cut the leading place shall be allocated to etiological and pathogenetic treatment.
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V. V. Amunts, V. G. Skrebitsky, V. N. Shelikhov; L. O. Badalyan (not BP.).