BASAL KERNELS [poses day of armor. basalis relating to the basis; synonym: central nodes, subcrustal kernels (nuclei subcorticales)] — the accumulations of gray matter in the thickness of big cerebral hemispheres participating in correction of the program of the difficult motive act and formation of emotional and affective reactions.
The first data on B.'s morphology I. meet in Burdakh's works (To. F. Burdach), 1819; I. P. Lebedeva, 1873; Anton, 1895; Kappersa (Page A. Kappers), 1908, etc. Big contribution to B.'s studying I. brought anatomic and kliniko-morphological researches S. Vogt and O. Vogt (S. of Vogt, O. of Vogt), 1920; M. O. Gurevich, 1930; Fua and Nikolesko (Foix, Nicolesco), 1925; E. Κ. Seppa, 1949; T. A. Leontovich, 1952, 1954; Η. P. Bekhtereva, 1963; E. I. Kandelia, 1961; L. A. Kukuyeva, 1968, etc.
B. I. along with the cerebral cortex (cortex cerebri) located on a surface of hemispheres make cellular substance of an end brain (telencephalon). Unlike the bark having the structure of the screen centers (which is characterized by certain cytovery tectonic signs: accurate allocation of layers, vertical orientation of the majority of neurons, their differentiation in a form and size depending on their situation in different layers), B. I. have a structure of the nuclear centers where the similar structural organization is absent. Quite often these kernels are called a subcortex. Treat them: kernel (nucleus caudatus) having a tail, lenticular kernel (nucleus lentiformis, s. nucleus lenticularis), fencing (claustrum) and amygdaloid nucleus (corpus amygdaloideum). To B. I. carry also basal complex of kernels located between the front made a hole substance (substantia perforata anterior) and the forefront of a pale sphere (globus pallidus), belonging septal area (see).
Researches of development of B. I. in fit and ontogenesis showed that the kernel having a tail and a shell of a lenticular kernel (putamen) develop from the ganglionic hillock located on the lower wall of a side ventricle. They represent uniform cellular weight, edges at the highest vertebrata is divided by fibers of a front leg of the internal capsule (crus anterior capsulae internae). In view of a community of an origin and the connection of a head of a kernel having a tail remaining during all life and front department of a shell the strips of gray matter alternating with white yarns of the internal capsule, a kernel having a tail and. the shell is combined under the name «striate body» (corpus striatum), or «striatum» (striatum). Since the striate body is phylogenetic later education, than medially located part of a lenticular kernel — the pale sphere consisting of outside and internal joints it is called by «neostriatum» (neostriatum), and a pale sphere — «paleostriatum» (paleostriatum). The last in a crust, time is allocated in special morphological unit under the name «pallidum» (pallidum).
L. A. Kukuyev (1968) researches show that outside and internal joints of a pale sphere have various origin. The outside joint, as well as a shell, develops from a ganglionic hillock of an end brain; the internal joint — from a diencephalon is also homologous to an entopedunkulyarny kernel of subprimacies (it is located at them in a brain over a visual tract, i.e. its topography is similar to topography of an internal joint of a pale sphere at early stages of development of a germ of the person). In process of both phylogenetic, and ontogenetic development there is a movement of an internal joint towards outside therefore they approach.
B. I. are variously presented in a brain of different classes of vertebrate animals. So, at fishes and B.'s amphibians I. are presented only by a pale sphere, the kernel having a tail and a shell appear for the first time at reptiles, especially well they are developed at birds. At mammals (predatory and rodents) the pale sphere is presented by uniform education, at the person it consists of two joints divided by a layer of white matter. The sizes of a striate body decrease in process of development of a brain in phylogenesis. At the lowest insectivorous it makes 8% of the size of all end brain of mammals, at tupaya and semi-monkeys — 7%, and at monkeys — 6%.
In ontogenesis the striate body can be differentiated at the beginning of the 2nd month of embryonic development. On the 3rd month of development the head of a kernel having a tail presses in a cavity of a side ventricle. Lateralny a kernel having a tail the shell forms, edges it is unsharply delimited from other parts of a hemisphere in the beginning. An amygdaloid nucleus I hold special position among B.; at early stages of embryonic development it separates from a striate body, the cytologic differentiation occurs in it later, than in a pale sphere, however slightly earlier, than in a striate body. Proceeding from onto-and phylogenetic development, it is also impossible to consider it and as the changed, reinforced part of bark of a temporal share or as result of its immersion inside and an otshnurovyvaniye. At a research of an amygdaloid nucleus in comparative and anatomic aspect noticeable reduction of its sizes at mammals — from the lowest insectivorous where it together with a paleocortex makes 31% of all size of an end brain, to the person was revealed, in a brain to-rogo an amygdaloid nucleus makes only 4% of all mass of an end brain. Researches of development of a fencing in onto-and phylogenesis (I. N. Filimonov) showed that it cannot be considered derivative a cortical plate or to connect by origin with a striate body. It represents transient formation between these ground cellular masses of an end brain.
Kernel having a tail has the pear-shaped form; its forefront is thickened and carries the name of a head of a kernel (caput nuclei caudati) having a tail. She is located in front department of a hemisphere and acts in a front horn of a side ventricle (cornu anterius ventriculi lateralis), forming its wall from below and lateralno. Kzadi from a head a kernel having a tail is narrowed and its this department is called a body of a kernel (corpus nuclei caudati) having a tail. The body of a kernel having a tail limits the central part of a side ventricle (pars centralis ventriculi lateralis) from the lateral party and describes a semicircle over a visual hillock (thalamus) and a lenticular kernel. The thinned back department of a kernel having a tail forming a part of a roof of the lower horn of a side ventricle (cornu inferius ventriculi lateralis) forms a tail of a kernel (cauda nuclei caudati) having a tail. The lateral surface of a kernel having a tail prilezhit to the internal capsule (capsula interna), its medial edge adjoins a final strip (stria terminalis).
Lenticular kernel has the form of a wedge, the basis to-rogo it is directed lateralno, and a top medially and down, adjoining podbugorny area. It lies lateralno and slightly lower (ventralny) from a kernel having a tail and a visual hillock, from to-rykh is separated by the internal capsule. In front and ventrally the lenticular kernel connects to a head of a kernel having a tail thin strips of gray matter. Its lateral surface a little convex is also located vertically, bordering on the outside capsule (capsula externa), edges represents the thin white brain plate limited lateralno to gray matter — a fencing (claustrum). The ventral surface of a lenticular kernel lies horizontally and in a middle part connects to bark in the field of the front made a hole substance. Two thin brain plates, medial and lateral (laminae medullares medialis et lateralis), divide it into three parts: the outside part more darkly painted is called a shell, two others represent more poorly painted outside and internal joints of a pale sphere. The fencing is a narrow plate of gray matter, edges is located lateralny a lenticular kernel and separated from it by the outside capsule. From bark of an island (insula) the fencing is separated by the layer of white matter forming the external capsule (capsula extrema).
Amygdaloid nucleus — it is a complex of the kernels located in the field of a hook of a parahippocampal crinkle (uncus gyri parahippocampalis), well differentiated and differing from each other cytologic and tsitoarkhitektonichesk (see area Amigdaloidnaya).
the kernel Having a tail and a shell are similar on a histologic structure. Gray matter of these kernels consists of two types of cellular elements: small and large cells. Small cells, the up to 15 — 20 microns, having short dendrites and thin axons, have gentle granularity and a big kernel with a kernel. Large cells, the sizes to 50 microns, generally triangular and polygonal, their kernel is often located excentricly, in protoplasm there are chromatinic kernels and in the neighbourhood with a kernel a large number of a yellow lipoid pigment. These cells are normal surrounded with satellites. The relation of large cells to small in a kernel having a tail and a shell averages 1: 20. Both small, and large cells have the long axons traced to other deep structures of a brain.
A certain relationship between cellular elements and fibers allowed Vogt (O. of Vogt) to point out similarity of a structure of a striate body to bark. In a kernel having a tail under an ependyma the zone poor in fibers is located; an outside part of this zone is poor in ganglionic cells, internal is richer with them. There is a layer of tangential fibers containing a small amount of ganglionic cells more deeply. On the basis of it Vogt developed the scheme of the strukturnofunktsionalny organization of a striate body (tsvetn. fig. 1): striopetalny fibers terminate on small cells, closely connected among themselves and with big cells, from to-rykh striofugalny fibers already begin. In small cells of fibrilla are not differentiated, in big — are distributed by bunches. Myelin fibers in a striate body are not enough, their most part arises in the most striate body and serves for connection with a pallidum; between bunches of myelin fibers the dense network of amyelinic is located. The rich network of a neuroglia surrounds nervous cells and nerve fibrils. In a pallidum there are cells only very large, various form — pyramidal, spindle-shaped, mul-tipolyarny with long dendrites (tsvetn. fig. 2 and 3). In protoplasm there is a lot of chromatophilous glybok. The surface of cells is covered with loop-shaped trailer little bodies — the terminations of the amyelinic fibers surrounding cells and myelin fibers. It is much more myelin fibers, than gray matter; pale color of a kernel is explained by it.
B.'s blood supply I. the hl is carried out. obr. from an average brain artery (a. cerebri media), the branches going to a striate body (rr. striati). Branches of a front brain artery (a. cerebri anterior) also take part in B.'s blood supply I. All B. I., especially striate body, are very rich with capillaries; distribution of capillaries in a striate body reminds distribution in bark; at defeats of vessels of a brain in a striate body the centers of a softening especially often appear.
Afferent fibers the striate body receives bonds of basal kernels from a visual hillock, from the kernels of a hypothalamus surrounding the III ventricle from a tire of a mesencephalon (tegmentum mesencepnali) and from black substance (substantia nigra). These fibers reach a limit about small cells of a striate body, from to-rykh axons preferential go to big cells, and already from these last fibers go to a pallidum as a part of a striopallidal bunch (fasciculus striopallidalis). Fibers of a kernel having a tail cross the internal capsule, enter a shell, and then, penetrating a brain plate, get into a pallidum. From a shell, from its large cells, fibers through a brain plate are also included into a pallidum. The last is the main place where fibers from a kernel having a tail and a shell go. Nek-ry authors do not deny a possibility of existence of the long fibers going directly from a shell to a trunk without being interrupted in a pallidum. The afferent fibers going to a pallidum consist of the fibers going: 1) directly from bark; 2) from bark through a visual hillock; 3) from a striate body; 4) from the central gray matter (substantia grisea centralis) of a diencephalon; 5) from a roof (tectum) and a tire (tegmentum) of a mesencephalon; 6) from black substance.
Efferent fibers B. I. depart from a pale sphere. The main bunch leaving it — a lentikulyarny loop (ansa lenticularis); its fibers begin in a kernel having a tail, take part in formation of brain plates (laminae medullares). The loop is interrupted in a pale sphere. The fibers which are coming out a pale sphere cross the internal capsule; on border with legs of a brain in a hypothalamus they are scattered fanlikely and come to an end in front and lateral kernels of a visual hillock, in a hypothalamus (hypothalamus), black substance, a podbugorny kernel (nucleus subthalamicus) and a red kernel (nucleus ruber). A part of fibers goes as a part of front decussation of a tire (decussatio tegmentalis anterior) on the opposite side where comes to an end in the educations of the same name. Other bunch leaving a pale sphere — a lentikulyarny bunch (fasciculus lenticularis). This bunch is located under zona incerta, includes the fibers going to a podbugorny kernel (around to-rogo form a bag), to a visual hillock, a red kernel, a kernel of the lower olive (nucleus olivaris), mesh substance (formatio reticularis), a chetverokholmiya [corpora quadrigemina (BNA); lamina tecti (PNA)], to periventrikulyarny kernels. A part of fibers through front decussation of a tire comes over to the opposite side and reaches a limit in the same educations. The ways connecting a striate body to area of a funnel (infundibulum) and which are located over zona incerta are described. From a red kernel, a chetverokholmiya peripheral Extrapyramidal fibers begin (tractus rubrospinalis, tractus tectospinalis). There are no exact data on communication of a fencing and amygdaloid nucleus yet. In literature there are instructions on communication for animals of a fencing with the fibers from an outside bag coming from piriformny area on its communication with an amygdaloid nucleus of opposite area and ventral area of a diencephalon. It is established also that the fencing is connected with bark of an island. Bonds of an amygdaloid nucleus — see. area Amigdaloidnaya .
Physiology of basal kernels
At the lowest steps of evolution (at fishes, reptiles, birds) B. I. are the highest coordination centers of difficult behavior. At the person and highest animal (primacies) difficult integrative activity is carried out by bark of big hemispheres, however B.'s role I. does not decrease, and only changes (E. K. Sepp, 1959).
At early stages of post-natal ontogenesis the main motive function of the newborn — the involuntary chaotic movements — is carried out generally at the expense of a pallidum. With development of a striatum in later terms of post-natal ontogenesis emotional manifestations (smile) are noted and statokinetic and tonic functions become complicated (the child holds a head, carries out the consensual movements). By consideration of a physiological role of B. I. it is necessary to proceed from features of bonds of these kernels with other departments of a brain (E. P. Kononova, 1959; I. N. Filimonov, 1959; O. Zager, 1962). B. I. are characterized by richness of afferent and efferent bonds with motor zones of a cerebral cortex (fig.,/), with kernels of a visual hillock (fig., II), between B. I. (fig., III), with kernels of a mesencephalon (fig., IV), and also with a hypothalamus, formations of limbic system and a cerebellum. Importance for understanding of physiology of B. I. has the accounting of the feed-backs going from them to a cerebral cortex. Such wide range of bonds causes complexity of functional value B. I. (combined in striopallidal system) in various neurophysiological and psychophysiological processes (V. A. Cherkes, 1963; E. Yu. Rivina, 1968; Η. P. Bekhtereva, 1971). B.'s participation I is established. in the following neurophysiological functions: a) difficult motive acts; b) vegetative functions; c) instinctive reflexes; d) touch processes; e) uslovnoreflektorny mechanisms; e) psychophysiological processes (emotions). B.'s role I. in implementation of difficult motive acts is that they cause myostatic reactions, optimum redistribution of a muscle tone (thanks to the modulating influences on underlying structures of c. the N of page defining regulation of movements).
So, the studying of function of a pallidum which is carried out in the conditions of chronic experience allowed to establish its important role in course of compound instinctive reflexes various biol, orientations — sexual, food, defensive, etc.
The method of direct electrostimulation of a pallidum showed ease of reproduction of motor and bioelectric manifestations of epileptiform reactions of tonic type. Among the most important functions of a kernel having a tail and a shell it should be noted their braking influence on a pallidum [Tilni and Rili (F. Tilney, H. And. Riley), 1921; J. W. Papez, 1942; A. M. Grinstein, 1946, etc.]. Effects of switching off of a neostriatum (striate body) affect functional activity of the pallidarny and srednemozgovy centers (black substance, a reticular formation of a trunk). There is their disinhibition that is followed by change of a muscle tone and emergence hyperkinesias (see). Numerous researches of influence of a kernel having a tail on uslovnoreflektorny activity and on purposive movements testify both about braking, and to the facilitating nature of these influences that led to the conclusion about existence of two ascending activating systems: neostriatal and reticular; neostriatal carries out influences on bark of big hemispheres both directly, and indirectly, through kernels of a visual hillock. In B. I. the phenomena of convergence of sound, visual, proprioceptive impulses are found. Apparently, B. I. are transfer instance of impulses from a reticular formation in bark of big hemispheres. The phenomena of a disorientation, a chaotic physical activity against the background of stimulation of a kernel having a tail and a shell are explained by it. The striate body is important in regulation of vegetative components of composite behavioural reactions. The irritation of a neostriatum is followed by emotionally expressive reactions (mimic reactions, the increased physical activity). At the treatment of patients in neurosurgical clinics which is carried out by means of the electrodes implanted on a long term the oppressing influence of stimulation of a kernel having a tail on performance of intellectual, speech activity, on memory state is shown (Η. P. Bekhtereva, 1971, etc.). The great value is attached by B. I. in the mechanism of development of hyperkinesias. At destruction of a pallidum or its pathology display of a hypertension of muscles, rigidity, a hyperkinesia is observed. However it is established that development of hyperkinesias is result of loss of function not of separate B. I., and it is accompanied by disturbance of functions of ventro-medial kernels of a visual hillock and the centers of a mesencephalon regulating a tone (V. A. Cherkes, 1963; Η. P. Bekhtereva, 1965, 1971).
Data of neurophysiological and kliniko-neurologic researches of functions B. I. allow to conclude that their physiological value needs to be considered in connection with other systems of a brain. Hartmann and Monakov (N. of Hartmann, To. Monakow, 1960) showed that during the difficult motive act of B. I. combine a continuous stream of impulses, to-rye extend on certain neural circles: a) a visual hillock — a striate body — a visual hillock; b) a visual hillock — bark of hemispheres — a striate body — a pale sphere — a visual hillock.
Functional relationship between B. I. still are up to the end not found out. Electrophysiologic researches showed that control of a striate body over a pale sphere is not only brake. In acute experiences on cats it was revealed as well the facilitating influence of a kernel having a tail on neural activity of a pale sphere, what increase of action potentials of separate elements of a pale sphere under the influence of irritation of a head of a kernel having a tail testifies to.
Studying of evoked potentials in B. I. showed a possibility of convergence of vozbuzhdeniye from various touch channels on the same neuron [Segundo and Makna (I. P. Segundo, X. Machne), 1956; Alb-Fessar et al. (D. Albe-Fessard) et al., 1960], and, in their opinion, in one of neural groups B. I. somatotopichesky localization is not presented.
Big specific weight of afferent morfo-functional linkages allows to consider that a physiological role of B. I. it is not exhausted by the motive sphere. Considering great value of feed-backs and close interaction of B. I. with other systems of a brain, it is possible to come to conclusion that B.'s role I. consists in checking of various afferent influences for performance of a final motive task. Proceeding from P. K. Anokhin's concept about functional system (1968), it is possible to consider that B. I. participate in formation of afferent synthesis, in correction of the program of the difficult motive act and in assessment of results of action. Besides, functional condition of B. I. affects also other functions of a brain, especially during the formation of emotional and affective reactions.
By methods of a research B. I. are: a) irritation (electro-and himiostimulyation); b) destruction — switching off (an extirpation, an electrolysis); c) electrophysiologic method (registration of EEG and evoked potentials); d) the analysis of dynamics of uslovnoreflektorny activity against the background of stimulation or B.'s switching off I.; e) the analysis of kliniko-neurologic syndromes, and also the psychophysiological observations in clinic made in the course of treatment by means of the electrodes implanted on a long term (Η. P. Bekhtereva, 1971). See also Extrapyramidal system .
Bibliogr. Anokhin P. K. Biology and neurophysiology of a conditioned reflex, M., 1968, bibliogr.; I. S berets. Nervous mechanisms of behavior of the highest vertebrate animals, M., 1961, bibliogr.; Bekhtereva Η. P. Neurophysiological aspects of mental activity of the person, L., 1971, bibliogr.; Belyaev F. P. Subcrustal mechanisms of compound motive reflexes, D., 1965, bibliogr.; Granite P. An electrophysiologic research of reception, the lane with English, M., 1957, bibliogr.; To about and A. B N. Electrophysiologic research of the central mechanisms of some compound reflexes, M., 1949, bibliogr.; Rozhansky N. A. Sketches on physiology of a nervous system, JI., 1957, bibliogr.; Sepp E. K. History of development of a nervous system of vertebrata. M, 1959, bibliogr.; Suvorov Η. T. Central mechanisms of vascular disorders, JI., 1967, bibliogr.; Filimonov I. H. Phylogeny and ontogenesis of a nervous system, Mnogotomn. the management on nevrol., under the editorship of N. I. Grashchenkov, t. 1, book 1, page 9, M., 1959; Circassian V. A. Ocherki on physiology basal gangliyev a brain, Kiev, 1963, bibliogr.; And 1 b e-Fessard D., Oswaldo-Cruz E. Rocha-M iranda C. Activity 6voqu6es dans le noyau caude du chat en rSponse h des types divers d'aff6rences, Electroenceph. clin. Neurophysiol., v. 12, p. 405, 1960; B u with at R. S. of The basal ganglia, the thalamus and hypothalamus, in book: Physiol, basis med. pract., ed. by of Page H. Best, p. 144, Baltimore, 1966, bibliogr.; Clara M. Das Nervensystem des Menschen, Lpz., 1959, Bibliogr.; The diseases of the basal ganglia, ed. by T. J. Putnam a. o., Baltimore, 1942, bibliogr.
H. H. Bogolepov, E. P. Kononova; F. P. Vedyaev (physical.).