ISSN 0362-1197, Human Physiology, 2007, Vol. 33, No. 6, pp. 701709. Pleiades Publishing, Inc., 2007.

. Original Russian Text D.P. Matyushkin, 2007, published in Fiziologiya Cheloveka, 2007, Vol. 33, No. 6, pp. 5059.

The Possible Neurophysiological Basis of the Inner Self1

D. P. Matyushkin
Institute of Biomedical Problems, Russian Academy of Sciences, Moscow, 123007 Russia
Received June 20, 2007

AbstractDifferent hypotheses on the nature of the inner self suggested by neurologists, neurophysiologists, and psychophysiologists are discussed, and a new hypothesis is put forward. This hypothesis is based on the structures of neural networks serving as mechanisms for making decisions (the active ego) and networks for introspection, self-control, and self-consciousness (the reexive ego). The hypotheses are compared with known facts, and their validity is estimated. DOI: 10.1134/S0362119707060060

The inner self is an extremely complicated psychological problem, only the basic aspects of which are considered here. Modern psychologists, e.g., Hjelle and Ziegler [1], Kon [2], and Raen [3], subdivide the ego into an active ego (self) and a reexive ego. Some authors further subdivide the reexive ego into a partly subconscious, broad ego image and an ego concept, which is a conscious sample from the ego image [4]. Neurologists, neurophysiologists, and psychophysiologists addressing this complicated subject attempt to apply their methods to the search for and study of the substrate and mechanism of the inner self. The philosopher Dubrovskii [5] logically demonstrated that the cerebral substrate of the inner self must constitute an ego system (hereinafter, the terms inner self and ego system (ES) are used as synonyms). Different hypotheses on the human ego have been put forward. Behaviorists have long since accepted a hypothesis that essentially denies that the ego has a special substrate and mechanism. More precisely, this hypothesis reduces them to known cerebral mechanisms and regards introspection of the self as an illusion [6]. This does not seem to agree with all factual data obtained by psychologists [2] and neurologists [7]. Popper and Eccles [8] and Eccles [9] put forward a dualistic hypothesis that the inner self is a self-conscious soul. The inner self comprises the ego, self, psyche, and will. The soul is World Two. It obtains, in some way or another, external and internal sensations from the physical World One, the connecting brain, or, to be precise, the neocortex of, predominantly, the left hemisphere. The brain, in turn, is controlled by the soul via modication of the probability elds for the positions of synaptic vesicles at nerve endings, which are inputs for cortical neurons. In this way, the soul allegedly can release neurotransmitters from these vesicles and control the brain function. This hypothesis is
1 This

article is published as a discussion paper.

difcult to test because we do not know what the soul is. Peneld and Jasper [10] and Peneld [11] assumed, on the basis of neurosurgical data, that the substrate of the ego was located in the centroencephalic system (the upper portion of the brainstem) because lesions in this region turn off consciousness. The authors believed that the centroencephalic system controls all cerebral functions; however, they did not determine the mechanism of this control. Peneld, a religious person, assumed that the soul could be involved in the function of this system [11]. Thus, Penelds hypothesis is also, in a sense, dualistic. The demonstration of the functions of the reticular formation (RF) and the cortical control of this structure [12] was interpreted against this hypothesis. Penelds hypothesis was accused of assuming that the brain contains a homunculus, in which the controlling coreanother homunculuswas to be found. This means searching for a homunculus in a homunculus and so on, along an absurd path of endless intra-action. Later, a few hypotheses arose that can be dened as informationalmaterialistic. They are based on the fact that information is neither matter nor energy but its existence requires a material carrier and energy expenditure. These hypotheses admit that there is a special material substrate of the ego and its introspection in the brain. Judging from some remarks of Edelman and Mountcastle [13], Edelman and Tononi [14], and Ivanitskii [15], they believe that the ego is a function of certain neuronal complexes distributed over the cortex and subcortex with repeated information inputs in this system, with the role of attention being emphasized. Edelman and Mountcastle [13] relate the self as a part of consciousness only with cortical associative areas, the limbic system, and the RF, rather than the whole brain. However, some hypotheses (from the same group) relate the ego only with the brain hemispheres. Rotenberg and Arshavskii [4] believe that the substrates of the ego image and ego concept are located in the right and left hemispheres, respectively, and the active ego is related to both hemispheres, but predominantly to the

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right one. Crick and Koch [16, 17] and Libet [18] assume that the apparatus of consciousness (and, hence, the ES) is a specic part of the brain. The neurologist Damasio [19], after a detailed analysis of the roles of different divisions of the brain in consciousness and self-consciousness (based on numerous clinical and literature data), concluded that the core of consciousness and the basis of the self (self-perception) are determined by the activity of phylogenetically old medial cerebral structures stretching from the base of the brainstem to the somatosensory and cingular cortices of both hemispheres. The author termed this formation the protoself. It does not include the cerebellum, hippocampus, or any cortical areas (except the somatosensory one) located on the convexial surface. According to Damasio, the protoself has no language or knowledge. They are introduced into the system of the self by structures that the author calls the autobiographic self. The protoself and autobiographic self are connected with each other via the core of the self (and consciousness). Earlier, we suggested an essentially similar hypothesis [2023]. Thus, the psychophysiological hypotheses considered here contradict one another and there is no consensus about this problem. We regard our previous works [2023] as preliminary attempts at solving the problem of the ES and now suggest a considerably more detailed hypothesis on the informationalmaterialistic nature of the ES. The hypothesis is based on previously known data on synapses, neural networks, and neuron interaction obtained in experiments on animals, studies in humans [2123], our own theoretical estimations and calculations in this area, and all available psychophysiological data. We tried to base the hypothesis on a notion of the tasks that the ES, the neurophysiological basis of the inner self, should fulll. It is reasonable to assume that the human ES should fulll several tasks: (1) uniting parts of the body into a whole, i.e., serving as a set of their representations (the basis of a persons integrity); (2) comparing the demands of the body with signals and states of the environment, mainly the social one, to choose a correct (adequate) vector of attention and behavior (the basis of inner struggle in a persons psyche and personal choices and decisions); and (3) performing introspection, i.e., self-control involving knowledge and memory (the basis of a persons self-consciousness). Humanity, with its knowledge, laws, religions, and traditions, forms the content-related part of a persons memory, thereby determining the customary subjective estimations and self-estimations. The soul of a people, nation, or religious confession fullls the function of Popper and Eccless [8] World Two. In the human brain, this world is represented by models, i.e., special neuronal chains or complex systems. Words (Pavlovs [24] second signal system) play a special role here. In addition, the words and acts that a person gen-

erates become part of the persons environment. Thus, environmental signals are complicated, and a psychophysiologist should begin by simplifying the situation, e.g., by considering ES responses to simple external signals. Like most psychophysiologists [1317], we proceed from the assumption that human subjective feelings or sensory experiences (qualia) are determined by the sensitivities and activities of neuronal structures serving as elements of the ES. Their activities are codes of subjective feelings, which are conscious or unconscious if elements encoding knowledge (notions) are, respectively, involved or not involved. For simplicity, let us regard neurons as elements of all ES structures. We assume that the ES consists of two components: (1) the lower self and (2) the higher self. The lower self is based on a network of monosynaptically interrelated elements representing various vectors of behavior. Since these vectors usually compete with one another, most connections between neurons in the network are mutually inhibiting. This network (referred to as the M network) is the basis for uniting the parts (systems) of the body. At the same time, it is also an arena of competition between representatives of different behavioral vectors. This is the apparatus that forms nal decisions, i.e., the choice of the behavioral dominant based on congenital and acquired connections of the network representatives with the respective sensory and central inputs (the basis of the active ego). The higher self is an apparatus consisting of neuronal chains and systems that contains the codes of acquired knowledge (memory). It serves as the basis for self-evaluation by introspection, i.e., the basis of selfconsciousness. In other words, this is both the reexive ego and the basis of the higher forms of a persons cerebral activity (except the nal decision on its vector). The higher and lower selves are closely related in a cyclic manner. The lower self informs the higher self about an arising vector of attention and orients the higher self toward the estimation of this vector (introspection), whereas the higher self gives an estimate of the vector, either positive (supporting) or negative (rejecting); in the latter case, it may offer the lower network another attention vector. The lower self, or, more precisely, the M network, serving as its basis, is assumed to be a network of not only inhibitorily linked, but also closely located neurons. It receives information on various internal motivations (hunger, thirst, sexual drive, etc.), the external situation, and exploratory and other programs (intentions) of both hemispheres of the brain. This information is distributed among the representatives of the corresponding attention and behavior vectors via congenital and acquired connections. Their competition based on mutual inhibition determines the choice of the dominant active element (a neuron or a group of consensual neurons), which is essentially the subjective choice of attention and behavior vectors. The outputs of
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the dominants of this network determine the attention vector, thereby permitting or forbidding instincts, conditioned reexes, awakening, emotions, and intellectual and other processes by affecting the central components of these mechanisms with high-frequency (500 1000 Hz) series of action potentials (APs). A permission on the part of an inhibitory neuron may be effected only via inhibition of some inhibitory interneurons upstream of the target, i.e., via disinhibition of the target. Apparently, the body in the waking state needs restraint of motor and other effector apparatuses (Ukhtomskiis operative rest [25]), which is necessary for quick and precise performance of behavioral programs. We assume that this network contains both permitting (yes) and forbidding (no) neurons for each behavioral vector. They may be called functional pairs. Decisions such as to be or not to be and to do or not to do are necessary for every function. However, no neurons are assumed to serve mainly as individual inhibitors of the respective yes neurons. The network must contain many such pairs linked with one another via permitting neurons (no neurons from different pairs are linked weakly with one another if at all). Yes neurons, strong competitors, have strong mutually inhibiting synapses. The yes neurons that consensually act in the framework of the same behavioral vector may have very weak inhibitory links; hence, given a common input, they may form dominant constellations [25]. Owing to summation of the inhibitory impulses from individual neurons to competitors, such a constellation may be a stronger dominant than a single neuron. It is assumed that yes neurons may support the system of awakening, and no neurons, the system of sleep. Technically, the inhibitory impulse from a yes neuron to a competitor yes neuron within this network corresponds to Pavlovs external inhibition, and the inhibitory impulse from a situationally activated no neuron to a yes neuron within a functional pair, to Pavlovs internal inhibition [24], which is in line with ideas put forward by Konorski [26] and Anokhin [27]. Pavlov [24] presented much evidence for the relationship of internal inhibition with sleep. The mechanism of the choice of the dominant yes neuron in the case of a common exciting input for two competing yes neurons may be very quick if the AP threshold of one of the yes neurons is drastically decreased (because of the effects of neural or humoral inputs). Then it can generate the AP at the very beginning of the development of the exciting postsynaptic potential. Given a small distance to the competitor, this AP will immediately reach the synapse, and the inhibitory neurotransmitter will be released within ~0.3 ms [28, 29]. The unprepared competitor could develop an AP in response to this common input only ~1.0 ms after the beginning of the exciting postsynaptic potential [28]. In this case, the prepared neuron inhibits the competitor beforehand, preventing it from developing its AP. Our calculations show that this preventive inhibition is possible if the neuron prepared to dominate has
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a threshold two or more times lower than the competitor has. If the AP thresholds of the two neurons differ by less than a factor of 2, both competitors are excited in parallel, forming a slow (because of mutual inhibition) AP rhythm. This may retrieve information, e.g., information decreasing the AP threshold of the dominant, from higher apparatuses of knowledge, which is necessary for solving the dilemma. If there are more than two competing instinctive or conscious programs/ideas, then the choice between them involves introspection, the least supported variants being sequentially excluded. In general, mutual inhibition is expected to restrict the number of simultaneously competing elements, which explains some psychophysiological phenomena [30]. The number () of yes neurons in a fully connected network can be calculated from the maximal number of synapses on the perikaryon of a large neuron (N), which is approximately 105 [28]. Note that, if every yes neuron in the network has N partners, then in a network of N + 1 yes neurons, all sites for synapses will be occupied by intranetwork synapses, which is impossible. Ideally, this network should have as many sites open for external inputs (S) as possible. S = M[N (M 1)], where M 1 is the number of sites per neuron occupied by intranetwork synapses in a network of M neurons. Note that the dependence of S on M is bellshaped (S increases with increasing M from zero and then decreases to zero at M = N + 1). The maximum of S can be found by equating its derivative to zero. The derivative of S is N 2M + 1; if it equals zero, we obtain the equation N 2M + 1 = 0. Then, 2M = N + 1; hence, S is maximum if there are (N + 1)/2 neurons in the network. This is approximately 5 104 neurons. Since the decision-making network has inputs from the skin, muscle, auditory, vestibular, visual, and olfactory sensory systems, it is conceivable that this network is extended. It may be a chain of networks connected via the sites that are open for external inputs involving interneurons, rather than a single global network. This chain comprises more than 5 104 elements (or pairs); however, the number of behavioral vectors is smaller than the number of elements in the chain because multiple duplication of elements for increasing reliability and grading the strength of inuence is assumed for each vector. Note that yes neurons of different vectors in these decision-making networks are essentially equivalent and there is no special commander or homunculus among them. This model also leaves few sites for any considerable number of new (temporary) connections, i.e., for functioning of the apparatus of new knowledge; however, these networks contain the hereditary, congenital experience of the species. We assume that the higher self of the ES is neuronal screens that are distributed over the cortex and limbic system and whose memory units store encoded [31] information on the person, the external world, and

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their possible interactions. This information is retrieved by a signal from a dominant of the lower network indicating the attention vector in which the higher self should act as an entity realizing the input sensory signals, estimating the dominant of the lower network (introspection) and the possible results of actualization of this dominant, and nding other activity vectors for the lower self. The part of the higher self that is related to speech may respond in the form of the inner voice code. Thus, the higher self plays the role of the reexive ego. Some of these screens may display the codes of all behavioral vectors, and others, the codes of all external factors. In this case, adequate programs of the persons responses to external factors must result from nding mutually corresponding elements on the screen of behavioral vectors and the screen of external factors. These temporary connections are assumed to be bidirectional [24]; therefore, intentions arising from the lower self or components of the higher self may lead to recollection of behaviors that will help to carry out this intention. If there are many such screen structures, then the input to them from the lower self network must occur through a special distributing apparatus and the numerous proposals arising in the screen structures may be transmitted to the lower network through an apparatus for preliminary selection (to prevent an overload of the nite network). Thus, our hypothesis predicts that the response of a person to an external stimulus may take one of two forms: (1) a subconscious response of the lower self if the stimulus is strong or (2) a conscious response programmed (upon a signal from the lower self) in higher apparatuses of the ES on the basis of the comparison of information on the dominant in the lower network (introspection) and the external situation. This higher program is transmitted to the lower network, where it supports, cancels, or replaces the existing dominant. The higher program, having become, in one way or another, the dominant in the lower network, determines the conscious behavioral response. Note that our hypothesis on the ES structure is similar to Damasios hypothesis [19], differing from it (and other related hypotheses) in that we suggest the main neuronal mechanisms of the lower self. Factual data demonstrate that there are both compact neural networks and neuronal screen structures in the brain. The brain contains an extended compact network similar to the hypothetical one. It stretches from the medulla oblongata to the septum pellucidum and is related to giant-cell nuclei, the interstitial nucleus of Cajal, the central gray matter (CGM), the centromedianparafascicular formations of the thalamus, the reticular nuclei of the thalamus, and the hypothalamus [32]. These structures contain neurons monosynaptically interconnected via quick glycinergic and GABAergic inhibitory synapses [3336]. Multidirec-

tional monosynaptic connections between neurons of the RF have been found using the horseradish peroxidase method [32, 37]. Sheibel and Sheibel [38] have conclusively demonstrated the existence of these connections. The RF is a compact system, and the speed of AP transmission in it may be as high as 138 or 150 m/s [39, 40]. This may permit the preventive inhibition of remote yes neurons by signals from the dominant yes neuron in the case of a synchronous exciting input. There are data on the development of dominant foci and competition between different inputs to the RF [41, 42]. Brainstem neural networks have very numerous inputs [12]. Information is transmitted to the RF from all sensory systems directly through their collaterals and via higher cerebral centers. Destruction of the cortical components of the visual or hearing analyzer in humans extremely simplies their visual (hearing) sensations [43, 44]. Yet these are conscious sensations, which suggests that the brainstem RF is involved in the mechanisms of consciousness [45]. Outputs from the reticular core of the brain are numerous and diverse. Various adrenergic, serotonergic, dopaminergic, cholinergic, and endorphinergic outputs are generally known. However, Limanskii [46] believes that some of these outputs are specially related to the organization of many functions. In the animal RF, relatively long, frequent (5001000 Hz) APs of the command type have been found in response to sensory signals [39]. The reward and punishment sites of the animal brain are the most numerous in the RF [47]. Focal electrical irritation of different regions of the human RF and CGM evoke, according to Bechtereva [31], clear feelings of pain, fear, orgasm, exhilaration, and surprise and, according to Delgado [48], also talkativeness. Note that focal irritation also affects neurons and the sensory pathway collaterals ending on them. These collaterals antidromically activate the sensory pathways and the corresponding sensory cortical areas. The irritated reticular neurons send signals to the memory (knowledge) apparatuses of the ES, mobilizing the attention to the codes of the corresponding images. This causes new sensory images (their codes) to meet ones stored in memory, which leads to recognition, i.e., realization of the sensation. Lesions in RF regions precluding the activation of attention result in neglect of sensory signals even if the specic sensory pathways are preserved [49], which agrees with our hypothesis. In animals, electrical irritation of different sites in the RF and CGM causes (1) quick permission of targeted acts, such as an attack on a present enemy [50], eating of existing food [51], and sexual behavior [52]; (2) immediate forbiddance of various acts, such as defense reexes [12, 53] and various forms of aggression [48]; or (3) sleep and awakening [12]. The RF contains the main systems activating and inactivating higher nervous activity. The human cortical programs of movements are transmitted to the motor cortex indirectly, via the thalamus (the upper region of the brainHUMAN PHYSIOLOGY Vol. 33 No. 6 2007

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stem) [10]. Temporary inactivation of the upper part of the human thalamus causes unconscious automatism [10]. Lesions in the upper midbrain cause coma in humans and animals [12]. Thus, the RF has the characteristics that are assumed for the decision-making network, which is the main component of the lower self and the substrate of the active ego. In terms of our hypothesis, this network may be termed the brainstem processor (BSP). We think that this is also the main mechanism of Damasios protoself [19]. Let us now consider higher cerebral regions, which can be supposed to be the location of the higher self, or higher ES apparatuses (HESAs). They may be termed the cortical processor (CP). Note that the convexial and limbic cortical areas and the hippocampus have a structure that morphologists refer to as a screen structure. The neuronal connections in each region of these structures are very complex [29], and their analysis is beyond the scope of this study. Apparently, the brain cortex (specically, its associative areas) contains the neuronal screens presumed in the framework of our hypothesis, where sensory inputs and various behavioral programs are represented [44]. It has been demonstrated that irritation of cortical associative areas, e.g., irritation of the temporal areas in humans during surgery, may evoke complex sensory experiences (bright images of people, landscapes, and events; sounds of musical instruments; etc.), which are superimposed onto the image of the operating room that the patient actually sees. It may be assumed that the mass of elements involved in the construction of such a picture encodes this subjective state, which is conscious if the picture contains elements of verbalconceptual estimation and the attention vector from the BSP is targeted at these elements. In this connection, the priority phenomenon described in [54] is noteworthy. The authors found that a conscious sensation of rhythmic irritation of the skin cortical area had a latent period of ~500 ms. However, a brief irritation of skin at about the 200th millisecond (against the background of the latent period of the cortical irritation) was consciously sensed so soon that this sensation appeared prior to the cortical sensation. In terms of our hypothesis on the ES, this is explained by the fact that the skin irritation activates the corresponding sensory cortical area simultaneously with the activation of the BSP and the sending of a signal for attention from the BSP to the cortex. Regarding direct irritation of the cortical projection of the skin, it can obtain this signal for attention from the BSP only by acting via corticoreticular pathways (i.e., less efcient, diffuse bypasses). In light of this explanation, we think that the attempts of advocates of the dualistic hypothesis [9] at relating the priority phenomenon to the uncertainty principle taken from quantum mechanics are incorrect and altogether unnecessary. We believe that sensations evoked by irritation of the sensory or associative cortex become conscious only if the BSP attention vector is directed toward these zones.
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Stevens [55] demonstrated both morphologically and by calculations that the cerebral cortex is not a network of intracortically multidirectionally linked neurons. Remote cortical elements (mini- and macrocolumns) are interconnected via white matter (i.e., associative and callosal nerve bers) and subcortical centers. Most of these connections involve interneurons. In most cases, some elements from the same or different hemispheres are located at long distances from one another. At moderate rates of AP transmission, the inhibiting impulses of the dominant may be delayed if columns with a common exciting input compete with one another. Preventive inhibition is excluded in this case. An element of such a construction may become dominant if the frequency of stimulation of its individual input is noticeably higher than those for other elements. Note that this dominant state must take a relatively long time to develop. Thus, the cerebral cortex is not the substrate for quickly deciding to be or not to be and to do or not to do. This, however, by no means makes cortical dominants less important as strategic programs to be transmitted to the nal, brainstem network. Analysis and synthesis of signals, associations, and memory are known to be the main functions of the cortex [24]. They serve as the basis for verbal and nonverbal thinking, intuition, etc. It has been demonstrated that complex forms of higher nervous (mental) activity occur in the cerebral cortex. However, the simplest motor and secretory conditioned reexes in response to visual and auditory irritants can be developed and maintained in dogs after a complete separation of the posterior (visual and auditory) and anterior (cutaneous and motor) areas by a deep cut across the cortex [56]; i.e., they are mediated by the subcortex (and, apparently, involve activation of the BSP). There are data on the organization of the main inputs to screens of the convexial cortex of the hemispheres from brainstem apparatuses. These are connections via limbic formations [57]. Of special importance here is probably the limbic cortex, particularly the gyrus cingularis and gyrus orbitalis, which, according to some authors [7, 19, 58], contain compact projections of the physical and spiritual/social human self. Apparently, these gyri integrate information transmitted from the BSP and various limbic regions to the convexial cortex to determine the vector of its activity. There are special systems for output from the screen cortical structures to the brainstem. The brain hemispheres have network structures; they are contained in the striata (caudate nuclei), which have networks of neurons with GABAergic inhibitory monosynaptic connections [59]. They may be the substrates for quick selection of one of the cortical programs in each hemisphere [6063] for its presentation to the lower decision-making network as an apparatus in which hemispheric (cortical cognitive and emotional) programs meet vital programs of the deep, lower self. The striatalpallidal systems of both hemispheres, as essential participants in the choice of adequate cortical motor

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programs [61], are necessary for normal voluntary movements. Let us consider some more factual data demonstrating the cooperation of the lower and higher selves. One example is introspection performed in the BSP HESAsBSP cycle. This cycle is unlikely to be a single act. Probably, this is a more or less continual self-control mechanism. It has been found that the introspection of the active ego may consist of two components, (1) cognitive (conscious) and (2) emotional (only partly conscious) [64]. These two components of introspection are relatively synchronous, but their substrates are mostly different. Cognitive introspection is mainly related to the speaking left hemisphere (the inner voice may serve as its estimation), and emotional introspection, with the right hemisphere. This applies to right-handed subjects; in left-handed ones, these relationships are different and more complex [65]. Kostandov [66] found that the response of the right hemisphere may sometimes be more rapid than the response of the left. A subconscious response to emotionally negative stimuli may activate mechanisms inhibiting the input of this stimulus to the conscious left hemisphere, which may be a defense according to Freud and his followers [1]. Possibly, this involves not only transcallosal inhibition, but also inhibition resulting from the activation of the defense focus in the BSP and supported by emotional memory mechanisms. Simonov [67, 68] demonstrated that the brainstem (hypothalamus), limbic structures, and cerebral cortex complexly interact in the mechanism of emotions. The left and right hemispheres are known to differ in some of their functions [69, 70]. This is evidenced by clinical data on patients with surgically separated cerebral hemispheres (commissurotomy). However, commissurotomized patients seem to be practically integrated subjects with a dominating left, speaking hemisphere. In these patients, hemispheric functions are integrated via the brainstem. For example, Sergent [71] demonstrated that both ipsilateral and contralateral hemispheres of commissurotomized patients could control the movement of a nger of either hand in response to a local visual (light) signal. Both hemispheres can use the BSP elements permitting this motor act, which agrees with our hypothesis. In the same study [71], both hemispheres of the patients were stimulated with green light signals of the same or different intensity (these different signals were easily distinguished by either hemisphere). In experiments where two such signals were presented (one signal to each hemisphere), the patient had to determine whether the signals presented to different hemispheres were the same or different. The patients could not answer this question correctly. In this case, the hemispheres made decisions at random. The competition of two random decisions in the BSP yielded a random result (the frequency of errors was 50%). The BSP could inform the hemispheres (initially and during introspection)

only about the vector of attention to the signals, and not about their comparative strengths. This agrees with characteristics of the BSP described above. Ivanitskii [15] demonstrated that a conscious visual sensation was ensured by a chain of excitation foci (evidenced by evoked potentials) in the analyzer, interpretative cortex, hippocampus, and hypothalamus. According to Ivanitskii, information is once more transmitted from the hypothalamus (i.e., the BSP) to the visual cortex. Before this, however, the limbic cortex (HESAs) is activated, which has been shown under similar conditions [16, 58]. It is conceivable that the signal from the hypothalamus (BSP) is a signal for attention to the irritant transmitted through the distributing component of the HESAs to the memory apparatuses of the visual system. As a result, reference patterns are retrieved from memory in order to compare them with the new sensory information, which is necessary for realizing the sensation. This agrees with both Ivanitskiis and our models (see also [72]). A study on the conscious attention to an intense sound (90 dB) in the human auditory system [73] demonstrated that the defense reex of the posterior auricular muscle in response to this sound was inhibited after a signal (<90 dB) attracting the subjects attention. Synchronously with the decrease in the reex, changes in evoked potentials from the midbrain to higher cerebral structures were observed. The authors assumed that conscious attention began to form at the level of the midbrain, with higher cerebral levels being rapidly activated after that. Earlier, we studied the electromyographic (EMG) responses of the human external auricular muscles as characteristics of acoustic attention [74]. Subjects had to discriminate between the pitches of two short acoustic signals (tones) presented in a pair. We recorded the EMG responses (an increase and decrease in the EMG signal intensity in the superior and posterior auricular muscles, respectively). The intensity of attention (estimated from the EMG recordings) was correlated with the difculty of the discrimination program and the percentage of errors (the subjects learned about their errors because they heard the correct estimate from a tape recorder after every response). Our ndings demonstrate that conscious acoustic attention involves higher nerve centers, which inuence the brainstem apparatus via, so to speak, the descending branch of the introspection loop. Data obtained with the use of positron emission tomography [75] conrm that some cortical centers are involved in conscious acoustic attention. All these data agree with our hypothesis. A complex electrophysiological and psychometric study in humans [76] showed that a voluntary movement of the human hand begins 450550 ms after EEG changes begin. Initially, however, an unconscious negative cortical readiness potential is observed in the EEG recording. Later (at the 350th millisecond), the intention to move the hand is realized, and the actual motor
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response occurs at the 450th to 550th millisecond. The subject can voluntarily cancel the movement during the 100- to 200-ms interval between the realization of the intention to move the hand and the movement itself. Apparently, this reects competition in the BSP, indicating that the permitting effect of HESAs can be suddenly suppressed by a new competing input. According to Evarts [77], the associative cortex affects the motor cortex indirectly, via the striatumpallidumthalamus (BSP). When Ukhtomskii [25] found an inhibitory effect of the subcortical dominant of the rhythmic swallow reex on excitation of the motor cortex caused by its electrical irritation (the response of an animals hind limb weakened as the swallow movement strengthened), he dealt with the same mechanism. Analysis of the EEG of the cat brainstem and cerebral cortex [78] demonstrated that a long-term waterless diet resulted in the development of a dominant focus (activation), rst in the lateral nuclei of the hypothalamus, then in the sensorimotor cortex, etc. In addition, treatment of the lateral hypothalamus with a NaCl solution induced a search for water [79] (interaction between the brainstem and cortex). Pacemakers of the , , and rhythms of the cortical EEG are located in the brain core. The rhythm pacemaker is assumed to be located in the septum [57]. The - and rhythm pacemakers have been located in the reticular nuclei of the thalamus [34]. Their function may be related to time quantization and elective ltration of information ows from receptive systems and from the thalamic RF to the cortex; however, this important function requires special consideration. CONCLUSIONS Thus, all the factual data considered above conrm that the presumed structure of the ES (the BSP + HESAs) actually exists in the brain and fullls all the aforementioned functions of the ES without the implication of otherworldly forces. In other words, all the materials analyzed here directly or indirectly support the informationalmaterialistic hypotheses on the ES. Regarding some particular issues, these data support ideas put forward by Crick and Koch, Ivanitskii, and Edelman and Tononi. In general, however, they agree best with the hypothesis suggested here and a similar hypothesis independently enunciated by the American researcher Damasio. I would like to emphasize that the hypothesis suggested in this paper not only agrees with the available factual data, but also combines the ideas of Pavlov, Peneld, and Ukhtomskii with those of modern researchers and explains some facts that could not be rationally interpreted before. REFERENCES
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