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Nervous System
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| Posted 6 months ago The reticular formation is an intricate and amazingly complex network of neurons forming a core of the brain stem and extending from the medulla up to the diencephalon. In the thalamus, fibers from the brain-stem reticular formation synapse in cell groups of nonspecific nuclei. These nonspecific nuclei—so called to distinguish them from the specific sensory nuclei of the thalamus—relay information to virtually all areas of the cerebral cortex. The thalamic nuclei and fibers which project reticular activity to the cortex are collectively known as the diffuse thalamocortical projection system. Brain-stem reticular activity is also projected to the cortex via the hypothalamus and certain limbic-system structures. The reticular formation has both descending and ascending fibers mediating activities which may be placed in four functional categories:(1) control of postural-reflex tonus by inhibition or facilitation of motor neurons in the spinal cord;(2) participation in the control of certain autonomic reflexes and respiration; (3) mediation of efferent activity in the central control of sensory input; and (4) regulation of the general state of excitation in the cerebral cortex. This last function is primarily executed by the ascending components of the reticular formation. Arousal, wakefulness, and sleep are to a large extent controlled by the reticular formation and may be measured by the electroencephalogram (EEG). The EEG is a recording of electrical activity in the cerebral cortex by the application of electrodes to the scalp, skull, or cortex itself. The potentials thus recorded are amplified about a million times for visual inspection on a cathode-ray oscilloscope or inkwriter, and each lead or channel represents the activity of a large number of cortical neurons. When a subject is in a highly aroused or excited state the EEG shows desynchronized low-amplitude (voltage), high-frequency activity (beta waves). A subject in a state of relaxed wakefulness whose eyes are closed displays a more synchronous rhythm that is of higher amplitude and lower frequency (alpha waves). As the individual falls into sleep, the frequency decreases further and the amplitude becomes even greater. This rhythm is most evident in deep sleep, in which the EEG is in a state of hypersynchrony (delta waves). A stage of sleep not recognized until the 1950s, called paradoxical sleep, is reflected in low-voltage, high-frequency activity that is virtually indistinguishable from the excited waking state. There is considerable controversy as to whether paradoxical sleep is deeper or lighter than traditional deep sleep of the delta variety, but there is little question that much dreaming occurs during this stage. [SeeDreamsandSleep.]
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| Posted 6 months ago The brain-stem reticular formation receives collateral fibers from the laterally placed ascending specific sensory pathways. Increased activity in these specific sensory tracts is crucial in producing a change in the pattern of firing of the brain-stem reticular-formation neurons, which in turn produces desynchronization (activation) of the cortical EEG rhythm. The appearance of this desynchronization of EEG activity is coincident with behavioral arousal and increased awareness. Thus, the sensory collaterals leading into the reticular formation form the anatomical pathway by which a sensory stimulus arouses a sleeping individual to wakefulness. Arousal from sleep can also be produced by direct electrical stimulation of the reticular formation, as well as by sensory stimulation. There is no question that the arousal function is normally mediated by the reticular formation rather than by the specific sensory pathways. If, in experimental animals, the reticular formation is destroyed at a level rostral to the point at which the sensory collaterals project into it, the subject enters an enduring state of coma, for although specific sensory information will reach the cortex over the lateral pathways and sensory systems will continue to fire into the reticular formation, ascending reticular discharges are blocked by the transection from reaching the cerebral cortex. In this situation, evoked potentials, representing specific sensory information provided to the cortex via the lateral afferent pathways, continue to be elicited; but generalized EEG desynchronization of the cortex is not sustained. These facts are of considerable importance, since without generalized cortical activation produced by reticular-formation activity, there can be little or no cortical integration of the sensory information which arrives over the specific pathways. Under these conditions information is probably neither retained by the central nervous system nor integrated with other incoming sensory data or stored perceptions and memories. If, on the other hand, the specific sensory pathways are cut at a level above the point at which their collaterals enter the reticular formation, cortical arousal and consciousness is still possible, but the animal is awkward in its movements and limited in its abilities, since the cortex is deprived of specific sensory information.
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| Posted 6 months ago III BRAIN STIMULATION One of the most powerful tools for the study of the interaction between brain function and behavior has been stimulation of the brain. Experimenters have employed a variety of techniques in order to induce an excitation or inhibition of the activity of the nervous system. Some of these include the introduction of chemicals and drugs into the brain. Others have employed heat and cold. However, by far the most widely used technique has been the application of electric currents to brain tissue. The use of these various methods of brain stimulation has resulted in an impressive body of data on the structure and function of the brain and its role in the control of behavior. Using these methods, experimenters have been able to stimulate the brain in one region and record the brain’s response in another region, thus determining the functional connections between brain areas. Others have stimulated various brain sites and observed the overt responses of the organism under study. These responses span the range of behaviors from simple reflexes to complex patterns of responses. Still other investigators have studied the effects of stimulation of specific brain areas on ongoing behavior in order to determine whether that particular brain area is involved in the maintenance of the behavior in question. Space does not permit us to summarize all of the various methods of stimulation that have been utilized. Therefore, discussion will be limited to the effects of electrical stimulation, since this has been the method most generally employed.
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| Posted 6 months ago History The study of the effects on behavior of electrical stimulation of the brain is as old as the study of electricity itself. Count Alessandro Volta (1745-1827), while observing the results of the application of electric currents to his various sense organs, passed the current through his own brain. Fortunately, he survived the experiment. Since that time, but particularly in the present century, many investigators have utilized electric currents to map pathways through the nervous system and to study the effects of stimulation of certain brain areas on behavior. Advances in electronic engineering have made available to the researcher more and more precise instrumentation, and it is now possible to stimulate tiny areas deep within the brain with only minimal damage to neural tissue. In 1870, G. Fritsch and E. Hitzig reported that an electrical stimulus applied to certain regions of the cerebral cortex could elicit movements of the face, arms, and legs of experimental animals. In 1899, Charles S. Sherrington utilized electrical stimulation to demonstrate the reciprocal innervation of flexor and extensor muscles. Although experiments such as these shed a good deal of light on the functioning of reflex pathways, little was learned about voluntary behavior since exposure of such large areas of the nervous system required extensive restraint and anesthesia. The work of W. R. Hess, however, represented a major technical advance with the introduction of the first practical technique for permanent implantation of electrodes into the brain. The development of these chronic electrodes made possible the study of stimulation effects involving deep structures in awake, unrestrained, unanesthetized animals.
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| Posted 6 months ago Methodology Hess’s electrode technique is still the basic method employed, although many laboratories have carried out modifications to suit their own particular needs. Generally, thin, rigid, stainless steel or platinum wires, insulated except at the tip, are lowered into the brain through a small hole in the skull. The animal is deeply anesthetized. Its head is held in a metal frame called a stereotaxic instrument. This frame, a modification of a device originally conceived by Horsely and Clark (1908), not only provides firm support for the head during surgery, but also allows the experimenter to locate deep cerebral structures with a fair degree of accuracy. This is accomplished by means of a threedimensional system of coordinates that locates any given brain structure by its position relative to some zero point. This is analogous to locating a region deep within the earth by means of its latitude, longitude, and distance from the surface. Stereotaxic atlases, or brain maps, of a number of common laboratory animals and of man are availble. Many of these may be found in a book edited by Sheer (1961) along with illustrations of stereotaxic instruments and detailed descriptions of electrode techniques. Because of the variability in skull dimensions among animals, the stereotaxic technique is not completely accurate. It is necessary, therefore, at the termination of the experiment to sacrifice the animal and verify the exact location of the electrode tip by microscopic examination of the brain. An important factor in the study of brain stimulation is the nature of the electrical stimulus. Direct current generally destroys nerve tissue. Indeed, prolonged application of direct current has often been used to produce lesions for experimental or clinical reasons. However, brief pulses of direct current, alternations of positive and negative pulses, and sine waves have all been used with good results in brain stimulation studies, provided that the current is not excessive.
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| Posted 6 months ago Sleep, wakefulness, and emotion Using the technique of chronic implanted electrodes, Hess systematically explored the diencephalon of the cat. He has summarized his findings in two recent works (1949; 1954). In brief, Hess observed that stimulation of the massa intermedia of the thalamus resulted in a progressive decrease of activity followed by sleep. The animal could be aroused from this sleep by some external stimulus, but once the external stimulus was removed, the animal returned to sleep. Stimulation of the posterior hypothalamus, however, resulted in immediate wakefulness and a state of excitation. With stronger stimulation in this area, the cat would hiss, bare its teeth and claws, arch its back, and show all the signs of rage and fear. Upon termination of the stimulation, the rage reaction ceased. Moreover, if the cat was provoked during this period of stimulation, a highly organized attack reaction would be directed toward the provocative object. Egger and Flynn (1962) have described a study, however, in which stimulation of the lateral nucleus of the amygdala suppressed the attack reaction elicited by hypothalamic stimulation. [seeEmotion and Sleep.]
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| Posted 6 months ago Motivation A variety of motivational effects have been evoked by electrical stimulation of the brain, particularly in the region of the hypothalamus. Perhaps the most striking of these are the effects on food and water intake. Food intake. Delgado and Anand (1953) have demonstrated that cats stimulated in the lateral hypothalamic area will perform gnawing and chewing movements and will increase their food intake. It is interesting to note that the increased eating is not necessarily directed toward edible objects. Miller (1960) reports that upon presentation of stimulation to the hypothalamic feeding area, rats will gnaw on blocks of wood, sticks, etc., when food objects are not available. This behavior has been described as “stimulus-bound” eating. Miller also described a study in which rats with electrodes in the lateral hypothalamus had been trained to press a lever for food. The animals were later foodsatiated and stimulated in the hypothalamus. Immediately following the onset of stimulation the animals began to press the food lever. When the stimulation was terminated, the rats immediately stopped lever-pressing. Suppression of food intake by stimulation has also been reported. An investigation by Wyrwicka and Dobrzecka (1960) demonstrated that stimulation of the ventromedial nucleus of the hypothalamus of hungry goats resulted in immediate cessation of eating. Water intake. Less work has been done in the area of central nervous system control of other motivational mechanisms by means of stimulation. Chemical stimulation of the hypothalamus has been demonstrated to control water intake effectively (Grossman 1960). There is also evidence that electrical stimulation can influence the thirst mechanism (Andersson & McCann 1955). Sexual behavior. Vaughan and Fisher (1962) have demonstrated that stimulation of the lateral anterior hypothalamus results in a marked increase in sexual capacity in male rats. MacLean and Ploog (1962) have reported that stimulation of a number of forebrain and diencephalic areas in monkeys results in erection of the penis and ejaculation. [SeeSexualBehavior, article onAnimalSexualBehavior.]
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| Posted 6 months ago Learning The application of the techniques of intracranial stimulation to the study of the neuropsychology of learning has been a fruitful one. The typical approach has been to compare the performance of experimental organisms on a learning task with and without stimulation of some neural areas. This method has advantages over the traditional lesion technique, which requires elaborate control groups to assess the possible extraneous effects of surgical shock, blood loss, intracranial pressure, etc. In a stimulation study, each animal frequently serves as its own control, since the effects of stimulation are almost always reversible. However, certain learning tasks may necessitate the use of a control group with nonstimulated electrodes. Experiments employing electrical stimulation of the brain to investigate learning are numerous. A few examples of recent studies may serve to illustrate some of the major trends in current research. Delayed alternation and visual discrimination. Rosvold and Delgado (1956) trained monkeys to perform a delayed-alternation task in which the animals were required to seek a peanut reward under the right food cup on one trial, under the left food cup on the next trial, under the right on the next, etc. A five-second delay was interposed between trials. Failure to alternate responses on successive trials was regarded as an error. The monkeys were also trained to perform a simple visual discrimination. Following stabilization of performance on these tasks, the monkeys were implanted with multiple-lead electrodes. Upon recovery from surgery, the animals were again tested on the delayed-alternation and visual discrimination problems. During a part of the testing session, electrical stimulation was delivered through one of the electrode leads. The experimenters found that performance declined markedly on the delayedalternation test when the lead stimulated was in the caudate nucleus. However, the same stimulation failed to interfere with the visual discrimination task. A later study by Buchwald and others (1961) revealed that stimulation of the caudate nucleus does interfere with acquisition of a visual discrimination task, but does not alter performance once the task has already been learned. Operant conditioning. In another type of experiment, Knott and others (1960) trained cats to press a lever for meat reward in a modified Skinner box, an apparatus provided with devices, e.g., levers or buttons, whose appropriate manipulation leads to some desired outcome. Electrodes were implanted in a number of deep cerebral structures. Continuous low-intensity stimulation was delivered during the lever-pressing task. The experimenters reported that stimulation of the hippocampus, caudate nucleus, and thalamus failed to alter the leverpressing rate. Stimulation in the septal area, however, resulted in a cessation of response during stimulation. The animals resumed pressing the lever following termination of the stimulation, but only after a prolonged delay. Hypothalamic stimulation generally had the same effect as septal stimulation, except that the post-stimulation delays were shorter. The authors concluded that these data support the notion that certain neural pathways critical to the mechanisms of learning and retention become “occluded” by the stimulation. The data could also be interpreted, however, in terms of interfering emotional responses evoked by stimulation of the hypothalamus and the septal area. Classical conditioning. A third type of investigation utilizes the classical Pavlovian conditioning paradigm except that both the conditioned stimulus (CS) and the unconditioned stimulus (US) are presented to the subject by means of implanted electrodes. Doty and Giurgea (1961) implanted electrodes in the motor area of the cerebral cortex of dogs, cats, and monkeys. Stimulation of these electrodes served as the US. The unconditioned response was the particular limb movement that resulted from the stimulation of a direct motorpathway. Stimulation in some other cortical area served as the CS. Six to ten pairings of the CS and the US were made daily. The experimenters were able to obtain clear-cut evidence of conditioned reflexes with cortical stimulation as the US. Aside from their implications for learning theory, the experiments of Doty and Giurgea are important because they suggest the possibility of studying the electrophysiology of the conditioning mechanisms in a simple form with known and well-defined inputs.
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| Posted 6 months ago Speech and memory A series of remarkable experiments have been carried out by Penfield and Roberts (1959). These investigations have involved the electrical stimulation of the cerebral cortex of awake humans. The studies were carried out during the course of neurosurgical procedures for the treatment of epilepsy. During the operation, the comfort of the patient was maintained at all times by the careful administration of local anesthesia. In addition to yielding highly detailed maps of the sensory and motor regions of the human cerebral cortex, this research has also indicated the location of areas that are involved in the mechanisms of speech and memory. Penfield has described regions on the temporal lobe in which electrical stimulation will result in the recall of vivid experiences of sights and sounds. The patients report that they feel as if they were reliving those events. Moreover, a specific memory can be repeated by interrupting the stimulation and then quickly reapplying it. Penfield suggests that memory is organized on the temporal lobe in somewhat the same manner in which electrical impulses representing visual and auditory patterns are stored on magnetic tape. Thus, whenever a particular region of the brain is made to yield up its stored memories, the memories are recalled with the same vividness and clarity as when they were originally stored. He further suggests the existence of mechanisms in the brain that inhibit the retrieval of those stored memories, the electrical stimulation somehow bypassing or otherwise nullifying the influence of these inhibitory mechanisms. Penfield and his coworkers have also mapped areas of the cortex that upon stimulation produce vocalization, hesitation and distortion of speech, repetition of vocal sounds, interference with speech, and complete arrest of speech. In addition, stimulation of certain areas results in an inability to name specific objects although the remainder of the speech mechanism seems to be unimpaired. These areas are mainly included in the parieto-temporaloccipital cortex, although the effects have also been obtained from several other well-defined cortical regions.
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| Posted 6 months ago Self-stimulation In an attempt to study the effects of subcortical stimulation on learning in rats, Olds and Milner (1954) placed a rat with a forebrain electrode in an open field maze. They observed that if the rat received stimulation in a particular place in the maze, it would spend more and more time in that place. The stimulation seemed to have a rewarding effect on the rat’s behavior. Next, they placed the rat in a T maze. The animal learned to go to that arm of the maze in which it was rewarded with brain stimulation. Microscopic examination of the rat’s brain revealed that the electrode tip was in the vicinity of the anterior commissure. Rats used subsequently were trained to deliver the stimulation to themselves by pressing a lever in a Skinner box. These animals would deliver several hundred stimulations per hour to themselves. The stimulation served as the sole reward for pressing the lever; food and water were never present in the box. Since the original report by Olds and Milner, many of the variables affecting the self-stimulation phenomenon have been explored. Some of the principal variables have been the animal species; the location of the electrodes in the brain; the intensity, frequency, and duration of stimulation; the motivational and emotional state of the organism; and the schedule of reward. In addition, the effects of drugs on self-stimulation have also been studied. Species. Self-stimulation of the brain has been clearly demonstrated in the following animals: rat., dog, cat, pigeon, monkey, dolphin, guinea pig, gold-fish, and man. At this point, experimenters would be more interested in learning of a vertebrate in which the phenomenon could not be demonstrated than in further additions to the list of positive instances. The demonstration of self-stimulation in man is of particular interest since this is the only species that can provide verbal reports about the subjective experience of brain stimulation. Sem-Jacobsen and Torkildsen (1960) have reported cases in which electrodes were implanted in the brains of human patients for several months in the course of treatment for Parkinson’s disease. During exploratory stimulation, the experimenters encountered several regions of the forebrain in which the patients seemed to enjoy the stimulation. They would smile or grin and express a desire for repeated stimulation. If given an opportunity to press a button to deliver the stimulation to themselves, the patients would press the button often. Their verbal reports ranged from descriptions of tickling sensations to expressions of satisfaction and euphoria. Anatomical variables. In general, self-stimulation has been demonstrated in those structures of the brain that form part of the limbic system or have strong anatomical connections with it (Olds 1956). Application of stimulation in purely sensory or motor areas does not appear to have any rewarding effect. Moreover, there are areas in which the effect of the stimulation appears to be punishing, and animals will learn to escape and avoid stimulation in these areas (Brown & Cohen 1959). Many of these negative regions correspond to the areas that, upon stimulation, produce rage and attack reactions. Within the positive reward system, rates of selfstimulation vary widely from area to area and from species to species. Thus, a rat may self-stimulate 800 to 1,000 times per hour when the locus of stimulation is the septal area, 2,000 times per hour when it is the hypothalamus, and 4,000 times per hour when it is the tegmentum. Furthermore, the reported rates of responding may vary considerably among laboratories because of differences in apparatus. For example, it is possible to obtain higher response rates with a telegraph key lever than with a microswitch lever. If an animal presses a lever 1,000 times per hour for septal stimulation and 2,000 per hour for hypothalamic stimulation, is it safe to conclude that the hypothalamic stimulation is more rewarding to the animal? An experiment by Hodos and Valenstein (1962) suggests that it may not necessarily be. Rats were given a choice, in a two-lever box, between septal stimulation and hypothalamic stimulation. These were presented at various intensities. When presented with high-intensity septal stimulation on one lever and low-intensity hypothalamic stimulation on the other lever, the animals showed a clear preference for the lever producing septal stimulation once they discovered it, even though their response rates were much higher on the lever producing hypothalamic stimulation. Considering these findings, it would seem hazardous to draw general conclusions about the relative reward values of stimulation in different neural areas based solely upon rate of responding. Directpreference tests at several intensities of stimulation provide data that are less subject to interpretive ambiguities resulting from the interfering influence of motor side-effects produced by the stimulation.
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| Posted 6 months ago Stimulation variables. Reynolds (1958) demonstrated that as intensity of stimulation increased, the rate of responding increased, passed through a maximum, and then declined. Keesey (1962) confirmed this finding and, in addition, reported similar effects with variation in stimulation frequency and duration. This decline in self-stimulation rate with high intensities of stimulation should not necessarily be interpreted as a decline in the rewarding properties of the stimulation. Hodos and Valenstein (1962) reported that when given the choice between two intensities of stimulation in the same area, rats consistently chose the higher of the two intensities, even though they self-stimulated at a lower rate for the higher intensity. Motivational factors. Brady and others (1957) observed that animals would self-stimulate faster when deprived of food or water than they would when they were satiated. This was later confirmed by Olds (1958), who further reported that injections of male sex hormones in castrated male rats also increased the rate of electrical self-stimulation. However, Hodos and Valenstein (1960) failed to find any effects on the septal self-stimulation rate in spayed female rats injected with female sex hormones. Recently, Hoebel and Teitelbaum (1962) have demonstrated an interesting correlation between the hypothalamic areas controlling feeding behavior and those yielding self-stimulation. They suggest that the feeding system may control selfstimulation in a manner similar to its control of food intake. Emotion. Brady and Conrad (1960b) demonstrated an interesting effect of self-stimulation on emotional behavior. Rats were trained to press a lever for either intracranial stimulation or water reward. Periodically, an auditory stimulus was presented for a period of five minutes. At the termination of this stimulus, a painful electric foot-shock was administered to the animals. When the animals were pressing the lever for water, presentation of the auditory stimulus produced a clear suppression in the rate of responding. This has been described as conditioned “fear” or “anxiety” in previous studies. No conditioned “fear” response to presentation of the auditory stimulus could be elicited, however, when the animals were leverpressing for brain stimulation in the medial forebrain area. Moreover, the possibility that the animals were unable to hear the auditory signal during the brain stimulation has been eliminated by Beer and Valenstein (1960), who showed that rats could, with little difficulty, make auditory discriminations when the auditory signals were presented simultaneously with brain stimulation. Reward schedules. Sidman and others (1955) presented data illustrating the point that behavior rewarded by brain stimulation on either a fixedratio or a variable-interval schedule generally has the same characteristics as food-rewarded behavior on the same schedule. However, Brodie and others (1960) reported that fixed ratios of more than 20 responses to each stimulation are difficult to maintain in monkeys unless very slow and gradual training is permitted. Fixed ratios of several hundred responses to each reward are not at all uncommon when animals are rewarded with food. Moreover, these researchers report far less resistance to extinction of performance when brain stimulation is the reward than has often been observed when the schedule is equivalent but the reward is food. In a study of timing behavior, Brady and Conrad (1960a) required monkeys to space their responses at least 20 seconds apart in order to receive a reward. In the case of food reward or anterior thalamic stimulation reward, the animals had no difficulty in delaying their responses for the required period of time and thereby received a large percentage of the possible rewards. The most frequent interresponse time was 20 seconds. However, in the case of stimulation of the globus pallidus, the animals were unable to delay their responses sufficiently and thereby received few rewards. The most frequent interresponse time was 10 seconds. There is some suggestion that the stimulation may have interfered with the mechanism of time perception in these animals. Pharmacological effects. Olds and others (1956) as well as other workers have demonstrated that some drugs can affect self-stimulation performance. Chlorpromazine and reserpine, both tranquilizing drugs, each depressed the self-stimulation rate in some areas, but not in others. However, phenobarbital was not observed to produce any specific effect on performance.
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| Posted 6 months ago Some general problems When an experimenter produces a change in behavior by stimulation of a brain area, there is a great temptation to speculate on the possible role of that area in the mechanisms underlying the behavior. However, such speculations should be made cautiously, because the electrical stimulus may not have the same effect on all neural areas. For example, we have seen that stimulation of the caudate nucleus results in a deficit in delayed-alternation performance and that stimulation of the amygdala suppresses rage reactions. These behavioral deficits are the same as the effects of lesions in those neural areas. Therefore, it seems likely that the electrical stimulation had a suppressing effect upon normal function. On the other hand, we have seen that electrical stimulation of the lateral hypothalamus yields eating responses while ventromedial hypothalamic stimulation results in suppression of eating. These effects are the opposite of those observed when lesions are made. Presumably, in this case, the stimulation was augmenting the activity of the feeding areas. Thus, the effects of stimulation studies alone are not sufficient for determining the role that a cerebral structure may play in behavior. A second and related problem is that of attempting to generalize from the highly unnatural type of stimulation that experimenters introduce into the brain to the normal types of physiological events present in the nervous system. The two may not necessarily have the same effect on neural tissue, and cautious interpretation of data is required. A detailed discussion of the problems of the interpretation of the behavioral effects of electrical stimulation of the brain may be found in a recent paper by Valenstein (1964). A third problem is that of the possible interaction between brain stimulation and other environmental variables that may be affecting the behavior under study. We have seen that caudate nucleus stimulation will markedly interfere with visual discrimination performance if the animal is still in the process of acquiring the discrimination. However, the effects will be scarcely detectable once the animal has mastered the problem. Similar difficulties may arise in the study of emotional behavior, memory, and perception. Therefore, a thorough knowledge of the environmental variables that influence behavior is essential before attempting to study the effects of stimulation on behavior.
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| Posted 6 months ago IV ELECTROENCEPHALOGRAPHY Electroencephalography derives its name from the fact that it provides a graph or recording of the electrical activity of the brain, or encephalon. In human subjects—except when the brain surface is exposed during a surgical operation—the electroencephalogram (EEG) is always recorded from electrodes distributed over the surface of the scalp. A variety of electrodes have been used in the past, but at the present time two principal types are in use. One type consists of silver dish-shaped or cupshaped electrodes, 5 millimeters in diameter, under which a conductive paste or electrode gel serves as an electrolyte between the surface of the scalp and the electrode. The other type consists of a finegauge stainless-steel needle, which is painlessly inserted through the scalp. The electrodes are connected by fine insulated wires to a plug-in box which identifies numbered locations on the surface of the head, usually according to a system agreed upon by the International Society of Electroencephalography (Jasper 1958) and known as the 10-20 system. The locations of the electrodes on the head are adjusted according to the over-all head dimensions by measuring from the nasion (bridge of the nose) to the inion (external protuberance at the back of the skull) and by defining electrode positions at intervals of 10 or 20 per cent of the total distance; measurements from the auditory meatuses over the top of the head are similarly fractionated. Finally, measurements of the circumference of the head enter into the placement of the electrodes. Such procedures insure that electroencephalographers will use a uniform system of deriving the potentials on which to base their reports. The number of electrodes placed on the head depends upon whether only a survey of representative areas is to be made or whether a detailed comparison of many areas is to be made in order to localize some relatively focal disorder, as in the cases of temporal lobe epilepsy, of a local region of trauma, or of a brain tumor. From the plug-in box, where electrodes representing the various areas converge, a larger, electrically shielded cable enclosing all of the individually insulated wires carries these to the electroencephalograph. The EEG recording unit consists of voltage amplifiers and power amplifiers and some type of write-out and/or display unit. Since the potential differences between any two electrodes on the surface of the scalp are of the order of millionths of a volt and usually range from 2 or 3 microvolts to 100 microvolts—except in the case of some pathological discharges which may reach 500 microvolts—these minute potentials must be amplified at least a millionfold in order to be recordable or visible on the display unit. The voltage amplifiers are responsible for the amplification; the power amplifiers, as their name implies, provide an increase in the power of the signals so that a relatively insensitive recording unit may be driven by the variations in the alternating current. The amplifiers themselves are usually of the resistancecapacity coupled type with a time constant suitable for amplifying and transmitting the potentials to the recording unit without appreciable distortion. The recording unit is typically an inkwriting oscillograph, although it may be a cathode-ray oscillograph or an optical oscillograph. The latter types require the recording of the deflections of a light beam on photographic film or paper. The inkwriting oscillograph consists of a pen motor which converts the electrical alternations from the brain into mechanical movements of the pen, tracing in the form of an arc the variations of potential as they occur. These oscillations of the tip of a stylus or pen upon paper moving at a uniform rate provide a tracing or recording upon the paper, which is a form of graph, with time represented on the abscissa and voltage on the ordinate. Thus, the frequency, magnitude or voltage, and the pattern of the waves so traced on the paper may be studied and measured relative to standards or norms of expectancy established for normal subjects. Abnormal or pathological deviations may be noted and contrasted with normal patterns.
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| Posted 6 months ago History Now let us return to the history of electrical activity of the brain. Caton (1875), an English physician, was the first to publish an account of the recording of electrical variations from the surface of the exposed brain of the rabbit. His findings were confirmed in a study of the dog by Russian and Polish investigators (see Brazier 1959), during the last quarter of the nineteenth century, but little further use of the phenomenon in interpreting the brain’s activities and functions was made until Hans Berger (1929), a German neuropsychiatrist, published the first report on the recording of the human EEG. Since Berger’s recordings showed predominantly wavelike oscillations, the classical neurophysiologists of this period, being acquainted only with spikelike deflections due to transient electrical potentials observed in peripheral nerves, looked upon his findings with doubt and felt that the waves might result from artifacts rather than brain activity. However, Adrian and Matthews (1934), distinguished electrophysiologists at Cambridge University, England, verified Berger’s findings and put their stamp of approval upon the work. Thus, for the first time it was recognized that at least two types of electrical activity exist in the nervous system: (1) when a nerve or neuron is excited and transmits an impulse, it* is accompanied by a sharp, spikelike negative potential and some wavelike after-potentials; (2) aggregates of neurons with their cell bodies and dendritic processes, such as are found in the cerebral cortex of the brain, give rise to continuous, autonomous electrical changes of wavelike nature. The spike discharge of the neuron is a discrete event and tends to occur only when the neuron is excited by impulses reaching it from a receptor or another neuron. In contrast the wavelike activity of large numbers of neurons closely arrayed structurally appears not to be dependent upon sensory input or excitation.
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| Posted 6 months ago Patterns of brain activity The EEG is composed principally of this wavelike activity, which in the human is continuously present in one form or another as long as the brain is alive. Typically, with the eyes closed and with the subject or patient relaxed and not thinking about anything in particular, Berger found, in the adult, that rhythmic waves, which he called alpha waves, were emitted at a rate of 10 per second. He noted that the frequency of these waves ranged from about 8 to 12 per second in different individuals, including older children and adults, but that very young children had slower alpha waves. We now know that alpha waves first appear in regular sequence over the sensory areas of the brain at about 3 months of age, at a frequency of 3 to 4 waves per second. By 1 year of age they have increased to 5 or 6 waves per second, and by 10 or 12 years they have reached the adult frequency, averaging about 10 per second. Such waves are remarkably constant in frequency, varying less than 1 cycle per second in a given individual, from one time to another—from moment to moment, from day to day, or from month to month. Anything which increases or decreases metabolic rate will shift the frequency, up or down respectively, 1 or 2 cycles per second. For example, in hypothyroidism, where metabolic rate is lowered, the alpha frequency tends to be 1 or more cycles slower than the norm for that individual or the average for a normal group; hyperthyroidism, with increased metabolic rate, tends to increase the alpha frequency. The administration of thyroxine, a thyroid extract used in the treatment of hypothyroidism, increases the alpha-wave frequency from its low level. Artificially increasing the body temperature by diathermy or other means usually increases alpha frequency accordingly; also, natural fevers tend to increase alpha frequency. Some people believe that the alpha rhythm constitutes a basic time clock in the nervous system, regulating or modulating other rhythmic processes of the brain and body and possibly playing a role in our estimation of timev]. v |
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| Posted 6 months ago It is normal to have an alpha rhythm when awake, relaxed, and not being stimulated. Also, it is normal for the alpha rhythm to be blocked by any type of stimulation which attracts the attention of the subject and alerts him or arouses him from his temporary quiescence or daydreaming. The alpha rhythm quickly recovers from this blocking and continues with its oscillations until another stimulus occurs. If the same, identical stimulus is repeated a number of times at more or less regular intervals, the subject tends to ignore it, and the brain’s response of blocking the alpha rhythm disappears. This state of adaptation or lack of responsiveness is sometimes referred to as habituation. As soon as the quality of the stimulus changes in either intensity, kind, or another characteristic, the blocking of the alpha rhythm will return, as it does in the case of any novel stimulus which attracts attention [seeAttention]. The alpha rhythm, or other characteristics of the EEG, are not significantly correlated with intelligence or personality traits. For references to numerous early studies of this relationship, as well as more detailed descriptions of the EEG, see extensive reviews by Jasper (1937) and Lindsley (1944).
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| Posted 6 months ago Sleep. The electroencephalogram has proven to be a useful correlate of psychological conditions. In the case of the transition from a waking state to sleep, in human subjects as well as lower animals, there are a series of fairly well-defined stages. There is an alpha rhythm of about 10 waves per second in normal, adult humans who are relaxed, awake, and free of special sensory stimulation. As drowsiness occurs the alpha rhythm slows slightly, decreases slightly in voltage, and shows brief periods of subsidence. As the drowsiness increases to the point where the subject is not aware of events in his environment, slower waves (3 to 8 per second) begin to appear, interspersed with declining alpha waves, and as the first true state of sleep occurs, there appears in the tracing periodic bursts of waves, called sleep spindles, at about 14 per second. In the next stage of sleep the spindles disappear and more and slower waves of higher voltage appear. Beyond this point, the waves become still slower and more prominent, ranging from 1 to 3 per second; these are called delta waves. At this time the state of sleep is deep, and it is more difficult to arouse the subject from his slumbers by various kinds of sensory stimuli. More recently, however, a still deeper stage of sleep has been described in which muscles of the limbs, body, and neck become more relaxed and the EEG shifts from large and very-low-frequency slow waves to fast, low-amplitude waves similar to those seen when a person is awake and alerted or excited. This state of sleep is sometimes called paradoxical sleep, because while the subject appears to be sleeping deeply and is difficult to arouse, his EEG appears to be that of a waking person. Such periods occur within an hour after going to sleep and last only a few minutes but recur again and again during the course of the night’s sleep, usually several times for durations of 5 to 15 minutes. It was discovered by Kleitman ([1939] 1963) and his associates, Aserinsky and Dement, that rapid eye movements occur during these comparatively brief periods in the course of the night’s sleep; more recently these periods have been referred to as REM-sleep, that is, rapid-eye-movement sleep. If a subject is awakened during one of these REM states, he usually reports that he has been dreaming, but he is less apt to have been dreaming if awakened during the slowwave stage of deep sleep. Consequently, REM-sleep is associated with dreaming, and there is some evidence (Dement 1965) that a person needs a certain amount of this type of sleep to remain mentally healthy. [ Contrary to widespread belief, fostered by advertising and commercial agencies selling “learn during sleep” apparatus or programs, no well-controlled and validated studies using the EEG to monitor the presence and depth of sleep show that learning does indeed occur during sleep. Simon and Emmons (1955) have reviewed this subject.
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| Posted 6 months ago Waking and anesthetic states. As we have seen in the instance of transitions from wakefulness to sleep, variations in the state of consciousness or awareness of one’s environment are correlated with marked changes in the EEG. When a person is alerted, excited, or aroused, EEG activity is low, fast, and not particularly rhythmic; when a person is awake, relaxed, and not very attentive—as in reverie or daydreaming—there are good rhythmic alpha waves; at the onset of true sleep, the alpha waves are displaced by slower waves and spindle bursts at the rate of 14 per second; in deep, quiet sleep without complete relaxation, large, billowy, slow waves are the rule; finally, in the most relaxed state, but with REM and dreaming, there are lowamplitude fast waves again. During the anesthetic state, in passing from one plane of anesthesia to another, similar sequences occur but vary with the type of anesthetic used. In general, it is believed that in both sleep and the anesthetic state a core structure of the lower brain stem, known as the reticular formation, is depressed or inhibited. The reticular formation can be activated by all types of sensory stimulation; in fact, it is generally referred to as a nonspecific sensory-arousal mechanism, which appears to keep the brain awake and alert, and perhaps even attentive, when properly sensitized to incoming sensory stimuli. It may also be affected by neurohumoral substances and endocrine secretions and by other constituents of the blood stream and cerebrospinal fluid. These physiological and biochemical factors are important in the functioning of all brain and nervous-system cells, and the activity of these neurons is reflected in their electrical activity, of which the EEG provides one measure.
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| Posted 6 months ago Neurological disorders Although in the past the EEG has not proved very useful in differentiating psychiatric patients from normal subjects or in distinguishing different personality characteristics, it has been a very valuable supplementary tool for the neurologist and neurosurgeon. It quickly proved its value in connection with a variety of neurological disorders. Perhaps most outstanding in this respect were the epilepsies. The minor attack, or petit mal, has a characteristic signature known as the spike-andslow-wave pattern, which recurs about three times per second. In the major attack, grand mal, where there are convulsions and loss of consciousness, the pattern of the EEG begins with increasedamplitude fast activity associated with stiffening and loss of consciousness, known as the tonic state, and shifts to grouped activity as the convulsive movements occur; finally, after the series of convulsions ceases and the EEG quiets to an almost isoelectric state, a period of coma and marked relaxation exists. Generally, this state of the EEG and behavior lasts only several seconds. Then the EEG develops a deep-sleeplike pattern with large, billowy waves, and the patient appears to sleep deeply for a number of minutes or even a few hours before awakening, usually with return of the EEG to a fairly normal status. A concussion produced by a blow on the head with temporary or prolonged loss of consciousness is accompanied by large, slow delta waves. A focal injury to the brain, caused by a rupture of a very small blood vessel (petechial hemorrhage) or a penetrating wound, gives rise to aperiodic spikelike discharges in that local region and, thus, helps to localize the source of the damage or injury. Encephalitis and other infections which damage tissue have toxic effects or create pressures which usually produce some kind of aberration in the EEG. Thus, the neurologist and the neurosurgeon find the EEG useful as a supplement or adjunct to their other techniques of detecting and localizing brain injuries or in diagnosing the source or cause of a neurologic disorder. The merit of the EEG is that it is a painless and harmless procedure and has no aftereffects, as do some neurologic test procedures. Almost every major hospital in the country and many lesser ones now have some facility for EEG recordings and for using these not only as a diagnostic tool but also as a tool measuring the effects of anesthetics in the operating room or measuring the effects of particular drugs or treatment procedures.
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| Posted 6 months ago Sensory processes The most recent development involving the EEG is the recording of average evoked potentials over sensory areas of the brain by means of computer summation of responses that are time-locked to their stimuli; the stimuli are repeated as many as a hundred times in order to build up a reliable average. These responses from sensory areas, and even from association areas, are very small in comparison to the ongoing EEG activity; their voltage generally is of the order of five to ten microvolts, whereas the alpha waves and other EEG activity may be several times larger. Using these computer-averaged responses, the senses of vision, audition, and bodily sensation have been studied (see Sensory Evoked Response in Man 1964). Thus, human sensory experience may be studied by judgmental or psychophysical methods and, at the same time, by electrical recording from the particular area of the brain concerned. Other sensory-related functions have been studied with interesting results. Haider and his colleagues (1964) demonstrated the usefulness of the computer-averaging methods in relation to an experiment on attention and vigilance maintained over time. They found that as attention diminished generally, so that a subject no longer detected certain dim flashes imbedded in a series of brighter flashes with the same percentage of accuracy, the magnitude of the evoked response diminished. As vigilance waned, so did the voltage of the evoked potentials; however, if a subject had a five-minute period in which he detected all interspersed signals, the magnitude of his evoked potentials for that period was as great as at the beginning of the task. In another experiment, Spong and his colleagues (1965) demonstrated that by directing the attention of a subject to a particular class of sensation—vision, for example—and by telling him to ignore audition, even though flashes and clicks alternated as stimuli throughout, the magnitude of the average evoked potential was always greater over the visual area when attention was directed toward visual stimuli and auditory stimuli were ignored; the opposite was true for responses over the auditory area. Thus, there is significant evidence that attention and vigilance have a central response which may now be recorded and studied in relation to psychological set or instruction. In addition, Walter has found by a similar method, using direct-current recording, that giving a subject an instruction and an “expectancy” leads to a build-up of potential until that expectancy is met, either correctly or incorrectly, at which time a discharge occurs (Walter et al. 1964). He calls this a “contingent negative variation” It is a brain process which undoubtedly plays a role in decision making and other higher cognitive functions, and will prove to be a valuable probe for getting at problem solving, thinking, and other functions of a higher level. The electroencephalogram has played a very useful role in the hands of the neurophysiologist and neuropharmacologist in the study of the electrical activity of the brain. It has proved to be a very considerable stimulus to brain study in animals and man, because it provided a parameter of measurement not previously possible. The psychologist and neurophysiologist today frequently combine their talents in the study of brain and behavior correlations, and through this work a better understanding of the mechanisms and organization of the brain and nervous system is coming about. Thus, the electroencephalogram is an experimental tool useful in research on the brain, as well as a clinical tool useful in the clinic and hospital. |
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| Posted 6 months ago
Vitamin B6 for regular nervous system function, production of normal red blood cells and for protein metabolism
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| Posted 6 months ago Treatment Of Epilepsy Natural Epilepsy is a neurological disease that disturbs the electrical functioning of the brain. Natural Epilepsy results in altered consciousness and seizures. The disease is a common one, with as many as 50 million people worldwide who are suffering from it. Though the condition is incurable, proper medication can control it effectively. There are different types of epilepsies, depending on the cause symptoms and the area of seizure. Some epilepsy occurs only during childhood.
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| Posted 6 months ago Nervous System? What Is the Role Of the Nervous System?
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