Ital J Neurol Sci (1999) 20:7-15 ORIGINAL

Springer-Verlag 1999

E. Niedermeyer

A concept of consciousness

Received: 15 December 1998 / Accepted: 11 January 1999

Abstract The term consciousness, so widely used in clinical settings, is considered extremely complex and practically undefinable. Historically, consciousness was ignored by the great classical philosophers, and was regarded as a basic condition of being. This changed with Descartes and during the ensuing centuries. A simplified, reductionistic and easily definable concept of consciousness is proposed; consciousness is proposed to consist of three main components: vigilance, mental contents, and selective attention. These three components can be investigated with modern neuroscientific methods - vigilance being the most readily explorable function. The striking differences between sleep and coma are pointed out (along with some observations on sleep initiation). Special attention is paid to epileptic impairment of consciousness and, in particular, to the spike-wave absence which is thought to be due to a temporay suspension of the working memory circuits within the frontal lobe. Key words Consciousness Vigilance Attention Selective consciousness Historical aspects Sleep Initiation Coma Epilepsy

What is consciousness? The term consciousness plays a major role in the daily routine work of most clinicians. Whenever acute cerebral involvement assumes serious character, the function of consciousness may become progressively impaired to the degree of profound coma. This so widely used term, however, is a muddled one and has always defied lucid analysis and clear definition. Consciousness has been described as the most obvious and the most mysterious feature of our minds by Dennett [1] who has been one of the most ardent and indefatigable workers in this field.

Attempted definitions of consciousness Attempts to define consciousness have yielded quite different results. Consciousness is the state of awareness of self and the environment [2] - a very terse statement which introduces a second unknown awareness. There is consciousness being defined as the sum of the following components: alertness, attention, sensation and perception, perceptual motor integration, self awareness, motivational systems and memory [3]. By far more complex are older definitions by Schiller (1952) [4] and Bumke (1941) [5], comprising huge portions of the entire field of psychology. Wundt (1911) [6] defined consciousness as the total con-

This paper was presented in part at Electrophysiology of the Brain. A Symposium in Honor of Ernst Niedermeyer, Johns Hopkins Medical Institutions, Department of Neurology, Baltimore, Maryland, 17 January 1998. It was also presented in part at the Simposio Interregionale Universit di Messina, Universit di Catania at Giardini Naxos (ME), Sicily, 24 October 1998

E. Niedermeyer () Departments of Neurology and Neurosurgery The Johns Hopkins University School of Medicine and Division of Neurology The Sinai Hospital 305 Colonial Court, 21204 Towson, Maryland, USA

E. Niedermeyer: A concept of consciousness

tent of our immediate experiences, thus emphasizing the perceptory component. All this indicates that the search for a clear concept of consciousness appears to be never ending. It is a term which cuts across the domains of clinical medicine, neurosciences, psychology, and philosophy. It is unlikely that a panel of specialists in these fields will ever agree on a generally acceptable definition. In spite of these seemingly insurmountable problems, a concept of consciousness will be attempted in the following - a very simple and reductionist concept.

A simplified concept Consciousness can be reduced to three main components: vigilance, mental contents and selective attention [7-10]. This is indeed a vastly reduced concept: it is consciousness without the genitive of. In other words, a state of being conscious - just conscious, period. Not being conscious of something or someone. Naturally, strong selective attention may target an object of which the person becomes aware. This is a major step which transcends beyond the confines of consciousness into the domain of awareness. The term awareness is linked to the genitive of, is by far more complex and - in accordance with the presented concept - has to be separated from the simple function of consciousness. I shall return to this basic concept after a discussion of the aforementioned main components: vigilance, mental contents and selective attention.

Fig. 1 A diffuse projection system connecting the ascending activating reticular formation with the cortex. This figure also shows specific afferent systems on route to the cortex, also branching off connections feeding into the brainstem reticular formation. (Modified from [12])

Sleep Sleep is, of course, a physiological state and the differences between sleep and coma (or abnormal subcomatose states) can hardly be overemphasized. The physiological criteria of sleep have been well established since the late 1950s [16]. The groundbreaking work of Jouvet [17] has demonstrated the dichotomy of NREM/REM sleep and their underlying pontine and pontomesencephalic substrata. In the wake of this work, sleep staging has become the mainstay of modern polysomnography. Even the clinical electroencephalographer in daily routine work has ample opportunity to observe the sleep stages and is constantly impressed by their powerful and trustworthy electrophysiological correlates especially in adulthood (Fig. 2). Physiological variants of these patterns are limited and seldom important.

Vigilance In order to be conscious, there has to be a system which governs the function of vigilance. The neurophysiological basis of vigilance has proved to be readily explorable by the neurosciences. The ascending activating pontomesodiencephalic reticular formation has been recognized as the principal substratum of vigilance since the pioneering work of Moruzzi and Magoun [11], Magoun [12] and numerous further investigators (Fig. l) who have also succeeded in the demonstration of cholinergic [13], noradrenergic [14] and serotonergic [15] mechanisms. Electroencephalography has proved to be a highly valuable method of exploration since desynchronization (acceleration and voltage decline) of the encephalogram (EEG) indicates cortical activation [11], whereas brain stem lesions result in EEG slowing and coma in the experimental animal. Even the routinely used clinical EEG constantly reflects the fluctuating action of the reticular activating system. Vigilance is the basic component for the function of consciousness: heightening to the state of alertness by excitement, danger or the use of stimulants but also slackening in drowsiness and depression (but not abolishment) in sleep.

The initiation of sleep The all-night architecture of sleep is apparently better known than the mechanisms of sleep initiation. Personal observations regarding sleep onset might support this statement. Readers may forgive me for this digression in the logical sequence of sections (adding to the subjective flavor of this presentation). Depth electroencephalography performed in adult epileptics (candidates for seizure surgery) has shown the presence of sleep spindles in mesio-superior frontal regions at a time when the patient was still awake, evidenced by the presence of posterior alpha rhythm in simul-

E. Niedermeyer: A concept of consciousness

a a b c d e f g
Fig. 2 Typical adult EEG patterns from the waking state to profound sleep. a Waking-relaxed. Typical posterior alpha rhythm, briefly interrupted by eye opening and a period of heightened vigilance. b Early drowsiness: alpha dropout, mixed low voltage activity. c Deep drowsiness/earliest sleep with a vertex sharp wave and spindle rudiments. d Light NREM sleep, stage 2. Well developed spindles over superior frontal region. e Deep sleep, NREM stage 3. Enhanced slow activity, spindles and K complexes. f Very deep sleep, NREM stage 4. Prominent delta activity and barely a trace of spindles. g REM sleep. Mixed voltage activity and large ocular artifacts caused by typical eye-movements (REM)

taneous scalp recordings (Fig. 3) [18]. In the further course of becoming drowsy and falling asleep, frontal spindle activity in the depth increases until spindles can be recorded from the frontocentral scalp regions. One is hence tempted to theorize that the frontomotor cortex knows that sleep is impending so that behavioral-motor activities are bound to decrease. Isolated spindle trains in the ventrobasal complex of the human thalamus may occur prior to the appearance of scalp EEG spindles (Figs. 4, 5) [19, 20]. There is also a peripheral approach to the study of sleep initiation. The periocular area, where the first subjective signs of sleepiness are experienced, can be investigated with thermography [21, 22]. For strange reasons, virtually no attention has been paid to this approach. Thermography shows evidence of relatively hot periocular zones in (EEGmonitored) states of drowsiness and early sleep (Fig. 6). This raises the question of peripheral biochemical changes (possibly tissue hormones) acting in concert with central sleep-inducing regions while sleep is in the offing.

Fig. 3a, b Combined depth-scalp EEG. Age 42 years, intractable left temporal lobe epilepsy - now in presurgical evaluation. Location of depth electrodes: Left and right temporal tresses. 1, fusiform gyrus; 2 and 3, amygdala; 4, corona radiata; 5 and 6, supplementary motor cortex. Left and right frontal tresses: 1 and 2, medial fronto-orbital cortex; 3-5, corona radiata; 6, supplementary motor cortex. a Left portion. Awake with well developed posterior 8-9/sec alpha rhythm on the scalp while sleep spindles are noted in shallow frontal depth contacts (LF, RF 5 and 6). Middle and right portion show decline of alpha rhythm and some slowing on the scalp while deep spindles persist. b Intensified spindles in shallow frontal depth leads are associated with spindles on the scalp: there is now behavioral evidence of NREM sleep

Sleep and consciousness As pointed out previously, vigilance is markedly depressed in sleep but not abolished. Dream activity, which may also

be present in the earliest periods of stage 2 NREM sleep (and not just limited to REM sleep), tends to be associated with a plethora of POSTS (positive occipital sharp transients of sleep) in the EEG - perhaps as a sign of enhanced excitability within the central visual system [23]. Figures 7 and 8 schematically indicate the actions of the three main components of consciousness in sleep: vigilance strongly attenuated and mental contents pale and relegated into the world of dreams, while selective attention is reduced to virtual nonexistence.

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E. Niedermeyer: A concept of consciousness

Fig. 4 Age 38 years, chronic pain due to spinal arachnoiditis. EEG from scalp and 4 ventrobasal thalamic leads. Note independent sleep spindles on scalp and in thalamus

Fig. 7 The main components of consciousness while awake. Squares denote mental contents - in the strength of full vigilance. Selective attention is shown grabbing a set of mental contents

Fig. 5 Spindle appearance on scalp, in cortex and in thalamus

Fig. 8 The main components of consciousness in sleep. Mental contents appear to be pale. Selective attention is inactivated. Vigilance is reduced but not abolished

Figure 9 shows an assortment of 10 different EEG patterns which can be encountered in comatose states. This number could be further increased. This variety of coma patterns is strikingly different from the orderly parade of EEG patterns found in the progression of sleep stages. Such a wide variety of EEG patterns is also found in deepening general anesthesia depending strongly on the type of anesthetic utilized. Thus, coma due to central nervous system (CNS) disorders and general anesthesia are dominated by (a) the type and locus of pathology, and (b) type of anesthetic and its mode of CNS action.
Fig. 6 Thermography: note temperature increase in periocular region in a state of drowsiness/sleepiness

Epileptic states and consciousness Effects of spikes and spike-waves on vigilance

Coma There is no orderly sequence of EEG patterns on the journey from lethargy to semicoma, coma and profound coma. It is common knowledge that some epileptic manifestations alter consciousness while others apparently leave this function intact. There is some controversy about the action of

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ated with immediate profound coma. Here the question arises: is grand mal an all-out standard reaction of the brain or are there certain nuances of variable coma depth? There has been some credible support for the latter possibility [27-29].

Petit mal absence and working memory The alteration of consciousness during the classic petit mal absence with generalized 3-second spike-waves shows unusual and even unique features. In every form of impairment of consciousness, the brain requires some recovery time before the pre-disturbance state of vigilance is reached. Even mild fainting spells give testimony to the gradual recovery of consciousness. This is not true for the petit mal absence during which, for instance, a child in school immediately stops reading a text but, precisely at the end of the seizure, is capable of continuing exactly at the point where the reading had stopped. This unique situation has been interpreted as brief suspension of the working memory [30]. According to Fuster [31, 32], the working memory (a term taken from the work of Baddeley [33] but vastly clarified by Fuster) should be perceived as a system which serves as a constant perceptionaction cycle and enables a person to check every bit of planned action against past experience. This system is of such crucial importance for mental functioning that its suspension would block intellectual activity. The frontal lobe and, in particular, its lateral-dorsal region, is considered the main substratum of the working memory [31, 32]. When one considers the superior frontal maximum of 3-sec spike-waves in absences [34-36], then one can easily understand the action of these discharges on the working memory. Only a suspended function - but not an impaired one is ready to bounce back immediately to full effectiveness with the termination of the seizure.

Fig. 9 Variety of EEG patterns in coma: ten different EEG patterns in comatose states (two adjacent channels for each pattern). Left Top: diffuse irregular slowing. Second from top: slowing with triphasic waves (note sharp component). Third from top: diffuse slowing with spindle trains (spindle coma). Fourth from top: intermittent rhythmical delta waves. Bottom: rhythmical delta waves precipitated by arousing stimulus (paradoxal delta activity). Right Top: burst-suppression activity (with pronounced spikes). Second from top: alpha coma (frontocentral rather than posterior alpha maximum). Third from top: mixed activity with declining voltage output. Fourth from top: recurrent intermittent spike discharges. Bottom: absence of EEG response to nociceptive stimuli. (With kind permission of S. Karger Publishers, Basel, Switzerland)

subclinical paroxysmal EEG patterns on consciousness. As far as subclinical, generalized 3-sec spike-wave complexes are concerned, pertinent studies can be traced back to the work of Schwab [24]. Certain momentary depressions of attention have been found to occur during the spike component but also during the slow wave component of the spikewave complex. Even ordinary single spikes have been reported capable of altering cerebral functioning in an extremely brief manner [25, 26]. Further work will be needed in order to substantiate such claims.

Complex partial seizures It is interesting to note that complex partial seizures may mildly tinge the level of vigilance by generating a dreamy state. In such states, the relationship of past and present becomes befuddled, resulting in the well known dj vu or similar type of attack. Such experienced seizures usually indicate a milder form of temporal lobe epilepsy (with fairly good response to appropriate medication). On the other hand, automatisms with more profound impairment of consciousness are usually found in more severe forms of epilepsy.

Neuroscientific approach to consciousness Vigilance The explorability of the neuroscientific underpinnings of vigilance and the wealth of studies in this field have been mentioned earlier.

Mental contents Electrophysiological investigations of the flow of mental contents require great sophistication of design and technology. The work of Gevins and Bressler [37] should be mentioned in this context. Lehmann and his coworkers [38] have been using a fascinating paradigm in their exploration of 13 young volunteers

Grand mal Generalized tonic-clonic seizures, also known as grand mal, are major convulsions which, by definition, are associ-

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E. Niedermeyer: A concept of consciousness

resulting in the analysis of event-related and spontaneous microstates. In a subsequent study, Lehmann and Koenig [39] have performed EEG studies with regard to spontaneous thoughts of (a) abstract and (b) visual imagery type. These authors feel that alpha rhythm consists of microstates as building blocks.

Selective attention Evoked potential studies have been used for the investigation of selective attention. A paradigm of Hillyard et al. [40] was employed for the demonstration of the effect of selective auditory attention on the N 100 complex (detection of a target tone slightly higher in pitch than the standard tones). Differences were found in the N 100 responses to target and nontarget tones [41, 42]. Heinze et al. [43] used event-related potentials and positron emission tomography (PET) in the study of selective attention to visual stimuli. There was evidence of special involvement of the fusiform gyrus in this process - confirmed with functional magnetic resonance imaging (MRI) and eventrelated potentials [44]. Studies of selective attention to somatosensory stimuli have shown an enhanced N 140 response with a maximum over the prefrontal cortex [45] along with an early (30 ms) response over the contralateral parietal lobe [45]. Earlier animal experiments in monkeys by Mountcastle and his coworkers [46] showed enhanced neuronal activity in the parietal region with selective visual attention. With the use of auditory evoked potential studies in humans, Knight and coworkers [47] were able to demonstrate the eminent role of the dorsolateral prefrontal cortex in attention control. It was found that damage to this area leaves the patients stimulus bound and severely impaired in their ability to interact with the environment.

switching to Aristotelianism under the strong influence of Thomas Aquinas (1225-1274). Throughout these centuries, the gigantic comprehensive work of these thinkers did not include any reflections on consciousness. This changed greatly when the emphasis on metaphysics (and especially on the notion of being) started to vanish with the work of Descartes (1596-1650). With this development, the term consciousness steadily gained in import-ance reaching a pinnacle in the nineteenth century with the work of Franz von Brentano (1838-1917) and William James (1842-1910). This spectacular advance of consciousness-oriented psychology, however, started to decline after 1900 under the impact of the psychoanalytic theories of Sigmund Freud (1856-1939) and the objective psychology of behaviorism (J.B. Watson, 1878-1958; B.F. Skinner, 1904-1990). Both the Freudian emphasis on subconscious mechanisms and the behavioristic obsession with objective psychology thwarted consciousness-oriented investigation. These shackles have been shaken off and a veritable cornucopia of new books dealing with consciousness has followed the demise of the oppressive ideologies.

The new wave of consciousness research and the lack of a guiding concept There is no doubt that guiding principles are badly needed. It is, however, quite interesting to note that the biomedical sciences have developed a strong aversion to concepts while most of the research emphasis is being placed on the acquisition of data. This development has been recognized and castigated as an impediment on the road to true progress [48, 49]. It has been pointed out that physics and chemistry have stressed their dependence on valuable concepts, theories or hypotheses [48]. In strange contrast to the aforementioned aversion to concepts in the biomedical domain, there has been a recent surge of mathematico-biophysical approaches to the neurosciences along with the development of models of brain function [50]. This tendency is bound to lead to lifeless and anemic views of the brain which, in reality, is a living organ of wonderful complexity and not just a system of functional models.

For a better understanding of the simplified concept of consciousness What historical development of human thought has turned consciousness into an undefinable term? During the classic Greek period (500 to 300 BC) when philosophy reached an all-time peak due to the unique talents of Greek thinkers in their search for philosophical truths, no attention was paid to consciousness (for which no word existed). This is true for the greatest of the Greek philosophers, Plato (428-348 BC) and Aristotle (384-322 BC) - considered the two great pillars of Western thought and culture (we have to leave aside the Eastern philosophies and religions in which consciousness may figure as a divine force and with a totally different meaning). The further development of philosophy remained based upon Platonic thought (Plotinus, about 205250 AD, and Augustin of Hippo, 354-430 AD), much later

Components of consciousness at work An earthy analogy, i.e. an example taken from American daily life, may help explain the working of vigilance, mental contents and selective attention. Figure 10 shows a Baltimorean reading The Sun newspaper. A schematic view of his vigilance, mental contents and selective attention is presented. A band of small multicolored circles indicates his flow of mental contents which remain mixed and unfocused. Brief actions of selective attention (thin vertical lines arising from the horizontal curve of vigilance) are very weak and amount to nothing. This situation changes when he reaches the sports pages where

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he finds the news of an impending baseball strike. The level of vigilance quickly rises (this is associated with a general sympathomimetic response - not indicated in this schematic view), the mental contents become nearly uniform and a powerful thrust of selective attention forms a core of uniform contents. This vigorous action of selective attention results in a new phenomenon which transcends the scope of consciousness: awareness of an imminent baseball strike. Note again: awareness, unlike consciousness, is necessarily tied to the genitive of! Out of this secondary phenomenon of awareness, a number of tertiary phenomena emerge, especially memories and emotions. According to the presently used criteria of definition, all of these phenomena would be regarded as manifestations of consciousness - with additional involvement of further neocortical and, in particular, limbic portions. To end our little story: we can see that this reader is not a sports fanatic; his mental contents become mixed again and the level of vigilance gradually approaches the baseline.

Fig. 11 The activities of life - along with emotional events - seen under the all-encompassing mantle of consciousness

Why was consciousness played down in old age and then overblown so that a reductionistic concept has become necessary? A pictorial illustration will help explain the philosophical background for the paunchy and the lean concept of consciousness. Figure 11 presents a view of daily life with its activities and their emotional overtones. Whatever this picture exhibits can be placed under the huge heading of consciousness. This is the view of the undefinable, all-encompassing consciousness as propagated in the nineteenth century and still felt to be valid in our days. We see the same activities of life pictorially demonstrated in Fig. 12 but, in this view, all actions are placed on a huge, double-layered podium consisting of the basic layer of life and, above that, a smaller layer of consciousness. This is the lean consciousness in the view of the great old philosophers who

conceived life and consciousness as the unalienable elements of being in an ontologically oriented philosophy which gradually vanished in the course of the past four centuries. In other words: the actions of life occur on the podium of being consisting of life and consciousness. Why bother about consciousness - it is a part of being! Life and Consciousness - the carriers of all activities are, of course, scientifically explorable: life being the topic of biology and consciousness, reduced to its three main components, the topic of cognitive neurosciences. Without such a reductionistic concept, investigators of consciousness are likely to move in circles like antarctic explorers without a compass in the whiteout of a blizzard. It is no disgrace to borrow ideas and principles from the great thinkers of the past so that modern neurosciences can proceed smoothly on the road to true progress.

The bottomline: the definition of consciousness Consciousness is the foundation of mental (neurocognitive) activities and consists of three main components: vigilance, mental contents and selective attention.

Fig. 10 A Baltimorean reads The Sun newspaper: his level of vigilance, mental contents and selective attention are schematically demonstrated. For further details, see text

Fig. 12 The activities of life - along with emotional events - conceived as occurring on a double-layered podium consisting of (a) life, and (b) consciousness: both forming the basic condition of being

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E. Niedermeyer: A concept of consciousness ascending serotonergic systems. In: Zschocke S, Speckmann E-J (eds) Basic mechanisms of the EEG. Birkhuser, Boston, pp 231-268 Dement W, Kleitman N (1957) Cyclic variations in EEG during sleep and their relation to eye movements, body motility and dreaming. Electroencephalogr Clin Neurophysiol 9:673-690 Jouvet M (1969) Biogenic amines and the states of sleep. Science 163:32-41 Caderas M, Niedermeyer E, Uematsu S, Long DM, Nastalski J (1982) Sleep spindles recorded from cerebral structures in man. Clin Electroencephalogr 13:216-225 Gcer G, Niedermeyer E, Long DM (1978) Thalamic EEG recordings in patients with chronic pain. J Neurol 219:47-61 Jankel WR, Niedermeyer E (1985) Sleep spindles. J Clin Neurophysiol 2:1-35 Niedermeyer E, Jankel WR (1984) Falling asleep: depth EEG and thermographic observations. Electroencephalogr Clin Neurophysiol 58:8P (abstract) Niedermeyer E, Jankel WR, Uematsu S (1986) Falling asleep: observations and thoughts Am J EEG Technol 26:165-175 Niedermeyer E, Lentz WJ (1976) Dreaming in non-REM sleep. Waking Sleeping l:49-51 Schwab RS (1939) Methods of measuring consciousness in attacks of petit mal. Arch Neurol Psychiatr 41:215-217 Hughes JR (1997) The significance of the interictal spike discharge: a review. Clin Electroencephalogr 28:60 (abstr) Niedermeyer E (1990) Clinical relevance of EEG signals in epilepsies. In: Wada JA, Ellingson RJ (eds) Clinical neurophysiology of epilepsy. Elsevier, Amsterdam, pp 237-261 (EEG handbook, revised series, vol. 4) Schmidt RP, Wilder BJ (1968) Epilepsy. Davis, Philadelphia Rodin EA, Gonzales S, Caldwell D, Laginess D (1966) Photic evoked responses during induced epileptic seizures. Epilepsia 7:202-214 Petsche H (1958) Epileptischer Anfall und kortikale Neuronenpopulation. Wien Z Nervenheilk 24:46-55 Niedermeyer E (1998) Frontal lobe functions and dysfunctions. Clin Electroencephalogr April 29:79-90 Fuster JM (1995) Memory and planning. In: Jasper HH, Riggio S, Goldman-Rakic PS (eds) Epilepsy and the functional anatomy of the frontal Lobe. Raven, New York, pp 9-19 Fuster JM (1995) Memory in the cerebral cortex. MIT, Cambridge Baddeley AD (1986) Working memory. Oxford University, Oxford Niedermeyer E (1972) The generalized epilepsies. Thomas, Springfield Niedermeyer E (1996) Primary (idiopathic) generalized epilepsy and underlying mechanisms. Clin Electroencephalogr 27:1-21 Rodin EA, Ancheta O (1987) Cerebral electrical fields during petit mal absences. Electroencephalogr Clin Neurophysiol 66:457-466 Gevins AS, Bressler SL (1988) Functional topography of the human brain. In: Pfurtscheller G, Lopes da Silva FH (eds) Functional brain imaging. Huber, Toronto, pp 99-116 Lehmann D, Kochi K, Koenig T, Koukkou M, Michel CM, Strik WK (1995) Microstates of the brain electric field and momentary mind states. In: Eiselt M, Zwiener U, Witte H (eds) Quantitative and topological EEG and MEG analysis. Universittsverlag, Jena, pp 139-146

Sommario Il termine coscienza, correttamente usato in clinica, indica una realt estremamente complessa e praticamente indefinibile. La coscienza stata ignorata dai grandi filosofi classici e considerata come inerente all essere. La situazione e cambiata con Descartes e nei secoli successivi. Viene proposto un concetto riduzionistico e di facile definizione della coscienza, secondo cui la coscienza consiste di tre componenti fondamentali: vigilanza, contenuti mentali e attenzione selettiva. Queste componenti possono essere studiate con le moderne metologie delle neuroscienze e la vigilanza risulta essere la funzione pi facilmente esplorabile. Vengono puntualizzate le importanti differenze tra sonno e coma (unitamente ad alcune osservazioni sulla fase di induzione del sonno). Speciale attenzione viene posta ai disturbi di coscienza associati alle crisi epilettiche e, in particolare, allassenza con punte-onda che riferita ad una temporanea sospensione dellattivit dei circuiti del lobo frontale che sottendono la memoria di lavoro.

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