3 Copyright 2002, Elsevier Science (USA). All rights reserved.The Cognitive Electrophysiology of Mind and Brain

C H A P T E R

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Cognitive Electrophysiology ofMind and Brain

Alberto Zani and Alice Mado Proverbio

INTRODUCTION

The event-related potentials (ERPs) ofthe brain are wave forms reflecting brainvoltage fluctuations in time. These waveforms consist of a series of positive andnegative voltage deflections relative tosome base line activity prior to the onset ofthe event. Under different conditions,changes may be observed in the morphol-ogy of the wave forms (e.g., the presence orabsence of certain peaks), the latency, dura-tion, or amplitude (size) of one or more ofthe peaks, or their distribution over thescalp. ERPs are useful measures for study-ing mind and brain functions because theyare continuous, multidimensional signals.Specifically, ERPs give a direct estimate ofwhat a significant part of the brain is doingjust before, during, and after an event ofinterest, even if this is prolonged. ERPs canindicate not only that two conditions aredifferent, but also whether, for example,there is a quantitative change in the timingand/or intensity of a process or a qualita-tive change as reflected by a different mor-phology or scalp distribution of the waveforms. For all these reasons, ERPs are wellestablished as powerful tools for studyingphysiological and cognitive functions ofthe brain.

ERPs AND COGNITIVE THEORY

The so-called cognitive revolution(Baars, 1986) that has permeated researchon the mind in psychology and the neuro-sciences has led to widespread recognitionthat cognition and the knowledge thatderives from it, rather than being anaccumulation of sensory experiences, is aconstructive process that requires theverification of hypotheses influenced byprevious knowledge, past experience, andcurrent aims, as well as emotional andmotivational states. Cognitive theory lednot only to the rejection of the mind–braindualism (Mecacci and Zani, 1982; Finger,1994), but also to firm establishment of thenotion that the nature of the mind is deter-mined to a large extent by the neuro-functional architecture of the brain. Animportant corollary of this concept is theidea that in order to understand the mindit is essential to study and understand thebrain (Gazzaniga, 1984, 1995; Posner andDiGirolamo, 2000).

Understanding the mind and brain does not in any way mean understandingconscious processes—quite the contrary,because to a large extent it means investi-gating nonconscious neural processes. Thisfact suggested to researchers of the stature

of Le Doux (1996) that the unconscious isreal, and the renown Gazzaniga (1998)stated that “many experiments highlighthow the brain acts earlier than we realize.”This occurs at different hierarchical levelswithin the complex entity of themind–brain, ranging from intra- and inter-cellular ion exchanges at the microcellularlevel to the flow of information, at themacrosystem level, along the differentfunctional circuits underlying the veryfunction of the brain and the mind. On theother hand, at a macrosystem level, uncon-scious function is manifested throughoutalmost all spheres of the mind, startingfrom the basic operations of analyzingphysical characteristics of stimuli by oursensory system, to recording past events ormaking decisions.

We do not believe that this is surprisingif we consider results of modern researchon the brain; contemporary studies demon-strate the existence of processes of uncon-scious or subliminal knowledge andperception that influence a manifestedbehavior, or the capacity of the brain to“filter” or suppress the processing ofstimuli (this argument is dealt with morefully in Section III of this volume, onprocesses of attention). This capacity tofilter, studied by Freud, who used the term“repression” to describe it, allows us to beconcious of specific thoughts and percep-tions, but not others, apparently under freewill and by choice.

In consideration of the relevance of thissubstantial unconscious component of themind and, indeed above all, the emotions,it can only be concluded that a heuristi-cally valid cognitive theory of the mind isone that considers the mind’s rational andcognitive aspects, which are maintained bythe activity of the neocortex, inseparablefrom the emotive and irrational aspects,expressed by the amygdala and the limbicanterior cingulate cortex (Bush et al., 2000;see also Chapter 8, this volume). Thisconclusion is also supported by the closerelationship existing between thought pro-

cesses and emotional processes, suggestedby authoritative researchers of the brainsuch as Le Doux (1996) and Damasio(1994). According to this logic, the brain isseen as a so-called living system. A system,despite being the sum of various parts,each with its specific function, acts as awhole in which each function inevitablyinfluences the other.

Furthermore, it must be rememberedthat whatever conception of the mind isadopted, it is not heuristically correct toconsider this latter as an immutable entity.In fact, the mind must be considered indynamic terms, that is, as undergoing con-tinuous variations on the basis of evolutiveprocesses and experience (Berlucchi andAglioti, 1997). It is essential to rememberthat the functional processes that distin-guish the mind vary as a function of theontogenetic development of the individual—depending, as a consequence, on thediversified maturation of cerebral structure—and as a function of the individual’slearning processes and specific experiencegained for the stage of developmentreached (Nelson and Luciana, 2001; seealso Chapter 9, this volume).

Cognitive electrophysiology is a verywell-established field of science (Heinze etal., 1994; Kutas and Dale, 1997). The newtechnologies used to pursue the investiga-tion of mind and brain, with the theoreticalbacking of the cognitive sciences, havedeveloped at a dizzying speed over therecent “decade of the brain.” As a researchtool, cognitive electrophysiology mayprovide relevant contributions to both cog-nitive and brain sciences, putting togethernew knowledge about humans as inte-grated sociobiological individuals. Thisambitious task implies an integration ofneurofunctional concepts and basic ormore complex cognitive concepts, such asthose proposed in cognitive sciences(Wilson and Keil, 1999). Unlike most elec-trophysiological research, mired down bydata collection and “correlation state-ments,” to the detriment of theorization,

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The relationships between these “tools”and cognitive processing are deduced bymeans of an “assumed” criterion thatlocates these physiological responses inaccordance with hypothesized constraintsabout their position and function withinongoing activity. These constraints aremediated by well-defined theories ofhuman cognition and information process-ing (Donchin, 1982, 1984a).

In seeking to clarify these proceduralsteps, let us take a concept such as learn-ing, viewed from the psychological orbehavioral level, and let us try to showhow this concept may fruitfully drive elec-trophysiological experiments. Both at thelevels of cognition and brain neurofunctionthree different, major principles of learninghave been coherently identified: (1) know-ing what is out in the world, to be used inlater recognition and recall (2) knowingwhat goes with, or follows, what, and (3) knowing how to respond or what to do,given the drive and the situation. Theinterweaving of these three kinds ofknowledge is manifested as complex vol-untary action and skilled performance. Inmany respects these principles underliethe acquisition and deployment of proce-dures that manipulate the knowledgestructures (Bransford et al., 1999). A veryrelevant topic relative to these proceduresis the distinction between so-called con-trolled and automatic procedures. Skilllearning is thought to be characterized bya slow transition from dominance by con-trolled processes to dominance by auto-matic processes. However, this transitionhas been shown to take place only fortasks for which consistent—i.e., repetitiveand predictable—information is available.Taking this theoretical framework as astarting point for psychophysiologicalresearch on learning, it may be predictedthat any spatiotemporal changes in ERPcomponents (amplitude and latency) thatmay occur with learning should only beobserved in tasks providing such consis-tent information (Kramer and Strayer,

the main assumption of cognitively ori-ented electrophysiological research is thatcognition is implemented in the brainthrough physiological changes. An implicitcorollary of this assumption is that electro-physiological measures, i.e., ERP compo-nents, may be taken as manifestations, andnot simply as correlates, of these interven-ing processes of the flow of informationprocessing (McCarthy and Donchin, 1979).

Indeed, arguments may be, and indeedoften are, raised against this theoreticalview in the name of “physiological objec-tivity.” However, we are aware that thesestatements arise from questionable adher-ence, in many cases without any aware-ness, to operationa1 meaning theory. Theprocedure of giving meaning to conceptsinductively on the basis of measures pro-vides an outmoded brand of operationismthat may have functioned well for theor-etically developed sciences, such as phy-sics, proceeding in the framework of thePopperian view of scientific progress, but which has been only detrimental toatheoretical electrophysiological research.Indeed, the difficulties often met indefining any intrinsic and immutable prop-erty of a physiological response, changingas a function of the conditions of its occur-rence, make the latter loosely defined inconceptual terms. This failure to find aspecific response definition is a problem-atic criterion for delineating a psychologi-cal process for the correlational approach.

To cope with the spatiotemporal overlapin scalp-recorded manifestations of under-lying cerebral processes, and with theproblems in determining their physiologi-cal generators, cognitive electrophysiolo-gists identify ERP components, i.e., thecerebral responses, as the portions of arecorded wave form that can be inde-pendently changed by experimental variables—task condition, state, subjectstrategy, etc. ERP components are notviewed as “structural markers” per se, butas “psychological tools,” as is any otherpsychological measure, e.g., reaction times.

1988; Sirevaag et al., 1989). In the pastdecades, evidence strongly supporting thisprediction has been accumulated in ERPliterature relative to all the best knowncomponents, especially the contigent nega-tive variation (CNV) and the so-called latepositive complex (LPC), i.e., N2, P300, andslow wave (see, Proulx and Picton, 1980;Kramer et al., 1986). Thanks to its inconsist-ent features, the “oddball” task is a simplebut flexible experimental task that hashelped to provide evidence, either as suchor in the context of a probe-based dual-taskparadigm, for the limited capacity of con-trolled processes and the spatiotemporalstability of ERP components.

ERPs AND THE BRAIN

Traditionally, for more than 100 yearscognitive and neurophysiological pro-cesses in humans have been studied bypsychophysical and behavioral methods.Modern neurosciences offer several hemo-dynamic, anatomofunctional, and elec-trophysiological methods to furtherinvestigations of the mind and brain.Nevertheless, only noninvasive whole-system procedures can be used to examinehumans (see Appendix A, this volume, fora synopsis of molecular and systemicresearch methods). Because neurophysio-logical processing takes place in fractionsof a second, one of the most feasible toolsis to record brain electrofunctional activity(see, e.g., Heinze et al., 1994; Rugg andColes, 1995). The advantages of electro-physiological signals, or ERPs, lie in theirvery high time resolution—in the order ofmilliseconds—and their reliable sensitivityin detecting functional changes of brainactivity. The high temporal resolution andnoninvasiveness of this method privilegeits use over brain imaging techniques suchas computed tomography (CT), positronemission tomography (PET), or functionalmagnetic resonance imaging (fMRI), aswell as over the behavioral measures most

used in traditional neuropsychologicalstudies. Thanks to these advantages, event-related brain potentials may reveal steps insensory–cognitive information processingoccurring very rapidly within the brain.Furthermore, unlike behavioral and neu-roimaging techniques, ERPs may revealdetails of functional organization, andtiming of the activation, of regional areasof anatomically distributed functionalsystems of the brain involved in cognitiveskills as well as in executive capacities.

Volume conduction and lack of three-dimensional reality do, however, mean thatthese brain signals are of more limited usethan neuroimaging techniques for examin-ing where in the brain processes take place.Nevertheless, localization processes carriedout using these signals may be made more sound through source-modelingalgorithms.

There is no doubt that modern neuro-imaging techniques have dramaticallyincreased our knowledge of the brain andthe mind (Posner and Raichle, 1994; Rugg,1998; Cabeza and Kingstone, 2001). Aswith ERPs, studies carried out with thesetechniques focus on an individual’s brainwhen it is involved in carrying out a partic-ular mental task: memorizing a list ofwords, distinguishing some objects fromothers that are similar but not the same,directing attention toward objects pre-sented in a particular part of the visualfield, etc. The theory underlying all thesestudies is that the areas of the brain that arefound to be most active during the tasksare those that are crucial for the varioustypes of mental activity.

However, simple mapping of the sites ofmental processes can indicate only wherein the brain a given functional activationtakes place but, at present, can in no wayexplain the mechanisms of the mind. Howdo we recognize objects and faces, how dowe recall the memory of experiences andthings, how do we direct our attention to objects and the surrounding space, etc.? These complex and extraordinary

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mental mechanisms still remain unchartedterritory.

No spatial or temporal resolution,however good, can localize something ofwhich we have only superficial know-ledge. In fact, in order to be able to “local-ize” a given cognitive state or mentalprocess (for example, “remembering some-thing”) in the brain we must know clearlywhat the state or process is and what thefunctional subprocesses are that invariablylead to one cognitive state and not another.If we do not know what these subprocessesare, or whether they vary in different con-ditions, we cannot reliably localize them inthe brain.

It is not at all difficult to find examplesin the literature to illustrate what we mean.The reader is referred to Chapter 7 in thisvolume for an impressive review showinghow different the cerebral localizations ofactivity can be during episodic mnemonicanalysis of figurative and linguistic infor-mation, according to the different type/state of analysis carried out by subjects(e.g., familiarity, encoding effort, recog-nition). Furthermore, there is no lack ofexamples of different localizations in dif-ferent studies by different scientists for asimilar form of mental activity, such asspatial attention. For example, Mangunand colleagues repeatedly reported activa-tion of the fusiform gyrus with PETimaging, and of this same gyrus togetherwith the medial occipital gyrus, whenimaged with fMRI, during attention to arelevant space location (see the exhaustivereview by Mangun in Chapter 10), whereasCorbetta and colleagues (see Corbetta,1998) reported a localized activity in theparietal lobe, a region of the cortex classi-cally associated with the control of spatialattention, in addition to the more dorsolat-eral occipital regions. It is not inconceiv-able that to cope with differences in thespatial tasks across these studies, differentcognitive processes, and thus, differentregions of the volunteers’ brains, musthave been activated during what was

reported by the authors as apparently thesame mental activity.

The difficulty in differentiating cog-nition from brain localization is not, however, unique to neuroimaging andelectrophysiological studies. Unfortun-ately, it is also difficult in most traditionalclinical neuropsychological research. Con-sider, for instance, research on hemineglector cortical blindness, or any other clinicalsyndrome. Although robust, direct post-mortem and neuroimaging evidence isavailable for the anatomical localization ofbrain lesions from which these syndromesderive, only controversial theories can beadvanced to explain which processes arelacking, compared to normal cognition, inthese patients’ cognitive processing andthus to explain their symptomatology.Examples of opposing theories can befound in Köhler and Moscovitch’s (1997)outstanding review on unconscious visualprocessing.

To complicate the picture further, local-ization research is often pushed to anextreme, frequently without being soundlybased on the theory of the mind or thefunctional architecture of the brain. Thereare now many authoritative investigatorsspeaking out against this approach toresearch, and it will probably emerge asmore of a hindrance than a help for under-standing the mind and brain. For example,according to Frith and Friston (1997), mostneuroimaging studies concentrate exclu-sively on subtraction techniques and onfunctional segregation to associate a givenarea with a given function. However,according to Frith and Friston, in order tobuild an accurate map of the mind, it iscrucial to understand the functional inter-connectivity of the centers and pathwaysof the brain by investigating the correla-tions between these different anatomo-functional entities.

This problem is felt, shared, and cre-atively developed in the excellent reviewby Cabeza and Nyberg in Chapter 3. Notby chance did they give their chapter the

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title “Seeing the Forest through the Trees:The Cross-Function Approach to ImagingCognition”; they identify “the trees” as thesingle cognitive functions on which manyimaging studies focus their concern, withloss of sight the whole—“the forest”––rep-resented by the fact that, on the one hand,many brain areas are involved in manycognitive functions, and, on the other, thatcognition is not actually subdivided intodistinct modular cognitive processes, asartificially proposed in cognitive sciencetextbooks for explanatory purposes.

Fuster (2000) is of the same opinion, andauthoritatively reports that “commonsense, psychophysics, and experimentalpsychology provide ample evidence thatall cognitive functions are interdependent.…Also interdependent must be, of course,their neural foundations.” And, cautioningthe reader about some of the problemswith the neuromodular principle of cog-nition, Fuster advances the concept of a“distributed cortical network” according towhich performance in cognitive tasks, or,more specifically, tasks of executive con-trol functions, is not solely mediated vialocalized areas of the brain, but by manyregional brain areas that are dispersedthroughout the brain, although beingstrictly linked to each other, and activatedin a divergent and convergent way at dif-ferent times. Again in Fuster’s (2000)words, “practically any cortical neuron orneuronal assembly, or module, can be partof many networks. A network can serveseveral cognitive functions, which consistsof neuronal interactions within andbetween cortical networks.” The closeresemblance between this carefully wordedand articulate definition and the nowadaysforgotten “functional system” theory ofbrain neurofunctional architecture, firstadvanced by the great Russian neuro-psychologist Alexander Lurija (Lurija,1962, 1976), will, we believe, have hardlyescaped anyone.

In the light of these considerations, webelieve that it is correct to think that the

moment has returned for researchers todedicate more of their forces to studyingthe mechanisms inherent to human cogni-tion in order to reach a fuller understand-ing not merely of the brain, of which in abroad sense we know quite a lot, but ratherthe mind, that is, its higher and morearcane product, of which we are still pro-foundly ignorant.

For decades, aware of the limited capac-ity of ERPs to localize intracerebralprocesses of cognition, cognitive electro-physiologists have continued their researchin the firm belief that the brain’s electro-magnetic signals spread over the scalpduring electrofunctional activation are pre-cious for understanding the ways withwhich the brain changes with experienceand knowledge. Furthermore, they haveshared the belief that the nature and mech-anisms of the neural processes of cognitiveand emotional reorganization are objec-tively and reliably codified by the differentcomponents of the ERPs and event-relatedfields (ERFs) (Donchin, 1979, 1984b;Hillyard and Picton, 1979; Zani, 1988;Hillyard, 1993; Näätänen and Ilmoniemi,1994; Rugg and Coles, 1995; Kutas andDale, 1997).

It was in this conceptual “framework”that the idea was advanced that ERPscould make an important contribution toour understanding of the cerebral mecha-nisms of knowledge (Kutas and Hillyard,1984; Heinze et al., 1994). And it is follow-ing this idea that the ERPs, rather thanbeing considered a now obsolete methodin comparison with the currently availabletechniques, are still used as a direct,quantifiable measure of processes ofknowledge, both conscious and uncon-scious, and as such are still used toproduce and validate models of the mindrather than to provide generic “correlates”of poorly defined psychological constructs.This will become extremely clear in the prestigious articles written byrenowned researchers collected together inthis book.

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In conclusion, it seems that using ERPs,in combination with other available tech-niques, as quantifiable measures of cog-nitive and affective processes of the brain,the cognitive electrophysiologist can helptest existing theories on the human mindand also can propose newer and moreheuristic ones. In order to be efficient inthis task of identifying mental processesarising from the brain, it is essential towork in the context of well-founded theor-ies and with sophisticated methodologycapable of distinguishing between thesetheories.

THIS BOOK—OVERVIEW

The chapters of this book—prepared bya panel of international neuroscientists and electrophysiologists—provide state-of-the-art reviews of the latest develop-ments in the study of the relationshipsbetween mind and brain as investigated byevent-related potentials and event-relatedfields. Some indications are explored ofhow these signals may be combined withthe high spatial resolution of the hemo-dynamic signals of the brain, such as thoseacquired through positron emission tomo-graphy and functional magnetic resonanceimaging, in order to come closer to the goalof localizing cognition within the brain.

The book is systematically organizedinto thematic sections. The three chaptersin the first section cover the theoretical andmethodological framework of investigat-ing the human mind through the recordingof electrical, magnetic, and hemodynamicsignals of the brain. In Chapter 1 we haveraised the point that the study of cognitioncan benefit enormously from the use of brain electrical and magnetic activity.Efforts are being made to demonstrate thatthese benefits will derive mostly fromtheoretically oriented electrophysiologicalresearch in the framework of cognitivesciences and neurosciences. Chapter 2focuses on the morphology of visual, audi-

tory, and somatosensory wave forms ofelectric potentials and magnetic fields ofthe brain, and also on the functionalsignificance of these electrophysiologicalindices in relation to the basic and higherdomains of cognition. Furthermore, theintracranial electro ionic origins of thesescalp-recorded physiological measures aredescribed, with indications for solving the“direct” and “inverse” problems of localiz-ing their electromagnetic dipoles withinthe brain.

Chapter 3 (Cabeza and Nyberg) offersan original theoretical cross-function frame-work for guiding hemodynamic functionalimaging of brain and cognition. Thisframework provides the foremost con-straints to functional interpretations, par-ticularly when assuming the so-calledsharing view, that is, the view that thesame brain region is recruited by differentcognitive functions. In the authors’ words,these constraints “help us overcome func-tion-chauvinism and see the ‘big picture.’In other words, cross-function compar-isons allow us to see the forest [what manyfunctional studies have in common]through the trees [the single cognitivedomains investigated by single studies].”

The second section (Chapters 4–9) sys-tematically covers electromagnetic researchon a representative sample of the neuraldomains of human cognition. Chapter 4(Skrandies) illustrates how the recordingof brain electrical activity in combinationwith knowledge on the human visualsystem may be employed to study visualinformation processing in healthy volun-teers as well as in patients with selectivevisual deficiencies. Data are presented ondifferent experimental questions related tohuman visual perception, including con-trast and stereoscopic vision as well asperceptual learning.

Chapter 5 (Aine and Stephen) dealswith magnetoencephalography (MEG)mapping of the ventral and dorsal streamsin human visual cortex. MEG cues, prov-ing that isoluminant, central field stimuli

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preferentially excite the ventral streamstructures and, alternatively, that periph-eral stimuli alternating at high rates prefer-entially activate the dorsal stream, aresystematically addressed. Furthermore, thefocus is on present progress in our under-standing of brain cortical areas involved inhigher visual processing, such as recogni-tion memory, as investigated by means ofMEG, and the future direction of MEGresearch in this field is discussed.

Chapter 6 (Federmeier, Kluender, andKutas) reviews ERP studies on language.Rather than simply presenting a collectionof various processes, throughout the chap-ter the authors illustrate the viewpoint thatthe goals of electrophysiological investiga-tions of language, as well as the goals ofresearch exploring language processingwith other tools, are to fashion an under-standing of how the various processesinvolved in language comprehension andproduction are coordinated to yield themessage-level apprehension we attain fromreading or listening to speech. As stated inChapter 6, “linguists, psycholinguists, andneurolinguists alike strive to understandhow the brain ‘sees’ language––because, inturn, language is such an important facetof how humans ‘see’ their world.”

Chapter 7 (Wilding and Sharpe) isdevoted to memory processes that con-stitute another very important domain ofhuman cognition. Indeed, memory hasbeen a subject of fascination to psycholo-gists and other brain scientists for over acentury. Recently, the study of the role ofdifferent brain areas in memory hasreceived a boost from new techniques andchanging pretheoretical orientations. Theauthors offer an original review of the bulkof electrophysiological studies on retrievaland encoding processes underlying epi-sodic memory. Commendably, they do notsimply share knowledge from ongoingresearch, but identify some acute out-standing problems in this field of inves-tigation, indicating its likely futuredevelopments.

Chapter 8 (Luu and Tucker) is intendedto close the misleading gap that exists inthe “cognitive” approach to brain functionand architecture between a pure cognitivefunctional processing of the brain and itsemotional counterpart, which is of suchimportance in producing thinking andbehavior, both in normal and emotionallydisordered people. With such a goal, theauthors deal specifically with mentalprocesses involved in emotion, motivation,and reward, reviewing studies in this field from a modern neuroscience-basedviewpoint.

Chapter 9 (Mitchell and Neville)addresses “neuroplasticity,” a dominantresearch theme in neuroscience at present.Neuroplasticity usually refers to somechange in the nervous system as a functionof age and/or experience. This contribu-tion reviews studies on the effects of ageand experience on the development of neu-rocognitive systems. A broad survey andsynthesis are provided of essential data onnormal brain and cognitive development,as well as on development after early deaf-ness, blindness, or following delays in lan-guage acquisition. The authors provideinsights into and ideas on the complexityand diversity of contemporary brain neu-roplasticity research in humans.

The three chapters gathered in the thirdsection are concerned with visual attention.Chapter 10 (Mangun) and Chapter 11 (DiRusso, Teder-Sälejärvi, and Hillyard)mostly address neural mechanisms ofspatial attention. Chapter 10 reviewsfindings indicating how human spatialattention involves top-down processes thatinfluence the gain of sensory transmissionearly in the visual cortex. Chapter 11 dealswith steady-state cortical processing of thebrain that reveals slow-rising changes incortical reactivity to the outer world. First,the authors provide an overview of thisprocessing mode in the visual modality.Then they present a review of experimentalfindings of modulations of this processingmode with selective attending of spatial

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and, to a lesser extent, nonspatial features(color, shape, etc.) of visual information, inline with the previous chapter. UnlikeChapters 10 and 11, Chapter 12 (Proverbioand Zani) concentrates on feature-based andobject-based selection mechanisms of thebrain as investigated with ERPs. An over-view is provided of studies showing theclose interconnections across the anteriorand posterior attention systems. In addition,a review is made of studies reporting the dif-ferential activation of the “Where” and“What” systems of the visual brain in con-ditions in which stimulus attributes have tobe separately and/or conjointly attended.Efforts are made to demonstrate the task-related relative segregation and complexinteractions of the aforementioned systemsduring the separate or conjoint processing ofstimulus attributes.

The final two chapters comprising thefourth section are concerned with clinicaland applied perspectives of ERP research.Chapter 13 (Verleger) provides an exhaus-tive overview of ERP studies on neuro-psychological syndromes. The detaileddescription given of these syndromes is sub-divided into three main categories. Theresult is a unique, up-to-date, and wide-ranging discussion of these disorders thatdraws on biology, genetics, neuropsychol-ogy, clinical presentation, and treatment.Chapter 14 (Näätänen, Brattico, and Tervan-iemi) introduces the mismatch negativity(MMN), a component of auditory ERPsreflecting the brain’s automatic response toany discriminable change in auditory stimu-lation. Because the MMN can be measuredeven in the absence of attention and withoutany task requirements, it is particularly suit-able for investigating several clinical popula-tions as well as infants. Moreover, the MMNprovides a unique index of the subject’saccuracy in the processing of speech andmusical sounds. It can be used, for example,to unravel the neural determinants of lan-guage skills and musical expertise.

In addition to the specialist review chap-ters, a fifth section of this book collects

together a number of appendixes contain-ing the primers of the theoretical andmethodological matters––including somesimple-level mathematical material—treated in the specialist chapters. Theseappendixes are intended for the benefit ofnonexperts (such as psychology andmedical students), as well as experts inother neighboring fields. These appendixeshave been included in order to clarify, insimple but detailed terms, the basics ofmolecular and systemic methods of inves-tigating the nervous system (Appendix A),as well as neuropsychological clinical prac-tice (Appendix B). They also provide thefundamentals of electromagnetic recordingand data analysis and laboratory setup(Appendixes C and D), and topographicand dipole mapping methods (Appendix E).Last but not least, the invasiveness and thespatial and temporal resolution of electro-magnetic techniques, as compared to othertechniques, are given (Appendix F).

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12 1. COGNITIVE ELECTROPHYSIOLOGY

I. A COGNITIVE FRAMEWORK