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Europe (approximately the fourth to fourteenthcenturies), the ventricles continued to be the focusof theories relating mind and brain.1 For example,according to fourth-century church fathers, theanterior ventricles were associated with percep-tion (later to be known as the sensorium com-mune), the middle with reason, and the posterior

with memory.2 It has been suggested that this fo-cus on the ventricles accorded better with the du-alism of Christian theology, as the hollow cavitiescould be said to contain the soul without hypothe-sizing an identity between mind and the physicalsubstrate of brain tissue.1 Figure 1-1 shows anearly illustration of the ventricular system.

During the Renaissance, the ventricular doc-

trine and the role of the rete mirabile began to losetheir influence on theories of mind-brain rela-tions.2,3,4 The seventeenth-century writings ofRene Descartes mark a transitional phase, inwhich the interaction between fluid in the ventri-cles and brain tissue itself was hypothesized to ex-plain intelligent action, as shown in Fig. 1-2. Forreflexive action, Descartes proposed a simpleloop, in which stimulated nerves caused the re-

lease of animal spirits in the ventricles, which, inturn, caused efferent nerves and muscles to act.For intelligent human action, this loop was modu-lated by the soul via its effects on the pineal gland.The pineal gland was chosen in part because it isunpaired and centrally located and also because itis surrounded by cerebrospinal fluid. It was alsomistakenly thought to be uniquely human. Of

course, the pineal gland was just the vehicle for

Chapter 1

A HISTORICAL

PERSPECTIVE ONCOGNITIVENEUROSCIENCE

Todd E. FeinbergMartha J. Farah

ANTIQUITY THROUGH THEEIGHTEENTH CENTURY

The most fundamental fact of cognitive neurosci-ence was established in ancient times, when theGreeks first determined that the brain was thephysical seat of the mind. Alcamaeon of Croton,

who may have been a student of Pythagoras, iscredited with making this basic advance in thefifth century B.C. on the basis of his observationsof human patients with brain damage. The alter-native hypothesis held the heart to be the organresponsible for sensation and thought. This wasthe accepted view among Egyptian writers andcontinued to attract adherents in ancient Greece

centuries after Alcamaeon. Although Hippocratesand Plato held a cerebrocentric view of the mind,no less a thinker than Aristotle remained in thecardiocentric camp.

Among the ancient cerebrocentrists, the na-ture of the mind-brain relation was poorly under-stood. The brain itself was considered by many tobe a mere package for the real substance ofthought, the cerebrospinal fluid, and the most im-

portant anatomical features of the brain weretherefore the ventricles. Although brain tissue it-self was considered important by some writers, in-cluding Galen, for many centuries the mind waspredominantly identified with cerebrospinal fluid.The present-day use of the word spirit to referboth to certain fluids and to the soul is vestige ofthis idea.

For the entire period of the middle ages in
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the minds influence on the body; Descartes the-ory still denied any form of identity between themind and neural tissue.3,5-8

Descartes theory was formulated at a timewhen neuroanatomic knowledge was quite primi-tive. This situation began to change with the workof such figures as Thomas Willis later in the seven-teenth century3,10 and Malpighi Pacchioni and Al-brecht von Haller in the eighteenth.11,12 For ex-ample, Von Haller stimulated the nerves of liveanimals in an effort to discover the pathways forperception and motor action, thus establishing the

experimental method in neurophysiology. Thiswork set the stage for the explosion of experimen-tal and clinical research of the nineteenth century,in which the brain organization underlying per-ception, action, language, and many other cogni-tive functions was revealed.

THE LOCALISM/HOLISM DEBATE OFTHE NINETEENTH CENTURY

One of the more notorious figures in the historyof behavioral neurology is Franz Josef Gall,shown in Fig. 1-3. In the late eighteenth and earlynineteenth centuries, he and his collaborator Jo-hann Spurzheim made a number of importantcontributions to functional neuroanatomy, includ-

ing proving by dissection the crossing of the pyra-mids and establishing the distinction between grayand white matter.1315 Gall is also credited withone of the earliest descriptions of aphasia linkedto a lesion of the frontal lobes.16 He is most fa-mous, however, for his general theory of cerebrallocalization, known today as phrenology. At theage of 9, Gall had noted that his schoolmates who

excelled at rote memory tasks had quite promi-nent eyes, les yeux a fleur de tete (cows eyes).He reasoned that this was the result of the overde-velopment of the subjacent regions of the brain,and speculated that these regions of the brainmight be particularly involved in language func-tions and especially verbal memory.

Gall identified 27 basic human faculties andassociated them with particular brain centers that

could affect the shape of the skull, as shown in Fig.

14. These included memory of things and facts,sense of spatial relations, vanity, God and reli-gion, and love for ones offspring.15 His theory wasbased on hundreds of skulls and casts of humansand beasts. For instance, the disposition to murder

and cruelty was based on a bump above the earpossessed by carnivorous animals. He located thesame feature in sadistic persons whom he had ex-amined personally,3 skulls of famous criminals,and the busts and paintings of famous murder-ers.12 Gall taught and practiced medicine in Vi-enna from 1781 to around 1802, until EmperorFrancis I banned Galls public lectures becausethey were materialistic and thus opposed to mo-

rality and religion.12 Gall then took to the road,

4 PART I/ HISTORY AND METHODS

Figure 1-1

The ventricular system according to Albertus Magnusfrom his Philosophia naturalis (1506).2


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ety of animal species to demonstrate the equipo-tentiality of cortex.

Galls status as a popularizer, and Flourenssempirical attacks, helped to push localism outof the mainstream of contemporary scientificthought in the early nineteenth century. When, in1825, Jean-Baptiste Bouillaud, shown in Fig. 1-5,

presented a large series of clinical cases of loss ofspeech following frontal lesions,12,18,19 his work waslargely ignored. This landmark work, in whichspeech per se was distinguished from nonspeechmovements of the mouth and tongue, is still rela-tively unknown.

Bouillaud was not the only one to suggest afrontal location for language functions. Duringthis period and lasting up to the 1860s, numerous

clinical reports of patients with frontal lobe

lecturing across Europe to enthusiastic popularaudiences. By the time he settled in Paris in 1807,he was hugely popular and internationally known.However, phrenology continued to create contro-versy in scientific circles.

The best-known critic of Gall was Marie-Jean-Pierre Flourens. Flourens mounted a scien-

tific research program to disprove Galls theory,but it appears to have been motivated at least asmuch by religious discomfort with the implica-tions of Galls straightforward mind-brain equiva-lences as by scientific considerations. Flourensviewed Galls theory as tantamount to denying theexistence of the soul, because it divided the mindand brain into functionally distinct parts andFlourens believed the soul to be unitary.12,1719 He

carried out extensive lesion experiments on a vari-

5CHAPTER 1 / A HISTORICAL PERSPECTIVE ON COGNITIVE NEUROSCIENCE

Figure 1-2

Descartes conception of sen-

sation and action as con-ceived in hisDe homine

(1662). Light was transferred

from the retina to the ventri-

cles, causing the release of

animal spirits. The pineal

gland modulated this mecha-

nism for voluntary action.9


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damage and loss of speech were recorded in Eu-

rope and America. Indeed, this idea had consid-erable historical precedence throughout antiq-uity.2022 However, intense interest in localizationof brain functions, particularly language, was nowdeveloping. It was during this time that Marc Daxnoted the association between left-hemisphericdamage, right hemiplegia, and aphasia, basedupon his examination of 40 patients over a 20-year

period. This paper was handwritten in 1836 andnot published at the time,23 but copies may havebeen distributed to friends and colleagues.3

It was not until 1861 that the field reconsid-ered localism with a more open mind. That yearthe Societe dAnthropologie in Paris held a seriesof debates between Pierre Gratiolet, arguing in fa-vor of holism or equipotentiality, and Ernest Au-bertin, the son-in-law of Bouillaud, arguing in fa-

vor of localism.12,24,25 Aubertin, shown in Fig. 1-6,

reported his clinical observations of a patientwhose frontal bone was removed following a sui-cide attempt. He reported that when the blade ofa spatula was applied to the anterior lobes,there was complete cessation of speech withoutloss of consciousness.3,12,25 Aubertin went on to de-scribe a patient of Bouillauds who had a speechdisturbance and was near death. Aubertin boldlyvowed if this patient lacked a frontal lesion hewould renounce his views.12,2427

The 1861 debate is best known not for thepresentations of Gratiolet and Aubertin but for

the eventual participation of the societys founderand secretary, Paul Broca, shown in Fig. 1-7. Al-though Broca did not initially take a strong posi-tion, his observations of a patient then under hiscare led him to play a pivotal role in the debate.His patient, Leborgne, suffered from epilepsy,

6 PARTI/ HISTORY AND METHODS

Figure 1-3

Franz Josef Gall (17581828).

Figure 1-4An example of a porcelain phrenology bust with

demarcations that demonstrate the reflection of the

human faculties on the skull. (Photograph courtesy of

Joseph A. Hefta.)


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right hemiplegia, and loss of speech, the last for aperiod of over 20 years. Leborgne had been insti-tutionalized for some 31 years and throughout thehospital was known by the name Tan, as this

was his only utterance along with a few obsceni-ties.3,9 In light of Aubertins declaration, Broca in-vited him to examine Tan, which Aubertin didand afterward concluded that indeed the patientmet the critieria of his prior challenge. Six dayslater Leborgne died; the following day, April 18,1961, Broca presented the brain to the societyalong with a brief statement but without firm con-

clusions.25

Figure 1-8 shows the brain of Leborgne.Four months later, at a meeting of the So-ciete Anatomique de Paris in August, Broca madea more extensive report. The brain of Leborgnehad demonstrated an egg-sized fluid-filled cavitylocated in the posterior second and third frontalconvolutions, with involvement of adjacent struc-tures as well, including the corpus striatum.36 Inthis report, Broca claimed that his findings would

support the ideas of M. Bouillaud on the seat of


Figure 1-5Jean-Baptiste Bouillaud (17961881).

Figure 1-6

Ernest Aubertin (18251865).

Figure 1-7

Paul Broca (18241880).


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the faculty for language;

25,28

he later suggested apossible localization of speech functions to thesecond or more probably third frontal convolu-tion. Later the same year, Broca presented an-other patient with speech disturbance, an 84-year-old laborer whose lesion also involved the leftsecond and third frontal convolutions. The lesionwas more circumscribed than that found in Le-borgne and strengthened the association of those

structures with speech localization.In the mid-1860s the issue of hemispheric

asymmetry entered the debate on localization.The previous cases strongly suggested that speechis localized to the left hemisphere, and an addi-tional series of eight cases published by Broca in1863 were exclusively left-sided.3,23,29 In spite ofthe strong lateralization of lesion locus in these

cases, Broca made note of this remarkable ob-servation but made no further claims.23 In thissame year and shortly before Brocas paper waspresented,3 Gustave Dax, son of Marc Dax, sent ahandwritten copy of his fathers manuscript to theAcademie de Medecine in Paris. In this document,Marc Dax had previously described his view onthe relation between speech and the left hemi-sphere. The paper was read before the Academie

in December 1864 and published in 1865.30,31 By

1865, Broca clearly expressed the opinion that theleft hemisphere played a dominant role in speechproduction.32,33 As far as the issue of priority ofdiscovery is concerned (a matter of controversyamong historians), most writers agree that theDax paper in its original form in 1836 had no in-fluence on Broca or the scientific communitywhen first written. This paper did, however, makeclear the association of language functions and the

left hemisphere. While Broca alone clarified therole of the second and third frontal convolutions,he apparently did not take a firm position on thespecific role of the left hemisphere until after theDax paper was read before the Academie deMedecine in Paris in December 1964.3 It appearsthat the reemergence of the Dax manuscriptand Brocas discovery were nearly simultaneous

events.

THE AFTERMATH OF 1861:THE EMERGENCE OF MODERNNEUROPSYCHOLOGY

The events in Paris in the 1860s constituted a turn-ing point in the history of ideas regarding brain

function. The concepts and methods developed in

8 PART I / HISTORY AND METHODS

Figure 1-8

Photograph of the brain of

Brocas first patient, Leborgne

(Tan). It is now housed in

the Musee Dupuytren.


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aphasia he called conduction aphasia, in whichcomprehension would be preserved but outputwould be as impaired as in sensory aphasia.3538

Wernicke had, in effect, proposed a model thatcould explain a number of different aphasic syn-dromes by lesions to different combination of cen-ters and connections between centers. This type oftheorizing came to be known as associationism,

because language use was viewed in terms of asso-ciating representations in different brain centers,or as connectionism, because of the emphasisthat view put on the connections between centers,as shown in Fig. 1-10.

The connectionist paradigm was quickly ex-tended to explain other disorders. Ludwig Licht-heim placed pure word deafness in this frame-work, predicting the critical lesion site as well as

noting that, given the connectionist explanation

the course of debating the localization of speechwere extended to a variety of different higherfunctions, experimental work on animals also de-veloped apace. From this period onward, it is im-possible to trace a single line of scientific develop-ment. Here we simply present a summary of someof the major advances seen in the behavioral neu-rology and neuropsychology of the late nineteenthand early twentieth centuries.

In the decade following Brocas contribu-tions, two important developments took place inGermany. First, Edward Hitzig and Gustav

Fritsch performed a series of experiments inwhich the cortex of a dog was stimulated while thedog lay on a dressing table in Hitzigs Berlinhome.4,9,34,42 These experiments established thatmotor functions are localized to anterior cortexand demonstrated experimentally the somato-topic organization of motor cortex inferred indi-rectly from previous clinical-anatomic correla-

tions in humans. In their report, the investigatorsspecifically noted that their results refuted the ho-lism of Flourens. Following their work, Sir DavidFerrier in England confirmed the findings of Hit-zig and Fritsch and improved upon their methodof stimulation to discover more detailed structure-function relationships.11

About the same time, the German neurolo-gist Carl Wernicke began to investigate language

functions other than speech. Wernicke, shown inFig. 1-9, documented a form of aphasia differentfrom the nonfluent variety that followed frontaldamage. In what he called sensory aphasia, a pos-terior lesion in the region of the first temporalgyrus caused a disturbance in auditory compre-hension, inappropriate word selection in sponta-neous speech, and impaired naming and writing.

In his landmark monograph Der aphasischeSymptomen-complex, Wernicke reasoned thatBrocas area was the center for the motor repre-sentation of speech, and the posterior first tempo-ral gyrus was the center for sound images. Wer-nicke also described global aphasia and explainedit as a result of destruction of both anterior andposterior language areas. He also made a predic-tion that a disturbance of the pathways between

these two areas would produce another variety of


Figure 1-9

Carl Wernicke (18481904).


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for this syndrome, a disturbance in repetitionshould accompany conduction aphasia.38,39 HugoLiepmann described the apraxias, including ideo-motor apraxia,40 and, with Maas, callosalapraxia,41 explaining them in terms of connec-tionist principles. Joseph Jules Dejerine, shown inFig. 1-11, also used the framework of centers and

connections in his explanation of alexia withoutagraphia.42

The nineteenth-century connectionist frame-work proved to have both parsimony and explana-tory power. Rather than hypothesizing a new cen-ter for every ability or every observed deficit, afterthe fashion of Gall, a relatively small number ofbasic centers (vision, sound images, motor out-puts) could be combined through connections to

explain a wide variety of higher functions and

their deficits. Connectionist explanations of apha-sia, apraxia, alexia, and other disorders survivedwell into the twentieth century; indeed, NormanGeschwind, one of the most influential behavioralneurologists of our time, championed themthroughout his career.43 Despite the current pro-liferation of theories and approaches in our field,the theories of Dejerine, Liepmann, Lichtheim,and Wernicke are still held to be correct by many.

Nevertheless, as successful as the connec-tionist framework was in late nineteenth centuryin explaining a variety of disorders, skeptics con-

tinued to reject the localism implicit in it. One ofthe most influential of these was the English neu-rologist John Hughlings Jackson, shown in Fig.1-12. He viewed the nervous system not as a seriesof centers connected by pathways but rather as ahierarchically organized and highly interactivewhole that could not be understood piecemeal.24

Figure 1-13 shows Pierre Marie, a Parisian studentof Broca and Charcot, who also took issue withthe connectionist theorizing of the late nineteenth


Figure 1-10

Wernickes model of the speech mechanism.36 The au-

ditory areas (a) project to centers subserving vocal out-

put (b) and areas which contain tactile (c) and visual

(d) images.

Figure 1-11

Joseph Jules Dejerine (18491917).


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toward holism continued into the early twentiethcentury, with Jackson and Marie followed by anumber of influential neurologists and psycholo-gists, including Henry Head in England,24 shownin Fig. 1-14, Kurt Goldstein in Germany,4547

shown in Fig. 1-15, and Karl Lashley in the UnitedStates.4851 This swing of the pendulum back to-ward holism has been explained by the waning ofGerman influence following World War I60 andthe growing influence of Gestalt psychology.61

While these workers emphasized the brainsunity, other researchers had pointed out the dif-

ference between brain regions in cellular mor-phology, cell densities, and lamination and pro-duced the first cytoarchitectonic maps. Oskar andCecile Vogt62,63 and Alfred W. Campbell64,65 pro-duced some of the earliest examples of these

century. His style was direct, to say the least. Oneof his articles was so offensive to Dejerine that itprovoked the latter to challenge Marie to a duel.His article questioning the empirical basis of theearly claims concerning speech localization wasentitled La troisieme circonvolution frontale

gauche ne joue aucun role special dans la fonctiondu langage (The third frontal convolution playsno special role at all in the function of lan-guage).44 Marie believed that there was just onebasic form of aphasia, a posterior aphasia, whichwas a type of general intellectual loss not specificto language per se. He held that the speech prob-lems of anterior aphasics were motoric in nature.When aphasia is viewed this way, a network of

specialized centers is superfluous. A movement


Figure 1-12

John Hughlings Jackson (18351911).

Figure 1-13

Pierre Marie (18531940).


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architectonic maps, followed by many others, in-cluding those of Korbinian Brodmann,58 whose

cortical maps of the human brain have had themost widespread application. While these workersdid not agree on the number and location of corti-cal areas (the Vogts counted over 200, Brodmannonly 523) it could not be contested that there wereclear regional neuroanatomic differences.

The late nineteenth century also saw the be-ginnings of the modern study of memory and vi-

sion. Theodule Ribot introduced the distinctionbetween anterograde and retrograde memory im-pairments and observed what is now known asRibots law, that the most recently laid downmemories were the most vulnerable to brain dam-age.5961 Ribot can also be credited with describingpreserved learning in amnesia, thus anticipatingthe distinction between declarative and nonde-clarative forms of memory that has been so inten-

sively investigated in our own recent times. An ad-

ditional contribution to memory research in thelatter nineteenth century was the description byWernicke and Korsakoff (shown in Fig. 1-16) ofthe syndrome that bears their names, includingKorsakoffs observations of what he calledpseudo-reminiscence, now known as confabu-lation.70,71

In 1881, Hermann Munk reported that whenhe ablated the occipital lobes of dogs, they seemedunable to recognize objects despite seeing wellenough to navigate the visual environment.72

Shortly thereafter, Lissauer presented one of the

earliest clinical descriptions of visual recognitionimpairment in a human and suggested the distinc-tion between apperceptive and associative impair-menta clinical dichotomy still in use today.73

Freud would later introduce the term agnosia todescribe these conditions.74 In the decades thatfollowed, the visuospatial functions of the righthemisphere finally attracted the attention of neu-


Figure 1-14

Henry Head (18611940).

Figure 1-15

Kurt Goldstein (18781965).


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took hold. Neurologists and neuropsychologistsbegan to design experiments patterned on re-search methods in experimental psychology.

Typical research designs in experimentalpsychology involved groups of normal subjectsgiven different experimental treatments (for ex-ample, different training or different stimulus ma-terials), and the effects of the treatments weremeasured in standardized protocols and com-pared using statistical methods such as analysis ofvariance. In neuropsychology, the treatmentswere, as a rule, naturally occurring brain lesions.

Groups of patients with different lesion sites orbehavioral syndromes were tested with standardprotocols, yielding quantitative measures of per-formance, and these performances were com-pared across patient groups and with non-brain-damaged control groups. Unlike the impairmentsstudied previously in single-case designs, whichwere so striking that control subjects would gener-ally have been superfluous, experimental neuro-psychology often focused on group differences ofa rather subtle nature, which required statisticalanalysis to substantiate.

The most common question addressed bythese studies concerned localization of function.Often the localization sought was no more precisethan left versus right hemisphere or one quadrantof the brain (which, in the days before computed

tomography, often amounted to left versus righthemisphere with presence or absence of visualfield defects and/or hemiplegia). Given the hugeamount of research done during this period onlanguage, memory, perception, attention, emo-tion, praxis, and so-called executive functions, itwould be hopeless even to attempt a summary.For those interested in some examples of this ap-

proach, we cite here some classic papers from avariety of the active laboratories of the period, ad-dressing the question Is the right hemispherespecialized for spatial perception of propertiessuch as location,7072 orientation,73,74 and large-scale topography?75,76

The influential research program of theMontreal Neurological Institute also began duringthis period. In the wake of William Scovilles

discovery that the bilateral medial temporal

rologists and neuropsychologists.6769 The rela-tively delayed entry of this realm of functioning

into the research arena is probably a result of thefields original focus on language and the lefthemisphere, reflected in the nineteenth-centuryterminology ofmajorand minor hemisphere.

THE RISE OF EXPERIMENTALNEUROPSYCHOLOGY

Most of the advances described so far in this chap-ter were made by studying individual patients, orat most a small series of patients with similar dis-orders. In many instances, particularly before themiddle of this century, patients behavior wasstudied relatively naturalistically, without plannedprotocols or quantitative measurements. In thenineteen sixties and seventies, a different ap-

proach to the study of brain-behavior relations


Figure 1-16

Sergei S. Korsakoff (18531900).


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resection he performed on epileptic patient H. M.resulted in permanent and dense amnesia, BrendaMilner and her colleagues investigated this patientand groups of other operated epileptic patients.This enabled them to address questions of func-tional localization with the anatomic precision ofknown surgical lesions (e.g., see Refs. 77 and 78for reviews of research from that period on frontallobe function and temporal lobe function, respec-tively). At the same time, another surgical inter-vention for epilepsy, callosotomy, also spawned

a productive and influential research program.Roger Sperry and his students and collaboratorswere able to address a wide variety of questionsabout hemispheric specialization by studying theisolated functioning of the human cerebral hemi-spheres.79

In addition to answering questions about lo-calization, the experimental neuropsychology ofthe sixties and seventies also uncovered aspects ofthe functional organization of behavior. By exam-ining patterns of association and dissociationamong abilities over groups of subjects, research-ers tried to determine which abilities depend onthe same underlying functional systems and whichare functionally independent. For example, thefrequent association of aphasia and apraxia hadbeen taken by some to support the notion that

aphasia was not language-specific but was just onemanifestation of a more pervasive loss of the abil-ity to symbolize or represent (asymbolia). Aclassic group study by Goodglass and Kaplan80 un-dermined this position by showing that severity ofapraxia and aphasia were uncorrelated in a largesample of left-hemisphere-damaged subjects. Asecond example of the use of dissociations be-tween groups of patients from this period is thedemonstration of the functional distinction, byNewcombe and Russell, within vision betweenpattern recognition and spatial orientation.81

By the end of the seventies, experimentalneuropsychology had matured to the point wheremany perceptual, cognitive, and motor abilitieshad been associated with particular brain regions,and certain features of the functional organization

of these abilities had been delineated. Accord-

ingly, it was at this time that first editions of someof the best-known neuropsychology texts ap-peared, such as those by Hecaen and Albert,82

Heilman and Valenstein,83 Kolb and Whishaw,84

Springer and Deutsch,85 and Walsh.86

Despite the tremendous progress of this pe-riod, experimental neuropsychology remaineddistinct from and relatively unknown within aca-demic psychology. Particularly in the UnitedStates, but also to a large extent in Canada andEurope (the three largest contributors to the

worlds psychology literature), experimental neu-ropsychologists tended to work in medical centersrather than university psychology departmentsand to publish their work in journals separatefrom mainstream experimental psychology. Animportant turning point in the histories of bothneuropsychology and the psychology of normalhuman function came when researchers in eacharea became aware of the other.

THE BIRTH OF COGNITIVENEUROSCIENCE

Patient-based cognitive neuroscience was bornwhen the theories and methods of cognitive psy-chology and neuropsychology were finally com-

bined. Both fields had strong incentives to over-come their isolation. Let us begin by reviewing thestate of cognitive psychology prior to the birth ofcognitive neuroscience.

The central tenet of cognitive psychology isthat cognition is information processing. Al-though the effects of damage to an information-processing mechanism might seem to be a goodsource of clues as to its normal operation, cogni-tive psychologists of the seventies were generallyquite ignorant of contemporary neuropsychology.

The reason that most cognitive psychologistsof the 1970s ignored neuropsychology stemmedfrom an overly narrow conception of informationprocessing, based on the digital computer. A basictenet of cognitive psychology was the computeranalogy for the mind: the mind is to the brain as

software is to hardware in a computer. Given that



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and measuring responses and response latencies,cognitive psychologists made inferences about theinformation processing that intervened betweenstimulus and responses. But such inferences wereindirect, and in some cases they were incapable ofdistinguishing between rival theories. In 1978 thecognitive psychologist John Anderson publishedan influential paper87 in which he called this theidentifiability problem and took as his examplethe debate over whether mental images weremore like perceptual representations or linguistic

representations. He argued that the fields inabil-ity to resolve this issue, despite many years of re-search, was due to the impossibility of uniquelyidentifying internal cognitive processes fromstimulus-response relations. He suggested that thedirect study of brain function could, in principle,make a unique identification possible, but he indi-cated that such a solution probably lay in the dis-tant future.

That distant future came to pass within thenext 10 years, as cognitive psychologists workingon a variety of different topics found that thestudy of neurologic patients provided a powerfulnew source of evidence for testing their theories.In the case of mental imagery, taken by Andersonto be emblematic of the identifiability problem,the finding that perceptual impairments after

brain damage were frequently accompanied byparallel imagery impairments strongly favored theperceptual hypothesis.88 The study of learning andmemory within cognitive psychology was revolu-tionized by the influx of ideas and findings on pre-served learning in amnesia, leading to the hypoth-esis of multiple memory systems.8991 In the studyof attention, cognitive psychologists had for yearsfocused on the issue of early versus late atten-tional selection without achieving a resolution,and here too neurologic disorders were crucial inmoving the field forward. The phenomena of ne-glect provided dramatic evidence of selectionfrom spatially formatted perceptual representa-tions, and the variability in neglects manifesta-tions from case to case helped to establish the pos-sibility of multiple loci for attentional selection as

opposed to a single early or late locus. The idea

the same computer can run different programsand the same program can be run on differentcomputers, this analogy suggests that hardwareand software are independent and that the brainis therefore irrelevant to cognitive psychology. Ifyou want to understand the nature of the programthat is the human mind, studying neuropsychologyis as pointless as trying to understand how a com-puter is programmed by looking at the circuitboards.

The problem with the computer analogy is

that hardware and software are independent onlyfor very special types of computational systems:those systems that have been engineered, throughgreat effort and ingenuity, to make the hardwareand software independent, enabling one computerto run many programs and enabling those pro-grams to be portable to other computers. Thebrain was designed by very different pressures,and there is no reason to believe that, in general,information-processing functions and the physicalsubstrate of those functions will be independent.In fact, as cognitive psychologists finally began tolearn about neuropsychology, it became apparentthat cognitive functions break down in character-istic and highly informative ways after brain dam-age. By the early 1980s, cognitive psychology andneuropsychology were finally in communication

with one another. Since then, we have seen an ex-plosion of meetings, books, and new journalsdevoted to so-called cognitive neuropsychology.Perhaps more important, existing cognitive psy-chology journals have begun to publish neuropsy-chological studies, and articles in existing neuro-psychology and neurology journals frequentlyinclude discussions of the cognitive psychologyliterature.

Let us take a closer look at the scientificforces that drove this change in disciplinaryboundaries. By 1980, both cognitive psychologyand neuropsychology had reached states of devel-opment that were, if not exactly impasses, pointsof diminishing returns for the concepts and meth-ods of their own isolated disciplines. In cognitivepsychology, the problem concerned methodologic

limitations. By varying stimuli and instructions



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of separate visual feature maps, supported bycases of acquired color, motion, and depth blind-ness, provided the inspiration for the most noveldevelopment in recent cognitive theories of atten-

tionnamely, feature integration theory.92

What did neuroscience gain from the rap-prochement with cognitive psychology? The mainbenefits were theoretical rather than methodo-logic. Traditionally, neuropsychologists studiedthe localization and functional organization ofabilities, such as speech, reading, memory, objectrecognition, and so forth. But few would doubt

that each of these abilities depends upon an or-chestrated set of component cognitive processes,and it seems far more likely that the underlyingcognitive components, rather than the task-defined abilities, are what is implemented in local-ized neural tissue. The theories of cognitive psy-chology therefore allowed neuropsychologists topose questions about the localization and func-tional organization of the components of the cog-nitive architecture, a level of theoretical analysisthat was more likely to yield clear and generaliz-able findings.

Among patients with reading disorders, forexample, some are impaired at reading nonwords(e.g., plif) while others are impaired at reading ir-regular words (e.g., yacht). Rather than attemptto localize nonword reading or irregular word

reading per se and delineate them as independentabilities, neuropsychologists have been able to usea theory of reading developed in cognitive psy-chology to interpret these disorders in terms ofdamage to a whole-word recognition system anda grapheme-to-phoneme translation system, re-spectively.93 This interpretation has the advantageof correctly predicting additional features of pa-tient behavior, such as the tendency to misreadnonwords as words of overall similar appearancewhen operating with only the whole-word system.

In recent years the neuroscience of everymajor cognitive system has adopted the theoreti-cal framework of cognitive psychology in a gen-eral way, and in some cases specific theories havebeen incorporated. This is reflected in the contentand organization of the present book. For the

most intensively studied areas of behavioral neu-

rology and neuropsychologynamely, visual at-tention, memory, language, frontal lobe function,and Alzheimers diseaseintegrated pairs ofchapters review the clinical and anatomic aspects

of the relevant disorders and their cognitive theo-retical interpretations. Chapters on other topicswill cover both the clinical and theoretical as-pects together.

COMPLEMENTARY METHODS INCOGNITIVE NEUROSCIENCE: PATIENT

STUDIES AND FUNCTIONAL IMAGING

Following its introduction in the 1970s, positronemission tomography (PET) was quickly em-braced by researchers interested in brain-behaviorrelations. This technique provides images of re-gional glucose utilization, blood flow, oxygen con-sumption, or receptor density in the brains of livehumans. Resting studies, in which subjects arescanned while resting passively, have provideda window on differences between normal andpathologic brain function in a number of neuro-logic and psychiatric conditions. With the use ofradioactive ligands, abnormalities can be localizedto specific neurotransmitter systems as well asspecific anatomic regions. Activation studies, inwhich separate images are collected while normal

subjects perform different tasks (typically oneor more active tasks and one resting baseline)yielded new insights on the localization of cogni-tive processes. These localizations were not stud-ied region by region, as necessitated by the lesiontechnique, but could be apprehended simultane-ously in a whole intact brain.

Positron emission tomography was soonjoined by other techniques for measuring regionalbrain activity, each of which has its own strengthsand weaknesses. Single photon emission com-puted tomography (SPECT) was quickly adaptedfor some of the same applications as PET, provid-ing a less expensive but also less quantifiable andspatially less accurate method for obtaining im-ages of regional cerebral blood flow. With new de-velopments in the measurement and analysis of

electromagnetic signals, the relatively old tech-



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neuroimaging has had a huge and salutary effecton cognitive neuroscience, significantly expandingthe range of questions that can be addressed, ithas not replaced research with neurological pa-

tients. A full discussion of the complementarystrengths and weaknesses of the two approachescould easily fill a chapter by itself, but a few of themost consequential differences can be summa-rized briefly here.

Lumping the different functional neuro-imaging modalities together, they generally offerbetter spatial resolution than can be obtained in

inferences from a few patients with focal brain le-sions. Imaging also allows us to study normalbrains, which lesion studies by definition do not.Furthermore, for some neurological conditionsmore than others there may be reason to suspecta degree of reorganization of remaining brain sys-tems in response to damage. These are probablythe greatest benefits of functional neuroimaging,although by no means the only ones.

The greatest weakness of neuroimaging is itsinability to settle any issue concerning what mightbe called mechanism. An important goal of cogni-tive neuroscience is to identify the causal chain ofneural events, or the mechanisms, underlying cog-nition. The data of functional neuroimaging arecorrelational: a certain area is activated when acertain cognitive process is occurring. Neuroimag-

ing can never disentangle correlation from causa-tion; in other words, it can never tell us whichbrain areas are causally involved in enabling acognitive process. Activated regions could play acausal role or could be activated in an optional oreven an epiphenomenal way. For this we mustturn to studying the effects of brain damage, theexperiments of nature that provide a direct testof the causal role of different brain areas by show-ing us how the system works in their absence.Given the complementary strengths of neuro-imaging and patient studies, we predict that themost successful cognitive neuroscience researchprograms of the twenty-first century will be thosethat combine the two approaches.

niques of electroencephalography (EEG) andevent-related potentials (ERPs), as well as mag-netoencephalography (MEG), joined the ranks offunctional imaging techniques allowing some de-

gree of anatomic localization of brain activity,with temporal resolution that is superior to theblood flow and metabolic techniques. Most re-cently, functional magnetic resonance imaging(fMRI) has provided a particularly attractivepackage of reasonably good anatomic and tempo-ral resolution, using techniques that are noninva-sive and can be implemented with equipment

available for clinical purposes in many hospitals.Much of the early work with functional neu-

roimaging could be considered a form of calibra-tion, in that researchers sought to confirm well-established principles of functional neuroanatomyusing the new techniquesfor example, demon-strating that visual stimulation activates visualcortex. As functional neuroimaging matured, re-searchers began to address new questions, towhich the answers were not already known in ad-vance. An important development in this secondwave of research was the introduction of theoriesand methods from cognitive psychology, whichspecified the component cognitive processes in-volved in performing complex tasks and provideda means of isolating them experimentally. In neu-roimaging studies of normal subjects, as with the

purely behavioral studies of patients, the entitiesmost likely to yield clear and consistent localiza-tions are these component cognitive processes andnot the tasks themselves. Starting in the mid-1980s, a collaboration between cognitive psychol-ogist Michael Posner and neurologist MarcusRaichle at Washington University led to a seriesof pioneering studies in which the neural circuitsunderlying language, reading, and attention werestudied by PET (see Ref. 94 for a review). Sincethen, researchers at Washington University and agrowing number of other centers around theworld have adapted neuroimaging techniques toall manner of topics in cognitive neuroscience.

To many psychologists and neuroscientists,cognitive neuroscience is equivalent to cognitiveneuroimaging. At the very least, we hope this

book shows this idea to be mistaken. Although



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