the history of neuroscience in autobiography - vol 2

8/12/2019 The History of Neuroscience in Autobiography - Vol 2

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T h e H is to r y o f N e u r o s c i e n c e

in A u to b io g r a p h y

VOLUME

Ed ited by Larry R Squire

C D E M I C P R E S S

London Bos ton New York Sydn ey Tokyoan Diego Toronto


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T h i s b o o k i s p r i n t e d o n a c i d - f r e e p a p e r . @

Copy right 9 1998 by The Socie ty for Neuroscience

Al l Rights R eserved .

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storage and retrieval system, without permission in writ ing from the publisher.

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Internat ional Standard Book Num ber: 0-12-660302-2

P R I N T E D I N T H E U N I T E D S T A T E S O F A M E R I C A

98 99 00 01 02 03 EB 9 8 7 6 5 4 3 2 1


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Contents

Lloyd M. Beidler 2

Arvid Carlsson 28

Donald R. Griffin 68

Roger Guillemin 94

Ray Guillery 132

Masao Ito 168

Martin G. Larrabee 192

Jerome Lettvin 222

Paul D. MacLean 244

Brenda Milner 276

Karl H. Pribram 306

Eugene Roberts 350

Gunther Stent 396


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K a r l H P r i b r a m

B O R N :

EDUCATION:

Vienna, Aust r ia

February 25, 1919

U niv ersit y of Chicago, B.S. (1938)

U niv ersi ty of Chicago, M .D. (1941)

APPOINTMENTS:

Yerkes La borator ies of Pr im ate Biology (1946)

Yale University (1948)

Ce nter for Ad vanced Studies , Stan ford U nive rsi ty (1958)

Stanford Universi ty (1959)

Professor Em eri tus , S tanford U nivers i ty (1989)

Distingu ished Professor, Radford U niv ersi ty (1989)

HONORS AND AWARDS (SELECTED):

NIH Life time Research Career Aw ard (1962)

International Neuropsychological Society (President, 1967)

A m erican Psychological Association

Division of Physiological and Co m parativ e P sychology

(President, 1967-1968)

Division of Theological an d Philosophical P sychology

(President , 1979-1980)

M enfred Sakel A ward, Society for Biological Ps ych iatry (1976)

Realia Honor, In st i tu te for Ad vanced Philosophic Research

(1986)

O uts tand ing Cont r ibu tions Award, Amer ican Board of

M edical Psy chot herap ists (1990)

H ono rary P h.D. in Psychology, U nive rsi ty of M ontreal ,

Canada (1992)

Neura l N e twork Leadersh ip Aw ard , In te rna t iona l Neu ra l

Ne tw ork Society (1994)

H onorary Ph.D. in Neuroscience, Un ivers i ty of Brem en,

G erm any (1996)

Karl Pribram was trained as a neurosurgeon and then devoted his

career to elucidating the structure and function of the cerebral cortex,

relating human clinical experience to his neurophysiological and

neurobehavioral studies on nonhuman primates. He discovered the visual

functions of the temporal lobe and the relationship of the anterior frontal

cortex to the limbic system. His theoretical writings include the topics of

perception, emotion, memory, and planning.

The History of Neuroscience in Autobiography, Volume 2

Copyright 9 1998 by The Society for Neuroscience


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K a r l H . P r i b r a m

P r e a m b l e

S

u m m e r 1918. The head of the bacterological services of the Austr ian

Army and a Dutch Red Cross nurse were swimming nude in the Dan-

ube somewhere between Vienna and Budapest. Some pigs pulled

their clothes from the bushes; retrieval entailed a considerable chase. The

child conceived on that occasion was often accused of "pigging out," and his

mann ers were attributed to that chase by the Danube.

February 25, 1919. I was born in Vienna, Austria at 8:00 PM encased

in the amniotic s ac~ t ak en by my mother to be a most propitious beginning.

A u t u m n 1923. I set off to a Kinderheim in Gstaadt, Switzerland to

protect me and my moth er from a stormy relationship.

S u m m e r 1924. Ernest August Pr ibram, my father, went to America to

save me from growing up in a Europe whose future he saw as torn with

political turmoil.

A u t u m n 1926. I joined my mother in Vienna and finished second grade

in Catholic school.

S u m m e r 1927. I went to a farm in Geneva, and learned French.

Au t u m n 1927. I and my mother arrived in the United States and my

reunited family settled in Chicago.

Th e S c i en t i s t a s a Y o u n g M an

It was Labor Day, 1932, when my father put me on a train in Chicago to

head for Culver Military Academy near Fort Wayne, Indiana. I had set my

sights on going to Culver once I had heard about it from my dentist, a

gentle, wonderful man who saved my adolescent cavitous molars which,

over the next 70 years, I have h ad to protect from the more rapac ious of the

dental profession. Tonsils I still also own, despite the medical fad that no

one should reach adulthood with such natural protective devices in place.

In each case battles with established practice had to be engaged and each

time I won: good tra ining for a career in research.

My attraction to Culver was simple: horses. My father could not afford

the additi onal fees to allow me to become a member of the famed Black

Horse Troop which was sent to Washington for each presidential inaugu-


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309

ration. But the field artillery was still horse-drawn in that long-ago time

when the caissons and French 75's went rolling along.

Before going to Culver, I had a dismal 6-year record in Chicago's public

schools, being repeatedly expelled for fighting with bullies who picked on

everyone, or from Catholic schools for asking the nuns simple questions

such as how God could be both all good and all just. Wha t I saw around me

during the depression of the 1930s didn't fit the picture of God th at those

good women tried to convey. When told to have faith, I lost all faith in what

the hooded ladies had to say, and expressed my opinion in no uncertain

terms. Over and over. My fath er had Jesuit friends who tried to reason with

me, but as I would not accept their premises, they taught me only that

reason can be totally reasonable and t hat what one needs to ascerta in tr uth

or falsity is to search for the premises, the axioms, from which the reasoning

takes off.

The public school teachers were being paid with script, so during the

last year in elementary school, I took over when teachers were absent--

which was often. I figured tha t each of the s tuden ts had fathers and th at

the fathers worked and could tell their children what they did. So, each

pupil came to class prepared to tell us about his father's occupation, and

what it entailed both intellectually and in practice. We were all fascinated

and much preferred our system to the su bstitut e teachers th at were foisted

upon us. The admini strat ion was not altogeth er pleased.

My father believed that a military environment was just what his son

needed. As we said good-bye I hugged him (he didn't like hugging much)

and I said that I was proud to be admitted to such an excellent school. My

fathe r replied, "Be sure to conduct yourself in such a mann er tha t C u l v e r

w i l l b e p r o u d o f y o u

when you leave." Those parting words stayed with me

for the res t of my life.

During my senior year science was to be taught for the first time in the

histor y of the Academy, one course in physics and ano the r in chemistry. I

immediately registered for both. Although the two courses were not to be

taken simultaneously, I pleaded that as a senior, I had been prevented from

having an adequate science education. I took the courses and received hon-

ors in both, as well as in history and in English literature. Each required a

special project that went beyond what had been covered in class. Only one

stud ent was to receive honors in a parti cular subject and each stud ent was

eligible to receive honors in only one subject. Fortunately, only the compet-

itive "one subject, one student" par t had been mad e explicit and my projects

were completed at graduation when the "one student, only one honor" be-

came elevated to everyone's consciousness, as we would say today. I gradu-

ated with all four honors.

The decision as to which university to attend was a difficult one. I had

been accepted at Oxford, Harvard, and the Univers ity of Chicago, but two

related factors favored Chicago: (1) I had decided to study biology and


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31 0 Karl H. Pribram

me di c i ne , a nd my f a th e r , a n e mi ne n t b i o l o g i s t ( ba c t e r i o l o g i s t , pa th o l o g i s t ,

a n d i m m u n o l o g i s t ) w i t h w h o m I h a r d l y e v e r h a d i n t e r a c t e d , w a s t h e r e ;

a n d ( 2 ) t h o u g h a d e c i s i o n f o r H a r v a r d o r O x f o r d w o u l d h a v e c a r r i e d m o r e

p r e s t i g e , i n 1 9 3 6 , C h i c a g o u n d e r M a y n a r d H u t c h i n s w a s m o r e i n t e l l e c t u -

a l l y a li ve a n d i nno v a t i ve . S o C h i c a go i t w a s .

M y f a t h e r a n d I m e t e v e r y S u n d a y . H e p o r t r a y e d t h e f a c t s a n d i d e a s

i m p o r t a n t t o p h y s i o l o g y a n d i m m u n o l o g y t o m e i n u n f o r g e t t a b l e f a s h i o n .

J u s t r e c e n t l y I j o i n e d i n w r i t i n g t w o p a p e r s t h a t d e a l w i t h t h e p o s s i b i li t y o f

s u p e r c o n d u c t i v i t y i n d e n d r i t e m e m b r a n e s b e c a u s e t h e f o r m u l a t i o n s a r e

c o n s o n a n t w i t h t h e s e e a r l y tu i t i on s .

F i ve ye a r s l a t e r , i n 1 9 41 , I r e c e i ve d m y M. D. A c c o r d i ng t o H u tc h i ns , i t

w a s n o t r e a l ly c o n s i d e r e d a d o c t o ra t e , r a t h e r s i m p l y a p e r m i t t o p r a c t ic e a

t r a d e . T h e C h i c ag o c ol or s w e r e w i t h h e l d f r o m t h e g o w n s w o r n t o t h e g r a d -

u a t i o n c e r e m o n i e s b y t h e i n c i p i e n t m e d ic s.

I h a d d o n e w e l l a t u n i v e r s i t y . I l o v e d m y u n d e r g r a d u a t e c o u r s e s i n

h i s to r y a nd e c o no mi c s , ph ys i c s a nd c h e mi s t r y , a nd b i o l o g i c a l d i s c o ve r y ,

m a d e s t r a i g h t A ' s , a n d t o o k c o m p r e h e n s i v e e x a m s i n s t r i d e , o f t e n w i t h o u t

h a v i n g a t t e n d e d c l as s es ( a n O .K . u n d e r H u t c h i n s ) . I t o o k c op i ou s n o te s. T h e

e n d o c r i n e s a n d t h e b r a i n w e r e e s p e ci a ll y i n t r i g u i n g b e c a u s e t h e y s e r v e d a s

i n t e g r a t o r s o f t h e f u n c t i o n s o f t h e b od y. V i e w i n g m y s e l f as a p o t e n t i a l e x-

p l o r er (I h a d s t e ep e d m y s e l f i n A m u n d s e n a n d A d m i r a l B y r d w h i le in m y

p r e t e e n s a n d i n P a u l D e K r u i f 's The Microbe Hun ters s o m e w h a t l a te r ) I sa w

e n d o c r i n o l o g y a n d b i o c h e m i s t r y a t o n e e x t r e m e ~ w h e r e t o o m u c h r e s e a r c h

h a d a l r e a d y b e e n d o n e t o c o n s id e r t h e m v i r g in f i e l d s ~ a n d b r a i n p h y s io lo g y

a t t h e o t h e r e x t r e m e , w h e r e t o o f e w t e c h n i q u e s s e e m e d t o b e a v a i l a b l e f o r

f r u i t f u l e xp l o r a t i o n .

A l l t h i s c h a n g e d o n c e I e n g a g e d t h e m e d i c a l c li n ic a l c u r r i c u l u m . S e e i n g

p a t i e n t s w a s w o n d e r f u l ly s a t is f y i n g f r om t h e h u m a n a n d d i a g n o st ic s t a n d -

p o i n t , b u t t h e l e c t u r e s a n d l a b o r a t o r y s e s s i o n s w e r e i n c r e d i b l y d u l l . W h e n

I a s k e d q u e s t i o n s o f " ho w " or "h o w co m e," I w a s s u m m a r i l y i n f o r m e d t h a t

t h e a n s w e r w a s , " b e c a u s e I s a y s o . " I w a s c h a l l e n g i n g a u t h o r i t y a n d t h a t

w a s a no - no . C a th o l i c s c h o o l a l l o ve r a ga i n . T h i s i n H u tc h i ns ' s C h i c a go ?

O bv i o us l y , H u tc h i ns w a s c o r r e c t i n h i s e va l ua t i o n : me d i c a l s c h o o l w a s a

t rade schoo l .

I n o n e cl a ss w e w e r e t o t y p e p n e u m o n i a s . T h i r t y - s i x t y p e s w e r e k n o w n

a n d d u r i n g t h e c o u r s e o f t h e c la ss , tw o m o r e w e r e d i s c o v er e d . S u l f a n o m i d e s

h a d a ls o j u s t b e e n d i s co v e r e d a n d w e r e o n t h e m a r k e t . W a s i t r e a l l y n ec e s-

s a r y t o l e a r n t h e t yp i ng p r o c e du r e ? T h e c l a s s de c i de d : No . T h e p r o f e s s o r

d e c l ar e d : Y es. T h e c la s s w a l k e d o u t a n d g o t a w a y w i t h it . S o m e o f H u t c h i n s ' s

i n f lu e n c e p e r m e a t e d e v e n i n t o t h e d e p t h s o f t h e m e d i c a l e s t a b l i s h m e n t .

O v e r t h e y e a r s m e d i c a l e d u c a t i o n h a s , t o so m e e x t e n t , r e m e d i e d t h i s i n to l-

e r a b l e t e a c h i n g s i t u a t i o n , b u t o n e s t i l l r e a d s a r t i c l e s b y s t u d e n t s , i n t e r n s ,

a nd r e s i de n t s w h o t e l l a s t o r y t h a t i s no t t o o d i f f e r e n t f r o m th e o ne I

e x p e r i e n c e d .


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Karl H. Pribra m 311

The only way to make the study of medicine outside of the clinic inter-

esting was to at tac h oneself to one or ano the r of the professors and aid them

in their research. One got to know them, the puzzles they were trying to

solve, and the broader biological perspective within which medical care

operates. I decided to apprentice in the depa rt ment of physiology. "Ajax"

Carlson was its head. He insisted that every student in the department

repeat the historically important experiments that led to current views of

how the human body functioned. This dictum had an enormous influence

on my research career. Over the years, many experiments done in my labo-

rator ies were init iated to see for ourselves the details of cur ren tly impor-

tant findings that can only come from first-hand experience in generating

data . Thus, when confronted in a discussion of ideas, I would always be able

to call up Carlson 's o ther dictum: "Wass iss die effidence?" And, to the ext ent

possible, I would have t ha t evidence first-hand.

I was privy to exploring with my professors the deter iorati on of the

activity of vit amin C in orange juice with storage; the effects in dogs of

denervat ion of the kidneys, an d a project I devised and performed myself:

How much blood flow does the liver need before deficiencies in function

would show up on laborato ry tests? I constr icted the flow of blood to the

liver by placing adjustable clamps on the hepatic artery and the vena

cava. The clamps had been devised by Goldbl att to do ju st this sort of

study on the kidneys. Goldblatt found that impairing kidney blood flow

produced persistently elevated blood pressure. The liver experiments had

no such dramatic effects. Turning the screws on the Goldblatt clamps had

no effect whatsoeve r on the then-ava ilable tests of liver fu nc ti on ~u nt il

the l ast one-eighth tur n of either screw (in the clamp on the vena cava

or the one on the hepatic artery). With this last turn, the animal, which

had to all intents been perfectly normal for weeks, died. Any small aggre-

gate of surviving liver cells could function in lieu of the ent ire liver, up to

a point.

It was a lesson I remember ed when, much later, I learn ed th at memory

storage in the brain shows considerable resistance to degradation when

large extents of th at organ are damaged.

Most exciting of these forays into research, however, was an exposure

to the work being done in Ralph Gerard's laboratory. On one occasion, an

electrical record from a bra in site energized a loudspeaker. Wheneve r a loud

clap was produced near the cat from which the record was being taken, the

loudspeaker would give out a distinct sound. A tap to the cat's paw also

produced a sharp sound on the loudspeaker. Brain electrical activity re-

flected the sensory input! A discussion ensued: why couldn't these results

be shown outside the laboratory? Electrocardiograms were being made

daily, why not electroencephalograms (EEGs)? The answer was that the

electrical changes produced by the brain were several magnitudes smaller

tha n those produced by the heart.


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312 Kar l H. Pr ib ram

That evening, across the table from me sat Frank Offner, a student

enginee r who became intri gued by the problem when I told him wh at I had

seen. He s tated th at the curr ent lack of sufficient amplification of the signal

with respect to the noise of the syste m should not be insurmo unta ble. I

introduced Offner to Gerard. Frank Offner spent his life making and mar-

keting EEG machines, the hard-copy electroencephalograms we sought

that evening at dinner. It was the first time I realized how many contribu-

tions to neurobehavioral science could be made in informal settings far

removed from the laboratory. There were to be many, many more.

Ralph Gerard was to play a most significant part in my education.

Gerard was an incisive thinker and a brilliant teacher. Whenever sloppy

reaso ning went on in the classroom he propelled a piece of chalk at the

perpetrator. On one such occasion, I was the target: my thinking had been

teleological, a process forbidden in Gerard's neurophysiology. Many years

later, neurophysio logists became aware of the ubiquit ous presence of feed-

back and feedforward processes in the nervous system. I laughed privately

at my former master and always cocksure friend. "Ha ha, how wrong you

were in your certainty," I exclaimed as I was composing

L a n g u a g e s o f

the Brain.

I have spent man y hours in the classroom pointing out how our ideas of

the functions of the brai n have evolved, and th at even what I have writ ten

and t au gh t has become obsolete. I am not sure th at my way of teachin g is

better than Gerard's. Perhaps uncertainty is too unsettling to a student.

Let st ude nts be misled and find out for themselves, allowing the m to expe-

rience t he glee of hav ing the "professor" shown to be wrong.

Of course I didn't always wait until later to challenge my professor. On

one occasion Gerard was providing the class with the criteria for classifying

mamma ls. Among these was hair. I stroked my long dense locks and asked:

"Really, do

all

mammals have hair?" The class roared. Gerard was as bald

as Yul Bryn ne r in "The King and I."

For me, the high point of Gerard 's classes came a t the t ime of a final

exam in an advanced course on neurophysiology. Gerard asked only one

question: Discuss the organiz ation of the nervous system. Fort una tely for

me, John Fulton's classic Physio logy o f the Nervo us S ys tem had just arrived

at the university bookstore the weekend before the exam. In preparation, I

had purchased a copy. The book was so fascinating that I could not put it

down and spent t he entire weekend reading, letti ng go the res t of my stud-

ies and preparation for finals. On Gerard's exam I filled eight blue books,

writing as fast as writing could be accomplished. Gerard stated it was the

very best he had ever received; how had I achieved such comprehensive

knowledge and superb organization? I told Gerard of Fulton's book. Fortu-

nately, Ge rard had not as yet seen it; the posing of his exam question was

not relat ed to the publication.

The first half of Fulton's

Physio logy o f the Nervo us Sys tem

was devoted

to sensory receptors, motor units, pe ripher al nerves, spinal cord, and brai n


13/49

Karl H. Pribram 313

stem. Microanatomy was presented with appropriate pictures. The second

hal f of the book did the same for the b rain's cortex and the fiber system

coursing to and from it and detailed the discovery of the functions of the

occipital (rear) lobes of the bra in's hemispheres . In a similar fashion, Fult on

reviewed the discovery of the connections of the parietal lobe to somatic

sensation and the temporal lobe to hearing. Nothing as yet was known of

the functions of the inferior pa rt of this lobe. This was to be my contribution

during the first years of work with Fulton.

The most fascinating and im port ant story for me, though I didn't realize

it at the time, was composed by the resul ts of damage to the frontal lobes of

the brain. Fulton's work had led to the procedure of frontal leukotomy, or

lobotomy, as it was commonly known. Severing the fibers connecting the

frontal lobes from structures in the brain stem was shown, on occasion, to

produce marked changes in personality. What caused these changes was

not known. After I had completed my training in neurosurgery (with Paul

Bucy, Eric Oddberg, Percival Bailey and Warren McCullock in Chicago,

Illinois; Eustace Simmes in Memphis, Tennessee; and Lyerly in Jackson-

ville, Florida), Fulton asked me to join him in finding out just what they

migh t be. But I am ahead of the story.

Another aspect of neurophysiology that I learned from Gerard and from

Fulton's book was that receptor-initiated signals course to the spinal cord

via nerves that reach it through a "root" that is separate from the one

leaving the cord to reach muscles. Experiments by Charles Bell in England

and Francois Magendie in France had shown that sectioning of the ent ering

root left the animal without sensation while its movements remained in-

tact. The reverse was true of the other, the outgoing root. Until these exper-

iments were completed, no one knew which root provided the input and

which the output. The results of the exper iment were heralded as a "law"

and were the basis for conceiving the basic unit of nervous activity as a

reflex arc.

Reflex arcs are segregated in segments which represent the fact that

the composition of our bodies is much like tha t of eart hworms. Each seg-

ment of the spinal cord is encapsulated by a vertebra. The vertebrae are

held together by sheaths which contain disc-shaped cartilaginous cushions.

When the sheaths rupture the cartilage oozes out to press on the nerve

roots, causing pain. When the compressed nerve roots carry signals from

the back of the leg, the pat ien t experiences sciatica. Much of the ordinary

practice of neur osurg ery is made up of removing such ru ptu red discs.

Many years later, just before joining Fulton in his laboratory at Yale

Univ ersi ty in 1948, one of the la st major hum an surgical operat ions I per-

formed was the removal of such a disc. I had recentl y read of an innova tive

technique, by which the operation was done with the patient lying on his

side, eliminating pressure on his abdominal blood vessels and thus mini-

mizing bleeding at the operative site. I was able to remove the disc before

the operating room nurse had fully completed setting up all the gear we


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314 Kar l H . Pr ib r am

usually needed, and performed the entire procedure in less than 20

minutes. The patient was eating steak a few hours later and had no recur-

rence of his problem.

However, in all his discussion of reflexes, Fu lto n did not me nti on a

major componen t of the out put root of the reflex: o ne-third of the nerves

composing this root end in th e receptors of muscles. Thu s muscle receptor

activity is regul ated not only by the s tretch ing of the muscle but by signals

coming to the receptor from the spinal cord. The spinal cord signals are in

tu rn controlled by signals coming from the brain. Althoug h known to exist

previously, the i mportan ce of these receptor regul atin g nerves came into

focus in the 1950s th rough the work of John Eccles and S tep hen Kuffler.

They inv estig ated the effects of sti mula tin g the nerves going to the recep-

tors after having removed the functions that make muscles contract.

The "law of Bell and Magendie" was, af ter all, not a law. The reflex arc

is not an arc but a mechanism akin to a thermostat that can be set to a

particular value which determines the operation (the on's and the off's) of

the system. Soon other receptors were found to be regulated in a similar

fashion. Years later, Spinelli and I would devote a decade to showing that

retinal processes were subject to such central control. In 1960 George Mil-

ler, Eugene Galanter, and I wrote a book (P lans and the S t r uc tu r e o f Behav-

ior)

in which we detailed the i mport of the new neurology and moved

psychology from stimulus-response, reflex-arc behaviorism to a cognitive

science which paid heed to the brain's control over its own input from

the senses.

In 1949 Jo hn Fulto n pres ented me with the thi rd edition of his book

inscribed, "In warm appreciation . . . . "My main contribution to this edition

was to rewrite the chapter on the brain's control over the autonomic ner-

vous system. The autonomic system is called this because it regulates the

functions of the viscera which, for the most p art, take care of their own

process and function autom atical ly withou t our awarenes s or conscious in-

tervention. However, I had by then established that the brain's cortex had

an in put to the hy poth alamu s of the brai nstem which, in Fulton's earlier

editions, was called the head ganglion of the autonomic system. F ulto n

himself had obtained, with Margaret Kennard, results tha t indicated the

possible ope ratio n of such cortical control and in fact gave me my appo int-

ment at Yale, in part, because of my findings. But I am again get tin g way

ahea d of the story as it has unfolded over the years.

Gerard's lectures and laboratory courses with the climactic final exam

had me hooked. The brain was to be the continent I was to explore. Many

years later, Paul MacLean inscribed a translation (for which he was re-

sponsible) of a book wri tte n by Ramon y Cajal: "To Karl, Magell an of the

Brain." I was delighted with his insight as to my motivation in choosing a

research career.

Over these years, Gerard became a close friend, referred to as Poppa

R a l p h , because when my father was killed driving an auto in which my


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315

b r i d e a n d I w e r e p a s s e n g e r s , G e r a r d t e l e g r a p h e d h is r e a d i n e s s t o f e tc h u s

f r o m M o n t a n a w h e r e t h e a c c i d e n t h a d o c c u r r e d . T h e C h i c a g o e x p e r i e n c e

w a s t h u s a w a r m a n d p e r s o n a l o n e a s w e l l a s a n i n t e l l e c t u a l f ea s t.

W o r ld W a r I I ca m e a s e x p ec t ed a n d t h e n e c e s si ti e s d e m a n d e d g r a d u a -

t i o n , i n t e r n s h i p , a n d r e s i d e n c i e s . I p a s s e d u p t h e o p p o r t u n i t y t o r e c e i v e a

P h .D . i n p h y s io l og y a l t h o u g h I h a d p a s s e d a ll t h e n e c e s s a r y e x a m i n a t i o n s .

T h i s w a s n o t im e t o p u r s u e t h e b a s i c r e s e a r c h n e c e s s a r y t o f i n is h a th e s is .

M y lif e a s a n e n t h u s i a s t i c , u n o r t h o d o x , a n d b r a s h y o u n g m a n h a d t o g i ve

w a y t o l i f e a m o n g m e d i c a l m e n , t h o s e a r c h c o n s e r v a t i v e s w h o t h r e w

S e m -

m e l w e i s s

o u t o f t h e i r p r o f e ss i o n f or s h o w i n g t h a t t h e y w e r e i n f e c t i n g m o t h -

e r s i n t h e h o s p i t a l d u r i n g c h i l d b i r t h a n d p u t P a s t e u r t e m p o r a r i l y i n t o a

j a i l f o r a d m i n i s t e r i n g l o c k j a w v a c c i n e t o a b o y w h o h a d b e e n b i t t e n b y a

r a b i d dog .

W h a t s u s t a i n e d m e d u r i n g t h o se y e a r s ( 1 9 4 1 - 1 9 4 8 ) o f t h e p r a c t ic e o f

m e d i c in e a n d s u r g e r y w e r e t h e r e w a r d i n g e x p e r i en c e s w i t h p a t i e n t s w h i c h

m a d e u p t h e p r a c t i c e a n d t h e c o n s u m i n g i n t e r e s t i n f i n d i n g o u t h o w t h e

b r a i n w o r k s w h i c h w a s f u e l l e d b y t h e s i g n s a n d s y m p t o m s p o r t r a y e d b y

t h e s e p a t i e n t s . D u r i n g m y e x t e r n s h i p s a n d i n t e r n s h i p I w a s f o r t u n a t e t o

h a v e a s a c o l l e a g u e , m y c o m p l e m e n t , J o s e p h R a n z a h o f f : h e a b h o r r e d n e u -

r ol og y a n d b r a i n s u r g e r y b u t l ov e d th e s m e l ly n e t h e r r e g io n s o f a b d o m i n a l

s u r g er y . T r a d e s o f p a t i e n t s w e r e t h e o r d e r o f t h e d a y :

c h a q u e u n a s o n g o u t .

I ro n ic a ll y , d u r i n g h i s m i l i t a r y s t i n t R a n z a h o f f w a s a s s i g n e d t o n e u r o s u r -

g e r y a n d a f t e r t h e w a r , h e b e c a m e a n e m i n e n t l y s u c c e s s f u l , t h o u g h s o m e -

w h a t g r u f f , p r a c t i t i o ne r o f t h e a r t i n N e w Y o r k ci t y ( s ee S h a i n be r g , 1 97 9) .

I n a d d i t i o n t o G e r a r d , t h e U n i v e r s i t y o f C h i c ag o w a s r i c h in o t h e r n e u -

r o sc ie n t is t s. S t e p h e n P o l y a k w a s w o r k i n g o n t h e a n a t o m y o f t h e r e t i n a a n d

v i s ua l s ys t e m . I w a s i n t r i g ue d b y t h e w o r k o f R o a f ( 1 927 , 1 9 30) o n co lo r

a f t e r i m a g e s a n d s a w i n P o l y a k ' s d e t a i li n g o f t h r e e s o r t s o f r e t i n a l b i p o l a r

c e l l s a m e c h a n i s m f o r a n a l y z i n g a n d f u r t h e r s e p a r a t i n g t h e H e l m h o l t z i a n

r e c e p to r p r o c e s s , a c c o u n t i n g f o r t h e e f fe c ts o f c o lo r a f t e r i m a ge s . I w r o t e up

t h e s e s u g g e s t io n s w i t h P o l y a k 's h e lp a n d s u b m i t t e d t h e r e s u l t a s a n u n p u b -

l i sh e d m e d i c a l s t u d e n t t h e s is .

P a u l W e i s s w a s t r a i n i n g R o g e r S p e r r y t o t r a n s p l a n t l i m b s o f

A m b l y s -

t o m a .

I b e c a m e w e ll a c q u a i n t e d w i t h b o t h o f t h e m w h e n W e is s a p p e a r e d o n

m y m e d i c a l s er v i ce d u r i n g m y i n t e r n s h i p . T h e f r i e n d s h i p s l a s t e d a l if e t im e

a n d c e n t e r e d o n th e p r o b l e m o f r e s o n a n c e : H o w c o u ld it b e t h a t a li m b

i n d u c e s i n t h e d e v e l o p i n g n e r v o u s s y s t e m a co d e t h a t a ll o w s t h e s y s t e m t o

r e c og n i z e t h e l im b i r r e s p e c t i v e o f i ts i n n e r v a t i o n ? S p e r r y ' s a n s w e r t o t h is

que s t i o n i nvo ke d s pe c i f i c c h e mi c a l c o de s ; mi ne , s ugge s t e d i n

L a n g u a g e s o f

t h e B r a i n ,

de vo l ve s o n t h e f i nd i ng by J .Z . Y o ung o f t h e i nd uc t i o n o f s pec if ic

ne r ve f i be r s i ze s pe c t r a by e a c h mus c l e . Mo s t l i ke l y t h e s pe c i f i c c h e mi s t r y

ind uc es spec if ic f ibe r s ize spe c t ra .

A . E a r l W a l k e r b e c a m e c h i e f o f n e u r o l o g i c al s u r g e r y w h e n P a u l B u c y

l ef t; I l e a r n e d t h e d e t a i l s o f t h a l a m i c a n a t o m y f r o m W a l k e r b e f or e j o i n i n g

B u cy . O v e r th e y e a r s t o co m e , t o g e t h e r w i t h K a o L i a n g C h o w a n d w i t h t h e


16/49

316 K a r l H . P r i b r a m

help of Jerzy Rose at Johns Hopkins University, I extended Walker's ana-

tomical resea rch to complete a classification ofthal amocor tica l connectivity.

Also during this period, Ward Halstead introduced me to what we now call

neuropsychological procedures, which are used to study the effects of bra in

injury in humans.

But most important to my future were Heinrich Klfiver and Paul Bucy,

pioneers in investigations of the functions of the temporal lobe of the brain.

In 1942, I became Bucy's first resident when he moved to the nearby Chi-

cago Memorial Hospital and wrote up our first 100 brain operat ions in order

to have the r esidency accredited. Bucy was editing a volume on the precen-

tral motor cortex at the time and I became privy to the controversies and

details of explorations of this research, as well as learn ing the techn iques

of surger y from a master.

My time with Bucy was exciting and fabulously enriching. Bucy would

tell stories as we made rounds. He had started in general practice and had

found that his patients were in fact patient and loyal even when he made

mis tak es or had to bumble t hrou gh because of his limited experience. What

counted, he found, was that he was really trying and that he was totally

honest with his patients and their families. On another occasion he re-

counted that, while in general practice, he had visited a mental hospital

only to find that almost all the patients were sedated with bromides. He

ordered the patien ts to be tak en off the drug. Within a fortnight more than

half of them were well enough to be considered for discharge. (We don't use

bromides today, but how will our current drugging practices be evaluated

by another generation?)

Most of all, Bucy taught me how to localize brain tumors and, in the

course of this, to learn about the localization of brain functions. I read avidly

during the few quiet moments while on emergency duty, including the book

Bucy had published with Buchanan on intracranial tumors in infancy and

childhood, and the section on brain tumors he wrote for Roy Grinker' s text-

book on neurology. In the section on treatment (p. 621), I saw once again

(as I had been t augh t in obstetrics) the admonition "we must follow the age-

old rule of surgery, p r i m u m n o n n o c e r e , and curb our enthusiasm to the

point where op timum result s in length of life, comfort, and happiness are

attained."

It was also the time I became acquainted with Percival Bailey's treatise

on intracranial tumors. After carefully and beautifully reviewing the evi-

dence, he unequivocally states (p. 69): "I merely wanted to impress upon

you that in the human brain the parts are not equipotential and that even

the defect of intelligence does not, as is somet imes sta ted (261), depend only

upon t he qua nti ty of cerebral t issue removed or destroyed." Reference (261)

is to Karl Lashley's 1929 monograph B r a i n M e c h a n i s m s a n d I n te ll ig e n ce ,

which I manage d to purch ase at a second-hand book store for a dime. Lash-

ley later became a major influence in my life and the tension between his


17/49

K a r l H . P r i b r a m 3 1 7

views and those I inheri ted from Bailey and Bucy formed the thr us t of my

research career.

Bailey could make his summary statement despite that in the text

(p. 67) leading up to it he had to re ma rk that:

The ana tomical correlates of such relatively simple functions as

sensation and voluntary motion are somewhat familiar to us.

We know also th at the centra l mechanism of the more compli-

cated function of language is usua lly c lustered closely around

the left lateral fissure, but when we attempt to discuss a higher

mental function such as intelligence, we are greatly hampered

by lack of consistent data. Yet certain areas of the b rain are

known, injury to which is peculiarly liable to disturb intelli-

gence. One of these is the left suprama rgi nal gyrus. Another is

the anterio r pa rt of the frontal lobe, although in this case the

disturbance of charac ter is predo minan t and I should be less

willing to indicate the exact area involved. It is significant that

these pa rts are just the ones in the hu ma n brain which are most

developed beyond those present in the higher apes.

Later, after I had been given techniques to study such general concepts

as intelligence and character by Karl Lashley, I made it my research busi-

ness to pin down more precisely the localization of the brain /behavior rela-

tions entailed. Only much later did I begin to understand what Lashley

meant by his dicta regarding equipotentiality and mass action in the stor-

age and retri eval of memories and in the processing of equivalence in per-

ceptions and actions.

In 1943, Bucy was editing a volume on the motor systems of the bra in.

I was privy to th at editing, chapter by chapter, as Bucy explained to me his

views and criticisms of wha t had been submitted. I found out th at Wilder

Penfield, Warren McCulloch, and Dus ser de Bar enne all tho ugh t of the

cerebral motor cortex more as a sensory cortex for movement than as the

final common path for all cortical activity, a view that Bucy shared. I

lea rned of how much scholarly activity goes into the wri ting and edi ting of

such a volume, the great care to provide the best currently available access

to knowledge.

My turn to become scholarly came when we admitted a 54-year-old

Greek woman who complained of bouts of twitch ing accompanied by local-

ized sweat ing over the left side of the face. While in the hospital she actua lly

experienced a grand mal epileptic seizure accompanied by sweating and by

flushing. When during surgery we found a small oligodendroglioma in the

precent ral motor cortex, a tumor which was readily removed with the result

of a complete cure for the patient, I suggested that we had come upon a

most important finding. Everyone in neurology

k n e w

that control over the


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318

Karl H. Pribram

autonomic nervous system was exercised by no higher station than the

hypothalamus. Cortical control would mean that the system was not as

autonomous, or automatic, as we had been taught to believe. But here was

a patient whose cortical tumor had produced epilepsy accompanied by lo-

calized sweat ing and flushing, definitely due to excitation of the autonomic

nervous system. I asked Bucy if this observation was worth publishing and

he agreed that, indeed, it was. I was eager to get something into print. I

was a lready 24 years old and most of my forebears had published in thei r

early 20's. I was about to become the laggard in the family.

The paper was accepted for publication in the Archives of Neurology &

Psychiatry (Bucy and Pribram, 1943) and Bucy received a letter from Earl

Walker that the Chicago Medical Society wished to have it presented at

their next meeting. Bucy showed me the letter and said "you do it." I did.

The other speak er tha t evening was Warren McCulloch, head of the re-

search t eam at the Neuropsychiatric Instit ute of the University of Illinois.

I did not und ers tan d a word of what he was trying to tell us and nei ther did

anyone else. It took me another 40 year s of listen ing to McCulloch (who was

at Mass ach use tts Inst itu te of Technology (MIT) when I was at Yale) and

discussion before I was able to grasp the "cybernetic" ideas that were to

tran sform our unders tan din g of the way the nervous system operated. Two

decades later, I was offered the headsh ip of the r esea rch t eam a t the Uni-

versity of Illinois. It was a most gratifying offer but, by then, I was en-

sconced in a most productive laboratory at Stanford and could not see

myself free to move.

A second research endeavor stemmed from the results of the surgery

performed on this interesting woman: I noted th at careful removals of cor-

tical tissue that minimally invaded white matter left the patient with no

perceptible aftereffects. During the 1950s, when Lawrence Weiskrantz was

a graduate student in my laboratory, discussing this insight led to his life-

long pursuit of careful remova ls of visual cortex and the devising of infi-

nitely sophisticated te sting procedures to determine the extent of residual

vision; these experiments resu lted in his discovery of blind-sight, the ability

to perform visual ta sks without conscious awaren ess of the visual stimuli

involved.

Within a few months of joining Bucy, I was so completely caught up in

neurosurgery (while still attending to all the other services in the small

hospital) t ha t I made a decision to pur sue the study of the nervous system,

as a neurologist, a psychiatrist, or a neurosurgeon. I had never been good

with my hands so I asked Bucy to tell me, after some months, whether I

could make it as a surgeon. His answer came in typical Bucy fashion. One

day he said, "Next month I am going on vacation and turning my practice

over to you." I asked if th at mean t th at he wan ted me to do the surgery. "Of

course," he said. That was all. I had set up a woodworking shop in my home

and had practiced using my hand s with the aid of machine shop workers


19/49


319

w h o w e r e m y n e i g h b o r s . A l l m y p a t i e n t s a n d I s u r v i v e d t h e m o n t h . T h a t

w a s i t; I b e c a m e a n e u r o s u r g e o n . B u c y a r r a n g e d f o r m e t o h a v e a r e s i d e n c y

w i t h E r i c O l d b e r g a t t h e n e i g h b o r i n g S t. L u k e ' s h o s p i t a l i f I w a n t e d i t.

O l d b e r g w a s h e a d o f t h e U n i v e r s i t y o f I ll in o is N e u r o p s y c h i a t r i c I n s t i t u t e

w h e r e P e r c i v a l B a i l e y , G e r h a r d t v o n B o n i n , a n d W a r r e n M c C u l l o c h w e r e

p u r s u i n g t h e i r o w n r e s e a r c h . I w a s t o b e p a r t o f t h i s t e a m .

T h u s , a f t e r m y y e a r w i t h B u c y , I b e c a m e O l d b e r g ' s r e s i d e n t ( a n d a l s o

t o o k o n t h e r e s i d e n c i e s i n n e u r o l o g y a n d p s y c h i a t r y f o r e x t e n d e d p e r i o d s

w h e n n e c e s s a r y b e c a u s e o f t h e w a r ) w i t h p r i v i l eg e d a c c e ss to t h is g r o u p .

B a i l e y t o o k o n a n o t h e r r e s i d e n t , J o h n G r e e n , a n d B a i l e y s a t w i t h u s o v e r a

s i x - m o n t h p e r io d d e t a i l i n g t h e h i s t o r y o f h i s t u t e l a g e w i t h H o r t e g a d e l Rio ,

w h o s e m e t h o d s a n d n e u r o e m b r y o l o g i c a l a p p r o a c h l ed t o B a i le y ' s p i o n e e r i n g

w o r k o n th e c l as s if i ca t io n o f b r a i n t u m o r s . E a c h s t o r y w a s i l l u s t r a t e d w i t h

m i c r o s c o p i c m a t e r i a l s e c t i o n e d f r o m b r a i n t u m o r s w h i c h w e e x a m i n e d t o -

g e t h e r i n g r e a t d e t a i l.

I o c c a s i o n a l l y p a r t i c i p a t e d i n t h e t h e n - o n g o i n g s t r y c h n i n i z a t i o n e x p e r -

i m e n t s o f c h i m p a n z e e c o r t ex a n d l i s t e n e d a t t e n t i v e l y t o B a il ey , v o n B o n i n ,

a nd Mc C ul l o c h d i s c us s t h e r e s u l t s . S o me ye a r s l a t e r , a t Y a l e U n i ve r s i t y , I

w a s a b l e t o p u t t o g o o d u s e m y s u r g i c a l s k i l l s a n d t h e k n o w l e d g e I h a d

a c q u i r e d f r o m t h e s e d i s c u s s i o n s t o c o m p l e t e t h e c h e m i c a l s t i m u l a t i o n e x -

p e r i m e n t s o n ca t a n d m o n k e y b y e x p l or a t io n s o f t h e m e d i a l a n d b a s a l s u r-

f a ce s of t h e b r a i n , w h i c h h a d r e m a i n e d i n a c c es s ib l e to t h e e a r l i e r r e s e a r c h .

A m o s t e x c i ti n g p a r t o f t h e r e s e a r c h g o i n g o n a t t h i s t i m e w a s t h e e x p lo -

r a t i o n o f t h e l a t e r a l s u r f ac e o f t h e h u m a n b r a i n f or s u p p r e s s i o n o f m o t o r

a c t i v i t y . A l t h o u g h t h e r e s u l t s o b t a i n e d w e r e h i g h l y c o n t r o v e r s i a l , t h e p r o -

c e ss o f c o rt ic a l s t i m u l a t i o n i n w h i c h B u c y a ls o p a r t i c i p a t e d , t h e e x a m i n a -

t i on o f t h e p a t i e n t ( s o m e t i m e s l e ft t o m e ) w h i l e t h is s t i m u l a t i o n w a s i n

p r o g r e s s , a n d t h e d i s c u s s i o n s w h i c h e n s u e d w e r e f a s c i n a t i n g . I r e m e m b e r

w e l l t h e o c c a si on d u r i n g o n e of t h e s e p r o c e d u r e s w h e n a t e l e g r a m a r r i v e d

f r om P a u l G l e es a t O x f o rd U n i v e r s i t y s t a t i n g t h a t h e h a d j u s t f o u n d n e r v e

f i b e r s c o n n e c t i n g t h e p r e c e n t r a l c o r t e x t o t h e c a u d a t e n u c l e u s , u s i n g h i s

n e w l y d e v e l o p e d s i l v e r s t a i n i n g t e c h n i q u e . M c C u l l o c h s u g g e s t e d t h a t t h e

t e r m nega t i ve f eedback b e a p p l i e d t o e x p l a i n t h e s u p p r e s s i o n o f m o t o r a c-

t i v i t y a n d t h a t G l e e s h a d f o u n d t h e a n a t o m i c a l b a s i s f o r s u c h f e e d b a c k .

K n o w l e d g e o f t h e s e f e e d b a c k c i r cu i ts , i n c o n j u n c t io n w i t h t h o s e o p e r a t i n g

o n t h e s p i n a l r ef le x, w e r e to p r od u c e t h e T e s t - O p e r a t e - T e s t s e q u e n c e as a

f u n d a m e n t a l p r o c e d u r e o p e r a t i n g i n th e f o r m a t i o n o f P l a n s i n P l a n s a n d

t he S t r u c t u r e o f B e h a v i o r (M il ler e t a l ., 1960) .

T h e U n i v e r s i t i e s o f C h i c a go a n d I l l ino i s w e r e no t t h e o n l y c e n t e r s f o r

n e u r o s c ie n c e r e s e a r c h i n C h i c ag o a t t h e t im e . H o r a c e M a g o u n a n d D o n a l d

L i n d s l e y a n d t h e i r c o l l a b o r a t o r s w e r e b e g i n n i n g t h e i r r e s e a r c h o n t h e m e s -

e n c e p h a l i c r e t i c u l a r f o r m a t i o n a t N o r t h w e s t e r n U n i v e r s i ty . I w a s t o p a r t ic -

i pa t e i n t h i s w o r k i n c o l l a bo r a t i o n w i th Pe r c i va l B a i l e y , h a v i ng r e c e i ve d a

f e l l o w s h i p t o do s o , bu t B a i l e y c h a nge d h i s p l a ns a nd w e n t o ve r s e a s f o r a


20/49


year. The proposed collaboration never took place, but my interest in the

project had been piqued so that I kept abreast of developments as they

occurred.

Exciting as all of these Chicago experiences were, they did not furnish

me wi th some of the basic tools I needed to accomplish my goals, which were

to explore the relationship between brain function and mental processes

such as emotion, cognition, and conation (the intention to act). In my search

for a hay fever-free location where I might ear n my living as a neurosur geon

and at the same time pursue these goals, I hea rd of the Yerkes Laboratories

of Primate Biology near Jacksonville, Florida, where Karl Lashley was di-

rector. Fortunately, there was a position open in Jacksonville with J.G.

Lyerly. Lyerly, as well as Poppen in Boston, had devised a superior incision

for frontal lobotomy which was safer than the classical (lateral) Fr ee ma n-

Watts procedure and left fewer unwanted side effects. The lateral incision

was shown by Fred Mettler and L. P. Rowland to invade Broca's speech

area. Although no language disturbances followed the lateral incision, fi-

bers from the medial and orbital cortex were more apt to be saved when

Lyerly's superior incision was used. Because of his innovative bent, I felt

th at Lyerly would be sympathet ic to my desire to work at Yerkes. In 1946, I

took my Florida Stat e Board examinations and began private practice.

Lyerly agreed to my working two half-days per week, plus any free time,

at my research at Yerkes. I called Lashley and he responded favorably,

stating that he had been looking for a neurosurgeon to assist him in his

primate research. Thus began a collaboration which was to prove most in-

fluential in shaping my subsequent research program.

Lash ley ta ught me the t echniques of experimental psychology, a field of

inquiry which I did not know existed. Paradoxically, although Lashley was

almost solipsistic, destructive in his research procedures and interpreta-

tions of any finding that would relate brain function to behavior, he pro-

vided many of the questions t hat needed to be answered and t hat led to

the discoveries which make up the substance of my career. Some of the

discoveries I made while he was still alive, such as the unique relationship

between the frontal cortex and the limbic forebrain, and the sensory speci-

ficity of various sectors of the posterior "association" cortex. He ignored or

played down these results, as they were contrary to his belief that the

mechanisms involved in organizing complex psychological processes were

distribu ted in the brain. But always, his critical wit sharpened my interpre-

tations and provided the basis for further observation and experiment.

The opportunity to work full time in resea rch came in 1948 when I was

asked by John Fulton to join him the department of physiology at Yale

University. My association with Yale lasted for a decade (1948-1958), dur-

ing which time I also directed the resea rch l aboratories of the Inst itu te of

Living, a mental hospital in nearby Hartford, Connecticut. The facilities at


21/49

Karl H. Pribram

321

Y a le a n d i n H a r t f o r d p r o v i d e d a m p l e s p a c e fo r a g r o u p o f y o u n g i n v e s t i g a -

t o r s d e d i c a t e d t o e x p l o r i n g t h e p o w e r o f c o m b i n i n g t h e t e c h n i q u e s o f e x p e r -

i m e n t a l p s y ch o lo g y w i t h t h o s e o f n e u r o p h y s i ol o g y a n d e x p e r i m e n t a l

n e u r o s u r g e r y . D o c t o ra l s t u d e n t s f r o m Y a le ( M u r i el B a g s h a w , M a r t h a H e l -

s o n W i l s o n ) , H a r v a r d ( L a w r e n c e W e i s k r a n t z ) , M c G i l l ( M o r t i m e r M i s h k i n ) ,

U n i v e r s i t y o f C a l i f o r n i a a t B e r k e l e y ( W i ll ia m W i ls on ) , a n d S t a n f o r d ( Je -

r o m e S c h w a r t z b a u m ) f o r m e d t h e n u c l e u s o f a m o s t p r o d u c t iv e t e a m , a ll of

w h o m r e ce iv e d t h e i r d e g r e e s w h i le w o r k i n g o n t h e p r o g r a m .

D u r i n g t h i s p e r i o d I s p e n t o n e m o n t h a y e a r a t t h e Y e r k e s L a b o r a t o r y ,

a n d K a o L i a n g C h ow , a n e a r l y s t u d e n t a n d c o l la b o r at o r, s p e n t a m o n t h w i t h

m e i n t h e n o r t h , r e e s t a b l i s h i n g a t l e a s t i n p a r t Y e r k e s ' o r i g i n a l v i s i o n , a

Y a l e U n i v e r s i t y - r e l a t e d p r i m a t e r e s e a r c h l a b o r a t o r y . T h i s c o n t i n u i n g c o l -

l a b o r a t i o n l ed t o a n i n v i t a t i o n t o su c c e ed L a s h l e y a s d i r e c t o r o f t h e l a b o r a -

to r ie s , a n d I f il le d t h i s p o s t un t i l t h e p r e s i d e n t o f Y a l e U n i v e r s i t y s o ld t h e

l a b o r a t o r i e s t o E m o r y U n i v e r s i t y i n A t l a n t a i n 1 95 6 .

A l so d u r i n g t h i s p e r io d , I b e g a n a n i n t i m a t e a s s o c ia t i o n w i t h p s y c ho lo -

g i s t s a t H a r v a r d U n i v e r s i t y . I t a u g h t s u m m e r s c h o o l t h e r e o n e y e a r , b u i l t

o p e r a n t e q u i p m e n t i n t h e H a r v a r d s h o p s , a n d l e a r n e d a g r e a t d e a l f r o m

S .S . S t e ve n s , G a r y B o r i ng , a n d G e o r g vo n B e ke sy . O nc e a m o n th , B e r t R o s -

n e r a n d I d r ov e u p to H a r v a r d ( a n d l a t e r M I T ) t o p e r f o r m e x p e r i m e n t s w i t h

W a l t e r R o s e n b l i t h o n m o n k e y s i n w h i c h w e e v o k e d e l e c t r i c a l p o t e n t i a l s i n

t h e c o r t e x b y a u d i t o r y s t i m u l a t i o n . S o m e w h a t l a te r , t h e s e s e s s i o n s w e r e

e x t e nd e d t o e xp l o re , w i th W o l f ga ng K o hl er , t h e e vo c a t i o n o f d i r e c t c u r r e n t

s h i f t s unde r s i mi l a r c o nd i t i o ns .

My i n t e r a c t i o ns w i th B . F . S k i n ne r a t H a r v a r d w e r e e s pe c ia l ly m e m o r a b l e

a n d l ed to a d e ca d e o f p r i m a t e o p e r a n t c o n d i t i o n i n g e x p e r i m e n t s , w h i c h

de ve l o pe d i n to s ubs e que n t r e s e a r c h i n c o gn i t i ve ne u r o ps yc h o l o gy . S h o r t l y ,

I w a s a b l e to a u t o m a t e a n d e x t e n d t h e o p e r a n t e q u i p m e n t t o re c o rd ( in c lu d -

i n g r e a c t i o n t i m e ) t h e r e s u l t s o f i n d i v i d u a l c h oi ce s a m o n g a d o z e n p o s si b le

p a n e l p r e s s es . L a t e r , o v e r m y t h r e e d e c a d e s a t S t a n f o r d ( 1 9 5 9 - 1 9 8 9 ) , t h e s e

r e s p o n s e s w e r e r e c o r d e d i n a l a r g e v a r i e t y o f p r o b l e m - s o l v i n g s it u a t i o n s .

T h e c o m p u t e r - c o n t r o l l e d t e s t i n g a p p a r a t u s w a s d u b b e d D i s c r i m i n a t i o n

A p p a r a t u s f or D i s c r e t e T r i a l A n a l y s i s ( D A D T A ).

A t o n e p o i n t in o u r i n t e r a c t io n , S k i n n e r a n d I ca m e to a n i m p a s s e o v e r

t h e p o s s ib l e m e c h a n i s m i n v o lv e d in t h e c h a i n i n g o f r e s p o n s e s . C h a i n i n g

w a s d i s r u p t e d b y r es e c t io n s o f t h e f a r f ro n t a l c o rt ex . S k i n n e r s u g g e s t e d t h a t

p r o p r i o c e p t i v e f e e d b a c k m i g h t h a v e b e e n d i s r u p t e d , b u t t h i s h y p o t h e s i s

w a s n o t s u p p o r t e d b y m y e x p e r i m e n t s . F u r t h e r m o r e , a s I i n d i c a te d t o S k in -

ne r , he , a s a Ph .D. in b io logy , could propose such an hypo thes i s , bu t I , a s a

l oy a l S k i n n e r i a n , h a d t o s e a r c h e l se w h e r e t h a n w i t h i n t h e " b l ac k b ox " f or

a n a n s w e r t o o u r q u e s t i on . G e o r g e M i l le r o v e r h e a r d s o m e o f o u r d i s c u s si o n s

a n d p o i n t e d o u t to m e t h a t h e h a d a v a i l ab l e a p r o c e d u r e t h a t m a d e c h a in -

i n g o f r e s p o n s e s e a sy : a c o m p u t e r p r o g r a m . M i ll e r e x p l a i n e d to m e t h e


22/49

322

Karl H. Pribram

p r i n c ip l e s o f l is t p r o g r a m m i n g w h i c h h e h a d j u s t l e a r n e d f r o m H e r b e r t S i m o n

a n d A l a n N e w e ll . T h e c u l m i n a t i o n o f t h e c o ll a b o r at i on b e g u n b y t h e s e

e n c o u n t e r s i n t h e h a ll s o f H a r v a r d w a s

Plans and the Structure of Behavior,

a b o o k i n f l u e n c e d a l s o b y o u r i n t e r a c t i o n s w i t h J e r o m e B r u n e r , w h o h a d

o r g a n i z e d a c o n f e r e n c e o n t h i n k i n g a t C a m b r i d g e U n i v e r s i t y i n 1 9 5 6 t o

w h i c h w e h a d b e e n i n v i te d . T h e b o ok w a s w r i t t e n i n 1 9 5 8 - 1 9 5 9 a t t h e

C e n t e r f or A d v a n c e d S t u d i e s i n t h e B e h a v i o r a l S c i en c es , a d j a c e n t t o t h e

c a m p u s o f S t a n f o r d U n i v e r s it y .

T h a n k s t o J a c k H i l g a r d a n d R o b e r t S e a r s o f t h e p s y c h ol og y d e p a r t -

m e n t , a n d t o T o m G o n d a ( t h e s on o f a n e u r o l o g i s t w h o h a d b e e n a f ri e n d o f

m y f a m i l y i n V i e n n a ) i n p s y c h ia t r y , I w a s g i v e n a n a p p o i n t m e n t a t S t a n f o r d

( s u p p o r t e d , i n it ia ll y , b y a g r a n t f r o m t h e M a r k e l F o u n d a t i o n f or So c ia l R e -

s e a r c h ) i n 1 9 59 . S o o n a f t e r w a r d , i n 1 9 62 , I r e c e i ve d a l i f e t i me r e s e a r c h

c a r e e r a w a r d f r o m t h e U .S. N a t i o n a l I n s t i t u t e s o f H e a l t h w h i c h , i n a d d i t i o n

t o s u b s t a n t i a l g r a n t s t o p u r s u e r e s e a r c h i n t er e s t s, s u s t a i n e d m e f or t h e

n e x t t h r e e d e c a d es .

A t S t a n f o r d , a n o t h e r g r o u p o f d o c t o ra l a n d p o s t d o c t o r a l a ss o c i a te s

jo i ne d t h e s e e nde a v o r s . ( A l to ge the r , s o me f i ft y do c to r a l a nd f if ty po s tdo c-

to r a l f e l l o w s w e r e t r a i ne d i n t h e ne u r o ps yc h o l o g i c a l l a bo r a to r i e s a t Y a l e

a n d S t a n f o r d u n d e r m y d i r e c t i o n . ) A t S t a n f o r d , R o b e r t A n d e r s o n , M u r i e l

B a g s h a w , B r u c e B r i d g e m a n , J a m e s D e w s o n , R o b e r t D o u g l a s , D a n i e l K i m -

b a ll , A b r a h a m S p e v ac k , a n d L e sl ie U n g e r l e i d e r w e r e a m o n g t h o se w h o

m a d e m a j o r c o n t r i b u t i o n s . N i c o S p i n e l l i b e c a m e a n i n t e g r a l a n d a l m o s t

i nd i s pe ns a b l e c o l l a bo r a to r .

W h e n I b e c a m e e m e r i t u s a t S t a n f o r d a t a g e 7 0, I w a s o f f er e d t h e o p p o r-

t u n i t y t o c o n t i n u e w o r k a t R a d f o r d U n i v e r s i t y i n V i r g i n i a . R a d f o r d , s i s t e r

u n i v e r s i t y t o V i r g i n i a T e c h , b u i l t a l a b o r a t o r y f o r m e , a n d I o r g a n i z e d a

C e n t e r f o r B r a i n R e s e a r c h a n d I n f o r m a t i o n a l S c i e n c e s ( B . R . A . I . N . S . ) w i t h

t h e h e l p o f A l a s t a i r H a r r i s , w h o c h a i r s t h e p s y ch o lo g y d e p a r t m e n t . T h e

a p p o i n t m e n t is s u p p o r t e d b y t h e e m i n e n t s c ho l ar s f u n d o f t h e C o m m o n -

w e a l t h o f V i r g i n ia a n d a n e n d o w m e n t f r om t h e J a m e s P. a n d A n n a K i n g

Fo unda t i o n . I de ve l o pe d a c l o s e a nd e f f e c t i ve c o l l a bo r a t i o n w i th J o s e ph

K i n g , w h o o b t a i n e d h i s P h. D . a t n e i g h b o r i n g V i r g i n i a T ec h u n d e r t h e d i re c-

t i o n o f A b e S p e v a c k , w h o h a d s p e n t s e v e r a l y e a r s w i t h m e a t S t a n f o r d a s a

po s td oc to ra l f el low.

R e s e a r c h T h e m e s

T h e r e s u l t s o f t h e r e s e a r c h c o m p l e t ed o v e r th e y e a r s a t t h e Y e r k e s la b o r a -

to r i e s , a t Y a l e , a t S t a n f o r d , a nd c u r r e n t l y i n V i r g i n i a , c a n be o r ga n i ze d

a c co r d in g t o o v e r l ap p i n g t h e m e s , e a c h t h e m e r e p r e s e n t i n g a p r o b le m a r e a

a n d t h e a p p l i c a t i o n o f t e c h n i q u e s a p p r o p r i a t e t o t h a t p r o b l e m a r e a . A d e -

s c r i p t i o n o f t h e t h e m e s f ol lo w s.


23/49

Kar l H . Pr ib r am 323

Theme I : Es ta b l i s h in g a Cor r e la t ion be tween Br a in Sys te ms and Spec if ic

Behav io r a l Ind ica to r s

B y t h e t i m e m y r e s e a r c h p r o g r a m b e g a n , l a rg e a r e a s o f t h e p r i m a t e c o rt ex

r e m a i n e d u n e x p l o r ed b y e x p e r i m e n t a l i n v es ti g at io n . I n h u m a n s , d a m a g e

t o t h e s e a r e a s r e s u l t e d i n a g n o s i a , a p h a s i a , a n d c h a n g e s i n c h a r a c t e r , a n d

t h u s i n i n t e r p e r s o n a l e m o t i o n a l i n t e ra c t io n s . B u t i t w a s n o t k n o w n w h e t h e r

t h e s e c h a n g e s i n c o m p e t e n ce a n d b e h a v i o r w e r e t h e r e s u l t o f d a m a g e a d d i-

t i o n a l t o t h a t i n f l i c t e d o n p r i m a r y s e n s o r y - m o t o r s y s t e m s , o r w h e t h e r t h e

c h a n g e s c o ul d o cc u r w i t h o u t s u c h d a m a g e . F u r t h e r m o r e , i t w a s n o t k n o w n

w h e t h e r t h e c h a n g e s w e r e s p e c i f i c t o o n e o r a n o t h e r l o c a t i o n w i t h i n t h e

s i l e n t ( kno w n a s t h e "a s s o c i a t i o n" ) c o r t e x .

B y u s i n g a b a t t e r y o f b e h a v i o r a l t e s t s a n d r e s e c t i n g l a r g e p a r t s o f t h e

t h e n - s i l e n t c o r te x of m o n k e y s w i t h o u t i n v a d i n g t h e p r i m a r y s e n s o r y - m o t o r

s ys t e ms , I f o und a ns w e r s t o t h e s e que s t i o ns r e l a t i ve l y ra p id l y . A m e th o d w a s

d e v i s e d w h i c h u s e d s u p e r i m p o s i t i o n s o f r e c o n s t r u c t i o n s o f r e s e c t e d c o r te x .

T h e n u m b e r o f t h e r e s e c t i o n s t h a t p r o d u c e d a p a r t i c u l a r b e h a v i o r a l d e fi ci t

w e r e s um m e d . T h e s um o f t h e r e s e c t i o ns w h i c h p r o d uc e d no de f ic it w e r e a l so

s u m m e d a n d t h e r e s u l t w a s s u b t r a c t e d f r o m th e s u m o f l es io n s t h a t p r o -

duc e d a def ic it . T h i s " i n t e r c e p t o f s um s " t e c h n i qu e w a s t h e o r i g i n o f t h e

"do ub le d i s so c ia t io n" t e c h n i qu e no w u s e d s o e x t e ns i ve l y i n c li n ic a l ne u r o p s y -

c h o l o g y a n d a l l o w e d m e t o m a k e m u l t i p l e d o u b l e d i s s o c i a t i o n s a m o n g t h e

va r iou s de f ici ts p roduc ed by the re sec t ions and to local ize the bra in sys tem in-

v o lv e d in t h e b e h a v i o r r e p r e s e n t e d b y e a c h t a s k ( r e v ie w e d b y P r i b r a m , 1 97 5) .

T h e r e s u l t s I o b t a i n e d a t Y ale i n t h e e a r l y 1 9 5 0 s w e r e u n e q u i v o c a l . O n e

t y p e o f d e fi ci t w a s p r o d u c e d w h e n t h e a n t e r i o r f r o n t a l , t h e c i n g u l a t e a n d

h i p p o c a m p a l c o rt ex , a n d t h e a m y g d a l a a n d a n t e r i o r t e m p o r a l c o rt ex w e r e

r e s e c t e d . A no th e r t yp e o f de f ic it f ol lo w e d r e s e c t i o ns o f t h e po s t e r i o r c o r t ic a l

c o n v e x i ty a n d c o u ld b e f u r t h e r s u b d i v i d e d i n t o s e n s o ry - s p ec if ic c o m p o n e n t s ,

e a c h o f w h i c h w a s r e l a t e d t o i t s o w n p o r t i o n o f t h e c o nve xa l c o r te x . I n no

i n s t a n c e d id i n v a s io n o f t h e a d j a c e n t p r i m a r y s e n s o r y - m o t o r s y s t em s p ro -

d u c e t h e d e f i c i t o r e v e n e n h a n c e i t . T h e s e f i n d i n g s w e r e p u b l i s h e d i n t h e

Jour n a l o f Com par a t i ve Neur o logy a n d Jour n a l o f Com par a t i ve and Phys i -

ological Psychology i n t h e e a r l y 1 9 50s , r e v i e w e d i n 1 9 54 i n Cur r en t Tr ends

in P sychology ( P r i b r a m , 1 9 5 4 ) , a n d r e p r i n t e d i n Behaviora l Sciences (see

P r ib ra m , 1969 , Vol. I) .

Theme H: De ter min ing the Behav io r a l Ca tegories Deno ted

by the Ind ica tors

H a v i ng i de n t i f i e d s pec if ic be h a v i o r a l i nd i c a to r s f o r t h e f un c t i o ns o f s pec if ic

a r e a s o f t h e c o rt ex , t h e n e x t p r o b l e m w a s t o d i s co v e r w h a t t h e i n d i c a t o r s

m e a n t . M u c h a s a B a b i n s k y s i g n s e rv e s a s a n i n d i c a t o r o f i m p r o p e r f u n c-

t i o n i n g o f t h e s p i n a l p y r a m i d a l m o t o r s y s t e m , s i g n s o f m a l f u n c t i o n o f b r a i n

c o gn i t i ve s ys t e ms w e r e no w a va i l a b l e t o u s .


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324 Karl H. Pribram

In order to define the mean ing of the behavioral indicators we had to

explore the effects of each brain resection with a wide range of behaviora l

tas ks related in one way or ano ther to the indicator. Limits were establi shed

by showing which tasks could be performed without any deficit. For ex-

ample, t he visual deficit produced by resections of the inferotemporal cortex

was observed du rin g discriminat ions of color, brightness, size, and two- and

three-dimensional shapes, but not when the animal was tracking even mi-

nute objects. Further, limits to the deficit on the b righ tnes s or size discrim-

ination were obtained when the difference between the brightness or size

of the cues was eithe r very large or very small. (In the la tte r case, normal

controls had as much difficulty discriminating size or brightness as did the

monkeys with brain damage.) I used response operator characteristic

curves (ROC) to check whe the r the deficiency in discr imination was a func-

tion of changes in detection threshold or in response bias.

Inte rpre tat ion was seldom straightforward, despite the wealth of data

accumulated. This was in large part due to the lack of agreement about the

construc ts used in experimental psychology. Just how does one compare the

results obtained in a fixed interval operant conditioning study with a result

obtained in an ROC decisional experiment? How does one compare either

of these with results obtained in a delayed alter nation situation tested in a

Yerkes box or with the DADTA apparatus? Interpretation had to be made

after much cross validation of techniques, often using the same subjects

and, of course, comparable resections. Nonetheless, some 80 publications in

Brain, Jou rna l of Neurophysiology, Jou rna l o f Com parative Neurology,

Jou rna l of Compa rative and Physiological Psychology, and Neuropsycholo-

gia presen ted the resul ts of these investigations, each in the technical lan-

guage appropriate to the behavioral methods used. But in most cases some

conceptual leaps were necessary in making the interpre tations; these leaps

were guided on one han d by findings on hu ma n neuropsychological pati ent s

and on the other by knowledge obtained about the anatomy and physiology

of the neura l systems being investigated.

Them e III: Determ ining the Physiological Processes M ediated

by the System s

Another line of research, which was made possible by the init ial findings of

Theme I, was an analysi s of the ana tomy and physiological processes of the

neu ral systems of which the critical cortical areas were a part. Chemical

and electrical stimulations in anesthetized or problem-solving monkeys

were performed. The effects of such stimulat ions on electrical recordings of

event-related, local field potentials were assessed while monkeys per-

formed in the DADTA. Also, such effects on the micro structure of recept ive

fields of single units in the visual somatosensory, somatosensory, and motor

systems were assayed.


25/49

Karl H. Pribram 325

O n c e a g a i n t h e r e s u l t s o f t h e s e e x p e r i m e n t s y i e ld e d a g o od d e a l o f d a t a

( some 40 pape r s ) , pub l i shed i n the

Journal of Neurophysiology, Brain Re-

search, Experimental Brain Research, Electroencephalography,

a n d

Clinical

Neurophysiology and Experimental Neurology,

w h i c h a r e i n t e r e s t i n g i n t h e i r

own r i gh t . Howeve r , a s i n Theme I I , interpretation ( a n d i n s o m e i n s t a n c e s

controversial interpretation) b e c a m e n e c e ss a r y . O n e m a j o r c o n t r o v e r s y ce n -

t e r s on w he th e r the s enso ry spec if i ci t y o f t he " a s soci a t ion" co r t ex o f the

pa r i e t a l , occ ip i t a l , and t empora l l obes i s due to i t s t r an sco r t i ca l i npu t v i a

c o n n e c t io n s f r o m t h e r e l a t e d p r i m a r y s e n s o r y c o rt ex , o r w h e t h e r t h e s pe ci -

f ic it y i s t o be a sc r ibed to an ou tp u t w h ich ope ra t e s do w ns t r ea m on the p r i -

m a r y s e n s o r y s y s t e m s . I w a s a b le to m a k e m a s s i v e d i s c o n n e ct i on s , s o m e o f

w h i c h a p p e a r e d t o b e co m p le te , b e t w e e n t h e p r i m a r y s e n s o ry s y s t e m s a n d

t h e i n f e r o t e m p o r a l c o r te x i n v o lv e d i n v i s u a l d i s c r i m i n a t i o n s . N o n e o f t h e s e

d i s c o n n e c ti o n s p r o d u c e d l a s t i n g d e f ic it s i n s e n s o r y d i s c r i m i n a t i o n s a n d t h i s

l e d m e t o p r o p o s e t h e o u t p u t h y p o t h e s i s . C o n t r o v e r s y h i n g e d o n w h e t h e r

t h e d i s c o n n e c ti o n s w e r e i n f ac t to t al : e v e n a s m a l l r e m n a n t o f c o n n e c t iv i t y

cou ld poss ib ly be su f f i c i en t t o med ia t e an i npu t . The f ac t s a r e r ev i ewed in

the pap e r " The Ro le o f Cor t i co -co r t ica l Connec t ions" (P r ib ram , 1986a ) .

Theme IV. Relevance of the Research Results to Humans

T h e r e s e a r c h p r o g r a m b e g a n w i t h th e a i m t o cl ar if y t h e b r a i n m e c h a n i s m s

invo lved i n cogn i t i ve , emo t iona l , and cona t i ve ( i nvo lv ing the i n t en t ion to

a ct ) p ro c e s se s i n h u m a n s . T h e f i n al r e s e a r c h p h a s e o f t h e p r o g r a m t h e r e f o r e

h a d t o a d d r e ss t h e r e l e v a n ce o f t h e r e s u l t s o f t h e n o n h u m a n p r i m a t e r e -

s e a r c h , i n w h i c h s o m e 1 5 0 0 m o n k e y s w e r e u s e d , t o h u m a n n e u r o p s y c h o l -

og ica l f i nd ings . S ince my ea r ly days i n the neu rosu rg i ca l c l i n i c , e l ec t r i ca l

r e c o r d in g s o f e v e n t - r e l a t e d s c al p p o t e n t ia l s , c o m p u t e r i z e d t o m o g r a p h y , a n d

n u c l e a r m a g n e t i c r e s o n a n c e i m a g i n g ( M R I ) t e c h n i q u e s h a v e b e e n d e v e l -

oped to a id i n the l oca l i z a t i on o f b r a in pa tho log ica l cond i t i ons. A ma jo r t a sk

a h e a d i s t o c o m p a r e t h e r e s u l t s o b t a i n e d w i t h t h e s e t e c h n i q u e s w i t h t h o s e

o b t a i n e d i n m o n k e y s .

Due to the p rod ig ious advances i n i n fo rma t ion p roces s ing t echno logy ,

r e c o r d i n g s o f t h e r u n n i n g e l e ct r ic a l b r a i n a c t iv i t y s h o w g r e a t p r o m i s e , a s

we l l . D i f f e r ences i n pa t t e rn s can r e f l ec t i nd iv idua l d i f f e r ences i n cha rac te r

t r a i t s and d i f f e r ences in consc ious s t a t e s . To the end o f exp lo i t i ng these

poss ib il i ti e s , my l ab o ra to ry was r e cen t l y f i t t ed w i th a 128 -e l ec t rode reco rd -

ing capab i l i t y . I n add i t i on , my co l l eagues and I have dev i sed seve ra l new

m e t h o d s f or q u a n t i f y i n g t h e s p a t i o t e m p o r a l d y n a m i c s o f E E G . D e v el op -

m e n t o f t h e s e m e t h o d s w a s m o t i v a t e d b y w a t c h i n g c o m p u t e r - g e n e r a t e d

a n i m a t i o n s o f E E G v o l t a g e r e c or d i n g s. T h e s e a n i m a t i o n s c o n t a i n a w e a l t h

o f i n f o r m a t i o n a b o u t t h e r a p i d i t y ( a b o u t 1 00 p e r s e c on d ) o f c h a n g e i n t h e

pa t t e rn s o f vo l t ages ob se rved ac ros s the su r f ace o f the s calp . W e quan t i f i ed

t h e s e s p a t i o - t e m p o r a l d y n a m i c s a s s c a l a r s , v e c t o r s , a n d c l u s t e r a n a l y t i c


26/49


p l ot s o f E E G a c t i v i ty a n d h a v e o b t a i n e d i n i t i a l f in d i n g s s u g g e s t i n g t h a t t h e

t ech n iqu es w i ll p rove use fu l ( P r ib r am e t a l. , 19 96).

Them e V: Theore ti ca l In t e rpre ta t ions o f the Research Resu l t s

T h e l a b o r a t o r y r e s e a r c h h a s y i e l d e d m a n y u n e x p e c t e d r e s u l t s . T h e s e r e -

s u l t s h a v e d r a m a t i c a l l y c h a n g e d m y v ie w s f ro m t i m e t o t i m e a n d p o s e d , as

c r i t i ca l t o fu r t h e r r e sea rch , p ro b lems wh ich I h ad t h o ugh t I co u ld igno re .

M u c h o f t h e t h e o r e ti c a l w o r k w h i ch h a s e n g a g e d m e h a s s t e m m e d f r om

t h e s e s u r p r i s e s .

D i s c o v e r i e s

K a r l P o p p e r c l a i m s t h a t s c ie n c e is b a s e d o n c o n je c t u r e a n d r e f u t a t i o n , a n d

K a r l L a s h l e y w a s a l w a y s c o m f o r t a b l e w h e n h e o p e r a t e d i n t h i s m o d e . M y

o w n r e s e a r c h h a s p r o c e e d e d i n a m o r e h a p h a z a r d f a s h i o n ( s e e P r i b r a m ,

1 982). Desp i t e t h e p lann ing re p re sen t ed by t h e t h em es desc r ibed ear l ie r , m y

a c t u a l r e s e a r c h h a s b e e n a s e a r c h w h ic h s t e m m e d f r om p r o b l em s a n d p a r a -

d o x es ( s u c h a s u n e x p e c t e d l y f i n d i n g r e l a ti v e l y d i r e c t s e n s o r y i n p u t s t o t h e

m o t o r c o rt ex ) r a t h e r t h a n f r o m w e l l - f o r m u l a t e d c o n j e c tu r e s o r h y p o t h e s e s .

T h e s e s t h e r e w e r e , b u t o n ly r a r e l y d i d I d e r i v e s in g l e, t e s t a b l e h y p o t h -

eses w i t h ex pe r im en t s d es igned t o co nf i rm o r d i sconf i rm . Ra t h e r , I fo llowed

t h e r u l e t h a t s e v e r a l m o r e o r l e s s c l e a r l y d e f i n e d a l t e r n a t i v e s p r e s e n t e d

t h e m s e l v e s w h e n t h e t h e s i s , t h a t i s , t h e r e a s o n s f o r p e r f o r m i n g t h e r e -

sea rch , bec am e c lea r . I des igned e xp er im en t s t o f ind ou t wh ich o f t h e a l t e r -

n a t i v e s f it t h e d a t a I h a d o b t a in e d . S o m e t i m e s t h e d a t a f it n o n e o f t h e

a l t e r n a t i v e s , t h e t h e s i s i t s e lf w a s f o u n d w a n t i n g , a n d n e w d i r e c ti o n s h a d t o

b e ta k e n . O f t en t h e s e n e w d i re c ti on s s t e m m e d f r om a t t e m p t s t o s y s t em a -

t i z e t h e d a t a a l r e a d y o b t a i n e d a n d t o d e v e l o p a n a p p r o p r i a t e f r a m e f o r

s o r t i n g a n d c l a ss i fy i n g th e m .

W h a t e v e r t h e m e r i t s o r d e f ic ie n ci es o f t h i s a p p r o a c h , i t i s s h a r e d b y

m a n y b i o l o g i s t s . C l a u d e B e r n a r d , w h e n a s k e d h o w h e p r o c e e d e d i n t h e

l a b o r a t o r y , a n s w e r e d t h a t h e s i m p l y a s k e d n a t u r e s o m e q u e s t i o n s . B y

a d o p t i n g t h i s p e r s p e c t iv e , t h e y i e ld o f m y r e s e a r c h h a s b e e n s u b s t a n t i a l

a n d I m a d e m a n y d is co v er ie s w h i c h m i g h t n o t h a v e b e e n u n c o v e r e d b y a

m o re r ig id app ro ach . So me o f t h ese d i sco ver ies a re de t a i l ed be lo w .

The Fu nc t ion s o f t he Fron to l imb ic Forebra in

T h e L i m b i c F o r e b r a in .

E a r l y r e s e a r c h r e s u l t s l e d m e t o r e d e f i n e t h e

bo u nd ar i e s o f t h e l imb ic fo reb ra in ( a lso ca ll ed th e o l fact o ry b ra in ) w h ich

h a d h i t h e r t o i n c l u d e d o n l y t h e h i p p o c a m p a l a n d c i n g u l a t e g y r i b y e s ta b l i s h -

i n g t h e r e l a t i o n s h i p b e t w e e n l i m b i c c o r t e x a n d v i s c e r o a u t o n o m i c a c t i v i t y

( see P r i b r am & Krug er , 1 954).


27/49

Karl H. Pribram

327

B a s e d o n t h e e a r l i e r w o r k o f W a r r e n M c C u l lo c h, P e r c i v a l B a il ey , a n d

G e r h a r d t v o n B o n i n , I e s t a b l i s h e d b y s t r y c h n i n e n e u r o n o g r a p h y a n d b y

e l e c t r i c a l s t i m u l a t i o n a n d h i s t o l o g i c a l e x a m i n a t i o n , t h e i n t e r r e l a t i o n s h i p

b e t w e e n t h e a m y g d a l o id c o m p le x a n d t h e s u r r o u n d i n g o r b it of r on t a l, a n t e -

r io r in su la r , an d t em po ra l p o la r co r tex and t h e d i rec t co nnec t io ns o f a l l o f

t h e s e to th e h y p o t h a l a m u s ( se e P r i b r a m e t al., 1 95 0; M a c L e a n a n d P r i b r a m ,

1 95 3 ; P r i b r a m a n d M a c L e a n , 1 95 3) .

T h e w o r k o f A r t h u r W a r d a n d R o b e r t L i v i n g s to n h a d s h o w n t h a t v i sc er o-

a u t o n o m i c r e s p o n s e s w e r e o b t a i n e d f r o m e l e c t r ic a l s t i m u l a t i o n o f t h e c i n-

g u l a t e g y r u s a n d o r b i t of r o n t a l c or te x . W i t h B. R . K a a d a a n d J . A. E p s t e i n

( see Kaada e t a l . , 1 9 49 ) I ex t ended t h ese r esu l t s t o t h e an t e r io r i n su la ,

t e m p o r a l p o l e , a n d a m y g d a l a . I n i t i a l l y , t h i s f i n d i n g w a s r e s i s t e d a s b e i n g

d u e to a r t i f a c t ~ a N o b el l a u r e a t e i n d i c a t e d to J o h n F u l t o n t h a t h e th o u g h t

o u r r e s u l t s w e r e d u e to i n a d v e r t e n t s t i m u l a t i o n o f t h e d u r a ~ a f t e r a ll, w e

k n e w t h a t t h e h y p o t h a l a m u s w a s t h e " h e a d g a n g li on " of t h a t s y s te m . F u l -

t o n s t u c k b y m e a n d p u b l i s h e d o u r f i n d in g s . W i t h i n 2 y e a r s , m o s t o f t h e

g r a d u a t e s t u d e n t s i n p h y s io l og y a t Y ale w e r e d o i n g th e i r t h e s e s o n l im b i c -

re la ted topics .

T h u s , t he a m y g d a l a a n d i t s s u r r o u n d i n g c o rt ex w e r e s h o w n to b e p a r t

o f t h e l imb ic fo reb ra in , wh ich , a s no t ed abo ve , h ad p rev io us ly inc luded o n ly

t h e h i p p o c a m p a l a n d c i n g u l a te s y s t e m s . F u r t h e r , a n e n t i r e e x t e n t o f m e -

d i o b as a l m o t o r c o r t e x ~ w h i c h i n c lu d e d t h e p e r i a m y g d a l o i d c or te x , t h e t e m -

p o r a l a n d t h e a d j a c e n t a n t e r i o r i n s u l a r , t h e o r b i t o f r o n t a l , m e d i a l f r o n t a l

a n d a n t e r i o r c i n g u l a te c o r t e x ~ w a s d i sc o v er e d w h o s e p r i m a r y f u n c ti o n is

t o r e g u l a t e v i s c e r o a u t o n o m i c f u n c ti o n s ( P r i b r a m , 1 96 1) .

T h e A n t e r i o r F r o n t a l C o r t e x a n d L i m b i c F o r e b r a i n . N e x t I

e s t a b -

l i sh ed t h e f ac t t h a t t h e f a r f ro n t a l co r t ex is t h e "asso c ia tio n" co r t ex fo r t h e

l imbic fo reb ra in . T h i s ac co un t ed fo r t h e psych o su rg ica l e f fec ts o f f ro n t a l

lo b ot om y . U s i n g t h e d e l a y e d r e s p o n s e a n d d e l a y e d a l t e r n a t i o n t e c h n i q u e s I

e x t e n d e d t h e w o r k o f C a r l y le J a c o b s e n a n d H e n r y N i s se n , w h o h a d s h o w n

t h a t r e s e c t io n s o f f a r f r o n t a l c o r te x d i s r u p t e d p e r f o r m a n c e o n t h e s e t a s k s . I

f o u n d t h a t r e s e c t io n s o f t h e v a r i o u s s t r u c t u r e s c o m p o s i n g t h e l im b i c fo re -

b r a i n ( h i p p o c a m p u s , a m y g d a l a , c i n g u l a t e c o r t e x ) a l s o d i s r u p t e d p e r f o r -

m ance o f de layed a l t e rna t io n (P r ib r am e t a l. , 19 62). By co n t ras t , r e sec t io ns

o f t h e co r t ex o f t h e po s t e r io r ce reb ra l co nv ex i t y f a i l ed t o d i s ru p t pe r fo rm -

the history of neuroscience in autobiography - vol 2

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